Effect of Road Traffic on Two Amphibian Species of Differing Vagility

Abstract: Vehicular traffic can be a major source of mortality for some species. Highly vagile organisms may be at a disadvantage in landscapes with roads because they are more likely to encounter roads and incur traffic mortality. To test this prediction, we assessed the population abundance of two anuran species of differing vagility, the leopard frog (    Rana pipiens , more vagile) and the green frog (    Rana clamitans , less vagile), at 30 breeding ponds. Traffic density, an index of the amount of potential traffic mortality, was measured in concentric circles radiating from the ponds out to 5 km. We conducted multiple linear regressions relating population abundance to traffic density, pond variables, and landscape habitat variables and found that leopard frog population density was negatively affected by traffic density within a radius of 1.5 km. There was no evidence that the presence of vehicular traffic affected green frog populations. These results suggest that traffic mortality can cause population declines and that more vagile species may be more vulnerable to road mortality than less vagile species.     

Metapopulation Dynamics and Amphibian Conservation

Abstract: In many respects, amphibian spatial dynamics resemble classical metapopulation models, in which subpopulations in breeding ponds blink in and out of existence and extinction and colonization rates are functions of pond spatial arrangement. This "ponds-as-patches" view of amphibian spatial dynamics is useful in several respects. First, it highlights the importance of regional and landscape processes in determining local patterns of abundance. Second, it offers a straightforward, pond-based approach to monitoring and managing amphibian populations. For many species, however, the ponds-as-patches view may be an oversimplification and metapopulation structure may be more apparent than real. Changes in distribution may be caused by processes other than extinction and recolonization, and most extinctions probably result from deterministic factors, not stochastic processes. In addition, the effects of pond isolation appear to be important primarily in disturbed environments, and in many cases these isolation effects may be better explained by the distribution of terrestrial habitats than by the distribution of breeding ponds. These complications have important implications for both researchers and managers. For researchers, future efforts need to determine the mechanisms underlying patterns of abundance and distributional change and patterns in amphibian populations. For managers, effective conservation strategies must successfully balance metapopulation considerations with careful attention to local habitat quality. Furthermore, translocations and active management may be indispensable tools for conserving amphibians in landscapes containing multiple breeding ponds.     

Habitat Matrix Effects on Pond Occupancy in Newts

Abstract: In farmlands, the population viability of many amphibians is suspected to depend on the resistance the matrix of crop fields presents to movements between ponds and terrestrial sites and movements among ponds. Over recent decades the increase in cereal growing at the expense of cattle breeding has caused a drastic change in habitat matrix in many European regions. We investigated the effect of such change on populations of three newt species (   Triturus helveticus , T. alpestris , and T. cristatus ) by comparing their abundances in sites that varied in amount of cultivated ground. A multivariate regression analysis of the relationship of newt abundance to both pond and landscape variables demonstrated the negative influence of cultivated ground on abundance. The width of the uncultivated sector linking the pond to the forest was a good predictor of abundance after the influences of both pond area and fish presence were removed. Moreover, newt presence was positively related to the number of ponds within that 50-ha surrounding area, highlighting the role of metapopulation functioning in newt occupancy of ponds. The relationship between newt abundance and width of uncultivated sectors agrees with present knowledge of the orientation mechanisms that underlie migration movements in urodeles. Such a relationship between connectedness and sector width shows that narrow, linear corridors such as hedgerows may not be useful in newt conservation. Our study also highlights the need to incorporate a behavioral component of habitat use into models of connectivity in conservation biology.     

Examining the relationship between local extinction risk and position in range

Over half of globally threatened animal species have experienced rapid geographic range loss. Identifying the parts of species’ distributions most vulnerable to local extinction would benefit conservation planning. However, previous studies give little consensus on whether ranges decline to the core or edge. We built on previous work by using empirical data to examine the position of recent local extinctions within species’ geographic ranges, address range position as a continuum, and explore the influence of environmental factors. We aggregated point‐locality data for 125 Galliform species from across the Palearctic and Indo‐Malaya into equal‐area half‐degree grid cells and used a multispecies dynamic Bayesian occupancy model to estimate rates of local extinctions. Our model provides a novel approach to identify loss of populations from within species ranges. We investigated the relationship between extinction rates and distance from range edge by examining whether patterns were consistent across biogeographic realm and different categories of land use. In the Palearctic, local extinctions occurred closer to the range edge than range core in both unconverted and human‐dominated landscapes. In Indo‐Malaya, no pattern was found for unconverted landscapes, but in human‐dominated landscapes extinctions tended to occur closer to the core than the edge. Our results suggest that local and regional factors override general spatial patterns of recent local extinction within species’ ranges and highlight the difficulty of predicting the parts of a species’ distribution most vulnerable to threat.     

