Using unstructured crowd-sourced data to evaluate urban tolerance of terrestrial native animal species within a California Mega-City.

In response to biodiversity loss and biotic community homogenization in urbanized landscapes, there are increasing efforts to conserve and increase biodiversity within urban areas. Accordingly, around the world, previously extirpated species are (re)colonizing and otherwise infiltrating urban landscapes, while other species are disappearing from these landscapes. Tracking the occurrence of traditionally urban intolerant species and loss of traditionally urban tolerant species should be a management goal of urban areas, but we generally lack tools to study this phenomenon. To address this gap, we first used species' occurrences from iNaturalist, a large collaborative dataset of species observations, to calculate an urban association index (UAI) for 967 native animal species that occur in the city of Los Angeles. On average, the occurrence of native species was negatively associated with our composite measure of urban intensity, with the exception of snails and slugs, which instead occur more frequently in areas of increased urban intensity. Next, we assessed 8,348 0.25 x 0.25 mile grids across the City of Los Angeles to determine the average grid-level UAI scores (i.e., a summary of the UAIs present in a grid cell, which we term Community Urban Tolerance Index or CUTI). We found that areas of higher urban intensity host more urban tolerant species, but also that taxonomic groups differ in their aggregate tolerance of urban areas, and that spatial patterns of tolerance vary between groups. The framework established here has been designed to be iteratively reevaluated by city managers of Los Angeles in order to track the progress of initiatives to preserve and encourage urban biodiversity, but can be rescaled to sample different regions within the city or different cities altogether to provide a valuable tool for city managers globally.

A cloudy forecast for species distribution models: Predictive uncertainties abound for California birds after a century of climate and land-use change.

Correlative species distribution models are widely used to quantify past shifts in ranges or communities, and to predict future outcomes under ongoing global change. Practitioners confront a wide range of potentially plausible models for ecological dynamics, but most specific applications only consider a narrow set. Here, we clarify that certain model structures can embed restrictive assumptions about key sources of forecast uncertainty into an analysis. To evaluate forecast uncertainties and our ability to explain community change, we fit and compared 39 candidate multi- or joint species occupancy models to avian incidence data collected at 320 sites across California during the early 20th century and resurveyed a century later. We found massive (>20,000 LOOIC) differences in within-time information criterion across models. Poorer fitting models omitting multivariate random effects predicted less variation in species richness changes and smaller contemporary communities, with considerable variation in predicted spatial patterns in richness changes across models. The top models suggested avian environmental associations changed across time, contemporary avian occupancy was influenced by previous site-specific occupancy states, and that both latent site variables and species associations with these variables also varied over time. Collectively, our results recapitulate that simplified model assumptions not only impact predictive fit but may mask important sources of forecast uncertainty and mischaracterize the current state of system understanding when seeking to describe or project community responses to global change. We recommend that researchers seeking to make long-term forecasts prioritize characterizing forecast uncertainty over seeking to present a single best guess. To do so reliably, we urge practitioners to employ models capable of characterizing the key sources of forecast uncertainty, where predictors, parameters and random effects may vary over time or further interact with previous occurrence states.

Demographic consequences of phenological asynchrony for North American songbirds.

Changes in phenology in response to ongoing climate change have been observed in numerous taxa around the world. Differing rates of phenological shifts across trophic levels have led to concerns that ecological interactions may become increasingly decoupled in time, with potential negative consequences for populations. Despite widespread evidence of phenological change and a broad body of supporting theory, large-scale multitaxa evidence for demographic consequences of phenological asynchrony remains elusive. Using data from a continental-scale bird-banding program, we assess the impact of phenological dynamics on avian breeding productivity in 41 species of migratory and resident North American birds breeding in and around forested areas. We find strong evidence for a phenological optimum where breeding productivity decreases in years with both particularly early or late phenology and when breeding occurs early or late relative to local vegetation phenology. Moreover, we demonstrate that landbird breeding phenology did not keep pace with shifts in the timing of vegetation green-up over a recent 18-y period, even though avian breeding phenology has tracked green-up with greater sensitivity than arrival for migratory species. Species whose breeding phenology more closely tracked green-up tend to migrate shorter distances (or are resident over the entire year) and breed earlier in the season. These results showcase the broadest-scale evidence yet of the demographic impacts of phenological change. Future climate change-associated phenological shifts will likely result in a decrease in breeding productivity for most species, given that bird breeding phenology is failing to keep pace with climate change.

