A general modeling framework for open wildlife populations based on the Polya tree prior.

Wildlife monitoring for open populations can be performed using a number of different survey methods. Each survey method gives rise to a type of data and, in the last five decades, a large number of associated statistical models have been developed for analyzing these data. Although these models have been parameterized and fitted using different approaches, they have all been designed to either model the pattern with which individuals enter and/or exit the population, or to estimate the population size by accounting for the corresponding observation process, or both. However, existing approaches rely on a predefined model structure and complexity, either by assuming that parameters linked to the entry and exit pattern (EEP) are specific to sampling occasions, or by employing parametric curves to describe the EEP. Instead, we propose a novel Bayesian nonparametric framework for modeling EEPs based on the Polya tree (PT) prior for densities. Our Bayesian nonparametric approach avoids overfitting when inferring EEPs, while simultaneously allowing more flexibility than is possible using parametric curves. Finally, we introduce the replicate PT prior for defining classes of models for these data allowing us to impose constraints on the EEPs, when required. We demonstrate our new approach using capture-recapture, count, and ring-recovery data for two different case studies.

Life span, growth, senescence and island syndrome: Accounting for imperfect detection and continuous growth.

Small vertebrates on islands are expected to attain a larger body size, and a greater survival than their mainland counterparts. Comparative studies have questioned whether lizards exhibit this set of adaptations, referred to as the 'island syndrome'. We collected data on 730 individuals the endemic Lilford's lizard Podarcis lilfordi throughout a 10-year period on a small island of the Balearic archipelago (Spain). We coupled a growth function with a capture-mark-recapture model to simultaneously estimate size- and sex-dependent growth rate and survival. To put our results into a wider context, we conducted a systematic review of growth, life span and age at maturity in different Podarcis species comparing insular and mainland populations. We found a low average growth coefficient (0.56 and 0.41 year-1 for males and females to reach an asymptotic size of 72.3 and 65.6 mm respectively), a high annual survival probability of 0.81 and 0.79 in males and females, and a large variability between individuals in growth parameters. Survival probability decreased with body size in both sexes, indicating a senescence pattern typical of long-lived species or in populations with a low extrinsic mortality. Assuming a constant survival after sexual maturity, at about 2 years old, the average life span was 6.18 years in males and 8.99 in females. The oldest animal was a male last captured at an estimated age of ≥13 years and still alive at the end of the study. Our results agree with the predictions of the 'island syndrome' for survival, life span and growth parameters. A comparative analysis of these values across 29 populations of 16 different species of Podarcis indicated that insular lizards grow slower and live longer than their mainland counterparts. However, our data differed from other island populations of the same species, suggesting that island-specific characteristics play an additional role to isolation. Within this study we developed an analytical approach to study the body size-dependent survival of small reptiles. We discuss its applicability to contrast hypotheses on senescence in different sexes of this species, and provide the code used to integrate the growth and capture-mark-recapture models.

Mesocarnivore community structuring in the presence of Africa's apex predator.

Apex predator reintroductions have proliferated across southern Africa, yet their ecological effects and proposed umbrella benefits of associated management lack empirical evaluations. Despite a rich theory on top-down ecosystem regulation via mesopredator suppression, a knowledge gap exists relating to the influence of lions (Panthera leo) over Africa's diverse mesocarnivore (less than 20 kg) communities. We investigate how geographical variation in mesocarnivore community richness and occupancy across South African reserves is associated with the presence of lions. An interesting duality emerged: lion reserves held more mesocarnivore-rich communities, yet mesocarnivore occupancy rates and evenness-weighted diversity were lower in the presence of lions. Human population density in the reserve surroundings had a similarly ubiquitous negative effect on mesocarnivore occupancy. The positive association between species richness and lion presence corroborated the umbrella species concept but translated into small differences in community size. Distributional contractions of mesocarnivore species within lion reserves, and potentially corresponding numerical reductions, suggest within-community mesopredator suppression by lions, probably as a result of lethal encounters and responses to a landscape of fear. Our findings offer empirical support for the theoretical understanding of processes underpinning carnivore community assembly and are of conservation relevance under current large-predator orientated management and conservation paradigms.

Decomposing fecundity and evaluating demographic influence of multiple broods in a migratory bird.

