Thermal and energetic constraints on ectotherm abundance: a global test using lizards

Population densities of birds and mammals have been shown to decrease with body mass at approximately the same rate as metabolic rates increase, indicating that energetic needs constrain endotherm population densities. In ectotherms, the exponential increase of metabolic rate with body temperature suggests that environmental temperature may additionally constrain population densities. Here we test simple bioenergetic models for an ecologically important group of ectothermic vertebrates by examining 483 lizard populations. We find that lizard population densities decrease as a power law of body mass with a slope approximately inverse to the slope of the relationship between metabolic rates and body mass. Energy availability should limit population densities. As predicted, environmental productivity has a positive effect on lizard density, strengthening the relationship between lizard density and body mass. In contrast, the effect of environmental temperature is at most weak due to behavioral thermoregulation, thermal evolution, or the temperature dependence of ectotherm performance. Our results provide initial insights into how energy needs and availability differentially constrain ectotherm and endotherm density across broad spatial scales.     

Rarity in Neotropical Forest Mammals Revisited

The identification of rare species is an important goal in conservation biology. Recent attempts to classify rare species have emphasized dichotomies in such characteristics as local population density, area of distribution, and degree of ecological specialization. In particular, Arita et al. (1990) dichotomized 100 Neotropical forest mammals according to local population density and area of distribution. Among these species of mammals, mean body mass was significantly associated with local population density and area of distribution. We argue that the effects of body mass should be removed before species are classified with respect to rarity. We re-evaluated the data on Neotropical mammal species, using regression analyses to remove the effects of body mass on population density and area of distribution, followed by analysis of residuals. This new method resulted in substantial changes in the dichotomous classification of rare species. We combined the analysis of regression residuals with a ranking procedure that assumed that local population density and area of distribution were equally important in their effects on rarity. The new ranking technique produced another different classification of the rarity of the Neotropical forest mammal species. A graphical analysis showed that ranked species differed substantially in their degree of rarity, and in the importance of local population density, area of distribution, or both, to their degree of rarity. The ranking method allows the species of greatest concern to be singled out, it can be modified to include additional variables such as niche breadth, and it should be helpful for making conservation decisions.     

Setting population targets for mammals using body mass as a predictor of population persistence

Conservation planning and biodiversity assessments need quantitative targets to optimize planning options and assess the adequacy of current species protection. However, targets aiming at persistence require population‐specific data, which limit their use in favor of fixed and nonspecific targets, likely leading to unequal distribution of conservation efforts among species. We devised a method to derive equitable population targets; that is, quantitative targets of population size that ensure equal probabilities of persistence across a set of species and that can be easily inferred from species‐specific traits. In our method, we used models of population dynamics across a range of life‐history traits related to species’ body mass to estimate minimum viable population targets. We applied our method to a range of body masses of mammals, from 2 g to 3825 kg. The minimum viable population targets decreased asymptotically with increasing body mass and were on the same order of magnitude as minimum viable population estimates from species‐ and context‐specific studies. Our approach provides a compromise between pragmatic, nonspecific population targets and detailed context‐specific estimates of population viability for which only limited data are available. It enables a first estimation of species‐specific population targets based on a readily available trait and thus allows setting equitable targets for population persistence in large‐scale and multispecies conservation assessments and planning.     

Litter reductions reveal a trade-off between offspring size and number in brown bears

Experimental manipulations have revealed positive effects of litter reduction on offspring mass in small mammals, but little is known about this trade-off in large mammals. We examined the determinants of natural litter size variation and quantified the effects of litter size, maternal characteristics, and litter composition on yearling mass using 24?years of data on marked brown bears (Ursus arctos) in Sweden. Infanticide by adult males is a major cause of cub-of-the-year mortality, leading to litter size reductions. Litter size (n?=?265) at den emergence ranged from one to four cubs (average, 2.7) and increased with maternal age. Litter size, however, appeared independent of maternal size, population density, interlitter interval, study area, or previous litter sex ratio. Yearling body mass increased with maternal body size but was independent of litter sex ratio. Litter size and yearling mass were negatively correlated, mostly because singletons were about 30?% heavier than yearlings from litters of two to four cubs. In reduced litters, survivors were on average 8?% heavier as yearlings than individuals from intact litters, suggesting that sibling competition reduces growth. Trade-offs between litter size and yearling mass in bears appear similar in magnitude to those found in small mammals.     

