Old-growth forest is what giant pandas really need

Giant pandas (Ailuropoda melanoleuca) are an iconic conservation species, but despite significant research effort, do we understand what they really need? Estimating and mapping suitable habitat play a critical role in conservation planning and policy. But if assumptions about ecological needs are wrong, maps with misidentified suitable habitat will misguide conservation action. Here, we use an information-theoretic approach to analyse the largest, landscape-level dataset on panda habitat use to date, and challenge the prevailing wisdom about panda habitat needs. We show that pandas are associated with old-growth forest more than with any ecological variable other than bamboo. Other factors traditionally used in panda habitat models, such as topographic slope, are less important. We suggest that our findings are disparate from previous research in part because our research was conducted over a larger ecological scale than previous research conducted over more circumscribed areas within individual reserves. Thus, extrapolating from habitat studies on small scales to conservation planning on large scales may entail some risk. As the Chinese government is considering the renewal of its logging ban, it should take heed of the panda''s dependency on old growth.     

岷山地区大熊猫(Ailuropoda melanoleuca)生境影响因子连通性及主导因子

岷山地区是大熊猫保护的关键地区.运用图论分析法研究了岷山地区大熊猫生境影响因子,及其相互作用关系.结果表明:在岷山地区,12个影响因子66对组合中,47.0%的影响因子间存在直接的联系,其中89.2%的影响具有增大效应,这说明人类活动对大熊猫生境的影响具有协同增大效应,并占主导地位.研究发现,在影响因子关系集中,存在强连通性的子集K={(TD);(RB);(MI);(HPL);(AP);(AD);(SR);(TH);(FC)},这表明近期进行的旅游景点开发,公路建设,采矿,高压电线走廊建设,以及传统的农业开发,畜牧业,薪柴采集等影响因子之间存在着强烈相互作用关系,这些人类活动还会将其他人类活动对大熊猫生境的影响放大,加剧其对大熊猫生境的不利影响,是影响岷山地区大熊猫生境的主导因素.研究还表明如果不能对主导因子进行有效的控制,很难有效地保护岷山地区大熊猫生境.研究表明图论分析法是研究大熊猫生境影响因子的一个有效工具,有助于明确影响熊猫生境的主导和关键因子,为制定有效的大熊猫保护策略提供科学依据.     

Hopes and challenges for giant panda conservation under climate change in the Qinling Mountains of China

One way that climate change will impact animal distributions is by altering habitat suitability and habitat fragmentation. Understanding the impacts of climate change on currently threatened species is of immediate importance because complex conservation planning will be required. Here, we mapped changes to the distribution, suitability, and fragmentation of giant panda habitat under climate change and quantified the direction and elevation of habitat shift and fragmentation patterns. These data were used to develop a series of new conservation strategies for the giant panda. Qinling Mountains, Shaanxi, China. Data from the most recent giant panda census, habitat factors, anthropogenic disturbance, climate variables, and climate predictions for the year 2050 (averaged across four general circulation models) were used to project giant panda habitat in Maxent. Differences in habitat patches were compared between now and 2050. While climate change will cause a 9.1% increase in suitable habitat and 9% reduction in subsuitable habitat by 2050, no significant net variation in the proportion of suitable and subsuitable habitat was found. However, a distinct climate change‐induced habitat shift of 11 km eastward by 2050 is predicted firstly. Climate change will reduce the fragmentation of suitable habitat at high elevations and exacerbate the fragmentation of subsuitable habitat below 1,900 m above sea level. Reduced fragmentation at higher elevations and worsening fragmentation at lower elevations have the potential to cause overcrowding of giant pandas at higher altitudes, further exacerbating habitat shortage in the central Qinling Mountains. The habitat shift to the east due to climate change may provide new areas for giant pandas but poses severe challenges for future conservation.     

Assessing vulnerability of giant pandas to climate change in the Qinling Mountains of China

Climate change might pose an additional threat to the already vulnerable giant panda (Ailuropoda melanoleuca). Effective conservation efforts require projections of vulnerability of the giant panda in facing climate change and proactive strategies to reduce emerging climate‐related threats. We used the maximum entropy model to assess the vulnerability of giant panda to climate change in the Qinling Mountains of China. The results of modeling included the following findings: (1) the area of suitable habitat for giant pandas was projected to decrease by 281 km² from climate change by the 2050s; (2) the mean elevation of suitable habitat of giant panda was predicted to shift 30 m higher due to climate change over this period; (3) the network of nature reserves protect 61.73% of current suitable habitat for the species, and 59.23% of future suitable habitat; (4) current suitable habitat mainly located in Chenggu, Taibai, and Yangxian counties (with a total area of 987 km²) was predicted to be vulnerable. Assessing the vulnerability of giant panda provided adaptive strategies for conservation programs and national park construction. We proposed adaptation strategies to ameliorate the predicted impacts of climate change on giant panda, including establishing and adjusting reserves, establishing habitat corridors, improving adaptive capacity to climate change, and strengthening monitoring of giant panda.     