Effects of Pond Draining on Biodiversity and Water Quality of Farm Ponds

Farm ponds have high conservation value because they contribute significantly to regional biodiversity and ecosystem services. In Japan pond draining is a traditional management method that is widely believed to improve water quality and eradicate invasive fish. In addition, fishing by means of pond draining has significant cultural value for local people, serving as a social event. However, there is a widespread belief that pond draining reduces freshwater biodiversity through the extirpation of aquatic animals, but scientific evaluation of the effectiveness of pond draining is lacking. We conducted a large‐scale field study to evaluate the effects of pond draining on invasive animal control, water quality, and aquatic biodiversity relative to different pond‐management practices, pond physicochemistry, and surrounding land use. The results of boosted regression‐tree models and analyses of similarity showed that pond draining had little effect on invasive fish control, water quality, or aquatic biodiversity. Draining even facilitated the colonization of farm ponds by invasive red swamp crayfish (Procambarus clarkii), which in turn may have detrimental effects on the biodiversity and water quality of farm ponds. Our results highlight the need for reconsidering current pond management and developing management plans with respect to multifunctionality of such ponds. Efectos del Drenado de Estanques sobre la Biodiversidad y la Calidad del Agua en Estanques de Cultivo     

A Concentric Analysis of the Impact of Urbanization on the Threatened European Tree Frog in an Agricultural Landscape

Abstract:  Pond-breeding amphibians are affected by site-specific factors and regional and landscape-scale patterns of land use. Recent anthropogenic landscape modifications (drainage, agriculture intensification, larger road networks, and increased traffic) affect species by reducing the suitable habitat area and fragmenting remaining populations. Using a robust concentric approach based on permutation tests, we evaluated the impact of recent landscape changes on the presence of the endangered European tree frog ( Hyla arborea ) in wetlands. We analyzed the frequency of 1 traffic and 14 land-use indices at 20 circular ranges (from 100-m up to 2-km radii) around 76 ponds identified in western Switzerland. Urban areas and road surfaces had a strong adverse effect on tree frog presence even at relatively great distances (from 100 m up to 1 km). When traffic measurements were considered instead of road surfaces, the effect increased, suggesting a negative impact due to a vehicle-induced effect. Altogether, our results indicate that urbanization and traffic must be taken into account when pond creation is an option in conservation management plans, as is the case for the European tree frog in western Switzerland. We conclude that our easy-to-use and robust concentric method of analysis can successfully assist managers in identifying potential sites for pond creation, where probability of the presence of tree frogs is maximized.     

快速城市化地区塘的消减过程研究--以广州市天河区为例

塘是我国南方地区常见的、分布广泛的景观类型,也是相对独立的水生生态系统,在生态系统服务和人类经济文化活动中扮演着非常重要的角色。国际学者对其在保护生物多样性、防涝排水、提升景观及改善人居环境等方面的生态功能研究较多,但国内对塘的形成、消减过程、生态环境功能效应等科学问题一直认识不足,在人口和土地的快速城市化胁迫下,为了增加建设用地使用面积而将水塘填埋的行为广泛存在。文章以广州市天河区为研究区,以1996、2000、2004、2007、2011年5期遥感影像为数据,通过目视解译获取研究区的土地利用特征,提取水塘信息,依据塘的主要功能对塘进行分类,通过缓冲区分析、主成份分析和转移矩阵分析探索快速城市化地区水塘消减过程的数量和空间分布特征。结果表明：（1）研究区内水塘景观在15 a间显著消减,数量由109个减至20个,面积由36.30 hm2降至13.18 hm2,个体减少率为81.65%,面积减少率为63.69%;（2）不同时段、不同功能类型的水塘消减过程存在明显差异,养殖塘、风水塘、山塘和灌溉塘面积急剧减少,景观塘面积先降后增;（3）各时段的塘转换主要去向为居住用地、商业及工矿用地等,周边土地利用格局有交通用地、荒草地的塘更易于被转换成建设用地等其他土地利用类型;（4）在城市人居环境建设中,水塘具有保持生物多样性、调节气温、改善气候、防旱排涝、改善人居环境等生态价值,又具有历史文化、景观美化等人文价值,但水塘已成为快速城市化地区居住、商业、工矿用地的主要侵占对象。作为冷岛的水塘生态系统,在城市化进程和各项规划中都应受到重视与保护。     

