Ecology of the Anthropocene signals hope for consciously managing the planetary ecosystem.

Human populations have grown to such an extent that our species has become a dominant force on the planet, prompting geologists to begin applying the term Anthropocene to recognize the present moment. Many approaches seek to explain the past and future of human population growth, in the form of narratives and models. Some of the most influential models have parameters that cannot be precisely known but are estimated by expert opinion. Here we apply a unified model of ecology to provide a macroscale summary of the net effects of many microscale processes, using a minimal set of parameters that can be known. Our models match estimates of historic and prehistoric global human population numbers and provide predictions that correspond to some of the more complicated current models. In addition to fitting the data well they reveal that, amidst enormous complexity in our human and prehuman past, three key ecological discontinuities have occurred in turn: 1) becoming dominant competitors of large predators rather than their prey, 2) becoming mutualists with food species rather than acting as predators upon them, and 3) changing from a regime of uncontrolled population growth to one of controlled fertility instead. All three processes have been interlinked with cultural evolution and all three ushered in developments of the Anthropocene. Understanding the trajectories that have delivered us to this stage can help guide prudent paths into the future.

Identifying mechanisms that structure ecological communities by snapping model parameters to empirically observed tradeoffs.

Theory predicts that interspecific tradeoffs are primary determinants of coexistence and community composition. Using information from empirically observed tradeoffs to augment the parametrisation of mechanism-based models should therefore improve model predictions, provided that tradeoffs and mechanisms are chosen correctly. We developed and tested such a model for 35 grassland plant species using monoculture measurements of three species characteristics related to nitrogen uptake and retention, which previous experiments indicate as important at our site. Matching classical theoretical expectations, these characteristics defined a distinct tradeoff surface, and models parameterised with these characteristics closely matched observations from experimental multi-species mixtures. Importantly, predictions improved significantly when we incorporated information from tradeoffs by 'snapping' characteristics to the nearest location on the tradeoff surface, suggesting that the tradeoffs and mechanisms we identify are important determinants of local community structure. This 'snapping' method could therefore constitute a broadly applicable test for identifying influential tradeoffs and mechanisms.

Impact of simian immunodeficiency virus infection on chimpanzee population dynamics.

Like human immunodeficiency virus type 1 (HIV-1), simian immunodeficiency virus of chimpanzees (SIVcpz) can cause CD4+ T cell loss and premature death. Here, we used molecular surveillance tools and mathematical modeling to estimate the impact of SIVcpz infection on chimpanzee population dynamics. Habituated (Mitumba and Kasekela) and non-habituated (Kalande) chimpanzees were studied in Gombe National Park, Tanzania. Ape population sizes were determined from demographic records (Mitumba and Kasekela) or individual sightings and genotyping (Kalande), while SIVcpz prevalence rates were monitored using non-invasive methods. Between 2002-2009, the Mitumba and Kasekela communities experienced mean annual growth rates of 1.9% and 2.4%, respectively, while Kalande chimpanzees suffered a significant decline, with a mean growth rate of -6.5% to -7.4%, depending on population estimates. A rapid decline in Kalande was first noted in the 1990s and originally attributed to poaching and reduced food sources. However, between 2002-2009, we found a mean SIVcpz prevalence in Kalande of 46.1%, which was almost four times higher than the prevalence in Mitumba (12.7%) and Kasekela (12.1%). To explore whether SIVcpz contributed to the Kalande decline, we used empirically determined SIVcpz transmission probabilities as well as chimpanzee mortality, mating and migration data to model the effect of viral pathogenicity on chimpanzee population growth. Deterministic calculations indicated that a prevalence of greater than 3.4% would result in negative growth and eventual population extinction, even using conservative mortality estimates. However, stochastic models revealed that in representative populations, SIVcpz, and not its host species, frequently went extinct. High SIVcpz transmission probability and excess mortality reduced population persistence, while intercommunity migration often rescued infected communities, even when immigrating females had a chance of being SIVcpz infected. Together, these results suggest that the decline of the Kalande community was caused, at least in part, by high levels of SIVcpz infection. However, population extinction is not an inevitable consequence of SIVcpz infection, but depends on additional variables, such as migration, that promote survival. These findings are consistent with the uneven distribution of SIVcpz throughout central Africa and explain how chimpanzees in Gombe and elsewhere can be at equipoise with this pathogen.

Diversifying bioenergy crops increases yield and yield stability by reducing weed abundance.

Relationships between species diversity, productivity, temporal stability of productivity, and plant invasion have been well documented in grasslands, and these relationships could translate to improved agricultural sustainability. However, few studies have explored these relationships in agricultural contexts where fertility and weeds are managed. Using 7 years of biomass yield and species composition data from 12 species mixture treatments varying in native species diversity, we found that species richness increased yield and interannual yield stability by reducing weed abundance. Stability was driven by yield as opposed to temporal variability of yield. Nitrogen fertilization increased yield but at the expense of yield stability. We show how relationships between diversity, species asynchrony, invasion, productivity, and stability observed in natural grasslands can extend into managed agricultural systems. Increasing bioenergy crop diversity can improve farmer economics via increased yield, reduced yield variability, and reduced inputs for weed control, thus promoting perennial vegetation on agricultural lands.

