Vegetation types of East Ladakh: species and growth form composition along main environmental gradients

Question: The vegetation of high mountain regions in Himalaya remains poorly assessed despite the expected vulnerability of these ecosystems to global change drivers. What are the main vegetation types in East Ladakh and which environmental factors influence the species composition and growth forms distribution? Location: The south‐westernmost extension of the Tibetan Plateau, an arid mountainous area, in East Ladakh, Jammu & Kashmir State, India. Methods: Species composition was recorded for 369 plots (each 100 m²). Plots, sampled from 4180 m a.s.l. (bottom of the Indus Valley) up to 6060 m a.s.l. (close to the snowline), covered a wide range of environmental conditions. Results: TWINSPAN clusters discriminated eight ecologically interpretable vegetation types, corresponding to the main habitats in the area: animal resting places, salt marshes, semi‐deserts and steppes, shrublands, alpine screes and boulder fields, alpine grasslands, water bodies and subnival zone. The most important environmental factors influencing the species composition were altitude, soil moisture and salinity. Screes and alpine grasslands were found to be the most species‐rich. The species were ranged into 20 growth forms with regard to life‐form and clonality, with growth forms showing different changes in proportion among vegetation types and along the different environmental gradients. Conclusion: The study summarizes the main vegetation types of East Ladakh in terms of species and growth form compositions. The results can have a heuristic value for designing future monitoring schemes and assess the effects of global change in these diverse, but poorly studied, regions.     

On the patterns of abundance and diversity of macrolichens of Chopta-Tunganath in the Garhwal Himalaya

A total of 3211 colonies of macrolichens, from twelve 50 m × 10 m plots distributed across four macrohabitat (vegetation) types between 1500 m–3700 m in the Chopta-Tunganath landscape of the Garhwal Himalaya, yielded 13 families with 15 genera and 85 species.Lobaria retigera stood out as a broad-niched generalist species with moderate levels of abundance in all the three major microhabitats, viz. rock, soil and wood across 83% of all the plots sampled, whereasUmbilicaria indica emerged as an abundantly occurring specialist confined to rock substrates.Heterodermia incana andLeptogium javanicum appeared to be rare members of the community as they were encountered only once during the field survey. Woody microhabitats turned out to be richer than rock and soil substrates for macrolichens. Amongst the macrohabitats, middle altitude (2500–2800 m)Quercus forest was richest in species and genera followed by high altitude (2900–3200 m)Rhododendron forest, higher altitude grasslands (3300–3700 m) and then the lower elevation (1500 m)Quercus forest. Species, genus and family level alphaas well as beta-diversities were significantly correlated with each other, implying that higher taxonomic ranks such as genera may be used as surrogates for species thus facilitating cost- and time-effective periodic monitoring of the biodiversity of macrolichens. Dynamics of the diversity of lichen communities in relation to various forms of environmental disturbance including livestock grazing and tourism as dominant land use activities in the higher Himalaya need further research.     

A diet rich in C3 plants reveals the sensitivity of an alpine mammal to climate change

Plant–herbivore interactions provide critical insights into the mechanisms that govern the spatiotemporal distributions of organisms. These interactions are crucial to understanding the impacts of climate change, which are likely to have an effect on the population dynamics of alpine herbivores. The Royle's pika (Ochotona roylei, hereafter pika) is a lagomorph found in the western Himalaya and is dependent on alpine plants that are at risk from climate change. As the main prey of many carnivores in the region, the pika plays a crucial role in trophic interactions. We examined topographical features, plant genera presence and seasonal dynamics as drivers of the plant richness in the pika's diet across an elevational gradient (2,600–4,450 m). We identified 79 plant genera in the faecal pellets of pikas, of which 89% were forbs, >60% were endemic to the Himalaya, and 97.5% of the diet plant genera identified followed the C₃ photosynthetic pathway. We found that, during the premonsoon season, the number of genera in the pika's diet decreased with increasing elevation. We demonstrate that a large area of talus supports greater plant diversity and, not surprisingly, results in higher species richness in the pika's diet. However, in talus habitat with deep crevices, pikas consumed fewer plant genera suggesting they may be foraging suboptimally due to predation risk. The continued increase in global temperature is expected to have an effect on the distribution dynamics of C₃ plants and consequently influence pika diet and distribution, resulting in a significant negative cascading effect on the Himalayan ecosystem.     