Relevance of breeding season fecundity as a driver of population dynamics has been highlighted by many studies. Despite that, knowledge about how brood type specific (i.e. first, second or replacement) fecundity affects demography of multiple-brooded species is limited. In fact, estimation of brood type specific fecundity is often challenging due to imperfect detection of nesting attempts. We examined the demographic contribution and the feedback on population density of different components of fecundity, along with other vital rates, in a facultative multiple-brooded migratory bird. We used a novel formulation of a fecundity model that allows incorporating reproductive data for which information on the type of brood was unknown in some cases, and embedded it into an integrated population model (IPM) to obtain consensual estimates of all demographic rates, including brood type specific fecundities, reproductive success probabilities and proportion of breeding pairs that performed a second or replacement brood. We then conducted transient life table response experiments on IPM estimates to account for non-stationary environments. We applied the model to two 20-year datasets collected in a Swiss and a German local population of wrynecks Jynx torquilla. Brood type specific fecundities and temporal patterns of brood type specific probabilities of success, number of successful and unsuccessful first broods, probability of starting a second or a replacement brood and proportion of pairs that performed a second or a replacement brood differed between the two populations. However, changes in immigration rate and apparent survival were the dominant contributors to temporal variation and large sequential changes in realized population growth rates in both populations. In the Swiss population we also found that second brood fecundity declined when population size increased. Our study provides insight into the reproductive processes that affect population dynamics and mediate density-dependent fecundity in a migratory bird. In addition, the analytical approach proposed can be used in other studies of multiple-brooded species to maximize the use of available fecundity data through the estimation of unknown brood types, thus favouring a better understanding of the demographic contribution of brood type specific fecundity.

Assessing the effect of density on population growth when modeling individual encounter data.

The relative role of density-dependent and density-independent variation in vital rates and population size remains largely unsolved. Despite its importance to the theory and application of population ecology, and to conservation biology, quantifying the role and strength of density dependence is particularly challenging. We present a hierarchical formulation of the temporal symmetry approach, also known as the Pradel model, that permits estimation of the strength of density dependence from capture-mark-reencounter data. A measure of relative population size is built in the model and serves to detect density dependence directly on population growth rate. The model is also extended to account for temporal random variability in demographic rates, allowing estimation of the temporal variance of population growth rate unexplained by density dependence. We thus present a model-based approach that enable to test and quantify the effect of density-dependent and density-independent factors affecting population fluctuations in a single modeling framework. More generally, we use this modeling framework along with simulated and empirical data to show the value of including density dependence when modeling individual encounter data without the need for auxiliary data.

Effects of anthropogenic disturbance on primate density at the landscape scale.

Accurate estimations of the abundance of threatened animal populations are required for assessment of species' status and vulnerability and conservation planning. However, density estimation is usually difficult and resource demanding, so researchers often collect data at local scales. However, anthropogenic pressures most often have landscape-level effects, for example, through habitat loss and fragmentation. We applied hierarchical distance sampling (HDS) to transect count data to determine the effect of habitat and anthropogenic factors on the density of 3 arboreal primate species inhabiting 5 distinct tropical forests across a landscape of 19,000 km2 in the Udzungwa Mountains of Tanzania. We developed a novel, multiregion extension of HDS that allowed us to model density and detectability jointly across forests without losing site-specific information. For all species, the effect of anthropogenic disturbance on density was overwhelmingly negative among metapopulations: -0.63 Angolan colobus (Colobus angolensis palliatus) (95% Bayesian CI -1.03 to -0.27), -0.54 Udzungwa red colobus (Procolobus gordonorum) (-0.89 to -0.22), and -0.33 Sykes' monkey (Cercopithecus mitis monoides) (-0.63 to -0.07). Some responses to habitat factors were shared, notably the negative effect of elevation and the positive effect of climber coverage. These results are important for conservation science and practice because: the among-populations negative responses to anthropogenic disturbance provides a foundation for development of conservation plans that hold at the landscape scale, which is a comprehensive and cost-efficient approach; the among-species consistency in responses suggests conservation measures may be generalized at the guild level, which is especially relevant given the functional importance of primates in tropical rainforests; and the greater primate densities in areas at low elevation, which are closer to human settlements, point to specific management recommendations, such as the creation of buffer zones and prioritization of areas for protection.