Traits shaping urban tolerance in birds differ around the world.

As human density increases, biodiversity must increasingly co-exist with urbanization or face local extinction. Tolerance of urban areas has been linked to numerous functional traits, yet few globally consistent patterns have emerged to explain variation in urban tolerance, which stymies attempts at a generalizable predictive framework. Here, we calculate an Urban Association Index (UAI) for 3,768 bird species in 137 cities across all permanently inhabited continents. We then assess how this UAI varies as a function of ten species-specific traits and further test whether the strength of trait relationships vary as a function of three city-specific variables. Of the ten species traits, nine were significantly associated with urban tolerance. Urban-associated species tend to be smaller, less territorial, have greater dispersal ability, broader dietary and habitat niches, larger clutch sizes, greater longevity, and lower elevational limits. Only bill shape showed no global association with urban tolerance. Additionally, the strength of several trait relationships varied across cities as a function of latitude and/or human population density. For example, the associations of body mass and diet breadth were more pronounced at higher latitudes, while the associations of territoriality and longevity were reduced in cities with higher population density. Thus, the importance of trait filters in birds varies predictably across cities, indicating biogeographic variation in selection for urban tolerance that could explain prior challenges in the search for global patterns. A globally informed framework that predicts urban tolerance will be integral to conservation as increasing proportions of the world's biodiversity are impacted by urbanization.

Multi-trophic occupancy modeling connects temporal dynamics of woodpeckers and beetle sign following fire.

In conifer forests of western North America, wildlife populations can change rapidly in the decade following wildfire as trees die and animals respond to concomitant resource pulses that occur across multiple trophic levels. In particular, black-backed woodpeckers (Picoides arcticus) show predictable temporal increases then declines following fire; this trajectory is widely believed to be a response to the woodpeckers' main prey, woodboring beetle larvae of the families Buprestidae and Cerambycidae, but we lack understanding of how abundances of these predators and prey may be associated in time or space. Here, we pair woodpecker surveys over 10 years with surveys of woodboring beetle sign and activity, collected at 128 survey plots across 22 recent fires, to ask whether accumulated beetle sign indicates current or past black-backed woodpecker occurrence, and whether that relationship is mediated by the number of years since fire. We test this relationship using an integrative multi-trophic occupancy model. Our results demonstrate that woodboring beetle sign is a positive indicator of woodpecker presence 1-3 years following fire, an uninformative indicator from 4-6 years after fire, and a negative indicator beginning 7 years following fire. Woodboring beetle activity, itself, is temporally variable and dependent on tree species composition, with beetle sign generally accumulating over time, particularly in stands with diverse tree communities, but decreasing over time in Pinus-dominated stands where faster bark decay rates lead to brief pulses of beetle activity followed by rapid degradation of tree substrate and accumulated beetle sign. Altogether, the strong connections of woodpecker occurrence to beetle activity support prior hypotheses of how multi-trophic interactions govern rapid temporal dynamics of primary and secondary consumers in burned forests. While our results indicate that beetle sign is, at best, a rapidly shifting and potentially misleading measure of woodpecker occurrence, the better we understand the interacting mechanisms underlying temporally dynamic systems, the more successfully we will be able to predict the outcomes of management actions.

Incorporating pyrodiversity into wildlife habitat assessments for rapid post-fire management: A woodpecker case study.

Spatial and temporal variation in fire characteristics-termed pyrodiversity-are increasingly recognized as important factors that structure wildlife communities in fire-prone ecosystems, yet there have been few attempts to incorporate pyrodiversity or post-fire habitat dynamics into predictive models of animal distributions and abundance to support post-fire management. We use the black-backed woodpecker-a species associated with burned forests-as a case study to demonstrate a pathway for incorporating pyrodiversity into wildlife habitat assessments for adaptive management. Employing monitoring data (2009-2019) from post-fire forests in California, we developed three competing occupancy models describing different hypotheses for habitat associations: (1) a static model representing an existing management tool, (2) a temporal model accounting for years since fire, and (3) a temporal-landscape model which additionally incorporates emerging evidence from field studies about the influence of pyrodiversity. Evaluating predictive ability, we found superior support for the temporal-landscape model, which showed a positive relationship between occupancy and pyrodiversity and interactions between habitat associations and years since fire. We incorporated the new temporal-landscape model into an RShiny application to make this decision-support tool accessible to decision-makers.

