Why are metapopulations so rare?

Roughly 40 years after its introduction, the metapopulation concept is central to population ecology. The notion that local populations and their dynamics may be coupled by dispersal is without any doubt of great importance for our understanding of population-level processes. A metapopulation describes a set of subpopulations linked by (rare) dispersal events in a dynamic equilibrium of extinctions and recolonizations. In the large body of literature that has accumulated, the term "metapopulation" is often used in a very broad sense; most of the time it simply implies spatial heterogeneity. A number of reviews have recently addressed this problem and have pointed out that, despite the large and still growing popularity of the metapopulation concept, there are only very few empirical examples that conform with the strict classical metapopulation (CM) definition. In order to understand this discrepancy between theory and observation, we use an individual-based modeling approach that allows us to pinpoint the environmental conditions and the life-history attributes required for the emergence of a CM structure. We find that CM dynamics are restricted to a specific parameter range at the border between spatially structured but completely occupied and globally extinct populations. Considering general life-history attributes, our simulations suggest that CMs are more likely to occur in arthropod species than in (large) vertebrates. Since the specific type of spatial population structure determines conservation concepts, our findings have important implications for conservation biology. Our model suggests that most spatially structured populations are panmictic, patchy, or of mainland-island type, which makes efforts spent on increasing connectivity (e.g., corridors) questionable. If one does observe a true CM structure, this means that the focal metapopulation is on the brink of extinction and that drastic conservation measures are needed.     

Antipredator strategies of house finches: are urban habitats safe spots from predators even when humans are around?

Urbanization decreases species diversity, but it increases the abundance of certain species with high tolerance to human activities. The safe-habitat hypothesis explains this pattern through a decrease in the abundance of native predators, which reduces predation risk in urban habitats. However, this hypothesis does not consider the potential negative effects of human-associated disturbance (e.g., pedestrians, dogs, cats). Our goal was to assess the degree of perceived predation risk in house finches (Carpodacus mexicanus) through field studies and semi-natural experiments in areas with different levels of urbanization using multiple indicators of risk (flock size, flight initiation distance, vigilance, and foraging behavior). Field studies showed that house finches in more urbanized habitats had a greater tendency to flock with an increase in population density and flushed at larger distances than in less urbanized habitats. In the semi-natural experiment, we found that individuals spent a greater proportion of time in the refuge patch and increased the instantaneous pecking rate in the more urbanized habitat with pedestrians probably to compensate for the lower amount of foraging time. Vigilance parameters were influenced in different ways depending on habitat type and distance to flock mates. Our results suggest that house finches may perceive highly urbanized habitats as more dangerous, despite the lower number of native predators. This could be due to the presence of human activities, which could increase risk or modify the ability to detect predators. House finches seem to adapt to the urban environment through different behavioral strategies that minimize risk.     

When are no-take zones an economically optimal fishery management strategy?

Discussions on the use of marine reserves (no-take zones) and, more generally, spatial management of fisheries are, for the most part, devoid of analyses that consider the ecological and economic effects simultaneously. To fill this gap, we develop a two-patch ecological-economic model to investigate the effects of spatial management on fishery profits. Because the fishery effects of spatial management depend critically on the nature of the ecological connectivity, our model includes both juvenile and adult movement, with density dependence in settlement differentiating the two types of dispersal. Rather than imposing a reserve on our system and measuring its effect on profits, we ask: "When does setting catch levels to maximize system-wide profits imply that a reserve should be created?" Closing areas to fishing is an economically optimal solution when the value derived from spillover from the reserve outweighs the value of fishing in the patch. The condition, while simple to state in summary form, is complex to interpret because it depends on the settlement success of the dispersing organisms, the nature of the costs of the fishing, the economic and ecological heterogeneity of the system, the discount rate, and growth characteristics of the fish population. The condition is more likely to be satisfied when the closed area is a net exporter of biomass and has higher costs of fishing, and for fish populations with density-independent settlement ("adult movement") than with density-dependent settlement ("larval dispersal"). Rather surprisingly, there are circumstances whereby closing low biological productivity areas, and even sometimes low cost areas to fish, can result in greater fishing profits than when both areas are open to fishing.     