Spatial variance in abundance and occupancy of corals across broad geographic scales

Species assemblages vary in structure due to a wide variety of processes operating at ecological and much broader biogeographical scales. Cross-scale studies of assemblage structure are necessary to fully understand this variability. Here, we evaluate the abundance and occupancy patterns of hierarchically sampled coral assemblages in three habitats (reef flat, crest, and slope) and five regions (Indonesia, Papua New Guinea, the Solomon Islands, American Samoa, and the Society Islands) across the west-central Pacific Ocean. Specifically, we compare two alternative models that unify spatial variance and occupancy via the negative binomial distribution. The first assumes a power-law scaling between the mean and variance of abundance; the second assumes a quadratic variance-mean relationship and a constant abundance-invariant aggregation parameter. Surprisingly, the well-established power-law model performs worse than the model assuming abundance-invariant aggregation, for both variance-mean and occupancy-abundance relationships. We also find strong evidence for regional and habitat variation in these relationships and in the levels of aggregation estimated by the abundance-invariant aggregation model. Among habitats, corals on reef flats exhibited lower occupancy and higher levels of aggregation compared to reef crests and slopes. Among regions, low occupancy and high aggregation were most pronounced across all habitats in American Samoa. These patterns may be related to habitat and regional differences in disturbance and recovery processes. Our results suggest that the spatial scaling of abundance and occupancy is sensitive to processes operating among these habitats and at regional scales. However, the consistency of these relationships across species within assemblages suggests that a theoretical unification of spatial variance and occupancy patterns is indeed possible.     

Metacommunity structure of pond macroinvertebrates: effects of dispersal mode and generation time

A recent challenge in community ecology is to understand under what conditions local and regional processes may be more important in shaping community structure. We investigated the role of dispersal mode and generation time for communities of macroinvertebrates in two sets of connected ponds during three consecutive years. We found no evidence that generation time affected metacommunity structure, possibly because statistical power was limited because the range of generation times present was small. In contrast, we found that the spatial structure of the macroinvertebrate metacommunities differed with dispersal mode in one of two sets of ponds. Passive dispersers showed positive distance-dissimilarity correlations, suggesting mass effects via the pond connections. These correlations did not stretch beyond the first pond downstream suggesting that, in this chain of connected ponds, intervening ponds effectively buffered dispersal. Active dispersers did not show any consistent spatial pattern, possibly because intensive over-land dispersal homogenized the metacommunity. Thus, although pond connections probably equally promoted dispersal of actively and passively dispersing macroinvertebrates, the implications for the structure of their metacommunities may depend on their dispersal mode. We conclude that dispersal mode has the potential to affect the mechanisms that are integral to metacommunity structure.     

Changes in Species Abundance, Distribution, and Diversity in a Central European Bird Community

We analyzed possible causes of changes in species abundance, range size, and diversity as well as extinctions and colonizations in a central European bird community. Using data from the semiquantitative "Lake Constance" breeding bird atlas, we demonstrated that changes in regional abundances from 1980–1981 to 1990–1992 of 151 coexisting bird species were influenced by breeding habitat and migratory status. Significant declines were found in populations of farmland species and long-distance migrants. Farmland species lost parts of their ranges but hardly changed in local abundance in sites where they still occurred. In contrast, declines in long-distance migrants were caused by significant declines in local abundance with only slight loss of occupied range. Regional extinctions and colonizations were predictable from overall population trends. For example, all species that went extinct were either farmland species or long-distance migrants. Avian community composition was influenced by disproportionate declines of abundant species. This led to declines in the total number of breeding pairs and in community biomass and to increases in community evenness, but to only slight declines in species richness. Future conservation efforts in Europe need to focus more on farmland species and on understanding causes for the declines of long-distance migrants.     

