A latitudinal gradient in Darwin's naturalization conundrum at the global scale for flowering plants.

Darwin's naturalization conundrum describes two seemingly contradictory hypotheses regarding whether alien species closely or distantly related to native species should be more likely to naturalize in regional floras. Both expectations have accumulated empirical support, and whether such apparent inconsistency can be reconciled at the global scale is unclear. Here, using 219,520 native and 9,531 naturalized alien plant species across 487 globally distributed regions, we found a latitudinal gradient in Darwin's naturalization conundrum. Naturalized alien plant species are more closely related to native species at higher latitudes than they are at lower latitudes, indicating a greater influence of preadaptation in harsher climates. Human landscape modification resulted in even steeper latitudinal clines by selecting aliens distantly related to natives in warmer and drier regions. Our results demonstrate that joint consideration of climatic and anthropogenic conditions is critical to reconciling Darwin's naturalization conundrum.

The poleward naturalization of intracontinental alien plants.

Plant introductions outside their native ranges by humans have led to substantial ecological consequences. While we have gained considerable knowledge about intercontinental introductions, the distribution and determinants of intracontinental aliens remain poorly understood. Here, we studied naturalized (i.e., self-sustaining) intracontinental aliens using native and alien floras of 243 mainland regions in North America, South America, Europe, and Australia. We revealed that 4510 plant species had intracontinental origins, accounting for 3.9% of all plant species and 56.7% of all naturalized species in these continents. In North America and Europe, the numbers of intracontinental aliens peaked at mid-latitudes, while the proportion peaked at high latitudes in Europe. Notably, we found predominant poleward naturalization, primarily due to larger native species pools in low-latitudes. Geographic and climatic distances constrained the naturalization of intracontinental aliens in Australia, Europe, and North America, but not in South America. These findings suggest that poleward naturalizations will accelerate, as high latitudes become suitable for more plant species due to climate change.

Evolutionary imbalance, climate and human history jointly shape the global biogeography of alien plants.

Human activities are causing global biotic redistribution, translocating species and providing them with opportunities to establish populations beyond their native ranges. Species originating from certain global regions, however, are disproportionately represented among naturalized aliens. The evolutionary imbalance hypothesis posits that differences in absolute fitness among biogeographic divisions determine outcomes when biotas mix. Here, we compile data from native and alien distributions for nearly the entire global seed plant flora and find that biogeographic conditions predicted to drive evolutionary imbalance act alongside climate and anthropogenic factors to shape flows of successful aliens among regional biotas. Successful aliens tend to originate from large, biodiverse regions that support abundant populations and where species evolve against a diverse backdrop of competitors and enemies. We also reveal that these same native distribution characteristics are shared among the plants that humans select for cultivation and economic use. In addition to influencing species' innate potentials as invaders, we therefore suggest that evolutionary imbalance shapes plants' relationships with humans, impacting which species are translocated beyond their native distributions.

The role of phylogenetic relatedness on alien plant success depends on the stage of invasion.

Darwin's naturalization hypothesis predicts successful alien invaders to be distantly related to native species, whereas his pre-adaptation hypothesis predicts the opposite. It has been suggested that depending on the invasion stage (that is, introduction, naturalization and invasiveness), both hypotheses, now known as Darwin's naturalization conundrum, could hold true. We tested this by analysing whether the likelihood of introduction for cultivation, as well as the subsequent stages of naturalization and spread (that is, becoming invasive) of species alien to Southern Africa are correlated with their phylogenetic distance to the native flora of this region. Although species are more likely to be introduced for cultivation if they are distantly related to the native flora, the probability of subsequent naturalization was higher for species closely related to the native flora. Furthermore, the probability of becoming invasive was higher for naturalized species distantly related to the native flora. These results were consistent across three different metrics of phylogenetic distance. Our study reveals that the relationship between phylogenetic distance to the native flora and the success of an alien species changes from one invasion stage to the other.