Understanding and predicting ecological dynamics: are major surprises inevitable?

Ecological surprises, substantial and unanticipated changes in the abundance of one or more species that result from previously unsuspected processes, are a common outcome of both experiments and observations in community and population ecology. Here, we give examples of such surprises along with the results of a survey of well-established field ecologists, most of whom have encountered one or more surprises over the course of their careers. Truly surprising results are common enough to require their consideration in any reasonable effort to characterize nature and manage natural resources. We classify surprises as dynamic-, pattern-, or intervention-based, and we speculate on the common processes that cause ecological systems to so often surprise us. A long-standing and still growing concern in the ecological literature is how best to make predictions of future population and community dynamics. Although most work on this subject involves statistical aspects of data analysis and modeling, the frequency and nature of ecological surprises imply that uncertainty cannot be easily tamed through improved analytical procedures, and that prudent management of both exploited and conserved communities will require precautionary and adaptive management approaches.     

Consumer resource matching in urbanizing landscapes: are synanthropic species over-matching?

Population responses of synanthropic species to urbanization may be explained by the resource-matching rule, which postulates that individuals should distribute themselves according to resource availability. According to the resource-matching rule, urban habitats will contain greater densities if they provide better resources than rural habitats. However, because resource availability is density dependent, individuals in urban areas would ultimately achieve fitness levels comparable to, but no better than, individuals in less urban areas. Some ecologists suggest that synanthropic birds may not conform to the resource-matching rule and may instead overmatch (i.e., overexploit) in urban habitats, ultimately leading to lower fitness despite greater resource levels. Using the Northern Cardinal (Cardinalis cardinalis) as a focal species, we evaluated if Cardinal populations in urban and rural habitats were consistent with predictions of consumer resource matching. During 2003-2006 we documented population density, adult body condition, apparent survival, and annual reproductive productivity of Cardinals in riparian forest stands within urban (n = 8 stands) and rural (n = 6 stands) landscapes in Ohio, USA. Density of Cardinals in urban forests was four times that found in more rural forests. Mark-resight data from 147 males and 125 females over four years indicated that apparent survival rates were similar between urban and rural landscapes (phi = 0.64, SE = 0.039 for males and phi = 0.57, SE = 0.04 for females). Similarly, body condition indices of 168 males, 142 females, and 118 nestlings did not differ significantly between landscapes. Annual reproductive productivity (mean number of fledglings per pair over breeding season) of 294 pairs was comparable for urban (2.4 +/- 0.18 [mean +/- SE] and rural (2.1 +/- 0.18) young birds. Thus, contrary to recent suggestions, we find that high densities of certain synanthropic species in urban landscapes are consistent with expectations of consumer resource matching.     

Traditional occupancy–abundance models are inadequate for zero-inflated ecological count data

Traditional occupancy–abundance and abundance–variance–occupancy models do not take into account zero-inflation, which occurs when sampling rare species or in correlated counts arising from repeated measures. In this paper we propose a novel approach extending occupancy–abundance relationships to zero-inflated count data. This approach involves three steps: (1) selecting distributional assumptions and parsimonious models for the count data, (2) estimating abundance, occupancy and variance parameters as functions of site- and/or time-specific covariates, and (3) modelling the occupancy–abundance relationship using the parameters estimated in step 2. Five count datasets were used for comparing standard Poisson and negative binomial distribution (NBD) occupancy–abundance models. Zero-inflated Poisson (ZIP) and zero-inflated negative binomial (ZINB) occupancy–abundance models were introduced for the first time, and these were compared with the Poisson, NBD, He and Gaston's and Wilson and Room's abundance–variance–occupancy models. The percentage of zero counts ranged from 45 to 80% in the datasets analysed. For most of the datasets, the ZINB occupancy–abundance model performed better than the traditional Poisson, NBD and Wilson and Room's model. He and Gaston's model performed better than the ZINB in two out of the five datasets. However, the occupancy predicted by all models increased faster than the observed as density increased resulting in significant mismatch at the highest densities. Limitations of the various models are discussed, and the need for careful choice of count distributions and predictors in estimating abundance and occupancy parameter are indicated.     

