Coevolutionary hotspots favour dispersal and fuel biodiversity in mutualistic landscapes under environmental changes.

Mutualistic interactions are key to sustaining Earth's biodiversity. Yet, we are only beginning to understand how coevolution in mutualistic assemblages can shape the distribution and persistence of species across landscapes. Here, we combine the geographic mosaic theory of coevolution with metacommunity dynamics to understand how geographically structured selection can shape patterns of richness, dispersal, extinction and persistence of mutualistic species. In this model, species may experience strong or weak reciprocal selection imposed by mutualisms within each patch (i.e. hotspots and coldspots, respectively). Using numerical simulations, we show that mutualistic coevolution leads to a concentration of species richness at hotspots. Such an effect occurs because hotspots sustain higher rates of colonization and lower rates of extinction than coldspots, whether the environment changes or not. Importantly, under environmental changes, coldspots fail to sustain a positive colonization-to-extinction balance. Rather, species persistence within coldspots relies on hotspots acting as biodiversity sources and enhancing population dispersal across the landscape. In fact, even a few hotspots in the landscape can fuel the spatial network of dispersal of populations in the metacommunity. Our study highlights that coevolutionary hotspots can act as biodiversity sources, favouring colonization and allowing species to expand their distribution across landscapes even in changing environments. This article is part of the theme issue 'Diversity-dependence of dispersal: interspecific interactions determine spatial dynamics'.

A global map of species at risk of extinction due to natural hazards.

An often-overlooked question of the biodiversity crisis is how natural hazards contribute to species extinction risk. To address this issue, we explored how four natural hazards, earthquakes, hurricanes, tsunamis, and volcanoes, overlapped with the distribution ranges of amphibians, birds, mammals, and reptiles that have either narrow distributions or populations with few mature individuals. To assess which species are at risk from these natural hazards, we combined the frequency and magnitude of each natural hazard to estimate their impact. We considered species at risk if they overlapped with regions where any of the four natural hazards historically occurred (n = 3,722). Those species with at least a quarter of their range subjected to a high relative impact were considered at high risk (n = 2,001) of extinction due to natural hazards. In total, 834 reptiles, 617 amphibians, 302 birds, and 248 mammals were at high risk and they were mainly distributed on islands and in the tropics. Hurricanes (n = 983) and earthquakes (n = 868) affected most species, while tsunamis (n = 272), and volcanoes (n = 171) affected considerably fewer. The region with the highest number of species at high risk was the Pacific Ring of Fire, especially due to volcanoes, earthquakes, and tsunamis, while hurricane-related high-risk species were concentrated in the Caribbean Sea, Gulf of Mexico, and northwestern Pacific Ocean. Our study provides important information regarding the species at risk due to natural hazards and can help guide conservation attention and efforts to safeguard their survival.

The reorganization of predator-prey networks over 20 million years explains extinction patterns of mammalian carnivores.

Linking the species interactions occurring at the scale of local communities to their potential impact at evolutionary timescales is challenging. Here, we used the high-resolution fossil record of mammals from the Iberian Peninsula to reconstruct a timeseries of trophic networks spanning more than 20 million years and asked whether predator-prey interactions affected regional extinction patterns. We found that, despite small changes in species richness, trophic networks showed long-term trends, gradually losing interactions and becoming sparser towards the present. This restructuring of the ecological networks was driven by the loss of medium-sized herbivores, which reduced prey availability for predators. The decrease in prey availability was associated with predator longevity, such that predators with less available prey had greater extinction risk. These results not only reveal long-term trends in network structure but suggest that prey species richness in ecological communities may shape large scale patterns of extinction and persistence among predators.

Climate change should drive mammal defaunation in tropical dry forests.

Human-induced climate change has intensified negative impacts on socioeconomic factors, the environment, and biodiversity, including changes in rainfall patterns and an increase in global average temperatures. Drylands are particularly at risk, with projections suggesting they will become hotter, drier, and less suitable for a significant portion of their species, potentially leading to mammal defaunation. We use ecological niche modelling and community ecology biodiversity metrics to examine potential geographical range shifts of non-volant mammal species in the largest Neotropical dryland, the Caatinga, and evaluate impacts of climate change on mammal assemblages. According to projections, 85% of the mammal species will lose suitable habitats, with one quarter of species projected to completely lose suitable habitats by 2060. This will result in a decrease in species richness for more than 90% of assemblages and an increase in compositional similarity to nearby assemblages (i.e., reduction in spatial beta diversity) for 70% of the assemblages. Small-sized mammals will be the most impacted and lose most of their suitable habitats, especially in highlands. The scenario is even worse in the eastern half of Caatinga where habitat destruction already prevails, compounding the threats faced by species there. While species-specific responses can vary with respect to dispersal, behavior, and energy requirements, our findings indicate that climate change can drive mammal assemblages to biotic homogenization and species loss, with drastic changes in assemblage trophic structure. For successful long-term socioenvironmental policy and conservation planning, it is critical that findings from biodiversity forecasts are considered.