卧龙自然保护区大熊猫生境评价

生物的生境是指生物生活繁衍的场所,由生物与非生物环境构成。近几个世纪以来,物种绝灭的速度加快,生物多样性丧失最重要的原因是生物生境的人为破坏。对保持生物的生境评价,是分析这些物种种群减少,濒危原因的重要手段,还能为制定合理的保护对策提供依据。根据大熊猫生境分布特点提出了大熊猫生境结构理论模型,将影响卧龙大熊猫生境质量的因素分为物理环境因素、生物环境因素和人类活动因素,探讨了生境评价的程序与卧龙大熊猫生境评价准则,运用地理信息系统技术与空间模拟方法分析了卧龙大熊猫生境质量。在人类活动影响下,卧龙自然保护区内适宜大熊猫生存的生境面积有57597.3hm^2,其中最适生境面积为6256.1hm^2,主要分布在海拔2300－2800m的平缓山坡与台地。     

Integrating population size analysis into habitat suitability assessment: implications for giant panda conservation in the Minshan Mountains,China

Compared to conventional approaches, the integration of population size analysis with habitat suitability assessment on a large scale can provide more evidence to explain the mechanisms of habitat isolation and fragmentation, and thus make regional conservation plans. In this paper, we analyzed the habitat suitability for giant pandas in the Minshan Mountains, China, using the ecological-niche factor analysis (ENFA) method, and then evaluated the current conservation status of this endangered species. The results showed that (1) giant pandas were distributed in a narrow altitudinal range in which vegetation cover was dominated by coniferous forest, mixed coniferous and deciduous broadleaf forest, and deciduous broadleaf forest with scattered bamboo understory, and (2) roads and human settlements had strong negative effects on the panda habitat selection. According to habitat analysis, the total habitat area of giant panda in the Minshan Mountains was 953,173 ha, which was fragmented into 12 habitat units by major roads, rivers, and human settlements. The habitat of the mid-Minshan was less fragmentized, but was seriously fragmented in the north. The panda population size estimation showed that 676 individuals inhabited the study area, and 94.53% of them were in the mid-Minshan, but small panda populations less than 30 individuals inhabited the isolated and fragmented habitat patches in the north. The nature reserves in the Minshan Mountains have formed three conservation groups, which covered 41.26% of panda habitat and protected 70.71% of panda population of the study area, but there still exists two conservation gaps, and the connectivity among these reserves is still weak. Due to habitat isolation and extensive human disturbances, giant pandas in the north (i.e., Diebu, Zhouqu, and Wudou) are facing threats of local extinction. In order to protect pandas and their habitats in this area, some effective conservation approaches, such as establishing new reserves in gap areas, creating corridors among patches, and seasonally controlling traffic flux in key roads, should be implemented in the future to link these isolated habitats together.     

Habitat requirements of the Himalayan red panda (Ailurus fulgens) and threat analysis in Jigme Dorji National Park,Bhutan

Understanding the influence of anthropogenic disturbances on species’ habitat use and distribution is critical to conservation managers in planning effective conservation strategies and mitigating the impact of development. Few studies have focused on the Himalayan red panda (Ailurus fulgens) in Bhutan. This study aimed to assess the habitat requirements and threats to this endangered species in the Khamaed subdistrict of the Jigme Dorji National Park, Bhutan. We employed a transect walk and plot‐sampling survey design across two seasons, that is, winter and spring. In total, we surveyed 84 × 50 m radius circular plots along 51 km of existing trails within a 25.4 km² study area. At 500 m intervals, we established plots at random distances and direction from the trail. We recorded direct sightings (n = 2) and indirect signs (n = 14), such as droppings and footprints as evidence of red panda presence within an altitudinal range of 2,414–3,618 m. We also noted 21 tree and 12 understory species within plots with red panda evidence; the dominant tree species was the Himalayan hemlock (Tsuga dumosa) and the Asian barberry (Berberis asiatica) as an understory species. Red panda presence showed a significant positive association with distance to water sources and fir forests. Plant disturbance and infrastructure, such as power transmission lines, were identified as prominent anthropogenic threats in the study area. Based on our findings, we recommend the development and implementation of local forest management plans, livestock intensification programs, and strict application of environmental impact assessment regulations to promote the conservation of the red panda in the region.     