Efectos de la Perturbación Antropogénica sobre la Densidad de Primates a Escala de Paisaje Resumen Se requieren estimaciones correctas del tamaño de las poblaciones de animales amenazados para evaluar el estado y la vulnerabilidad de las especies y así planear su conservación. Sin embargo, la estimación de la densidad es usualmente complicada y demanda recursos, así que los investigadores con frecuencia recolectan la información a escala local, a pesar de que la presión antropogénica suele tener efectos a nivel del paisaje; por ejemplo, la pérdida del hábitat y la fragmentación. Aplicamos un muestro jerárquico a distancia (HDS, en inglés) para contar datos por transecto y determinar el efecto de los factores del hábitat y los factores antropogénicos sobre la densidad de tres especies de primates arbóreos en un paisaje de 19,000 km2 en las montañas de Udzungwa en Tanzania. Desarrollamos una novedosa extensión multiregión de HDS que nos permitió modelar en conjunto la densidad y la facilidad de detección en los bosques sin perder información específica del sitio. Para todas las especies, el efecto de la perturbación antropogénica sobre la densidad fue abrumadoramente negativa entre las metapoblaciones: −0.63 para el colobo angoleño (Colobus angolensis palliatus) (95% bayesiano CI de −1.03 a −0.27), −0.54 para el colobo rojo de Udzungwa (Procolobus gordonorum) (de −0.89 a −0.22), y −0.33 para el mono de Syke (Cercopithecus mitis monoides) (de −0.63 a −0.07). Los monos compartieron algunas respuestas a los factores de hábitat, notoriamente los efectos negativos de la elevación y el efecto positivo de la cobertura de plantas trepadoras. Estos resultados son importantes para la ciencia y práctica de la conservación ya que: las respuestas negativas a las perturbaciones antropogénicas entre las poblaciones proporcionan los cimientos para el desarrollo de planes de conservación que serán válidos a nivel de paisaje, la cual es una estrategia completa y rentable; la consistencia de las respuestas entre las especies sugiere que las medidas de conservación pueden estar generalizadas a nivel de gremio, lo cual es de suma relevancia dada la importancia funcional de los primates en los bosques tropicales; y la densidad mayor de primates en áreas con elevación baja, las cuales están próximas a los asentamientos humanos, señalan hacia recomendaciones específicas de manejo, como la creación de zonas de amortiguamiento y la priorización de áreas para proteger.

Conspecific and not performance-based attraction on immigrants drives colony growth in a waterbird.

Local recruitment and immigration play an important part in the dynamics and growth of animal populations. However, their estimation and incorporation into open population models is, in most cases, problematic. We studied factors affecting the growth of a recently established colony of Eurasian spoonbill (Platalea leucorodia) and assessed the contribution of local recruits, i.e. birds born in the colony, and immigrants, i.e. birds of unknown origin, to colony growth. We applied an integrated population model that accounts for uncertainty in breeding state assignment and merges population surveys, local fecundity and individual longitudinal data of breeding and non-breeding birds, to estimate demographic rates and the relative role of recruitment and immigration in driving the local dynamics. We also used this analytical framework to assess the degree of support for the 'performance-based' and 'conspecific attraction' hypotheses as possible mechanisms of colony growth. Among the demographic rates, only immigration was positively and significantly correlated with population growth rate. In addition, the number of immigrants settling in the colony was positively correlated with colony size in the previous and current year, but was not correlated with fecundity of the previous year. Our results suggest that the variation in immigration affected colony dynamics and that conspecific attraction likely triggered the relevant role of immigration in the growth of a recently formed waterbird colony, supporting the need of including immigration in population analysis.

Climate-driven vital rates do not always mean climate-driven population.

Current climatic changes have increased the need to forecast population responses to climate variability. A common approach to address this question is through models that project current population state using the functional relationship between demographic rates and climatic variables. We argue that this approach can lead to erroneous conclusions when interpopulation dispersal is not considered. We found that immigration can release the population from climate-driven trajectories even when local vital rates are climate dependent. We illustrated this using individual-based data on a trans-equatorial migratory seabird, the Scopoli's shearwater Calonectris diomedea, in which the variation of vital rates has been associated with large-scale climatic indices. We compared the population annual growth rate λi , estimated using local climate-driven parameters with ρi , a population growth rate directly estimated from individual information and that accounts for immigration. While λi varied as a function of climatic variables, reflecting the climate-dependent parameters, ρi did not, indicating that dispersal decouples the relationship between population growth and climate variables from that between climatic variables and vital rates. Our results suggest caution when assessing demographic effects of climatic variability especially in open populations for very mobile organisms such as fish, marine mammals, bats, or birds. When a population model cannot be validated or it is not detailed enough, ignoring immigration might lead to misleading climate-driven projections.