Human Population Density and Prediction of Local Plant Extinction in Britain

Throughout the world, and particularly in densely populated countries like Britain, human activities exert a dominant influence on the abundance of both plants and animals. The commonness and rarity of plants in Britain has been plausibly linked to human land use. In Western Europe the identity of increasing and decreasing plants appears to depend on human population density, which is itself a crude measure of human impact on the landscape. The publication of new data on the changing distributions of scarce British plants allowed us to investigate the relationship between loss of scarce plants and human population density in Britain. Our results confirm that a direct effect of human population density on local plant extinctions can be detected at the regional scale in Britain. Although intensive agriculture is conventionally regarded as the greatest threat to British wildlife, our analysis suggests that urbanization may be at least as significant a danger.     

Quantifying the importance of regional and local filters for community trait structure in tropical and temperate zones

The influence of regional and local processes on community structure is a major focus of ecology. Classically, ecologists have used local-regional richness regressions to evaluate the role of local and regional processes in determining community structure, an approach that has numerous flaws. Here, we implemented a novel trait-based approach that treats local and regional influences as a continuum, rather than a dichotomy. Using hylid frogs (Hylidae), we compared trait dispersion among members of local species assemblages to the trait dispersion in the regional assemblage from which they were drawn. Similarly, we compared trait dispersion in the regional assemblages to dispersion in the continental species pool. We estimated the contributions of local and regional filters, and we compared their strength in temperate and tropical zones. We found that regional and local filters explained 80% of the total variation among local assemblages in community body size dispersion. Overall, regional filters reduced trait dispersion, and local filters increased it, a pattern driven by particularly strong antagonistic effects in temperate zones that reduced the realized total variation by more than 40%. In contrast, local and regional filters acted in concert in tropical regions. Patterns within the tropics did not differ from the random expectation based on a null model, but within the temperate zone, local community filtering was stronger than expected by chance. Furthermore, in temperate regions, antagonistic regional and local filtering masked from 76% to 90% of the total variation in trait dispersion. Together, these results suggest that there are fundamental differences in the scale and identity of the processes determining community structure in temperate and tropical regions.     

Interactions of an Insecticide with Larval Density and Predation in Experimental Amphibian Communities

Abstract: This study examines the effects of the short-lived insecticide carbaryl, a neurotoxin, on amphibian communities experiencing natural stresses of competition and predation. Tadpoles of three species (  Woodhouse's toad [ Bufo woodhousii ], gray treefrog [ Hyla versicolor ], and green frog [ Rana clamitans ]), representing a commonly encountered assemblage in Missouri, were reared in outdoor polyethylene pond mesocosms. We determined the effects of initial tadpole density ( low or high), predation (newts [  Notophthalmus viridescens ] absent or present), chemical exposure (0, 3.5, or 7.0 mg /L carbaryl), and their interactions on body mass, larval period, and survival to metamorphosis. Green frogs in high-density ponds did not reach metamorphosis, but metamorphs in low-density ponds and tadpoles in high-density ponds were not significantly affected by treatments or their interaction. Carbaryl reduced survival to metamorphosis in toads and treefrogs and increased mass at metamorphosis in treefrogs. Effects of carbaryl varied with predator environment and initial larval density. Interactions of carbaryl with predator and with density may result in an indirect effect of carbaryl causing increased food resources through the elimination of zooplankton populations that may compete for similar resources. Our results indicate that differences in biotic conditions influenced the potency of carbaryl and that even low concentrations induce changes that may alter community dynamics in ways not predicted from single-factor, laboratory-based studies.     

Habitat Loss and Changes in the Species-Area Relationship

Abstract: The species-area relationship (SAR) has been used successfully to predict extinction from extent of habitat reduction. These extinction estimates assume that species have uniformly distributed range requirements and a minimum abundance level required for persistence; how many species are lost depends solely on how much habitat is removed, not on where it is removed. We consider another limiting case in which range requirements, rather than abundances, determine extinctions. We used a new method for constructing SARs based on assumptions about geographic ranges of species. Our results show that habitat destruction can change the SAR and consequently the number of species predicted to be lost due to habitat destruction. Our method generates SARs that vary in shape according to the specific distributions of geographic range and occupancy but that have the common feature of being described by a power law with an exponent of <1. When the geographic range of species was included in the SAR, the way habitat was lost became important. Although the SAR before habitat destruction is often used to predict species loss after habitat destruction, assumptions must be clearly stated. To predict the damage caused by habitat loss with our model, it is necessary to know the fraction of aggregated species, the distribution of geographic ranges, the form of habitat destruction, and the sampling protocol. The remaining theoretical challenge is to develop a full theory that links abundance and range.     

