The geography of climate and the global patterns of species diversity.

Climate's effect on global biodiversity is typically viewed through the lens of temperature, humidity and resulting ecosystem productivity1-6. 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 size7,8. 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.

When cheating turns into a stabilizing mechanism of plant-pollinator communities.

Mutualistic interactions, such as plant-mycorrhizal or plant-pollinator interactions, are widespread in ecological communities and frequently exploited by cheaters, species that profit from interactions without providing benefits in return. Cheating usually negatively affects the fitness of the individuals that are cheated on, but the effects of cheating at the community level remains poorly understood. Here, we describe 2 different kinds of cheating in mutualistic networks and use a generalized Lotka-Volterra model to show that they have very different consequences for the persistence of the community. Conservative cheating, where a species cheats on its mutualistic partners to escape the cost of mutualistic interactions, negatively affects community persistence. In contrast, innovative cheating occurs with species with whom legitimate interactions are not possible, because of a physiological or morphological barrier. Innovative cheating can enhance community persistence under some conditions: when cheaters have few mutualistic partners, cheat at low or intermediate frequency and the cost associated with mutualism is not too high. Under these conditions, the negative effects of cheating on partner persistence are overcompensated at the community level by the positive feedback loops that arise in diverse mutualistic communities. Using an empirical dataset of plant-bird interactions (hummingbirds and flowerpiercers), we found that observed cheating patterns are highly consistent with theoretical cheating patterns found to increase community persistence. This result suggests that the cheating patterns observed in nature could contribute to promote species coexistence in mutualistic communities, instead of necessarily destabilizing them.

Landscape dynamics and diversification of the megadiverse South American freshwater fish fauna.

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.

Land-use intensity influences European tetrapod food webs.

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.

Linking human impacts to community processes in terrestrial and freshwater ecosystems.

Human impacts such as habitat loss, climate change and biological invasions are radically altering biodiversity, with greater effects projected into the future. Evidence suggests human impacts may differ substantially between terrestrial and freshwater ecosystems, but the reasons for these differences are poorly understood. We propose an integrative approach to explain these differences by linking impacts to four fundamental processes that structure communities: dispersal, speciation, species-level selection and ecological drift. Our goal is to provide process-based insights into why human impacts, and responses to impacts, may differ across ecosystem types using a mechanistic, eco-evolutionary comparative framework. To enable these insights, we review and synthesise (i) how the four processes influence diversity and dynamics in terrestrial versus freshwater communities, specifically whether the relative importance of each process differs among ecosystems, and (ii) the pathways by which human impacts can produce divergent responses across ecosystems, due to differences in the strength of processes among ecosystems we identify. Finally, we highlight research gaps and next steps, and discuss how this approach can provide new insights for conservation. By focusing on the processes that shape diversity in communities, we aim to mechanistically link human impacts to ongoing and future changes in ecosystems.

Morphology and niche evolution influence hummingbird speciation rates.

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.

Diel niche partitioning of a plant-hummingbird network in the Atlantic forest of Brazil.

Niche partitioning is an important mechanism that allows species to coexist. Within mutualistic interaction networks, diel niche partitioning, i.e., partitioning of resources throughout the day, has been neglected. We explored diel niche partitioning of a plant-hummingbird network in the Brazilian Atlantic forest for nine months. To evaluate diel patterns of hummingbird visits and nectar production, we used time-lapse cameras on focal flowers and repeated nectar volume and concentration measures, respectively. Additionally, we measured flower abundance around focal flowers and flower morphological traits. We did not observe diel partitioning for either hummingbirds or plants. Instead, hummingbirds appeared to specialize in different plant species, consistent with trophic niche partitioning, potentially resulting from competition. In contrast, plant species that co-flowered and shared hummingbird visits produced nectar during similar times, consistent with facilitation. Our focus on the fine-scale temporal pattern revealed that plants and hummingbirds appear to have different strategies for promoting co-existence.

Hummingbird community structure and nectar resources modulate the response of interspecific competition to forest conversion.