Males are sensitive — sex-dependent effect of rearing conditions on nestling growth

The sex-dependent effect of environmental conditions on nestlings has been extensively studied in size dimorphic birds. Whether males or females are more sensitive to poor conditions is not yet clear; however, the degree of sexual size-dimorphism, brood size and their interactions seem to influence the pattern. Much less is known about sex-dependent environmental sensitivity in size-monomorphic species, even though it may result in biased sex allocation. We altered the rearing conditions by brood size manipulation in the size-monomorphic collared flycatcher and then examined the sex-specific development of the nestlings. In all analyses, we controlled for the effect of paternity, because one may expect extra-pair young to be of better genetic quality and perform better at least under poor conditions. However, this was not the case, because we did not find any difference in growth rate or fledging size between extra- and within-pair young. We found that male nestlings had the potential for faster growth under favourable conditions, but suffered more under poor conditions. We found no sex × environment interaction for fledging size probably because the growth curves level off before fledging, and the disadvantaged nestlings can catch up with their siblings. The larger sensitivity of males does not explain the previously found seasonal shift in brood sex ratios and contradicts previous findings in another size-monomorphic species where females were more sensitive. This suggests that even in size-monomorphic species, no general rule exists, which determines the more sensitive sex.     

Surplus territory contenders in male red-winged blackbirds: where are the desperados?

Summary We tested whether payoff asymmetries could explain why floater red-winged blackbirds seldom escalate contests to acquire territories. We removed territorial male \ldowners\rd and allowed floater replacements to claim territories. We then released owners to see how three currencies of payoff asymmetries (replacement occupancy) owners' likelihood and speed of reclaiming their territories (owner success). Neither the duration of an owner's internment nor the amount of time that a floater replacement had held the territory affected owner success (Figs. 1 and 2). Owner success was also not affected by the number of neighbors that they had (Fig. 3). Finally, owner success was the same irrespective of whether or not he was the likely sire of offspring on the territory, or the size of his harem (Fig. 4). Although these results are consistent with the proposal that payoff asymmetries will be irrelevant in contests for valuable resources, they are inconsistent with the proposal's corollary that excluded individuals should become \lddesperados\rd and escalate or even fight to the death in contests for those resources. Expected payoffs for passive acquisition of territories in this species may be higher than from a more aggressive desperado strategy.Correspondence to: P.J. Weatherhead     

Avian predators are less abundant during periodical cicada emergences, but why?

Despite a substantial resource pulse, numerous avian insectivores known to depredate periodical cicadas (Magicicada spp.) are detected less commonly during emergence years than in either the previous or following years. We used data on periodical cicada calls collected by volunteers conducting North American Breeding Bird Surveys within the range of cicada Brood X to test three hypotheses for this observation: lower detection rates could be caused by bird calls being obscured by cicada calls ("detectability" hypothesis), by birds avoiding areas with cicadas ("repel" hypothesis), or because bird abundances are generally lower during emergence years for some reason unrelated to the current emergence event ("true decline" hypothesis). We tested these hypotheses by comparing bird detections at stations coincident with calling cicadas vs. those without calling cicadas in the year prior to and during cicada emergences. At four distinct levels (stop, route, range, and season), parallel declines of birds in groups exposed and not exposed to cicada calls supported the true decline hypothesis. We discuss several potential mechanisms for this pattern, including the possibility that it is a consequence of the ecological and evolutionary interactions between predators of this extraordinary group of insects.     

Begging and short-term need in cowbird nestlings: how different are brood parasites?