Habitat correlates of the red panda in the temperate forests of Bhutan

Anthropogenic activities and associated global climate change are threatening the biodiversity in the Himalayas against a backdrop of poor knowledge of the region's threatened species. The red panda (Ailurus fulgens) is a threatened mammal confined to the eastern Himalayas, and because of Bhutan's central location in the distributional range of red pandas, its forests are integral to the long-term viability of wild populations. Detailed habitat requirements of the red panda are largely speculative, and there is virtually no ecological information available on this species in Bhutan. Between 2007 and 2009, we established 615 presence/absence plots in a systematic sampling of resident habitat types within Jigme Dorji and Thrumshingla National Parks, Bhutan, to investigate broad and fine-scale red panda habitat associations. Additional locality records of red pandas were obtained from interviewing 664 park residents. Red pandas were generally confined to cool broadleaf and conifer forests from 2,110-4,389 m above sea level (asl), with the majority of records between 2,400-3,700 m asl on south and east-facing slopes. At a finer scale, multivariate analysis revealed that red pandas were strongly associated with old growth Bhutan Fir (Abies densa) forest dominated by a dense cover of Yushania and Arundanaria bamboo with a high density of fallen logs and tree stumps at ground level; a high density of trees, dead snags, and rhododendron shrubs in the mid-storey; and locations that were close to water. Because Bhutan's temperate forests that encompass prime red panda habitat are also integral to human subsistence and socio-economic development, there exists an inadvertent conflict between the needs of people and red pandas. As such, careful sustainable management of Bhutan's temperate forests is necessary if a balance is to be met between the socioeconomic needs of people and the conservation goals for red pandas.     

Identifying core habitats and corridors for giant pandas by combining multiscale random forest and connectivity analysis

Habitat loss and fragmentation are widely acknowledged as the main driver of the decline of giant panda populations. The Chinese government has made great efforts to protect this charming species and has made remarkable achievements, such as population growth and habitat expansion. However, habitat fragmentation has not been reversed. Protecting giant pandas in a large spatial extent needs to identify core habitat patches and corridors connecting them. This study used an equal‐sampling multiscale random forest habitat model to predict a habitat suitability map for the giant panda. Then, we applied the resistant kernel method and factorial least‐cost path analysis to identify core habitats connected by panda dispersal and corridors among panda occurrences, respectively. Finally, we evaluated the effectiveness of current protected areas in representing core habitats and corridors. Our results showed high scale dependence of giant panda habitat selection. Giant pandas strongly respond to bamboo percentage and elevation at a relatively fine scale (1 km), whereas they respond to anthropogenic factors at a coarse scale (≥2 km). Dispersal ability has significant effects on core habitats extent and population fragmentation evaluation. Under medium and high dispersal ability scenarios (12,000 and 20,000 cost units), most giant panda habitats in the Qionglai mountain are predicted to be well connected by dispersal. The proportion of core habitats covered by protected areas varied between 38% and 43% under different dispersal ability scenarios, highlighting significant gaps in the protected area network. Similarly, only 43% of corridors that connect giant panda occurrences were protected. Our results can provide crucial information for conservation managers to develop wise strategies to safeguard the long‐term viability of the giant panda population.     

Spatial genetic structure and dispersal of giant pandas on a mountain-range scale

Understanding the spatial genetic structure of populations can provide insight into the ecological or evolutionary processes of the species, and enable wise conservation decisions. We examined the spatial genetic structure of a giant panda (Ailuropoda melanoleuca) population in a heterogeneous mountainous landscape using noninvasive genetic sampling and 12 microsatellite loci. Nonrandom genetic structure was detected through spatial autocorrelation analysis, demonstrating a significantly positive autocorrelation over closer distances. Additional spatial analyses showed significantly positive genetic correlation among spatially-proximate males, and no correlation among females and among male–female pairs. These findings suggest a female-biased dispersal pattern and cryptic family grouping among giant pandas on a large mountain-range scale. The spatial extent of genetic structure occurred within 12.5 km, measured by a least-cost path distance model integrating information of habitat quality and habitat preferences of this species. Using the bearing analysis of PASSAGE, we found that directional genetic autocorrelations were in agreement with habitat structure, and habitat heterogeneity may affect the direction of giant panda dispersal. The characterization of spatial genetic structure can provide potentially valuable information for the conservation and management of giant pandas and their habitat.     

