森林的风/雪灾害研究综述

风／雪灾害不仪极大影响木材生产,同时对森林生态系统的稳定性也造成很大影响。森林风／雪危害的主要类型有树干弯曲、干(冠)折、掘根以及后续危害等;其发生主要依赖于气象条件、立地因子、树木和林分特征及其之间的相互作用。其中．林木尖削度(胸径／树高)和林分结构特征(树种、组成、密度等)是控制树木和林分对风／雪荷载抵抗的主要特征量。因此．通过造林、调整林分结构．加强林分管理如间伐、施肥等措施一直是用来减少林木的风／雪灾害的主要措施。另外．林木或林分发生风／雪害的模型分析研究也取得了很大进展,但由于森林风／雪害受诸如地形、天气等多种因素影响、目前所建立的模型系统在实际应用中普适性较芹。通过综述以往研究结果认为：在气象和立地条件难以控制的情况下．通过改变可控因子林分结构来减少森林风／雪害是可行的。因此．研究如何加强森林经营管理,尤其是不同形式的间伐技术和不同处理的造林措施与风／雪灾害发生的关系、如何增加林木和林分抵抗风／雪灾害的能力等是今后该研究领域的重点和难点。与此同时．应加强风／雪灾害危险率评估研究．进而对森林进行风／雪灾害危险率管理;并注重对受灾前后林地内生态效应的研究,以便为灾后的森林经营管理和调控提供坚实的理论依据。     

Effects of sika deer on tree seedlings in a warm temperate forest on Yakushima Island,Japan

High-density herbivore species often play an important role in forest regeneration. Native sika deer (Cervus nippon yakushimae) inhabit a high density (51.5–63.8head/km², estimated by a pellet count method) area in the western part of a lowland natural forest on Yakushima Island, Japan. To test experimentally the impact of sika deer on the mortality and the survivability of current-year seedlings, which are at a more vulnerable stage than the later stages, we constructed fenced exclosures, planted seeds of nine sapfruit tree species and compared the mortality and the survivability of current-year seedlings between fenced and unfenced quadrats. Large seeded species had significantly greater survivability in fenced quadrats than in unfenced quadrats. However, the survivability disagreed with feeding preferences. Sika deer activity increased seedling mortality of large-seeded species more than that of small-seeded species, and did not decrease much seedling survivability of not-preferred species. We found that the physical disturbance by the high density of sika deer resulted in mortality for both preferred and not-preferred species, and that deer herbivory was important for preferred species.     

Land abandonment and changes in snow cover period accelerate range expansions of sika deer

Ongoing climate change and land‐use change have the potential to substantially alter the distribution of large herbivores. This may result in drastic changes in ecosystems by changing plant–herbivore interactions. Here, we developed a model explaining sika deer persistence and colonization between 25 years in terms of neighborhood occupancy and habitat suitability. We used climatic, land‐use, and topographic variables to calculate the habitat suitability and evaluated the contributions of the variables to past range changes of sika deer. We used this model to predict the changes in the range of sika deer over the next 100 years under four scenario groups with the combination of land‐use change and climate change. Our results showed that both climate change and land‐use change had affected the range of sika deer in the past 25 years. Habitat suitability increased in northern or mountainous regions, which account for 71.6% of Japan, in line with a decrease in the snow cover period. Habitat suitability decreased in suburban areas, which account for 28.4% of Japan, corresponding to land‐use changes related to urbanization. In the next 100 years, the decrease in snow cover period and the increase in land abandonment were predicted to accelerate the range expansion of sika deer. Comparison of these two driving factors revealed that climate change will contribute more to range expansion, particularly from the 2070s onward. In scenarios that assumed the influence of both climate change and land‐use change, the total sika deer range increased by between +4.6% and +11.9% from the baseline scenario. Climate change and land‐use change will require additional efforts for future management of sika deer, particularly in the long term.     