Because brood parasitic nestlings are usually unrelated to their nestmates and to the provisioning adult, they are free from indirect costs of begging. Consequently, they are predicted to beg more intensely than host nestlings, and some models predict they will beg at an invariantly high level, regardless of short-term need. Previous work has shown intense begging by parasitic cowbirds, but short-term need was not controlled. In this study, we manipulated short-term need and measured begging intensity in two species pairs of host and parasitic nestlings: shiny cowbirds (Molothrus bonariensis) parasitizing larger hosts than themselves (rufous-bellied thrushes, Turdus rufiventris), and brown-headed cowbirds (M. ater) parasitizing smaller hosts than themselves (yellow warblers, Dendroica petechia). All four species increased their begging intensity with short-term need, though the change was much less pronounced between food-deprived and control treatments than between control and hand-feeding treatments. Shiny cowbirds begged more intensely than rufous-bellied thrushes following each treatment. In contrast, brown-headed cowbirds did not beg significantly more intensely than yellow warblers under any of the treatments. Intense begging by both species of parasites was as effective as host begging in stimulating the adults to make provisioning visits, but shiny cowbirds were less successful at acquiring food from adult thrushes. A wide array of factors may underlie the apparent differences in the begging behavior and success of brown-headed and shiny cowbirds, including relative size, experience, and local risk of nest predation. Our experiments clearly demonstrate, however, that these two species of parasitic cowbirds adjust their begging intensity based on short-term need.Communicated by H. Kokko     

Radon and lung cancer in Finland: are there signs of radiation hormesis?

Prolonged exposure to any level of radon in indoor air is generally thought to be hazardous to human health and to give rise to lung cancer. On the other hand, low or moderate levels of radon in indoor air may, according to the radiation hormesis hypothesis, be beneficial to human health and even inhibit the growth of cancerous cells. An attempt is made here to test these contrasting views by comparing mortality rates from lung cancer during the 10-year period 1986–1995 with average indoor radon levels in the 435 communes (municipalities) of Finland (excluding Åland). The results show a very low, statistically non-significant correlation between radon and lung cancer mortality (r=–0.0331), which in turn implies that at least in this case neither the possible beneficial hormetic effect of radon nor its cancer-provoking hazardous effect can make itself apparent against the dominant and masking effect of cigarette smoking, the main cause of lung cancer.     

Flight speeds observed with radar,a correction: slow “birds” are insects

Summary Based upon new data and re-analysis of previous data, some results concerning migration of birds at night are now attributed to insects. In particular, slowly flying radar targets over land (Larkin and Thompson 1980) and, by analogy, over the ocean (Larkin et al. 1979) are shown to lack wing beat patterns of birds and the appearance of birds when viewed with a radar-controlled high-power telescope and spotlamp. The insect targets overlap with birds in the amount of radar echo returned, are abundant in warmer months, and fly even in March and November in temperate North America. Other previous results are largely substantiated; however, a previous report (Larkin 1980) that migrating birds can detect the wind direction over the ocean is not substantiated.     

Diversity and host range of foliar fungal endophytes: are tropical leaves biodiversity hotspots?

Fungal endophytes are found in asymptomatic photosynthetic tissues of all major lineages of land plants. The ubiquity of these cryptic symbionts is clear, but the scale of their diversity, host range, and geographic distributions are unknown. To explore the putative hyperdiversity of tropical leaf endophytes, we compared endophyte communities along a broad latitudinal gradient from the Canadian arctic to the lowland tropical forest of central Panama. Here, we use molecular sequence data from 1403 endophyte strains to show that endophytes increase in incidence, diversity, and host breadth from arctic to tropical sites. Endophyte communities from higher latitudes are characterized by relatively few species from many different classes of Ascomycota, whereas tropical endophyte assemblages are dominated by a small number of classes with a very large number of endophytic species. The most easily cultivated endophytes from tropical plants have wide host ranges, but communities are dominated by a large number of rare species whose host range is unclear. Even when only the most easily cultured species are considered, leaves of tropical trees represent hotspots of fungal species diversity, containing numerous species not yet recovered from other biomes. The challenge remains to recover and identify those elusive and rarely cultured taxa with narrower host ranges, and to elucidate the ecological roles of these little-known symbionts in tropical forests.     

