Potential warm-stage microrefugia for alpine plants: Feedback between geomorphological and biological processes

During interglacial stages, microrefugia are sites that support locally favorable climates within larger areas with unfavorable warmer climates. Despite recent theoretical representations of microrefugia, an appropriate ecological characterization is still lacking, mostly for warm periods. Across mountain/alpine areas, cold-adapted plant species could adopt different strategies to manage the effects of climate warming: (A) migration toward higher elevations and summits; (B) in situ resilience of communities and species populations within microrefugia; and C) adaptation and evolution by genetic differentiation. This review aims to distinguish and characterize from an ecological perspective glacial, nival, periglacial and composite landforms and deposits that may function as potential microrefugia during interglacial warm periods.We conducted a literature screening related to the geomorphological processes and landforms associated with vegetation and plant communities in alpine/mountain environments of Europe. They include glacial deposits rock glaciers, debris-covered glaciers, composite cones and channels. In Alpine regions, geomorphologic niches that constantly maintain cold-air pooling and temperature inversions are the main candidates for microrefugia. Within such microrefugia, microhabitat diversity modulates the responses of plants to disturbances caused by geomorphologic processes and supports their aptitude for surviving under extreme conditions on unstable surfaces in isolated patches. Currently, European marginal mountain chains may be considered as examples of macrorefugia where relict boreo-alpine species persist within peculiar geomorphological niches that act as microrefugia.This review contributes to identifying potential warm-stage microrefugia areas across alpine and mountain regions and determining certain landforms that play or may play such role under global-change scenarios. The occurrence of warm-stage microrefugia within these locations may be of great importance for the modeling of future distributions of species and assessing the risk of extinction for alpine species. Microrefugia may have important implications in micro-evolutionary processes that occur across alternating climatic phases.     

A climatic basis for microrefugia: the influence of terrain on climate

There is compelling evidence from glacial and interglacial periods of the Quaternary of the utilization of microrefugia. Microrefugia are sites that support locally favorable climates amidst unfavorable regional climates, which allow populations of species to persist outside of their main distributions. Knowledge of the location of microrefugia has important implications for climate change research as it will influence our understanding of the spatial distribution of species through time, their patterns of genetic diversity, and potential dispersal rates in response to climate shifts. Indeed, the implications of microrefugia are profound and yet we know surprisingly little about their climatic basis; what climatic processes can support their subsistence, where they may occur, their climatic traits, and the relevance of these locations for climate change research. Here I examine the climatic basis for microrefugia and assert that the interaction between regional advective influences and local terrain influences will define the distribution and nature of microrefugia. I review the climatic processes that can support their subsistence and from this climatic basis: (1) infer traits of the spatial distribution of microrefugia and how this may change through time; (2) review assertions about their landscape position and what it can tell us about regional climates; and (3) demonstrate an approach to forecasting where microrefugia may occur in the future. This synthesis highlights the importance of landscape physiography in shaping the adaptive response of biota to climate change.     

Fine‐grain,large‐domain climate models based on climate station and comprehensive topographic information improve microrefugia detection

Large‐domain species distribution models (SDMs) fail to identify microrefugia, as they are based on climate estimates that are either too coarse or that ignore relevant topographic climate‐forcing factors. Climate station data are considered inadequate to produce such estimates, a viewpoint we challenge here. Using climate stations and topographic data, we developed three sets of large‐domain (450 000 km²), fine‐grain (50 m) temperature grids accounting for different levels of topographic complexity. Using these fine‐grain grids and the Worldclim data, we fitted SDMs for 78 alpine species over Sweden, and assessed over‐ versus underestimations of local extinction and area of microrefugia by comparing modelled distributions at species' rear edges. Accounting for well‐known topographic climate‐forcing factors improved our ability to model fine‐scale climate, despite using only climate station data. This approach captured the effect of cool air pooling, distance to sea, and relative humidity on local‐scale temperature, but the effect of solar radiation could not be accurately accounted for. Predicted extinction rate decreased with increasing spatial resolution of the climate models and with increasing number of topographic climate‐forcing factors accounted for. About half of the microrefugia detected in the most topographically complete models were not detected in the coarser SDMs and in the models calibrated from climate variables extracted from elevation only. Although major limitations remain, climate station data can potentially be used to produce fine‐grain topoclimate grids, opening up the opportunity to model local‐scale ecological processes over large domains. Accounting for the topographic complexity encountered within landscapes permits the detection of microrefugia that would otherwise remain undetected. Topographic heterogeneity is likely to have a massive impact on species persistence, and should be included in studies on the effects of climate change.     