Estimates of minimum patch size depend on the method of estimation and the condition of the habitat

Minimum patch size for a viable population can be estimated in several ways. The density-area method estimates minimum patch size as the smallest area in which no new individuals are encountered as one extends the arbitrary boundaries of a study area outward. The density-area method eliminates the assumption of no variation in density with size of habitat area that accompanies other methods, but it is untested in situations in which habitat loss has confined populations to small areas. We used a variant of the density area method to study the minimum patch size for the gopher tortoise (Gopherus polyphemus) in Florida, USA, where this keystone species is being confined to ever smaller habitat fragments. The variant was based on the premise that individuals within populations are likely to occur at unusually high densities when confined to small areas, and it estimated minimum patch size as the smallest area beyond which density plateaus. The data for our study came from detailed surveys of 38 populations of the tortoise. For all 38 populations, the areas occupied were determined empirically, and for 19 of them, duplicate surveys were undertaken about a decade apart. We found that a consistent inverse density area relationship was present over smaller areas. The minimum patch size estimated from the density-area relationship was at least 100 ha, which is substantially larger than previous estimates. The relative abundance of juveniles was inversely related to population density for sites with relatively poor habitat quality, indicating that the estimated minimum patch size could represent an extinction threshold. We concluded that a negative density area relationship may be an inevitable consequence of excessive habitat loss. We also concluded that any detrimental effects of an inverse density area relationship may be exacerbated by the deterioration in habitat quality that often accompanies habitat loss. Finally, we concluded that the value of any estimate of minimum patch size as a conservation tool is compromised by excessive habitat loss.     

Savviness of prey to introduced predators

The prey naivety hypothesis posits that prey are vulnerable to introduced predators because many generations in slow gradual coevolution are needed for appropriate avoidance responses to develop. It predicts that prey will be more responsive to native than introduced predators and less responsive to introduced predators that differ substantially from native predators and from those newly established. To test these predictions, we conducted a global meta-analysis of studies that measured the wariness responses of small mammals to the scent of sympatric mammalian mesopredators. We identified 26 studies that met our selection criteria. These studies comprised 134 experiments reporting on the responses of 36 small mammal species to the scent of six introduced mesopredators and 12 native mesopredators. For each introduced mesopredator, we measured their phylogenetic and functional distance to local native mesopredators and the number of years sympatric with their prey. We used predator and prey body mass as a measure of predation risk. Globally, small mammals were similarly wary of the scent of native and introduced mesopredators; phylogenetic and functional distance between introduced mesopredators and closest native mesopredators had no effect on wariness; and wariness was unrelated to the number of prey generations, or years, since first contact with introduced mesopredators. Small mammal wariness was associated with predator-prey body mass ratio, regardless of the nativity. The one thing animals do not seem to recognize is whether their predators are native.     

Density-related reproductive trade-offs in the green sea urchin, Strongylocentrotus droebachiensis