Evaluating mortality rates with a novel integrated framework for nonmonogamous species.

The conservation of wildlife requires management based on quantitative evidence, and especially for large carnivores, unraveling cause-specific mortalities and understanding their impact on population dynamics is crucial. Acquiring this knowledge is challenging because it is difficult to obtain robust long-term data sets on endangered populations and, usually, data are collected through diverse sampling strategies. Integrated population models (IPMs) offer a way to integrate data generated through different processes. However, IPMs are female-based models that cannot account for mate availability, and this feature limits their applicability to monogamous species only. We extended classical IPMs to a two-sex framework that allows investigation of population dynamics and quantification of cause-specific mortality rates in nonmonogamous species. We illustrated our approach by simultaneously modeling different types of data from a reintroduced, unhunted brown bear (Ursus arctos) population living in an area with a dense human population. In a population mainly driven by adult survival, we estimated that on average 11% of cubs and 61% of adults died from human-related causes. Although the population is currently not at risk, adult survival and thus population dynamics are driven by anthropogenic mortality. Given the recent increase of human-bear conflicts in the area, removal of individuals for management purposes and through poaching may increase, reversing the positive population growth rate. Our approach can be generalized to other species affected by cause-specific mortality and will be useful to inform conservation decisions for other nonmonogamous species, such as most large carnivores, for which data are scarce and diverse and thus data integration is highly desirable.

Seabirds pecking polystyrene items in offshore Adriatic Sea waters.

A number of seabird species have been known to peck, displace, and ingest various plastic items including expanded polystyrene, for reasons that remain largely conjectural. Ingestion of polystyrene parts potentially causes lethal or sublethal effects on birds. Pecking can also result in the damage of polystyrene items, resulting in increased market turnover and environmental build-up, or economic consequences for stakeholders. In January and February, 2022, fishers in a portion of the western Adriatic Sea coast reported pecking damage caused by gulls (Laridae) to polystyrene buoys used to float, signal, and retrieve static fishing nets and traps. We investigated the magnitude of this phenomenon in four fishing harbours of Italy by scoring damage to 470 buoys and interviewing 29 fishers (encompassing 42% of the relevant fleet). Information was complemented by opportunistic observations at sea. Our preliminary assessment suggests that offshore polystyrene pecking increases in winter months, and it occurs sporadically among years. The overall economic damage to the static net fishery appeared generally modest (approximately 3-4 Euro to replace one buoy), with wide variations in the extent of reported damage. We reviewed the hypotheses behind polystyrene pecking, but none of them provide a clear explanation for the observed behaviour. Finally, we discuss potential effects on seabirds and advocate monitoring to investigate causal factors and mitigate damage to seabirds, fisheries, and marine environment.

Immigration as the main driver of population dynamics in a cryptic cetacean.

Empirical evidence about the role and interaction of immigration with local demographic processes in shaping population dynamics is still scarce. This knowledge gap limits our capability to derive a conceptual framework that can be used to inform conservation actions. Populations exposed to nonstationary environment do not converge to a stable stage distribution, implying the need for evaluating the demographic role of both vital rates and stage distribution using appropriate tools. This is particularly important for species with larger generation times like cetaceans. We explored the relative demographic role of vital rates and population structure of a poorly known cetacean, the Mediterranean Cuvier's beaked whale, while accounting for the exposure to nonstationary environments. We performed a retrospective analysis through transient life table response experiments (tLTRE) using demographic rates and population structure of both sexes obtained from an integrated population model. The contribution of immigration to variation in realized population growth rates was 4.2, 7.6, and 12.7 times larger than that of female apparent survival, proportional abundance of breeding females with a 2-year-old calf, and proportional abundance of breeding females with a 3-year-old calf, respectively. Immigration rate and proportional abundance of breeding females with a 2- or 3-year-old calf explained, respectively, 65% and 20% of total temporal variability in realized population growth rates. Changes in realized population growth rate between successive years were mainly driven by changes in immigration and population structure, specifically the proportional abundance of breeding females with a 2-year-old calf. Our study provides insight into the demographic processes that affect population dynamics and in a cryptic cetacean. We presented an analytical approach for maximizing the use of available data through the integration of multiple sources of information for individuals of different distinctiveness levels.