Bridging the gap between microclimate and microrefugia: A bottom-up approach reveals strong climatic and biological offsets

In the context of global warming, a clear understanding of microrefugia—microsites enabling the survival of species populations outside their main range limits—is crucial. Several studies have identified forcing factors that are thought to favor the existence of microrefugia. However, there is a lack of evidence to conclude whether, and to what extent, the climate encountered within existing microrefugia differs from the surrounding climate. To investigate this, we adopt a “bottom-up” approach, linking marginal disconnected populations to microclimate. We used the southernmost disconnected and abyssal populations of the circumboreal herbaceous plant Oxalis acetosella in Southern France to study whether populations in sites matching the definition of “microrefugia” occur in particularly favorable climatic conditions compared to neighboring control plots located at distances of between 50 to 100 m. Temperatures were recorded in putative microrefugia and in neighboring plots for approximately 2 years to quantify their thermal offsets. Vascular plant inventories were carried out to test whether plant communities also reflect microclimatic offsets. We found that current microclimatic dynamics are genuinely at stake in microrefugia. Microrefugia climates are systematically colder compared to those found in neighboring control plots. This pattern was more noticeable during the summer months. Abyssal populations showed stronger offsets compared to neighboring plots than the putative microrefugia occurring at higher altitudes. Plant communities demonstrate this strong spatial climatic variability, even at such a microscale approach, as species compositions systematically differed between the two plots, with species more adapted to colder and moister conditions in microrefugia compared to the surrounding area.     

Hiding from the climate: Characterizing microrefugia for boreal forest understory species

Climate warming is likely to shift the range margins of species poleward, but fine‐scale temperature differences near the ground (microclimates) may modify these range shifts. For example, cold‐adapted species may survive in microrefugia when the climate gets warmer. However, it is still largely unknown to what extent cold microclimates govern the local persistence of populations at their warm range margin. We located 99 microrefugia, defined as sites with edge populations of 12 widespread boreal forest understory species (vascular plants, mosses, liverworts and lichens) in an area of ca. 24,000 km² along the species' southern range margin in central Sweden. Within each population, a logger measured temperature eight times per day during one full year. Using univariate and multivariate analyses, we examined the differences of the populations' microclimates with the mean and range of microclimates in the landscape, and identified the typical climate, vegetation and topographic features of these habitats. Comparison sites were drawn from another logger data set (n = 110), and from high‐resolution microclimate maps. The microrefugia were mainly places characterized by lower summer and autumn maximum temperatures, late snow melt dates and high climate stability. Microrefugia also had higher forest basal area and lower solar radiation in spring and autumn than the landscape average. Although there were common trends across northern species in how microrefugia differed from the landscape average, there were also interspecific differences and some species contributed more than others to the overall results. Our findings provide biologically meaningful criteria to locate and spatially predict potential climate microrefugia in the boreal forest. This opens up the opportunity to protect valuable sites, and adapt forest management, for example, by keeping old‐growth forests at topographically shaded sites. These measures may help to mitigate the loss of genetic and species diversity caused by rear‐edge contractions in a warmer climate.     

Climate refugia for Pinus spp. in topographic and bioclimatic environments of the Madrean sky islands of México and the United States