Green sea urchin, Strongylocentrotus droebachiensis (O.F. Müller), populations are being depleted rapidly in the Gulf of Maine and there is justified concern that the potential of this free-spawner to produce larvae may be severely inhibited. We evaluated the opposing effects of different population densities on gonad development and fertilization success, using population surveys and fertilization experiments. We determined gonad indices (gonad mass/body mass) over a range of population densities (0.1 to 250 ind. m⁻²) at seven sites in coastal Maine, USA, sampled at two depths (5 and 15 m). At shallow sites, we found that gonad indices declined by 50% over the 1500-fold range in adult population density. At 15 m deep locations, gonad mass was consistently low and did not vary significantly with density. Patterns of macroalgal abundance suggest urchins at high density and in deeper water were food limited. Because macroalgal cover co-varies inversely with sea urchin density, we designed field experiments to determine the interaction between sea urchin density and kelp canopy on fertilization success. On square arrays we manipulated the spacing of simulated urchins, but held their numbers constant (five sperm-filled syringes interspersed with four Nitex mesh egg containers permeable to sperm). These experiments, simulating the observed range of natural density, suggested that (1) fertilization rates decreased many times faster than individual gamete production increased over the same range in density, and (2) kelp increased fertilization success at high density when eggs were within 25 cm of a sperm source, but not when spaced 1 m apart. Additional laboratory fertilization experiments at ambient temperatures (3 to 5 °C) indicated that diluted sperm were viable for <1 h, but egg viability was virtually unchanged for >8 h. In short, to the individual the reproductive benefits of aggregating appear to outweigh the costs; and while sperm may be limiting at low population density, eggs may remain viable long enough to be fertilized by sperm from more distant males. Received: 10 February 1998 / Accepted: 22 January 1999     

Effects of different management strategies on long-term trends of Australian threatened and near-threatened mammals

Monitoring is critical to assess management effectiveness, but broadscale systematic assessments of monitoring to evaluate and improve recovery efforts are lacking. We compiled 1808 time series from 71 threatened and near-threatened terrestrial and volant mammal species and subspecies in Australia (48% of all threatened mammal taxa) to compare relative trends of populations subject to different management strategies. We adapted the Living Planet Index to develop the Threatened Species Index for Australian Mammals and track aggregate trends for all sampled threatened mammal populations and for small (<35 g), medium (35–5500 g), and large mammals (>5500 g) from 2000 to 2017. Unmanaged populations (42 taxa) declined by 63% on average; unmanaged small mammals exhibited the greatest declines (96%). Populations of 17 taxa in havens (islands and fenced areas that excluded or eliminated introduced red foxes [Vulpes vulpes] and domestic cats [Felis catus]) increased by 680%. Outside havens, populations undergoing sustained predator baiting initially declined by 75% but subsequently increased to 47% of their abundance in 2000. At sites where predators were not excluded or baited but other actions (e.g., fire management, introduced herbivore control) occurred, populations of small and medium mammals declined faster, but large mammals declined more slowly, than unmanaged populations. Only 13% of taxa had data for both unmanaged and managed populations; index comparisons for this subset showed that taxa with populations increasing inside havens declined outside havens but taxa with populations subject to predator baiting outside havens declined more slowly than populations with no management and then increased, whereas unmanaged populations continued to decline. More comprehensive and improved monitoring (particularly encompassing poorly represented management actions and taxonomic groups like bats and small mammals) is required to understand whether and where management has worked. Improved implementation of management for threats other than predation is critical to recover Australia's threatened mammals.     

Patterns of Genetic Diversity and Its Loss in Mammalian Populations

Abstract:  Policy aimed at conserving biodiversity has focused on species diversity. Loss of genetic diversity, however, can affect population persistence, evolutionary potential, and individual fitness. Although mammals are a well-studied taxonomic group, a comprehensive assessment of mammalian genetic diversity based on modern molecular markers is lacking. We examined published microsatellite data from populations of 108 mammalian species to evaluate background patterns of genetic variability across taxa and body masses. We tested for loss of genetic diversity at the population level by asking whether populations that experienced demographic threats exhibited lower levels of genetic diversity. We also evaluated the effect of ascertainment bias (a reduction in variability when microsatellite primers are transferred across species) on our assessment of genetic diversity. Heterozygosity did not vary with body mass across species ranging in size from shrews to whales. Differences across taxonomic groupings were noted at the highest level, between populations of marsupial and placental mammals. We documented consistently lower heterozygosity, however, in populations that had experienced demographic threats across a wide range of mammalian species. We also documented a significant ( p = 0.01) reduction in heterozygosity as a result of ascertainment bias. Our results suggest that populations of both rare and common mammals are currently losing genetic diversity and that conservation efforts focused above the population level may fail to protect the breadth of persisting genetic diversity. Conservation policy makers may need to focus their efforts below the species level to stem further losses of genetic resources.     