Suppression of Regeneration in New Zealand Mountain Beech Forests is Dependent on Species of Introduced Deer

We compared the impacts on forest regeneration of introduced sika (Cervus nippon) and red (Cervus elaphus) deer in New Zealand. Plot data were used to compare mountain beech (Nothofagus solandri var. cliffortioides) regeneration between a region with sika deer, and four regions without sika deer. All regions surveyed had red deer present. In the region where sika deer had been present for more than a decade, there was evidence of poor mountain beech seedling regeneration. In the four regions without sika deer, there was evidence of a strong regenerative response at stands with low occupancy by trees. When compared to larger deer species, sika deer have a digestive morphology allowing greater dietary versatility, which may result in them impeding forest regeneration where red deer do not. In contrast to mountain beech, some small-leaved shrub species may have been competitively advantaged by intensive browsing from sika deer. This is contra to a current view that small-leaved shrub species with interlacing branches were able to tolerate browsing from extinct ratite birds, but not introduced deer. Sika deer have been introduced into countries where other deer species are indigenous, such as Canada, Denmark, Great Britain, Czech Republic, Ireland and the United States of America. Because of their dietary advantage, sika deer may have a greater potential to impede forest regeneration and competitively exclude larger deer species, particularly at low basal area sites where impacts on tree regeneration are likely to be greatest.     

Theileria infection with severe anemia and unhealed fracture in a sika deer Cervus nippon aplodontus (Cervidae: Cetartiodactyla)

An adult female sika deer (Cervus nippon aplodontus) inhabiting Nara Park, Nara, Japan, had broken bone injuries from a car accident. During its treatment, we found that the sika deer had severe anemia and the fracture remained unhealed throughout. Peripheral blood smear revealed piroplasms in the erythrocytes, which were identified as merozoites of undescribed Theileria species, widely found in sika deer in Japan. This is the report of a clinical case of Theileria infection, accompanied by severe anemia in a sika deer.     

台风“布拉万”对东北近天然落叶松云冷杉试验林的影响

台风是重要的森林干扰因子之一,会对森林生态系统的结构和功能产生较大的影响。2012年的台风"布拉万"对我国东北地区局部森林造成了严重的破坏。以受灾最重的吉林省汪清林业局的近天然落叶松云冷杉林为对象,采用方差分析和相关分析方法,研究林分结构和地形条件对林木株数损伤率的影响。结果表明:(1)林木损伤类型可分为折断、连根拔起、搭挂、压弯4种,其中连根拔起为最主要的损伤类型,占总损伤株数的52%,台风灾害造成的林木株数损伤率平均为14.09%。(2)径级大小对林木株数损伤率的影响显著。损伤主要发生于径级较小林分处,径级越大,其株数损伤率越小。(3)林木株数损伤率随林分密度的增加有减小的趋势,但在统计学上它们的关系不显著。(4)不同树种间的林木株数损伤率差异显著,落叶松、冷杉等针叶树种损伤株数最多。(5)林分的树种多样性指数与林木株数损伤率无显著的相关性。(6)海拔、坡度和坡位对林木株数损伤率的影响不显著,但坡向的影响显著,东北坡向林分的林木株数损伤率最大。研究结果可以为灾后森林恢复和减少风灾影响的森林培育措施提供依据。     

Female Sexual Preferences Toward Conspecific and Hybrid Male Mating Calls in Two Species of Polygynous Deer, <Emphasis Type="Italic">Cervus elaphu</Emphasis>s and <Emphasis Type="Italic">C. nippon</Emphasis>

The behavioral processes at the basis of hybridization and introgression are understudied in terrestrial mammals. We use a unique model to test the role of sexual signals as a reproductive barrier to introgression by investigating behavioral responses to male sexual calls in estrous females of two naturally allopatric but reproductively compatible deer species, red deer and sika deer. Previous studies demonstrated asymmetries in acoustic species discrimination between these species: most but not all female red deer prefer conspecific over sika deer male calls while female sika deer exhibit no preference differences. Here, we extend this examination of acoustic species discrimination to the role of male sexual calls in introgression between parent species and hybrids. Using two-speaker playback experiments, we compared the preference responses of estrous female red and sika deer to male sexual calls from conspecifics versus red × sika hybrids. These playbacks simulate early secondary contact between previously allopatric species after hybridization has occurred. Based on previous conspecific versus heterospecific playbacks, we predicted that most female red deer would prefer conspecific calls while female sika deer would show no difference in their preference behaviors toward conspecific and hybrid calls. However, results show that previous asymmetries did not persist as neither species exhibited more preferences for conspecific over hybrid calls. Thus, vocal behavior is not likely to deter introgression between these species during the early stages of sympatry. On a wider scale, weak discrimination against hybrid sexual signals could substantially contribute to this important evolutionary process in mammals and other taxa.     