Climate refugia, or places where habitats are expected to remain relatively buffered from regional climate extremes, provide an important focus for science and conservation planning. Within high-priority, multi-jurisdictional landscapes like the Madrean sky islands of the United States and México, efforts to identify and manage climate refugia are hindered by the lack of high-quality and consistent transboundary datasets. To fill these data gaps, we assembled a bi-national field dataset (n?=?1416) for five pine species (Pinus spp.) and used generalized boosted regression to model pine habitats in relation to topographic variability as a basis for identifying potential microrefugia at local scales in the context of current species’ distribution patterns. We developed additional models to quantify climatic refugial attributes using coarse scale bioclimatic variables and finer scale seasonal remote sensing indices. Terrain metrics including ruggedness, slope position, and aspect defined microrefugia for pines within elevation ranges preferred by each species. Response to bioclimatic variables indicated that small shifts in climate were important to some species (e.g., P. chihuahuana, P. strobiformis), but others exhibited a broader tolerance (e.g., P. arizonica). Response to seasonal climate was particularly important in modeling microrefugia for species with open canopy structure and where regular fires occur (e.g., P. engelmannii and P. chihuahuana). Hotspots of microrefugia differed among species and were either limited to northern islands or occurred across central or southern latitudes. Mapping and validation of refugia and their ecological functions are necessary steps in developing regional conservation strategies that cross jurisdictional boundaries. A salient application will be incorporation of climate refugia in management of fire to restore and maintain pine ecology. Una versión en español de este artículo está disponible como descarga.

     

Fine-Scale Habitat Selection by Breeding White-Tailed Ptarmigan in Colorado

Species distributions are influenced by climate and topography in alpine ecosystems, yet resource selection studies of alpine species are uncommon. Basic characteristics of habitats used by alpine-endemic white-tailed ptarmigan (Lagopus leucura) have been described to explain foraging behavior, morphology, and survival in many alpine regions; however, there is a lack of information about fine-scale habitat selection for nesting and brood-rearing, particularly in the southern extent of the species’ range. Few studies have tested whether nest and brood-site selection by white-tailed ptarmigan are influenced by fine-scale components such as vegetation and arthropod communities. We assessed these fine-scale habitat characteristics analyzing paired use-available resource selection for nest (n = 61) and brood (n = 54) sites. We used conditional logistic regression for data collected in 2 alpine areas along the Front Range of Colorado, USA, during 2014 and 2015. We evaluated resource selection at larger (patch) and finer (nest site) scales. Nest-site selection at the patch scale was best predicted by cover (%) of forage forbs, rock and gravel, and shrubs. Forage forb cover explained more variation in our top nest model at the patch scale when compared to models with specific vegetation species. Females placed their nests along elevational gradients but more so at lower elevations and selected for less graminoid cover at the nest-site scale. Brood habitat selection at the patch level was influenced by cover (%) of rock and gravel and proximity to shrubs (m). Analysis of a subset of our brood data (n = 34) revealed females selected brood habitat that contained high arthropod abundance (e.g., Cicadellidae) over high vegetation cover, likely as a response to meet dietary requirements of chicks. Our results demonstrate how and where white-tailed ptarmigan are currently selecting these different breeding sites in Colorado's alpine, giving us insight into consequences this alpine-endemic bird may face if their breeding habitat is altered. © 2019 The Wildlife Society.     

Topographically controlled thermal‐habitat differentiation buffers alpine plant diversity against climate warming

Aim We aim to: (1) explore thermal habitat preferences in alpine plant species across mosaics of topographically controlled micro‐habitats; (2) test the predictive value of so‐called ‘indicator values’; and (3) quantify the shift in micro‐habitat conditions under the influence of climate warming. Location Alpine vegetation 2200–2800 m a.s.l., Swiss central Alps. Methods High‐resolution infra‐red thermometry and large numbers of small data loggers were used to assess the spatial and temporal variation of plant‐surface and ground temperatures as well as snow‐melt patterns for 889 plots distributed across three alpine slopes of contrasting exposure. These environmental data were then correlated with Landolt indicator values for temperature preferences of different plant species and vegetation units. By simulating a uniform 2 K warming we estimated the changes in abundance of micro‐habitat temperatures within the study area. Results Within the study area we observed a substantial variation between micro‐habitats in seasonal mean soil temperature (ΔT = 7.2 K), surface temperature (ΔT = 10.5 K) and season length (>32 days). Plant species with low indicator values for temperature (plants commonly found in cool habitats) grew in significantly colder micro‐habitats than plants with higher indicator values found on the same slope. A 2 K warming will lead to the loss of the coldest habitats (3% of current area), 75% of the current thermal micro‐habitats will be reduced in abundance (crowding effect) and 22% will become more abundant. Main conclusions Our results demonstrate that the topographically induced mosaics of micro‐climatic conditions in an alpine landscape are associated with local plant species distribution. Semi‐quantitative plant species indicator values based on expert knowledge and aggregated to community means match measured thermal habitat conditions. Metre‐scale thermal contrasts significantly exceed IPCC warming projections for the next 100 years. The data presented here thus indicate a great risk of overestimating alpine habitat losses in isotherm‐based model scenarios. While all but the species depending on the very coldest micro‐habitats will find thermally suitable ‘escape’ habitats within short distances, there will be enhanced competition for those cooler places on a given slope in an alpine climate that is 2 K warmer. Yet, due to their topographic variability, alpine landscapes are likely to be safer places for most species than lowland terrain in a warming world.     