The global loss of floristic uniqueness.

Regional species assemblages have been shaped by colonization, speciation and extinction over millions of years. Humans have altered biogeography by introducing species to new ranges. However, an analysis of how strongly naturalized plant species (i.e. alien plants that have established self-sustaining populations) affect the taxonomic and phylogenetic uniqueness of regional floras globally is still missing. Here, we present such an analysis with data from native and naturalized alien floras in 658 regions around the world. We find strong taxonomic and phylogenetic floristic homogenization overall, and that the natural decline in floristic similarity with increasing geographic distance is weakened by naturalized species. Floristic homogenization increases with climatic similarity, which emphasizes the importance of climate matching in plant naturalization. Moreover, floristic homogenization is greater between regions with current or past administrative relationships, indicating that being part of the same country as well as historical colonial ties facilitate floristic exchange, most likely due to more intensive trade and transport between such regions. Our findings show that naturalization of alien plants threatens taxonomic and phylogenetic uniqueness of regional floras globally. Unless more effective biosecurity measures are implemented, it is likely that with ongoing globalization, even the most distant regions will lose their floristic uniqueness.

Dimensions of invasiveness: Links between local abundance, geographic range size, and habitat breadth in Europe's alien and native floras.

Understanding drivers of success for alien species can inform on potential future invasions. Recent conceptual advances highlight that species may achieve invasiveness via performance along at least three distinct dimensions: 1) local abundance, 2) geographic range size, and 3) habitat breadth in naturalized distributions. Associations among these dimensions and the factors that determine success in each have yet to be assessed at large geographic scales. Here, we combine data from over one million vegetation plots covering the extent of Europe and its habitat diversity with databases on species' distributions, traits, and historical origins to provide a comprehensive assessment of invasiveness dimensions for the European alien seed plant flora. Invasiveness dimensions are linked in alien distributions, leading to a continuum from overall poor invaders to super invaders-abundant, widespread aliens that invade diverse habitats. This pattern echoes relationships among analogous dimensions measured for native European species. Success along invasiveness dimensions was associated with details of alien species' introduction histories: earlier introduction dates were positively associated with all three dimensions, and consistent with theory-based expectations, species originating from other continents, particularly acquisitive growth strategists, were among the most successful invaders in Europe. Despite general correlations among invasiveness dimensions, we identified habitats and traits associated with atypical patterns of success in only one or two dimensions-for example, the role of disturbed habitats in facilitating widespread specialists. We conclude that considering invasiveness within a multidimensional framework can provide insights into invasion processes while also informing general understanding of the dynamics of species distributions.

Economic use of plants is key to their naturalization success.

Humans cultivate thousands of economic plants (i.e. plants with economic value) outside their native ranges. To analyze how this contributes to naturalization success, we combine global databases on economic uses and naturalization success of the world's seed plants. Here we show that naturalization likelihood is 18 times higher for economic than non-economic plants. Naturalization success is highest for plants grown as animal food or for environmental uses (e.g. ornamentals), and increases with number of uses. Taxa from the Northern Hemisphere are disproportionately over-represented among economic plants, and economic plants from Asia have the greatest naturalization success. In regional naturalized floras, the percentage of economic plants exceeds the global percentage and increases towards the equator. Phylogenetic patterns in the naturalized flora partly result from phylogenetic patterns in the plants we cultivate. Our study illustrates that accounting for the intentional introduction of economic plants is key to unravelling drivers of plant naturalization.

Local species diversity, ß-diversity and climate influence the regional stability of bird biomass across North America.

