Geographic patterns and ecological causes of phylogenetic structure in mosses along an elevational gradient in the central HimalayaOA

Understanding the underlying mechanisms driving species assembly along elevational gradients is of great interest in ecology and biogeography. The Himalaya is one of the world's hotspots of biodiversity,and the elevational gradient of the central Himalaya in Nepal is one of the longest elevational gradients in the world. Mosses are important constituents of vegetation in the Himalaya, and knowledge of geographic patterns and ecological causes of phylogenetic structure of mosses along elevational gradients in the Himalaya is critical to understanding the assembly of plant diversity in general, and moss diversity in particular, in the Himalaya. Here, we investigate the relationships of phylogenetic structure metrics reflecting different evolutionary depths with elevation and climatic variables representing mean temperature and precipitation conditions, climate extremes, and climate seasonality for mosses distributed along an elevational gradient spanning about 5000 m in the central Himalaya, Nepal. For a given climatic variable, different metrics of phylogenetic structure show different spatial and climatic patterns, but all phylogenetic metrics standardized for species richness show that phylogenetic dispersion in moss assemblages tend to increase with increasing elevation and decreasing temperature. The standardized effect size of mean pairwise distance of moss assemblages shows a triphasic(zig-zag)pattern, which is generally consistent with the triphasic patterns previously found in angiosperms and ferns along the same elevational gradient. Our study shows that temperature-related variables and climate seasonality variables are more important drivers of phylogenetic dispersion in mosses in Nepal,compared with precipitation-related variables and climate extreme variables, respectively.     

Replicated hybrid zones of Xiphophorus swordtails along an elevational gradient

Natural hybrid zones provide opportunities to study a range of evolutionary phenomena from speciation to the genetic basis of fitness-related traits. We show that widespread hybridization has occurred between two neo-tropical stream fishes with partial reproductive isolation. Phylogenetic analyses of mitochondrial sequence data showed that the swordtail fish Xiphophorus birchmanni is monophyletic and that X. malinche is part of an independent monophyletic clade with other species. Using informative single nucleotide polymorphisms in one mitochondrial and three nuclear intron loci, we genotyped 776 specimens collected from twenty-three sites along seven separate stream reaches. Hybrid zones occurred in replicated fashion in all stream reaches along a gradient from high to low elevation. Genotyping revealed substantial variation in parental and hybrid frequencies among localities. Tests of F(IS) and linkage disequilibrium (LD) revealed generally low F(IS) and LD except in five populations where both parental species and hybrids were found suggesting incomplete reproductive isolation. In these locations, heterozygote deficiency and LD were high, which suggests either selection against early generation hybrids or assortative mating. These data lay the foundation to study the adaptive basis of the replicated hybrid zone structure and for future integration of behaviour and genetics to determine the processes that lead to the population genetic patterns observed in these hybrid zones.     

Flowering phenology and reproduction of the Solidago virgaurea L. complex along an elevational gradient on Mt Norikura,central Japan

This study examined the flowering phenology and reproductive traits of the Solidago virgaurea complex at four elevations in the subalpine zone in Japan using a bagging experiment. Flowering started earlier at higher elevations. Syrphid flies mainly visited flowers of the S. virgaurea complex, and the number of visits was considerably lower at the lowest elevation than at the three other elevations. Although the number of seeds per individual did not differ among the four elevations, total achene weight per individual was lower at the lowest elevation than at the three other elevations. The weight of an achene and seed germination rate of the control were much greater at higher elevations than at the lowest elevation. The weight of an achene and germination rate for the control were as low as the bagging treatment at the lowest elevation with infrequent flower visitors, which indicates that the S. virgaurea complex is a facultative outcrosser. The S. virgaurea complex is thought to produce seeds during a short growing season at high elevations by starting to flower earlier, and the large seed size is advantageous for seedling establishment at high elevations. Therefore, elevational changes in flowering phenology and reproductive traits are thought to be an adaptation to the short growing season at high elevations.     