Experimental evaluation of seed limitation in alpine snowbed plants

Background

The distribution and abundance of plants is controlled by the availability of seeds and of sites suitable for establishment. The relative importance of these two constraints is still contentious and possibly varies among species and ecosystems. In alpine landscapes, the role of seed limitation has traditionally been neglected, and the role of abiotic gradients emphasized.

Methodology/Principal Findings

We evaluated the importance of seed limitation for the incidence of four alpine snowbed species (Achillea atrata L., Achillea clusiana Tausch, Arabis caerulea L., Gnaphalium hoppeanum W. D. J. Koch) in local plant communities by comparing seedling emergence, seedling, juvenile and adult survival, juvenile and adult growth, flowering frequency as well as population growth rates λ of experimental plants transplanted into snowbed patches which were either occupied or unoccupied by the focal species. In addition, we accounted for possible effects of competition or facilitation on these rates by including a measure of neighbourhood biomass into the analysis. We found that only A. caerulea had significantly lower seedling and adult survival as well as a lower population growth rate in unoccupied sites whereas the vital rates of the other three species did not differ among occupied and unoccupied sites. By contrast, all species were sensitive to competitive effects of the surrounding vegetation in terms of at least one of the studied rates.

Conclusions/Significance

We conclude that seed and site limitation jointly determine the species composition of these snowbed plant communities and that constraining site factors include both abiotic conditions and biotic interactions. The traditional focus on abiotic gradients for explaining alpine plant distribution hence appears lopsided. The influence of seed limitation on the current distribution of these plants casts doubt on their ability to readily track shifting habitats under climate change unless seed production is considerably enhanced under a warmer climate.     

Formation of disjunct plant distributions in Northeast Asia: a case study of <Emphasis Type="Italic">Betula davurica</Emphasis> using a species distribution model

Repeated climate change during glacial and interglacial periods of the Quaternary led to mass migrations that resulted in disjunct distributions for many species. However, few studies have examined the processes that form disjunct distributions in Northeast Asia (NEA). In this study, we examined the disjunct distribution of Betula davurica Pall. in the Japanese archipelago. This species is a dominant canopy tree found in cool-temperate deciduous broad-leaved forests of continental NEA. We hypothesized that Quaternary climate change caused the present disjunct distribution pattern of this species. To test this hypothesis, we adopted a species distribution model and examined a series of potential habitats in the Last Glacial Maximum (LGM), the mid-Holocene, and the present. We generated models in MaxEnt with B. davurica presence as the response variable and six bioclimatic variables as predictor variables. During the LGM, projected potential habitats were distributed around the Korean Peninsula, East China, and the Japanese archipelago, excluding Hokkaido. In the mid-Holocene, habitats retreated both from East China and western Japan, remained unchanged in the Korean Peninsula and central Honshu mountains, and expanded to northern China, the Russian Far East, as well as northern Japan (Hokkaido). Thus, post-LGM global warming led to an expansion of B. davurica distribution to northern parts of continental NEA, along with a retreat in the Japanese archipelago. This shift in populations formed the present disjunct distribution.     

