Capturing the dynamics of small populations: A retrospective assessment using long-term data for an island reintroduction.

The art of population modelling is to incorporate factors essential for capturing a population's dynamics while otherwise keeping the model as simple as possible. However, it is unclear how optimal model complexity should be assessed, and whether this optimal complexity has been affected by recent advances in modelling methodology. This issue is particularly relevant to small populations because they are subject to complex dynamics but inferences about those dynamics are often constrained by small sample sizes. We fitted Bayesian hierarchical models to long-term data on vital rates (survival and reproduction) for the toutouwai Petroica longipes population reintroduced to Tiritiri Matangi, a 220-ha New Zealand island, and quantified the performance of those models in terms of their likelihood of replicating the observed population dynamics. These dynamics consisted of overall growth from 33 (±0.3) to 160 (±6) birds from 1992-2018, including recoveries following five harvest events for further reintroductions to other sites. We initially included all factors found to affect vital rates, which included inbreeding, post-release effects (PRE), density-dependence, sex, age and random annual variation, then progressively removed these factors. We also compared performance of models where data analysis and simulations were done simultaneously to those produced with the traditional two-step approach, where vital rates are estimated first then fed into a separate simulation model. Parametric uncertainty and demographic stochasticity were incorporated in all projections. The essential factors for replicating the population's dynamics were density-dependence in juvenile survival and PRE, i.e. initial depression of survival and reproduction in translocated birds. Inclusion of other factors reduced the precision of projections, and therefore the likelihood of matching observed dynamics. However, this reduction was modest when the modelling was done in an integrated framework. In contrast, projections were much less precise when done with a two-step modelling approach, and the cost of additional parameters was much higher under the two-step approach. These results suggest that minimization of complexity may be less important than accounting for covariances in parameter estimates, which is facilitated by integrating data analysis and population projections using Bayesian methods.

Using long-term data for a reintroduced population to empirically estimate future consequences of inbreeding.

Inbreeding depression is an important long-term threat to reintroduced populations. However, the strength of inbreeding depression is difficult to estimate in wild populations because pedigree data are inevitably incomplete and because good data are needed on survival and reproduction. Predicting future population consequences is especially difficult because this also requires projecting future inbreeding levels and their impacts on long-term population dynamics, which are subject to many uncertainties. We illustrate how such projections can be derived through Bayesian state-space modeling methods based on a 26-year data set for North Island Robins (Petroica longipes) reintroduced to Tiritiri Matangi Island in 1992. We used pedigree data to model increases in the average inbreeding level (F) over time based on kinship of possible breeding pairs and to estimate empirically Ne /N (effective/census population size). We used multiple imputation to model the unknown components of inbreeding coefficients, which allowed us to estimate effects of inbreeding on survival for all 1458 birds in the data set while modeling density dependence and environmental stochasticity. This modeling indicated that inbreeding reduced juvenile survival (1.83 lethal equivalents [SE 0.81]) and may have reduced subsequent adult survival (0.44 lethal equivalents [0.81]) but had no apparent effect on numbers of fledglings produced. Average inbreeding level increased to 0.10 (SE 0.001) as the population grew from 33 (0.3) to 160 (6) individuals over the 25 years, giving a Ne/N ratio of 0.56 (0.01). Based on a model that also incorporated habitat regeneration, the population was projected to reach a maximum of 331-1144 birds (median 726) in 2130, then to begin a slow decline. Without inbreeding, the population would be expected stabilize at 887-1465 birds (median 1131). Such analysis, therefore, makes it possible to empirically derive the information needed for rational decisions about inbreeding management while accounting for multiple sources of uncertainty.

