Species distribution models for conservation-oriented studies in Switzerland : filling data and tools gaps

SUMMARYSpecies distribution models (SDMs) represent nowadays an essential tool in the research fields of ecology and conservation biology. By combining observations of species occurrence or abundance with information on the environmental characteristic of the observation sites, they can provide information on the ecology of species, predict their distributions across the landscape or extrapolate them to other spatial or time frames. The advent of SDMs, supported by geographic information systems (GIS), new developments in statistical models and constantly increasing computational capacities, has revolutionized the way ecologists can comprehend species distributions in their environment. SDMs have brought the tool that allows describing species realized niches across a multivariate environmental space and predict their spatial distribution. Predictions, in the form of probabilistic maps showing the potential distribution of the species, are an irreplaceable mean to inform every single unit of a territory about its biodiversity potential. SDMs and the corresponding spatial predictions can be used to plan conservation actions for particular species, to design field surveys, to assess the risks related to the spread of invasive species, to select reserve locations and design reserve networks, and ultimately, to forecast distributional changes according to scenarios of climate and/or land use change.By assessing the effect of several factors on model performance and on the accuracy of spatial predictions, this thesis aims at improving techniques and data available for distribution modelling and at providing the best possible information to conservation managers to support their decisions and action plans for the conservation of biodiversity in Switzerland and beyond. Several monitoring programs have been put in place from the national to the global scale, and different sources of data now exist and start to be available to researchers who want to model species distribution. However, because of the lack of means, data are often not gathered at an appropriate resolution, are sampled only over limited areas, are not spatially explicit or do not provide a sound biological information. A typical example of this is data on 'habitat' (sensu biota). Even though this is essential information for an effective conservation planning, it often has to be approximated from land use, the closest available information. Moreover, data are often not sampled according to an established sampling design, which can lead to biased samples and consequently to spurious modelling results. Understanding the sources of variability linked to the different phases of the modelling process and their importance is crucial in order to evaluate the final distribution maps that are to be used for conservation purposes.The research presented in this thesis was essentially conducted within the framework of the Landspot Project, a project supported by the Swiss National Science Foundation. The main goal of the project was to assess the possible contribution of pre-modelled 'habitat' units to model the distribution of animal species, in particular butterfly species, across Switzerland. While pursuing this goal, different aspects of data quality, sampling design and modelling process were addressed and improved, and implications for conservation discussed. The main 'habitat' units considered in this thesis are grassland and forest communities of natural and anthropogenic origin as defined in the typology of habitats for Switzerland. These communities are mainly defined at the phytosociological level of the alliance. For the time being, no comprehensive map of such communities is available at the national scale and at fine resolution. As a first step, it was therefore necessary to create distribution models and maps for these communities across Switzerland and thus to gather and collect the necessary data. In order to reach this first objective, several new developments were necessary such as the definition of expert models, the classification of the Swiss territory in environmental domains, the design of an environmentally stratified sampling of the target vegetation units across Switzerland, the development of a database integrating a decision-support system assisting in the classification of the relevés, and the downscaling of the land use/cover data from 100 m to 25 m resolution.The main contributions of this thesis to the discipline of species distribution modelling (SDM) are assembled in four main scientific papers. In the first, published in Journal of Riogeography different issues related to the modelling process itself are investigated. First is assessed the effect of five different stepwise selection methods on model performance, stability and parsimony, using data of the forest inventory of State of Vaud. In the same paper are also assessed: the effect of weighting absences to ensure a prevalence of 0.5 prior to model calibration; the effect of limiting absences beyond the environmental envelope defined by presences; four different methods for incorporating spatial autocorrelation; and finally, the effect of integrating predictor interactions. Results allowed to specifically enhance the GRASP tool (Generalized Regression Analysis and Spatial Predictions) that now incorporates new selection methods and the possibility of dealing with interactions among predictors as well as spatial autocorrelation. The contribution of different