Dispersal and microsite limitation of a rare alpine plant

Knowledge on the limitation of plant species’ distributions is important for preserving alpine biodiversity, particularly when the loss of alpine habitats because of global warming or land use changes is faster than colonization of new habitats. We investigated the potential of the rare alpine plant Campanula thyrsoides L. to colonize grassland sites of different suitability on a small mountain plateau in the Swiss Alps. A total of 15 experimental sites were selected according to their differences in habitat suitability for adult C. thyrsoides, which was measured by the Beals index. At each site we applied a disturbance treatment, added seeds at different densities and monitored the survival of seedlings over two consecutive years. The number of surviving seedlings was not positively related to habitat suitability for adult C. thyrsoides. Furthermore, C. thyrsoides appears to be strongly dispersal limited at the regional scale because seed addition to unoccupied habitats resulted in successful germination and survival of seedlings. Since an increase of seed density in already occupied sites did not affect the number of seedlings, we suggest that C. thyrsoides is microsite limited at the local scale. Microsite limitation is supported by the result that seedling survival of the species was enhanced in vegetation gaps created by disturbance. We conclude that C. thyrsoides may become endangered in the future if environmental changes cause local extinction of populations. An appropriate management, such as a disturbance regime for enhancing recruitment in existing populations, may ensure the long-term survival of this rare alpine plant species.     

Testing the environmental warming responses of Brachyscome daisy species using a common garden approach

As temperatures increase in a warming world, there will be different responses among related plant species, with some species able to increase growth rate under warmer conditions and others less likely. Here, we identify survival and growth parameters in a group of 19 related Australian daisies from the genera Brachyscome and Pembertonia when exposed to higher soil temperature, focusing particularly on species from the alpine environment. We used a common garden approach to measure growth and survival under warming. We tested for the effects of evolutionary history by investigating phylogeny and testing for a phylogenetic signal, and for the effects of ecological history by considering climatic variables associated with species distributions in their native range. Evolutionary history did not have a detectable effect on warming responses. While there was a moderate signal for plant growth in the absence of warming, there was no signal for growth changes in response to warming, despite variability among species to warming that ranged from positive to negative growth responses. There was no strong effect of climate context, as species that showed a positive response to warming did not necessarily originate from hotter environments. In fact, several species from hot environments grew relatively poorly when exposed to higher soil temperature. However, species endemic to alpine areas were less likely to benefit from warming than widespread species. We found a strong phylogenetic signal for climate history, in that closely related species tend to occur in areas with similar annual variability in precipitation. Species differences in response to soil warming were variable and difficult to link to climate conditions except for the poor response of alpine endemics. There was no significant association between survival and warming responses of species. However, as some species showed weak growth responses, this may reduce their fitness into the future.     

Interglacial microrefugia and diversification of a cactus species complex: phylogeography and palaeodistributional reconstructions for Pilosocereus aurisetus and allies

The role of Pleistocene climate changes in promoting evolutionary diversification in global biota is well documented, but the great majority of data regarding this subject come from North America and Europe, which were greatly affected by glaciation. The effects of Pleistocene changes on cold‐ and/or dry‐adapted species in tropical areas where glaciers were not present remain sparsely investigated. Many such species are restricted to small areas surrounded by unfavourable habitats, which may represent potential interglacial microrefugia. Here, we analysed the phylogeographic structure and diversification history of seven cactus species in the Pilosocereus aurisetus complex that are restricted to rocky areas with high diversity and endemism within the Neotropical savannas of eastern South America. We combined palaeodistributional estimates with standard phylogeographic approaches based on two chloroplast DNA regions (trnT‐trnL and trnS‐trnG), exon 1 of the nuclear gene PhyC and 10 nuclear microsatellite loci. Our analyses revealed a phylogeographic history marked by multiple levels of distributional fragmentation, isolation leading to allopatric differentiation and secondary contact among divergent lineages within the complex. Diversification and demographic events appear to have been affected by the Quaternary climatic cycles as a result of isolation in multiple patches of xerophytic vegetation. These small patches presently harbouring P. aurisetus populations seem to operate as microrefugia, both at present and during Pleistocene interglacial periods; the role of such microrefugia should be explored and analysed in greater detail.     

