Simon A. Levin

Effective leadership and decision-making in animal groups on the move

For animals that forage or travel in groups, making movement decisions often depends on social interactions among group members. However, in many cases, few individuals have pertinent information, such as knowledge about the location of a food source, or of a migration route. Using a simple model we show how information can be transferred within groups both without signalling and when group members do not know which individuals, if any, have information. We reveal that the larger the group the smaller the proportion of informed individuals needed to guide the group, and that only a very small proportion of informed individuals is required to achieve great accuracy. We also demonstrate how groups can make consensus decisions, even though informed individuals do not know whether they are in a majority or minority, how the quality of their information compares with that of others, or even whether there are any other informed individuals. Our model provides new insights into the mechanisms of effective leadership and decision-making in biological systems.

Intertidal Landscapes: Disturbance and the Dynamics of Pattern

The mussel Mytilus californianus is a competitive dominant on wave—swept rocky intertidal shores. Mussel beds may exist as extensive monocultures; more often they are an everchanging mosaic of many species which inhabit wave—generated patches or gaps. This paper describes observations and experiments designed to measure the critical parameters of a model of patch birth and death, and to use the model to predict the spatial structure of mussel beds. Most measurements were made at Tatoosh Island, Washington, USA, from 1970—1979. Patch size ranged at birth from a single mussel to 38 m2; the distribution of patch sizes approximates the lognormal. Birth rates varied seasonally and regionally. At Tatoosh the rate of patch formation varied during six winters from 0.4—5.4% of the mussels removed per month. The disturbance regime during the summer and at two mainland sites was 5—10 times less. Annual disturbance patterns tended to be synchronous within 11 sites on one face of Tatoosh over a 10—yr interval, and over larger distances (16 km) along the coastline. The pattern was asynchronous, however, among four Tatoosh localities. Patch birth rate, and mean and maximum size at birth can be used as adequate indices of disturbance. Patch disappearance (death) occurs by three mechanisms. Very small patches disappear almost immediately due to a leaning response of the border mussels (0.2 cm/d). Intermediate—sized patches (<3.0 m2) are eventually obliterated by lateral movement of the peripheral mussels: estimates based on 94 experimental patches yield a mean shrinking rate of 0.05 cm/d from each of two principal dimensions. Depth of the adjacent mussel bed accounts for much of the local variation in closing rate. In very large patches, mussels must recruit as larvae from the plankton. Recovery begins at an average patch age of 26 mo; rate of space occupation, primarily due to individual growth, is 2.0—2.5%/mo. Winter birth rates suggest a mean turnover time (rotation period) for mussel beds varying from 8.1—34.7 yr, depending on the location. The minimal value is in close agreement with both observed and calculated minimal recovery times. Projections of total patch area, based on the model, are accurate to within 5% of the observed. Using a method for determining the age of patches, based on a growth curve of the barnacle Balanus cariosus, the model permits predictions of the age—size structure of the patch population. The model predicts with excellent resolution the distribution of patch area in relation to time since last disturbance. The most detailed models which include size structure within age categories are inconclusive due to small sample size. Predictions are food for large patches, the major determinants of environmental patterns, but cannot deal adequately with smaller patches because of stochastic effects. Colonization data are given in relation to patch age, size and intertidal position. We suggest that the reproductive season of certain long—lived, patch—dependent species is moulded by the disturbance regime. The necessary and vital connection between disturbance which generates spatial pattern and species richness in communities open to invasion is discussed.

Ecosystems and the Biosphere as Complex Adaptive Systems

ABSTRACT Ecosystems are prototypical examples of complex adaptive systems, in which patterns at higher levels emerge from localized interactions and selection processes acting at lower levels. An essential aspect of such systems is nonlinearity, leading to historical dependency and multiple possible outcomes of dynamics. Given this, it is essential to determine the degree to which system features are determined by environmental conditions, and the degree to which they are the result of self-organization. Furthermore, given the multiple levels at which dynamics become apparent and at which selection can act, central issues relate to how evolution shapes ecosystems properties, and whether ecosystems become buffered to changes (more resilient) over their ecological and evolutionary development or proceed to critical states and the edge of chaos.

