Biology subjects

Bugalho, M. N.

Publications and source records attributed to Bugalho, M. N..

8 recordsLinked to original sources

bioRxiv2026

Nutrient enrichment and herbivore exclusion disrupt the climate-driven balance between C3 and C4 plants in grasslands

The distribution of plants with different photosynthetic pathways is strongly structured by climate, with C3 plants favoured in cooler temperate regions and C4 plants in hotter, high-light conditions. The relative abundance of C3 and C4 plants across the world has cascading impacts on local food webs, decomposition, productivity and other vital ecosystem processes. Human impacts, including climate change, changes to herbivore assemblages, and increased nutrient availability, are shifting the optimal conditions for important C3 and C4-dominated ecosystems and crops. Using 3,184 plot-level observations from 112 sites across six continents, we reveal how chronic nutrient enrichment disrupts the climate-driven balance between C3 and C4 plants in grasslands. We found that, consistent with expectations, the global distribution of C4 plants was strongly related to climate. However, experimental nutrient addition reduced the relative cover of C4 species, with the strongest declines found when nitrogen and phosphorus were added together. Herbivore exclusion had no consistent effect on C4 plants. Our results provide global experimental evidence that elevated nutrients, particularly nitrogen, alter competitive outcomes among plant functional types to suppress C4 grasses, even in climatically optimal conditions. This has major implications for predicting vegetation responses to global change, with consequences for carbon cycling, primary productivity, herbivore dynamics, and food security.

ecology↗

bioRxiv2025

Carnivore-mediated seed crossings are more likely on roadkill hotspots

Roadkill hotspots concentrate animal movement and mortality, but whether they also interfere with animal-mediated seed dispersal--a conservation-relevant process underpinning plant regeneration and landscape connectivity--remains unknown. In Mediterranean oak woodlands of southern Portugal, we used seed mimics to test how road type (paved vs. unpaved) and road-forest context (edge vs. non-edge) shape carnivore-mediated seed crossings, and whether crossings coincide with roadkill hotspots detected by kernel density analysis. Bayesian logistic models indicated that seed-crossing probability was about twice as high on unpaved as on paved roads (predicted means 0.28 vs. 0.14), with weaker evidence for a negative edge effect. Crossing probability also tended to increase with carnivore abundance and distance to streams, and decrease with rodent density, albeit with some uncertainty. Crucially, paved sections intersecting roadkill hotspots showed nearly threefold higher predicted crossing probabilities than non-hotspot sections (0.51 vs. 0.18). These results yield clear conservation implications: prioritize paved segments that overlap carnivore roadkill hotspots. Implement precision measures--short-segment speed management, directional fencing that funnels animals to existing culverts/underpasses, and verge management that removes carcasses and limits small-mammal/scavenging attractants--to cut mortality while maintaining seed-mediated connectivity. A light, repeatable monitor-act-reassess workflow (hotspot mapping + seed-mimic trials + camera checks) enables rapid deployment and evaluation, and is transferable to other human-dominated forest mosaics. Hotspot-guided mitigation offers a pragmatic path to reconcile mobility infrastructure with living connectivity.

ecology↗

bioRxiv2025

Roads shape seed dispersal by rodents and carnivores at forest edges

The influence of road networks on seed dispersal remains poorly understood. We conducted a field experiment in Mediterranean oak woodlands of southern Portugal to assess how road type (paved vs. unpaved) and road-forest context (edge vs. non-edge) shape seed dispersal by rodents and carnivores. Using labeled acorns and seed mimics, we tracked dispersal distances, number of dispersals, road crossings, and dispersal directions. Rodents dispersed seeds farther in forest edges and along paved roadsides, predominantly moving them parallel to roads, but rarely across--although crossings were more frequent on unpaved roads. In contrast, carnivores mediated long-distance dispersal, primarily perpendicular to roads, and although seed road crossings were rare, carnivore-dispersed seeds crossed roads--especially unpaved ones--nine times more frequently than rodent-dispersed seeds. Shrub cover increased rodent-mediated dispersal events, heavier acorns traveled farther, and water patches decreased carnivore-mediated dispersal events near roads. These findings highlight the dual role of roads as barriers and of roadsides as corridors for seed dispersal, with implications for forest regeneration and landscape connectivity. Roadside management should prioritize moderate shrub cover along paved roads to support rodent dispersal while balancing ecological benefits with potential trade-offs. Unpaved roads, in turn, should be managed as functional corridors for carnivores, enhancing connectivity between fragmented forests. By integrating species-specific dispersal processes into road planning, landscapes can be designed to minimize road barrier effects while promoting natural regeneration in human-modified forests.