Alpine community recruitment potential is determined by habitat attributes in the alpine ecosystems of the Himalaya‐Hengduan Mountains,SW China

The fragility and sensitivity to climate change of alpine ecosystems make it difficult to maintain the stability of their plant communities. Thus, it is important to determine which plant propagules are stored in the soils in order to understand community recruitment potential, especially under different environmental conditions. Based on a soil seed germination and seedling cultivation experiment, we aimed to identify differences in the soil seed attributes between three typical habitat types in the alpine subnival ecosystems of the Himalaya‐Hengduan Mountains and hence to predict the community recruitment potential of each of these different communities. We found that the seed assemblages in the soils differed between habitats. The most abundant taxa were from the genera Saxifraga, Kobresia, Arenaria, Polygonum, Draba, and Viola, while the taxa with lowest abundance were Apiaceae, Campanulaceae, Circaea, Crassulaceae, and Gentiana. Different habitats exhibited variable soil seed richness, diversity, and density. However, the patterns differed between study sites. Specifically, at Baima (BM) and Shika (SK) snow mountains, soil seed richness, diversity, and density were generally highest in grassland, followed by rock bed and bare ground. In contrast, on Jiaozi (JZ) snow mountain, the rock bed supported the highest soil seed richness and density, followed by grassland and bare ground. These results suggest that the attributes of habitats and communities can both affect the accumulation of soil seeds. Bare ground supports the lowest seed diversity and density but also harbors the most empty niches. We, therefore, predict that, once the thermal conditions become suitable as a result of global warming, this habitat has the potential to see greater changes than grassland and rock bed in terms of community recruitment.     

Do clonal growth form and habitat origin affect resource-induced plasticity in Tibetan alpine herbs?

To investigate how growth form and habitat origin affect phenotypic plasticity to resource supply in the Tibetan alpine herbs, the phalanx-type species Stipa capillacea and the guerilla-type species Carex montis-everestii were sampled from two different habitats (alpine steppe and alpine scrubland) and grown under three levels of light intensity and two levels of nutrient supply. Interspecific differences in light-induced plasticity were detected only in number of ramets, specific leaf area and leaf sheath length. Plasticity in plant biomass, number of ramets and rhizome length in response to light intensity differed between the two habitats. Stipa plants were more plastic than Carex plants in number of ramets and specific leaf area in response to light intensity. Carex plants from the alpine scrubland expressed greater light-induced plasticity in plant biomass and number of ramets than those from the alpine steppe, and Stipa plants showed less interhabitat differences in plasticity, which may be closely related to their contrasting growth forms. Clonal growth form and habitat origin affected nutrient-induced plasticity in none of the measured traits. It may be the guerilla growth form that makes Carex plants more efficiently adapted to highly heterogeneous light conditions in scrubland, and less habitat-dependent plasticity contributes to success of the phalanx-type Stipa plants in alpine habitats. The results are discussed in the context of foraging for heterogeneously distributed essential resources and adaptation to habitat origin.     

Ecological and evolutionary responses of an arctic plant to variation in microclimate and soil

The arctic and alpine regions are predicted to experience some of the highest rates of climate change, and the arctic vegetation is expected to be especially sensitive to such changes. Understanding the ecological and evolutionary responses of arctic plant species to changes in climate is therefore a key objective. Geothermal areas, where natural temperature gradients occur over small spatial scales, and without many of the confounding environmental factors present in latitudinal and other gradient studies, provide a natural experimental setting in which to examine the response of arctic–alpine plants to increasing temperatures. To test the ecological and evolutionary response of the circumpolar alpine bistort Persicaria vivipara to temperature, we collected plant material and soil from areas with low, intermediate and high soil temperatures and grew them at three different temperatures in a three-factorial growth chamber experiment. At higher experimental soil temperatures, sprouting was earlier and plants had more leaves. Sprouting was earlier in soil originating from intermediate temperature and plants had more leaves when grown in soil originating from low temperatures. We did not find evidence of local adaptation or genetic variation in reaction norms among plants originating from areas with low, intermediate and high soil temperature. Our findings suggest that the alpine bistort has a strong plastic response to warming, but that differences in soil temperature have not resulted in genetic differentiation. The lack of an observed evolutionary response may, for example, be due to the absence of temperature-mediated selection on P. vivipara, the low rate of sexual recombination, or high levels of gene flow balancing differences in selection. When placed within the context of other studies, we conclude that arctic–alpine plant species often show strong plastic responses to spring warming, while evidence of evolutionary responses varies among species.     