Soil carbon sequestration accelerated by restoration of grassland biodiversity.

Agriculturally degraded and abandoned lands can remove atmospheric CO2 and sequester it as soil organic matter during natural succession. However, this process may be slow, requiring a century or longer to re-attain pre-agricultural soil carbon levels. Here, we find that restoration of late-successional grassland plant diversity leads to accelerating annual carbon storage rates that, by the second period (years 13-22), are 200% greater in our highest diversity treatment than during succession at this site, and 70% greater than in monocultures. The higher soil carbon storage rates of the second period (years 13-22) are associated with the greater aboveground production and root biomass of this period, and with the presence of multiple species, especially C4 grasses and legumes. Our results suggest that restoration of high plant diversity may greatly increase carbon capture and storage rates on degraded and abandoned agricultural lands.

Fluctuating Selection in the Moran.

Contrary to classical population genetics theory, experiments demonstrate that fluctuating selection can protect a haploid polymorphism in the absence of frequency dependent effects on fitness. Using forward simulations with the Moran model, we confirm our analytical results showing that a fluctuating selection regime, with a mean selection coefficient of zero, promotes polymorphism. We find that increases in heterozygosity over neutral expectations are especially pronounced when fluctuations are rapid, mutation is weak, the population size is large, and the variance in selection is big. Lowering the frequency of fluctuations makes selection more directional, and so heterozygosity declines. We also show that fluctuating selection raises dn /ds ratios for polymorphism, not only by sweeping selected alleles into the population, but also by purging the neutral variants of selected alleles as they undergo repeated bottlenecks. Our analysis shows that randomly fluctuating selection increases the rate of evolution by increasing the probability of fixation. The impact is especially noticeable when the selection is strong and mutation is weak. Simulations show the increase in the rate of evolution declines as the rate of new mutations entering the population increases, an effect attributable to clonal interference. Intriguingly, fluctuating selection increases the dn /ds ratios for divergence more than for polymorphism, a pattern commonly seen in comparative genomics. Our model, which extends the classical neutral model of molecular evolution by incorporating random fluctuations in selection, accommodates a wide variety of observations, both neutral and selected, with economy.

Conventional functional classification schemes underestimate the relationship with ecosystem functioning.

Studies linking the functional diversity of a biota to ecosystem functioning typically employ a priori classifications of species into hypothetically complementary groups. However, multiple alternate classifications exist in which the number of functional groups, the number of species per functional group, and the grouping of species differ from the a priori scheme. Without assessing the relative precision, or ability of an a priori scheme to accurately predict ecosystem functioning relative to its many alternatives, the validity and utility of analyses based on a single a priori classification scheme remains unclear. We examine the precision of a priori classifications used in 10 experimental grassland systems in Europe and the United States that have found evidence for a significant role of functional plant diversity in governing ecosystem function. The predictive precision of the a priori classifications employed in these studies was seldom significantly higher than the precision of random classifications. Post-hoc classification schemes that performed well in predicting ecosystem function resembled each other more with regard to species composition than average classifications, but there was still considerable variability in the manner in which these classification schemes grouped species. These results suggest that we need a more nuanced understanding of how the diversity of functional traits of species in an assemblage affects ecosystem functioning.

Biodiversity, Stability, and Productivity in Competitive Communities.

Three markedly different models of multispecies competition-one mechanistic, one phenomenological, and one statistical-all predict that greater diversity increases the temporal stability of the entire community, decreases the temporal stability of individual populations, and increases community productivity. We define temporal stability as the ratio of mean abundance to its standard deviation. Interestingly, the temporal stability of entire communities is predicted to increase fairly linearly, without clear saturation, as diversity increases. Species composition is predicted to be as important as diversity in affecting community stability and productivity. The greater temporal stability of more diverse communities is caused by higher productivity at higher diversity (the "overyielding" effect), competitive interactions (the "covariance" effect), and statistical averaging (the "portfolio" effect). The relative contribution of each cause of temporal stability changes as diversity increases, but the net effect is that greater diversity stabilizes the community even though it destabilizes individual populations. This theory agrees with recent experiments and provides a degree of resolution to the diversity-stability debate: both sides of the longstanding debate were correct, but one addressed population stability and the other addressed community stability.

Modeling of bovine spongiform encephalopathy in a two-species feedback loop.