Appropriate vegetation indices for measuring the impacts of deer on forest ecosystems

Increasing deer density can cause serious degradation of forests in the Americas, Europe, and Asia. To manage deer impacts, evaluating their current impacts on forest ecosystems is necessary, usually via vegetation indices. However, the relationship between vegetation indices and absolute deer density, while taking into account tree size, snow depth, light condition, and the type of understory vegetation, has never been investigated. We examined the relationship between various vegetation indices and absolute deer density in 344 study plots in the deciduous broad-leaved forest of Yamanashi Prefecture, central Japan. In each plot, debarking and browsing, along with the coverage and maximum height of understory vegetation, were surveyed. Estimated deer densities for 82 5 × 5-km mesh units ranged from 0.8 deer/km² to 32.7 deer/km². The percentages of debarked trees within a plot ranged from 0 to 84%. Debarking was promoted by high deer density, small tree size, and thick snow. The effect of tree size on debarking was stronger than that of deer density. Occurrence of browsing on understory vegetation was higher at higher deer densities, and where understory vegetation was dominated by evergreen dwarf bamboo. Coverage and maximum height of understory vegetation were unaffected by deer density but increased with canopy openness and the dominance of dwarf bamboo in the understory. Overall, we predict that debarking of small trees living in heavy snow areas should occur even at low deer densities (<10 deer/km²). Browsing on dwarf bamboo should occur at intermediate deer densities (10–30 deer/km²), while debarking of thick trees living in low snow areas should occur only at high deer densities (≥30 deer/km²). Our study shows that debarking and browsing on understory vegetation are appropriate indices for evaluating deer impacts on forest ecosystems, but that tree size, snow depth, and the type of understory vegetation should also be considered.     

Bottleneck effects on the sika deer Cervus nippon population in Hokkaido, revealed by ancient DNA analysis

The population size of the sika deer Cervus nippon on Hokkaido Island of Japan had been remarkably reduced because of heavy hunting pressure since the beginning of Meiji Period and effects of heavy snow in 1879 and 1881. After that, the number of sika deer in Hokkaido has increased gradually due to the protection by the Hokkaido government. In the present study, in order to investigate the bottleneck effects, we analyzed ancient mitochondrial DNA (mtDNA) on sika deer bones excavated from archaeological sites just before Meiji Period. On 86 of 113 bones from 13 archaeological sites of Ainu Culture Period (17-19th centuries), 602 base-pair fragments of the mtDNA control region were successfully sequenced. Consequently, we found three new haplotypes (g-, h- and i-types) which had not been identified in modern sika deer. In addition, four haplotypes (a-, b-, c- and d-types) identified from modern sika deer were also found in the archaeological deer. The new haplotypes and previously reported hapoltypes from sika deer of Hokkaido were phylogenetically much closer to each other, compared with those of modern sika deer from Honshu, Kyushu and the Chinese continent. Geographical distribution patterns of haplotypes of the ancient population were different from those of the modern population in Hokkaido. Our findings indicated that their genetic diversity was reduced through the bottleneck and that population structures of sika deer were changed widely in Hokkaido due to genetic drift.     

辽东山区天然次生林雪／风灾害成因及分析

对2003年春发生在辽东山区的森林雪/风灾害的成因、过程、受灾情况、造成的危害、与林分结构特征的关系以及对未来次生林生态系统的影响等进行了调查分析。结果表明,雪/风的发生是在一个大的降水天气过程基础上,由于气温的异常变化形成的,受灾严重区多分布于海拔高、坡度大,林型比较单一的桦树、柞树、色树、胡桃树和杨树等林分,林分密度和受灾率及土层厚度和受害株数均呈显著的线性负相关;受灾数量与径级和树高分别呈指数负相关和指数正相关,同时探讨了雪/风害对天然次生林生态系统内病虫害发生、林下植被、生境因子和建群种变化产生的可能影响。     

Molecular detection of tick-borne protozoan parasites in sika deer (Cervus nippon) from western regions of Japan

