The history of natural history and race: Decolonizing human dimensions of ecology.

Natural history, loosely defined as the observational study of organisms in the habitats where they occur, is recognized at the roots of ecology. Although the centrality of natural history in ecology has shifted over time, natural history is currently in resurgence: many again consider it to be the foundation of ecological and evolutionary inquiry and advocate the value of organism-centered approaches to address contemporary ecological challenges. Educators identify natural history as the foundational entryway into the practice of ecology, for example in the Ecological Society of America's Four-Dimensional Ecology Education (4DEE) framework. A strong natural history foundation can help generate testable hypotheses to refine mechanistic understanding of the drivers regulating species distributions and abundances and to inform restoration and conservation efforts. Given the resurgence of natural history as the foundation for ecological knowledge and practice, it is important to recognize that natural history has a long history of racism that has impacted ecological thought and priorities. This history shapes not only who conducts ecological science but also foundational ecological concepts. For example, natural history's emphasis on pristine nature untouched by humans disregards or appropriates stewardship and knowledge of most of the world's population. Because of the legacy of chattel slavery, this exclusion is particularly strong for people of African descent. This exclusion narrows ecological inquiry, limits the capacity to find solutions to ecological problems, and risks interventions that perpetuate the relation between eugenics, ecological knowledge, and natural systems. If ecology is to become an inclusive, responsive, and resilient discipline, this knowledge gap must be addressed. We here present the colonial and racist underpinnings of natural history and offer strategies to expand inclusion in the study of nature. Natural history was steeped in racism, providing a hierarchy of cultures and a taxonomy of races. Complementing growing interest in traditional and Indigenous ecological knowledge, we focus on Black ecological knowledge, for example in the study of "maroon ecologies." Addressing the racist history of natural history is necessary for removing structural and racist barriers to diverse participation and expanding ecological knowledge bases in service of better and more just science.

The identity crisis of ecological diversity.

Developing the ecological scientist mindset among underrepresented students in ecology fields (Bowser and Cid, this Forum) provides timely and compelling strategies to broaden inclusion in ecology and environmental biology. Chronic underrepresentation of minorities in ecology and environmental disciplines (EE) is a crisis that is surprising to many, and even more surprising that, for African-Americans, this underrepresentation is more severe compared to other STEM disciplines. It is beyond irony that a discipline that values diversity as a cornerstone of ecological practice continues to struggle to achieve diversity in the ranks of its practitioners.

Overcoming Blind Spots to Promote Environmental Justice Research.

Ecological research includes social-ecological-evolutionary processes, but the intersectionality and feedbacks between ecology and environmental justice (EJ) remain low. We here present opportunities for ecological research contributions to EJ. Ultimately, such work can reduce racial and social disparities in environmental provisioning and improve global environmental sustainability.

Advancing an interdisciplinary framework to study seed dispersal ecology.

Although dispersal is generally viewed as a crucial determinant for the fitness of any organism, our understanding of its role in the persistence and spread of plant populations remains incomplete. Generalizing and predicting dispersal processes are challenging due to context dependence of seed dispersal, environmental heterogeneity and interdependent processes occurring over multiple spatial and temporal scales. Current population models often use simple phenomenological descriptions of dispersal processes, limiting their ability to examine the role of population persistence and spread, especially under global change. To move seed dispersal ecology forward, we need to evaluate the impact of any single seed dispersal event within the full spatial and temporal context of a plant's life history and environmental variability that ultimately influences a population's ability to persist and spread. In this perspective, we provide guidance on integrating empirical and theoretical approaches that account for the context dependency of seed dispersal to improve our ability to generalize and predict the consequences of dispersal, and its anthropogenic alteration, across systems. We synthesize suitable theoretical frameworks for this work and discuss concepts, approaches and available data from diverse subdisciplines to help operationalize concepts, highlight recent breakthroughs across research areas and discuss ongoing challenges and open questions. We address knowledge gaps in the movement ecology of seeds and the integration of dispersal and demography that could benefit from such a synthesis. With an interdisciplinary perspective, we will be able to better understand how global change will impact seed dispersal processes, and potential cascading effects on plant population persistence, spread and biodiversity.

Employing plant functional groups to advance seed dispersal ecology and conservation.