Indirect effects shape species fitness in coevolved mutualistic networks.

Ecological interactions are one of the main forces that sustain Earth's biodiversity. A major challenge for studies of ecology and evolution is to determine how these interactions affect the fitness of species when we expand from studying isolated, pairwise interactions to include networks of interacting species1-4. In networks, chains of effects caused by a range of species have an indirect effect on other species they do not interact with directly, potentially affecting the fitness outcomes of a variety of ecological interactions (such as mutualism)5-7. Here we apply analytical techniques and numerical simulations to 186 empirical mutualistic networks and show how both direct and indirect effects alter the fitness of species coevolving in these networks. Although the fitness of species usually increased with the number of mutualistic partners, most of the fitness variation across species was driven by indirect effects. We found that these indirect effects prevent coevolving species from adapting to their mutualistic partners and to other sources of selection pressure in the environment, thereby decreasing their fitness. Such decreases are distributed in a predictable way within networks: peripheral species receive more indirect effects and experience higher reductions in fitness than central species. This topological effect was also evident when we analysed an empirical study of an invasion of pollination networks by honeybees. As honeybees became integrated as a central species within networks, they increased the contribution of indirect effects on several other species, reducing their fitness. Our study shows how and why indirect effects can govern the adaptive landscape of species-rich mutualistic assemblages.

Identifying climate change refugia for South American biodiversity.

Refugia-based conservation offers long-term effectiveness and minimize uncertainty on strategies for climate change adaptation. We used distribution modelling to identify climate change refugia for 617 terrestrial mammals and to quantify the role of protected areas (PAs) in providing refugia across South America. To do so, we compared species potential distribution across different scenarios of climate change, highlighting those regions likely to retain suitable climatic conditions by year 2090, and explored the proportion of refugia inside PAs. Moist tropical forests in high-elevation areas with complex topography concentrated the highest local diversity of species refugia, although regionally important refugia centers occurred elsewhere. Andean-Amazon forests contained climate change refugia for more than half of the continental species' pool and for up to 87 species locally (17 × 17 km2 grid cell). The highlands of the southern Atlantic Forest also included megadiverse refugia for up to 76 species per cell. Almost half of the species that may find refugia in the Atlantic Forest will do so in a single region-the Serra do Mar and Serra do Espinhaço. Most of the refugia we identified, however, were not in PAs, which may contain <6% of the total area of climate change refugia, leaving 129-237 species with no refugia inside the territorial limits of PAs of any kind. Our results reveal a dismal scenario for the level of refugia protection in some of the most biodiverse regions of the world. Nonetheless, because refugia tend to be in high-elevation, topographically complex, and remote areas, with lower anthropogenic pressure, formally protecting them may require a comparatively modest investment.

Identificación de refugios para la biodiversidad de Sudamérica ante el cambio climático Resumen Las estrategias de conservación basadas en refugios ofrecen efectividad a largo plazo y minimizan la incertidumbre sobre las estrategias de adaptación al cambio climático. Utilizamos modelos de distribución para identificar los refugios del cambio climático de 617 especies de mamíferos terrestres y cuantificar el papel de las áreas protegidas en la provisión de refugios en Sudamérica. Para esto, comparamos la distribución potencial de las especies en diferentes escenarios de cambio climático, destacando las regiones que probablemente conservarán las condiciones climáticas adecuadas para el año 2090, y exploramos la proporción de refugios dentro de las áreas protegidas. Los bosques tropicales húmedos de zonas de gran altitud y topografía compleja concentraron la mayor diversidad local de refugios de especies, aunque también hubo centros de refugio de importancia regional en otras localidades. Los bosques amazónicos andinos albergaron los refugios ante el cambio climático de más de la mitad del conjunto de especies continentales y para hasta 87 especies a escala local (celda cuadriculada de 17 × 17 km2 ). Las tierras altas del sur del Bosque Atlántico también incluyeron refugios megadiversos para hasta 76 especies por celda. Casi la mitad de las especies que pueden refugiarse en el Bosque Atlántico lo harán en una sola región: la Serra do Mar y la Serra do Espinhaço. Sin embargo, la mayoría de los refugios que identificamos no estaban en áreas protegidas, las cuales pueden contener <6% del área total de refugios del cambio climático, dejando entre 129 y 237 especies sin refugio dentro de los límites territoriales de las áreas protegidas de cualquier tipo. Nuestros resultados revelan un panorama desolador para el nivel de protección de los refugios en algunas de las regiones con mayor biodiversidad del mundo. No obstante, dado que los refugios suelen encontrarse en zonas remotas de gran altitud con topografía compleja y menor presión antropogénica, protegerlos formalmente puede requerir una inversión comparativamente modesta.