On-going land-use change has profound impacts on biodiversity by filtering species that cannot survive in disturbed landscapes and potentially altering biotic interactions. In particular, how land-use change reshapes biotic interactions remains an open question. Here, we used selectivity experiments with nectar feeders in natural and converted forests to test the direct and indirect effects of land-use change on resource competition in Andean hummingbirds along an elevational gradient. Selectivity was defined as the time hummingbirds spent at high resource feeders when feeders with both low and high resource values were offered in the presence of other hummingbird species. Selectivity approximates the outcome of interspecific competition (i.e., the resource intake across competing species); in the absence of competition, birds should exhibit higher selectivity. We evaluated the indirect effect of forest conversion on selectivity, as mediated by morphological dissimilarity and flower resource abundance, using structural equation models. We found that forest conversion influenced selectivity at low and mid-elevations, but the influence of morphological dissimilarity and resource availability on selectivity varied between these elevations. At mid-elevation, selectivity was more influenced by the presence of morphologically similar competitors than by resource abundance while at low-elevation resource abundance was a more important predictor of selectivity. Our results suggest that selectivity is influenced by forest conversion, but that the drivers of these changes vary across elevation, highlighting the importance of considering context-dependent variation in the composition of resources and competitors when studying competition.

Global plant-frugivore trait matching is shaped by climate and biogeographic history.

Species interactions are influenced by the trait structure of local multi-trophic communities. However, it remains unclear whether mutualistic interactions in particular can drive trait patterns at the global scale, where climatic constraints and biogeographic processes gain importance. Here we evaluate global relationships between traits of frugivorous birds and palms (Arecaceae), and how these relationships are affected, directly or indirectly, by assemblage richness, climate and biogeographic history. We leverage a new and expanded gape size dataset for nearly all avian frugivores, and find a positive relationship between gape size and fruit size, that is, trait matching, which is influenced indirectly by palm richness and climate. We also uncover a latitudinal gradient in trait matching strength, which increases towards the tropics and varies among zoogeographic realms. Taken together, our results suggest trophic interactions have consistent influences on trait structure, but that abiotic, biogeographic and richness effects also play important, though sometimes indirect, roles in shaping the functional biogeography of mutualisms.

Seasonal structure of interactions enhances multidimensional stability of mutualistic networks.

Community ecologists have made great advances in understanding how natural communities can be both diverse and stable by studying communities as interaction networks. However, focus has been on interaction networks aggregated over time, neglecting the consequences of the seasonal organization of interactions (hereafter 'seasonal structure') for community stability. Here, we extended previous theoretical findings on the topic in two ways: (i) by integrating empirical seasonal structure of 11 plant-hummingbird communities into dynamic models, and (ii) by tackling multiple facets of network stability together. We show that, in a competition context, seasonal structure enhances community stability by allowing diverse and resilient communities while preserving their robustness to species extinctions. The positive effects of empirical seasonal structure on network stability vanished when using randomized seasonal structures, suggesting that eco-evolutionary dynamics produce stabilizing seasonal structures. We also show that the effects of seasonal structure on community stability are mainly mediated by changes in network structure and productivity, suggesting that the seasonal structure of a community is an important and yet neglected aspect in the diversity-stability and diversity-productivity debates.

A heterothermic spectrum in hummingbirds.

Many endotherms use torpor, saving energy by a controlled reduction of their body temperature and metabolic rate. Some species (e.g. arctic ground squirrels, hummingbirds) enter deep torpor, dropping their body temperature by 23-37°C, while others can only enter shallow torpor (e.g. pigeons, 3-10°C reduction). However, deep torpor in mammals can increase predation risk (unless animals are in burrows or caves), inhibit immune function and result in sleep deprivation, so even for species that can enter deep torpor, facultative shallow torpor might help balance energy savings with these potential costs. Deep torpor occurs in three avian orders, but the trade-offs of deep torpor in birds are unknown. Although the literature hints that some bird species (mousebirds and perhaps hummingbirds) can use both shallow and deep torpor, little empirical evidence of such an avian heterothermy spectrum within species exists. We infrared imaged three hummingbird species that are known to use deep torpor, under natural temperature and light cycles, to test whether they were also capable of shallow torpor. All three species used both deep and shallow torpor, often on the same night. Depending on the species, they used shallow torpor for 5-35% of the night. The presence of a heterothermic spectrum in these bird species indicates a capacity for fine-scale physiological and genetic regulation of avian torpid metabolism.