ecology↗

bioRxiv2024

Meta-analysis of livestock effects on tree regeneration in oak agroforestry systems

Livestock grazing occupies over a quarter of terrestrial land and is prevalent to agroforestry ecosystems, potentially affecting the survival, growth, and density of trees early developmental stages, such as seeds, seedlings, and saplings. To address the effects of livestock on tree recruitment in the face of ongoing debates about their impacts, we conducted a 33-year meta-analysis in Quercus-dominated agroforestry systems. Our analysis revealed a consistently negative effect of livestock on oak acorns, seedlings, and saplings. Significantly, livestock body size influenced oak regeneration, with small-sized livestock, notably sheep and goats, having a more pronounced negative impact compared to mixed-size systems, mainly involving cattle and sheep. The effects of small-sized livestock were markedly detrimental on acorn survival and seedling/sapling density, although no studies eligible for meta-analysis examined large livestock impacts on acorns. Overall, mixed-size livestock systems, often involving cattle and sheep, lessen the negative effects. Our findings indicate that the body size and foraging behaviors of livestock should be considered for the ecological sustainability of the tree component in agroforestry systems. While protective measures have long been integral to well-managed agroforestry systems, our results underscore the importance of integrating diverse livestock sizes and applying specific protective strategies, particularly for acorns and saplings, to further refine these practices. Future research should expand to underrepresented regions and livestock types to refine global agroforestry management practices. HighlightsO_LILivestock reduces oak tree regeneration at acorn, seedling, and sapling stages. C_LIO_LISmall livestock effect more negative, especially acorn survival and young oak density. C_LIO_LIMixed-size livestock agroforestry systems reduce harm more than small single-size ones. C_LIO_LIWe challenge perceptions that replacing cattle with sheep is necessarily less harmful. C_LIO_LIWe advocate for diverse livestock sizes in Quercus-dominated agroforestry systems. C_LI

ecology↗

bioRxiv2023

Multidimensional responses of ecological stability to eutrophication in grasslands

Eutrophication usually impacts biodiversity, species composition, and functioning of grassland communities. Whether such effects propagate to influence the stability of these community aspects is unknown. Using standardized experiments across 55 global grasslands, we quantified the effects of nutrient addition on five stability facets (i.e., temporal invariability and resistance during and recovery after dry and wet growing seasons) for three community aspects (i.e., aboveground biomass, community composition, and species richness). Nutrient addition reduced the temporal invariability and resistance of species richness and community composition, but not biomass, during dry and wet growing seasons. Temporal invariability and resistance during, but not recovery after, dry and wet growing seasons were strongly positively correlated in both ambient and eutrophic conditions. This indicates that maintaining and restoring the stability of plant communities requires increasing resistance rather than recovery. Harnessing the complexity of ecological stability provides new insights for grassland ecosystem sustainability in a changing world.

ecology↗

bioRxiv2023

Cattle grazing alters how cork oak crop failure leads to faster and farther rodent-mediated acorn dispersal

