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.     

Molecular analysis of dispersal in giant pandas

Although dispersal in the giant panda (Ailuropoda melanoleuca) is a demographic mechanism which can potentially counteract the negative effect of habitat fragmentation, little is known about dispersal in this species because of difficulties in observing individuals. Using data from faecal microsatellite genotyping, we compared the spatial distribution of giant pandas in two populations and the proximity of relatives in one key population to infer their dispersal pattern. We conclude that giant pandas exhibit female-biased dispersal because: (i) vAIc (variance of assignment index) for females was significantly larger than for males, suggesting that females comprise both 'local' and 'foreign' genotypes; (ii) the average spatial distance of related female dyads was significantly larger than that of males; (iii) larger r (relatedness), F(ST) (genetic variance among populations) and mAIc (mean of assignment index) values were found in males using the software FSTAT, although the differences were not significant; (iv) males set up territories neighbouring to their birth place; (v) significant population structure using microsatellites with a concomitant lack of mitochondrial structure was found in a previous study, possibly indicating more extensive female dispersal; and (vi) female-biased dispersal was strongly supported by evidence from concomitant ecological studies. Considering previous ecological data and life-history characteristics of the giant panda, female-biased dispersal is most likely to be due to competition for birth dens among females, inbreeding avoidance and enhancing inclusive fitness among related males.     

Synergistic effects of anthropogenic disturbances on giant pandas

Livestock grazing and the collection of bamboo shoots are the main threats to giant panda (Ailuropoda melanoleuca) habitat in the Liangshan Mountains in China. It is important to clarify the effect of these disturbances to the giant panda to formulate targeted management policies. Based on species distribution models and daily activity models, we investigated the effects of livestock grazing and bamboo shoot collection on giant pandas from May 2021 to July 2022. Our results indicated the giant panda's suitable habitat in the reserve covered 51.83 km² (15.02% of the reserve area). Grazing and bamboo shoot collection led to losses of 19.08 km² and 7.68 km² of suitable habitat, respectively. Together, the 2 activities resulted in a loss of 28.35 km² of suitable habitat, which was more than half of the area of panda habitat. The areas of suitable habitat for giant pandas significantly overlapped with the areas affected by both disturbances. Giant pandas did not show significant differences in daily activity rhythms under a single disturbance, but the daily activity rhythms of giant pandas differed when we compared the area combining the 2 disturbances with the undisturbed area. Our study reveals that the anthropogenic disturbances in the reserve have varying effects on the suitable habitat range and daily activity rhythm of giant pandas and evidence of a synergistic effect. Therefore, when formulating relevant conservation policies, it is important to fully evaluate the extent and characteristics of anthropogenic disturbances in shaping the population distribution and habitat preferences of the giant panda and other wildlife to enhance the efficacy of conservation management practices.     

四川省大熊猫保护地生态安全及其时空演变

大熊猫是生态保护的旗舰物种,保护大熊猫及其栖息地具有重要的示范作用和科学价值。中国致力于建设以国家公园为主体的自然保护地体系,分析大熊猫保护地生态安全,为建立大熊猫国家公园提供理论依据,具有重要意义。以四川省为例,选取被划入或将来可能被划入大熊猫保护地的典型县域,构建大熊猫保护地生态安全评价指标体系,采用熵权法、层次分析法进行综合评价,并结合空间相关性分析,从时间和空间两个维度对大熊猫保护地生态安全状况进行时空动态分析。结果表明：（1）2003-2017年四川省大熊猫保护地生态安全综合指数呈波动性上升趋势,生态环境改善显著,但仍有部分县域出现不同程度的环境恶化现象。（2）各县域生态安全状况表现出较强的差异性,并且相邻县域生态安全指数存在显著的空间正相关性和空间集聚性。（3）大熊猫保护地生态安全与周边社会经济发展之间矛盾依然尖锐,相邻县域之间的经济发展和生态保护状况相互影响的作用明显,需联合治理,共同促进生态平衡,提高大熊猫及其栖息地保护效果。     

大熊猫与同域动物在海拔分布上的生态位分化

大熊猫（Ailuropoda melanoleuca）是中国的特有物种,其成功放归可能受放归地气候、栖息地、同域动物等多种因素的影响。本研究在野外调查的基础上,基于生态位理论等方法,探讨了栗子坪国家级自然保护区大熊猫与其大中型同域动物的空间关系。调查结果表明,该保护区内大熊猫同域动物共20种,分别属于2纲5目,其中东洋型分布物种占优势（30%）。大熊猫同域动物分布海拔显著低于大熊猫的分布海拔（P＜0.05）,大熊猫分别与灵长目等5个目的动物在海拔分布上均存在显著的生态位分化（P＜0.001）。灵长目动物在空间分布上与大熊猫分布相异,而其他目的动物与大熊猫空间分布类似。偶蹄目动物痕迹数量占比最高（54.14 %）,而食肉目的小熊猫与大熊猫样线共同遇见率达到了43.75%。本研究结果表明,该地大熊猫同域动物较为丰富,在海拔分布上与大熊猫存在生态位分化。该研究可为圈养大熊猫的放归提供参考依据。     

