Human geography of New Orleans’ high-lead geochemical setting

Previous soil lead studies in New Orleans focused on the geochemical footprint and its health impacts. This study examines the human geography of race, income, and age in pre-Katrina metropolitan New Orleans within the context of lead accumulation in soils. Sample points of soil lead data (n = 5,467) collected in 1998–2000 were mapped in a geographic information system (GIS), binned into 9 ranges, and queried by (1) 2000 Census racial demographic data, (2) 1999 median household income, and (3) 2000 age data. The absolute population generally declines as lead levels increase except at lead levels from 200–400 to 400–1,000 mg/kg when population increases; the African–American population comprises a disproportionate share of this cohort. The high-lead areas occur in the inner city, home to the largest populations of African-Americans in New Orleans. The mean household income curve indicates that lower economic groups are at risk to higher levels of lead. A total of 44,701 children under the age of 5 years, plus 123,579 children aged 5–17, lived in census block groups containing at least one sample point with over 100 mg/kg lead, and these include 23,124 and 64,064 young people, respectively, who live near at least one point over 400 mg/kg. Lead exposure affects a panoply of outcomes that influence the health and welfare of the community. Unless corrected, children are likely to return to the same or, because of lack of lead-safe practices during renovation, even higher exposure risks than before the flooding of New Orleans.     

Using a sub-grid scale modeling approach to simulate the transport and fate of toxic air pollutants

Several air toxics are emitted from mobile sources on roadways and these emissions account for a significant fraction of the health risks to the population. In addition, health effect studies are now becoming more comprehensive and some account for the spatial heterogeneities of air pollutant concentration fields (as is the case near roadways). Standard models can simulate either the near-source concentration fields or the urban background, but no model can handle both spatial scales in the vicinity of roadways in a coherent and scientifically correct manner. Here, we present a model that provides such an integrated treatment by combining a grid-based air quality model of the urban background with a plume-in-grid representation of roadway emissions. The model is applied to simulate near-roadway concentrations due to emissions from a busy interstate highway in New York City. Qualitative comparisons with typical measured concentration profiles show that the model captures the observed features of toxic air pollutant concentrations near roadways.     

Chemical contamination of soils in the New York City area following Hurricane Sandy

This paper presents a unique data set of lead, arsenic, polychlorinated biphenyl (PCB), and polycyclic aromatic hydrocarbon (PAH) concentrations in soil samples collected from the metropolitan New York City area in the aftermath of Hurricane Sandy. Initial samples were collected by citizen scientists recruited via social media, a relatively unusual approach for a sample collection project. Participants in the affected areas collected 63 usable samples from basements, gardens, roads, and beaches. Results indicate high levels of arsenic, lead, PCBs, and PAHs in an area approximately 800 feet south of the United States Environmental Protection Agency (US EPA) Superfund site at Newtown Creek. A location adjacent to the Gowanus Canal, another Superfund site, was found to have high PCB concentrations. Areas of high PAH contamination tended to be near high traffic areas or next to sites of known contamination. While contamination as a direct result of Hurricane Sandy cannot be demonstrated conclusively, the presence of high levels of contamination close to known contamination sites, evidence for co-contamination, and decrease in number of samples containing measureable amounts of semi-volatile compounds from samples collected at similar locations 9 months after the storm suggest that contaminated particles may have migrated to residential areas as a result of flooding.     

Multiscale Plume Transport from the Collapse of the World Trade Center on September 11, 2001

The collapse of the world trade center (WTC) produced enhanced levels of airborne contaminants in New York City and nearby areas on September 11, 2001 through December, 2001. This catastrophic event revealed the vulnerability of the urban environment, and the inability of many existing air monitoring systems to operate efficiently in a crisis. The contaminants released circulated within the street canyons, but were also lifted above the urban canopy and transported over large distances, reflecting the fact that pollutant transport affects multiple scales, from single buildings through city blocks to mesoscales. In this study, ground-and space-based observations were combined with numerical weather forecast fields to initialize fine-scale numerical simulations. The effort is aimed at reconstructing pollutant dispersion from the WTC in New York City to surrounding areas, to provide means for eventually evaluating its effect on population and environment. Atmospheric dynamics were calculated with the multi-grid Regional Atmospheric Modeling System (RAMS), covering scales from 250 m to 300 km and contaminant transport was studied using the Hybrid Particle and Concentration Transport (HYPACT) model that accepts RAMS meteorological output. The RAMS/HYPACT results were tested against PM2.5 observations from the roofs of public schools in New York City (NYC), Landsat images, and Multi-angle Imaging SpectroRadiometer (MISR) retrievals. Calculations accurately reproduced locations and timing of PM2.5 peak aerosol concentrations, as well as plume directionality. By comparing calculated and observed concentrations, the effective magnitude of the aerosol source was estimated. The simulated pollutant distributions are being used to characterize levels of human exposure and associated environmental health impacts.     