Effects of Human Activity on Global Extinction Risk

Both natural and anthropogenic factors are important in determining a species' risk of extinction. Little work has been done, however, to quantify the magnitude of current anthropogenic influences on the extinction process. The purpose of this study is to determine the extent to which measures of the intensity of human activity are related to the global variability of two measures of species' susceptibility to extinction. We observed six indices of human activities in 90 countries, and we tested their relationships to the proportion of threatened bird and mammal species in each country, as well as to mammalian population density. After correcting for area effects, latitudinal diversity gradients, and body size (for population density), 28 to 50% of the remaining variation was statistically attributable to anthropogenic variables. Different measures of anthropogenic influence were most closely related to extinction risk in birds and mammals. Human population density was the variable most closely related to the proportion of threatened bird species per country, whereas per capita GNP was more important for mammal species. Mammalian population density strongly correlates with the extent of protected area per country. Contrary to suggestions in earlier literature, our work does not support the hypothesis that habitat loss is a prime contributor to species loss because frequencies of threatened birds and mammals are not closely related to patterns of land use.     

Conservation of Mammals in Eastern North American Wildlife Reserves: How Small Is Too Small?

Abstract: A common objective of methods of systematic reserve selection has been to maximize conservation benefits—frequently current species richness—while reducing the costs of acquiring and maintaining reserves. But the probability that a reserve will lose species in the future is frequently not known because the minimum area requirements for most species have not been estimated empirically. For reserves within the Alleghenian-Illinoian mammal province of eastern North America, we empirically estimated the minimum area requirement of terrestrial mammals such that reserves should not lose species because of insularization. We compared this estimate to the actual size of 2355 reserves and reserve assemblages within the mammal province. The estimated minimum area requirement was 5037 km² (95% CI: 2700–13,296 km²). Fourteen reserves and reserve assemblages were> 2700 km², 9 were> 5037 km², and 3 were> 13,296 km². These 14 reserves accounted for 73% of the total area of reserves and 10% of the total area of the mammal province. Few reserves appear large enough to avoid loss of some mammal species without the additional cost of active management of habitat or populations. Immigration corridors and buffer zones that combine small reserves into assemblages totaling at least 2700 km² may be the most efficient means of conserving mammals in these reserves.     

Correlates of bushmeat in markets and depletion of wildlife

We used data on number of carcasses of wildlife species sold in 79 bushmeat markets in a region of Nigeria and Cameroon to assess whether species composition of a market could be explained by anthropogenic pressures and environmental variables around each market. More than 45 mammal species from 9 orders were traded across all markets; mostly ungulates and rodents. For each market, we determined median body mass, species diversity (game diversity), and taxa that were principal contributors to the total number of carcasses for sale (game dominance). Human population density in surrounding areas was significantly and negatively related to the percentage ungulates and primates sold in markets and significantly and positively related to the proportion of rodents. The proportion of carnivores sold was higher in markets with high human population densities. Proportion of small‐bodied mammals (<1 kg) sold in markets increased as human population density increased, but proportion of large‐bodied mammals (>10 kg) decreased as human population density increased. We calculated an index of game depletion (GDI) for each market from the sum of the total number of carcasses traded per annum and species, weighted by the intrinsic rate of natural increase (r_max) of each species, divided by individuals traded in a market. The GDI of a market increased as the proportion of fast‐reproducing species (highest r_max) increased and as the representation of species with lowest r_max (slow‐reproducing) decreased. The best explanatory factor for a market's GDI was anthropogenic pressure—road density, human settlements with >3000 inhabitants, and nonforest vegetation. High and low GDI were significantly differentiated by human density and human settlements with >3000 inhabitants. Our results provided empirical evidence that human activity is correlated with more depleted bushmeat faunas and can be used as a proxy to determine areas in need of conservation action.     