sources of remotely sensed information to species distribution models was also assessed. The second paper (to be submitted) explores the combined effects of sample size and data post-stratification on the accuracy of models using data on grassland distribution across Switzerland collected within the framework of the Landspot project and supplemented with other important vegetation databases. For the stratification of the data, different spatial frameworks were compared. In particular, environmental stratification by Swiss Environmental Domains was compared to geographical stratification either by biogeographic regions or political states (cantons). The third paper (to be submitted) assesses the contribution of pre- modelled vegetation communities to the modelling of fauna. It is a two-steps approach that combines the disciplines of community ecology and spatial ecology and integrates their corresponding concepts of habitat. First are modelled vegetation communities per se and then these 'habitat' units are used in order to model animal species habitat. A case study is presented with grassland communities and butterfly species. Different ways of integrating vegetation information in the models of butterfly distribution were also evaluated. Finally, a glimpse to climate change is given in the fourth paper, recently published in Ecological Modelling. This paper proposes a conceptual framework for analysing range shifts, namely a catalogue of the possible patterns of change in the distribution of a species along elevational or other environmental gradients and an improved quantitative methodology to identify and objectively describe these patterns. The methodology was developed using data from the Swiss national common breeding bird survey and the article presents results concerning the observed shifts in the elevational distribution of breeding birds in Switzerland.The overall objective of this thesis is to improve species distribution models as potential inputs for different conservation tools (e.g. red lists, ecological networks, risk assessment of the spread of invasive species, vulnerability assessment in the context of climate change). While no conservation issues or tools are directly tested in this thesis, the importance of the proposed improvements made in species distribution modelling is discussed in the context of the selection of reserve networks.RESUMELes modèles de distribution d'espèces (SDMs) représentent aujourd'hui un outil essentiel dans les domaines de recherche de l'écologie et de la biologie de la conservation. En combinant les observations de la présence des espèces ou de leur abondance avec des informations sur les caractéristiques environnementales des sites d'observation, ces modèles peuvent fournir des informations sur l'écologie des espèces, prédire leur distribution à travers le paysage ou l'extrapoler dans l'espace et le temps. Le déploiement des SDMs, soutenu par les systèmes d'information géographique (SIG), les nouveaux développements dans les modèles statistiques, ainsi que la constante augmentation des capacités de calcul, a révolutionné la façon dont les écologistes peuvent comprendre la distribution des espèces dans leur environnement. Les SDMs ont apporté l'outil qui permet de décrire la niche réalisée des espèces dans un espace environnemental multivarié et prédire leur distribution spatiale. Les prédictions, sous forme de carte probabilistes montrant la distribution potentielle de l'espèce, sont un moyen irremplaçable d'informer chaque unité du territoire de sa biodiversité potentielle. Les SDMs et les prédictions spatiales correspondantes peuvent être utilisés pour planifier des mesures de conservation pour des espèces particulières, pour concevoir des plans d'échantillonnage, pour évaluer les risques liés à la propagation d'espèces envahissantes, pour choisir l'emplacement de réserves et les mettre en réseau, et finalement, pour prévoir les changements de répartition en fonction de scénarios de changement climatique et/ou d'utilisation du sol. En évaluant l'effet de plusieurs facteurs sur la performance des modèles et sur la précision des prédictions spatiales, cette thèse vise à améliorer les techniques et les données disponibles pour la modélisation de la distribution des espèces et à fournir la meilleure information possible aux gestionnaires pour appuyer leurs décisions et leurs plans d'action pour la conservation de la biodiversité en Suisse et au-delà. Plusieurs programmes de surveillance ont été mis en place de l'échelle nationale à l'échelle globale, et différentes sources de données sont désormais disponibles pour les chercheurs qui veulent modéliser la distribution des espèces. Toutefois, en raison du manque de moyens, les données sont souvent collectées à une résolution inappropriée, sont échantillonnées sur des zones limitées, ne sont pas spatialement explicites ou ne fournissent pas une information écologique suffisante. Un exemple typique est fourni par les données sur 'l'habitat' (sensu biota). Même s'il s'agit d'une information essentielle pour des mesures de conservation efficaces, elle est souvent approximée par l'utilisation du sol, l'information qui s'en approche le plus. En outre, les données ne sont souvent pas échantillonnées selon un plan d'échantillonnage établi, ce qui biaise les échantillons et par conséquent les résultats de la modélisation. Comprendre les sources de variabilité liées aux différentes phases du processus de modélisation s'avère crucial afin d'évaluer l'utilisation des cartes de distribution prédites à des fins