The sika deer (Cervus nippon) is one of the most common species of wildlife in Japan. This study aimed to reveal the prevalence of tick-borne protozoan parasites in wild sika deer living in western Japan. We used nested polymerase chain reaction (PCR) to detect the 18S rRNA gene of tick-borne apicomplexan parasites (Babesia, Theileria, and Hepatozoon spp.) from 276 blood and liver samples from sika deer captured in the Yamaguchi, Oita, Kagoshima, Okayama, Ehime, Kochi, and Tokushima Prefectures. In total, 259 samples (259/276; 93.8%) tested positive in the nested PCR screening. Gene sequencing revealed that 99.6% (258/259) of positive samples contained Theileria sp. (sika 1), while Theileria sp. (sika 2), another Theileria species, was detected in only 3 samples. We also found that one sample from a sika deer captured in Kagoshima contained the gene of an unidentified Babesia sp. related to Babesia sp. Kh-Hj42, which was previously collected from tick in western Siberia. In conclusion, we found a high prevalence of piroplasms in sika deer from western Japan, and DNA analysis revealed that Theileria sp. (sika 1) had the highest infection rate.     

黄泥河自然保护区狍冬季卧息地选择

2009年12月到2010年1月,在黄泥河自然保护区采用样线法对狍冬季卧息地选择进行研究.在研究中共设置了47条样线,调查了72个狍利用样方和109个对照样方,评价了15类生态因子对狍冬季卧息地选择的影响.研究结果表明:在黄泥河自然保护区冬季,狍在卧息时喜欢选择平均海拔在591 m,位于阳坡中坡位上雪被较浅、食物丰富度、灌丛盖度、郁闭度和隐蔽水平都较高的针阔混交林生境,尤其喜欢在针阔混交林中的针叶树下卧息,避开选择阴坡、针叶林和裸岩.逻辑斯蒂回归分析结果表明:食物丰富度、针叶树、雪深、裸岩和海拔是影响黄泥河自然保护区冬季狍卧息地选择的主导因子,林型、坡向和隐蔽水平是次要因子.由这7个变量组成的回归模型为:Z=32.628+11.675×坡向(1) +9.741×坡向(2)-5.486×林型(1)-7.933×林型(2)-7.496×裸岩(1)-9.906×针叶树(1)-0.043×海拔+0.170×隐蔽水平+0.220×食物丰富度-0.429×雪深.模型选择利用概率为P(z)=ez/1+ez,整体正确预测率为96.1％.     

东北虎豹国家公园梅花鹿的空间遗传格局及其影响因素

东北梅花鹿是东北虎豹国家公园主要的大型食草动物之一,是东北虎豹的主要猎物,对针阔混交林群落的维持有关键的作用,探究其遗传多样性及空间遗传格局对东北梅花鹿的保护以及国家公园生态系统的健康至关重要。在国家公园珲春保护区内,通过非损伤方法获得遗传样本,利用微卫星标记,研究该梅花鹿种群的空间遗传格局及其影响因素。结果表明：本研究区梅花鹿种群平均期望杂合度为0.721,遗传多样性较为丰富。有限的扩散能力常常导致种群在遗传距离上具有显著的空间自相关模式,本研究区梅花鹿种群在0-1km距离等级内在遗传距离上具有显著的空间自相关现象,据此可推测,该地区梅花鹿扩散距离为1km左右。STRUCTURE分析表明,珲春地区梅花鹿种群不存在明显的遗传分化。各种空间变量可以显著影响物种的遗传分化。本研究选取海拔、坡度、坡向、地表起伏率、人类干扰5个变量,研究其对梅花鹿种群遗传结构的影响,这5个变量多被认为与大中型哺乳动物扩散阻碍相关。依据5个变量建立了336个阻力模型,并进行偏曼特尔检验。其中,依据海拔、坡向、地表起伏率、人类干扰假设建立的246个阻力模型与遗传距离之间的关系并不显著,综合所有变量的15个生境适宜性模型阻力模型与遗传距离的关系也都不显著。在依据坡度假设建构的75个阻力模型中,只有1个模型与遗传距离有显著的正相关关系,该模型同时也是在控制空间自相关影响后,在所有模型中与遗传距离相关性最高的模型。根据该模型推测,最适宜梅花鹿扩散的坡度为10°,梅花鹿可能倾向于利用缓坡进行扩散。结果对东北虎豹国家公园梅花鹿种群的保护具有重要意义。     

Landscape structure affects food quality of sika deer (<Emphasis Type="Italic">Cervus nippon</Emphasis>) evidenced by fecal nitrogen levels