Distribution of bats along an elevational gradient in the Andes of south-eastern Peru

At least 193 species of mammals are known to occur within the Manu Biosphere Reserve in south-eastern Peru, contributing to its stature as one of the world's richest protected areas. Bats (Order Chiroptera) comprise more than 42% (82 species) of this diversity. Analyses of bat capture records over a transect extending more than 3 km in elevation show that most bat species at Manu are widely distributed in the Amazon Basin. Few are montane endemics or are localized in south-eastern Peru, although exceptions to this generalization include two species new to science. Highland bat faunas tend to be attenuated versions of those found below, and the elevational zonation of bat communities is weak. Species turnover with elevation is monotonic and more-or-less smooth, with Jaccard's similarity values falling to 0.5 for sites differing by 750m in elevation. Subtle and orderly change in species composition with elevation is also reflected in the nested-subset structure of these communities; over 19 different levels, this pattern of hierarchical structure is both striking and highly significant. Elevational ranges of species generally increase with elevation, in accordance with Stevens' extension of 'Rapoport's rule' of range amplitude. However, support for 'Stevens' rule' may be trivial, given Amazonian richness and Andean impoverishment. Reduced richness and poorly developed endemism in Andean bat communities contrast with patterns shown by sympatric rodent faunas, which are diverse and strongly endemic on the Altiplano and markedly zoned along the Eastern Versant. Contrasts are less sharp with bird communities, which nevertheless exhibit stronger zonation and higher endemicity. Factors responsible for these distinctive distributional patterns are discussed.     

Variations in sexual and asexual reproduction of Scirpus mariqueter along an elevational gradient

In order to assess the importance of sexual and asexual reproduction during the life history of Scirpus mariqueter, its reproductive and growth characters were concurrently examined along an elevational gradient (from low elevation to high elevation). The proportions of flowering shoot and inflorescence mass, seed : flower ratio and seed weight were used to quantify the investment in sexual reproduction. The proportions of current-year shoot and rhizome mass were used to quantify the investment in asexual reproduction, and the proportion of corm mass was used for growth, respectively. It was found that vegetative propagation predominated at low elevation, whereas sexual reproduction predominated at high elevation; and that sexual reproduction increased with declining asexual reproduction along the gradient. The results suggest that asexual reproduction is relatively favored in the early life stage, whereas sexual reproduction is favored when the population becomes mature and aged, probably because of the functional differentiation between the two reproductive types. Sexual productive characters (i.e. the proportions of flowering shoot and inflorescence mass) were negatively correlated to both growth and asexual reproductive characters along the gradient, indicating there might exist some trade-offs among growth, sexual and asexual reproduction during the life history. However, no obvious pattern was found between asexual reproductive characters and growth characters along the elevational gradient, possibly because of the varied relationships between them at different life stages. The variations in sexual and asexual reproduction in the species and the relationship between them are thought to be of great significance for local population growth, species persistence and evolution.     

Plant diversity patterns in semi-natural grasslands along an elevational gradient in southern Norway

Leaf demography and growth of six common, co-occurring woody plant species that varied in stature (tree vs. shrub) and leaf texture (sclerophyllous, coriaceous, malacophyllous) were examined in a subtropical savanna parkland in southern Texas, USA. We tested the hypotheses that, (a) leaves of plants with evergreen canopies would have longer life spans than those of deciduous species; (b) supplementation of soil moisture would decrease leaf life span in both evergreen and deciduous species; (c) species responses to increased soil moisture availability would be inversely related to leaf longevity; and (d) deciduous growth forms would exhibit a greater growth response to increased soil moisture availability than their evergreen counterparts.A variety of seasonal leaf habits (evergreen, winter-deciduous and summer-deciduous canopies) and leaf life spans (median = 66 to 283 days) were represented by the targeted species, but there was no clear relationship between seasonal leaf habit and leaf longevity. Among species with evergreen canopies, median leaf longevity ranged from short (Zanthoxylum fagara = 116 days; Condalia hookeri = 158 days) to long (Berberis trifoliolata = 283 days) but did not exceed 1 yr. In fact, leaf longevity in evergreen shrubs was often comparable to, or shorter than, that of species with deciduous canopies (Ziziphus obtusifolia = 66 days; Diospyros texana = 119 days; Prosopis glandulosa = 207 days). Augmentation of surface soil moisture had no detectable effect on median leaf life span in any species and there was no clear relationship between leaf longevity and species growth responses to irrigation. Contrary to expectations, species with evergreen canopies responded to irrigation by producing more leaf biomass, longer shoots and more leaf cohorts/year than did deciduous species.Species differences in the annual cycle of leaf initiation, leaf longevity and canopy development, combined with contrasts in root distributions and a highly variable climate, may allow for spatial and temporal partitioning of resources and hence, woody species coexistence and diversity in this system. However, the lack of expected relationships between leaf longevity, leaf habit and plant responses to resource enhancement suggests that structure-function relationships and functional groupings developed in strongly seasonal environments cannot be applied with confidence to these subtropical savannas and thorn woodlands.     