de conservation.La recherche présentée dans cette thèse a été essentiellement menée dans le cadre du projet Landspot, un projet soutenu par le Fond National Suisse pour la Recherche. L'objectif principal de ce projet était d'évaluer la contribution d'unités 'd'habitat' pré-modélisées pour modéliser la répartition des espèces animales, notamment de papillons, à travers la Suisse. Tout en poursuivant cet objectif, différents aspects touchant à la qualité des données, au plan d'échantillonnage et au processus de modélisation sont abordés et améliorés, et leurs implications pour la conservation des espèces discutées. Les principaux 'habitats' considérés dans cette thèse sont des communautés de prairie et de forêt d'origine naturelle et anthropique telles que définies dans la typologie des habitats de Suisse. Ces communautés sont principalement définies au niveau phytosociologique de l'alliance. Pour l'instant aucune carte de la distribution de ces communautés n'est disponible à l'échelle nationale et à résolution fine. Dans un premier temps, il a donc été nécessaire de créer des modèles de distribution de ces communautés à travers la Suisse et par conséquent de recueillir les données nécessaires. Afin d'atteindre ce premier objectif, plusieurs nouveaux développements ont été nécessaires, tels que la définition de modèles experts, la classification du territoire suisse en domaines environnementaux, la conception d'un échantillonnage environnementalement stratifié des unités de végétation cibles dans toute la Suisse, la création d'une base de données intégrant un système d'aide à la décision pour la classification des relevés, et le « downscaling » des données de couverture du sol de 100 m à 25 m de résolution. Les principales contributions de cette thèse à la discipline de la modélisation de la distribution d'espèces (SDM) sont rassemblées dans quatre articles scientifiques. Dans le premier article, publié dans le Journal of Biogeography, différentes questions liées au processus de modélisation sont étudiées en utilisant les données de l'inventaire forestier de l'Etat de Vaud. Tout d'abord sont évalués les effets de cinq méthodes de sélection pas-à-pas sur la performance, la stabilité et la parcimonie des modèles. Dans le même article sont également évalués: l'effet de la pondération des absences afin d'assurer une prévalence de 0.5 lors de la calibration du modèle; l'effet de limiter les absences au-delà de l'enveloppe définie par les présences; quatre méthodes différentes pour l'intégration de l'autocorrélation spatiale; et enfin, l'effet de l'intégration d'interactions entre facteurs. Les résultats présentés dans cet article ont permis d'améliorer l'outil GRASP qui intègre désonnais de nouvelles méthodes de sélection et la possibilité de traiter les interactions entre variables explicatives, ainsi que l'autocorrélation spatiale. La contribution de différentes sources de données issues de la télédétection a également été évaluée. Le deuxième article (en voie de soumission) explore les effets combinés de la taille de l'échantillon et de la post-stratification sur le la précision des modèles. Les données utilisées ici sont celles concernant la répartition des prairies de Suisse recueillies dans le cadre du projet Landspot et complétées par d'autres sources. Pour la stratification des données, différents cadres spatiaux ont été comparés. En particulier, la stratification environnementale par les domaines environnementaux de Suisse a été comparée à la stratification géographique par les régions biogéographiques ou par les cantons. Le troisième article (en voie de soumission) évalue la contribution de communautés végétales pré-modélisées à la modélisation de la faune. C'est une approche en deux étapes qui combine les disciplines de l'écologie des communautés et de l'écologie spatiale en intégrant leurs concepts de 'habitat' respectifs. Les communautés végétales sont modélisées d'abord, puis ces unités de 'habitat' sont utilisées pour modéliser les espèces animales. Une étude de cas est présentée avec des communautés prairiales et des espèces de papillons. Différentes façons d'intégrer l'information sur la végétation dans les modèles de répartition des papillons sont évaluées. Enfin, un clin d'oeil aux changements climatiques dans le dernier article, publié dans Ecological Modelling. Cet article propose un cadre conceptuel pour l'analyse des changements dans la distribution des espèces qui comprend notamment un catalogue des différentes formes possibles de changement le long d'un gradient d'élévation ou autre gradient environnemental, et une méthode quantitative améliorée pour identifier et décrire ces déplacements. Cette méthodologie a été développée en utilisant des données issues du monitoring des oiseaux nicheurs répandus et l'article présente les résultats concernant les déplacements observés dans la distribution altitudinale des oiseaux nicheurs en Suisse.L'objectif général de cette thèse est d'améliorer les modèles de distribution des espèces en tant que source d'information possible pour les différents outils de conservation (par exemple, listes rouges, réseaux écologiques, évaluation des risques de propagation d'espèces envahissantes, évaluation de la vulnérabilité des espèces dans le contexte de changement climatique). Bien que ces questions de conservation ne soient pas directement testées dans cette thèse, l'importance des améliorations proposées pour la modélisation de la distribution des espèces est discutée à la fin de ce travail dans le contexte de la sélection de réseaux de réserves.