Uso de Datos a Largo Plazo de una Población Reintroducida para Estimar Empíricamente las Consecuencias Futuras de la Endogamia Resumen La depresión endogámica es una amenaza importante a largo plazo para las poblaciones reintroducidas. Sin embargo, es complicado estimar la fuerza de la depresión endogámica en las poblaciones silvestres porque los datos sobre el linaje sin duda estarán incompletos y porque se necesitan datos sólidos sobre la supervivencia y la reproducción. Es especialmente difícil predecir las consecuencias poblacionales a futuro pues esto requiere proyectar a futuro los niveles de endogamia y sus impactos sobre las dinámicas poblacionales a largo plazo, las cuales están sujetas a muchas incertidumbres. Ilustramos cómo dichas proyecciones pueden derivarse mediante métodos de modelado bayesiano de estado-espacio basados en un conjunto de datos obtenidos durante 26 años para los tordos de la Isla del Norte (Petroica longipes) reintroducidos a la isla Tiritiri Matangi en 1992. Usamos datos de linaje para modelar los incrementos en el nivel promedio de endogamia ( F̲ ) a lo largo del tiempo con base en el parentesco de las posibles parejas reproductoras y para estimar empíricamente Ne/N (tamaño poblacional efectivo/por censo). Usamos una imputación múltiple para modelar los componentes desconocidos de los coeficientes de endogamia, lo que nos permitió estimar los efectos de la endogamia sobre la supervivencia para todas las aves (1458) incluidas en el conjunto de datos a la vez que modelamos la dependencia de la densidad y la estocasticidad ambiental. Este modelado indicó que la endogamia redujo la supervivencia juvenil (1.83 equivalentes letales [SE 0.81]) y podría haber reducido la subsecuente supervivencia adulta (0.44 equivalentes letales [0.81]) pero no tuvo un efecto aparente sobre los números de polluelos producidos. El nivel promedio de endogamia incrementó a 0.10 (SE 0.001) conforme la población creció de 33 (0.3) a 160 (6) individuos a lo largo de los 25 años, lo que resultó en una proporción Ne/N de 0.56 (0.01). Con base en un modelo que también incorporó la regeneración del hábitat, se proyectó que la población alcanzaría un máximo de 331-1144 aves (mediana: 726) para 2130 y después comenzaría una lenta disminución. Sin la endogamia, se esperaría que la población se estabilizaría con 887-1465 (mediana: 1131) aves. Por lo tanto, dicho análisis hace posible la derivación empírica de la información necesaria para las decisiones racionales sobre el manejo de la endogamia a la vez que considera a varias fuentes de incertidumbre.

A modelling framework for integrating reproduction, survival and count data when projecting the fates of threatened populations.

A key goal of ecological research is to obtain reliable estimates of population demographic rates, abundance and trends. However, a common challenge when studying wildlife populations is imperfect detection or breeding observation, which results in unknown survival status and reproductive output for some individuals. It is important to account for undetected individuals in population models because they contribute to population abundance and dynamics, and can have implications for population management. Promisingly, recent methodological advances provide us with the tools to integrate data from multiple independent sources to gain insights into the unobserved component of populations. We use data from five reintroduced populations of a threatened New Zealand bird, the hihi (Notiomystis cincta), to develop an integrated population modelling framework that allows missing values for survival status, sex and reproductive output to be modelled. Our approach combines parallel matrices of encounter and reproduction histories from marked individuals, as well as counts of unmarked recruits detected at the start of each breeding season. Integrating these multiple data types enabled us to simultaneously model survival and reproduction of detected individuals, undetected individuals and unknown (never detected) individuals to derive parameter estimates and projections based on all available data, thereby improving our understanding of population dynamics and enabling full propagation of uncertainty. The methods presented will be especially useful for management programmes for populations that are intensively monitored but where individuals are still imperfectly detected, as will be the case for most threatened wild populations.

Predicting reintroduction outcomes for highly vulnerable species that do not currently coexist with their key threats.

Predicting reintroduction outcomes before populations are released is inherently challenging. It becomes even more difficult when the species being considered for reintroduction no longer coexists with the key threats limiting its distribution. However, data from other species facing the same threats can be used to make predictions under these circumstances. We used an integrated Bayesian modeling approach to predict growth of a reintroduced population at a range of predator densities when no data are available for the species in the presence of that predator. North Island Saddlebacks (Philesturnus rufusater) were extirpated from mainland New Zealand by exotic mammalian predators, particularly ship rats (black rats [Rattus rattus]) but are now being considered for reintroduction to sites with intensive predator control. We initially modeled data from previous saddleback reintroductions to predator-free sites to predict population growth at a new predator-free site while accounting for random variation in vital rates among sites. We then predicted population growth at different rat-tracking rates (an index of rat density) by incorporating a previously modeled relationship between rat-tracking and vital rates of another predator-sensitive species, the North Island Robin (Petroica longipes), and accounted for greater vulnerability of saddlebacks to rat predation based on information on historical declines of both species. The results allowed population growth to be predicted as a function of management effort while accounting for uncertainty, which means formal decision analysis could be used to decide whether to proceed with a reintroduction. Similar approaches could be applied to other situations where data on the species of interest are limited and provide an alternative to decision making based solely on expert judgment.

