High-altitude Species Research Articles

Population Trends of Central European Montane Birds Provide Evidence for Adverse Impacts of Climate Change on High-Altitude Species.

Climate change is among the most important global threats to biodiversity and mountain areas are supposed to be under especially high pressure. Although recent modelling studies suggest considerable future range contractions of montane species accompanied with increased extinction risk, data allowing to test actual population consequences of the observed climate changes and identifying traits associated to their adverse impacts are very scarce. To fill this knowledge gap, we estimated long-term population trends of montane birds from 1984 to 2011 in a central European mountain range, the Giant Mountains (Krkonoše), where significant warming occurred over this period. We then related the population trends to several species' traits related to the climate change effects. We found that the species breeding in various habitats at higher altitudes had more negative trends than species breeding at lower altitudes. We also found that the species moved upwards as a response to warming climate, and these altitudinal range shifts were associated with more positive population trends at lower altitudes than at higher altitudes. Moreover, long-distance migrants declined more than residents or species migrating for shorter distances. Taken together, these results indicate that the climate change, besides other possible environmental changes, already influences populations of montane birds with particularly adverse impacts on high-altitude species such as water pipit (Anthus spinoletta). It is evident that the alpine species, predicted to undergo serious climatically induced range contractions due to warming climate in the future, already started moving along this trajectory.

Arabis alpina and Arabidopsis thaliana have different stomatal development strategies in response to high altitude pressure conditions

The altitudinal gradient involves changes of the partial pressures of atmospheric gases such as CO2. This omnipresent phenomenon likely represents an evolutionary selective agent. We asked whether high altitude plant species had evolved specific response strategies to cope with high altitude pressure conditions. Plants of the high altitude species Arabis alpina and the low altitude species Arabidopsis thaliana were cultivated in growth chambers with high altitude pressure conditions (corresponding to 3000 m a.s.l.) and low altitude conditions (560 m). In both species, high altitude conditions resulted in the narrowing of stomatal aperture as well as a decrease in leaf area and weight. A. alpina produced significantly more stomata under high altitude conditions compared to low altitude conditions, while A. thaliana did not. Under low altitude conditions, however, stomatal density of A. alpina was smaller compared to A. thaliana. The increase in stomatal density of A. alpina was strongly related to the decrease in the partial pressure of CO2 under high altitude conditions. Thus, the adaptation of the high altitude plant A. alpina to high altitude pressure conditions does not consist in a genetically fixed elevated stomatal density but in a different response strategy of stomatal development to environmental factors compared to the lowland plant A. thaliana. A. alpina developed stomata largely uncoupled from other environmental factors than CO2. The increased stomatal density of A. alpina may ensure an optimal CO2 supply during the periods of favourable weather conditions for photosynthesis that are relatively rare and short in the alpine life zone.

Altitudinal patterns of abundances and parasitism in frugivorous drosophilids in west Java, Indonesia

Abundances and parasitism rates of frugivorous drosophilid flies were studied in three localities at altitudes of 250–1350 m in and near Bogor, Indonesia. The drosophilid and parasitoid species were classified into four groups: low-altitude species, high-altitude species, species abundant at a mid-altitude location and species occurring rather evenly from low to high locations. The ananassae and immigrans species groups were major drosophilids collected. All species of the ananassae species group were more abundant at lower altitudes, and the parasitism rate in this species group decreased with increasing altitude. Thus, the host abundance seems to affect the parasitism rate. On the other hand, the rate of parasitism in the immigrans species group showed no apparent relation with altitude or density, possibly due to the fact that species of this species group varied in altitudinal distribution. It is also suggested that the diversity of drosophilid species affects the composition of parasitoid species.