How much biodiversity is concealed in the word 'biodiversity'?

Amidst a global biodiversity crisis1, the word 'biodiversity' has become indispensable for conservation and management2. Yet, biodiversity is often used as a buzzword in scientific literature. Resonant titles of papers claiming to have studied 'global biodiversity' may be used to promote research focused on a few taxonomic groups, habitats, or facets of biodiversity - taxonomic, (phylo)genetic, or functional. This usage may lead to extrapolating results outside the target systems of these studies with direct consequences for our understanding of life on Earth and its practical conservation. Here, we used a random sample of papers with the word 'biodiversity' in their title to take a long view of the use of this term. Despite improvements in analytical tools, monitoring technologies, and data availability3,4, we found that the taxonomic scope of research articles has not increased in recent years. We also show that studies with a wider taxonomic scope attract more citations and online attention. Our results have broad ramifications for understanding how extrapolating from studies with narrow taxonomic scope affects our view of global biodiversity and conservation.

When parks work: Effect of anthropogenic disturbance on occupancy of tropical forest mammals.

Protected areas (PAs) in the tropics are vulnerable to human encroachment, and, despite formal protection, they do not fully mitigate anthropogenic threats to habitats and biodiversity. However, attempts to quantify the effectiveness of PAs and to understand the status and changes of wildlife populations in relation to protection efficiency remain limited. Here, we used camera-trapping data collected over 8 consecutive years (2009-2016) to investigate the yearly occurrences of medium-to-large mammals within the Udzungwa Mountains National Park (Tanzania), an area of outstanding importance for biological endemism and conservation. Specifically, we evaluated the effects of habitat and proxies of human disturbance, namely illegal hunting with snares and firewood collection (a practice that was banned in 2011 in the park), on species' occurrence probabilities. Our results showed variability in species' responses to disturbance: The only species that showed a negative effect of the number of snares found on occurrence probability was the Harvey's duiker, a relatively widespread forest antelope. Similarly, we found a moderate positive effect of the firewood collection ban on only the suni, another common antelope, and a negative effect on a large opportunistic rodent, the giant-pouched rat. Importantly, we found evidence of temporal stability in occurrence probability for all species over the 8-year study period. Our findings suggest that well-managed PAs can sustain mammal populations in tropical forests. However, variability among species in their responses to anthropogenic disturbance necessitates consideration in the design of conservation action plans for multiple taxa.

Reserve size and anthropogenic disturbance affect the density of an African leopard (Panthera pardus) meta-population.

Determining correlates of density for large carnivores is important to understand their ecological requirements and develop conservation strategies. Of several earlier density studies conducted globally, relatively few addressed a scale (usually >1000 km2) that allows inference on correlates of density over heterogeneous landscapes. We deployed 164 camera trap stations covering ~2500 km2 across five areas characterized by broadly different vegetation cover in the Udzungwa Mountains, Tanzania, to investigate correlates of density for a widespread and adaptable carnivore, the leopard (Panthera pardus). We modelled data in a spatially explicit capture-recapture framework, with both biotic and abiotic covariates hypothesised to influence density. We found that leopard density increased with distance to protected area boundary (mean±SE estimated effect = 0.44±0.20), a proxy for both protected area extent and distance from surrounding human settlements. We estimated mean density at 4.22 leopards/100 km2 (85% CI = 3.33‒5.35/100 km2), with no variation across habitat types. Results indicate that protected area extent and anthropogenic disturbance limit leopard populations whereas no support was found for prey availability and trap array as drivers of leopard density. Such vulnerability is relevant to the conservation of the leopard, which is generally considered more resilient to human disturbance than other large cats. Our findings support the notion that protected areas are important to preserve viable population of leopards, increasingly so in times of unprecedented habitat fragmentation. Protection of buffer zones smoothing the abrupt impact of human activities at reserve edges also appears of critical conservation relevance.

Data integration for inference about spatial processes: A model-based approach to test and account for data inconsistency.