Fire management, climate change and their interacting effects on birds in complex Mediterranean landscapes: dynamic distribution modelling of an early-successional species - the near-threatened Dartford Warbler (Sylvia undata)

The current challenge in a context of major environmental changes is to anticipate the responses of species to future landscape and climate scenarios. In the Mediterranean basin, climate change is one the most powerful driving forces of fire dynamics, with fire frequency and impact having markedly increased in recent years. Species distribution modelling plays a fundamental role in this challenge, but better integration of available ecological knowledge is needed to adequately guide conservation efforts. Here, we quantified changes in habitat suitability of an early-succession bird in Catalonia, the Dartford Warbler (Sylvia undata) ― globally evaluated as Near Threatened in the IUCN Red List. We assessed potential changes in species distributions between 2000 and 2050 under different fire management and climate change scenarios and described landscape dynamics using a spatially-explicit fire-succession model that simulates fire impacts in the landscape and post-fire regeneration (MEDFIRE model). Dartford Warbler occurrence data were acquired at two different spatial scales from: 1) the Atlas of European Breeding Birds (EBCC) and 2) Catalan Breeding Bird Atlas (CBBA). Habitat suitability was modelled using five widely-used modelling techniques in an ensemble forecasting framework. Our results indicated considerable habitat suitability losses (ranging between 47% and 57% in baseline scenarios), which were modulated to a large extent by fire regime changes derived from fire management policies and climate changes. Such result highlighted the need for taking the spatial interaction between climate changes, fire-mediated landscape dynamics and fire management policies into account for coherently anticipating habitat suitability changes of early succession bird species. We conclude that fire management programs need to be integrated into conservation plans to effectively preserve sparsely forested and early succession habitats and their associated species in the face of global environmental change.

Declines of Aerial Insectivores in North America Follow a Geographic Gradient

North American birds that feed on aerial insects are experiencing widespread population declines. An analysis of the North American Breeding Bird Survey trend estimates for 1966 to 2006 suggests that declines in this guild are significantly stronger than in passerines in general. The pattern of decline also shows a striking geographical gradient, with aerial insectivore declines becoming more prevalent towards the northeast of North America. Declines are also more acute in species that migrate long distances compared to those that migrate short distances. The declines become manifest, almost without exception, in the mid 1980s. The taxonomic breadth of these downward trends suggests that declines in aerial insectivore populations are linked to changes in populations of flying insects, and these changes might be indicative of underlying ecosystem changes.

Birding 2.0: Citizen Science and Effective Monitoring in the Web 2.0 World

The amateur birding community has a long and proud tradition of contributing to bird surveys and bird atlases. Coordinated activities such as Breeding Bird Atlases and the Christmas Bird Count are examples of "citizen science" projects. With the advent of technology, Web 2.0 sites such as eBird have been developed to facilitate online sharing of data and thus increase the potential for real-time monitoring. However, as recently articulated in an editorial in this journal and elsewhere, monitoring is best served when based on a priori hypotheses. Harnessing citizen scientists to collect data following a hypothetico-deductive approach carries challenges. Moreover, the use of citizen science in scientific and monitoring studies has raised issues of data accuracy and quality. These issues are compounded when data collection moves into the Web 2.0 world. An examination of the literature from social geography on the concept of "citizen sensors" and volunteered geographic information (VGI) yields thoughtful reflections on the challenges of data quality/data accuracy when applying information from citizen sensors to research and management questions. VGI has been harnessed in a number of contexts, including for environmental and ecological monitoring activities. Here, I argue that conceptualizing a monitoring project as an experiment following the scientific method can further contribute to the use of VGI. I show how principles of experimental design can be applied to monitoring projects to better control for data quality of VGI. This includes suggestions for how citizen sensors can be harnessed to address issues of experimental controls and how to design monitoring projects to increase randomization and replication of sampled data, hence increasing scientific reliability and statistical power.