Biodiversity often stabilizes aggregate ecosystem properties (e.g. biomass) at small spatial scales. However, the importance of species diversity within communities and variation in species composition among communities (ß-diversity) for stability at larger scales remains unclear. Using a continental-scale analysis of 1657 North American breeding-bird communities spanning 20-years and 35 ecoregions, we show local species diversity and ß-diversity influence two components of regional stability: local stability (stability of bird biomass within sites) and spatial asynchrony (asynchronous fluctuations in biomass among sites). We found spatial asynchrony explained three times more variation in regional stability of bird biomass than did local stability. This result contrasts with studies at smaller spatial scales-typically plant metacommunities under 1 ha-that find local stability to be more important than spatial asynchrony. Moreover, spatial asynchrony of bird biomass increased with bird ß-diversity and climate heterogeneity (temperature and precipitation), while local stability increased with species diversity. Our study reveals new insights into the scale-dependent processes regulating ecosystem stability, providing evidence that both local biodiversity loss and homogenization can destabilize ecosystem processes at biogeographic scales.

Cultural transmission and ecological opportunity jointly shaped global patterns of reliance on agriculture.

The evolution of agriculture improved food security and enabled significant increases in the size and complexity of human groups. Despite these positive effects, some societies never adopted these practices, became only partially reliant on them, or even reverted to foraging after temporarily adopting them. Given the critical importance of climate and biotic interactions for modern agriculture, it seems likely that ecological conditions could have played a major role in determining the degree to which different societies adopted farming. However, this seemingly simple proposition has been surprisingly difficult to prove and is currently controversial. Here, we investigate how recent agricultural practices relate both to contemporary ecological opportunities and the suitability of local environments for the first species domesticated by humans. Leveraging a globally distributed dataset on 1,291 traditional societies, we show that after accounting for the effects of cultural transmission and more current ecological opportunities, levels of reliance on farming continue to be predicted by the opportunities local ecologies provided to the first human domesticates even after centuries of cultural evolution. Based on the details of our models, we conclude that ecology probably helped shape the geography of agriculture by biasing both human movement and the human-assisted dispersal of domesticates.

Alternative ecological strategies lead to avian brain size bimodality in variable habitats.

The ecological contexts that promote larger brains have received considerable attention, but those that result in smaller-than-expected brains have been largely overlooked. Here, we use a global sample of 2062 species to provide evidence that metabolic and life history tradeoffs govern the evolution of brain size in birds and play an important role in defining the ecological strategies capable of persisting in Earth's most thermally variable and unpredictable habitats. While some birds cope with extreme winter conditions by investing in large brains (e.g., greater capacity for planning, innovation, and behavioral flexibility), others have small brains and invest instead in traits that allow them to withstand or recover from potentially deadly events. Specifically, these species are restricted to large body sizes, diets consisting of difficult-to-digest but readily available foods, and high reproductive output. Overall, our findings highlight the importance of considering strategic tradeoffs when investigating potential drivers of brain size evolution.

Big brains stabilize populations and facilitate colonization of variable habitats in birds.

The cognitive buffer hypothesis posits that environmental variability can be a major driver of the evolution of cognition because an enhanced ability to produce flexible behavioural responses facilitates coping with the unexpected. Although comparative evidence supports different aspects of this hypothesis, a direct connection between cognition and the ability to survive a variable and unpredictable environment has yet to be demonstrated. Here, we use complementary demographic and evolutionary analyses to show that among birds, the mechanistic premise of this hypothesis is well supported but the implied direction of causality is not. Specifically, we show that although population dynamics are more stable and less affected by environmental variation in birds with larger relative brain sizes, the evolution of larger brains often pre-dated and facilitated the colonization of variable habitats rather than the other way around. Our findings highlight the importance of investigating the timeline of evolutionary events when interpreting patterns of phylogenetic correlation.

Reconsidering alternative transportation systems to reach academic conferences and to convey an example to reduce greenhouse gas emissions.