Masting, the synchronized production of variable quantities of seeds, occurs in many grazed systems and can be interspersed with years of extreme crop failure, whose frequency and unpredictability are increasing with climate change. Yet, the combined impact of crop failure and grazing on seed dispersal and seed-to-seedling transition remains poorly understood. To address this concern, we investigated rodent-mediated cork-oak (Quercus suber) acorn predation, dispersal, and seedling emergence in cattle grazed and non-grazed areas in central Portugal during years with contrasting masting seasons. We found that extreme crop failure led to six times longer and faster acorn dispersal, with 83% more dispersal events than during a year of reproductive success. The percentage of predated acorns also increased by 84%. However, the higher acorn predation was offset by a 2.4-fold higher percentage of unpredated dispersed acorns recruiting into seedlings. Both years ended up recruiting a similar number of seedlings. Acorns emerged seedlings 3.4 times farther in the crop failure year than in the crop success year. Cattle grazing was the main constraint on seed dispersal distance, reducing it by 51% during the extreme crop failure year, while having no noticeable effect during the successful crop year. Our study provides empirical evidence that cattle grazing modulates how an extreme crop failure year can surprisingly be an opportunity for trees remaining fecund to have seedlings established farther apart than in a crop success year. If we are to better manage and preserve the high conservation and socio-economic value of Mediterranean cork oak woodlands in the face of climate change, we must prioritize fecund trees and carefully manage seed dispersal factors such as cattle grazing, particularly during years of crop failure.

ecology↗

bioRxiv2021

Multiple resource limitations explain biomass-precipitation relationships in grasslands

Interannual variability in grassland primary production is strongly driven by precipitation, nutrient availability and herbivory, but there is no general consensus on the mechanisms linking these variables. If grassland biomass is limited by the single most limiting resource at a given time, then we expect that nutrient addition will not affect biomass production at arid sites. We conducted a distributed experiment manipulating nutrients and herbivores at 44 grassland sites in 8 regions around the world, spanning a broad range in aridity. We estimated the effects of 5-11 years of nutrient addition and herbivore exclusion treatments on precipitation sensitivity of biomass (proportional change in biomass relative to proportional change in rainfall among years), and the biomass in the driest year (to measure treatment effects when water was most limiting) at each site. Grazer exclusion did not interact with nutrients to influence driest year biomass or sensitivity. Nutrient addition increased driest year biomass by 74% and sensitivity by 0.12 (proportional units), and that effect did not change across the range of aridity spanned by our sites. Grazer exclusion did not interact with nutrients to influence sensitivity or driest year biomass. At almost half of our sites, the previous year's rainfall explained as much variation in biomass as current year precipitation. Overall, our distributed fertilization experiment detected co-limitation between nutrients and water governing grasslands, with biomass sensitivity to precipitation being limited by nutrient availability irrespective of site aridity and herbivory. Our findings refute the classical ideas that grassland plant performance is limited by the single most limiting resource at a site. This suggests that nutrient eutrophication will destabilize grassland ecosystems through increased sensitivity to precipitation variation.

ecology↗

bioRxiv2018

Unravelling associations between tree seedling performance, herbivory, competition, and facilitation in high nature value farmlands

Herbivory, plant facilitation, and competition have complex impacts on tree regeneration which are seldom investigated together. Grazing exclosure experiments have allowed to quantify the effects of large herbivores on tree regeneration dynamics but have often ignored the effect of herbivorous insects. We experimentally tested how folivory (percentage of leaf damaged by insects), and microenvironment (tree-canopy cover and herbs) jointly alter performance (growth and survival) of seedlings of two dominant Mediterranean oak-species within ungulate exclosures. An agroforestry system dominated by cork oak (Quercus suber) and holm oak (Q. rotundifolia) was assessed in south-east Portugal. We aimed also to determine whether the two oak-species differ in the interdependences between folivory, microenvironment, covaring factors, and seedling performance. Unexpectedly, under the low-moderate insect defoliation occurred in our 3-year field study, growth and survival of cork and holm oak-seedlings were positively associated with herbivory damage. Herb removal increased oak folivory by 1.4 times. Herb removal was also positively associated with growth, directly and indirectly through its negative effect on oak folivory. Tree-canopy favored insect folivory upon cork oak seedlings directly and upon holm oak indirectly via decreasing light availability. Folivory was threefold greater upon cork than upon holm oak-seedlings. Our study shows that tree-canopy, herbs, and covarying factors can affect cork and holm oak-seedling performances through complex pathways, which markedly differ for the two species. The combined effect of insect herbivory and positive and negative plant-plant interactions need to be integrated into future tree regeneration efforts toward tackling the regeneration crisis of oak agroforestry systems of the Mediterranean.

ecology↗