Standard energetics of phyllostomid bats: the inadequacies of phylogenetic-contrast analyses

The basal rates of metabolism (BMR) of bats belonging to the family Phyllostomidae are re-examined after an earlier correlation with food habits was rejected because it did not take phylogeny into consideration. This rejection was based on an erroneous attribution of food habits and on an analytical method, phylogenetic contrasts, that ignores interactions that occur among character states and preferentially attributes responsibility for character states to phylogeny. The re-examination made here was based on analysis of covariance, which makes no a priori assumptions on the relative impact of factors that influence character states and permits factor interactions to be identified. A resulting model, based on variation in body mass, food habits, occurrence with respect to elevation, and residence on islands or continents, accounts for 99.4% of the variation in the BMR of 30 species of phyllostomids. Basal rate is also correlated with subfamily, but only if food habits are excluded because they are correlated with subfamily affiliation, as is residence on islands and continents, two examples of factor interaction. The preference to assign the effects of food habits and island residence on basal rate to subfamily affiliation (and phylogeny) is not justified. The concept that quantitative physiological characters can be transmitted via phylogeny without regard to the habits of animals and the characteristics of their environments cannot be defended. Phylogeny is the historical context in which the evolution of character states occurs, not the 'cause' of their evolution.     

影响食肉目动物能量学的生态因子

在食肉目的62种动物中,体重的变异可以解释基础代谢率86.8%的变化。当栖息基底、食性、生境和纬度等4个因素与体重合起来一起分析,则可以解释基础代谢率98.7%的变化,即这些生态和行为因子可以解释代谢率残差变异的81.1%。身体成分也是影响基础代谢率的另一个因素,可以解释一些大型树栖种类的较低的代谢率。除体重因素外,导致真兽类基础代谢率变异的主要原因是:当生态因素适合时,高水平的能量消耗可以促进动物的高繁殖输出,而动物的某些习性和生存环境则会要求低能量消耗,从而使繁殖率降低。当以科为单元进行分析时,对结果没有影响。生理参数与分类单元之间大多数的相关性反映了生态和行为因素与分类系统之间的一种粗略的相关关系。除非系统学可以反映动物的生态学和行为学,否则系统学不能决定动物适应性特征的状态     

The comparative energetics of 'caviomorph' rodents.

The energetics of 11 species of New World hystricognath ('caviomorph') rodents are presented and compared with data from the literature on 19 additional species. Log(10) body mass alone accounts for 94% of the variation in the log(10) basal rate of metabolism in caviomorphs. The residual variation in basal rate is correlated with the stratum on which species live: arboreal species have low basal rates; terrestrial and fossorial species have intermediate basal rates; and aquatic species have high basal rates. When stratum is not included in the analysis, folivores, especially those that are arboreal, have lower basal rates than species with other food habits when combined with log(10) body mass. Small island endemics, all of which are folivores, have basal rates that are 61% of continental species. Log(10) basal rate correlates with family affiliation when combined with log(10) mass, but only if no other factor is included. Therefore, caviomorphs with low basal rates are arboreal, folivorous, live on small islands and belong to the Capromyidae, whereas other character combinations are associated with higher basal rates. These observations demonstrate that the basal rates of caviomorphs reflect many factor interactions. No differences in basal rate were found to reflect climate. Log(10) mass, the only factor to correlate with conductance, accounts for 82% of the variation in log(10) minimal thermal conductance. Mean interspecific body temperature was 36.9 degrees C; it was lowest in aquatic and fossorial species.     

Analysis of factors that influence energy expenditure in honeyeaters (Meliphagidae)

The energy expenditure of the tūī (Prosthemadera novaeseelandiae), a meliphagid endemic to New Zealand, was measured and compared with 20 species of honeyeaters (family Meliphagidae) to determine whether its expenditure is influenced either by life in a moist, temperate climate or an island residence. Body mass in the honeyeaters accounted for 91.5% of the variation in basal rate. The combination of body mass, climate and the maximal limit to an altitudinal distribution explained 98.6% of the variation in basal rate with tropical, low-altitude species having the highest mass-independent rate. The basal rates of meliphagids in tropical highlands are similar to those in temperate lowlands, which may reflect similar food supplies. The tūī mass-independent expenditure appears to reflect an active lifestyle in a temperate climate with no evidence that an island residence influenced its rate, whereas sedentary birds on New Zealand have responded to island life with a depressed basal rate. An effective analysis of the variation in energy expenditure requires the inclusion of the ecological and behavioural characteristics that distinguish species.     

The evolution of energetics in eutherian “insectivorans”: an alternate approach

An analysis of standard energetics in 57 species of “insectivorans”, small eutherians that preferentially feed on soil invertebrates, indicated that a combination of climate, the use of torpor, substrate, food habits, and log₁₀ body mass accounted for 92.5% of the variation in log10 basal rate of metabolism in insectivorans, whereas log₁₀ body mass alone accounted for 76.7% of the variation. With the addition of subfamily affiliation, this analysis accounted for 95.5% of the variation in log₁₀ basal rate, the most distinctive subfamilies being Soricinae and the Talpinae, which have equally high basal rates.Sorex species have basal rates that average 2.5 times those of tropical crocidurines, reflecting an approach bySorex to life in cold climates that does not include the use of torpor, a stratagem widely used by crocidurines in warm-temperate and tropical climates. The absence of torpor inSorex may facilitate a high reproductive rate through a high basal rate of metabolism, a combination that may be incompatible with a small mass, insectivorous food habits, and life in the lowland tropics, but required in cold-temperate environments. Insectivorans other than shrews, moles, and cold-temperature hedgehogs have low basal rates principally in association with tropical distributions and the use of torpor. Basal rate of metabolism in insectivorans also correlated with ordinal, familial, subfamilial, and tribal affiliations. The suggestion that phylogeny is an important determinant of performance characters like rate of metabolism ignores the requirement that performance must be compatible with conditions in the environment and with a species’ other characteristics. The principal reason why performance characters are correlated with taxonomic affiliation is that many clades consist of species that share a common approach to the environment. Thus, clades not only represent evolutionary histories, they also are crude measures of physiological and behavioral performances.     