The role of GA3 in the germination process of high-mountain endemic and threatened species: Leontopodium nivalePinguicula fiorii and Soldanella minima subsp. samnitica (central Apennines, Italy)

The ex situ conservation of biodiversity is an essential tool for environmental protection interventions. Germination studies of seeds that belong to endangered species are essential for ex situ conservation strategies. In this study, we investigate the germination responses of three high-altitude endemic and vulnerable species (Leontopodium nivale, Pinguicula fiorii and Soldanella minima subsp. samnitica). Specifically we identified potential dormancy mechanisms by investigating the responses of germination percentage and rate to different concentrations of gibberellic acid (GA3), by performing a general linear model. L. nivale reached a germination percentage of 98.0 ± 2.0% (mean ± SE) under control conditions (20°C; 12/12 photoperiod; no GA3 addition). P. fiorii showed the highest germination percentage (78.0 ± 2.0%) in the treatment with GA3 500 ppm. S. minima subsp. samnitica did not show sensitivity to GA3 but responded positively to cold stratification (6 month at 5°C) with a germination percentage of 90 ± 6%. This study made it possible to acquire important information on the germination process of threatened and rare endemic taxa.

A taxonomic revision of the genus Pauridia (Hypoxidaceae) in southern Africa

Pauridia, a genus confined to southern Africa, Australia, Tasmania and New Zealand, is distinguished from other genera of Hypoxidaceae by the synapomorphic mucilage canals above the vascular bundles of the leaves and the disulcate, micro-echinate pollen grains, apart from the secondarily derived trisulcate state in one of the Australian species. The majority of species (86%) are endemic to southern Africa, mostly in seasonally damp sites within the winter rainfall and all-year-round rainfall regions of Namibia and South Africa. The Australian, Tasmanian and New Zealand taxa were revised by Henderson (1987) and the African taxa were revised by Nel in 1914. The difficulties of applying Nel’s classification to the wealth of new material collected within the last century duly prompted a revision of Pauridia in southern Africa. Thirty species and three subspecies are recognised in this work, which includes a key to the species and full descriptions of all the southern African taxa, together with the updated nomenclature, geographic distribution, discussion on diagnostic characters and putative relationships of each. Four new species, P. breviscapa, P. maryae, P. monticola, and P. pudica and two new subspecies, P. monticola subsp. nubigena, P. gracilipes subsp. speciosa are described and illustrated, adding to the four other southern African species described subsequent to Nel’s revision. P. acida, a high altitude species that appears to flower after fire, remains poorly understood and one taxon, though almost certainly distinct, has been left unnamed until more complete material becomes available. The study of morphology, leaf anatomy and seed micromorphology suggests that the floral bracts and seeds are most helpful in revealing putative relationships among species. These characters are nevertheless insufficiently discrete to allow the recognition of formal infrageneric taxa. The northwestern and southwestern Cape has the highest concentration of species and according to IUCN criteria is home to three Endangered species (P. canaliculata, P. maximiliani, P. pygmaea) and two Vulnerable species (P. alba, P. linearis), found mainly in the lowland areas where natural habitats have been extensively fragmented.

Testing the efficiency of nested barriers to dispersal in the Mediterranean high mountain plant Edraianthus graminifolius (Campanulaceae).

Due to strong spatial heterogeneity and limited Pleistocene glaciation, the Balkan Peninsula is a major European biodiversity hot spot. Surprisingly little, however, is known about patterns and processes of intraspecific diversification of its biota in general and of high-altitude species in particular. A well-suited system to test hypotheses with respect to various isolating factors acting at different geographic scales and to explore full-range phylogeographic patterns on the Balkan Peninsula is Edraianthus graminifolius (Campanulaceae), distributed in the western Balkan mountain systems, the southwestern Carpathians and the Apennine Peninsula. To this end, we used a dense population sampling and employed amplified fragment length polymorphism (AFLP) markers and plastid DNA sequences supplemented by ecological niche modelling. The strongest splits were inferred to separate southern and northern Balkan populations from the central ones, from where range extension occurred to the Carpathians and, in more recent times, once or twice to the Apennine Peninsula. The three genetic groups in the western Balkan Peninsula were remarkably congruent among molecular markers, suggesting that the barriers to gene flow acted over long time periods facilitating allopatric differentiation. Each main group of Balkan populations contained genetically and geographically distinct subgroups, which likely are the result of local refugia during warmer periods. Evidently, the topographically highly complex and during the Last Glacial Maximum only locally glaciated Balkan Peninsula is a hot spot of species richness and endemism as well as a sanctuary of intraspecific genetic diversity, even if the underlying causes remain insufficiently understood.