Mammalian species richness and morphological complexity along an elevational gradient in the arid south-west

Aim We examined the relationship between species richness and morphological complexity of terrestrial mammal communities along an elevational gradient. Location The gradient is in the Sonoran Desert in Southern California and extends from a sand dune habitat near sea level to coniferous forest ending at >2600 m. Methods Morphological diversity, characterized by both size and shape of coexisting mammal species, was estimated within and between sites from projections of variables on principal components axes. Similarities among species were calculated as Euclidean distances. To tease apart size and shape, we constructed two principal component analyses: one based on log-transformed original measurements, the other on log-transformed proportional shape variables. To test whether species number accounted for the morphological diversity at each site we designed two null models. The models generated were random communities generated from the forty-two-species pool. Indices of morphological diversity for real communities were compared with the results of 500 simulations of each null model. Results Species richness varied along the gradient, peaking in the mid-elevation agave-ocotillo habitat. Morphological diversity of shapes and sizes correlated strongly with species richness. Locomotor, tooth, and skull traits were all important in distinguishing among species. Main conclusions Two important patterns emerged: (1) diversity of both sizes and of shapes of species within communities correlated positively with species number, and both sets of variables behaved similarly across this gradient; (2) the most species rich sites were not composed of specialists on these best places, but rather, a community of species derived from overlapping faunal groups.     

Superficial lizards in cold climates: Nest site choice along an elevational gradient

Embryonic conditions may limit the distributions of egg‐laying ectotherms, and recent research suggests that nesting mothers of wide‐ranging species may use a number of factors to compensate for differing climates. However, while variation in temporal factors across environmental gradients are common or pervasive (i.e. seasonal timing of nesting), similar evidence for spatial factors is rare (e.g. aspect, openness and depth of nest sites). I tested the idea that a wide‐ranging lizard, the Australia water dragon (Physignathus lesueurii), uses nest depth to counter climate differences along a temperature cline at their cold‐end range margin. Two measures of nest depth were significantly, inversely related to elevation across six populations spanning 700 m. Elevation explained 83–86% of the variation in nest depth. These findings support a thermal compensatory mechanism for this pattern, although soil moisture compensation is plausible. My results directly support a recent, untested prediction that the evolution of viviparity in reptiles is preceded by a behavioural shift towards increasingly superficial nest sites in cold climates, followed by selection for increased egg retention to avoid temperature extremes. However, in the present study egg desiccation rates increased with increasing elevation in a dry year, suggesting that increased egg retention may evolve in response to lethal hydric conditions, rather than lethal temperatures. When considered alongside recent research, the present study indicates that water dragons possess several mechanisms for adjusting to climate change.     

Grasshopper community response to climatic change: variation along an elevational gradient

Background

The impacts of climate change on phenological responses of species and communities are well-documented; however, many such studies are correlational and so less effective at assessing the causal links between changes in climate and changes in phenology. Using grasshopper communities found along an elevational gradient, we present an ideal system along the Front Range of Colorado USA that provides a mechanistic link between climate and phenology.

Methodology/Principal Findings

This study utilizes past (1959–1960) and present (2006–2008) surveys of grasshopper communities and daily temperature records to quantify the relationship between amount and timing of warming across years and elevations, and grasshopper timing to adulthood. Grasshopper communities were surveyed at four sites, Chautauqua Mesa (1752 m), A1 (2195 m), B1 (2591 m), and C1 (3048 m), located in prairie, lower montane, upper montane, and subalpine life zones, respectively. Changes to earlier first appearance of adults depended on the degree to which a site warmed. The lowest site showed little warming and little phenological advancement. The next highest site (A1) warmed a small, but significant, amount and grasshopper species there showed inconsistent phenological advancements. The two highest sites warmed the most, and at these sites grasshoppers showed significant phenological advancements. At these sites, late-developing species showed the greatest advancements, a pattern that correlated with an increase in rate of late-season warming. The number of growing degree days (GDDs) associated with the time to adulthood for a species was unchanged across the past and present surveys, suggesting that phenological advancement depended on when a set number of GDDs is reached during a season.

Conclusions

Our analyses provide clear evidence that variation in amount and timing of warming over the growing season explains the vast majority of phenological variation in this system. Our results move past simple correlation and provide a stronger process-oriented and predictive framework for understanding community level phenological responses to climate change.     

