Gender, Environment and Population

The relative scarcity of studies at the intersection of behavioral and population ecology is surprising given the presumed importance of behavior in density-dependent population regulation. Here we tested whether North American red squirrels (Tamiasciurus hudsonicus) adjust their behavior in response to local population density and whether they use rates of territorial vocalizations in their local neighborhood to assess density. We examined these relationships using 18 years of live trapping and 20 years of behavioral data that were collected across natural variation in local population density. To disentangle the effects of population density on behavior from those due to changes in per capita food abundance or changes in the frequency of antagonistic interactions with neighbors, we also experimentally manipulated population density with long-term food supplementation as well as perceived population density with long-term playbacks of territorial vocalizations. The frequency with which squirrels emitted territorial vocalizations was positively associated with local population density. In contrast, antagonistic physical interactions observed between squirrels and territorial intrusions were rare and the frequency of intrusions was weakly and negatively, not positively, associated with population density. Squirrels experiencing naturally and experimentally high density conditions spent less time in the nest and feeding but more time being vigilant. Similar density-dependent changes in behavior were observed in response to our manipulations of perceived population density, indicating that vocalization rates and not physical interactions or food abundance were the mechanism by which squirrels assessed and responded behaviorally to changes in local density.

This paper first reviews the descriptions and structural explanations for variations in urban population densities in the United States over several decades. The research reviewed focuses on shifts in the technology and economics of travel as being the major influences on urban density distributions. These variables appear to be somewhat less adequate as explanations of the process in recent decades. After reviewing these findings and suggesting several research questions, we attempt to assess the evidence relating variations in population density to several other aspects of human behavior. A review of the research relating high population densities to various indicates that little of the variance in rates of these pathologies can be attributed to density independently of other social structural variables. Some important consequences of population density have been reported, however, and possible research directions are outlined.

In 1994, a “Pan-European Programme for Intensive and Continuous Monitoring of Forest Ecosystems” started to contribute to a better understanding of the impact of air pollution, climate change and natural stress factors on forest ecosystems. The programme today counts approximately 760 permanent observation plots including near 500 plots with data on both air quality and forest ecosystem impacts. This paper first presents impacts of air pollution and climate on forests ecosystems as reported in the literature on the basis of laboratory and field research. Next, results from monitoring studies, both at a European wide scale and related national studies, are presented in terms of trends and geographic variations in nitrogen and sulphur deposition and ozone concentrations and the impacts of those changes in interaction with weather conditions on (i) water and element budgets and nutrient-acidity status, (ii) forest crown condition, (iii) forest growth and carbon sequestration and (iv) species diversity of the ground vegetation. The empirical, field based forest responses to the various drivers are evaluated in view of available knowledge. Analyses of large scale monitoring data sets show significant effects of atmospheric deposition on nutrient-acidity status in terms of elevated nitrogen and sulphur or sulphate concentrations in forest foliage and soil solution and related soil acidification in terms of elevated aluminium and/or base cation leaching from the forest ecosystem. Relationships of air pollution with crown condition, however, appear to be weak and limited in time and space, while climatic factors appear to be more important drivers. Regarding forest growth, monitoring results indicate a clear fertilization effect of N deposition on European forests but the field evidence for impacts of ambient ozone exposure on tree growth is less clear.

The existence of patterns in population dynamics across species geographic ranges and climatic niches is a pervasive idea in ecology. Population variability (i.e. temporal variability in population density) should hypothetically increase near range edges or niche limits because of less suitable environments in these areas, but the occurrence of such patterns remains largely unexplored. Further, fluctuations in temperature could pose demographic constraints on populations and also influence their variability. We used Breeding Bird Survey data to show that the population variability of 97 resident North American birds consistently increases towards their niche limits and in areas with more variable temperatures, but not towards their geographic range edges. However, our model has limited explanatory power, and phylogenetic history and species traits could not explain these results. These findings suggest that other factors, such as biotic interactions and resource availability, might be more important drivers of population variability in resident North American birds.