Geographic distribution of the chytrid pathogen Batrachochytrium dendrobatidis among mountain amphibians along the Italian peninsula

The amphibian chytrid pathogen Batrachochytrium dendrobatidis (Bd) is considered a major cause of amphibian population declines, particularly in montane areas. Here, we investigated the presence and distribution of Bd among populations of 3 mid- to high-altitude species spanning the entire Italian peninsula (486 individuals from 39 sites overall): the stream frog Rana italica, the fire salamander Salamandra salamandra gigliolii, and the alpine newt Mesotriton alpestris apuanus. We found Bd in all of the analyzed species. Despite the widespread distribution of the pathogen, its overall prevalence (6, 9 and 19%, respectively) was lower than previously reported for the endangered Apennine yellow-bellied toad Bombina pachypus (62.5%). Moreover, several populations of the species studied here were not infected, even at sites where Bd has been detected in other host species. When coupled with the lack of evidence for Bd-related mortalities in these species in peninsular Italy, these results suggest that mechanisms of resistance and/or tolerance are protecting populations of these species from the pathogenic activity of Bd. Nevertheless, in light of the dynamic pattern of Bd-host interactions reported in other studies, of Bd-related mortalities in at least 1 study species (S. s. salamandra) in other areas, and the ongoing climate changes in montane environments, we suggest that the occurrence of Bd should be considered a potential threat to the long-term persistence of these species, and urge the implementation of monitoring and conservation plans.

The Impact of Global Warming on Floral Traits That Affect the Selfing Rate in a High-Altitude Plant

Premise of research. Changes in the abiotic environment can influence plant mating systems through phenotypic changes in floral traits that affect selfing. Herkogamy (spatial separation of male and female functions within a flower), dichogamy (temporal separation), and total flower number affect within- (autogamy) and/or among- (geitonogamy) flower selfing. We examined the potential effects of environmental changes associated with global warming on herkogamy, dichogamy, and number of flowers per plant and discussed the impacts on the mating system in a high-altitude species, Aquilegia coerulea.Methodology. In the high-altitude habitats in which this species grows, global warming is expected to increase both temperature and spring precipitation in the form of rain. We grew plants from 17 families from each of three natural populations of A. coerulea at two temperature and two water treatments, leading to 692 experimental plants (2 temperature × 2 water × 3 populations × 17 families × 3–4 individuals per fa...

Long-term vegetation dynamic in the Northwestern Caucasus: which communities are more affected by upward shifts of plant species?

We studied long-term (25–31 years) dynamics of alpine communities at the Teberda Reserve, NW Caucasus, Russia, to test the following hypotheses: (1) lower altitude species increase and high altitude species decrease their abundance as a consequence of climate warming; (2) such changes in abundance are more significant in communities with short growth season (due to persistent snow cover) compared to exposed communities; (3) species with similar changes in abundance have similar functional traits. Four alpine communities with different positions in relief were considered in order of winter snow cover: alpine lichen heaths (ALH), Festuca varia grasslands (FVG), Geranium-Hedysarum meadows (GHM), and snowbed communities (SBC).

Influence of HbCO Structure of the Bar-Headed Goose on Photolysis Thermodynamics as Studied by the Nanosecond Laser-Ultrasonic Technique

The bar-headed goose, a specialized high-altitude species, has a capacity for high oxygen uptake from a hypoxic environment. It thus has a higher oxygen affinity than other bird species of lower-altitude environments. Oxygen affinity is determined by molecular structures and genetic mutations of hemoglobin (Hb), which can also influence the coordinating structures and dynamics of oxygen-Hb. To explore the structural differences in Hbs as between high and low altitude species, photolysis dynamic parameters, including quantum yield, enthalpy, and conformational volume changes in carboxy-Hbs (HbCO) for the bar-headed goose and low altitude counterparts (the Chinese goose and chicken) were investigated by the laser pumping-probing technique and photoacoustic calorimetry. Comparing the photolysis results for HbCO of the three species, the enthalpy and conformational volume changes of the bar-headed goose were much smaller than those of the others, although the quantum yields of all three species are similar. To explain the possible mechanisms of these differences, modifications of salt bridges and key residue mutations at the α β subunit interfaces of the proteins are described and discussed briefly.