Area, isolation and climate explain the diversity of mammals on islands worldwide.

Insular biodiversity is expected to be regulated differently than continental biota, but their determinants remain to be quantified at a global scale. We evaluated the importance of physical, environmental and historical factors on mammal richness and endemism across 5592 islands worldwide. We fitted generalized linear and mixed models to accommodate variation among biogeographic realms and performed analyses separately for bats and non-volants. Richness on islands ranged from one to 234 species, with up to 177 single island endemics. Diversity patterns were most consistently influenced by the islands' physical characteristics. Area positively affected mammal diversity, in particular the number of non-volant endemics. Island isolation, both current and past, was associated with lower richness but greater endemism. Flight capacity modified the relative importance of past versus current isolation, with bats responding more strongly to current and non-volant mammals to past isolation. Biodiversity relationships with environmental factors were idiosyncratic, with a tendency for greater effects sizes with endemism than richness. The historical climatic change was positively associated with endemism. In line with theory, we found that area and isolation were among the strongest drivers of mammalian biodiversity. Our results support the importance of past conditions on current patterns, particularly of non-volant species.

Spatial variation in direct and indirect effects of climate and productivity on species richness of terrestrial tetrapods.

AIM: We aimed to dissect the spatial variation of the direct and indirect effects of climate and productivity on global species richness of terrestrial tetrapods. LOCATION: Global. TIME PERIOD: Present. MAJOR TAXA STUDIED: Terrestrial tetrapods. METHODS: We used a geographically weighted path analysis to estimate and map the direct and indirect effects of temperature, precipitation and primary productivity on species richness of terrestrial tetrapods across the globe. RESULTS: We found that all relationships shift in magnitude, and even in direction, among taxonomic groups, geographical regions and connecting paths. Direct effects of temperature and precipitation are generally stronger than both indirect effects mediated by productivity and direct effects of productivity. MAIN CONCLUSIONS: Richness gradients seem to be driven primarily by effects of climate on organismal physiological limits and metabolic rates rather than by the amount of productive energy. Reptiles have the most distinct relationships across tetrapods, with a clear latitudinal pattern in the importance of temperature versus water.

Phenological synchronization of seasonal bird migration with vegetation greenness across dietary guilds.

The seasonal movement of animals has been linked to seasonal variation in ecological productivity, and it has been hypothesized that primary consumers synchronize migration with vegetation phenology. Within temperate regions of the Northern Hemisphere, herbivorous bird species often track the phenology of vegetation greenness during spring migration. Phenological synchronization with vegetation greenness by migratory birds in other dietary guilds, across the full extent of their annual distributions during both spring and autumn migration, has not been explored. Here, we document population-level associations with a remotely sensed measure of vegetation greenness for 230 North American migratory bird species in seven dietary guilds across the full annual cycle using eBird occurrence information for the combined period 2006-2018. Evidence of phenological synchronization was strongest for omnivores, herbivores, herbivore-granivores and granivores during spring and autumn migration, except for omnivores in the west during spring migration. Strong evidence of synchronization was also observed for insectivores during spring migration and carnivores during spring and autumn migration that migrated across the entire breadth of the continent. The level of evidence declined for insectivores in the west and east during spring migration, and for nectarivores in the west during spring and autumn migration. Limited evidence was also found for insectivores in the east during autumn migration, insectivores in the west and the centre of the continent during spring and autumn migration, and carnivores in the west during spring migration. Carnivores in the west during autumn migration showed the weakest evidence of synchronization. We found broad support across an array of dietary guilds for phenological coupling between vegetation greenness and seasonal bird migration within North America. Our results highlight the potential for many migratory bird species to encounter phenological mismatches as vegetation phenology responds to climate change. Our findings emphasize the need to better understand the environmental cues that regulate migratory behaviour across dietary guilds, consumer levels and migration tactics.

