Ochotona Curzoniae Research Articles

The complete mitochondrial genome of Daurian pika (Ochotona dauurica)

In this study, we sequenced and uploaded the complete mitochondrial genome of Daurian pika (Ochotona dauurica) for the first time using tissue obtained from a wild female sample captured from Inner Mongolia, China. This mitochondrial genome is a circular molecule of 17,293 bp in length, consisting of two ribosomal RNA genes, 22 transfer RNA genes, 13 protein-coding genes, 1 control region, and 1 rep_origin region. The phylogenetic analysis basis of 12 protein-coding genes except for ND6 gene shows that Ochotona dauurica clusters with Ochotona curzoniae.

Functional-trait ecology of the plateau pika Ochotona curzoniae in the Qinghai-Tibetan Plateau ecosystem.

Understanding a species' functional traits allows for a directed and productive perspective on the role a species plays in nature, and thus its relative importance to conservation planning. The functional-trait ecology of the plateau pika Ochotona curzoniae is examined to better understand the resilience and sustainability of the high alpine grasslands of the Qinghai-Tibetan Plateau (QTP). The key functional traits of plateau pikas are their abundance and behavior of digging extensive burrow systems. Plateau pikas have been poisoned over a significant part of their original geographic distribution across the QTP, allowing comparison of ecological communities with and without pikas. Nearly all mammalian and avian carnivores, most of which are obligate predators on pikas, have been lost in regions where pikas have been poisoned. Most endemic birds on the QTP nest in pika burrows; when pikas are poisoned, burrows collapse, and these birds are greatly reduced in number. Due to the biopedturbation resulting from their burrows, regional plant species richness is higher in areas with pikas than without. The presence of pika burrows allows higher rates of infiltration during heavy monsoon rains compared to poisoned areas, possibly mitigating runoff and the potential for serious downslope erosion and flooding. Thus, the functional traits of plateau pikas enhance native biodiversity and other important ecosystem functions; these traits are irreplaceable. As plateau pikas are not natural colonizers, active re-introduction programs are needed to restore pikas to areas from which they have been poisoned to restore the important functional ecological traits of pikas.

Dynamics of a predator-prey model with state-dependent carrying capacity

Vegetation and plateau pika are two key species in alpine meadow ecosystems on the Tibetan Plateau. It is frequently observed on the field that plateau pika reduces the carrying capacity of vegetation and the mortality of plateau pika increases along with the increasing height of vegetation. This motivates us to propose and study a predator-prey model with state-dependent carrying capacity. Theoretical analysis and numerical simulations show that the model exhibits complex dynamics including the occurrence of saddle-node bifurcation, transcritical bifurcation and Hopf bifurcation, and the coexistence of two stable equilibria.

Genetic Diversity in Echinococcus multilocularis From the Plateau Vole and Plateau Pika in Jiuzhi County, Qinghai Province, China.

The Qinghai-Tibet Plateau is a highly endemic area of alveolar echinococcosis where a series of intermediate hosts, especially voles and pikas, are infected with Echinococcus multilocularis. The metacestodes of E. multilocularis are fluid-filled, asexually proliferating cysts, and they are mainly found in the host's liver in the form of tumor-like growths. In this study, we investigated the genetic variations of E. multilocularis in four mitochondrial (mt) genes, namely, NADH dehydrogenase subunit 5 (nad5), adenosine triphosphate subunit 6 (atp6), cytochrome c oxidase subunit 1 (cox1), and NADH dehydrogenase subunit 1 (nad1). The complete nad5, atp6, cox1, and nad1 genes were amplified separately from each hydatid cyst isolate using polymerase chain reaction (PCR) and then sequenced. Phylogenetic trees were then generated based on the combined mt genes using MrBayes 3.1.2 and PAUP version 4.0b10. The results showed that thirty of 102 voles and two of 49 pikas were infected with E. multilocularis. The genetic variation distances among all E. multilocularis samples were 0.1–0.4%, 0.2–0.4%, 0.1–0.6%, and 0.1–0.4% for nad5, atp6, nad1, and cox1, respectively. Compared to previous studies of the genetic diversity of E. multilocularis based on the cox1 gene, the genetic distances within the same group were 1.3–1.7% (Mongolia strain), 0.6–0.8% (North American strain), 0.3–0.6% (European strain), and 0.1–0.4% (Asian strain). Based on concatenated sequences of the nad5, atp6, cox1, and nad1 genes all haplotypes were divided into two clusters. In conclusion, the genetic diversity of E. multilocularis based on mt genes on a small local area is at low level but between different regions with long distance and different ecological environment each other, the genetic diversity is at relatively high level; genetic variation is higher in the nad1 gene than that in the other three mt genes. The results on a local scale provide basic information for further study of the molecular epidemiology, genetic differences and control of E. multilocularis in Qinghai Province, China.