Population feedback after successful invasion leads to ecological suicide in seasonal environments

For most consumer species, winter represents a period of harsh food conditions in addition to the physiological strain that results from the low ambient temperatures. In size-structured populations, larger-bodied individuals do better during winter as they have larger energy reserves to buffer starvation periods. In contrast, smaller-bodied individuals do better under growing conditions, as they have lower maintenance costs. We study how the interplay between size-dependent life-history processes and seasonal changes in temperature and food availability shape the long-term dynamics of a size-structured consumer population and its unstructured resource. We show that the size dependence of maintenance requirements translates into a minimum body size that is needed for surviving starvation when consumers can adapt only to a limited extent to the low food densities in winter. This size threshold can lead to population extinction because adult individuals suffer only a little during winter and hence produce large numbers of offspring. Due to population feedback on the resource and intense intra-cohort competition, newborn consumers then fail to reach the size threshold for survival. Under these conditions, small numbers of individuals can survive, increase in density, and build up a population, which will subsequently go extinct due to its feedback on the resource. High juvenile mortality may prevent this ecological suicide from occurring, as it releases resource competition among newborns and speeds up their growth. In size-structured populations, annual fluctuations in temperature and food availability may thus lead to a conflict between individual fitness and population persistence.     

The interplay between environmental conditions and allee effects during the recovery of stressed zooplankton communities

Many important ecological phenomena depend on the success or failure of small introduced populations. Several factors are thought to influence the fate of small populations, including resource and habitat availability, dispersal levels, interspecific interactions, mate limitation, and demographic stochasticity. Recent field studies suggest that Allee effects resulting from mate limitation can prevent the reestablishment of sexual zooplankton species following a disturbance. In this study, we explore the interplay between Allee effects and local environmental conditions in determining the population growth and establishment of two acid-sensitive zooplankton species that have been impacted by regional anthropogenic acidification. We conducted a factorial design field experiment to test the impact of pH and initial organism densities on the per capita population growth (r) of the sexual copepod Epischura lacustris and the seasonally parthenogenetic cladoceran Daphnia mendotae. In addition, we conducted computer simulations using r values obtained from our experiments to determine the probability of extinction for small populations of acid-sensitive colonists that are in the process of colonizing recovering lakes. The results of our field experiment demonstrated that local environmental conditions can moderate the impacts of Allee effects for E. lacustris: Populations introduced at low densities had a significantly lower r at pH 6 than at pH 7. In contrast, r did not differ between pH 6 and 7 environments when E. lacustris populations were introduced at high densities. D. mendotae was affected by pH levels, but not by initial organism densities. Results from our population growth simulations indicated that E. lacustris populations introduced at low densities to pH 6 conditions had a higher probability of extinction than those introduced at low densities to a pH 7 environment. Our study indicates that environmental conditions and mate limitation can interact to determine the fate of small populations of sexually reproducing zooplankton species. If a more rapid recovery of acid-damaged zooplankton communities is desired, augmentation of dispersal levels may be needed during the early phases of pH recovery in order to increase the probability of establishment for mate-limited zooplankton species.     

Bycatch of Marine Mammals in U.S. and Global Fisheries

Abstract:  Fisheries bycatch poses a significant threat to many populations of marine mammals, but there are few published estimates of the magnitude of these catches. We estimated marine mammal bycatch in U.S. fisheries from 1990 to 1999 with data taken from the stock assessment reports required by the U.S. Marine Mammal Protection Act. The mean annual bycatch of marine mammals during this period was 6215 ± 448 (SE). Bycatch of cetaceans and pinnipeds occurred in similar numbers. Most cetacean (84%) and pinniped (98%) bycatch occurred in gill-net fisheries. Marine mammal bycatch declined significantly over the decade, primarily because of a reduction in the bycatch of cetaceans. Total marine mammal bycatch was significantly lower after the implementation of take reduction measures in the latter half of the decade. We derived a crude first estimate of marine mammal bycatch in the world's fisheries by expanding U.S. bycatch with data on fleet composition from the Food and Agriculture Organization. The global bycatch of marine mammals is in the hundreds of thousands. Bycatch is likely to have significant demographic effects on many populations of marine mammals. Better data are urgently needed to fully understand the impact of these interactions.     