Evaluating Landscape Options for Corridor Restoration between Giant Panda Reserves

The establishment of corridors can offset the negative effects of habitat fragmentation by connecting isolated habitat patches. However, the practical value of corridor planning is minimal if corridor identification is not based on reliable quantitative information about species-environment relationships. An example of this need for quantitative information is planning for giant panda conservation. Although the species has been the focus of intense conservation efforts for decades, most corridor projects remain hypothetical due to the lack of reliable quantitative researches at an appropriate spatial scale. In this paper, we evaluated a framework for giant panda forest corridor planning. We linked our field survey data with satellite imagery, and conducted species occupancy modelling to examine the habitat use of giant panda within the potential corridor area. We then conducted least-cost and circuit models to identify potential paths of dispersal across the landscape, and compared the predicted cost under current conditions and alternative conservation management options considered during corridor planning. We found that due to giant panda''s association with areas of low elevation and flat terrain, human infrastructures in the same area have resulted in corridor fragmentation. We then identified areas with high potential to function as movement corridors, and our analysis of alternative conservation scenarios showed that both forest/bamboo restoration and automobile tunnel construction would significantly improve the effectiveness of corridor, while residence relocation would not significantly improve corridor effectiveness in comparison with the current condition. The framework has general value in any conservation activities that anticipate improving habitat connectivity in human modified landscapes. Specifically, our study suggested that, in this landscape, automobile tunnels are the best means to remove current barriers to giant panda movements caused by anthropogenic interferences.     

An assessment of human impacts on endangered red pandas (Ailurus fulgens) living in the Himalaya

Anthropogenic factors play an important role in shaping the distribution of wildlife species and their habitats, and understanding the influence of human activities on endangered species can be key to improving conservation efforts as well as the implementation of national strategies for sustainable development. Here, we used species distribution modeling to assess human impacts on the endangered red panda (Ailurus fulgens) in high‐altitude regions of Nepal. We found that the distance to paths (tracks used by people and animals), livestock density, human population density, and annual mean temperature were the most important factors determining the habitat suitability for red pandas in Nepal. This is the first study that attempts to use comprehensive environmental and anthropogenic variables to predict habitat suitability for the red pandas at a national level. The suitable habitat identified by this study is important and could serve as a baseline for the development of conservation strategies for the red panda in Nepal.     

野生大熊猫现状、面临的挑战及展望

截至2003年底,我国野生大熊猫种群数量达1596只,分布在陕西、四川和甘肃3省的45个县境内,总栖息地面积达2304991hm^2。与第2次大熊猫调查相比,野生大熊猫生存状况已得到改善,分布范围扩大、栖息地面积增加、种群数量进一步增长。本文在第3次大熊猫调查的基础上,就野生大熊猫种群及栖息地现状进行了分析,指出未来保护大熊猫所面临的3个方面的挑战,即来自物种自身生物学特性的挑战、栖息地破碎化及隔离小种群未来命运的挑战以及大熊猫保护与社区经济发展需求相冲突的挑战。作者还就我国大熊猫保护前景进行了展望,即自然保护区数量将进一步增加,栖息地状况将进一步改善;种群数量在总体保持稳定的基础上将逐步增长,但局部小种群灭绝风险将加剧;圈养种群将形成能自我维持的种群,圈养个体通过培训将逐步放归到隔离野生小种群中以改变其命运。     

Potential habitat distribution of Himalayan red panda and their connectivity in Sakteng Wildlife Sanctuary,Bhutan

Survival of endangered Himalayan red panda is threatened by ever‐growing anthropogenic activities leading to an unprecedented rate of habitat degradation and loss. However, limited studies have been conducted in the context of the spatial distribution of habitats and habitat connectivity for the species in the landscape of Sakteng Wildlife Sanctuary (SWS). Lack of such information remains a challenge while implementing effective and holistic conservation initiatives. Therefore, this study identifies the distribution of potential habitats and their connectivity using maxent and linkage mapper, respectively. Precipitation‐related predictor variables exhibited a significant influence on the prediction of habitat distribution. The model predicted 27.7% of the SWS as a potential habitat (fundamental niche). More than 75% of the predicted habitats fall outside the existing core zones where anthropogenic disturbance is relatively high, indicating the need to reassess existing management options. In SWS, 15 core habitats (CH) are predicted which are connected by a least‐cost corridor (length µ = 2.91 km) with several pinch points in it. Centrally located CH5 and CH11 are identified as the most important habitat in maintaining overall connectivity within SWS. However, CH located in the peripheries could be equally important in facilitating the transboundary movement of the species. Overall, SWS can play a critical role as a connecting link between the larger landscape of Bhutan and the adjacent Indian state of Arunachal Pradesh in the conservation of Himalayan red panda that exhibits narrow dispersal with special habitat needs. Based on our findings, we recommend initiating GPS/satellite telemetry of the species to enable SWS to understand the precise interaction of Himalayan red panda to widespread herder communities, livestock, and free‐roaming dogs dwelling in the same landscape. It will also help to evaluate the functionality of the predicted habitats, linkages, and feasibility of transboundary conservation initiatives.     