Variation in home range size along an elevational gradient in the iguanid lizard Sceloporus merriami

Summary Variation in home range size among three populations of the saxicolous iguanid lizard Sceloporus merriami was studied in relation to density, food availability, behavioral patterns, and daily activity periods. These populations, separated by a maximum distance of 50 km, occur along an elevational gradient which spans the elevational range of the species in west Texas. The population with the smallest home ranges had the highest density, the lowest food availability, and the most restricted daily activity period. Within populations, males consistently had larger home ranges than females. No significant between-year variation in home range size was noted within a given area or sex. Intraspecific variation in home range size is not explained by any single factor, such as food availability or density, but appears to be complexly determined by the interaction of energetic, breeding, and activity constraints.     

Stomatal responses and water relations of Eucalyptus pauciflora in summer along an elevational gradient

Summary The spatial and temporal variation of lead conductance (g) in Eucalyptus pauciflora was analysed with respect to photon flux area density (I), temperature (T), water vapour concentration deficit (w), and leaf water potential () at four different sites between 940 m and 2,040 m altitude in the Snowy Mountains of south-eastern Australia. Along this altitudinal gradient the precipitation/evaporation ratio increases from 1 to 4. The results show that gas diffusion in this tree species is primarily controlled by I and w at all sites, independently of the specific soil moisture regime. Even under dry midsummer conditions with predawn leaf water potentials of-1 MPa at the lowest altitude, had no striking effect on g.The humidity threshold for the onset of stomatal closure does not vary greatly between the study sites (12.2±1.3 Pa kPa^-1). The highest and lowest values observed for , the osmotic potential at water saturation (from pressure/volume curves), the mean and maximum g and stomatal dentity, all increase with elevation. The highest (least negative) osmotic potentials were obtained at all sites in midsummer. It therefore appears that there is no osmotic adjustment to drought in the seasonal course. The maximum difference between osmotic potentials obtained at the lowest and highest sites is 0.46 MPa. In general osmotic potential varies less than has been reported for other plant species exposed to varying water regimes. This may be the consequence of the pronounced feed-forward response of the stomata to evaporative demand, which led to only moderate tissue desiccation, never exceeding the turgor loss point. E. pauciflora is a tree species with a very conservative utilisation of soil water, which adjusts to drought via stomatal control of water loss, rather than via osmotic properties.These results explain previous reports of the comparatively high susceptibility of E. pauciflora to severe drought and its positive influence on the hydrological balance of mountain ecosystems in the Australian Alps.     

Response to mowing of coastal brackish meadow plant communities along an elevational gradient

Over a six year period, 1979–1985, eight stands of till then ungrazed and unmown brackish meadow plant communities were treated by mowing once a year (August), in order to detect the response of the vegetation. The stands were distributed along a local elevation gradient, which was found to influence the composition of the vegetation through a complex-gradient, composed by unidirectional gradients in inundation, in ground water level and in Na as per cent of Ca+Mg+Na+K.
The mowing was carried out within 5 ×5 m treatment plots and the effect was indicated by quantitative analysis of the vegetation. A natural succession progressing at the locality, most distinctly manifested by expansion of Phragmites australis , was traced during the study period by means of unmown control plots.
The mowing caused increase in species diversity in all stands, mainly due to increase in evenness among the species.
The resistance of the stands to the mowing was found to differ in relation to elevation and to the initial composition of the vegetation. In upper geolittoral Festuca rubra stands the resistance was high and the mowing did not affect presence or colonisation of Phragmites . In upper-middle Festuca rubra stands resistance was lower, mainly indicated by suppression of Phragmites . Low elevation Scirpus maritimus reed beds on waterlogged soil was rather resistant, but four additional years of mowing led to a change into Agrostis stolonifera-Puccinellia maritima meadow. The resistance of low elevation Phragmites reed bed was low, as the mowing within a few years changed the reed bed into Agrostis stolonifera meadow.
The results were discussed in relation to the use of mowing as a nature conservation management tool in salt meadows.     

Contrasting nitrogen and phosphorus dynamics across an elevational gradient for subarctic tundra heath and meadow vegetation

Aims

This study explores soil nutrient cycling processes and microbial properties for two contrasting vegetation types along an elevational gradient in subarctic tundra to improve our understanding of how temperature influences nutrient availability in an ecosystem predicted to be sensitive to global warming.

Methods

We measured total amino acid (Amino-N), mineral nitrogen (N) and phosphorus (P) concentrations, in situ net N and P mineralization, net Amino-N consumption, and microbial biomass C, N and P in both heath and meadow soils across an elevational gradient near Abisko, Sweden.