Sociogenetic responses to ecological variation in the ant Myrmica punctiventris are context dependent

Models of social evolution predict a strong relationship between ecological factors and sociogenetic organization in social insects (e.g. queen number, nestmate relatedness and population structure). Despite a large body of coherent theory, empirical support for these predictions is weak. Here we report the results of an experiment that manipulated two ecological parameters, food and nestsite availability, thought to be important for a population of the forest ant Myrmica punctiventris. Earlier work had shown that the sociogenetic structure varies between two populations of this species, and an ecological experiment in one of the populations (in Vermont) revealed that food supplementation had the strongest effects on nestmate relatedness. We repeated the experiment in the second population (in New York) and obtained strikingly different results. We show that nestsite supplementation had the strongest effect in the New York population, and that adding both food and nestsites affected nestmate relatedness in a direction opposite to that reported from the Vermont study. These results show that the ecological context is critically important for understanding the determinants of colony structure in ants. Furthermore, comparison of our data with that of a previous study shows that social organization in New York is temporally unstable. Thus, not only do ecological factors strongly influence social organization, but their influence can vary over time. Our study underscores the need for detailed information on the natural history and ecology of social species.     

Effects of crowding and wave exposure on penis morphology of the acorn barnacle, Semibalanus balanoides

Wave action and low population density can strongly reduce the ability of sessile acorn barnacles to find mates and copulate. For Semibalanus balanoides, penis morphology varies with wave exposure and with characteristics of the mating neighborhood. Field experiments were conducted at five intertidal sites on Long Island, New York, USA from July to December 2005 to determine how wave exposure and aggregation structure influence the length, diameter, mass, and number of annulated folds of the penis. Sparsely crowded barnacles had more annulations in the penis and are inferred to have greater ability to stretch. At higher wave exposure, the diameter of the penis was greater, but the mass was not. This study identifies density of crowding as the most important cue that barnacles respond to when perceiving their mating group and details how penis morphology varies in response to wave exposure.     

Temperature ecotypes near the southern boundary of the kelp Laminaria saccharina

Effects of temperature on survival, growth, and photosynthesis were compared for two USA populations of Laminaria saccharina Lamour. One population was located in New York State, near the southern latitudinal boundary of the species in the western North Atlantic. This southern boundary population was exposed to ambient temperatures 20°C for about 6 wk each summer. The second population was located in Maine, toward the center of the latitudinal range of the species, and was rarely exposed to temperatures>17°C. sporophytes from the New York (NY) population exhibited greater tolerance of high temperature than plants from the Maine (ME) site. Juvenile sporophytes from the two sites had similar rates of survivorship and growth at temperatures below 20°C, but showed different responses at 20°C in laboratory experiments. NY plants survived and grew for 6 wk at 20°C. ME plants showed negative growth during wk 2 and 100% mortality during wk 3. NY and ME plants held in situ at the NY site during June to September, 1985, also exhibited differential survivorship when ambient temperatures exceeded 20°C. Results of photosynthesis and dark respiration measurements on NY and ME plants grown at various temperatures suggested that the high-temperature tolerance of NY plants was attributable to their ability to maintain positive daily net C-fixation at 20°C. The high-temperature tolerance of the NY plants appeared to be due to genetic adaptation and is probably crucial to the persistence of the species near its southern boundary.     