Morphological Adaptation of Inflorescences in Plants that Develop at Low Temperatures in Early Spring: The Convergent Evolution of "Downy Plants"

Abstract: A new category of plants that exhibit convergent evolution, namely "downy plants", is described and discussed here on the bais of natural selection. So-called snowball plants can be represented by Saussurea gossypiphora D. Don (Compositae), which has extremely dense trichomes on well-developed bracts that are tightly packed around floral buds. Plants whose morphology is similar to that of S. gossypiphora are found at high elevations of alpine zones in the Nepalese Himalayas, where temperatures are low and precipitation is high (frequent rain) in summer. Nonetheless, we noticed that plants with a morphology similar to that of Himalaya snowball plants are commonly distributed from temperature to Arctic zones, and are even found in Alaska where precipitation is very limited. Willows ( Salix spp.: Salicaceae) and deciduous magnolias (Magnoliaceae) are typical examples of such plants. Measurements of temperature inside and outside the inflorescences of Salix (pussy willow or catkin) and of Magnolia suggested that the pubescent bracts might play a role in keeping the interior of buds warm, but that the effect depends on light intensity. Our examination of such species led us to extend the concept of "snowball plants" to a larger group of plants, namely "downy plants", that are characterized by very dense trichomes on tightly packed bracts of inflorescences. Downy plants are thereby considered to represent a convergent adaptation that allows blooming at low temperatures.     

Succession in a New Zealand alpine cushion community: a Markovian model

A permanent plot in a low-alpine cushion/tussock/shrub community was mapped at ca. decade intervals from 1953 to 1984. Transition probabilities between vegetation types were calculated, and a Markovian model was fitted.In spite of non-stationarity a Markov model was consistent with the observations, and produced reasonable predictions, especially for the major components of the vegetation. The model predictions were conservative in terms of future changes, in that future changes would be smaller than those observed, but such predictions could be supported by independent evidence, such as the nature of the non-stationarity, plant counts and seedling numbers.The general pattern was of a decrease in the cover of both Chionochloa rigida tussocks and mixed turf, and an increase in the cover of cushion bog species, especially Donatia novae-zelandiae. Shrubs, the suggested climax, showed no tendency to increase. Known climatic changes cannot explain the trend, which therefore seems to be a local retrogressive succession. There may be a cycle of alternating cushion and turf.     

不同氮素形态对向日葵生长和光合功能的影响

利用砂培试验研究了不同氮素形态对向日葵生长和光合功能的影响。结果表明：NO^3-N(NN)处理的株高、叶面积、干物质重显著高于NH4^ -N(AA)处理,施用NO3^--N的植株有较高的净光合速率(Pn)和较低的气孔导度(C8)、细胞间隙CO2浓度(Ci)。不同处理之间Fv／Fm没有显著差异,NN处理的植株PSⅡ线性电子传递量子效率ФPSⅡ、光化学荧光猝灭系数qP和表观光合电子传递效率ETR都显著高于AA处理的植株;而非光化学荧光猝灭系数qN则明显低于AA处理的植株。碳同化受到抑制是NH4^ -N条件下净光合速率下降的重要原因。     

Establishment and growth of Quercus floribunda seedlings after a mast year

Abstract. Population dynamics, mortality factors, growth and dry weight changes in Quercus floribunda seedlings were studied to determine the implication of sun and shade microsites and the role of mast years on regeneration and maintenance of oak forests in Kumaun Himalaya. The number of seedlings was significantly greater at the sun microsites, but seedling survival was not, as ground herbage clearance at sun microsites accounted for 53.5% mortality. Insect herbivores (e.g. acorn worm) were also one of the major mortality factors. At both microsites, most mortality was encountered between June and August. The initial seedling height at sun microsites was significantly lower compared to shade microsites. Seedlings at sun microsites were superior in terms of number of leaves and peak dry weight. Dry mass allocation in seedling components and root:shoot ratio was similar at both microsites. The study shows the importance of sun microsites and of masting in the potential for regeneration and expansion of Q. floribunda forests.     