Geomorphological disturbance affects ecological driving forces and plant turnover along an altitudinal stress gradient on alpine slopes

The altitudinal gradient is considered as a stress gradient for plant species because the development and fitness of plant communities tend to decrease as a result of the extreme environmental conditions present at high elevations. Abiotic factors are predicted to be the primary filter for species assemblage in high alpine areas, influencing biotic interactions through both competition for resources and positive interactions among species. We hypothesised that the relative importance of the ecological driving forces that affect the biotic interactions within plant communities changes along an elevation gradient on alpine debris slopes. We used multiple gradient analyses of 180 vegetation plots along an altitudinal range from ~1,600 to 2,600 m and single 100 m-bands in the Adamello-Presanella Group (Central Alps) to investigate our hypothesis; we measured multiple environmental variables related to different ecological driving forces. Our results illustrate that resource limitations at higher elevations affect not only the shift from competition to facilitation among species. A geomorphological disturbance regime along alpine slopes favours the resilience of the high-altitude species within topographic/geomorphological traps. An understanding of the ecological driving forces and positive interactions as a function of altitude may clarify the mechanisms underlying plant responses to present and future environmental changes.

Thermospheric neutral temperatures derived from charge-exchange produced N<sub>2</sub><sup>+</sup> Meinel (1,0) rotational distributions

Abstract. Thermalized rotational distributions of neutral and ionized N2 and O2 have long been used to determine neutral temperatures (Tn) during auroral conditions. In both bright E-region (≲150 km) auroras, and in higher-altitude auroras, spectral distributions of molecular emissions employed to determine Tn in the E-region cannot likewise be used to obtain Tn in the F-region. Nevertheless, charge-exchange reactions between high-altitude (≳130 km) species provide an exception to this situation. In particular, the charge-exchange reaction O+(2D) + N2(X) → N+2(A2Πu, ν' = 1 + O(3P) yields thermalized N2+ Meinel (1,0) emissions, which, albeit weak, can be used to derive neutral temperatures at altitudes of ~130 km and higher. In this work, we present N2+ Meinel (1,0) rotational temperatures and brightnesses obtained at Svalbard, Norway, during various auroral conditions. We calculate Tn at thermospheric altitudes of 130–180 km from thermalized rotational populations of N2+ Meinel (1,0); these emissions are excited by soft electron (≲1 keV) impact and charge-exchange reactions. We model the contributions of the respective excitation mechanisms, and compare derived brightnesses to observations. The agreement between the two is good. Emission heights obtained from optical data, modeling, and ISR data are consistent. Obtaining thermospheric Tn from charge-exchange excited N2+ Meinel (1,0) emissions provides an additional means of remotely sensing the neutral atmosphere, although certain limiting conditions are necessary. These include precipitation of low-energy electrons, and a non-sunlit emitting layer.

Local adaptation to altitude underlies divergent thermal physiology in tropical killifishes of the genus Aphyosemion.

In watersheds of equatorial West Africa, monophyletic groups of killifish species (genus Aphyosemion) occur in discrete altitudinal ranges, low altitude species (LA, sea level to ∼350 m) or high altitude species (HA, 350 to 900 m). We investigated the hypothesis that local adaptation to altitude by the LA and HA species would be revealed as divergent effects of temperature on their physiological energetics. Two species from each group (mass ∼350 mg) were acclimated to 19, 25 and 28°C, with 19 and 28°C estimated to be outside the thermal envelope for LA or HA, respectively, in the wild. Wild-caught animals (F0 generation) were compared with animals raised in captivity at 25°C (F1 generation) to investigate the contribution of adaptation versus plasticity. Temperature significantly increased routine metabolic rate in all groups and generations. However, LA and HA species differed in the effects of temperature on their ability to process a meal. At 25°C, the specific dynamic action (SDA) response was completed within 8 h in all groups, but acclimation to temperatures beyond the thermal envelope caused profound declines in SDA performance. At 19°C, the LA required ∼14 h to complete the SDA, whereas the HA required only ∼7 h. The opposite effect was observed at 28°C. This effect was evident in both F0 and F1. Reaction norms for effects of temperature on SDA therefore revealed a trade-off, with superior performance at warmer temperatures by LA being associated with inferior performance at cooler temperatures, and vice-versa in HA. The data indicate that divergent physiological responses to temperature in the LA and HA species reflect local adaptation to the thermal regime in their habitat, and that local adaptation to one thermal environment trades off against performance in another.