Urban afforestation and infant health: Evidence from MillionTreesNYC

This paper examines the impact of urban afforestation on infant health outcomes by exploiting a quasi-experimental setting where one million new trees were planted in New York City (NYC), but not in counties surrounding NYC over the same time period. Using a near-universal birth record of NYC and surrounding counties over 2004–2015 and employing both the synthetic control method and a difference-in-differences model, we find that an approximately 20% increase in urban forest cover decreased prematurity and low birth weight among mothers in NYC by 2.1 and 0.24 percentage points, respectively, relative to similar mothers outside of NYC. The low birth weight finding is equivalent to getting a mother smoking two cigarettes a day during pregnancy to quit. An internal validity test suggests that changes in the composition of NYC mothers cannot explain the observed effects. Additionally, we find evidence that declines in PM_2.5 concentrations and increases in outdoor walks are potential causal mechanisms. Results suggest that urban afforestation may be able to complement existing policies aimed at improving infant health.     

Assessing citizen contributions to butterfly monitoring in two large cities

Citizen science may be especially effective in urban landscapes due to the large pool of potential volunteers. However, there have been few evaluations of the contributions of citizen scientists to knowledge of biological communities in and around cities. To assess the effectiveness of citizen scientists' monitoring of species in urban areas, we compared butterfly data collected over 10 years in Chicago, Illinois (U.S.A.), and New York City, New York (U.S.A.). The dates, locations, and methods of data collection in Chicago were standardized, whereas data from New York were collected at any location at any time. For each city, we evaluated whether the number of observers, observation days (days on which observations were reported), and sampling locations were associated with the reported proportion of the estimated regional pool of butterfly species. We also compared the number of volunteers, duration of volunteer involvement, and consistency of sampling efforts at individual locations within each city over time. From 2001 to 2010, there were 73 volunteers in Chicago and 89 in New York. During this period, volunteers observed 86% and 89% of the estimated number of butterfly species present in Chicago and New York, respectively. Volunteers in New York reported a greater proportion of the estimated pool of butterfly species per year. In addition, more species were observed per volunteer and observation day in New York, largely due to the unrestricted sampling season in New York. Chicago volunteers were active for more years and monitored individual locations more consistently over time than volunteers in New York. Differences in monitoring protocol--especially length of sampling season and selection protocol for monitoring locations--influenced the relationship between species accrual and sampling effort, which suggests these factors are important in volunteer-based species-monitoring programs.     

Nest site limitation and facultative polygyny in the ant Leptothorax longispinosus

Summary The ant L. longispinosus displays geographic variation in its pattern of facultative polygyny (Fig. 2). In nature, nest density and frequency of multiple queening are positively associated over three sites. A putative causal relation between availability of vacant nest sites and polygyny was examined in New York, where a plot was seeded with additional nest sites and monitored for 24 months. Both queen number and worker number per nest on the experimental plot were reduced relative to controls (Fig. 4, Fig. 5), indicating that scarcity of available nest sites influences the pattern of polygyny in this species. The observed demographic changes resulted from fractionation of existing colonies; adding nest sites induced polydomy. Although numbers of adult ants changed with addition of nest sites, the numbers of immatures were no different after 2 years (Table 1), suggesting that the population was undergoing growth to expand into the additional sites. These results are the first direct experimental evidence linking polygyny to an ecological parameter for any ant species.     

Trihalomethanes formation in water treatment plants and distribution lines: a monitoring and modeling scheme

Monitoring and modeling of the concentrations of trihalomethanes (THMs) within four water treatment plants (WTPs) and distribution lines in Fayoum City, Egypt, were studied. Sampling sites for raw and treated waters were determined by global positioning system and the Arc geographic information system software was used for mapping. THMs were monitored using 25 sampling points located at the plant exit and through the distribution lines up to the plant extremity. Results indicated that the THMs concentration varied significantly but it was very rarely higher than the allowed maximum contaminant level of 100 μg L^?1. However, at the dead zones in the distribution lines, the THMs increased by about 160% compared to in-plant level. Furthermore, the level of THMs in hot months was about 1.3 times higher than in cold ones. The influence of some independent variables such as temperature, pH, ammonia concentration, total organic carbon and color were investigated by Pearson model to find their correlation with the rate of THMs formation in WTPs. The resulted R values ranged from 0.81 to 0.98. Successful application of the model to a selected region on the distribution lines has resulted in a correlation coefficient of 0.98.     