Testosterone-retinoic acid signaling directs spermatogonial differentiation and seasonal spermatogenesis in the Plateau pika (Ochotona curzoniae)

During evolution, animals optimize their reproductive strategies to increase offspring survival. Seasonal breeders reproduce only during certain times of the year. In mammals, reproduction is tightly controlled by hypothalamus-pituitary-gonad axis. Although pathways regulating gametogenesis in non-seasonal model species have been well established, molecular insights into seasonal reproduction are severely limited. Using the Plateau pika (Ochotona curzoniae), a small rodent animal species native to the Qinghai-Tibetan plateau, as a model, here we report that seasonal spermatogenesis is governed at the level of spermatogonial differentiation. In testis of the reproductively dormant animals, undifferentiated spermatogonia failed to differentiate and accumulated in the seminiferous tubules. RNA-seq analyses of the active and dormant testes revealed that genes modulating retinoic acid biogenesis and steriodogenesis were differentially regulated. A single injection of all-trans retinoic acid (ATRA) reinitiated spermatogenesis and inhibition the function of RA-degrading enzyme CYP26B1 for 10 days induced spermatogonial differentiation. Strikingly, testosterone injection reinitiated spermatogenesis in short day adapted animals. Testosterone provides a permissive environment of RA biogenesis and actions in testis, therefore, indirectly controls spermatogonial differentiation. Collectively, these findings provide a key mechanistic insight regarding the molecular regulation of seasonal reproduction in mammals.

Impact of plateau pikas (Ochotona curzoniae) on soil properties and nitrous oxide fluxes on the Qinghai-Tibetan Plateau

This paper demonstrates the impact of an endemic fossorial animal, plateau pika (Ochotona curzoniae), on soil properties and N2O flux at the Zoige Wetland. Pika burrow and control sites without disturbance by pika were selected to measure the soil water content, bulk density, soil organic matter (SOM), NH4-N content and NO3-N content in August 2012. N2O fluxes were measured with static opaque chambers at these sites in June and August 2012. Pika burrowing altered soil aeration by transferring deeper soil to the surface and by constructing underground burrows, which significantly increased bulk density, and reduced soil water content, SOM and NH4-N content at 0–10 cm and 10–20 cm soil depth. N2O flux had a significant correlation with bulk density, SOM and NH4-N content. Pika burrowing significantly influenced N2O flux by increasing N2O flux at the control site from near zero to 0.063±0.011 mg m-2 h-1. Our findings described how pika burrowing influences the soil traits and significantly increases the principal greenhouse gas N2O emission. As plateau pika was commonly considered as a pest, our findings give a novel clue to effectively manage populations of plateau pika on the Qinghai-Tibet Plateau from the perspective of greenhouse gas emission.