Long-term drivers of persistence and colonization dynamics in spatially structured amphibian populations

Many organisms live in networks of local populations connected by dispersing individuals, called spatially structured populations (SSPs), where the long-term persistence of the entire network is determined by the balance between 2 processes acting at the scale of local populations: extinction and colonization. When multiple threats act on an SSP, a comparison of the different factors determining local extinctions and colonizations is essential to plan sound conservation actions. We assessed the drivers of long-term population dynamics of multiple amphibian species at the regional scale. We used dynamic occupancy models within a Bayesian framework to identify the factors determining persistence and colonization of local populations. Because connectivity among patches is fundamental to SSPs dynamics, we considered 2 measures of connectivity acting on each focal patch: incidence of the focal species and incidence of invasive crayfish. We used meta-analysis to summarize the effect of different drivers at the community level. Persistence and colonization of local populations were jointly determined by factors acting at different scales. Persistence probability was positively related to the area and the permanence of wetlands, whereas it was negatively related to occurrence of fish. Colonization probability was highest in semipermanent wetlands and in sites with a high incidence of the focal species in nearby sites, whereas it showed a negative relationship with the incidence of invasive crayfish in the landscape. By analyzing long-term data on amphibian population dynamics, we found a strong effect of some classic features commonly used in SSP studies, such as patch area and focal species incidence. The presence of an invasive non-native species at the landscape scale emerged as one of the strongest drivers of colonization dynamics, suggesting that studies on SSPs should consider different connectivity measures more frequently, such as the incidence of predators, especially when dealing with biological invasions.     

Effects of a larval stage on isolation of populations of the trochid<Emphasis Type="Italic"> Austrocochlea constricta</Emphasis> in tidal ponds

To test the importance of special environments for local genetic subdivision in species with a larval phase, we examined allozyme variation among populations of the intertidal snail Austrocochlea constricta, in 18 tidal ponds in the Houtman Abrolhos Islands, Western Australia. Levels of genetic divergence between pond populations were correlated with those of parallel analyses among adjacent shore populations. However, divergence among the isolated ponds, which lack surface connection to the sea, were generally substantially higher, with an overall F_ST of 0.408, compared with 0.274 among the shore sites. The pond populations had less genetic variation than their shore counterparts, and the reduction of heterozygosity was correlated with the isolation of the pond population, as measured by F_ST. Both the degree of isolation and the reduction of heterozygosity were greater in deeper ponds, where snails can produce a local pool of larvae. In contrast, ponds that dry out frequently are less likely to allow production of local recruits, and these appear to be better connected genetically to adjacent shore populations. These patterns contrast sharply with those previously documented at the same sites for the direct-developing snail Bembicium vittatum, which shows greater isolation in ponds that are often dry. The comparison between the two species shows significant interaction between intrinsic and extrinsic impediments to gene flow, and highlights the importance of characteristics of the life history in determining which circumstances favour isolation of local populations.Electronic Supplementary Material Supplementary material is available in the online version of this article at http://dx.doi.org/10.1007/s00277-005-1553-5.Communicated by G.F. Humphrey, Sydney     

Historical processes constrain patterns in global diatom diversity

There is a long-standing belief that microbial organisms have unlimited dispersal capabilities, are therefore ubiquitous, and show weak or absent latitudinal diversity gradients. In contrast, using a global freshwater diatom data set, we show that latitudinal gradients in local and regional genus richness are present and highly asymmetric between both hemispheres. Patterns in regional richness are explained by the degree of isolation of lake districts, while the number of locally coexisting diatom genera is highly constrained by the size of the regional diatom pool, habitat availability, and the connectivity between habitats within lake districts. At regional to global scales, historical factors explain significantly more of the observed geographic patterns in genus richness than do contemporary environmental conditions. Together, these results stress the importance of dispersal and migration in structuring diatom communities at regional to global scales. Our results are consistent with predictions from the theory of island biogeography and metacommunity concepts and likely underlie the strong provinciality and endemism observed in the relatively isolated diatom floras in the Southern Hemisphere.     