Simulated potential effects of ecological factors on a hypothetical population of Chiricahua leopard frog (Rana chiricahuensis)

Simulations provide an opportunity to examine how single or multiple perturbations may impact a specific species. The objectives of this study were to identify thresholds at which changes in stream peak flow, stream base flow, and/or chytrid fungus presence alter long-term Rana chiricahuensis populations. We used scenarios with varying peak flow mortality rates, base flow mortality rates, and chytrid fungus mortality rates. Sensitivity analysis was also conducted. Over 50 years, populations in six scenarios increased and 13 scenarios decreased. Eight scenarios resulting with fewer than 100 individuals included stochastic effects for at least two of three perturbations and the remaining scenarios included chronic effects of 30% or higher. Scenarios with population increases had either no chytrid fungus effect or chronic effects from perturbations totaling less than 30%. In the absence of chytrid fungus, populations increased and became stable. At a 10% annual death rate caused by chytrid fungus, the R. chiricahuensis population decreased 46.8%. At a 20% death rate, the population decreased 98.6%. Model scenarios were sensitive to peak flow death rates. As peak flow mortality increased to 10 and 20%, extinction rates increased to 91.7 and 99.9%, respectively. With model parameters and the no base flow mortality, R. chiricahuensis populations declined by 92% with a 3.2% extinction rate at 50 years. Models with base flow mortality rates of 10 and 20% resulted in population extinction rates of 48.7 and 96.1%, respectively. Scenario analysis of perturbations on a hypothetical R. chiricahuensis population provided a framework in which to view combined effects on a species. Analysis supports supposition that chytrid fungus is the proximate cause of many amphibian declines, but the added effect of base flow and peak flow has the potential to hasten declines.     

Mitochondrial DNA Variability of the Gray Wolf: Genetic Consequences of Population Decline and Habitat Fragmentation

The gray wolf is a large, highly mobile predator whose original geographic range included most of the Northern Hemisphere. High rates of genetic exchange probably characterized even distantly-separated populations in the past, but recent population declines and habitat fragmentation have isolated previously contiguous populations, especially in the Old World. We examine mitochondrial DNA (mtDNA) variability among twenty-six populations of wolves from throughout their geographic range. We find eighteen mtDNA genotypes in gray wolves, seven of these are derived from hybridization with coyotes, four are confined to the New World, six are confined to the Old World and one is shared by both areas. Genetic differentiation among wolf populations is significant but small in magnitude. In the Old World, most localities have a single unique genotype, whereas in the New World several genotypes occur at most localities and three of the five genotypes are nearly ubiquitous. The pattern of genetic differentiation in the gray wolf contrasts with that of another large, highly vagile canid, the coyote, in which genetic differentiation among populations is not significant even among widely separated localities. We suggest that the difference between these two species reflects the rapid, recent increase in coyote numbers and expansion of their geographic range, and the coincident decline in gray wolf populations. Apparent genetic differences among extant wolf populations may be a recent phenomenon reflecting population declines and habitat fragmentation rather than a long history of genetic isolation.     

Height-growth response to climatic changes differs among populations of Douglas-fir: a novel analysis of historic data

Projected climate change will affect existing forests, as substantial changes are predicted to occur during their life spans. Species that have ample intraspecific genetic differentiation, such as Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), are expected to display population-specific growth responses to climate change. Using a mixed-effects modeling approach, we describe three-year height (HT) growth response to changes in climate of interior Douglas-fir populations. We incorporate climate information at the population level, yielding a model that is specific to both species and population. We use data from provenance tests from previous studies that comprised 236 populations from Idaho, Montana, and eastern Washington, USA. The most sensitive indicator of climate was the mean temperature of the coldest month. Population maximum HT and HT growth response to changes in climate were dependent on seed source climate. All populations had optimum HT growth when transferred to climates with warmer winters; those originating in sites with the warmest winters were taller across sites and had highest HT growth at transfer distances closest to zero; those from colder climates were shortest and had optimum HT growth when transferred the farthest. Although this differential response damped the height growth differences among populations, cold-climate populations still achieved their maximum growth at lower temperatures than warm-climate populations. The results highlight the relevance of understanding climate change impacts at the population level, particularly in a species with ample genetic variation among populations.     