Decoupled responses of biodiversity facets driven from anuran vulnerability to climate and land-use changes.

Biodiversity loss not only implies the loss of species but also entails losses in other dimensions of biodiversity, such as functional, phylogenetic and interaction diversity. Yet, each of those facets of biodiversity may respond differently to extinctions. Here, we examine how extinction, driven by climate and land-use changes may affect those different facets of diversity by combining empirical data on anuran-prey interaction networks, species distribution modelling and extinction simulations in assemblages representing four Neotropical ecoregions. We found a mismatch in the response of functional, phylogenetic and interaction diversity to extinction. In spite of high network robustness to extinction, the effects on interaction diversity were stronger than those on phylogenetic and functional diversity, declining linearly with species loss. Although it is often assumed that interaction patterns are reflected by functional diversity, assessing species interactions may be necessary to understand how species loss translates into the loss of ecosystem functions.

Mutualistic coevolution and community diversity favour persistence in metacommunities under environmental changes.

Linking local to regional ecological and evolutionary processes is key to understand the response of Earth's biodiversity to environmental changes. Here we integrate evolution and mutualistic coevolution in a model of metacommunity dynamics and use numerical simulations to understand how coevolution can shape species distribution and persistence in landscapes varying in space and time. Our simulations show that coevolution and species richness can synergistically shape distribution patterns by increasing colonization and reducing extinction of populations in metacommunities. Although conflicting selective pressures emerging from mutualisms may increase mismatches with the local environment and the rate of local extinctions, coevolution increases trait matching among mutualists at the landscape scale, counteracting local maladaptation and favouring colonization and range expansions. Our results show that by facilitating colonization, coevolution can also buffer the effects of environmental changes, preventing species extinctions and the collapse of metacommunities. Our findings reveal the mechanisms whereby coevolution can favour persistence under environmental changes and highlight that these positive effects are greater in more diverse systems that retain landscape connectivity.

Differences in prey availability across space and time lead to interaction rewiring and reshape a predator-prey metaweb.

Space and time promote variation in network structure by affecting the likelihood of potential interactions. However, little is known about the relative roles of ecological and biogeographical processes in determining how species interactions vary across space and time. Here we study the spatiotemporal variation in predator-prey interaction networks formed by anurans and arthropods and test for the effects of prey availability in determining interaction patterns, information that is often absent and limits the understanding of the determinants of network structure. We found that network dissimilarity between ecoregions and seasons was high and primarily driven by interaction rewiring.We also found that species turnover was positively related to geographical distance. Using a null model approach to disentangle the effect of prey availability on the spatial and temporal variation, we show that differences in prey availability were important in determining the variation in network structure between seasons and among areas. Our study reveals that fluctuations in prey abundance, alongside the limited dispersal abilities of anurans and their prey, may be responsible for the spatial patterns that emerged in our predator-prey metaweb. These findings contribute to our understanding of the assembly rules that maintain biotic processes in metacommunities and highlight the importance of prey availability to the structure of these systems.

The effect of past defaunation on ranges, niches, and future biodiversity forecasts.