The effect of insect food availability on songbird reproductive success and chick body condition: Evidence from a systematic review and meta-analysis.

Reports of declines in abundance and biomass of insects and other invertebrates from around the world have raised concerns about food limitation that could have profound impacts for insectivorous species. Food availability can clearly affect species; however, there is considerable variation among studies in whether this effect is evident, and thus a lack of clarity over the generality of the relationship. To understand how decreased food availability due to invertebrate declines will affect bird populations, we conducted a systematic review and used meta-analytic structural equation modelling, which allowed us to treat our core variables of interest as latent variables estimated by the diverse ways in which researchers measure fecundity and chick body condition. We found a moderate positive effect of food availability on chick body condition and a strong positive effect on reproductive success. We also found a negative relationship between chick body condition and reproductive success. Our results demonstrate that food is generally a limiting factor for breeding songbirds. Our analysis also provides evidence for a consistent trade-off between chick body condition and reproductive success, demonstrating the complexity of trophic dynamics important for these vital rates.

Geomagnetic disturbance associated with increased vagrancy in migratory landbirds.

Rare birds known as "accidentals" or "vagrants" have long captivated birdwatchers and puzzled biologists, but the drivers of these rare occurrences remain elusive. Errors in orientation or navigation are considered one potential driver: migratory birds use the Earth's magnetic field-sensed using specialized magnetoreceptor structures-to traverse long distances over often unfamiliar terrain. Disruption to these magnetoreceptors or to the magnetic field itself could potentially cause errors leading to vagrancy. Using data from 2 million captures of 152 landbird species in North America over 60 years, we demonstrate a strong association between disruption to the Earth's magnetic field and avian vagrancy during fall migration. Furthermore, we find that increased solar activity-a disruptor of the avian magnetoreceptor-generally counteracts this effect, potentially mitigating misorientation by disabling the ability for birds to use the magnetic field to orient. Our results link a hypothesized cause of misorientation to the phenomenon of avian vagrancy, further demonstrating the importance of magnetoreception among the orientation mechanisms of migratory birds. Geomagnetic disturbance may have important downstream ecological consequences, as vagrants may experience increased mortality rates or facilitate range expansions of avian populations and the organisms they disperse.

Adult male birds advance spring migratory phenology faster than females and juveniles across North America.

Advances in spring migratory phenology comprise some of the most well-documented evidence for the impacts of climate change on birds. Nevertheless, surprisingly little research has investigated whether birds are shifting their migratory phenology equally across sex and age classes-a question critical to understanding the potential for trophic mismatch. We used 60 years of bird banding data across North America-comprising over 4 million captures in total-to investigate both spring and fall migratory phenology for a total of 98 bird species across sex and age classes, with the exact numbers of species for each analysis depending on season-specific data availability. Consistent with protandry, in spring (n = 89 species), adult males were the first to arrive and immature females were the last to arrive. In fall (n = 98), there was little difference between sexes, but adults tended to depart earlier than juveniles. Over 60 years, adult males advanced their phenology the fastest (-0.84 days per decade, 95 CrI = -1.22 to -0.47, n = 36), while adult and immature females advanced at a slower pace, causing the gap in male and female arrival times to widen over time. In the fall, there was no overall trend in phenology by age or sex (n = 57), driven in part by high interspecific variation related to breeding and molt strategies. Our results indicate consistent and predictable age- and sex-based differences in the rates at which species' springtime phenology is shifting. The growing gap between male and female migratory arrival indicates sex-based plasticity in adaptation to climate change that has strong potential to negatively impact current and future population trends.

Abiotic conditions shape spatial and temporal morphological variation in North American birds.

