Prof. Dr. Catherine Graham

The presence in ecological communities of unfeasible species interactions, termed forbidden links, due to physiological or morphological exploitation barriers has been long debated, but little direct evidence has been found. Forbidden links are likely to make ecological communities less robust to species extinctions, stressing the need to assess their prevalence. Here, we used a dataset of plant–hummingbird interactions, coupled with a Bayesian hierarchical model, to assess the importance of exploitation barriers in determining species interactions. We found evidence for exploitation barriers between flowers and hummingbirds across the 32 studied communities; however, the proportion of forbidden links changed drastically among communities because of changes in trait distributions. The higher the proportion of forbidden links, the more they decreased network robustness because of constraints on interaction rewiring. Our results suggest that exploitation barriers are not rare in plant–hummingbird communities and have the potential to limit the rescue of species experiencing partner extinction.

Ecological and evolutionary questions addressing diversity-environment relationships have been evaluated almost entirely in geographic space, yet most hypotheses are formulated in terms of environmental conditions. Recent examples evaluating macroecological patterns directly in environmental space suggest that such refocusing provides different perspectives on the mechanisms driving broad-scale patterns of diversity. Yet, we lack both conceptual frameworks and targeted studies to fully evaluate the potential contribution of such a refocus. Here, we focus on the concept of environmental space by briefly reviewing its use in ecology and evolution and suggesting avenues for further development. We encourage a re-evaluation of hypotheses and frameworks that have dominated ecological theory since the foundations of ecology with a very simple shift in the lens, that is, from geographical to environmental space. Focusing on environmental space also provides a crucial lens for climate change research, enabling a comprehensive evaluation of biodiversity dynamics and offering a holistic view of the interplay between species and their evolving environments. This shift enhances our ability to predict and adapt to future changes, enriching our understanding of biodiversity beyond more commonly done geographic analyses.

Pollinator specialization, crucial for the stability of plant-pollinator interactions, can be affected by habitat loss and fragmentation; however, the pathways that induce these changes are poorly understood. We aim to understand the underlying factors by which habitat loss, fragmentation, and vegetation structure change species specialization of hummingbirds and consequently influence community robustness in tropical montane forests of southern Ecuador. We used a piecewise structural equation model to simultaneously analyze the influence of functional traits of pollinators and resource availability as pathways driving the changes in specialization, resulting from an increase in habitat loss, fragmentation, and simplification of vegetation structure. We found that fragmentation reduces hummingbird specialization by filtering out hummingbirds with long bills. In addition, we observed that forest edges have greater floral diversity associated with a decrease in hummingbird specialization. The observed shift towards generalization in hummingbird assemblages, related to the transformation of landscapes, appears to promote more robust communities capable of sustaining hummingbird populations. However, morphologically specialized pollinators can be highly sensitive to fragmentation and even become locally extinct when fragmentation increases in a landscape. We conclude that while fragmented habitats may maintain robust communities, specialist species, often with unique ecological functions, may be lost.

Tropical mountains often harbour high species richness. Yet the mechanisms behind such high richness remain poorly understood. One prominent hypothesis for high species richness is niche partitioning, which reduces competition and promotes coexistence. Here, we evaluate niche structure and specialisation of plant species based on the floral traits related to pollination interactions across an elevational gradient in the northern Andes. Niche structure can vary among sites, either expanding or contracting, or becoming more or less packed. We sampled eleven communities of hummingbird-visited plants along an elevation gradient in the Ecuadorian Andes and measured a series of functional traits associated with hummingbird pollination. We used the traits of all co-occurring species to calculate community weighted means, functional richness, and mean nearest neighbour distances and evaluated how they varied across elevation. Additionally, we measured specialisation based on plant–hummingbird interaction records to assess if packing is associated with narrower resource use or greater niche overlap. Species and functional richness were constant along the elevation gradient; however, niche packing was stronger at mid-elevation. We found changes in network specialisation, where the least specialised communities were those with higher niche packing. These results suggest that traits related to pollination and plant reproduction help to explain species co-occurrence and specialisation.