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 Assessment for Giant Pandas in the Qinling Mountain Region of China

ABSTRACT Because habitat loss and fragmentation threaten giant pandas (Ailuropoda melanoleuca), habitat protection and restoration are important conservation measures for this endangered species. However, distribution and value of potential habitat to giant pandas on a regional scale are not fully known. Therefore, we identified and ranked giant panda habitat in Foping Nature Reserve, Guanyinshan Nature Reserve, and adjacent areas in the Qinling Mountains of China. We used Mahalanobis distance and 11 digital habitat layers to develop a multivariate habitat signature associated with 247 surveyed giant panda locations, which we then applied to the study region. We identified approximately 128 km²of giant panda habitat in Foping Nature Reserve (43.6% of the reserve) and 49 km²in Guanyinshan Nature Reserve (33.6% of the reserve). We defined core habitat areas by incorporating a minimum patch-size criterion (5.5 km²) based on home-range size. Percentage of core habitat area was higher in Foping Nature Reserve (41.8% of the reserve) than Guanyinshan Nature Reserve (26.3% of the reserve). Within the larger analysis region, Foping Nature Reserve contained 32.7% of all core habitat areas we identified, indicating regional importance of the reserve. We observed a negative relationship between distribution of core areas and presence of roads and small villages. Protection of giant panda habitat at lower elevations and improvement of habitat linkages among core habitat areas are important in a regional approach to giant panda conservation.     

Population genetics reveals high connectivity of giant panda populations across human disturbance features in key nature reserve

The giant panda is an example of a species that has faced extensive historical habitat fragmentation, and anthropogenic disturbance and is assumed to be isolated in numerous subpopulations with limited gene flow between them. To investigate the population size, health, and connectivity of pandas in a key habitat area, we noninvasively collected a total of 539 fresh wild giant panda fecal samples for DNA extraction within Wolong Nature Reserve, Sichuan, China. Seven validated tetra‐microsatellite markers were used to analyze each sample, and a total of 142 unique genotypes were identified. Nonspatial and spatial capture–recapture models estimated the population size of the reserve at 164 and 137 individuals (95% confidence intervals 153–175 and 115–163), respectively. Relatively high levels of genetic variation and low levels of inbreeding were estimated, indicating adequate genetic diversity. Surprisingly, no significant genetic boundaries were found within the population despite the national road G350 that bisects the reserve, which is also bordered with patches of development and agricultural land. We attribute this to high rates of migration, with four giant panda road‐crossing events confirmed within a year based on repeated captures of individuals. This likely means that giant panda populations within mountain ranges are better connected than previously thought. Increased development and tourism traffic in the area and throughout the current panda distribution pose a threat of increasing population isolation, however. Maintaining and restoring adequate habitat corridors for dispersal is thus a vital step for preserving the levels of gene flow seen in our analysis and the continued conservation of the giant panda meta‐population in both Wolong and throughout their current range.     

No evidence for MHC‐based mate choice in wild giant pandas

Major histocompatibility complex genes (MHC), a gene cluster that controls the immune response to parasites, are regarded as an important determinant of mate choice. However, MHC‐based mate choice studies are especially rare for endangered animals. The giant panda (Ailuropoda melanoleuca), a flagship species, has suffered habitat loss and fragmentation. We investigated the genetic variation of three MHC class II loci, including DRB1, DQA1, and DQA2, for 19 mating‐pairs and 11 parent‐pairs of wild giant pandas based on long‐term field behavior observations and genetic samples. We tested four hypotheses of mate choice based on this MHC variation. We found no supporting evidence for the MHC‐based heterosis, genetic diversity, genetic compatibility and “good gene” hypotheses. These results suggest that giant pandas may not use MHC‐based signals to select mating partners, probably because limited mating opportunities or female‐biased natal dispersal restricts selection for MHC‐based mate choice, acknowledging the caveat of the small sample size often encountered in endangered animal studies. Our study provides insight into the mate choice mechanisms of wild giant pandas and highlights the need to increase the connectivity and facilitate dispersal among fragmented populations and habitats.     