Anticipating Critical Transitions

All Change Research on early warning signals for critical transitions in complex systems such as ecosystems, climate, and global finance systems recently has been gathering pace. At the same time, studies on complex networks are starting to reveal which architecture may cause systems to be vulnerable to systemic collapse. Scheffer et al. (p. 344) review how previously isolated lines of work can be connected, conclude that many critical transitions (such as escape from the poverty trap) can have positive outcomes, and highlight how the new approaches to sensing fragility can help to detect both risks and opportunities for desired change. Tipping points in complex systems may imply risks of unwanted collapse, but also opportunities for positive change. Our capacity to navigate such risks and opportunities can be boosted by combining emerging insights from two unconnected fields of research. One line of work is revealing fundamental architectural features that may cause ecological networks, financial markets, and other complex systems to have tipping points. Another field of research is uncovering generic empirical indicators of the proximity to such critical thresholds. Although sudden shifts in complex systems will inevitably continue to surprise us, work at the crossroads of these emerging fields offers new approaches for anticipating critical transitions.

Are We Consuming Too Much

This paper articulates and applies frameworks for examining whether consumption is excessive. We consider two criteria for the possible excessiveness (or insufficiency) of current consumption. One is an intertemporal utility-maximization criterion: actual current consumption is deemed excessive if it is higher than the level of current consumption on the consumption path that maximizes the present discounted value of utility. The other is a sustainability criterion, which requires that current consumption be consistent with non-declining living standards over time. We extend previous theoretical approaches by offering a formula for the sustainability criterion that accounts for population growth and technological change. In applying this formula, we find that some poor regions of the world are failing to meet the sustainability criterion: in these regions, genuine wealth per capita is falling as investments in human and manufactured capital are not sufficient to offset the depletion of natural capital.

Optimal nitrogen-to-phosphorus stoichiometry of phytoplankton

Redfield noted the similarity between the average nitrogen-to-phosphorus ratio in plankton (N:P = 16 by atoms) and in deep oceanic waters (N:P = 15; refs 1, 2). He argued that this was neither a coincidence, nor the result of the plankton adapting to the oceanic stoichiometry, but rather that phytoplankton adjust the N:P stoichiometry of the ocean to meet their requirements through nitrogen fixation, an idea supported by recent modelling studies. But what determines the N:P requirements of phytoplankton? Here we use a stoichiometrically explicit model of phytoplankton physiology and resource competition to derive from first principles the optimal phytoplankton stoichiometry under diverse ecological scenarios. Competitive equilibrium favours greater allocation to P-poor resource-acquisition machinery and therefore a higher N:P ratio; exponential growth favours greater allocation to P-rich assembly machinery and therefore a lower N:P ratio. P-limited environments favour slightly less allocation to assembly than N-limited or light-limited environments. The model predicts that optimal N:P ratios will vary from 8.2 to 45.0, depending on the ecological conditions. Our results show that the canonical Redfield N:P ratio of 16 is not a universal biochemical optimum, but instead represents an average of species-specific N:P ratios.

Global antibiotic consumption 2000 to 2010: an analysis of national pharmaceutical sales data

BACKGROUND Antibiotic drug consumption is a major driver of antibiotic resistance. Variations in antibiotic resistance across countries are attributable, in part, to different volumes and patterns for antibiotic consumption. We aimed to assess variations in consumption to assist monitoring of the rise of resistance and development of rational-use policies and to provide a baseline for future assessment. METHODS With use of sales data for retail and hospital pharmacies from the IMS Health MIDAS database, we reviewed trends for consumption of standard units of antibiotics between 2000 and 2010 for 71 countries. We used compound annual growth rates to assess temporal differences in consumption for each country and Fourier series and regression methods to assess seasonal differences in consumption in 63 of the countries. FINDINGS Between 2000 and 2010, consumption of antibiotic drugs increased by 36% (from 54 083 964 813 standard units to 73 620 748 816 standard units). Brazil, Russia, India, China, and South Africa accounted for 76% of this increase. In most countries, antibiotic consumption varied significantly with season. There was increased consumption of carbapenems (45%) and polymixins (13%), two last-resort classes of antibiotic drugs. INTERPRETATION The rise of antibiotic consumption and the increase in use of last-resort antibiotic drugs raises serious concerns for public health. Appropriate use of antibiotics in developing countries should be encouraged. However, to prevent a striking rise in resistance in low-income and middle-income countries with large populations and to preserve antibiotic efficacy worldwide, programmes that promote rational use through coordinated efforts by the international community should be a priority. FUNDING US Department of Homeland Security, Bill & Melinda Gates Foundation, US National Institutes of Health, Princeton Grand Challenges Program.