Hunting for Sustainability in Tropical Secondary Forests

Abstract:  The interaction between land-use change and the sustainability of hunting is poorly understood but is critical for sustaining hunted vertebrate populations and a protein supply for the rural poor. We investigated sustainability of hunting in an Amazonian landscape mosaic, where a small human population had access to large areas of both primary and secondary forest. Harvestable production of mammals and birds was calculated from density estimates. We compared production with offtake from three villages and used catch-per-unit-effort as an independent measure of prey abundance. Most species were hunted unsustainably in primary forest, leading to local depletion of the largest primates and birds. The estimated sustainable supply of wild meat was higher for primary (39 kg · km⁻²· yr⁻¹) than secondary forest (22 kg · km⁻²· yr⁻¹) because four species were absent and three species at low abundance in secondary forests. Production of three disturbance-tolerant mammal species was 3 times higher in secondary than in primary forest, but hunting led to overexploitation of one species. Our data suggest that an average Amazonian smallholder would require ≥3.1 km² of secondary regrowth to ensure a sustainable harvest of forest vertebrates. We conclude that secondary forests can sustainably provide only 2% of the required protein intake of Amazonian smallholders and are unlikely to be sufficient for sustainable hunting in other tropical forest regions.     

Sink populations in carnivore management: cougar demography and immigration in a hunted population.

Carnivores are widely hunted for both sport and population control, especially where they conflict with human interests. It is widely believed that sport hunting is effective in reducing carnivore populations and related human-carnivore conflicts, while maintaining viable populations. However, the way in which carnivore populations respond to harvest can vary greatly depending on their social structure, reproductive strategies, and dispersal patterns. For example, hunted cougar (Puma concolor) populations have shown a great degree of resiliency. Although hunting cougars on a broad geographic scale (> 2000 km2) has reduced densities, hunting of smaller areas (i.e., game management units, < 1000 km2), could conceivably fail because of increased immigration from adjacent source areas. We monitored a heavily hunted population from 2001 to 2006 to test for the effects of hunting at a small scale (< 1000 km2) and to gauge whether population control was achieved (lambda < or = 1.0) or if hunting losses were negated by increased immigration allowing the population to remain stable or increase (lambda > or = 1.0). The observed growth rate of 1.00 was significantly higher than our predicted survival/fecundity growth rates (using a Leslie matrix) of 0.89 (deterministic) and 0.84 (stochastic), with the difference representing an 11-16% annual immigration rate. We observed no decline in density of the total population or the adult population, but a significant decrease in the average age of independent males. We found that the male component of the population was increasing (observed male population growth rate, lambda(OM) = 1.09), masking a decrease in the female component (lambda(OF) = 0.91). Our data support the compensatory immigration sink hypothesis; cougar removal in small game management areas (< 1000 km2) increased immigration and recruitment of younger animals from adjacent areas, resulting in little or no reduction in local cougar densities and a shift in population structure toward younger animals. Hunting in high-quality habitats may create an attractive sink, leading to misinterpretation of population trends and masking population declines in the sink and surrounding source areas.     