Independence between coping style and stress reactivity in plateau pika

The concept of coping style represents the way individual animals react to a stressful situation, both behaviourally and neurophysiologically. Over the last decades coping style has been linked to the development of research on animal personality. Based on this concept, we should find a proactive-reactive continuum in animal populations, with proactive individuals being fast explorer, bold, aggressive, and show high sympathetic reactivity (higher heart rate), as well as low hypothalamus-pituitary-adrenocortical (PHA) axis reactivity to external stressor (higher plasma glucocorticoid level). At the other extreme, shy, lowly aggressive, reactive individuals should be slow in their exploration, and show a low sympathetic reactivity and a high HPA axis reactivity. However, a recent two-tier model proposed that coping style and stress reactivity should be independent of each other. In this study, we tested the two-tier model in a wild plateau pika (Ochotona curzoniae) population on the Tibetan Plateau, by quantifying the associations between several behavioural and physiological traits at the among- and within-individual levels. We repeatedly measured exploration, docility, boldness, heart rate and plasma cortisol concentration in individuals between April and September of 2013. All traits tested were repeatable. At the among-individual level, all behavioural traits were correlated with each other and with heart rate, but were independent of both basal level and variation of plasma cortisol concentration. Most correlations were negligible at the within-individual level. In support of the two-tier model, these results suggest that coping style (i.e. behaviour and heart rate associations) is independent of stress reactivity (i.e. glucocorticoid reactivity) in that species.

Diet simplification selects for high gut microbial diversity and strong fermenting ability in high-altitude pikas.

The gut microbiota in mammals plays a key role in host metabolism and adaptation. However, relatively little is known regarding to how the animals adapts to extreme environments through regulating gut microbial diversity and function. Here, we investigated the diet, gut microbiota, short-chain fatty acid (SCFA) profiles, and cellulolytic activity from two common pika (Ochotona spp.) species in China, including Plateau pika (Ochotona curzoniae) from the Qinghai-Tibet Plateau and Daurian pika (Ochotona daurica) from the Inner Mongolia Grassland. Despite a partial diet overlap, Plateau pikas harbored lower diet diversity than Daurian pikas. Some bacteria (e.g., Prevotella and Ruminococcus) associated with fiber degradation were enriched in Plateau pikas. They harbored higher gut microbial diversity, total SCFA concentration, and cellulolytic activity than Daurian pikas. Interestingly, cellulolytic activity was positively correlated with the gut microbial diversity and SCFAs. Gut microbial communities and SCFA profiles were segregated structurally between host species. PICRUSt metagenome predictions demonstrated that microbial genes involved in carbohydrate metabolism and energy metabolism were overrepresented in the gut microbiota of Plateau pikas. Our results demonstrate that Plateau pikas harbor a stronger fermenting ability for the plant-based diet than Daurian pikas via gut microbial fermentation. The enhanced ability for utilization of plant-based diets in Plateau pikas may be partly a kind of microbiota adaptation for more energy requirements in cold and hypoxic high-altitude environments.

Echinococcus multilocularis and Echinococcus shiquicus in a small mammal community on the eastern Tibetan Plateau: host species composition, molecular prevalence, and epidemiological implications

BackgroundThe eastern part of the Tibetan Plateau is now recognized as an endemic region with the highest reported human infection rates in the world of human alveolar echinococcosis (AE) caused by Echinococcus multilocularis. Existing epidemiological studies on AE have mainly focused on the synanthropic environment, while basic parasitological and ecological aspects in wildlife host species remain largely unknown, especially for small mammal hosts. Therefore, we examined small mammal host species composition, occurrence, and the prevalence of both E. multilocularis and E. shiquicus in Shiqu County (Sichuan Province, China), eastern Tibetan Plateau.ResultsIn total, 346 small mammals from five rodent and one pika species were trapped from four randomly set 0.25 ha square plots. Two vole species, Lasiopodomys fuscus (n = 144) and Microtus limnophilus (n = 44), and the plateau pika (Ochotona curzoniae) (n = 135), were the three most-dominant species trapped. Although protoscoleces of E. multilocularis and E. shiquicus were only observed in L. fuscus and O. curzoniae, respectively, cox1 and nad1 gene DNA of E. shiquicus was detected in all the small mammal species except for Neodon irene, whereas E. multilocularis was detected in the three most-dominant species. The overall molecular prevalence of Echinococcus species was 5.8 (95% CI: 3.3–8.2%) ~ 10.7% (95% CI: 7.4–14.0%) (the conservative prevalence to the maximum prevalence with 95% CI in parentheses), whereas for E. multilocularis it was 4.3 (95% CI: 2.2–6.5%) ~ 6.7% (95% CI: 4.0–9.3%), and 1.5 (95% CI: 0.2–2.7%) ~ 4.1% (95% CI: 2.0–6.1%) for E. shiquicus. The prevalence of both E. multilocularis and E. shiquicus, was significantly higher in rodents (mainly voles) than in pikas. Phylogenetic analyses revealed that Echinococcus haplotypes of cox1 from small mammal hosts were actively involved in the sylvatic and anthropogenic transmission cycles of E. multilocularis in the eastern Tibetan Plateau.ConclusionsIn contrast to previous studies, the current results indicated that rodent species, rather than pikas, are probably more important natural intermediate hosts of E. multilocularis and E. shiquicus in the eastern Tibetan Plateau. Thus, understanding interspecific dynamics between rodents and pikas is essential to studies of the echinococcosis transmission mechanism and human echinococcosis prevention in local communities.