Black and white and read all over: the past,present and future of giant panda genetics

Few species attract much more attention from the public and scientists than the giant panda (Ailuropoda melanoleuca), a popular, enigmatic but highly endangered species. The application of molecular genetics to its biology and conservation has facilitated surprising insights into the biology of giant pandas as well as the effectiveness of conservation efforts during the past decades. Here, we review the history of genetic advances in this species, from phylogeny, demographical history, genetic variation, population structure, noninvasive population census and adaptive evolution to reveal to what extent the current status of the giant panda is a reflection of its evolutionary legacy, as opposed to the influence of anthropogenic factors that have negatively impacted this species. In addition, we summarize the conservation implications of these genetic findings applied for the management of this high‐profile species. Finally, on the basis of these advances and predictable future changes in genetic technology, we discuss future research directions that seem promising for giant panda biology and conservation.     

野生大熊猫种群数量调查方法研究进展

大熊猫(Ailuropodamelanoleuca)是中国特有珍稀野生动物,被誉为野生动物保护领域的"旗舰物种"。大熊猫种群数量的变化可以实时反应大熊猫种群动态,提供大熊猫分布区域、栖息地质量等最直接的信息,是制定大熊猫保护方案的基础,也是有效实施大熊猫保护措施的前提。综述了几种传统野生大熊猫种群数量调查方法(包括直接计数法、数学模型法、距离-咬节分析法及分子生物学方法),以及近年来最新应用于野生动物种群数量调查的红外相机技术、足迹鉴定法,讨论了传统方法中可能存在的问题、分析新方法的应用前景,并针对今后的野生大熊猫种群数量调查提出了一些建议与展望。     

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.     

Impact of livestock on giant pandas and their habitat

Livestock production is one of the greatest threats to biodiversity worldwide. However, impacts of livestock on endangered species have been understudied, particularly across the livestock–wildlife interface in forested protected areas. We investigated the impact of an emerging livestock sector in China's renowned Wolong Nature Reserve for giant pandas. We integrated empirical data from field surveys, remotely sensed imagery, and GPS collar tracking to analyze (1) the spatial distribution of horses in giant panda habitat, (2) space use and habitat selection patterns of horses and pandas, and (3) the impact of horses on pandas and bamboo (panda's main food source). We discovered that the horse distribution overlapped with suitable giant panda habitat. Horses had smaller home ranges than pandas but both species showed similarities in habitat selection. Horses consumed considerable amounts of bamboo, and may have resulted in a decline in panda habitat use. Our study highlights the need to formulate policies to address this emerging threat to the endangered giant panda. It also has implications for understanding livestock impacts in other protected areas across the globe.     

汶川地震对大熊猫栖息地的影响与恢复对策

 大熊猫是生物多样性保护的旗舰种, 保护大熊猫及其栖息地是保护生物多样性和生态系统功能完整性与稳定性的重要保障体现。汶川地震灾区位于大熊猫重点分布区岷山-邛崃山, 地震及其次生灾害导致该区27个大熊猫自然保护区不同程度受损, 8.3%的大熊猫栖息地因地震而被破坏。地震及其次生灾害对大熊猫栖息地的影响主要表现在: 1) 地震埋没和砸毁大熊猫赖以生存的主食竹, 地震可能诱发主食竹开花, 威胁到大熊猫的健康和食物安全; 2) 地震及其诱发的土壤和山石运动显著影响森林的动态特征, 森林大面积丧失或质量下降; 3) 地震改变大熊猫栖息地生境特征, 大熊猫个体交流的廊道阻断, 形成“生殖孤岛”, 遗传多样性降低, 栖息地破碎化进程加快。应对震后大熊猫栖息地恢复的主要对策有: 1) 重新评估震后大熊猫栖息地质量, 并重新规划现有大熊猫保护区群的布局; 2) 应用地理信息系统、遥感及数学模型等手段与野外实地实证研究相结合的方法, 全面查清震后大熊猫栖息地主食竹资源状况及分布规律并及时监测其动态, 复壮更新大熊猫主食竹; 3) 利用天然植被自然恢复和人工重建等措施恢复因地震而退化或丧失的大熊猫栖息地。     

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.     

Inbreeding and inbreeding avoidance in wild giant pandas

Inbreeding can have negative consequences on population and individual fitness, which could be counteracted by inbreeding avoidance mechanisms. However, the inbreeding risk and inbreeding avoidance mechanisms in endangered species are less studied. The giant panda, a solitary and threatened species, lives in many small populations and suffers from habitat fragmentation, which may aggravate the risk of inbreeding. Here, we performed long‐term observations of reproductive behaviour, sampling of mother–cub pairs and large‐scale genetic analyses on wild giant pandas. Moderate levels of inbreeding were found in 21.1% of mating pairs, 9.1% of parent pairs and 7.7% of panda cubs, but no high‐level inbreeding occurred. More significant levels of inbreeding may be avoided passively by female‐biased natal dispersal rather than by breeding dispersal or active relatedness‐based mate choice mechanisms. The level of inbreeding in giant pandas is greater than expected for a solitary mammal and thus warrants concern for potential inbreeding depression, particularly in small populations isolated by continuing habitat fragmentation, which will reduce female dispersal and increase the risk of inbreeding.     