Impact of replanting on regeneration of the medicinal plantNardostachys grandiflora DC. (Valerianaceae)

Commercial collection of alpine medicinal plants in Nepal is widely believed to destroy the resource base. This study explores the impact of three different collection methods on the regeneration of a frequently collected and traded plant species, Nardostachys grandiflora DC (Valerianaceae), listed in CITES Appendix II. The collected product is the plant rhizome from which essential oil is extracted. An experiment with 209 2X2 m plots on both south- and north-facing slopes of a valley in Gorkha District was conducted over two years. Harvesting 100% of the plants in plots followed by replanting of upper plant parts and 2 cm of the rhizome provided the fastest regeneration and rhizome biomass growth.     

Vegetation patterns,regeneration rates and divergence in an old-field succession of the high tropical Andes

Vegetation restoration during old-field succession was studied in an alpine Andean ecosystem (paramo). 123 plots with different fallow times (1 to 12 years) and 8 plots under natural vegetation were sampled. The results indicate that secondary succession in the paramo, like in other extreme environments, can be interpreted as an autosuccession: there are mainly changes in species relative abundance and little floristic relay (i.e. species turnover). Only a few herbaceous species, mostly introduced (e.g. Rumex acetosella), act as strict pioneers and strongly dominate the early stages. Then, they undergo a progressive decline, while native forbs (e.g. Lupinus meridanus) and grasses (e.g. Vulpia myuros) have their peak abundance in intermediate stages. The characteristic paramo life forms, sclerophilous shrubs (e.g. Baccharis prunifolia, Hypericum laricifolium) and giant rosettes (e.g. Espeletia schultzii), appear very early and gradually increase in abundance during succession, becoming dominant in the late stages and showing a dual behaviour, both as ruderal and stress tolerant species. The 1st axis of a Detrended Correspondence Analysis arranges the sites according to their fallow time. The 2nd and 3rd axes, associated with diverging pathways of regeneration, are correlated with topographic factors and physio-chemical soil characteristics. Hence, structural divergence between plots increases along succession as community composition starts to reflect the conditions of each site. We found evidence of a constant rate of succession during the first 12 years, contradicting the generally accepted hypothesis in the succession literature of a continous slow down up to the climax. Regeneration of vegetation physiognomy is relatively fast, questioning the prevailing idea of slow restoration in alpine ecosystems. However, 12 years of fallow are insufficient to attain the species richness of the natural paramo. Under the current trend of fallow length reduction observed in traditional potato cultivation in the Andes, our results raise doubts about the conservationist value of this management strategy.     

Postweaning mass gain in juvenile alpine marmots Marmota marmota

The effects of several environmental factors on the postweaning growth of wild Alpine marmots were investigated. Factors considered were year of birth, sun exposure in the home range, litter size, and sex of young. Components of growth were juvenile mass at emergence from the natal burrow (as a result of preweaning growth) and postweaning growth rate. We also considered the length of the active season during which growth occurs. Mass at emergence and postweaning growth rate varied according to year of birth, were higher in south-facing than in north-facing home ranges, and were higher in small litters. Mass at emergence was higher for males than for females. We suggest that environmental factors affected the juvenile growth pattern through influences on maternal body condition. Our results support Trombulak's hypothesis that mothers maintain as many young as physiologically possible. We suggest that mothers in poor condition sacrificed the mass of their offspring rather than their number. A body mass sexual dimorphism of juveniles occurred at emergence, suggesting that mothers may provide more care for their male than their female offspring. Received: 9 June 1997 / Accepted: 22 September 1997     

Vascular plant diversity and climate change in the alpine belt of the central Apennines (Italy)

The aim of this study is to analyse the vascular flora and the local climate along the altitude gradient in the largest alpine belt of the central Apennines (Majella National Park), and to contribute to the evaluation of the possible effects of global climate changes on the biodiversity of the alpine ecosystem. For this purpose floristic-quantitative analyses and temperature records on three different summits have been carried out by using the methodological protocol of the UE-GLORIA project (2001 2003); the project aims toward a standardised monitoring of flora and temperature in the alpine environment of the main European chains. From the analysis of the changes in species richness along the altitude gradient (2405 m versus 2730 m a.s.l.), it emerged that 70% of species do not reach the highest summit and only 11% of the overall flora is shared by all of the summits examined; a drop in mean temperature has been observed at soil level, along the same gradient from 3.11 to 0.03 °C. Floristic-quantitative and climatic analyses have been carried out even along the horizontal gradient (principal exposures), highlighting a great species richness and vegetation cover in eastward aspects. We singled out some endangered rare species and we proved that the slopes facing east will be the first to be affected by the coming of subalpine species from below, whereas northward exposures will be the most conservative, showing greater inertia toward the invasive process caused by the climate warming.     