A survey on the geographical distribution of Chnootriba similis (Thunberg), formerly known as Epilachna similis, was undertaken from 2001 to 2003 in the main cereal-growing areas of Ethiopia. The population density of the insect was estimated by sweep netting (100 sweeps/farm). In addition, more detailed studies on seasonal and annual fluctuations in population density of C. similis were carried out in southern Ethiopia between 2002 and 2003 by sampling 10 quadrants of 0.25m2 per farmer’s field. The damage levels were determined using scores of 0–9. The survey revealed that C. similis is widely distributed in different parts of the cereal-growing regions of Ethiopia. It was observed that the beetles had spread to altitudes ranging from 1217 to around 3000m asl. However, the variations in population densities of C. similis cannot be directly attributed to altitude, as highly differing population densities in similar altitudes of different locations were recorded. Annual and seasonal variations in population density were observed in different areas of the southern part of Ethiopia. These results suggest that C. similis has a sporadic nature and can occasionally build up to outbreak levels. Beetles fed on both cereals (barley, wheat, tef, maize, sorghum and finger millet) and wild grasses (Leersia hexandra (Sw.), Avena abyssinica (Hoechst) and Lolium temulentum (L.)). Population densities and damage levels on the cereal crops varied considerably with the highest densities and damage levels on barley.

Temporal variability in population density has been shown to increase with increasing length of a study. This has serious implications for ecological studies and suggests that obtaining an objective measure of temporal variability may not be possible. To test this hypothesis more generally, I calculated inter-annual variability, using the standard deviation of the logarithm of annual mean population density (STD(log D)), for 70 populations of phytoplankton, zooplankton and fish which have been sampled over 10 to 51 consecutive yr in lakes around the world. STD(log D) fluctuates most widely during the first 3-6 yr of a study, but these changes do not necessarily represent long-term trends in temporal variability. Median changes in STD(log D) over time are small («0.1 unit, even over a logarithmic time scale) and decrease as more years are included in the analysis. The inter-annual variability of most freshwater populations appears to reach an asymptotic value. Several populations, however, show an alternation of plateaux and of sudden increases in STD(log D), suggesting that the asymptotic values found in most populations may be temporary and may not extrapolate beyond the temporal scale of this study (10-50 yr). Overall, the increases in inter-annual variability are expected to be small (< 0.1 units, on average, per 25 yr). The pattern and the magnitude of these changes did not differ significantly between algae, zooplankton and fish, and were similar to those reported for birds and mammals. Increases in temporal variability may only be important for questions addressed over long periods of time.

1. In many groups of species, range size and population density are positively correlated (the ‘distribution−abundance relationship’) and range size is positively correlated with latitude (‘Rapoport's Rule’). This study investigated these two relationships among Australian forest mammals, and also tested for the existence of a relationship between population density and latitude in the same fauna.2.All three relationships were significant, with the distribution–abundance relationship the strongest pattern of the three, and Rapoport's Rule the weakest. Partial correlation analysis showed that: (i) range size and population density remained positively correlated when latitude was held constant; and (ii) population density and latitude remained positively correlated when range size was held constant; but (iii) latitude and range size were not significantly correlated when population density was held constant. For (i) and (ii), partial correlation did not reveal stronger relationships than were revealed by simple correlation. These results suggest that the distribution–abundance relationship is conditioned by latitude to produce small ranges and low population densities in the tropics, and that tropical species tend to have low population densities relative to range size.3.Body mass accounted for little variation in population density, and for no variation in range size and latitudinal position. The small ranges and low population densities of tropical species were not related to a latitudinal gradient in species richness, which did not occur in this fauna.4.The latitudinal gradient in population density held within widespread species as well as among species. The effect of Bergmann's Rule (the tendency for organisms in cool environments to be larger bodied than organisms in warm environments) meant that in some species, population energy use rose more steeply with latitude than did population density.5.I argue that all three relationships might be due to a tendency for the foraging efficiency of animals to be lower in tropical than in temperate ecosystems, as a result of a lower concentration of nutrients in tropical ecosystems. For resource‐limited species, foraging efficiency should influence both range size and population density. Differences in foraging efficiency may therefore produce the distribution−abundance relationship as well as latitudinal gradients in range size and population density.

Large variations in population densities of Sericothrips staphylinus were observed between gorse bushes at a field site in an earlier Tasmanian study. A replicated potted plant experiment was conducted to assess whether genetic variation between gorse bushes could affect densities of S. staphylinus. There were significant differences in the number of S. staphylinus recovered from the individual bushes sampled within sites and individual bushes across all sites. This provides evidence that genetic variation between gorse bushes is at least partly responsible for the observed variability in field populations. The result validates the current release strategy used for the field establishment of S. staphylinus in south-eastern Australia. This strategy involves making multiple small releases (ca. 250 adults per bush per site) instead of a small number of large releases (ca. 1000 per bush per site), thereby reducing variation in numbers due to genetic affects. Genetic variation between individual plants of the same species and associated effects on herbivore densities should be considered when devising release strategies for biological control agents.