Bovine spongiform encephalopathy, otherwise known as mad cow disease, can spread when an individual cow consumes feed containing the infected tissues of another individual, forming a one-species feedback loop. Such feedback is the primary means of transmission for BSE during epidemic conditions. Following outbreaks in the European Union and elsewhere, many governments enacted legislation designed to limit the spread of such diseases via elimination or reduction of one-species feedback loops in agricultural systems. However, two-species feedback loops-those in which infectious material from one-species is consumed by a secondary species whose tissue is then consumed by the first species-were not universally prohibited and have not been studied before. Here we present a basic ecological disease model which examines the rôle feedback loops may play in the spread of BSE and related diseases. Our model shows that there are critical thresholds between the infection's expansion and decrease related to the lifespan of the hosts, the growth rate of the prions, and the amount of prions circulating between hosts. The ecological disease dynamics can be intrinsically oscillatory, having outbreaks as well as refractory periods which can make it appear that the disease is under control while it is still increasing. We show that non-susceptible species that have been intentionally inserted into a feedback loop to stop the spread of disease do not, strictly by themselves, guarantee its control, though they may give that appearance by increasing the refractory period of an epidemic's oscillations. We suggest ways in which age-related dynamics and cross-species coupling should be considered in continuing evaluations aimed at maintaining a safe food supply.

Energy potential of biomass from conservation grasslands in Minnesota, USA.

Perennial biomass from grasslands managed for conservation of soil and biodiversity can be harvested for bioenergy. Until now, the quantity and quality of harvestable biomass from conservation grasslands in Minnesota, USA, was not known, and the factors that affect bioenergy potential from these systems have not been identified. We measured biomass yield, theoretical ethanol conversion efficiency, and plant tissue nitrogen (N) as metrics of bioenergy potential from mixed-species conservation grasslands harvested with commercial-scale equipment. With three years of data, we used mixed-effects models to determine factors that influence bioenergy potential. Sixty conservation grassland plots, each about 8 ha in size, were distributed among three locations in Minnesota. Harvest treatments were applied annually in autumn as a completely randomized block design. Biomass yield ranged from 0.5 to 5.7 Mg ha(-1). May precipitation increased biomass yield while precipitation in all other growing season months showed no affect. Averaged across all locations and years, theoretical ethanol conversion efficiency was 450 l Mg(-1) and the concentration of plant N was 7.1 g kg(-1), both similar to dedicated herbaceous bioenergy crops such as switchgrass. Biomass yield did not decline in the second or third year of harvest. Across years, biomass yields fluctuated 23% around the average. Surprisingly, forb cover was a better predictor of biomass yield than warm-season grass with a positive correlation with biomass yield in the south and a negative correlation at other locations. Variation in land ethanol yield was almost exclusively due to variation in biomass yield rather than biomass quality; therefore, efforts to increase biomass yield might be more economical than altering biomass composition when managing conservation grasslands for ethanol production. Our measurements of bioenergy potential, and the factors that control it, can serve as parameters for assessing the economic viability of harvesting conservation grasslands for bioenergy.

Entrepreneurs, chance, and the deterministic concentration of wealth.

In many economies, wealth is strikingly concentrated. Entrepreneurs--individuals with ownership in for-profit enterprises--comprise a large portion of the wealthiest individuals, and their behavior may help explain patterns in the national distribution of wealth. Entrepreneurs are less diversified and more heavily invested in their own companies than is commonly assumed in economic models. We present an intentionally simplified individual-based model of wealth generation among entrepreneurs to assess the role of chance and determinism in the distribution of wealth. We demonstrate that chance alone, combined with the deterministic effects of compounding returns, can lead to unlimited concentration of wealth, such that the percentage of all wealth owned by a few entrepreneurs eventually approaches 100%. Specifically, concentration of wealth results when the rate of return on investment varies by entrepreneur and by time. This result is robust to inclusion of realities such as differing skill among entrepreneurs. The most likely overall growth rate of the economy decreases as businesses become less diverse, suggesting that high concentrations of wealth may adversely affect a country's economic growth. We show that a tax on large inherited fortunes, applied to a small portion of the most fortunate in the population, can efficiently arrest the concentration of wealth at intermediate levels.

Carbon-negative biofuels from low-input high-diversity grassland biomass.

Biofuels derived from low-input high-diversity (LIHD) mixtures of native grassland perennials can provide more usable energy, greater greenhouse gas reductions, and less agrichemical pollution per hectare than can corn grain ethanol or soybean biodiesel. High-diversity grasslands had increasingly higher bioenergy yields that were 238% greater than monoculture yields after a decade. LIHD biofuels are carbon negative because net ecosystem carbon dioxide sequestration (4.4 megagram hectare(-1) year(-1) of carbon dioxide in soil and roots) exceeds fossil carbon dioxide release during biofuel production (0.32 megagram hectare(-1) year(-1)). Moreover, LIHD biofuels can be produced on agriculturally degraded lands and thus need to neither displace food production nor cause loss of biodiversity via habitat destruction.