Results

For the meadow, NH₄ ⁺ concentrations and net N mineralization were highest at high elevations and microbial properties showed variable responses; these variables were largely unresponsive to elevation for the heath. Amino-N concentrations sometimes showed a tendency to increase with elevation and net Amino-N consumption was often unresponsive to elevation. Overall, PO₄-P concentrations decreased with elevation and net P immobilization mostly occurred at lower elevations; these effects were strongest for the heath.

Conclusions

Our results reveal that elevation-associated changes in temperature can have contrasting effects on the cycling of N and P in subarctic soils, and that the strength and direction of these effects depend strongly on dominant vegetation type.     

Reproductive limits of a late-flowering high-mountain Mediterranean plant along an elevational climate gradient

Mountain plants are particularly sensitive to climate warming because snowmelt timing exerts a direct control on their reproduction. Current warming is leading to earlier snowmelt dates and longer snow-free periods. Our hypothesis is that high-mountain Mediterranean plants are not able to take advantage of a lengthened snow-free period because this leads to longer drought that truncates the growing season. However, reproductive timing may somewhat mitigate these negative effects through temporal shifts. We assessed the effects of flowering phenology on the reproductive success of Silene ciliata, a Mediterranean high-mountain plant, across an altitudinal gradient during two climatically contrasting years. The species showed a late-flowering pattern hampering the use of snowmelt water. Plant fitness was largely explained by the elapsed time from snowmelt to onset of flowering, suggesting a selective pressure towards early flowering caused by soil moisture depletion. The proportion of flowering plants decreased at the lowest population, especially in the drier year. Plants produced more flowers, fruits and seeds at the highest population and in the mild year. Our results indicate that water deficit in dry years could threaten the lowland populations of this mountainous species, while high-altitude environments are more stable over time.     

Biomass and structure of planktonic communities along an air temperature gradient in subarctic Sweden

1. Air temperature will probably have pronounced effects on the composition of plankton communities in northern lake ecosystems, either via indirect effects on the export of essential elements from catchments or through direct effects of water temperature and the ice‐free period on the behaviour of planktonic organisms. 2. We assessed the role of temperature by comparing planktonic communities in 15 lakes along a 6 °C air temperature gradient in subarctic Sweden. 3. We found that the biomass of phytoplankton, bacterioplankton and the total planktonic biomass were positively related to air temperature, probably as a result of climatic controls on the export of nitrogen from the catchment (which affects phytoplankton biomass) and dissolved organic carbon (affecting bacterioplankton biomass). 4. The structure of the zooplankton community, and top down effects on phytoplankton, were apparently not related to temperature but mainly to trophic interactions ultimately dependent on the presence of fish in the lakes. 5. Our results suggest that air temperature regimes and long‐term warming can have strong effects on the planktonic biomass in high latitude lakes. Effects of temperature on the structure of the planktonic community might be less evident unless warming permits the invasion of fish into previous fishless lakes.     

Carbon isotope composition of N2-fixing and N-fertilized legumes along an elevational gradient

Dinitrogen-fixing legumes are frequently assumed to be less water-use efficient than plants utilizing soil mineral N, because of the high respiratory requirements for driving N₂ fixation. However, since respiration is assumed not to discriminate against ¹³C, any differences in water-use efficiency exclusively due to respiration should not be apparent in carbon isotope discrimination () values. Our objective was to determine if the source of N (N₂ fixation versus soil N) had any effect on of field-grown grain legumes grown at different elevations. Four legume species, Glycine max, Phaseolus lunatus, P. vulgaris, and Vigna unguiculata, were grown on five field sites spanning a 633 m elevational gradient on the island of Maui, Hawaii. The legumes were either inoculated with a mixture of three effective strains of rhizobia or fertilized weekly with urea at 100 kg N ha^-1 in an attempt to completely suppress symbiotic N₂-fixing activity. In 14 of 20 analyses of stover and 12 of 15 analyses of seed values were significantly higher (p=0.10) in the inoculated plants than the N-fertilized plants. Nitrogen concentrations were generally higher in the fertilized treatments than the inoculated treatments. The different values obtained depending on N-source may have implications in using as an indicator of water-use efficiency or yield potential of legumes.     

Intraspecific variability of leaf cuticle alkanes in Sedum lanceolatum along an elevational gradient

Characterization of variability patterns in leaf n-alkanes isolated from Sedum lanceolatum field populations indicates little correlation with elevation gradients. Inherent variability is better explained by genetic constitution than phenotypic plasticity, thus suggesting that leaf wax samples isolated from field-collected specimens are suitable for use in elucidation of systematic relationships.