Stochastic processes govern bacterial communities from the blood of pikas and from their arthropod vectors.

Vector-borne microbes influence pathogen transmission and blood microbiomes, thereby affecting the emergence of infectious diseases. Thus, understanding the relationship between host and vector microbiomes is of importance. In this study, we investigated the bacterial community composition, diversity and assembly of the flea (Rhadinopsylla dahurica vicina), torsalo (Hypoderma curzonial), and the blood and gut of their shared pika host, Ochotona curzoniae. Bartonella, Sphingomonas and Bradyrhizobium were enriched in blood, while Wolbachia and Fusobacterium were more abundant in fleas and torsaloes. Most of potential pathogenic microbes (belonging to Fusobacterium, Rickettsia, Kingella, Porphyromonas, Bartonella and Mycoplasma) were present in the blood of pikas and their vectors. Blood communities were more similar to those from fleas than other sample types and were independent of host factors or geographical sites. Notably, blood microbes originate mainly from fleas rather than gut or torsaloes. Interestingly, the community assembly of blood, fleas or torsaloes was primarily governed by stochastic processes, while the gut microbiome was determined by deterministic processes. Ecological drift plays a dominant role in the assembly of blood and flea microbiomes. These results reflect the difficulty for predicting and regulating the microbial ecology of fleas for the prevention of potential microbiome-associated diseases.

Effects of plateau pika disturbance levels on the plant diversity and biomass of an alpine meadow

AbstractThe plateau pika (Ochotona curzoniae) is known to influence the plant diversity and biomass of the alpine meadow, and it is regarded as a pest. The species has been subject to extensive controls without a clear target density in the Qinghai‐Tibetan Plateau. We investigated the effects of different plateau pika disturbance levels on plant composition, species diversity and biomass. The density of active burrow entrances was used as an index of disturbance levels I, II, III and IV (corresponding to 8, 19, 27 and 39 active burrow entrances per 625 m2). We found that the plant community composition differed with different disturbance levels. The plant replacements were mainly related to associate plants rather than dominant plants. With the increase in the plateau pika disturbance levels, the plant cover decreased, and the plant height first increased and then decreased. The plant evenness index was higher in the levels II and III disturbance conditions, while the plant diversity index and plant richness index were highest in the level IV disturbance condition. The disturbance levels had no effect on the total plant biomass; however, the intermediate disturbance levels (II and III) increased the palatable plant biomass. The total plant biomass was mainly dependent on the unpalatable plant biomass. The total plant biomass showed a positive correlation with the richness index and a negative correlation with the evenness index. The palatable plant biomass showed a positive correlation with the evenness index and a negative correlation with the richness index. These results imply that the intermediate disturbance levels are beneficial to livestock production as they increase the alpine meadow quality, and the higher disturbance level is better for plant diversity conservation in the alpine meadows of the Qinghai‐Tibetan Plateau.