Studies on comparative population growth of some species of the rotifer Lecane (Rotifera)

We compared the population growth patterns of 5 species of the rotifer genus Lecane [(L. quadridentata (Ehrenberg, 1830), L. comuta (Muller, 1786), L. papuana (Murray, 1913), L. unguitata (Fadeev, 1925) and L. pyriformis (Daday, 1905)] ranging in adult average body size from 30 to 140 microm. All species were cultured under laboratory conditions for 25-30 days using the green alga Scenedesmus acutus as the exclusive diet, at a density of 1.0 x 10(6) cells ml(-1) at 24 degrees C. Regardless of the species, lecanids reached their peak population densities after 4 weeks. Peak population densities ranged from 15 to 320 ind. ml(-1), depending on body size. There was an inverse curvilinear relation between body lengths and peak population abundances (densities) of the Lecane species. Egg ratios (eggs per female) for the tested species were < 0.6 during the exponential phase but declined to 0.1 (or lower) as the population density increased. The rates of population increase for the lecanids were in general lower(0.10 to 0.21 day (-1)) than other well-studied rotifer species including members of Brachionidae.     

Genetic structure,lack of sex-biased dispersal and behavioral flexibility in the pair-living fat-tailed dwarf lemur, <Emphasis Type="Italic">Cheirogaleus medius</Emphasis>

Mating system and dispersal patterns influence the spatio-genetic structure within and between populations. Among mammals, monogamy is rare, and its socio-genetic consequences have not been studied in detail before. The goal of our study was to investigate population history, demographic structure, and dispersal patterns in a population of pair-living fat-tailed dwarf lemurs, Cheirogaleus medius, a small, nocturnal primate from western Madagascar, and to infer their underlying behavioral mechanisms. Tissue samples for DNA extraction were obtained from a total of 140 individuals that were captured in two subpopulations about 3 km apart. Analyses of mtDNA variability at the population level revealed very low levels of genetic variability combined with high haplotype diversity, which is indicative of a recent population bottleneck. We found no evidence for spatial clustering of same-sexed individuals with identical haplotypes within each of two subpopulations but significant clustering between them. Thus, a high level of local subpopulation differentiation was observed (F _ST = 0.230). The sexes showed equal variances in the number of individuals representing each haplotype, as well as equal levels of aggregation of identical haplotypes. Hence, both sexes disperse from their natal area, one pattern expected in a pair-living mammal. There is a possibility of behavioral and social flexibility in this species, however, because we documented pronounced differences in density and sex ratio between the two subpopulations, suggesting that single study sites or populations may not be representative of a given local population or even species.     

Spatial heterogeneity in distribution and ecology of Western Palearctic birds

Species vary in abundance and heterogeneity of spatial distribution, and the ecological and evolutionary consequences of such variability are poorly known. Evolutionary adaptation to heterogeneously distributed resources may arise from local adaptation with individuals of such locally adapted populations rarely dispersing long distances and hence having small populations and small overall ranges. We quantified mean population density and spatial heterogeneity in population density of 197 bird species across 12 similarly sized regions in the Western Palearctic. Variance in population density among regions differed significantly from a Poisson distribution, suggesting that random processes cannot explain the observed patterns. National estimates of means and variances in population density were positively correlated with continental estimates, suggesting that means and variances were maintained across spatial scales. We used Morisita's index of population abundance as an estimate of heterogeneity in distribution among regions to test a number of predictions. Heterogeneously distributed passerine bird species as reflected by Morisita's index had small populations, low population densities, and small breeding ranges. Their breeding populations had been consistently maintained at low levels for considerable periods of time, because the degree of genetic variation in a subsample of non-passerines and passerines was significantly negatively related to heterogeneity in distribution. Heterogeneously distributed passerine species were not more often habitat specialists than homogeneously distributed species. Furthermore, heterogeneously distributed passerine species had high annual adult survival rates but did not differ in annual fecundity from homogeneously distributed species. Heterogeneously distributed passerine species rarely colonized urban habitats. Finally, homogeneously distributed bird species were hosts to a greater diversity of blood parasite species than heterogeneously distributed species. In conclusion, small breeding ranges, population sizes, and population densities of heterogeneously distributed passerine bird species, combined with their low degree of genetic variability, and their inability to colonize urban areas may render such species particularly susceptible to human-influenced global climatic changes.