Biased representation of disturbance rates in the roadside sampling frame in boreal forests: implications for monitoring design

The North American Breeding Bird Survey (BBS) is the principal source of data to inform researchers about the status of and trend for boreal forest birds. Unfortunately, little BBS coverage is available in the boreal forest, where increasing concern over the status of species breeding there has increased interest in northward expansion of the BBS. However, high disturbance rates in the boreal forest may complicate roadside monitoring. If the roadside sampling frame does not capture variation in disturbance rates because of either road placement or the use of roads for resource extraction, biased trend estimates might result. In this study, we examined roadside bias in the proportional representation of habitat disturbance via spatial data on forest “loss,” forest fires, and anthropogenic disturbance. In each of 455 BBS routes, the area disturbed within multiple buffers away from the road was calculated and compared against the area disturbed in degree blocks and BBS strata. We found a nonlinear relationship between bias and distance from the road, suggesting forest loss and forest fires were underrepresented below 75 and 100 m, respectively. In contrast, anthropogenic disturbance was overrepresented at distances below 500 m and underrepresented thereafter. After accounting for distance from road, BBS routes were reasonably representative of the degree blocks they were within, with only a few strata showing biased representation. In general, anthropogenic disturbance is overrepresented in southern strata, and forest fires are underrepresented in almost all strata. Similar biases exist when comparing the entire road network and the subset sampled by BBS routes against the amount of disturbance within BBS strata; however, the magnitude of biases differed. Based on our results, we recommend that spatial stratification and rotating panel designs be used to spread limited BBS and off-road sampling effort in an unbiased fashion and that new BBS routes be established where sufficient road coverage exists.

Conservation in tropical landscape mosaics: the case of the cacao landscape of southern Bahia, Brazil

A recent debate has contrasted two conservation strategies in agricultural landscapes; either ""land sparing`` farm development combining intensive production practices with forest set-asides, or ""wildlife-friendly`` farming with greater on-farm habitat value but lower yields. We argue that in established mosaic landscapes including old cacao production regions where natural forest has already been reduced to relatively small fragments, a combination of both strategies is needed to conserve biodiversity. After reviewing the evidence for the insufficiency of either strategy alone if applied to such landscapes, the paper focuses on the cacao production landscape of southern Bahia, Brazil, once the world`s second largest cacao producer. Here, small remaining areas of Atlantic Forest are embedded in a matrix dominated by traditional cacao agroforests, resulting in a landscape mosaic that has proven favorable to the conservation of the region`s high biodiversity. We show that current land use dynamics and public policies pose threats but also offer opportunities to conservation and describe a three- pronged landscape conservation strategy, consisting of (i) expansion of the protected areas system, (ii) promotion of productive yet biodiversity-friendly cacao farming practices, and (iii) assistance to land users to implement legally mandated on-farm reserves and voluntary private reserves. We discuss recent experiences concerning the implementation of this strategy, discuss likely future scenarios, and reflect on the applicability of the Bahian experience to biodiversity rich cacao production regions elsewhere in the tropics.

Density and population size of mammals introduced on a land-bridge island in southeastern Brazil

The introduction of alien species is one of the main threats to the conservation of native species, especially in island ecosystems. Here, we report on the population growth of 15 species of mammals introduced in 1983 on the island of Anchieta, an 828 ha land-bridge island in southeastern Brazil. We estimated the density of mammals through 296 km of line transect census. Five species introduced became extinct (coypu, brocket deer, six-banded armadillo, nine-banded armadillo, maned three-toed sloth); six became over-abundant (marmoset, coati, agouti, seven-banded armadillo, and capybara); one has a stable population (capuchin monkey). Anchieta Island has the highest density of mammals in the entire Atlantic forest (486.77 ind/km(2)), especially nest predators (232.83 ind/km(2)) and herbivores (253.58 ind/km(2)). Agoutis (Dasyprocta spp.) and marmosets (Callithrix penicillata) were, by far, the species with the highest population growth. The high density of mammals in this island may have strong consequences for plant recruitment and bird diversity.

Fragmentação florestal: efeitos em múltipla escala sobre a diversidade de aves em remanescentes florestais no noroeste do Estado de São Paulo

Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES)

Recommendations for monitoring avian populations with point counts: a case study in southeastern Brazil