Chronic cold exposure results in subcutaneous adipose tissue browning and altered global metabolism in Qinghai-Tibetan plateau pika (Ochotona curzoniae)

The plateau pika (Ochotona curzoniae), one of the indigenous animals of the Qinghai-Tibet Plateau, is adapted to life in a cold and hypoxic environment. We conducted a series of genomic, proteomic and morphological studies to investigate whether changes in energy metabolism contribute to adaptation of the plateau pika to cold stress by analyzing summer and winter cohorts. The winter group showed strong morphological and histological features of brown adipose tissue (BAT) in subcutaneous white adipose tissue (sWAT). To obtain molecular evidence of browning of sWAT, we performed reverse transcription and quantitative real-time PCR, which revealed that BAT-specific genes, including uncoupling protein 1 (UCP-1) and PPAR-γ coactivator 1α (PGC-1α), were highly expressed in sWAT from the winter group. Compared with the summer group, Western blot analysis also confirmed that UCP-1, PGC-1α and Cox4 protein levels were significantly increased in sWAT from the winter group. Increased BAT mass in the inter-scapular region of the winter group was also observed. These results suggest that the plateau pika adapts to cold by browning sWAT and increasing BAT in order to increase thermogenesis. These changes are distinct from the previously reported adaptation of highland deer mice. Understanding the regulatory mechanisms underlying this adaptation may lead to novel therapeutic strategies for treating obesity and metabolic disorders.

The first report of Cryptosporidium spp. in Microtus fuscus (Qinghai vole) and Ochotona curzoniae (wild plateau pika) in the Qinghai-Tibetan Plateau area, China.

Cryptosporidium is one of the most important genera of intestinal zoonotic pathogens, which can infect various hosts and cause diarrhoea. There is little available information about the molecular characterisation and epidemiological prevalence of Cryptosporidium spp. in Microtus fuscus (Qinghai vole) and Ochotona curzoniae (wild plateau pika) in the Qinghai-Tibetan Plateau area of Qinghai Province, Northwest China. Therefore, the aim of this study was to determine Cryptosporidium species/genotypes and epidemiological prevalence in these mammals by detecting the SSU rRNA gene by PCR amplification. The Cryptosporidium spp. infection rate was 8.9% (8/90) in Qinghai voles and 6.25% (4/64) in wild plateau pikas. Positive samples were successfully sequenced, and the following Cryptosporidium species were found: C. parvum, C. ubiquitum, C. canis and a novel genotype in Qinghai voles and C. parvum and a novel genotype in wild plateau pikas. This is the first report of Cryptosporidium infections in M. fuscus and wild O. curzoniae in Northwest China. The results suggest the possibility of Cryptosporidium species transmission among these two hosts, the environment, other animals and humans and provide useful molecular epidemiological data for the prevention and control of Cryptosporidium infections in wild animals and the surrounding environments. The results of the present study indicate the existence of Cryptosporidium species infections that have potential public health significance. This is the first report of Cryptosporidium multi-species infections in these animal hosts.

The disturbance and disturbance intensity of small and semi-fossorial herbivores alter the belowground bud density of graminoids in alpine meadows

Do semi-fossorial herbivores influence the trajectory of plant community succession by altering belowground bud density in alpine meadows? Field surveys were carried out to investigate the effect of disturbance and disturbance intensity of the plateau pika (Ochotona curzoniae) on the belowground bud density in alpine meadows at two sites. The Linear Mixed Model (LMM) test and Linear Model (LM) test were used to analyze the relative effect of plateau pika disturbance on bud density and to clarify the response of belowground bud density to disturbance intensity. Our results showed that disturbance by the plateau pika increased belowground bud density of graminoids and plants from the grass and sedge families, including Kobresia pygmaea and Elymus nutans, but had no effect on forbs bud density. The bud density of graminoids, grass and sedge plants, and K. pygmaea and E. nutans showed a downward parabolic trend in response to increased disturbance intensity of the plateau pika, implying that there was an optimal level of disturbance intensity that maximized belowground bud density of graminoids. These results suggest that disturbance by the plateau pika has different effects on belowground bud densities of graminoids versus forbs. Additionally, disturbance by plateau pika up to the optimal intensity may improve grazing quality of alpine meadow in the long term through increasing graminoid bud density, resulting in a higher proportion of graminoids in pasture, while higher levels of disturbance by plateau pika may deteriorate alpine meadows through a reduction in graminoid bud density.