Humans have reshaped the distribution of biodiversity across the globe, extirpating species from regions otherwise suitable and restricting populations to a subset of their original ranges. Here, we ask if anthropogenic range contractions since the Late Pleistocene led to an under-representation of the realized niches for megafauna, an emblematic group of taxa often targeted for restoration actions. Using reconstructions of past geographic distributions (i.e., natural ranges) for 146 extant terrestrial large-bodied (>44 kg) mammals, we estimate their climatic niches as if they had retained their original distributions and evaluate their observed niche dynamics. We found that range contractions led to a sizeable under-representation of the realized niches of several species (i.e., niche unfilling). For 29 species, more than 10% of the environmental space once seen in their natural ranges has been lost due to anthropogenic activity, with at least 12 species undergoing reductions of more than 50% of their realized niches. Eighteen species may now be confined to low-suitability locations, where fitness and abundance are likely diminished; we consider these taxa 'climatic refugees'. For those species, conservation strategies supported by current ranges risk being misguided if current, suboptimal habitats are considered baseline for future restoration actions. Because most climate-based biodiversity forecasts rely exclusively on current occurrence records, we went on to test the effect of neglecting historical information on estimates of species' potential distribution - as a proxy of sensitivity to climate change. We found that niche unfilling driven by past range contraction leads to an overestimation of sensitivity to future climatic change, resulting in 50% higher rates of global extinction, and underestimating the potential for megafauna conservation and restoration under future climate change. In conclusion, range contractions since the Late Pleistocene have also left imprints on megafauna realized climatic niches. Therefore, niche truncation driven by defaunation can directly affect climate and habitat-based conservation strategies.

How deregulation, drought and increasing fire impact Amazonian biodiversity.

Biodiversity contributes to the ecological and climatic stability of the Amazon Basin1,2, but is increasingly threatened by deforestation and fire3,4. Here we quantify these impacts over the past two decades using remote-sensing estimates of fire and deforestation and comprehensive range estimates of 11,514 plant species and 3,079 vertebrate species in the Amazon. Deforestation has led to large amounts of habitat loss, and fires further exacerbate this already substantial impact on Amazonian biodiversity. Since 2001, 103,079-189,755 km2 of Amazon rainforest has been impacted by fires, potentially impacting the ranges of 77.3-85.2% of species that are listed as threatened in this region5. The impacts of fire on the ranges of species in Amazonia could be as high as 64%, and greater impacts are typically associated with species that have restricted ranges. We find close associations between forest policy, fire-impacted forest area and their potential impacts on biodiversity. In Brazil, forest policies that were initiated in the mid-2000s corresponded to reduced rates of burning. However, relaxed enforcement of these policies in 2019 has seemingly begun to reverse this trend: approximately 4,253-10,343 km2 of forest has been impacted by fire, leading to some of the most severe potential impacts on biodiversity since 2009. These results highlight the critical role of policy enforcement in the preservation of biodiversity in the Amazon.

Morphophysiological analysis of the salivary glands of Rhipicephalus sanguineus sensu lato (Acari: Ixodidae) exposed to ozonated water: A control strategy.

The tick Rhipicephalus sanguineus sensu lato has great medical and veterinary importance, mainly because the ability to transmit many diseases, causing harm to pets but also risks to public health. The blood spoliation and transmission of pathogens occur because of the immunosuppressive action of these ticks' saliva, a potent mixture of bioactive substances that is secreted by the salivary glands, one of the organs responsible for their biological success, and hence the target of studies for their control. Ozone has promise for use as an alternative acaricide, due to its proven efficiency in controlling agricultural and food pests, besides posing no risk of environmental contamination or to animal and human health. Therefore, this study evaluated the acaricidal potential of exposure of females of R. sanguineus s.l. to ozonated water at many concentrations and analysed the morphophysiological alterations of the salivary glands, employing histological and light microscopic techniques. The results demonstrated that the ozonated water at the concentrations investigated caused severe alterations in the salivary glands, bringing a new perspective for control of R. sanguineus s.l., through an ecologically correct method due to the absence of harm to non-target organisms and the environment.

Climate and land-use change will lead to a faunal "savannization" on tropical rainforests.