Scientists are typically responsible for greater greenhouse gas emissions than the general population. These 'extra' emissions are largely due to frequent travel, often by airplane, to professional and academic meetings. In the following commentary, we explore how employing mixed modes of transportation, particularly by prioritizing train travel, can significantly reduce the environmental costs associated with attending conferences. Estimating travel distances for attendants to recent meetings, we demonstrate that the proposed strategy has the potential to decrease emissions, even when considering exotic, remote (and potentially enticing) locations. Our suggestions are easy to implement, can be combined with previous proposals for reducing emissions, and provide an opportunity for scientists to convey an important message to the general public. This message is not confined to our particular example and we hope that it will encourage ecologists, sociologists and other scientists, historians and philosophers to pursue additional creative behavioural solutions in order to reduce our impact on an increasingly stressed ecosphere.

Metabolic heat production and thermal conductance are mass-independent adaptations to thermal environment in birds and mammals.

The extent to which different kinds of organisms have adapted to environmental temperature regimes is central to understanding how they respond to climate change. The Scholander-Irving (S-I) model of heat transfer lays the foundation for explaining how endothermic birds and mammals maintain their high, relatively constant body temperatures in the face of wide variation in environmental temperature. The S-I model shows how body temperature is regulated by balancing the rates of heat production and heat loss. Both rates scale with body size, suggesting that larger animals should be better adapted to cold environments than smaller animals, and vice versa. However, the global distributions of ∼9,000 species of terrestrial birds and mammals show that the entire range of body sizes occurs in nearly all climatic regimes. Using physiological and environmental temperature data for 211 bird and 178 mammal species, we test for mass-independent adaptive changes in two key parameters of the S-I model: basal metabolic rate (BMR) and thermal conductance. We derive an axis of thermal adaptation that is independent of body size, extends the S-I model, and highlights interactions among physiological and morphological traits that allow endotherms to persist in a wide range of temperatures. Our macrophysiological and macroecological analyses support our predictions that shifts in BMR and thermal conductance confer important adaptations to environmental temperature in both birds and mammals.

Macroecology Meets Macroeconomics: Resource Scarcity and Global Sustainability.

The current economic paradigm, which is based on increasing human population, economic development, and standard of living, is no longer compatible with the biophysical limits of the finite Earth. Failure to recover from the economic crash of 2008 is not due just to inadequate fiscal and monetary policies. The continuing global crisis is also due to scarcity of critical resources. Our macroecological studies highlight the role in the economy of energy and natural resources: oil, gas, water, arable land, metals, rare earths, fertilizers, fisheries, and wood. As the modern industrial technological-informational economy expanded in recent decades, it grew by consuming the Earth's natural resources at unsustainable rates. Correlations between per capita GDP and per capita consumption of energy and other resources across nations and over time demonstrate how economic growth and development depend on "nature's capital". Decades-long trends of decreasing per capita consumption of multiple important commodities indicate that overexploitation has created an unsustainable bubble of population and economy.

The Malthusian-Darwinian dynamic and the trajectory of civilization.

Two interacting forces influence all populations: the Malthusian dynamic of exponential growth until resource limits are reached, and the Darwinian dynamic of innovation and adaptation to circumvent these limits through biological and/or cultural evolution. The specific manifestations of these forces in modern human society provide an important context for determining how humans can establish a sustainable relationship with the finite Earth.

The macroecology of sustainability.

The discipline of sustainability science has emerged in response to concerns of natural and social scientists, policymakers, and lay people about whether the Earth can continue to support human population growth and economic prosperity. Yet, sustainability science has developed largely independently from and with little reference to key ecological principles that govern life on Earth. A macroecological perspective highlights three principles that should be integral to sustainability science: 1) physical conservation laws govern the flows of energy and materials between human systems and the environment, 2) smaller systems are connected by these flows to larger systems in which they are embedded, and 3) global constraints ultimately limit flows at smaller scales. Over the past few decades, decreasing per capita rates of consumption of petroleum, phosphate, agricultural land, fresh water, fish, and wood indicate that the growing human population has surpassed the capacity of the Earth to supply enough of these essential resources to sustain even the current population and level of socioeconomic development.