No hemisfério norte, o censo de aves é fundamental para gerar informações que auxiliam na compreensão de tendências populacionais. Tais censos, devido à marcada sazonalidade deste hemisfério, são realizados durante dois momentos distintos: na estação reprodutiva (aves residentes) e no inverno (quando as aves migratórias deixam determinadas regiões). Na região neotropical, porém, dependendo da localidade, as aves podem se reproduzir durante qualquer ou vários períodos do ano; podem ou não migrar, e aquelas que o fazem podem apresentar um padrão assincrônico. Em contraste com o hemisfério norte, tendências populacionais são desconhecidas, bem como o impacto das taxas rápidas de urbanização e desmatamento, que também são pouco monitoradas. Para melhor entender padrões temporais de riqueza e abundância de aves, e avaliar como um censo similar pode ser implementado na América tropical, foram utilizados pontos de escuta ao longo de 12 meses em uma localidade no Estado de São Paulo, sudeste do Brasil. Os censos ocorreram duas vezes por dia (manhãs/tardes) em uma floresta semidecidual ao longo de transecções com 10 pontos (20 pontos por dia) distantes 200 m entre si e com raio de detecção limitado em 100 m. Ambas as riquezas e abundâncias de aves foram maiores durante as manhãs, mas as curvas de acumulação sugerem que os censos vespertinos com maior esforço amostral podem fornecer resultados similares aos censos matutinos. Riqueza e abundância das aves não variam de acordo com estações (i.e., sem padrão aparente entre reprodução e migração), enquanto espécies exclusivas foram encontradas todos os meses e relativamente poucas espécies (20%) foram registradas em todos os meses do ano. Durante este ano, 84% de todas as aves florestais da área estudada foram registradas. Sugerimos que a metodologia de pontos de escuta pode ser utilizada à semelhança dos censos do hemisfério norte. Recomendamos ainda que o esforço amostral em transecções deva incluir ao menos 20 pontos, e que o início da contagem das aves deva ser sazonal, utilizando o período de migração das espécies austrais (e os seis meses seguintes) para coordenar pontos de escuta. Por último, sugerimos que os censos no Brasil e até mesmo na América Latina podem ajudar no entendimento de tendências populacionais, mas também demandam maior esforço do que o observado em latitudes temperadas, devido à maior riqueza de espécies e diferenças nas dinâmicas de reprodução e migração. Por meio do uso de censos de aves coordenados poderá ser desenvolvida uma técnica para os trópicos que irá gerar informações que permitam acompanhar tendências populacionais, com benefícios para a conservação das aves, similarmente aos censos realizados em países do hemisfério norte.

As influências da paisagem urbana em grupos funcionais e riqueza de aves

Pós-graduação em Biologia Animal - IBILCE

Ácaros de penas (Acari: Astigmata) em aves não passeriformes do Brasil

Pós-graduação em Ciências Biológicas (Zoologia) - IBRC

Retention of canopy trees as biological legacies for balancing woody biomass production and biodiversity in managed aspen forests of the Great Lakes Region

Green-tree retention under the conceptual framework of ecological forestry has the potential to provide both biomass feedstock for industry and maintain quality wildlife habitat. I examined the effects of retained canopy trees as biological legacies (“legacy trees”) in aspen (Populus spp.) forests on above-ground live woody biomass, understory plant floristic quality, and bird diversity. Additionally, I evaluated habitat quality for a high conservation priority species, the Golden-winged Warbler (Vermivora chrysoptera). I selected 27 aspen-dominated forest stands in northern Wisconsin with nine stands in each of three legacy tree retention treatments (conifer retention, hardwood retention, and clearcuts or no retention) across a chronosequence (4-36 years post-harvest). Conifer retention stands had greater legacy tree and all tree species biomass but lower regenerating tree biomass than clearcuts. Coniferous but not hardwood legacy trees appeared to suppress regenerating tree biomass. I evaluated the floristic quality of the understory plant assemblage by estimating the mean coefficient of conservatism (C). Mean C was lower in young stands than in middle-age or old stands; there was a marginally significant (p=0.058) interaction effect between legacy tree retention treatment and stand age. Late-seral plant species were positively associated with stand age and legacy tree diameter or age revealing an important relationship between legacy tree retention and stand development. Bird species richness was greatest in stands with hardwood retention particularly early in stand development. Six conservation priority bird species were indicators of legacy tree retention or clearcuts. Retention of legacy trees in aspen stands provided higher quality nest habitat for the Golden-winged Warbler than clearcuts based on high pairing success and nesting activity. Retention of hardwoods, particularly northern red oak (Quercus rubra), yielded the most consistent positive effects in this study with the highest bird species richness and the highest quality habitat for the Golden-winged Warbler. This treatment maintained stand biomass comparable to clearcuts and did not suppress regenerating tree biomass. In conclusion, legacy tree retention can enhance even-aged management techniques to produce a win-win scenario for the conservation of declining bird species and late-seral understory plants and for production of woody biomass feedstock from naturally regenerating aspen forests.