Effects of plateau pika (Ochotona curzoniae) on alpine meadow phytocoenosium and analysis of the strategy it uses to expand habitat

High-density plateau pikas (Ochotona curzoniae) disturbance is one of the main causes of alpine meadow degradation. The response of phytocoenosium to the disturbance of plateau pika may reflect its habitat expanding strategy. We used quadrat sampling method to investigate vegetation height, coverage, and aboveground biomass of non-pika area (NA), transition area (TA), and pika-active area (AA) distributed continuously in alpine meadow located in Guoluo Tibetan Autonomous County of Qinghai Province in 2011 and 2012. The results showed that, from NA to AA (i) vegetation coverage, height, and above-ground biomass decreased (P<0.01). (ii) While the number of plant species decreased from 41 to 30, and species diversity decreased significantly (P<0.05), evenness of the alpine meadow showed no significant variations. (iii) Dominant species changed from grasses to weeds. (iv) While plants in TA showed no obvious difference in height between 2011 and 2012, coverage reduced significantly (P<0.05). These results demonstrate that (i) plateau pikas disturbance on alpine meadow will change plant community structure and composition, directly leading to degeneration of alpine meadow. Additionally, (ii) plateau pikas expand their habitat by altering plant coverage firstly.

Microclimate response of soil to plateau pika's disturbance in the northeast Qinghai‐Tibet Plateau

SummaryThe plateau pika (Ochotona curzoniae) is one of the main native soil faunas on the Qinghai‐Tibet Plateau and plays a key role in the terrestrial ecosystem there. To understand how and why the soil microclimate changes after the plateau pika's disturbance, this study measured soil, vegetation, hydrologic and thermal properties and investigated soil moisture and soil temperature dynamics from 2014 to 2015 at the plot scale of four types of land surface: original grassland, new mound, old mound and bald patch. Our results showed that the average coefficients of surface runoff of original grassland, new mound, old mound and bald patch were 0.6, 3.0, 4.3 and 10.0%, respectively (P &lt; 0.05). Evapotranspiration was largest for original grassland, especially under wet conditions, and was similar among the other three types of land surface during the growing season (P &lt; 0.01). During varying precipitation events, the soil moisture content of new mound increased first, followed by old mound, original grassland and bald patch. Meanwhile, the increasing magnitude of soil moisture content had the same order. At the seasonal timescale, soil moisture content at 5‐cm depth was largest for old mound and smallest for bald patch (P &lt; 0.01). The mean daily soil temperature at 5‐cm depth of new mound was approximately 0.8°C higher than that of old mound (P &lt; 0.01) because of the smaller land surface reflectance of the former. The daily range of soil temperature at 5‐cm depth of original grassland and bald patch was about 2.7 and 4.7°C higher than the average value of new and old mounds, respectively, because of the larger soil thermal conductivity of the former two. A conceptual framework is suggested in this study to synthesize the evolution of soil microclimate under disturbance by the plateau pika. Overall, results indicated that new and old mounds accelerate soil hydrologic processes and have a better soil temperature buffer.Highlights Change in soil microclimate of different types of land surface under disturbance by the plateau pika. New and old mounds accelerate soil hydrologic processes and have a better soil temperature buffer. New and old mounds provide advantageous environments for plateau pika. Bald patch had the smallest soil moisture content and largest daily range of soil temperature.

Summer habitat use of plateau pikas (Ochotona curzoniae) in response to winter livestock grazing in the alpine steppe Qinghai-Tibetan Plateau

ABSTRACT Plateau pikas (Ochotona curzoniae), burrowing lagomorphs endemic to the Qinghai-Tibetan Plateau (QTP), are considered pests by Chinese authorities because of their association with grasslands that are characterized as degraded. Officials typically blame pikas for causing the degraded conditions, whereas studies increasingly suggest that sparse vegetation encourages population growth and high densities of pikas. Correlational investigations that document pika density while simultaneously describing grassland conditions can quantify this association, but are uninformative regarding causation. We used livestock exclosures and pika reductions in partially controlled experiments to examine how pika habitat use responds to changes in vegetation structure caused by wintertime livestock grazing. Mean counts of pikas were higher on grazed than ungrazed plots. Linear models accounting for experimental pika reduction as well as seasonal phenology indicated that grazing significantly increased pika use compared with ungrazed plots. These results show that moderate livestock density is likely consistent with both biodiversity conservation and economic activity, and that high pika populations alone are not responsible for observed degradation of the QTP alpine grasslands. Moderate levels of livestock grazing and pika presence are consistent with maintaining the integrity of the alpine steppe ecosystem of the QTP.