Balance between climate change mitigation benefits and land use impacts of bioenergy: conservation implications for European birds

Both climate change and habitat modification exert serious pressure on biodiversity. Although climate change mitigation has been identified as an important strategy for biodiversity conservation, bioenergy remains a controversial mitigation action due to its potential negative ecological and socio-economic impacts which arise through habitat modification by land use change. While the debate continues, the separate or simultaneous impacts of both climate change and bioenergy on biodiversity have not yet been compared. We assess projected range shifts of 156 European bird species by 2050 under two alternative climate change trajectories: a baseline scenario, where the global mean temperature increases by 4 °C by the end of the century, and a 2 degrees scenario, where global concerted effort limits the temperature increase to below 2 °C. For the latter scenario, we also quantify the pressure exerted by increased cultivation of energy biomass as modelled by IMAGE2.4, an integrated land use model. The global bioenergy use in this scenario is in the lower end of the range of previously estimated sustainable potential. Under the assumptions of these scenarios, we find that the magnitude of range shifts due to climate change is far greater than the impact of land conversion to woody bioenergy plantations within the European Union, and that mitigation of climate change reduces the exposure experienced by species. However, we identified potential for local conservation conflict between priority areas for conservation and bioenergy production. These conflicts must be addressed by strict bioenergy sustainability criteria that acknowledge biodiversity conservation needs beyond existing protected areas and apply also to biomass imported from outside the European Union.     

圈养大熊猫野化培训期的生境选择特征

大熊猫(Ailuropoda melanoleuca)是我国特有的珍稀物种,也是世界上最濒危的野生动物之一。为了将人工繁育的部分大熊猫个体重引入其历史分布区或复壮野生种群,中国保护大熊猫研究中心从2003年开始进行圈养大熊猫的野外放归工作,通过野化培训以提高圈养大熊猫适应和选择野外环境的能力。对野化培训大熊猫"淘淘"的生境选择研究表明:该野化培训大熊猫幼仔经常活动于新笋密度较大的区域[生境与对照:(2.68±1.14)对(1.58±0.66)],却避开成竹密度过大[(9.91±2.51)对(12.18±4.68)]、竹子较高[(4.57±1.09) m对(4.98±0.66) m]以及枯死竹过多[(2.52±0.86)对(3.39±1.33)]的区域;喜欢活动于离水源[(1.59±0.67)对(2.19±0.87)]和隐蔽场所较近[(5.37±2.14) m对(8.35±7.76)m],以及距离乔木较远[(3.09±0.69) m对(2.70±0.42) m]和郁闭度较低[(1.85±0.57)对(2.10±0.47)]的区域(P < 0.05),新笋密度大小是该栖息地在整个野化培训期间是否被利用的最重要因素。该野化培训大熊猫幼仔保持着与带仔母兽相近的生境选择特征,对竹子环境的选择也与卧龙野生大熊猫相似,野化培训对该大熊猫幼仔产生了积极的作用。野化培训大熊猫幼仔形成的家域和核域面积分别为9.21 hm² 和1.93 hm²,占野化培训圈面积的51.95%和10.89%,其中家域面积仅有卧龙野生大熊猫的1.4%-2.4%,所以在以后的野化培训过程中需要采取增加野化培训圈中环境丰富度等方式,促进野化培训大熊猫形成较大的家域面积。     

Conservation and human livelihoods at the crossroads: Local needs and knowledge in the management of Arabuko Sokoke Forest

Arabuko Sokoke Forest is the largest remaining single block of indigenous dry coastal tropical forest in Eastern Africa. Households within a 5 km buffer zone depend heavily on the forest for their livelihood needs, and the pressure on forest resources is on the increase. In May 2015, 109 households were interviewed on resources they obtain from the forest, in terms of the self‐reported level of monthly income. We found household income and farm size significantly positively correlated with benefits from the forest, highlighting the possible influence of household wealth in exploiting forest resources. A large proportion of households (32%) had limited knowledge of local birds, while human–bird conflict was reported by 44% of the households. While many households were keen to participate in conservation projects that maintain the forest, 44% had no knowledge of the forest management plan, and 60% of those interviewed had no idea of how forest zones were designated for particular activities. Drivers for local community participation in conservation projects appear to be sustainable income and fulfilment of basic household needs.     