Traits influencing range contraction in New Zealand's endemic forest birds.

Understanding vulnerability of endemic taxa to predation is clearly important for conservation management. In New Zealand, predation by introduced mammals such as rats and mustelids is widely recognized as the primary factor responsible for declines of indigenous fauna. The aim of our study was to evaluate the vulnerability of New Zealand's surviving endemic forest bird species to impacts of introduced mammalian predators, and identify key life history attributes underlying this vulnerability. We measured range contraction following the introduction of exotic mammalian predators for 23 endemic forest bird species using information on both pre-human and current distributions. We used Bayesian modeling techniques to analyze whether variation in range contraction was associated with life history traits potentially influencing species' predation vulnerability, while accounting for phylogenetic relatedness. Our results showed that the extent of range contraction varied greatly among species, with some species remaining in available forest habitat throughout most of their pre-human range, and others having disappeared completely from the main islands. Cavity nesting was the key trait associated with more extensive range decline, suggesting that cavity-nesting species are more vulnerable to predation than species that nest in more open sites.

An integrated approach for predicting fates of reintroductions with demographic data from multiple populations.

We devised a novel approach to model reintroduced populations whereby demographic data collected from multiple sites are integrated into a Bayesian hierarchical model. Integrating data from multiple reintroductions allows more precise population-growth projections to be made, especially for populations for which data are sparse, and allows projections that account for random site-to-site variation to be made before new reintroductions are attempted. We used data from reintroductions of the North Island Robin (Petroica longipes), an endemic New Zealand passerine, to 10 sites where non-native mammalian predators are controlled. A comparison of candidate models that we based on deviance information criterion showed that rat-tracking rate (an index of rat density) was a useful predictor of robin fecundity and adult female survival, that landscape connectivity and a binary measure of whether sites were on a peninsula were useful predictors of apparent juvenile survival (probably due to differential dispersal away from reintroduction sites), and that there was unexplained random variation among sites in all demographic rates. We used the two best supported models to estimate the finite rate of increase (λ) for populations at each of the 10 sites, and for a proposed reintroduction site, under different levels of rat control. Only three of the reintroduction sites had λ distributions completely >1 for either model. At two sites, λ was expected to be >1 if rat-tracking rates were <5%. At the other five reintroduction sites, λ was predicted to be close to 1, and it was unclear whether growth was expected. Predictions of λ for the proposed reintroduction site were less precise than for other sites because distributions incorporated the full range of site-to-site random variation in vital rates. Our methods can be applied to any species for which postrelease data on demographic rates are available and potentially can be extended to model multiple species simultaneously.

Links between personality, early natal nutrition and survival of a threatened bird.

There is growing recognition that variation in animal personality traits can influence survival and reproduction rates, and consequently may be important for wildlife population dynamics. Despite this, the integration of personality research into conservation has remained uncommon. Alongside the establishment of personality as an important source of individual variation has come an increasing interest in factors affecting the development of personality. Recent work indicates the early environment, including natal nutrition, may play a stronger role in the development of personality than previously thought. In this study, we investigated the importance of three personality metrics (activity, boldness and acclimation time) for estimating survival of a threatened species, the hihi (Notiomystis cincta), and evaluated the influence of early natal nutrition on those metrics. Our results showed that boldness (as measured from a one-off cage test) had a positive effect on the probability of juvenile hihi surviving to adulthood. There was also a tendency for juveniles that received carotenoid supplementation in the nest to be bolder than those that did not, suggesting that the early environment had some influence on the expression of boldness in juvenile hihi. Linking the development of personality traits with ultimate effects on vital rates may benefit conservation management, as it could enable developmentally targeted management interventions. To our knowledge, this study is the first to identify potential linkages between early natal nutrition, personality and fitness in a wild-living population. This article is part of the theme issue 'Linking behaviour to dynamics of populations and communities: application of novel approaches in behavioural ecology to conservation'.