Humans have fragmented, reduced or altered the biodiversity in tropical forests around the world. Climate and land-use change act synergistically, increasing drought and fire frequencies, converting several tropical rainforests into derived savannas, a phenomenon known as "savannization." Yet, we lack a full understanding of the faunal changes in response to the transformation of plant communities. We argue that the composition of vertebrate assemblages in ecotone regions of forest-savanna transitions from South America will be increasingly replaced by open savanna species, a phenomenon we name "faunal savannization." We combined projections from ecological niche models, habitat filter masks and dispersal simulations to forecast the distribution of 349 species of forest- and savanna-dwelling mammal species across South America. We found that the distribution of savanna species is likely to increase by 11%-30% and spread over lowland Amazon and Atlantic forests. Conversely, forest-specialists are expected to lose nearly 50% of their suitable ranges and to move toward core forest zones, which may thus receive an influx of more than 60 species on the move. Our findings indicate that South American ecotonal faunas might experience high rates of occupancy turnover, in a process parallel to that already experienced by plants. Climate-driven migrations of fauna in human-dominated landscapes will likely interact with fire-induced changes in plant communities to reshape the biodiversity in tropical rainforests worldwide.

The indirect paths to cascading effects of extinctions in mutualistic networks.

Biodiversity loss is a hallmark of our times, but predicting its consequences is challenging. Ecological interactions form complex networks with multiple direct and indirect paths through which the impacts of an extinction may propagate. Here we show that accounting for these multiple paths connecting species is necessary to predict how extinctions affect the integrity of ecological networks. Using an approach initially developed for the study of information flow, we estimate indirect effects in plant-pollinator networks and find that even those species with several direct interactions may have much of their influence over others through long indirect paths. Next, we perform extinction simulations in those networks and show that although traditional connectivity metrics fail in the prediction of coextinction patterns, accounting for indirect interaction paths allows predicting species' vulnerability to the cascading effects of an extinction event. Embracing the structural complexity of ecological systems contributes towards a more predictive ecology, which is of paramount importance amid the current biodiversity crisis.

Associated evolution of fruit size, fruit colour and spines in Neotropical palms.

Understanding how ecological interactions have shaped the evolutionary dynamics of species traits remains a challenge in evolutionary ecology. Combining trait evolution models and phylogenies, we analysed the evolution of characters associated with seed dispersal (fruit size and colour) and herbivory (spines) in Neotropical palms to infer the role of these opposing animal-plant interactions in driving evolutionary patterns. We found that the evolution of fruit colour and fruit size was associated in Neotropical palms, supporting the adaptive interpretation of seed-dispersal syndromes and highlighting the role of frugivores in shaping plant evolution. Furthermore, we revealed a positive association between fruit size and the presence of spines on palm leaves, bracteas and stems. We hypothesize that interactions between palms and large-bodied frugivores/herbivores may explain the evolutionary relationship between fruit size and spines. Large-bodied frugivores, such as extinct megafauna, besides consuming the fruits and dispersing large seeds, may also have consumed the leaves or damaged the plants, thus simultaneously favouring the evolution of large fruits and defensive structures. Our findings show how current trait patterns can be understood as the result of the interplay between antagonistic and mutualistic interactions that have happened throughout the evolutionary history of a clade.

Efficacy of carvacrol on Rhipicephalus (Boophilus) microplus engorged female ticks (Canestrini, 1887) (Acari: Ixodidae): effects on mortality and reproduction.

The activity of carvacrol was evaluated in R. microplus female ticks by estimating the mean lethal concentration (LC50) and the acaricidal efficacy. Ticks were subjected to the Adult Immersion Test (AIT) to calculate LC50. From this value, the AIT was performed again at the concentrations of 20%, 40%, 60%, 80% and 100% of the LC50. Two control groups were established: one treated with distilled water and one with the solvent (ethanol 50%). The following parameters were evaluated: female weight before oviposition, egg mass weight, pre-oviposition and incubation period, hatching percentage, egg production index, fecundity rate, estimated reproduction, reduction in oviposition and hatching, and product efficacy. The obtained LC50 was 20.11 mg/mL. Egg mass weight, egg production index, pre-oviposition and fecundity rates of groups treated with 80% and 100% of LC50 were statistically different when compared to control groups. The product efficacy in groups treated with 100% of LC50 was 61.10%.

Revealing biases in the sampling of ecological interaction networks.