Mitogenomic analysis of the genus Pseudois: Evidence of adaptive evolution of morphological variation in the ATP synthase genes

The genus Pseudois includes two variable taxa, blue sheep (Pseudois nayaur) and dwarf blue sheep (Pseudois schaeferi), that exhibit notable geographic variation in morphology and ecological niche, suggesting the potential for significant adaptive differentiation between these two goats. Blue sheep are broadly distributed in the Tibetan Plateau and peripheral mountains through Central Asia, while dwarf blue sheep are only found in the gorges of the upper Yangtze River (Jinsha River) near Batang county, Sichuan province and adjacent mountains. Although they are all adapted to high altitude environments, endangered dwarf blue sheep show unique morphological variation and niche shifts compared to blue sheep. Mitochondria play important roles in oxygen usage and energy metabolism. The energetically demanding lifestyles of these high altitude species may have altered the selective regimes on mitochondrial genes encoding proteins related to cellular respiration. Here, we compared the sequences of 13 protein-coding genes in the mitochondrial genome of dwarf blue sheep with those of blue sheep to understand the genetic basis of morphological variation. Using neighbor-joining, maximum-likelihood and Bayesian approaches, we estimated rates of synonymous (dS) and nonsynonymous (dN) substitutions. Independent analyses showed that no ω ratio was larger than 1, suggesting that all mitochondrial 13 genes were under the purifying selection. Surprisingly, we found that the ω ratio (dN/dS) of the ATP synthase complex (ATP6 and ATP8) in blue sheep is sixteen times that of dwarf blue sheep (0.340 compared to 0.021). This result was confirmed by a separate analysis of ATP synthase genes from two additional P. schaeferi individuals and two P. nayaur individuals. We hypothesize that the large body size and diverse feeding styles are factors influencing the nonsynonymous substitutions in the ATP synthase complex of blue sheep.

Small sample, substantial contribution: additions to the Honduran hawkmoth (Lepidoptera: Sphingidae) fauna based on collections from a mountainous protected area (Cusuco National Park)

With the largest part of diversity in the world absorbed by invertebrates, ignoring invertebrates in biodiversity surveys and monitoring of areas under conservation would give a strongly incomplete image. The poor knowledge of most invertebrate taxa and their enormous diversity limits most surveys to the better-studied groups. Hawkmoths (Lepidoptera: Sphingidae) are one of the more charismatic and well-known groups among the Lepidoptera and hence a valuable group commonly used in biodiversity research. In this small-scale study, 42 museum specimens of sphingids from Cusuco National Park (Cortes, Honduras) were identified and compared to recent published accounts. This yielded three new country records and, in addition, four new regional records for the park. Some of the additions to the Honduran fauna probably result from recent taxonomic changes. However, the several contributions using a small collection of this well-studied group in an area which has attracted previous research interest, demonstrate the incomplete data availability and the necessity for more rigorous surveying. Several new records concern high altitude species, indicating the data gap in mountains. As elevation is an important determinant of sphingid community structure, sampling across an altitudinal range is recommended. This study also underpins the usefulness of a reference collection-based approach in particular, as many hawkmoth species are identified using subtle diagnostic characters.

Recent Plant Diversity Changes on Europe’s Mountain Summits

In mountainous regions, climate warming is expected to shift species' ranges to higher altitudes. Evidence for such shifts is still mostly from revisitations of historical sites. We present recent (2001 to 2008) changes in vascular plant species richness observed in a standardized monitoring network across Europe's major mountain ranges. Species have moved upslope on average. However, these shifts had opposite effects on the summit floras' species richness in boreal-temperate mountain regions (+3.9 species on average) and Mediterranean mountain regions (-1.4 species), probably because recent climatic trends have decreased the availability of water in the European south. Because Mediterranean mountains are particularly rich in endemic species, a continuation of these trends might shrink the European mountain flora, despite an average increase in summit species richness across the region.