Same-sex pair-bonds are equivalent to male–female bonds in a life-long socially monogamous songbird

Same-sex sexual behaviors are well documented in both captive and wild animals. In monogamous species, these behaviors are often exclusive, each individual having only one same-sex partner. A bias in sex ratio has been proposed as a social context yielding same-sex pair-bonding, but this hypothesis has rarely been tested. Focusing on a life-long pair-bonding songbird, the zebra finch Taeniopygia guttata, we tested whether same-sex pairing results from a shortage of individuals of the opposite sex. By experimentally skewing the sex ratio towards members of one sex, we observed a greater proportion of same-sex pair-bonds of that sex. Moreover, we assessed whether the quality and stability of social interactions were equivalent in same-sex and male–female pairs. Male–male and female–female same-sex bonds display the same behavioral characteristics as male–female ones: they are intense, highly selective, and stable affinitive relationships involving the same behavioral displays already described in wild birds. Moreover, same-sex male bonds were sufficiently strong not to split up when individuals were given the opportunity to reproduce with females. Because the pair-bond in socially monogamous species represents a partnership that may give advantages for survival (e.g., resources defense, fighting against predators, etc.), we propose that same-sex pairing in the zebra finch may result from the pressure to find a social partner.     

How sensitive are elasticities of long-run stochastic growth to how environmental variability is modelled?

A variable environment leaves a signature in a population's dynamics. Deriving statistical and mathematical models of how environmental variability affects population projections has - in the wake of reports of substantial climatic fluctuations - received much recent attention. If the model changes, then so too does the population projection. This is because a different model of environmental variability changes estimates of long-run stochastic growth, which is a function of demographic rates and their temporal sequence. Decomposing elasticities of long-run stochastic growth into constituent parts can assess the relative influence of different components. Here, we investigate the consequences of changing the environmental state definition, and therefore altering the shape of demographic rate distributions and their temporal sequence, by using age-structured matrix models to project vertebrate populations into the future under a range of environmental scenarios. The identity of the most influential demographic rate was consistent among all approaches that perturbed only the mean, but was not when only the variance was perturbed. Furthermore, the influence of each demographic rate fluctuated among projections by up to factors of six and two for changes to the variance and mean, respectively. These changes in influence depend in part upon how environmental variability - in particular, the color of environmental noise - is incorporated. In the light of predictions of increasing climatic variability in the future, these results suggest caution when drawing quantitative conclusions from stochastic population projections.     

Space use in territorial amberwing dragonflies: are residents female maximizers or neighbor minimizers?

Residents of mating territories interact with different categories of conspecifics: females, nonneighbor males, and neighbor males. Interaction with these different types of conspecifics is likely to have different costs and benefits; for example, interactions with females will be beneficial, while interactions with neighbors are more likely to be costly. In this study, we investigated patterns of intrusions and space use in territorial male amberwing dragonflies (Perithemis tenera) to test the idea that residents will adjust their use of space to maximize their beneficial interactions with conspecifics while minimizing their costly interactions with conspecifics. Because territories were arranged linearly around the edge of a pond, each resident had two neighbors, one of which was often closer to the focal resident than the other. Residents experienced more intrusions by neighbors and fewer intrusions by females on the side of their closer neighbor. Residents generally perched on the side of their territory that experienced the fewest intrusions by neighbors and the most intrusions by females, but the pattern was more strongly related to neighbor intrusions than female intrusions. Subsequent to pursuits of neighbors and females, residents tended to shift their perches away from where they pursued neighbors but toward where they pursued females. Nonneighbor intrusions were not affected by neighbor proximity, nor did residents adjust their space use in response to nonneighbor intrusions. Our results suggest that residents do adjust their space use in response to intrusions by conspecifics, that their adjustment depends on the type of conspecific that intruded, and that residents may be using a simple decision rule such as "move away from male intrusions, move closer to female intrusions" to adjust their within-territory space use.Communicated by D. Gwynne     

Mortality and predation in ecosystem models: is it important how these are expressed?