Age structure,growth, and morphometric variations in the Atlantic surf clam,Spisula solidissima,from estuarine and inshore waters

The surf clam, Spisula solidissima (Dillwyn), population in the estuarine waters of Long Island Sound, New York, USA, was characterized in 1984 and again in 1988 by an age structure restricted to just two age-classes, and an apparent lifespan of only about 10 yr. In the inshore coastal waters off Fire Island, New York, a wide age range from 2 to 22 yr old was present. The age structure at Long Beach, New York, a third geographic region which is coastal but influenced by the Hudson River estuary, was similar to Long Island Sound. Juvenile surf clams grew at similar rates in all three geographic regions. However, adults from the Long Island Sound population grew significantly slower and reached an asymptotic maximum size which was 37% smaller than Fire Island adults. Long Beach adults had intermediate growth rates and maximum sizes. The shells of Long Island Sound clams were also 25% thinner than those from the other two regions. Density dependent effects on growth, evaluated over abundances ranging from 0.5 to 294 ind. m^–2, were present but were too small to account for observed regional differences. Results suggest that adult surf clams may be physiologically stressed by the reduced salinity and more extreme temperatures found in estuarine waters.     

Recolonizing wolves and mesopredator suppression of coyotes: impacts on pronghorn population dynamics.

Food web theory predicts that the loss of large carnivores may contribute to elevated predation rates and, hence, declining prey populations, through the process of mesopredator release. However, opportunities to test predictions of the mesopredator release hypothesis are rare, and the extent to which changes in predation rates influence prey population dynamics may not be clear due to a lack of demographic information on the prey population of interest. We utilized spatial and seasonal heterogeneity in wolf distribution and abundance to evaluate whether mesopredator release of coyotes (Canis latrans), resulting from the extirpation of wolves (Canis lupus) throughout much of the United States, contributes to high rates of neonatal mortality in ungulates. To test this hypothesis, we contrasted causes of mortality and survival rates of pronghorn (Antilocapra americana) neonates captured at wolf-free and wolf-abundant sites in western Wyoming, USA, between 2002 and 2004. We then used these data to parameterize stochastic population models to heuristically assess the impact of wolves on pronghorn population dynamics due to changes in neonatal survival. Coyote predation was the primary cause of mortality at all sites, but mortality due to coyotes was 34% lower in areas utilized by wolves (P < 0.001). Based on simulation modeling, the realized population growth rate was 0.92 based on fawn survival in the absence of wolves, and 1.06 at sites utilized by wolves. Thus, wolf restoration is predicted to shift the trajectory of the pronghorn population from a declining to an increasing trend. Our results suggest that reintroductions of large carnivores may influence biodiversity through effects on prey populations mediated by mesopredator suppression. In addition, our approach, which combines empirical data on the population of interest with information from other data sources, demonstrates the utility of using simulation modeling to more fully evaluate ecological theories by moving beyond estimating changes in vital rates to analyses of population-level impacts.     

The importance of space, time, and stochasticity to the demography and management of Alliaria petiolata

As population modeling is increasingly called upon to guide policy and management, it is important that we understand not only the central tendencies of our study systems, but the consequences of their variation in space and time as well. The invasive plant Alliaria petiolata (garlic mustard) is actively managed in the United States and is the focus of a developing biological control program. Two weevils (Coleoptera: Curculionidae: Ceutorhynchus) that reduce fecundity (C. alliariae) and rosette survival plus fecundity (C. scrobicollis) are under consideration for release pending host specificity testing. We used a demographic modeling approach to (1) quantify variability in A. petiolata growth and vital rates and (2) assess the potential for single- or multiple-agent biocontrol to suppress growth of 12 A. petiolata populations in Illinois and Michigan studied over three plant generations. We used perturbation analyses and simulation models with stochastic environments to estimate stochastic growth rates (lambda(S)) and predict the probability of successful management using either a single biocontrol agent or two agent species together. Not all populations exhibited invasive dynamics. Estimates of lambda(S) ranged from 0.78 to 2.21 across sites, while annual, deterministic growth (lambda) varied up to sevenfold within individual sites. Given our knowledge of the biocontrol agents, this analysis suggests that C. scrobicollis alone may control A. petiolata at up to 63% of our study sites where lambda >1, with the combination of both agents predicted to succeed at 88% of sites. Across sites and years, the elasticity rankings were dependent on lambda. Reductions of rosette survival, fecundity, or germination of new seeds are predicted to cause the greatest reduction of lambda in growing populations. In declining populations, transitions affecting seed bank survival have the greatest effect on lambda. This contrasts with past analyses that varied parameters individually in an otherwise constant matrix, which may yield unrealistic predictions by decoupling natural parameter covariances. Overall, comparisons of stochastic and deterministic growth rates illustrate how analyses of individual populations or years could misguide management or fail to characterize complex traits such as invasiveness that emerge as attributes of populations rather than species.     