Seed dispersal enables plants to reach hospitable germination sites and escape natural enemies. Understanding when and how much seed dispersal matters to plant fitness is critical for understanding plant population and community dynamics. At the same time, the complexity of factors that determine if a seed will be successfully dispersed and subsequently develop into a reproductive plant is daunting. Quantifying all factors that may influence seed dispersal effectiveness for any potential seed-vector relationship would require an unrealistically large amount of time, materials and financial resources. On the other hand, being able to make dispersal predictions is critical for predicting whether single species and entire ecosystems will be resilient to global change. Building on current frameworks, we here posit that seed dispersal ecology should adopt plant functional groups as analytical units to reduce this complexity to manageable levels. Functional groups can be used to distinguish, for their constituent species, whether it matters (i) if seeds are dispersed, (ii) into what context they are dispersed and (iii) what vectors disperse them. To avoid overgeneralization, we propose that the utility of these functional groups may be assessed by generating predictions based on the groups and then testing those predictions against species-specific data. We suggest that data collection and analysis can then be guided by robust functional group definitions. Generalizing across similar species in this way could help us to better understand the population and community dynamics of plants and tackle the complexity of seed dispersal as well as its disruption.

Growth and fecundity of fertile Miscanthus × giganteus ("PowerCane") compared to feral and ornamental Miscanthus sinensis in a common garden experiment: Implications for invasion.

Perennial grasses are promising candidates for bioenergy crops, but species that can escape cultivation and establish self-sustaining naturalized populations (feral) may have the potential to become invasive. Fertile Miscanthus × giganteus, known as "PowerCane," is a new potential biofuel crop. Its parent species are ornamental, non-native Miscanthus species that establish feral populations and are sometimes invasive in the USA. As a first step toward assessing the potential for "PowerCane" to become invasive, we documented its growth and fecundity relative to one of its parent species (Miscanthus sinensis) in competition with native and invasive grasses in common garden experiments located in Columbus, Ohio and Ames, Iowa, within the targeted range of biofuel cultivation. We conducted a 2-year experiment to compare growth and reproduction among three Miscanthus biotypes-"PowerCane," ornamental M. sinensis, and feral M. sinensis-at two locations. Single Miscanthus plants were subjected to competition with a native grass (Panicum virgatum), a weedy grass (Bromus inermis), or no competition. Response variables were aboveground biomass, number of shoots, basal area, and seed set. In Iowa, all Miscanthus plants died after the first winter, which was unusually cold, so no further results are reported from the Iowa site. In Ohio, we found significant differences among biotypes in growth and fecundity, as well as significant effects of competition. Interactions between these treatments were not significant. "PowerCane" performed as well or better than ornamental or feral M. sinensis in vegetative traits, but had much lower seed production, perhaps due to pollen limitation. In general, ornamental M. sinensis performed somewhat better than feral M. sinensis. Our findings suggest that feral populations of "PowerCane" could become established adjacent to biofuel production areas. Fertile Miscanthus × giganteus should be studied further to assess its potential to spread via seed production in large, sexually compatible populations.

Contemporary evolution and the dynamics of invasion in crop-wild hybrids with heritable variation for two weedy life-histories.

Gene flow in crop-wild complexes between phenotypically differentiated ancestors may transfer adaptive genetic variation that alters the fecundity and, potentially, the population growth (λ) of weeds. We created biotypes with potentially invasive traits, early flowering or long leaves, in wild radish (Raphanus raphanistrum) and F5 crop-wild hybrid (R. sativus × R. raphanistrum) backgrounds and compared them to randomly mated populations, to provide the first experimental estimate of long-term fitness consequences of weedy life-history variation. Using a life table response experiment design, we modeled λ of experimental, field populations in Pellston, MI, and assessed the relative success of alternative weed strategies and the contributions of individual vital rates (germination, survival, seed production) to differences in λ among experimental populations. Growth rates (λ) were most influenced by seed production, a trait altered by hybridization and selection, compared to other vital rates. More seeds were produced by wild than hybrid populations and by long-leafed than early-flowering lineages. Although we did not detect a biotype by selection treatment effect on lambda, lineages also exhibited contrasting germination and survival strategies. Identifying life-history traits affecting population growth contributes to our understanding of which portions of the crop genome are most likely to introgress into weed populations.

Demographic consequences of greater clonal than sexual reproduction in Dicentra canadensis.