Quantifying environment-morphology relationships is important not only for understanding the fundamental processes driving phenotypic diversity within and among species but also for predicting how species will respond to ongoing global change. Despite a clear set of expectations motivated by ecological theory, broad evidence in support of generalizable effects of abiotic conditions on spatial and temporal intraspecific morphological variation has been limited. Using standardized data from >250,000 captures of 105 landbird species, we assessed intraspecific shifts in the morphology of adult male birds since 1989 while simultaneously measuring spatial morphological gradients across the North American continent. We found strong spatial and temporal trends in average body size, with warmer temperatures associated with smaller body sizes both at more equatorial latitudes and in more recent years. The magnitude of these thermal effects varied both across and within species, with results suggesting it is the warmest, rather than the coldest, temperatures that drive both spatial and temporal trends. Stronger responses to spatial-rather than temporal-variation in temperature suggest that morphological change may not be keeping up with the pace of climate change. Additionally, as elevation increases, we found that body size declines as relative wing length increases, probably due to the benefits that longer wings confer for flight in thin air environments. Our results provide support for both existing and new large-scale ecomorphological 'rules' and highlight how the response of functional trade-offs to abiotic variation drives morphological change.

Decadal-scale phenology and seasonal climate drivers of migratory baleen whales in a rapidly warming marine ecosystem.

Species' response to rapid climate change can be measured through shifts in timing of recurring biological events, known as phenology. The Gulf of Maine is one of the most rapidly warming regions of the ocean, and thus an ideal system to study phenological and biological responses to climate change. A better understanding of climate-induced changes in phenology is needed to effectively and adaptively manage human-wildlife conflicts. Using data from a 20+ year marine mammal observation program, we tested the hypothesis that the phenology of large whale habitat use in Cape Cod Bay has changed and is related to regional-scale shifts in the thermal onset of spring. We used a multi-season occupancy model to measure phenological shifts and evaluate trends in the date of peak habitat use for North Atlantic right (Eubalaena glacialis), humpback (Megaptera novaeangliae), and fin (Balaenoptera physalus) whales. The date of peak habitat use shifted by +18.1 days (0.90 days/year) for right whales and +19.1 days (0.96 days/year) for humpback whales. We then evaluated interannual variability in peak habitat use relative to thermal spring transition dates (STD), and hypothesized that right whales, as planktivorous specialist feeders, would exhibit a stronger response to thermal phenology than fin and humpback whales, which are more generalist piscivorous feeders. There was a significant negative effect of western region STD on right whale habitat use, and a significant positive effect of eastern region STD on fin whale habitat use indicating differential responses to spatial seasonal conditions. Protections for threatened and endangered whales have been designed to align with expected phenology of habitat use. Our results show that whales are becoming mismatched with static seasonal management measures through shifts in their timing of habitat use, and they suggest that effective management strategies may need to alter protections as species adapt to climate change.

Anna's hummingbird (Calypte anna) physiological response to novel thermal and hypoxic conditions at high elevations.

Many species have not tracked their thermal niches upslope as predicted by climate change, potentially because higher elevations are associated with abiotic challenges beyond temperature. To better predict whether organisms can continue to move upslope with rising temperatures, we need to understand their physiological performance when subjected to novel high-elevation conditions. Here, we captured Anna's hummingbirds - a species expanding their elevational distribution in concordance with rising temperatures - from across their current elevational distribution and tested their physiological response to novel abiotic conditions. First, at a central aviary within their current elevational range, we measured hovering metabolic rate to assess their response to oxygen conditions and torpor use to assess their response to thermal conditions. Second, we transported the hummingbirds to a location 1200 m above their current elevational range limit to test for an acute response to novel oxygen and thermal conditions. Hummingbirds exhibited lower hovering metabolic rates above their current elevational range limit, suggesting lower oxygen availability may reduce performance after an acute exposure. Alternatively, hummingbirds showed a facultative response to thermal conditions by using torpor more frequently and for longer. Finally, post-experimental dissection found that hummingbirds originating from higher elevations within their range had larger hearts, a potential plastic response to hypoxic environments. Overall, our results suggest lower oxygen availability and low air pressure may be difficult challenges to overcome for hummingbirds shifting upslope as a consequence of rising temperatures, especially if there is little to no long-term acclimatization. Future studies should investigate how chronic exposure and acclimatization to novel conditions, as opposed to acute experiments, may result in alternative outcomes that help organisms better respond to abiotic challenges associated with climate-induced range shifts.

Extinction of biotic interactions due to habitat loss could accelerate the current biodiversity crisis.