The effects of the form of the grazing and mortality terms used in plankton models are well known. The same cannot be said for ecosystem models. As ecosystem models become more popular more needs to be known about the effects of model formulation on model behaviour and performance. The impact of the form of the grazing response function and mortality terms used in a biogeochemical ecosystem model are considered here. We show that in the large and inter-linked webs used in ecosystem models, model behaviour is far more sensitive to the form of the grazing term than to that of the mortality terms that close the modelled food web.When using biogeochemical ecosystem models in shallow marine ecosystems, the most dynamic and sophisticated functional responses describing grazing require more parameters and validation than the simpler Holling disk equation, but usually still lead to the same general conclusions about the system state and the effects of changes in forcing functions. Thus, the use of more complex functional responses is not necessarily warranted in many cases. Similarly, the extra effort and data required to explicitly represent the top predators (sharks, mammals and birds) is not necessary if they are not the focus of the study. A quadratic mortality term applied to intermediate predators (such as piscivores) is sufficient to achieve plausible model behaviour. It should be noted, however, that some degree of sophistication is required in the grazing and mortality terms. Use of simple linear functional responses and mortality terms is unsuitable for models used to consider a range of nutrient loading or harvesting scenarios.     

Nonparametric estimation of Shannon’s index of diversity when there are unseen species in sample

A biological community usually has a large number of species with relatively small abundances. When a random sample of individuals is selected and each individual is classified according to species identity, some rare species may not be discovered. This paper is concerned with the estimation of Shannons index of diversity when the number of species and the species abundances are unknown. The traditional estimator that ignores the missing species underestimates when there is a non-negligible number of unseen species. We provide a different approach based on unequal probability sampling theory because species have different probabilities of being discovered in the sample. No parametric forms are assumed for the species abundances. The proposed estimation procedure combines the Horvitz–Thompson (1952) adjustment for missing species and the concept of sample coverage, which is used to properly estimate the relative abundances of species discovered in the sample. Simulation results show that the proposed estimator works well under various abundance models even when a relatively large fraction of the species is missing. Three real data sets, two from biology and the other one from numismatics, are given for illustration.     

Ecological literacy and socio-demographics: who are the most eco-literate in our community,and why?

Ecological literacy is founded upon knowledge and understanding about how the ecological systems of the Earth function and support life. A widespread concern that levels of ecological literacy within many contemporary human communities are inadequate to enable effective decision-making about sustainable ways of living led to a rigorous and carefully designed assessment of over 1000 South Australian adults from diverse sectors including industry, government, nongovernment, business and community organisations. This assessment tested for local and global knowledge, including an understanding of the interconnection of natural systems with human society. Quantitative analyses revealed significant relationships between scores and a range of socio-demographic characteristics. Scores varied significantly with gender, age, education, employment, place of growing up and time spent in South Australia (SA). Highest average scores were found in males, middle-age groups, the most highly educated, those with science-related educational backgrounds, those working in environment-related fields, those who grew up in small town environments rather than in large towns or cities and in those who had spent more than 10 years in SA. These findings contribute to a broader study of factors associated with ecologically literate individuals and groups within the western and industrialised adult community of SA. They form a valuable foundation on which to build efforts to further understand and develop ecological literacy within our citizenry and governing bodies.     

Air pollutant levels are 12 times higher than guidelines in Varanasi,India. Sources and transfer

Air quality in an urban atmosphere is regulated by both local and distant emission sources. For air quality management in urban areas, identification of sources and their relationships with local meteorology and air pollutants are essential. The critical condition of air quality in Indo-Gangetic plain is well known, but lack of data on both local and distant emission sources limits the scope of improving air quality in this region. Concentrations of particulate matter of size lower than 10 μm (PM₁₀) were assessed in the highly urbanized Varanasi city situated in middle Indo-Gangetic plain of India from 2014 to 2017, to identify the distant air pollution sources based on trajectory statistical models and local sources by conditional bivariate probability function. Modifying effects of meteorology and air pollutants on PM₁₀ were also explored. Mean PM₁₀ concentration for the study period was 244.8 ± 135.8 μg m^?3, which was 12 times higher than the WHO annual guideline. Several distinct sources of traffic as the major source of PM₁₀ were identified in the city. Trajectory statistical models like cluster analysis, potential source contribution function and concentration-weighted trajectory showed significant contributions from north-west and eastern directions in the transport of polluted air masses to the city. Dew point, wind speed, temperature and ventilation coefficient are the major factors in PM₁₀ formation and dispersion.