Clonality is a widespread life history trait in flowering plants that may be essential for population persistence, especially in environments where sexual reproduction is unpredictable. Frequent clonal reproduction, however, could hinder sexual reproduction by spatially aggregating ramets that compete with seedlings and reduce inter-genet pollination. Nevertheless, the role of clonality in relation to variable sexual reproduction in population dynamics is often overlooked. We combined population matrix models and pollination experiments to compare the demographic contributions of clonal and sexual reproduction in three Dicentra canadensis populations, one in a well-forested landscape and two in isolated forest remnants. We constructed stage-based transition matrices from 3 years of census data to evaluate annual population growth rates, λ. We used loop analysis to evaluate the relative contribution of different reproductive pathways to λ. Despite strong temporal and spatial variation in seed set, populations generally showed stable growth rates. Although we detected some pollen limitation of seed set, manipulative pollination treatments did not affect population growth rates. Clonal reproduction contributed significantly more than sexual reproduction to population growth in the forest remnants. Only at the well-forested site did sexual reproduction contribute as much as clonal reproduction to population growth. Flowering plants were more likely to transition to a smaller size class with reduced reproductive potential in the following year than similarly sized nonflowering plants, suggesting energy trade-offs between sexual and clonal reproduction at the individual level. Seed production had negligible effects on growth and tuber production of individual plants. Our results demonstrate that clonal reproduction is vital for population persistence in a system where sexual reproduction is unpredictable. The bias toward clonality may be driven by low fitness returns for resource investment in sexual reproduction at the individual level. However, chronic failure in sexual reproduction may exacerbate the imbalance between sexual and clonal reproduction and eventually lead to irreversible loss of sex in the population.

Increasing aridity reduces soil microbial diversity and abundance in global drylands.

Soil bacteria and fungi play key roles in the functioning of terrestrial ecosystems, yet our understanding of their responses to climate change lags significantly behind that of other organisms. This gap in our understanding is particularly true for drylands, which occupy ∼41% of Earth´s surface, because no global, systematic assessments of the joint diversity of soil bacteria and fungi have been conducted in these environments to date. Here we present results from a study conducted across 80 dryland sites from all continents, except Antarctica, to assess how changes in aridity affect the composition, abundance, and diversity of soil bacteria and fungi. The diversity and abundance of soil bacteria and fungi was reduced as aridity increased. These results were largely driven by the negative impacts of aridity on soil organic carbon content, which positively affected the abundance and diversity of both bacteria and fungi. Aridity promoted shifts in the composition of soil bacteria, with increases in the relative abundance of Chloroflexi and α-Proteobacteria and decreases in Acidobacteria and Verrucomicrobia. Contrary to what has been reported by previous continental and global-scale studies, soil pH was not a major driver of bacterial diversity, and fungal communities were dominated by Ascomycota. Our results fill a critical gap in our understanding of soil microbial communities in terrestrial ecosystems. They suggest that changes in aridity, such as those predicted by climate-change models, may reduce microbial abundance and diversity, a response that will likely impact the provision of key ecosystem services by global drylands.

Climate and soil attributes determine plant species turnover in global drylands.

AIM: Geographic, climatic, and soil factors are major drivers of plant beta diversity, but their importance for dryland plant communities is poorly known. This study aims to: i) characterize patterns of beta diversity in global drylands, ii) detect common environmental drivers of beta diversity, and iii) test for thresholds in environmental conditions driving potential shifts in plant species composition. LOCATION: 224 sites in diverse dryland plant communities from 22 geographical regions in six continents. METHODS: Beta diversity was quantified with four complementary measures: the percentage of singletons (species occurring at only one site), Whittake's beta diversity (ß(W)), a directional beta diversity metric based on the correlation in species occurrences among spatially contiguous sites (ß(R2)), and a multivariate abundance-based metric (ß(MV)). We used linear modelling to quantify the relationships between these metrics of beta diversity and geographic, climatic, and soil variables. RESULTS: Soil fertility and variability in temperature and rainfall, and to a lesser extent latitude, were the most important environmental predictors of beta diversity. Metrics related to species identity (percentage of singletons and ß(W)) were most sensitive to soil fertility, whereas those metrics related to environmental gradients and abundance ((ß(R2)) and ß(MV)) were more associated with climate variability. Interactions among soil variables, climatic factors, and plant cover were not important determinants of beta diversity. Sites receiving less than 178 mm of annual rainfall differed sharply in species composition from more mesic sites (> 200 mm). MAIN CONCLUSIONS: Soil fertility and variability in temperature and rainfall are the most important environmental predictors of variation in plant beta diversity in global drylands. Our results suggest that those sites annually receiving ~ 178 mm of rainfall will be especially sensitive to future climate changes. These findings may help to define appropriate conservation strategies for mitigating effects of climate change on dryland vegetation.

Decoupling of soil nutrient cycles as a function of aridity in global drylands.