Recently-developed methods that integrate multiple data sources arising from the same ecological processes have typically utilized structured data from well-defined sampling protocols (e.g., capture-recapture and telemetry). Despite this new methodological focus, the value of opportunistic data for improving inference about spatial ecological processes is unclear and, perhaps more importantly, no procedures are available to formally test whether parameter estimates are consistent across data sources and whether they are suitable for integration. Using data collected on the reintroduced brown bear population in the Italian Alps, a population of conservation importance, we combined data from three sources: traditional spatial capture-recapture data, telemetry data, and opportunistic data. We developed a fully integrated spatial capture-recapture (SCR) model that included a model-based test for data consistency to first compare model estimates using different combinations of data, and then, by acknowledging data-type differences, evaluate parameter consistency. We demonstrate that opportunistic data lend itself naturally to integration within the SCR framework and highlight the value of opportunistic data for improving inference about space use and population size. This is particularly relevant in studies of rare or elusive species, where the number of spatial encounters is usually small and where additional observations are of high value. In addition, our results highlight the importance of testing and accounting for inconsistencies in spatial information from structured and unstructured data so as to avoid the risk of spurious or averaged estimates of space use and consequently, of population size. Our work supports the use of a single modeling framework to combine spatially-referenced data while also accounting for parameter consistency.

Primates decline rapidly in unprotected forests: evidence from a monitoring program with data constraints.

Growing threats to primates in tropical forests make robust and long-term population abundance assessments increasingly important for conservation. Concomitantly, monitoring becomes particularly relevant in countries with primate habitat. Yet monitoring schemes in these countries often suffer from logistic constraints and/or poor rigor in data collection, and a lack of consideration of sources of bias in analysis. To address the need for feasible monitoring schemes and flexible analytical tools for robust trend estimates, we analyzed data collected by local technicians on abundance of three species of arboreal monkey in the Udzungwa Mountains of Tanzania (two Colobus species and one Cercopithecus), an area of international importance for primate endemism and conservation. We counted primate social groups along eight line transects in two forest blocks in the area, one protected and one unprotected, over a span of 11 years. We applied a recently proposed open metapopulation model to estimate abundance trends while controlling for confounding effects of observer, site, and season. Primate populations were stable in the protected forest, while the colobines, including the endemic Udzungwa red colobus, declined severely in the unprotected forest. Targeted hunting pressure at this second site is the most plausible explanation for the trend observed. The unexplained variability in detection probability among transects was greater than the variability due to observers, indicating consistency in data collection among observers. There were no significant differences in both primate abundance and detectability between wet and dry seasons, supporting the choice of sampling during the dry season only based on minimizing practical constraints. Results show that simple monitoring routines implemented by trained local technicians can effectively detect changes in primate populations in tropical countries. The hierarchical Bayesian model formulation adopted provides a flexible tool to determine temporal trends with full account for any imbalance in the data set and for imperfect detection.

Demographic consequences of poison-related mortality in a threatened bird of prey.

Evidence for decline or threat of wild populations typically comes from multiple sources and methods that allow optimal integration of the available information, representing a major advance in planning management actions. We used integrated population modelling and perturbation analyses to assess the demographic consequences of the illegal use of poison for an insular population of Red Kites, Milvus milvus. We first pooled into a single statistical framework the annual census of breeding pairs, the available individual-based data, the average productivity and the number of birds admitted annually to the local rehabilitation centre. By combining these four types of information we were able to increase estimate precision and to obtain an estimate of the proportion of breeding adults, an important parameter that was not directly measured in the field and that is often difficult to assess. Subsequently, we used perturbation analyses to measure the expected change in the population growth rate due to a change in poison-related mortality. We found that poison accounted for 0.43 to 0.76 of the total mortality, for yearlings and older birds, respectively. Results from the deterministic population model indicated that this mortality suppressed the population growth rate by 20%. Despite this, the population was estimated to increase, albeit slowly. This positive trend was likely maintained by a very high productivity (1.83 fledglings per breeding pair) possibly promoted by supplementary feeding, a situation which is likely to be common to many large obligate or facultative European scavengers. Under this hypothetical scenario of double societal costs (poisoning of a threatened species and feeding programs), increasing poison control would help to lower the public cost of maintaining supplementary feeding stations.