Pervasive Genomic Signatures of Local Adaptation to Altitude Across Highland Specialist Andean Hummingbird Populations.

Populations along steep environmental gradients are subject to differentiating selection that can result in local adaptation, despite countervailing gene flow, and genetic drift. In montane systems, where species are often restricted to narrow ranges of elevation, it is unclear whether the selection is strong enough to influence functional differentiation of subpopulations differing by a few hundred meters in elevation. We used targeted capture of 12 501 exons from across the genome, including 271 genes previously implicated in altitude adaptation, to test for adaptation to local elevations for 2 highland hummingbird species, Coeligena violifer (n = 62) and Colibri coruscans (n = 101). For each species, we described population genetic structure across the complex geography of the Peruvian Andes and, while accounting for this structure, we tested whether elevational allele frequency clines in single nucleotide polymorphisms (SNPs) showed evidence for local adaptation to elevation. Although the 2 species exhibited contrasting population genetic structures, we found signatures of clinal genetic variation with shifts in elevation in both. The genes with SNP-elevation associations included candidate genes previously discovered for high-elevation adaptation as well as others not previously identified, with cellular functions related to hypoxia response, energy metabolism, and immune function, among others. Despite the homogenizing effects of gene flow and genetic drift, natural selection on parts of the genome evidently optimizes elevation-specific cellular function even within elevation range-restricted montane populations. Consequently, our results suggest local adaptation occurring in narrow elevation bands in tropical mountains, such as the Andes, may effectively make them "taller" biogeographic barriers.

Temporal stability in species richness but reordering in species abundances within avian assemblages of a tropical Andes conservation hot spot.

As the pace of environmental change increases, there is an urgent need for quantitative data revealing the temporal dynamics of local communities in tropical areas. Here, we quantify the stability of avian assemblages in the highly threatened, but poorly studied, Andean biodiversity hot spot. We evaluated the temporal variation in species richness and community composition of local bird assemblages in three habitat types (native forest, introduced forest, native shrub) using a unique, relatively long-term data series from Cajas National Park and Mazán Reserve in the southern Andes of Ecuador. We sampled birds with mist nets using a standardized protocol over 11 years, from 2006 to 2016. Species richness remained stable over time across habitats, but community composition changed in the native forest. In particular, we observed taxonomic reordering in the native forest, in which the evenness in the distribution of abundances of taxa decreased over time. This finding is consistent with other studies where species richness remained constant over time while community composition changed. Our study highlights the value of long-term studies in the tropical Andes as we show that species composition of birds in a montane forest is changing, consistent with global trends in biodiversity change.

La tasa acelerada del incremento en el cambio ambiental crea la necesidad urgente de datos cuantitativos que expongan la dinámica temporal de comunidades locales en áreas tropicales. En este manuscrito, cuantificamos la estabilidad de ensambles de aves un área altamente amenazada pero poco estudiada, el hot­spot de biodiversidad Andino. Evaluamos la variación temporal en riqueza y composición del ensamble de aves local en tres tipos de hábitat (bosque nativo, bosque introducido, matorrales nativos) usando una serie de datos de largo plazo relativo, obtenidos en el Parque Nacional Cajas y la reserva Mazan, en los Andes del sur del Ecuador. Las aves se estudiaron con redes de neblina por medio de protocolos estandarizados por 11 años, desde 1996 al 2016. La riqueza de especies permaneció estable en el tiempo en todos los hábitats, pero la composición de especies cambió en el bosque nativo. En particular, observamos un reorden taxonómico en el bosque nativo, en el cual existió una diminución en la equitatividad de las distribuciones de las especies en el tiempo. Este resultado es consistente con otros estudios que encuentran que la riqueza de especies permanece constante en el tiempo mientras que la composición de la comunidad cambia. Nuestro trabajo resalta el valor de los estudios de largo tiempo en los Andes tropicales al demostrar que la composición de aves está cambiando, y concuerda con los patrones globales de cambio de la biodiversidad.