Scale-dependent changes in the importance of larval supply and habitat to abundance of a reef fish

While there is great interest in the degree to which local interactions "scale-up" to predict regional patterns of abundance, few studies in marine systems have simultaneously examined patterns of abundance at both the large scale (tens of kilometers) typical of larval movement and the small scale (meters) typical of post-settlement interactions. We addressed this gap by monitoring larval supply, adult survivorship, and giant kelp (Macrocystis pyrifera, a primary habitat-forming species) abundance for 13 populations of kelp bass (Paralabrax clathratus) spread over approximately 200 km in the Santa Barbara Channel, California, USA. At the small, within-site scale, both recruitment and adult survivorship of kelp bass were density-dependent and positively related to kelp abundance. At the larger, among-site scale, the spatial pattern of adult kelp bass abundance was predicted well by the pattern of kelp bass larval supply, but there was a consistent negative spatial relationship between kelp abundance and kelp bass larval supply despite the positive effects of kelp on kelp bass at the smaller spatial scale. This large-scale negative relationship was likely a product of a channel-wide spatial mismatch between oceanographic conditions that favor kelp survival and those that concentrate and distribute fish larvae. These results generally support the recruit-adult hypothesis: kelp bass populations are limited by recruitment at low recruit densities but by density-dependent competition for food resources and/or predator refuges at high recruit densities. At the same time, spatial variation in kelp abundance produced substantial spatiotemporal heterogeneity in kelp bass demographics, which argues for a multispecies, metacommunity approach to predicting kelp bass dynamics.     

A heuristic approach to predicting water beetle diversity in temporary and fluctuating waters

An understanding of the causal mechanisms and processes that shape macroinvertebrate communities at a local scale has important implications for the management and conservation of freshwater biodiversity. Here we compare the performance of linear and non-linear statistics to explore diversity-environment relationships using data from 76 temporary and fluctuating ponds in two regions of southern England. We focus on aquatic beetle assemblages, which have been shown to be excellent surrogates of wider freshwater macroinvertebrate diversity. Ponds in the region contained a rich coleopteran fauna, totaling 68 species, which provided an excellent model system with which to compare the performance of two non-linear procedures (artificial neural networks—ANNs and generalised additive models—GAMs) and one more traditional linear approach (Multiple linear regression—MLR) to modelling diversity-environment relationships. Of all approaches employed, the best fit was obtained using an ANN model with only four input variables (conductivity, turbidity, magnesium concentration and depth). This model accounted for 82% of the observed variability in Shannon diversity index across ponds. In contrast, the best GAM and MLR models only explained 50% and 14% of this variation, respectively. Contribution profile analysis of conductivity, turbidity, magnesium concentration and depth, obtained from the best fit ANN through a hierarchical cluster analysis, allowed the identification of direct and proxy effects in relation to the environmental variables measured in this study. In each case, distinct clusters of ponds were identified in contribution profile analysis, suggesting that ponds across the two regions fall into a number of discrete groups, whose beetle faunas respond in subtly yet significantly different ways to key environmental variables. Aquatic coleopteran diversity in ponds in the two regions appears to be driven at a local scale by changes in relatively few physicochemical gradients, which are related to diversity in a clearly non-linear manner.     

Modeling Abundance Index Data from Anuran Calling Surveys

Abstract:  Evaluation of anuran populations is commonly based on calling surveys that report categorical abundance index data. I present a statistical model for abundance index data that are observations representing ordered abundance classes (e.g., none, some, many). The proposed model provides a formal treatment of detection probability, factors that affect detection, and variation in abundance. The model can be viewed as a generalization of that proposed by MacKenzie et al. (2002) for estimating site-occupancy rates in that it allows for more than two abundance classes. Because the abundance distribution is characterized by multiple abundance classes, it may be more sensitive to subtle changes in the underlying abundance that may go undetected with simple occupancy estimates under which sites are characterized merely as occupied or not. The method is most immediately applicable to surveys of anurans in which index data related to the intensity of calling activity are collected. I applied the proposed method to calling index data from the green frog (  Rana clamitans ) collected as part of the North American Amphibian Monitoring Program. The best model indicated considerable variation in detectability over time and in response to temperature. The resulting adjusted (for detectability) abundance-state distribution demonstrates the negative bias in abundance state obtained from simplistic summaries of calling index data that disregard these sources of variation in detectability.