Noninvasive sampling reveals population genetic structure in the Royle’s pika,Ochotona roylei,in the western Himalaya

Understanding population genetic structure of climate‐sensitive herbivore species is important as it provides useful insights on how shifts in environmental conditions can alter their distribution and abundance. Herbivore responses to the environment can have a strong indirect cascading effect on community structure. This is particularly important for Royle's pika (Lagomorpha: Ochotona roylei), a herbivorous talus‐dwelling species in alpine ecosystem, which forms a major prey base for many carnivores in the Himalayan arc. In this study, we used seven polymorphic microsatellite loci to detect evidence for recent changes in genetic diversity and population structure in Royle's pika across five locations sampled between 8 and 160 km apart in the western Himalaya. Using four clustering approaches, we found the presence of significant contemporary genetic structure in Royle's pika populations. The detected genetic structure could be primarily attributed to the landscape features in alpine habitat (e.g., wide lowland valleys, rivers) that may act as semipermeable barriers to gene flow and distribution of food plants, which are key determinants in spatial distribution of herbivores. Pika showed low inbreeding coefficients (F_IS) and a high level of pairwise relatedness for individuals within 1 km suggesting low dispersal abilities of talus‐dwelling pikas. We have found evidence of a recent population bottleneck, possibly due to effects of environmental disturbances (e.g., snow melting patterns or thermal stress). Our results reveal significant evidence of isolation by distance in genetic differentiation (F_ST range = 0.04–0.19). This is the first population genetics study on Royle's pika, which helps to address evolutionary consequences of climate change which are expected to significantly affect the distribution and population dynamics in this talus‐dwelling species.     

Effects of macrophyte growth forms on invertebrate communities in saline lakes of the Wyoming High Plains

In saline lakes, areal cover and both species and structural diversity of macrophytes often decline as salinity increases. To assess effects of the loss of certain macrophyte growth forms, we characterized benthic and epiphytic invertebrates in three growth forms (thin-stemmed emergents, erect aquatics, and low macroalgae) in oligosaline lakes (0.8–4.2 mS cm⁻¹) of the Wyoming High Plains, USA. We also measured the biomass and taxonomic composition of epiphytic and benthic invertebrates in two erect aquatics with very similar structure that are found in both oligosaline (Potamogeton pectinatus) and mesosaline (9.3–23.5 mS cm⁻¹) (Ruppia maritima) lakes. Although total biomass of epiphytic invertebrates varied among oligosaline lakes, the relative distribution of biomass among growth forms was similar. For epiphytic invertebrates, biomass per unit area of lake was lowest in emergents and equivalent in erect aquatics and low macroalgae; biomass per unit volume of macrophyte habitat was greatest in low macroalgae. For benthic invertebrates, biomass was less beneath low macroalgae than other growth forms. Taxonomic composition did not differ appreciably between growth forms for either benthic or epiphytic invertebrates, except that epiphytic gastropods were more abundant in erect aquatics. Total biomass of epiphytic and benthic invertebrates for the same growth form (erect aquatic) did not differ between oligosaline (Potamogeton pectinatus) and mesosaline (Ruppia maritima) lakes, but taxonomic composition did change. In the oligosaline to mesosaline range, direct toxic effects of salinity appeared important for some major taxa such as gastropods and amphipods. However, indirect effects of salinity, such as loss of macrophyte cover and typically higher nutrient levels at greater salinities, probably have larger impacts on total invertebrate biomass lake-wide.     