The allometry of daily energy expenditure in hummingbirds: An energy budget approach.

Within-clade allometric relationships represent standard laws of scaling between energy and size, and their outliers provide new avenues for physiological and ecological research. According to the metabolic-level boundaries hypothesis, metabolic rates as a function of mass are expected to scale closer to 0.67 when driven by surface-related processes (e.g. heat or water flux), while volume-related processes (e.g. activity) generate slopes closer to one. In birds, daily energy expenditure (DEE) scales with body mass (M) in the relationship logDEE=2.35+0.68×logM , consistent with surface-level processes driving the relationship. However, taxon-specific patterns differ from the scaling slope of all birds. Hummingbirds have the highest mass-specific metabolic rates among all vertebrates. Previous studies on a few hummingbird species, without accounting for the phylogeny, estimated that the DEE-body mass relationship for hummingbirds was logDEE=1.72+1.21×logM . In Contrast to the theoretical expectations, this slope >1 indicates that larger hummingbirds are less metabolically efficient than smaller hummingbirds. We collected DEE and mass data for 12 hummingbird species, which, combined with published data, represented 17 hummingbird species in eight of nine hummingbird clades over a sixfold size range of body size (2.7-17.5 g). After accounting for phylogenetic relatedness, we found DEE scales with body mass as logDEE=2.04+0.95×logM . This slope of 0.95 is lower than previously estimated for hummingbirds, but much higher than the slope for all birds (0.68). The high slopes of torpor, hovering and flight potentially explain the high interspecific DEE slope for hummingbirds compared to other endotherms.

Global priorities for conservation across multiple dimensions of mammalian diversity.

Conservation priorities that are based on species distribution, endemism, and vulnerability may underrepresent biologically unique species as well as their functional roles and evolutionary histories. To ensure that priorities are biologically comprehensive, multiple dimensions of diversity must be considered. Further, understanding how the different dimensions relate to one another spatially is important for conservation prioritization, but the relationship remains poorly understood. Here, we use spatial conservation planning to (i) identify and compare priority regions for global mammal conservation across three key dimensions of biodiversity-taxonomic, phylogenetic, and traits-and (ii) determine the overlap of these regions with the locations of threatened species and existing protected areas. We show that priority areas for mammal conservation exhibit low overlap across the three dimensions, highlighting the need for an integrative approach for biodiversity conservation. Additionally, currently protected areas poorly represent the three dimensions of mammalian biodiversity. We identify areas of high conservation priority among and across the dimensions that should receive special attention for expanding the global protected area network. These high-priority areas, combined with areas of high priority for other taxonomic groups and with social, economic, and political considerations, provide a biological foundation for future conservation planning efforts.

Twenty-million-year relationship between mammalian diversity and primary productivity.

At global and regional scales, primary productivity strongly correlates with richness patterns of extant animals across space, suggesting that resource availability and climatic conditions drive patterns of diversity. However, the existence and consistency of such diversity-productivity relationships through geological history is unclear. Here we provide a comprehensive quantitative test of the diversity-productivity relationship for terrestrial large mammals through time across broad temporal and spatial scales. We combine >14,000 occurrences for 690 fossil genera through the Neogene (23-1.8 Mya) with regional estimates of primary productivity from fossil plant communities in North America and Europe. We show a significant positive diversity-productivity relationship through the 20-million-year record, providing evidence on unprecedented spatial and temporal scales that this relationship is a general pattern in the ecology and paleo-ecology of our planet. Further, we discover that genus richness today does not match the fossil relationship, suggesting that a combination of human impacts and Pleistocene climate variability has modified the 20-million-year ecological relationship by strongly reducing primary productivity and driving many mammalian species into decline or to extinction.