The lichens of high ground in the English Lake District

Abstract: A general account of the lichens of high-level acid habitats in the Lake District is provided. This is followed by detailed studies of base-rich sites in the Helvellyn range, acid outcrops in Langdale, and three upland tarns, one of which is of outstanding importance. The lichen vegetation is richer than in Snowdonia or the Northern Pennines, but its diversity is considerably less than that of certain individual mountains in the Scottish Highlands. The montane element is interpreted as representing the last remnants of communities that have declined gradually through the Postglacial Period. Many species are extremely rare. Current restraints on the alpine lichen flora include the climate, the predominance of wet, acid rock and heavy sheep grazing. Rhizocarpon simillimum is reported as new to the British lichen flora.     

Altitudinal distribution of the entire invasive small mammal guild in the eastern dryland zone of New Zealand’s Southern Alps

New Zealand faces significant ecological problems caused by the introduction of a variety of invasive small mammal species. Many of these species originate from temperate to subarctic climates and occur across wide elevations in their native range, and so arrived predisposed to adapt to a variety of habitats and bioclimatic zones in their new environs, including the alpine zone. Almost all of New Zealand’s invasive small mammal species have been recorded in the country’s alpine zones, yet neither the altitudinal distribution nor the extent to which such species use high elevation areas has been clearly defined. We conducted extensive camera trap surveys in summer to autumn periods of 2019 and 2020 across an elevation range of 500–2250 m above sea level, and used detection rates and occupancy modelling to reveal the altitudinal distributions and habitat associations of all 10 invasive small mammals that occur in the dryland zone of the central South Island. We found altitudinal distributions varied greatly across species, and that while most exhibited decreasing detection rates and site occupancy probabilities with increasing elevation, some used the subalpine and alpine zones to a greater degree than adjacent lower elevations. There were clear habitat associations, as well as interspecific associations that helped to explain the altitudinal distribution of some species. Understanding how such factors influence the distribution of invasive small mammals has both broad implications for invasive species management, and direct applications in evaluating threats to native taxa, advancing management strategies, and benchmarking distributions in a changing climate.

     

Vegetation of rock clefts and ledges in the Pamir Alai Mts, Tajikistan (Middle Asia)

The paper presents the results of phytosociological research conducted on the vegetation of rock clefts and ledges in the Pamir-Alai Mountains (Tajikistan, Middle Asia). During the field studies, done in 2010–2012, 101 phytosociological relevés were taken. Plant species were recorded according to the Braun-Blanquet cover-abundance scale. Communities of calcareous rock clefts and ledges with small soil amounts from several ranges (Zeravshan Mts, Hissar Mts, Hazratishokh Mts, Darvaz Mts, Rushan Mts and Vanch Mts), inhabiting mainly the alpine and subnival zone, have been described. A synopsis of the rock communities of the Pamir-Alai is proposed. In the examined vegetation plots 77 vascular plant and 6 moss species were noted. The most frequent were: Achoriphragma pinnatifidum, Artemisia rutifolia, Asperula albiflora, Campanula incanescens, C. lehmanniana, Parietaria judaica, Pentanema albertoregelia, Poa relaxa and Stipa zeravshanica among vascular plants and Brachythecium albicans and Bryum caespiticum among mosses. Most of them are narrow endemics of Tajikistan or Middle Asia. The collected material presents most of the variability among the phytocoenoses of large crevices and rock ledges in limestone massifs in the alpine and subnival zones. As a result of field research and numerical analyses, 7 associations have been distinguished: Achoriphragmetum pinnatifidi, Asperulo albiflorae-Stipetum zeravshanicae, Inuletum glaucae, Paraquilegietum anemonoidis, Pentanemetum albertoregeliae, Rhinactinidietum popovii and Saussureaetum ovatae. The distinctiveness of habitat and species composition of Middle Asiatic rock communities makes it necessary to distinguish a new suballiance, Pentanemenion albertoregeliae, within the Asperulo albiflorae-Poion relaxae alliance. The main factors determining the species composition of classified associations seem to be the elevation above sea level and exposition. Alpine rock communities are one of the most unique and interesting plant formations in the moutainous areas of Pamir-Alai. Despite not being species-rich they often harbour many specialists adapted to harsh and extreme environments, especially in areas of Mediterranean-like climate.