The biogeochemical cycles of carbon (C), nitrogen (N) and phosphorus (P) are interlinked by primary production, respiration and decomposition in terrestrial ecosystems. It has been suggested that the C, N and P cycles could become uncoupled under rapid climate change because of the different degrees of control exerted on the supply of these elements by biological and geochemical processes. Climatic controls on biogeochemical cycles are particularly relevant in arid, semi-arid and dry sub-humid ecosystems (drylands) because their biological activity is mainly driven by water availability. The increase in aridity predicted for the twenty-first century in many drylands worldwide may therefore threaten the balance between these cycles, differentially affecting the availability of essential nutrients. Here we evaluate how aridity affects the balance between C, N and P in soils collected from 224 dryland sites from all continents except Antarctica. We find a negative effect of aridity on the concentration of soil organic C and total N, but a positive effect on the concentration of inorganic P. Aridity is negatively related to plant cover, which may favour the dominance of physical processes such as rock weathering, a major source of P to ecosystems, over biological processes that provide more C and N, such as litter decomposition. Our findings suggest that any predicted increase in aridity with climate change will probably reduce the concentrations of N and C in global drylands, but increase that of P. These changes would uncouple the C, N and P cycles in drylands and could negatively affect the provision of key services provided by these ecosystems.

Plant species richness and ecosystem multifunctionality in global drylands.

Experiments suggest that biodiversity enhances the ability of ecosystems to maintain multiple functions, such as carbon storage, productivity, and the buildup of nutrient pools (multifunctionality). However, the relationship between biodiversity and multifunctionality has never been assessed globally in natural ecosystems. We report here on a global empirical study relating plant species richness and abiotic factors to multifunctionality in drylands, which collectively cover 41% of Earth's land surface and support over 38% of the human population. Multifunctionality was positively and significantly related to species richness. The best-fitting models accounted for over 55% of the variation in multifunctionality and always included species richness as a predictor variable. Our results suggest that the preservation of plant biodiversity is crucial to buffer negative effects of climate change and desertification in drylands.

Twenty years of changes in spatial association and community structure among desert perennials.

I present results from analyses of 20 years of spatiotemporal dynamics in a desert perennial community. Plants were identified and mapped in a 1-ha permanent plot in Joshua Tree National Park (California, USA) in 1984. Plant size, mortality, and new seedlings were censused every five years through 2004. Two species, Ambrosia dumosa and Tetracoccus hallii, were dominant based on their relative abundance and ubiquitous distributions. Spatial analysis for distance indices (SADIE) identified regions of significantly high (patches) or low (gaps) densities. I used SADIE to test for (1) transience in the distribution of patches and gaps within species over time and (2) changes in juvenile-adult associations with conspecific adults and adults of the two dominant species over time. Plant performance was quantified in patches and gaps to determine plant responsiveness to local spatial associations. Species identity was found to influence associations between juveniles and adults. Juveniles of all species showed significant positive spatial associations with the dominant A. dumosa but not with T. hallii. The broad distribution of A. dumosa may increase the spatial extent of non-dominant species that are facilitated by this dominant. The spatial location of patches and gaps was generally consistent over time for adults but not juveniles. Observed variability in the locations of juvenile patches and gaps suggested that suitable locations for establishment were broad relative to occupied regions of the habitat, and that conditions for seed germination were independent of conditions for seedling survival. A dramatic change in spatial distributions and associations within and between species occurred after a major drought that influenced data from the final census. Positive associations between juveniles and adults of all species were found independent of previous associations and most species distributions contracted to areas that were previously characterized by low density. By linking performance to spatial distribution, results from this study offer a spatial context for plant-plant interactions within and among species. Community composition could be influenced both by individual species tolerances of abiotic conditions and by the competitive or facilitative interactions individuals exert over neighbors.

Episodic death across species of desert shrubs.

Extreme events shape population and community trajectories. We report episodic mortality across common species of thousands of long-lived perennials individually tagged and monitored for 20 years in the Colorado Desert of California following severe regional drought. Demographic records from 1984 to 2004 show 15 years of virtual stasis in populations of adult shrubs and cacti, punctuated by a 55-100% die-off of six of the seven most common perennial species. In this episode, adults that experienced reduced growth in a lesser drought during 1984-1989 failed to survive the drought of 2002. The significance of this event is potentially profound because population dynamics of long-lived plants can be far more strongly affected by deaths of adults, which in deserts potentially live for centuries, than by seedling births or deaths. Differential mortality and rates of recovery during and after extreme climatic events quite likely determine the species composition of plant and associated animal communities for at least decades. The die-off recorded in this closely monitored community provides a unique window into the mechanics of this process of species decline and replacement.