Plant–hummingbird interactions are considered a classic example of coevolution, a process in which mutually dependent species influence each other's evolution. Plants depend on hummingbirds for pollination, whereas hummingbirds rely on nectar for food. As a step towards understanding coevolution, this review focuses on the macroevolutionary consequences of plant–hummingbird interactions, a relatively underexplored area in the current literature. We synthesize prior studies, illustrating the origins and dynamics of hummingbird pollination across different angiosperm clades previously pollinated by insects (mostly bees), bats, and passerine birds. In some cases, the crown age of hummingbirds pre-dates the plants they pollinate. In other cases, plant groups transitioned to hummingbird pollination early in the establishment of this bird group in the Americas, with the build-up of both diversities coinciding temporally, and hence suggesting co-diversification. Determining what triggers shifts to and away from hummingbird pollination remains a major open challenge. The impact of hummingbirds on plant diversification is complex, with many tropical plant lineages experiencing increased diversification after acquiring flowers that attract hummingbirds, and others experiencing no change or even a decrease in diversification rates. This mixed evidence suggests that other extrinsic or intrinsic factors, such as local climate and isolation, are important covariables driving the diversification of plants adapted to hummingbird pollination. To guide future studies, we discuss the mechanisms and contexts under which hummingbirds, as a clade and as individual species (e.g. traits, foraging behaviour, degree of specialization), could influence plant evolution. We conclude by commenting on how macroevolutionary signals of the mutualism could relate to coevolution, highlighting the unbalanced focus on the plant side of the interaction, and advocating for the use of species-level interaction data in macroevolutionary studies.

Land use intensification favours particular trophic groups which can induce architectural changes in food webs. These changes can impact ecosystem functions, services, stability and resilience. However, the imprint of land management intensity on food-web architecture has rarely been characterized across large spatial extent and various land uses. We investigated the influence of land management intensity on six facets of food-web architecture, namely apex and basal species proportions, connectance, omnivory, trophic chain lengths and compartmentalization, for 67,051 European terrestrial vertebrate communities. We also assessed the dependency of this influence of intensification on land use and climate. In addition to more commonly considered climatic factors, the architecture of food webs was notably influenced by land use and management intensity. Intensification tended to strongly lower the proportion of apex predators consistently across contexts. In general, intensification also tended to lower proportions of basal species, favoured mesopredators, decreased food webs compartmentalization whereas it increased their connectance. However, the response of food webs to intensification was different for some contexts. Intensification sharply decreased connectance in Mediterranean and Alpine settlements, and it increased basal tetrapod proportions and compartmentalization in Mediterranean forest and Atlantic croplands. Besides, intensive urbanization especially favoured longer trophic chains and lower omnivory. By favouring mesopredators in most contexts, intensification could undermine basal tetrapods, the cascading effects of which need to be assessed. Our results support the importance of protecting top predators where possible and raise questions about the long-term stability of food webs in the face of human-induced pressures.

Forests are under pressure and going through rapid changes. However, current inventorying and monitoring (IM) programs are often either disjointed, too narrow in their scope and/or do not operate at fine enough temporal resolutions, which may hinder scientific understanding, the timely supply of information, fast decision making, and may result in the sub-optimal use of resources. For these reasons, there is an urgent need for Advanced Forest Inventorying and Monitoring (AIM) programs to (i) achieve expanded relevance (by augmenting data/information across ecosystem properties and trophic levels), (ii) have increased temporal resolution (by tailored data collection frequency), and (iii) make use of technological advances (by incorporating novel tools and technologies). The Advanced Inventorying and Monitoring for Swiss Forests (SwissAIM) initiative was launched in 2020 to address these needs. SwissAIM builds upon the foundation offered by the existing programs (e.g., national forest inventory, long-term forest ecosystem research, biodiversity monitoring). It aims to offer a collaborative and adaptive framework to enable integrated data collection, evaluation, interpretation, analysis, and modeling. Ideally, it will result in a more responsive system with respect to current and predicted biotic/abiotic stressors that will challenge Swiss forests. Developing such a system implies identifying the information needs of different stakeholders (e.g., science, policy, practice), related technical requirements, and governance frameworks. Here, we present (i) the main features of the SwissAIM initiative (vision, scientific questions and variables, governance and engagement), (ii) the main outcomes of the participatory design process (measurements, sampling, and plot design), (iii) the potential transferability of AIM initiatives outside Switzerland (timing, relevance, practicability), and (iv) the key messages that emerged (i.e., need for advancement, integration and transdisciplinarity, statistical underpinning). Since similar needs related to forest inventorying and monitoring are emerging throughout Europe and elsewhere, the objective of this opinion paper is to share our experience and promote a dialog with those interested in developing AIM initiatives in other countries and regions.

Rising temperatures are leading to increased prevalence of warm-affinity species in ecosystems, known as thermophilisation. However, factors influencing variation in thermophilisation rates among taxa and ecosystems, particularly freshwater communities with high diversity and high population decline, remain unclear. We analysed compositional change over time in 7123 freshwater and 6201 terrestrial, mostly temperate communities from multiple taxonomic groups. Overall, temperature change was positively linked to thermophilisation in both realms. Extirpated species had lower thermal affinities in terrestrial communities but higher affinities in freshwater communities compared to those persisting over time. Temperature change’s impact on thermophilisation varied with community body size, thermal niche breadth, species richness and baseline temperature; these interactive effects were idiosyncratic in the direction and magnitude of their impacts on thermophilisation, both across realms and taxonomic groups. While our findings emphasise the challenges in predicting the consequences of temperature change across communities, conservation strategies should consider these variable responses when attempting to mitigate climate-induced biodiversity loss.