Habitat loss disrupts species interactions through local extinctions, potentially orphaning species that depend on interacting partners, via mutualisms or commensalisms, and increasing secondary extinction risk. Orphaned species may become functionally or secondarily extinct, increasing the severity of the current biodiversity crisis. While habitat destruction is a major cause of biodiversity loss, the number of secondary extinctions is largely unknown. We investigate the relationship between habitat loss, orphaned species, and bipartite network properties. Using a real seed dispersal network, we simulate habitat loss to estimate the rate at which species are orphaned. To be able to draw general conclusions, we also simulate habitat loss in synthetic networks to quantify how changes in network properties affect orphan rates across broader parameter space. Both real and synthetic network simulations show that even small amounts of habitat loss can cause up to 10% of species to be orphaned. More area loss, less connected networks, and a greater disparity in the species richness of the network's trophic levels generally result in more orphaned species. As habitat is lost to land-use conversion and climate change, more orphaned species increase the loss of community-level and ecosystem functions. However, the potential severity of repercussions ranges from minimal (no species orphaned) to catastrophic (up to 60% of species within a network orphaned). Severity of repercussions also depends on how much the interaction richness and intactness of the community affects the degree of redundancy within networks. Orphaned species could add substantially to the loss of ecosystem function and secondary extinction worldwide.

Woody encroachment happens via intensification, not extensification, of species ranges in an African savanna.

Widespread woody encroachment is a prominent concern for savanna systems as it is often accompanied by losses in productivity and biodiversity. Extensive ecosystem-level work has advanced our understanding of its causes and consequences. However, there is still debate over whether local management can override regional and global drivers of woody encroachment, and it remains largely unknown how encroachment influences woody community assemblages. Here, we examined species-level changes in woody plant distributions and size structure from the late 1980s to the late 2000s based on spatially intensive ground-based surveys across Kruger National Park, South Africa. This study region spans broad gradients in rainfall, soil texture, fire frequency, elephant density, and other topographic variables. Species-level changes in frequency of occurrence and size class proportion reflected widespread woody encroachment primarily by Dichrostachys cinerea and Combretum apiculatum, and a loss of large trees mostly of Sclerocarya birrea and Acacia nigrescens. Environmental variables determining woody species distributions across Kruger varied among species but did not change substantially between two sampling times, indicating that woody encroachers were thickening within their existing ranges. Overall, more areas across Kruger were found to have an increased number of common woody species through time, which indicated an increase in stem density. These areas were generally associated with decreasing fire frequency and rainfall but increasing elephant density. Our results suggest that woody encroachment is a widespread but highly variable trend across landscapes in Kruger National Park and potentially reflects an erosion of local heterogeneity in woody community assemblages. Many savanna managers, including in Kruger, aim to manage for heterogeneity in order to promote biodiversity, where homogenization of vegetation structure counters this specific goal. Increasing fire frequency has some potential as a local intervention. However, many common species increased in commonness even under near-constant disturbance conditions, which likely limits the potential for managing woody encroachment in the face of drivers beyond the scope of local control. Regular field sampling coupled with targeted fire management will enable more accurate monitoring of the rate of encroachment intensification.

Migratory strategy drives species-level variation in bird sensitivity to vegetation green-up.

Animals and plants are shifting the timing of key life events in response to climate change, yet despite recent documentation of escalating phenological change, scientists lack a full understanding of how and why phenological responses vary across space and among species. Here, we used over 7 million community-contributed bird observations to derive species-specific, spatially explicit estimates of annual spring migration phenology for 56 bird species across eastern North America. We show that changes in the spring arrival of migratory birds are coarsely synchronized with fluctuations in vegetation green-up and that the sensitivity of birds to plant phenology varied extensively. Bird arrival responded more synchronously with vegetation green-up at higher latitudes, where phenological shifts over time are also greater. Critically, species' migratory traits explained variation in sensitivity to green-up, with species that migrate more slowly, arrive earlier and overwinter further north showing greater responsiveness to earlier springs. Identifying how and why species vary in their ability to shift phenological events is fundamental to predicting species' vulnerability to climate change. Such variation in sensitivity across taxa, with long-distance neotropical migrants exhibiting reduced synchrony, may help to explain substantial declines in these species over the last several decades.

Juvenile survival of a burned forest specialist in response to variation in fire characteristics.