Back-and-forth shifts in habitat selection by giant pandas over the past two decades in the Daxiangling Mountains,southwestern China

Habitat selection is a complex process, and animals may change their habitat selection over time. However, we have a poor understanding of temporal variation in habitat selection of endangered giant pandas (Ailuropoda melanoleuca), which has limited the development of conservation strategies for them. This study examined giant panda habitat selection in three stages of the past two decades (2001–2002, 2011–2012, and 2019–2020) in the Daxiangling Mountains, southwestern China. We applied the resource selection function to estimate the probability of species occurrence with five limiting habitat factors, i.e., elevation, slope, aspect, tree diameter at breast height, and bamboo cover. We found that giant pandas consistently selected for south- and west-facing slopes with mid-elevations and large trees in the three stages. However, there were shifts back-and-forth in the choice between higher and lower elevations and between higher and lower bamboo cover across the three stages. We suggest that the giant panda habitat selection had important consequences for gaining the high availability of palatable bamboo and good shelters. The back-and-forth shifts might be the result of changes in the type, spatial distribution, and intensity of human disturbance. Our study provides insights for improving our understanding and approaches for quantifying temporal variation in habitat selection in giant pandas and other species, suggesting that adaptive conservation and management in response to changing environments is needed.     

圈养繁殖在大熊猫保护生物学中的作用

栖息地的日益恶化、片段化和种群孤岛化已成为保护大熊猫的严重障碍。尽管就地保护濒危物种是最有希望的措施,但对于大熊猫而言,研究表明圈养繁殖也是保护这一极危物和中的有效手段之一,对保护遗传多样性起以了重要作用,是延缓绝灭速率的保证。本文阐述了大圈养繁殖的必要性及其在保护生物学中的作用。     

Impacts of temperature on giant panda habitat in the north Minshan Mountains

Understanding the impacts of meteorological factors on giant pandas is necessary for future conservation measures in response to global climate change. We integrated temperature data with three main habitat parameters (elevation, vegetation type, and bamboo species) to evaluate the influence of climate change on giant panda habitat in the northern Minshan Mountains using a habitat assessment model. Our study shows that temperature (relative importance = 25.1%) was the second most important variable influencing giant panda habitat excepting the elevation. There was a significant negative correlation between temperature and panda presence (ρ = −0.133, P < 0.05), and the temperature range preferred by giant pandas within the study area was 18–21°C, followed by 15–17°C and 22–24°C. The overall suitability of giant panda habitats will increase by 2.7%, however, it showed a opposite variation patterns between the eastern and northwestern region of the study area. Suitable and subsuitable habitats in the northwestern region of the study area, which is characterized by higher elevation and latitude, will increase by 18007.8 hm² (9.8% habitat suitability), while the eastern region will suffer a decrease of 9543.5 hm² (7.1% habitat suitability). Our results suggest that increasing areas of suitable giant panda habitat will support future giant panda expansion, and food shortage and insufficient living space will not arise as problems in the northwest Minshan Mountains, which means that giant pandas can adapt to climate change, and therefore may be resilient to climate change. Thus, for the safety and survival of giant pandas in the Baishuijiang Reserve, we propose strengthening the giant panda monitoring program in the west and improving the integrity of habitats to promote population dispersal with adjacent populations in the east.     

Application of ecological-niche factor analysis in habitat assessment of giant pandas

Ecological-niche factor analysis (ENFA) is a multivariate approach to study geographic distribution of species on a large scale with only “presence” data. It has been widely applied in many fields including wildlife management, habitat assessment and habitat prediction. In this paper, this approach was applied in habitat suitability assessment for giant pandas in Pingwu County, Sichuan Province, China. With “presence” data of giant pandas and remote sensing data, habitat suitability of pandas in this county was evaluated based on ENFA model, and spatial distribution pattern of nature reserves and conservation gaps were then evaluated. The results show that giant pandas in this county prefer high-elevation zones (> 2128 m) dominated by coniferous forest, and mixed coniferous and deciduous broadleaf forest, and avoid deciduous broadleaf forest and shrubs. Pandas avoid staying at habitats with human disturbances. Farmland is a major factor threatening panda habitat. Panda habitat is mainly distributed in north and west of Pingwu with a total area of 234033 hm², 106345hm² for suitable habitat and 127688 hm² for marginally suitable habitat). 3 nature reserves were located in Pingwu, covering over 49.2% of total suitable habitat and 45.6% of total marginally suitable habitat. Although 47.2% of panda habitat was in reserves under protection, connectivity between reserves was weak and a conservation gap existed in the north part of Pingwu. Thus, a new nature reserve in Baima and Mupi should be established to link the isolated habitats.