Species traits greatly influence interactions between plants and pollinators where floral nectar is the primary energy source fostering this mutualism. However, very little is known about how nectar traits mediate interactions in pollination networks compared with morphological traits. Here, we evaluated the role of morphological and nectar traits in shaping plant–hummingbird interaction networks along an elevation gradient. For this, we assessed patterns in floral phenotypic traits and network properties of plant species across elevations in Costa Rica. We also analysed whether plant species with generalized flower traits are ecological generalists and how morphological trait matching versus nectar traits affect interactions. We found marked variation in floral phenotypic traits and flower abundance of hummingbird-visited plant species across 10 sites along the elevation gradient. We did not find evidence for a relationship between flower morphology and nectar traits or between morphological and ecological generalization of plant species. Plant–hummingbird interaction frequency increased when the lengths of hummingbird bill and flower corolla were similar, indicating morphological matching, whereas nectar traits were unrelated to interactions. While nectar may play a difficult-to-detect secondary role within plant–hummingbird networks, our results reinforce the idea that morphological matching is an important factor in structuring ecological communities.

Europe's semi-natural grasslands support notably high levels of temperate biodiversity across multiple taxonomic groups. However, these ecosystems face unique conservation challenges. Contemporary agricultural practices have replaced historical traditional low-intensity agriculture in many regions, resulting in a spectrum of management intensities within these ecosystems, ranging from highly intensive methods to complete abandonment. Paradoxically, both extremes along this spectrum of management intensity can be detrimental to biodiversity of semi-natural grasslands. Moreover, while anthropogenic climate change is an overarching threat to these ecosystems, rapid changes in land use and its intensity often present more immediate pressures. Often occurring at a faster rate than climate change itself, these land-use changes have the potential to rapidly impact the biodiversity of these grasslands. Here, we divide the ecological processes, threats, and developments to semi-natural grasslands into three sections. First, we examine the different impacts of agricultural intensification and abandonment on these ecosystems, considering their different consequences for biodiversity. Second, we review seminal works on various evidence-based management practices and offer a concise summary that provides support for various conservation and management strategies. However, the socio-economic factors that drive both abandonment and intensification in semi-natural grasslands can also be used to develop solutions through strategic governmental and non-governmental interventions. Accordingly, we conclude with a way forward by providing several key policy recommendations. By synthesizing existing knowledge and identifying research gaps, this essay aims to provide valuable insights for advancing the sustainable management of semi-natural grasslands.

Human land-use intensification threatens arthropod (for example, insect and spider) biodiversity across aquatic and terrestrial ecosystems. Insects and spiders play critical roles in ecosystems by accumulating and synthesizing organic nutrients such as polyunsaturated fatty acids (PUFAs). However, links between biodiversity and nutrient content of insect and spider communities have yet to be quantified. We relate insect and spider richness to biomass and PUFA-mass from stream and terrestrial communities encompassing nine land uses. PUFA-mass and biomass relate positively to biodiversity across ecosystems. In terrestrial systems, human-dominated areas have lower biomass and PUFA-mass than more natural areas, even at equivalent levels of richness. Aquatic ecosystems have consistently higher PUFA-mass than terrestrial ecosystems. Our findings reinforce the importance of conserving biodiversity and highlight the distinctive benefits of aquatic biodiversity.

How traits affect speciation is a long-standing question in evolution. We investigate whether speciation rates are affected by the traits themselves or by the rates of their evolution, in hummingbirds, a clade with great variation in speciation rates, morphology and ecological niches. Further, we test two opposing hypotheses, postulating that speciation rates are promoted by trait conservatism or, alternatively, by trait divergence. To address these questions, we analyse morphological (body mass and bill length) and niche traits (temperature and precipitation position and breadth, and mid-elevation), using a variety of methods to estimate speciation rates and correlate them with traits and their evolutionary rates. When it comes to the traits, we find faster speciation in smaller hummingbirds with shorter bills, living at higher elevations and experiencing greater temperature ranges. As for the trait evolutionary rates, we find that speciation increases with rates of divergence in the niche traits, but not in the morphological traits. Together, these results reveal the interplay of mechanisms through which different traits and their evolutionary rates (conservatism or divergence) influence the origination of hummingbird diversity.

Quantifying the vulnerability of ecosystems to global change requires a better understanding of how trophic ecosystem functions emerge. So far, trophic ecosystem functions have been studied from the perspective of either functional diversity or network ecology. To integrate these two perspectives, we propose the interaction functional space (IFS) a conceptual framework to simultaneously analyze the effects of traits and interactions on trophic functions. We exemplify the added value of our framework for seed dispersal and wood decomposition and show how species interactions influence the relationship between functional trait diversity and trophic functions. We propose future applications for a range of functions where the IFS can help to elucidate mechanisms underpinning trophic functions and facilitate understanding of functional changes in ecosystems amidst global change.