Pyrodiversity, defined as variation in fire history and characteristics, has been shown to catalyse post-fire biodiversity in a variety of systems. However, the demographic and behavioural mechanisms driving the responses of individual species to pyrodiversity remain largely unexplored. We used a model post-fire specialist, the black-backed woodpecker (Picoides arcticus), to examine the relationship between fire characteristics and juvenile survival while controlling for confounding factors. We radio-tracked fledgling black-backed woodpeckers in burned forests of California and Washington, USA, and derived information on habitat characteristics using ground surveys and satellite data. We used hierarchical Bayesian mixed-effects models to determine the factors that influence both fledgling and annual juvenile survival, and we tested for effects of fledgling age on movement rates. Burn severity strongly affected fledgling survival, with lower survival in patches created by high-severity fire compared to patches burned at medium to low severity or left unburned. Time since leaving the nest was also a strong predictor of fledgling survival, annual juvenile survival and fledgling movement rates. Our results support the role of habitat complementation in generating species-specific benefits from variation in spatial fire characteristics-one axis of pyrodiversity-and highlight the importance of this variation under shifting fire regimes. High-severity fire provides foraging and nesting sites that support the needs of adult black-backed woodpeckers, but fledgling survival is greater in areas burned at lower severity. By linking breeding and foraging habitat with neighbouring areas of reduced predation risk, pyrodiversity may enhance the survival and persistence of animals that thrive in post-fire habitat.

Nests in the cities: adaptive and non-adaptive phenotypic plasticity and convergence in an urban bird.

Phenotypic plasticity plays a critical role in adaptation to novel environments. Behavioural plasticity enables more rapid responses to unfamiliar conditions than evolution by natural selection. Urban ecosystems are one such novel environment in which behavioural plasticity has been documented. However, whether such plasticity is adaptive, and if plasticity is convergent among urban populations, is poorly understood. We studied the nesting biology of an 'urban-adapter' species, the dark-eyed junco (Junco hyemalis), to understand the role of plasticity in adapting to city life. We examined (i) whether novel nesting behaviours are adaptive, (ii) whether pairs modify nest characteristics in response to prior outcomes, and (iii) whether two urban populations exhibit similar nesting behaviour. We monitored 170 junco nests in urban Los Angeles and compared our results with prior research on 579 nests from urban San Diego. We found that nests placed in ecologically novel locations (off-ground and on artificial surfaces) increased fitness, and that pairs practiced informed re-nesting in site selection. The Los Angeles population more frequently nested off-ground than the San Diego population and exhibited a higher success rate. Our findings suggest that plasticity facilitates adaptation to urban environments, and that the drivers behind novel nesting behaviours are complex and multifaceted.

Fire and biodiversity in the Anthropocene.

Fire has been a source of global biodiversity for millions of years. However, interactions with anthropogenic drivers such as climate change, land use, and invasive species are changing the nature of fire activity and its impacts. We review how such changes are threatening species with extinction and transforming terrestrial ecosystems. Conservation of Earth's biological diversity will be achieved only by recognizing and responding to the critical role of fire. In the Anthropocene, this requires that conservation planning explicitly includes the combined effects of human activities and fire regimes. Improved forecasts for biodiversity must also integrate the connections among people, fire, and ecosystems. Such integration provides an opportunity for new actions that could revolutionize how society sustains biodiversity in a time of changing fire activity.

The importance of accounting for imperfect detection when estimating functional and phylogenetic community structure.

Incorporating imperfect detection when estimating species richness has become commonplace in the past decade. However, the question of how imperfect detection of species affects estimates of functional and phylogenetic community structure remains untested. We used long-term counts of breeding bird species that were detected at least once on islands in a land-bridge island system, and employed multi-species occupancy models to assess the effects of imperfect detection of species on estimates of bird diversity and community structure by incorporating species traits and phylogenies. Our results showed that taxonomic, functional, and phylogenetic diversity were all underestimated significantly as a result of species' imperfect detection, with taxonomic diversity showing the greatest bias. The functional and phylogenetic structure calculated from observed communities were both more clustered than those from the detection-corrected communities due to missed distinct species. The discrepancy between observed and estimated diversity differed according to the measure of biodiversity employed. Our study demonstrates the importance of accounting for species' imperfect detection in biodiversity studies, especially for functional and phylogenetic community ecology, and when attempting to infer community assembly processes. With datasets that allow for detection-corrected community structure, we can better estimate diversity and infer the underlying mechanisms that structure community assembly, and thus make reliable management decisions for the conservation of biodiversity.