Response of leaf traits of common plants in alpine meadow to plateau pika disturbance

Leaf traits have been proven to reflect the adaptation of individual plants to disturbance environments in a grassland ecosystem. A field survey was conducted to investigate the effects of the disturbance intensity of plateau pika on the leaf traits of a dominant (Kobresia pygmaea) and two common plants (Elymus nutans and Anemone rivularis var. flore-minore) in an alpine meadow. This study indicated that the plateau pika disturbance enables the individuals of three plants to exhibit respective plasticity because the three plants had different leaf indices (LI) as the disturbance intensity increased. K. pygmaea, E. nutans and A. rivularis var. flore-minore had high specific leaf area (SLA), leaf dry mass content (LDMC), and leaf nitrogen content (LNC) at relatively low, moderate, and high disturbance intensities of plateau pika, respectively. K. pygmaea, E. nutans and A. rivularis var. flore-minore suffered low nutrient stress at low, moderate and high disturbance intensities due to high N : P at corresponding disturbance intensities. These results indicated that K. pygmaea, E. nutans and A. rivularis var. flore-minore grew well at relatively low, moderate, and high disturbance intensity conditions, respectively, which contributed to the improvement of alpine meadows with a higher proportion of E. nutans at a moderate disturbance intensity or the deterioration of alpine meadows with a higher proportion of A. rivularis var. flore-minore at a high disturbance intensity. Our findings suggest that leaf traits are effective tools to explain how small burrowing herbivore disturbances often lead to the improvement or deterioration of alpine meadows under different disturbance intensities.

Architecture Characteristics of Burrow System of Plateau Pika, Ochotona curzoniae

Burrows serve many functions for animals by providing a place for rearing young, sleeping, hibernation, food storage, protection from predators and extreme environment, and the function of burrows varies with the degree of burrow complexity. We studied the burrow architecture (length, internal dimensions, fractal dimension of tunnel systems, number of nesting chambers and surface holes) by excavating the tunnels of 24 burrow systems of the plateau pika (Ochotona curzoniae). Pikas have two types of burrow systems, namely temporary and permanent burrows. The number of surface holes and length of tunnels were significantly different between the temporary and permanent burrows. Tunnel width and tunnel height were not significantly different between the temporary and permanent burrows. The permanent burrow had a nesting chamber but temporary did not have any. No hoarding chamber was found in either the permanent or the temporary burrow. The temporary burrow was used to avoid predators in emergency situations, and probably was the initial stage of construction of more complex burrows of some dispersing pikas. Permanent burrows were used for avoiding predators, resting and rearing offsprings.

A PREY-PREDATOR MODEL WITH HOLLING Ⅱ FUNCTIONAL RESPONSE AND THE CARRYING CAPACITY OF PREDATOR DEPENDING ON ITS PREY

One prey-predator model is formulated and the global behavior of its solution is analyzed. In this model, the carrying capacity of predator depends on the amount of its prey, and the Holling Ⅱ functional response is involved. This model may have four classes of positive equilibriums and limit cycle. The positive equilibriums may be stable, or a saddle-node, or a saddle, or a degenerate singular point. In alpine meadow ecosystem, the dynamics of vegetation and plateau pika can be described by this model. Through simulating with virtual parameters, the cause of alpine meadow degradation and effective recovery strategy is investigated. Increasing grazing rate or decreasing plateau pika mortality may cause alpine meadow degradation. Correspondingly, reducing grazing rate and increasing plateau pika mortality may recover the degraded alpine meadow effectively.