The structure of ecological interactions is commonly understood through analyses of interaction networks. However, these analyses may be sensitive to sampling biases with respect to both the interactors (the nodes of the network) and interactions (the links between nodes), because the detectability of species and their interactions is highly heterogeneous. These ecological and statistical issues directly affect ecologists' abilities to accurately construct ecological networks. However, statistical biases introduced by sampling are difficult to quantify in the absence of full knowledge of the underlying ecological network's structure. To explore properties of large-scale ecological networks, we developed the software EcoNetGen, which constructs and samples networks with predetermined topologies. These networks may represent a wide variety of communities that vary in size and types of ecological interactions. We sampled these networks with different mathematical sampling designs that correspond to methods used in field observations. The observed networks generated by each sampling process were then analyzed with respect to the number of components, size of components and other network metrics. We show that the sampling effort needed to estimate underlying network properties depends strongly both on the sampling design and on the underlying network topology. In particular, networks with random or scale-free modules require more complete sampling to reveal their structure, compared to networks whose modules are nested or bipartite. Overall, modules with nested structure were the easiest to detect, regardless of the sampling design used. Sampling a network starting with any species that had a high degree (e.g., abundant generalist species) was consistently found to be the most accurate strategy to estimate network structure. Because high-degree species tend to be generalists, abundant in natural communities relative to specialists, and connected to each other, sampling by degree may therefore be common but unintentional in empirical sampling of networks. Conversely, sampling according to module (representing different interaction types or taxa) results in a rather complete view of certain modules, but fails to provide a complete picture of the underlying network. To reduce biases introduced by sampling methods, we recommend that these findings be incorporated into field design considerations for projects aiming to characterize large species interaction networks.

Interaction paths promote module integration and network-level robustness of spliceosome to cascading effects.

The functionality of distinct types of protein networks depends on the patterns of protein-protein interactions. A problem to solve is understanding the fragility of protein networks to predict system malfunctioning due to mutations and other errors. Spectral graph theory provides tools to understand the structural and dynamical properties of a system based on the mathematical properties of matrices associated with the networks. We combined two of such tools to explore the fragility to cascading effects of the network describing protein interactions within a key macromolecular complex, the spliceosome. Using S. cerevisiae as a model system we show that the spliceosome network has more indirect paths connecting proteins than random networks. Such multiplicity of paths may promote routes to cascading effects to propagate across the network. However, the modular network structure concentrates paths within modules, thus constraining the propagation of such cascading effects, as indicated by analytical results from the spectral graph theory and by numerical simulations of a minimal mathematical model parameterized with the spliceosome network. We hypothesize that the concentration of paths within modules favors robustness of the spliceosome against failure, but may lead to a higher vulnerability of functional subunits, which may affect the temporal assembly of the spliceosome. Our results illustrate the utility of spectral graph theory for identifying fragile spots in biological systems and predicting their implications.

Estimating interaction credit for trophic rewilding in tropical forests.

Trophic rewilding has been suggested as a restoration tool to restore ecological interactions and reverse defaunation and its cascading effects on ecosystem functioning. One of the ecological processes that has been jeopardized by defaunation is animal-mediated seed dispersal. Here, we propose an approach that combines joint species distribution models with occurrence data and species interaction records to quantify the potential to restore seed-dispersal interactions through rewilding and apply it to the Atlantic Forest, a global biodiversity hotspot. Using this approach, we identify areas that should benefit the most from trophic rewilding and candidate species that could contribute to cash the credit of seed-dispersal interactions in a given site. We found that sites within large fragments bearing a great diversity of trees may have about 20 times as many interactions to be cashed through rewilding as small fragments in regions where deforestation has been pervasive. We also ranked mammal and bird species according to their potential to restore seed-dispersal interactions if reintroduced while considering the biome as a whole and at finer scales. The suggested approach can aid future conservation efforts in rewilding projects in defaunated tropical rainforests.This article is part of the theme issue 'Trophic rewilding: consequences for ecosystems under global change'.

Diversification dynamics of mammalian clades during the K-Pg mass extinction.

The Cretaceous/Palaeogene (K-Pg) episode is an iconic mass extinction, in which the diversity of numerous clades abruptly declined. However, the responses of individual clades to mass extinctions may be more idiosyncratic than previously understood. Here, we examine the diversification dynamics of the three major mammalian clades in North America across the K-Pg. Our results show that these clades responded in dramatically contrasting ways to the K-Pg event. Metatherians underwent a sudden rise in extinction rates shortly after the K-Pg, whereas declining origination rates first halted diversification and later drove the loss of diversity in multituberculates. Eutherians experienced high taxonomic turnover near the boundary, with peaks in both origination and extinction rates. These findings indicate that the effects of geological episodes on diversity are context dependent and that mass extinctions can affect the diversification of clades by independently altering the extinction regime, the origination regime or both.