The importance of integrating behavioural ecology into experimental design when censusing Himalayan Galliformes

The Himalayan species of Galliformes are scantily studied, and their present status is not properly documented. Spring surveys conducted in some areas usually rely on dawn counting to estimate population abundance and distribution. In the upper Khumbu–Himal region of Sagarmatha National Park (Nepal), we conducted point count surveys of three high-altitude species: Himalayan monal, Blood pheasant and Tibetan snowcock. Our study area was between 3300 and 5000 m a.s.l, and it was characterised by subalpine and alpine vegetation. We recorded bird distribution in relation to altitude and habitats (forest, shrub and high-altitude grassland). Two different patterns emerged in the species studied: Tibetan snowcock and Blood pheasant counts gave optimal results at dawn. Contrarily, Himalayan monal counts at dawn were far less representative of species abundance than dusk counts. This implies the need to fully understand the behavioural ecology of a species before planning a field study for population estimates.

Seeds Germination and Seedlings Analysis of Saussurea Costus Royle Ex Benth. In High and Low Altitudinal Villages of District Uttarkashi (Uttarakhand)

Saussurea costus (Falc.) Lipschitz, syn. Saussurea lappa C. B. Clarke is a well known and important medicinal plant widely used in several indigenous systems of medicine for the treatment of various ailments, viz. asthma, inflammatory diseases, ulcer and stomach problems. Different pharmacological experiments in a number of invitro and invivo models have convincingly demonstrated the ability of Saussurea costus to exhibit anti-inflammatory, anti-ulcer, anti-cancer and hepatoprotective activities, lending support to the rationale behind several of its traditional uses. The present review is an up-to-date and comprehensive analysis of seed germination, survival percentage and seeding analysis of Saussurea costus in lower and higher Himalayan altitudinal villages of district Uttarkashi. The present study deals with successful seeds germination, survival percentage and seedlings analysis of this species using both field and within the polyhouse techniques in low and high altitudinal villages. The materials were collected from the Besaj Sang Ikai, Uttarkashi. Vegetative propagation was achieved by rooting runner cuttings. Seedlings were measured by seed germination and survival percentage, root and shoot length. A significant increment in root length was recorded in high altitudinal Polyhouse condition as compared to low altitude. In low altitudinal village Genwala, open field condition seed germination percentage and survival percentage ranged from 5.00% 18.18% and 18.18% to 25.00% while in low altitudinal village seed germination percentage and survival percentage within Polyhouse were ranged from 23.00% to 45.00% and 40.45% to 66.70% respectively. Highest percentage of seed germination and survival percentage were noticed in high altitudinal village, Bagori (Within Polyhouse) from 40.50% to 60.00% (seed germination), 60.70% to 76.50% (survival percentage) respectively. The leaves are produced in a dense basal rosette, and spiral up the flowering stem. The woolly hairs in high altitude species of the family are the densest, and aid in thermoregulation of the flowers, minimizing frost damage at night, and also preventing ultraviolet light damage from the intense high altitude sunlight.

Woody plant diversity along Tons Riverine ecosystem in Uttarakhand (India)

The present paper incorporates the woody plant diversity along the Tons river forest ecosystem from Naitwar to Tiuni (42km). This study is restricted to 100m from either side of the river. Tons riverine forest ecosystem supports 156 species, 122 genera and 62 families. The growth habit of the woody flora indicates high percentage of shrubs 50% followed by large trees 39%, woody climber 6% and small trees 5%. The study site indicates rich woody vegetation showing interconnectivity of high altitude and low altitude specific plant species.

Last Word on Point:Counterpoint: High altitude is/is not for the birds!

to the editor: We would like to thank the scientists who commented on this Point:Counterpoint. We originally argued that high altitude is for the birds because of several unique features that should improve their ability to exercise at extremely high altitudes ([4][1]). Many authors agreed with our