Evaluating the quality of wildlife habitat is essential for understanding and predicting population dynamics in heterogeneous environments. We used fecal nitrogen levels as an indicator of habitat quality of sika deer (Cervus nippon) and explored important landscape elements influencing nitrogen levels, taking deer density into account. We established 92 plots differing in deer density and landscape structure on the Boso Peninsula, central Japan, and collected fecal samples along a 1-km transect at each plot. The regression models involving two independent variables, i.e., deer density and the length of forest edge within an area of 100 or 200 m from the transect, were selected based on the Akaike’s Information Criterion (AIC). Levels of fecal nitrogen were positively correlated with the length of the forest edge and negatively correlated with population density of deer. The area of 100 or 200 m from the transect most likely reflected the behavioral scale of the deer. Coverage of palatable understory vegetation increased with proximity to forest edge and decreased with deer density. Variability in the level of fecal nitrogen could thus be explained by food availability in the landscape. These results suggest that landscape alterations increase the carrying capacity of sika deer and thereby increase impacts upon the ecosystem.     

Forest edge creates small-scale variation in reproductive rate of sika deer

Habitat edges are considered to have an important role in determining the abundance of deer in forest landscapes, but to our knowledge there are few lines of evidence indicating that forest edge enhances the vital rate of deer. We examined pregnancy of female sika deer in Boso peninsula, central Japan, and explored how forest edges, food availability in forests, and local population density influence the pregnancy rate of sika deer. Local deer density was estimated by the number of fecal pellets, and food availability in forests was estimated by combining GIS data of vegetation distribution and the relationship between vegetation biomass and local deer density. Forest edge length was also determined by GIS data. Model selection was performed with multiple logistic regression analyses using the AIC to find the best model for accounting for the observed variation in pregnancy rates of the deer. Multiple logistic regression analysis showed that the length of forest edge had a positive effect on the pregnancy rate of females, whereas food availability in forests and local deer density had little effect. This forest edge effect was detected in a 100–200-m radius from deer captured locations, indicating that deer pregnancy is primarily determined by habitat quality within a 10-ha area. This result was confirmed by tracking females with GPS telemetry, which found that the core areas of the home range were less than 12 ha. The positive effect of edges and the lack of density dependence could be a result of high plant productivity in open environments that produces forages not depleted by high deer densities. Our results support the view that land management is the cause of the current problem of deer overabundance. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Red and sika deer in the British Isles,current management issues and management policy

The red deer (Cervus elaphus) is one of the most widely distributed species of deer in Europe. Due to its economic value as game species or its negative impacts on forestry, agriculture and conservation areas, most populations are currently managed, with strategies and intensity of the management varying between countries. In Britain, and less certainly in Ireland, red deer have been continuously present since the end of the last glaciation and constitute the largest population of red deer in Europe. Although they thrived in the past when forests were abundant, the current distribution of red deer in the British Isles is uneven, with the largest numbers being found in Scotland and few and more localised populations in England, Wales and the Republic of Ireland. In the British Isles, as in many other parts of Europe, there is a long history of man interacting with deer populations including local extinctions, multiple translocations and introductions of exotic species of deer. Among introduced exotic species of deer, the Japanese sika (Cervus nippon) is the one of most concern. After introduction of small numbers at multiple locations in Britain and Ireland from 1860 onwards, sika have increased in population number and range in areas with good forest cover, and where they overlap with red deer there is a risk of hybridisation. Due to recent increases in numbers and range of red and sika deer, both species pose a range of management challenges which are not easy to solve. In this review we summarise the history and status of these two species in Great Britain and Ireland, describe current management and discuss management options for the future.     

Tree and stand-scale factors affecting richness and composition of epiphytic bryophytes and lichens in deciduous woodland key habitats

Conservation and sustainable forestry are essential in a multi-functional landscape. In this respect, ecological studies on epiphytes are needed to determine abiotic and biotic factors associated with high diversity. The aim of the present study was to evaluate relative sensitivity of conservation targets (epiphytic bryophytes and lichens) in relation to contrasting environmental variables (tree species, tree diameter at breast height, bark crevice depth, pH, tree inclination, pH, forest stand age, area and type) in boreo-nemoral forests. The study was conducted in Latvian 34 woodland key habitat (WKH) boreo-nemoral forest stands. Generalized linear mixed models and canonical correspondence analysis showed that tree species and tree bark pH were the most important variables explaining epiphytic bryophyte and lichen composition and richness (total, Red-listed, WKH indicator species). Forest stand level factors, such as stand size and habitat type, had only minor influence on epiphytic species composition and richness. The results of the present study indicate a need to maintain the diversity of tree species and large trees, particularly Acer platanoides, Carpinus betulus, Fraxinus excelsior, Populus tremula, Tilia cordata, Ulmus glabra and Ulmus laevis in conservation of epiphytic bryophyte and lichen communities in the future.     