In animal-pollinated plants, fitness is affected by pollinator attraction and plant-pollinator trait matching. Plant-pollinator interactions are often characterized using metrics derived from mutualistic networks, such as centrality and interaction selectivity. This approach often focuses on the species level, thus neglecting significant among-individual variation in attracting foraging pollinators and associated fitness consequences for plants. Here, we tested if centrality and interaction selectivity were related to fitness at the species and the individual level for 14 hummingbird-pollinated plant species in the Atlantic Forest. In addition, we tested if plant-pollinator trait-matching influenced plant fitness. At the species-level, we did not find a relationship between plant fitness and centrality nor interaction selectivity; we found instead that plant fitness increased with higher trait-matching between bill and corolla lengths. At the individual-level we found that individuals with higher centrality values produced less seeds, likely due to lower quality in pollen deposition. Moreover, individuals with a lower interaction selectivity also produced more seeds, probably because of higher conspecific pollen flow. Our results show that individual position and interaction selectivity within pollination networks may have important implications for pollination effectiveness and hence plant fitness.

Landscape dynamics are widely thought to govern the tempo and mode of continental radiations, yet the effects of river network rearrangements on dispersal and lineage diversification remain poorly understood. We integrated an unprecedented occurrence dataset of 4,967 species with a newly compiled, time-calibrated phylogeny of South American freshwater fishes-the most species-rich continental vertebrate fauna on Earth-to track the evolutionary processes associated with hydrogeographic events over 100 Ma. Net lineage diversification was heterogeneous through time, across space, and among clades. Five abrupt shifts in net diversification rates occurred during the Paleogene and Miocene (between 30 and 7 Ma) in association with major landscape evolution events. Net diversification accelerated from the Miocene to the Recent (c. 20 to 0 Ma), with Western Amazonia having the highest rates of in situ diversification, which led to it being an important source of species dispersing to other regions. All regional biotic interchanges were associated with documented hydrogeographic events and the formation of biogeographic corridors, including the Early Miocene (c. 23 to 16 Ma) uplift of the Serra do Mar and Serra da Mantiqueira and the Late Miocene (c. 10 Ma) uplift of the Northern Andes and associated formation of the modern transcontinental Amazon River. The combination of high diversification rates and extensive biotic interchange associated with Western Amazonia yielded its extraordinary contemporary richness and phylogenetic endemism. Our results support the hypothesis that landscape dynamics, which shaped the history of drainage basin connections, strongly affected the assembly and diversification of basin-wide fish faunas.

Speciation rates vary greatly among taxa and regions and are shaped by both biotic and abiotic factors. However, the relative importance and interactions of these factors are not well understood. Here we investigate the potential drivers of speciation rates in South American freshwater fishes, the most diverse continental vertebrate fauna, by examining the roles of multiple biotic and abiotic factors. We integrate a dataset on species geographic distribution, phylogenetic, morphological, climatic, and habitat data. We find that Late Neogene-Quaternary speciation events are strongly associated with body-size evolution, particularly in lineages with small body sizes that inhabit higher elevations near the continental periphery. Conversely, the effects of temperature, area, and diversity-dependence, often thought to facilitate speciation, are negligible. By evaluating multiple factors simultaneously, we demonstrate that habitat characteristics associated with elevation, as well as body size evolution, correlate with rapid speciation in South American freshwater fishes. Our study emphasizes the importance of integrative approaches that consider the interplay of biotic and abiotic factors in generating macroecological patterns of species diversity.

Climate’s effect on global biodiversity is typically viewed through the lens of temperature, humidity and resulting ecosystem productivity. However, it is not known whether biodiversity depends solely on these climate conditions, or whether the size and fragmentation of these climates are also crucial. Here we shift the common perspective in global biodiversity studies, transitioning from geographic space to a climate-defined multidimensional space. Our findings suggest that larger and more isolated climate conditions tend to harbour higher diversity and species turnover among terrestrial tetrapods, encompassing more than 30,000 species. By considering both the characteristics of climate itself and its geographic attributes, we can explain almost 90% of the variation in global species richness. Half of the explanatory power (45%) may be attributed either to climate itself or to the geography of climate, suggesting a nuanced interplay between them. Our work evolves the conventional idea that larger climate regions, such as the tropics, host more species primarily because of their size. Instead, we underscore the integral roles of both the geographic extent and degree of isolation of climates. This refined understanding presents a more intricate picture of biodiversity distribution, which can guide our approach to biodiversity conservation in an ever-changing world.