ABSTRACTThe environmental context of the nitrogen-fixing mutualism between leguminous plants and rhizobial bacteria varies over space and time. Variation in resource availability, population density, and composition likely affect the ecology and evolution of rhizobia and their symbiotic interactions with hosts. We examined how host genotype, nitrogen addition, rhizobial density, and community complexity affected selection on 68 rhizobial strains in the Sinorhizobium meliloti–Medicago truncatula mutualism. As expected, host genotype had a substantial effect on the size, number, and strain composition of root nodules (the symbiotic organ). The understudied environmental variable of rhizobial density had a stronger effect on nodule strain frequency than the addition of low nitrogen levels. Higher inoculum density resulted in a nodule community that was less diverse and more beneficial but only in the context of the more selective host genotype. Higher density resulted in more diverse and less beneficial nodule communities with the less selective host. Density effects on strain composition deserve additional scrutiny as they can create feedback between ecological and evolutionary processes. Finally, we found that relative strain rankings were stable across increasing community complexity (2, 3, 8, or 68 strains). This unexpected result suggests that higher-order interactions between strains are rare in the context of nodule formation and development. Our work highlights the importance of examining mechanisms of density-dependent strain fitness and developing theoretical predictions that incorporate density dependence. Furthermore, our results have translational relevance for overcoming establishment barriers in bioinoculants and motivating breeding programs that maintain beneficial plant-microbe interactions across diverse agroecological contexts.IMPORTANCE Legume crops establish beneficial associations with rhizobial bacteria that perform biological nitrogen fixation, providing nitrogen to plants without the economic and greenhouse gas emission costs of chemical nitrogen inputs. Here, we examine the influence of three environmental factors that vary in agricultural fields on strain relative fitness in nodules. In addition to manipulating nitrogen, we also use two biotic variables that have rarely been examined: the rhizobial community's density and complexity. Taken together, our results suggest that (i) breeding legume varieties that select beneficial strains despite environmental variation is possible, (ii) changes in rhizobial population densities that occur routinely in agricultural fields could drive evolutionary changes in rhizobial populations, and (iii) the lack of higher-order interactions between strains will allow the high-throughput assessments of rhizobia winners and losers during plant interactions.

Urban Density and Covid-19

How demographic factors lead to variation or change in growth rates can be investigated using life table response experiments (LTRE) based on structured population models. Traditionally, LTREs focused on decomposing the asymptotic growth rate, but more recently decompositions of annual 'realized' growth rates using 'transient' LTREs have gained in popularity. Transient LTREs have been used particularly to understand how variation in vital rates translate into variation in growth for populations under long-term study. For these, complete population models may be constructed to investigate how temporal variation in environmental drivers affect vital rates. Such investigations have usually come down to estimating covariate coefficients for the effects of environmental variables on vital rates, but formal ways of assessing how they lead to variation in growth rates have been lacking. We extend transient LTREs to further partition the contributions from vital rates into contributions from temporally varying factors that affect them. The decomposition allows one to compare the resultant effect on the growth rate of different environmental factors, as well as density dependence, which may each act via multiple vital rates. We also show how realized growth rates can be decomposed into separate components from environmental and demographic stochasticity. The latter is typically omitted in LTRE analyses. We illustrate these extensions with an integrated population model (IPM) for data from a 26 years study on northern wheatears (Oenanthe oenanthe), a migratory passerine bird breeding in an agricultural landscape. For this population, consisting of around 50-120 breeding pairs per year, we partition variation in realized growth rates into environmental contributions from temperature, rainfall, population density and unexplained random variation via multiple vital rates, and from demographic stochasticity. The case study suggests that variation in first year survival via the unexplained random component, and adult survival via temperature are two main factors behind environmental variation in growth rates. More than half of the variation in growth rates is suggested to come from demographic stochasticity, demonstrating the importance of this factor for populations of moderate size.