Seed Dispersal Spectrum of Woody Species in South Ecuadorian Dry Forests: Environmental Correlates and the Effect of Considering Species Abundance

This study examines the seed dispersal spectrum of the tropical dry forests of Southern Ecuador, in an effort to contribute to the knowledge of the complex dynamics of tropical dry forests. Seed dispersal spectrum was described for a total number of 160 species. Relationships of dispersal syndromes with plant growth form and climatic seasonality were explored. For a subset of 97 species, we determined whether dispersal spectrum changes when species abundance, in addition to species number, is taken into account. The same subset was used to relate dispersal syndromes with the environmental conditions. Zoochorous species dominated in the studied community. When considering the individual abundance of each species, however, anemochory was the prevalent dispersal syndrome. We found a significant difference in the frequency of dispersal syndromes among plant growth forms, with epizoochory only occurring in shrub species. The dispersal spectrum was dependent on climatic seasonality. The largest proportion of anemochorous species fructified during the dry season, while zoochorous diaspores dominated during the rainy season. A fourth‐corner analysis indicated that the seed dispersal spectrum of Southern Ecuador dry forests is controlled by environmental conditions such as annual precipitation, annual temperature range or topography. Our results suggest that spatio‐temporal changes in the environmental conditions may affect important ecological processes for dispersal. Thus, the predominance of one syndrome or another may depend on the spatial variation of environmental conditions. Abstract in Spanish is available at http://www.blackwell‐synergy.com/loi/btp .     

Plant height as a simple predictor of the root to shoot ratio: Evidence from alpine grasslands on the Tibetan Plateau

Question: Since increases in altitude and grazing intensity generally result in decreases in height growth of alpine grasslands, plant height may integrate effects of environmental stress and grazing disturbance and provide better assessments of the variation in root: shoot (R: S) biomass ratio than other variables. However, it is unclear if there is a general relationship between plant height and R:S ratio across grassland ecosystems. Such knowledge would be helpful for root biomass estimation in grasslands. Location: An altitudinal transect in the Gonghe Basin (2880–4040 m a.s.l.), northeast Tibetan plateau. Methods: We measured standing biomass both above‐ground and below‐ground, maximum plant height (MPH) and soil variables across 43 plots. Results: Climatic variables explained the variations in MPH and R: S ratio of undegraded grasslands better than soil variables (46–50% vs < 19%), while those of degraded grasslands generally showed insignificant correlations with climatic and soil variables. There was a general relationship between R: S ratio and MPH (negative, R²= 0.76, P< 0.001) across degraded and undegraded grasslands. The relationship was used to predict R: S ratio in 13 additional plots in steppe grasslands of Inner Mongolia, and good agreement of expected and observed values has been found (R²= 0.87, P < 0.001). Conclusions: MPH, that is relatively easy to measure, can be used to predict R:S ratio at plot to regional scales. It is promising to develop a new method for large‐scale estimation of root biomass in grasslands using MPH and shoot biomass avoiding tedious procedures of physical measuring of above and below‐ground biomass.     

Effect of temperature on the progamic phase in high-mountain plants

Progamic processes are particularly temperature-sensitive and, in lowland plants, are usually drastically reduced below 10 °C and above 30 °C. Little is known about how effectively sexual processes of mountain plants function under the large temperature fluctuations at higher altitudes. The present study examines duration and thermal thresholds for progamic processes in six common plant species (Cerastium uniflorum, Gentianella germanica, Ranunculus alpestris, R. glacialis, Saxifraga bryoides, S. caesia) from different altitudinal zones in the European Alps. Whole plants were collected from natural sites shortly before anthesis and kept in a climate chamber until further processing. Flowers with receptive stigmas were hand-pollinated with allopollen and exposed to controlled temperatures between -2 and 40 °C. Pollen performance (adhesion to the stigma, germination, tube growth, fertilisation) was quantitatively analysed, using the aniline blue fluorescence method. Pollen adhesion was possible from -2 to 40 °C. Pollen germination and tube growth occurred from around 0 to 35 °C in most species. Fertilisation was observed from 5 to 30-32 °C (0-35 °C in G. germanica). The progamic phase was shortest in G. germanica (2 h at 30 °C, 12 h at 5 °C, 24 h at 0 °C), followed by R. glacialis (first fertilisation after 2 h at 30 °C, 18 h at 5 °C). In the remaining species, first fertilisation usually occurred after 4-6 h at 30 °C and after 24-30 h at 5 °C. Thus, mountain plants show remarkably flexible pollen performance over a wide temperature range and a short progamic phase, which may be essential for successful reproduction in the stochastic high-mountain climate.