Wind damage and response in New Zealand forests: a review

The literature on wind damage in New Zealand forests is reviewed to investigate how abiotic and biotic factors influence damage severity, damage type, and forest recovery. Winds that damage forests tend to result from extra-tropical depressions or from topographically enhanced westerly air flows. Severe wind damage can occur when wind speeds exceed c. 0 km/hr, although investigating the relationship between damage and wind speeds is difficult, as gusts, for which speed is usually unrecorded, are important. Damage is often quantified by estimates of area affected, with some authors detailing the size and species of damaged trees within a given area. Key abiotic factors that influence damage patterns are topographical position, edaphic conditions, and disturbance history. Important biotic factors are tree height, tree health, position of the tree within the stand, and species. Damage type (uprooting or breakage) is primarily controlled by canopy position and rooting depth. Forest responses to wind damage include sprouting, recruitment, release, and suppression, with the dominant mode of forest recovery being strongly influenced by the severity of damage, and the species composition of the stand. As noted in international literature on wind damage, a lack of consistent methods, combined with poor species and spatial coverage, makes identifying general trends difficult. Investigating the role of wind damage in New Zealand forests has focused to date on Nothofagus forests and plantations of exotic trees and few studies have investigated long term dynamics following wind disturbance events.     

Evaluation of introgressive hybridization among Cervidae in Japan's Kinki District via two novel genetic markers developed from public NGS data

Hybridization and backcrossing of native populations with introduced species can lead to introgression and genetic alteration. In this study, we evaluated introgression in 43 deer from a potential hybrid zone around Okinoshima Island, Kinki District, Japan. This region witnessed the migration of a hybrid population (cross between the Formosan sika deer [Cervus nippon taiouanus] and other deer species) that could potentially breed with the native Japanese sika deer (C. n. centralis). We used an existing genetic marker for the mitochondrial cytochrome b gene and two novel markers for nuclear DNA, developed using publicly available next‐generation sequencing data. We identified one mainland deer with a mitochondrial haplotype identical to that of the Formosan sika deer as well as nuclear heterozygous sequences identical to those of Formosan and Japanese sika deer. This suggests that the mainland deer is a hybrid offspring of the Okinoshima population and native deer. However, only Japanese sika deer sequences were found in the other 42 samples, indicating limited introgression. Nevertheless, hybridization pre‐ and postintroduction in the Okinoshima population could cause multispecies introgression among Japanese sika deer, negatively affecting genetic integrity. We developed a simple test based on polymerase chain reaction–restriction fragment length polymorphism to detect introgression in natural populations. Our method can accelerate genetic monitoring of Japanese sika deer in Kinki District. In conclusion, to prevent further introgression and maintain genetic integrity of Japanese sika deer, we recommend establishing fences around Okinoshima Island to limit migration, besides a continued genetic monitoring of the native deer.     

黄泥河自然保护区狍冬季栖息地选择

2009年12月—2010年1月,在黄泥河自然保护区采用样线法对狍冬季栖息地选择进行了研究。共设置了47条样线,调查了109个狍利用样方和109个对照样方,测定了海拔、坡度、坡向、坡位、隐蔽级、郁闭度、食物丰富度、灌丛盖度、树胸径、人为干扰距离、林型、裸岩、雪深、倒木和针叶树15类栖息地因子。Baileys法和Mann-WhiteyU检验表明:在黄泥河自然保护区冬季,狍喜欢在坡度平缓、平均海拔602m,位于阳坡或半阴半阳坡的中坡位上雪被较浅,食物丰富度、灌丛盖度和隐蔽级都较高的针阔混交林生境中活动,对倒木因子表现出明显的选择性,避开选择阴坡、裸岩和针叶林生境。在主成分分析中,前5个主成分中的累积贡献率达到88.6%,第一主成分贡献率为30.8%,主要反映利用样方的林型、海拔和隐蔽级的影响,第二主成分贡献率为26.5%,主要反映利用样方的食物丰富度和灌丛盖度影响。