Echinoderms play a key role in structuring many marine ecosystems and are notorious for large population density variations in so‐called “outbreak” or “dieoff” events. In a review of this phenomenon, we assess the causal factors and ecological and evolutionary consequences. We identified 28 species (6 Asteroidea, 8 Echinoidea, 10 Holothuroidea, 4 Ophiuroidea) that exhibit large (more than two population doublings or halvings) population density changes. Three generalized patterns were identified and named for exemplary species: (1) rapid decreases followed by no or slow recovery (Diadema–Paracentrotus Model), (2), rapid increase and apparent stability at a new population density (Amperima–Amphiura Model), and (3) population density fluctuations (Acanthaster–Asterias Model). Echinoderms identified were distributed from the shallow intertidal to the deep sea, and from tropical to temperate regions. In most cases, significant impacts on the respective ecosystems were observed. The most striking similarity among all species identified was possession of the ancestral‐type planktotrophic larva. This larval type was significantly overrepresented in species identified within the Asteroidea, Echinoidea, Holothuroidea, and for the combined data set. We suggest three main factors that render a life history with planktotrophic larvae a high‐risk–high‐gain strategy: (1) a strong nonlinear dependency of larval production on adult densities (Allee effects), (2) a low potential for compensatory feedback mechanisms, and (3) an uncoupling of larval and adult ecology. The alternative (derived) lecithotrophic larva occurs in 68% of recent echinoderm species, suggesting an evolutionary trend toward this larval type. Lecithotrophic development represents a more buffered life history because compensatory feedback between adult densities and larval output is likely to be more efficient. For lecithotrophic developers, direct nutritive coupling from adult to larva to the early benthic juvenile provides a buffer against starvation. Lecithotrophic larvae are independent of the vagaries of planktonic food supply, and their short planktonic duration may promote local recruitment. Anthropogenic influences contributed to the population density variations in most cases, including increased primary productivity through eutrophication or global change, disease, overfishing, and species introductions. We suggest that anthropogenic disturbance, through its influence on the frequency and/or amplitude of echinoderm population density changes, may go beyond present ecosystem impacts and alter future evolutionary trends.

Urbanization processes modulate the immunological challenges faced by animals. Urban habitat transformations reshape pathogen diversity and abundance, while high population density-common in urban exploiter species-promotes disease transmission. Responses to urbanization may include adaptive adjustments of constitutive innate immune defenses (e.g. complement system and natural antibodies [NAbs]), which serve as first-line protection against infections. Here, we investigated associations of habitat urbanization and host population density with complement and NAbs in an urban bird, the feral pigeon Columba livia domestica. To do so, we employed the hemolysis-hemagglutination assay to analyze nearly 200 plasma samples collected across urbanization and pigeon population density gradients in five major cities in Poland. We found a negative association between urbanization score and hemagglutination (i.e. NAbs activity), but not hemolysis (i.e. complement activity), indicating either immunosuppression or adaptive downregulation of this immune defense in highly transformed urban landscape. Population density was not significantly related to either immune parameter, providing no evidence for density-dependent modulation of immune defenses. At the same time, there was a negative association of hemolysis with condition (scaled mass index), suggesting resource allocation trade-offs or contrasting effects of the urban environment on immune defenses and body condition. The results demonstrate that habitat structure can be an important factor shaping the immune defenses of the feral pigeon, although these associations were not mediated by variation in population density. Our study highlights the complexity of the links between immune defenses in wildlife and urbanization and reinforces the need for comprehensive ecoimmunological studies on urban animals.

Population densities, size and condition factor of juvenile brown trout and variation in environmental factors (discharge, temperature, and food availability) were examined over 2.5 years (28 months) in an upstream and downstream reach of Silverstream, a lowland river in New Zealand. Population densities of juvenile brown trout varied between the upstream and downstream reach, with the latter showing considerably higher temporal variation in fish density than the upstream reach. Juvenile brown trout from the downstream reach were larger and had higher condition factors than those from the upstream reach over the study period. Discharge had a negative effect on fish densities in 1 year in the downstream reach, and high water temperatures were recorded in one summer, with possible negative effects on fish densities. However, the different effects of environmental variables on populations between the two reaches could not explain the higher temporal variation in fish densities in the downstream reach as spatial variation in population densities also occurred in years without high discharge events or high summer water temperatures. Variability in food availability over time was higher in the upstream reach, but differences between the two reaches were small. These results suggest that factors other than condition of the local environment are important in controlling brown trout population dynamics in Silverstream.

Climate and socio-economic factors drive the spatio-temporal dynamics of HFRS in Northeastern China