Habitat availability for multiple avian species under modeled alternative conservation scenarios in the Two Hearted River watershed in Michigan,USA

Due to differences in the responses of species to changing landscape patterns, developing a conservation plan with an optimal outcome of supporting contrasting habitat needs can be difficult. Landscape scenario modeling can provide a means to compare alternative conservation strategies and can reveal tradeoffs of managing for one objective versus another. In order to evaluate the impacts of alternative conservation strategies in a 53,653 ha landscape in Michigan's Upper Peninsula, four scenarios of alternative conservation strategies were modeled 100 years into the future using the VDDT^®/TELSA^® spatial model suite, and habitat availability was evaluated for five target bird species of local conservation concern under each scenario. The target species were Dendroica fusca (Blackburnian Warbler), Picoides arcticus (Black-backed Woodpecker), Dendroica kirtlandii (Kirtland's Warbler), Buteo lineatus (Red-shouldered Hawk), and Scolopax minor (American Woodcock). Scenarios were ranked based on relative performance of three habitat metric results (total primary habitat area, average size of habitat patches, and average distance to the nearest neighboring habitat patch) for each species. The final overall rank for each scenario was generally related to harvest intensity; the scenario with the smallest total area of even-aged management ranked the highest. Ranks were not consistent across all response variables. Relative species sensitivity was also evaluated, and the ranks did not match expectations, with the more habitat generalist species showing the highest sensitivity and the most specialist species showing the lowest. The approach here provides a means of projecting and comparing potential long-term impacts of alternative landscape strategies on diverse wildlife habitats. These results, when considered with budget considerations and species’ habitat area and population goals, can assist local managers and stakeholders in conservation planning by identifying tradeoffs and compromises aimed at optimizing protection for a variety of target species.     

‘Mitigation banks’ for wetland conservation: a major success or an unmitigated disaster?

First developed in the USA in the early 1970s, ‘wetland mitigation banks’ provide a framework for conservation activities that are designed to offset residual, unavoidable damage to the natural environment caused by development activities. The concept is now a worldwide phenomenon. In this paper I consider the level of success of wetland mitigation banks in the USA for biodiversity conservation with a view to informing ‘best practice’ in Australia. I conclude that although the concept has merit, even in the USA where the processes have been evolving for over 30 years, the outcomes frequently fall short of the target of a ‘like for like’ swap of habitat. While the outcome for wetland mitigation may not be an ‘unmitigated disaster’ it is, at best, apparently only modestly successful.     

Drivers and socioeconomic impacts of tourism participation in protected areas

Nature-based tourism has the potential to enhance global biodiversity conservation by providing alternative livelihood strategies for local people, which may alleviate poverty in and around protected areas. Despite the popularity of the concept of nature-based tourism as an integrated conservation and development tool, empirical research on its actual socioeconomic benefits, on the distributional pattern of these benefits, and on its direct driving factors is lacking, because relevant long-term data are rarely available. In a multi-year study in Wolong Nature Reserve, China, we followed a representative sample of 220 local households from 1999 to 2007 to investigate the diverse benefits that these households received from recent development of nature-based tourism in the area. Within eight years, the number of households directly participating in tourism activities increased from nine to sixty. In addition, about two-thirds of the other households received indirect financial benefits from tourism. We constructed an empirical household economic model to identify the factors that led to household-level participation in tourism. The results reveal the effects of local households' livelihood assets (i.e., financial, human, natural, physical, and social capitals) on the likelihood to participate directly in tourism. In general, households with greater financial (e.g., income), physical (e.g., access to key tourism sites), human (e.g., education), and social (e.g., kinship with local government officials) capitals and less natural capital (e.g., cropland) were more likely to participate in tourism activities. We found that residents in households participating in tourism tended to perceive more non-financial benefits in addition to more negative environmental impacts of tourism compared with households not participating in tourism. These findings suggest that socioeconomic impact analysis and change monitoring should be included in nature-based tourism management systems for long-term sustainability of protected areas.