The Global Extent and Determinants of Savanna and Forest as Alternative Biome States

Savanna and forest are alternative states governed by fire at intermediate rainfall levels. Theoretically, fire–tree cover feedbacks can maintain savanna and forest as alternative stable states. However, the global extent of fire-driven discontinuities in tree cover is unknown, especially accounting for seasonality and soils. We use tree cover, climate, fire, and soils data sets to show that tree cover is globally discontinuous. Climate influences tree cover globally but, at intermediate rainfall (1000 to 2500 millimeters) with mild seasonality (less than 7 months), tree cover is bimodal, and only fire differentiates between savanna and forest. These may be alternative states over large areas, including parts of Amazonia and the Congo. Changes in biome distributions, whether at the cost of savanna (due to fragmentation) or forest (due to climate), will be neither smooth nor easily reversible.

Mechanisms of long-distance dispersal of seeds by wind

Long-distance dispersal (LDD) is central to species expansion following climate change, re-colonization of disturbed areas and control of pests. The current paradigm is that the frequency and spatial extent of LDD events are extremely difficult to predict. Here we show that mechanistic models coupling seed release and aerodynamics with turbulent transport processes provide accurate probabilistic descriptions of LDD of seeds by wind. The proposed model reliably predicts the vertical distribution of dispersed seeds of five tree species observed along a 45-m high tower in an eastern US deciduous forest. Simulations show that uplifting above the forest canopy is necessary and sufficient for LDD, hence, they provide the means to define LDD quantitatively rather than arbitrarily. Seed uplifting probability thus sets an upper bound on the probability of long-distance colonization. Uplifted yellow poplar seeds are on average lighter than seeds at the forest floor, but also include the heaviest seeds. Because uplifting probabilities are appreciable (as much as 1–5%), and tree seed crops are commonly massive, some LDD events will establish individuals that can critically affect plant dynamics on large scales.

Comparing classical community models: theoretical consequences for patterns of diversity.

Mechanisms proposed to explain the maintenance of species diversity within ecological communities of sessile organisms include niche differentiation mediated by competitive trade‐offs, frequency dependence resulting from species‐specific pests, recruitment limitation due to local dispersal, and a speciation‐extinction dynamic equilibrium mediated by stochasticity (drift). While each of these processes, and more, have been shown to act in particular communities, much remains to be learned about their relative importance in shaping community‐level patterns. We used a spatially‐explicit, individual‐based model to assess the effects of each of these processes on species richness, relative abundance, and spatial patterns such as the species‐area curve. Our model communities had an order‐of‐magnitude more individuals than any previous such study, and we also developed a finite‐size scaling analysis to infer the large‐scale properties of these systems in order to establish the generality of our conclusions across system sizes. As expected, each mechanism can promote diversity. We found some qualitative differences in community patterns across communities in which different combinations of these mechanisms operate. Species‐area curves follow a power law with short‐range dispersal and a logarithmic law with global dispersal. Relative‐abundance distributions are more even for systems with competitive differences and trade‐offs than for those in which all species are competitively equivalent, and they are most even when frequency dependence (even if weak) is present. Overall, however, communities in which different processes operated showed surprisingly similar patterns, which suggests that the form of community‐level patterns cannot in general be used to distinguish among mechanisms maintaining diversity there. Nevertheless, parameterization of models such as these from field data on the strengths of the different mechanisms could yield insight into their relative roles in diversity maintenance in any given community.