Spatial selection and inheritance: applying evolutionary concepts to population dynamics in heterogeneous space

Organisms in highly suitable sites generally produce more offspring, and offspring can inherit this suitability by not dispersing far. This combination of spatial selection and spatial inheritance acts to bias the distribution of organisms toward suitable sites and thereby increase mean fitness (i.e., per capita population increase). Thus, population growth rates in heterogeneous space change over time by a process conceptually analogous to evolution by natural selection, opening avenues for theoretical cross-pollination between evolutionary biology and ecology. We operationally define spatial inheritance and spatial selective differential and then combine these two factors in a modification of the breeder's equation, derived from simple models of population growth in heterogeneous space. The modified breeder's equation yields a conservative criterion for persistence in hostile environments estimable from field measurements. We apply this framework for understanding gypsy moth population persistence amidst abundant predators and find that the predictions of the modified breeder's equation match initial changes in population growth rate in independent simulation output. The analogy between spatial dynamics and natural selection conceptually links ecology and evolution, provides a spatially implicit framework for modeling spatial population dynamics, and represents an important null model for studying habitat selection.     

Population structure and life history of orange roughy (Hoplostethus atlanticus) in the SW Pacific: inferences from otolith chemistry

We examined site differences in the elemental composition of the primordium and ontogenetic variability of Sr in otoliths of fish from Australia and New Zealand and, as an out-group, the North Atlantic. Differences among sites in primordium composition are slight, but significant for all five elements assayed (Sr, Pb, Cu, Zn and Hg), but principally reflect differences between the North Atlantic and SW Pacific specimens, do not replicate for independent samples in the SW Pacific and constitute a poor “natural tag” in roughy, with <25% of fish successfully assigned to source location. However, mean Sr weight-fractions at the primordium showed similar latitudinal variation across sites in Australia, New Zealand and the Tasman Sea, indicating both spatially structured populations and a common structuring process across the region. Comparisons of ontogenetic variability of Sr in otoliths from juveniles and young adults within and between sites in the SW Pacific strongly support the hypothesis that variability in this element is site-specific and environmentally sensitive, although the environmental factors involved are not obvious. The otolith analysis confirms previous suggestions that juvenile and adult Hoplostethus atlanticus are relatively sedentary, but also indicates that the population sub-structuring by age within sites is more complex and there are likely to be more spawning areas in Australian waters than previously thought. More broadly, although single point analysis of otolith composition at the primordium resolves a population structure in roughy, alone it is not precise enough to test hypotheses about the processes causing this structure. Ontogenetic variability in Sr provides better resolution of spatial structure, even in a relatively homogenous marine environment like the deep ocean, and also provides insight into behavioural and ecological factors. Ontogenetic analyses of Sr in otoliths are expensive to obtain, require more effort in specimen preparation than single point analyses, and are difficult to compare statistically, but the increased information they yield warrants their broader consideration in marine species.     

Behavioral transitions with the evolution of cooperative nest founding by harvester ant queens

Research on the evolution of cooperative groups tends to explore the costs and benefits of cooperation, with less focus on the proximate behavioral changes necessary for the transition from solitary to cooperative living. However, understanding what proximate changes must occur, as well as those pre-conditions already in place, is critical to understanding the origins and evolution of sociality. The California harvester ant Pogonomyrmex californicus demonstrates population-level variation in colony founding over a close geographic range. In adjacent populations, queens either found nests as single individuals (haplometrosis) or form cooperative groups of nonrelatives (pleometrosis). We compared aggregation, aggression, and tolerance of queens from one pleometrotic and two haplometrotic populations during nest initiation, to determine which behaviors show an evolutionary shift and which are present at the transition to pleometrosis. Surprisingly, within-nest aggregative behavior was equally present among all populations. In nesting boxes with multiple available brood-rearing sites, both queen types readily formed and clustered around a single common brood pile, suggesting that innate attraction to brood (offspring) facilitates the transition to social aggregation. In contrast, queens from the three populations differed in their probabilities of attraction on the ground to nest sites occupied by other queens and in levels of aggression. Our results suggest that some key behavioral mechanisms facilitating cooperation in P. californicus are in place prior to the evolution of pleometrosis and that the switch from aggression to tolerance is critical for the evolution of stable cooperative associations.