Flightless rails endemic to islands have lower energy expenditures and clutch sizes than flighted rails on islands and continents

Data are presented on the standard energetics of six flighted and five flightless species of rails (Aves: Rallidae). The factors influencing these data and those from three additional species available from the literature, one of which was flightless, are examined. Basal rate of metabolism correlates with body mass, residency on islands or continents, volant condition, pectoral muscle mass, and food habits, but not with climate. The greatest capacity (96.2%) to account for the variation in basal rate of metabolism in 15 populations that belong to the 14 species occurs when body mass, volant condition, and food habits are combined. Then flighted species have basal rates that average 1.38 times those of flightless species and herbivorous rails have basal rates that are 1.37 times those of omnivorous species, which means that, independent of body mass, flighted gallinules have basal rates that are 1.9 times those of flightless, omnivorous rails. Distribution, pectoral muscle mass, and flight ability cannot be combined in the same analysis because they code for similar information. The evolution of a flightless condition in rails requires the absence of eutherian predators, but has occurred in the presence of marsupial predators. Each of the six studied flightless rails independently evolved a flightless condition and a low basal rate, whereas the evolution of herbivory and an associated high basal rate evolved at least twice in these species. Flightless rails on islands have clutch sizes that are only about one-half those of flighted rails living on continents, the reduction in clutch size correlating with a reduction in basal rate of metabolism. Thermal conductance in rails is correlated with body mass and food habits: herbivorous rails had conductances that were 1.43 times those of omnivores, i.e., conductances are highest in species with the highest basal rates.     

The comparative energetics of rigid endothermy: the Arvicolidae

Rate of metabolism and body temperature were examined in 24 species of arvicolid rodents, for 15 of which data are presented here, to determine the factors that influence their level of energy expenditure. Arvicolids are characterized by a high, precisely-regulated body temperature, a high basal rate of metabolism by general mammalian standards, and a standard thermal conductance, except at large masses, when low conductances can occur. No evidence of entrance into torpor is known for any arvicolid; its absence is associated with especially high basal rates at masses smaller than 52 g. Arvicolids that live in cold climates, i.e. at high altitudes and latitudes, have higher basal rates than species that live in other environments. Basal rates, however, appear to be independent of food habits, presumably because of the small mass of most species and possibly because all species are herbivorous. A small size in combination with herbivory has permitted arvicolids to maintain continuous endothermy, which in turn has permitted them to exploit cool-to-cold environments by means of a high rate of production. The use of daily or seasonal torpor might well have prevented arvicolids from attaining the pivotal position in energy transfer that they presently occupy in many high-latitude communities.     

Energetics of East African Pycnonotids1

Rate of oxygen consumption was measured in five bulbuls (Family Pycnonotidae) from western Uganda to evaluate whether this group is indeed characterized by the very low basal rates of metabolism previously reported. For three of these species, body temperature and rate of metabolism were measured as a function of ambient temperature from 10°C to 35°C. In these species body temperature was highly variable, and declined with ambient temperature in Andropadus virens. Such variation, in conjunction with behavioral adjustments, may reduce heat loss at low ambient temperatures. Body mass accounted for 98 percent of the variation in the basal rates of metabolism presented here. Basal rates in these species ranged from 81 to 90 percent of values predicted by the Aschoff–Pohl relationship for passerines, whereas previous measurements ranged from 56 to 72 percent of predicted values. This difference may reflect differences in species or measurement techniques, which, if the latter, suggests that the reduction in metabolic rate in this family may be less than originally thought. These data underline the importance of continued data collection on the metabolism of tropical birds, few of which have been measured to date.     

Physiological convergence amongst ant-eating and termite-eating mammals

Ant- and termite-eating are among the few food habits common to monotremes, marsupials, and eutherians. Data are reported on the rate of metabolism and temperature regulation of 14 species of mammals having these food habits, including two monotremes, one marsupial and 11 eutherians. Small mammals with these habits have comparatively high body temperatures and high basal rates of metabolism, but ant- and termite-eaters that weigh more than 1 kg generally have low body temperatures and low basal rates of metabolism. The higher basal rates in small species ensure effective temperature regulation. Low body temperatures in large species principally result from low rates of metabolism. Rates of metabolism are low in these mammals because they use a food that has a limited availability and a low energy density, the density being further decreased in large species by the ingestion of non-nutritive material during feeding. Burrowing habits in some large species also contribute to low rates of metabolism. The combination of body size, food habits, and presence or absence of burrowing behaviour can account for all but about 6% of the range in basal rate in ant- and termite-eaters. Ants and termites, because of their locally clumped distributions, permit a larger mass in terrestrial predators than do other invertebrate prey. The reason why so many "primitive" mammals feed on ants and termites is that, once evolved, mammals with these habits are nearly impossible to displace ecologically, because much of ecological replacement is associated with high rates of reproduction, which are themselves correlated with high rates of metabolism in eutherians. Consequently, the ecological replacement of ant- and termite-eaters is inhibited, because this food habit does not permit high rates of metabolism, except at small masses.     

Food habits and the basal rate of metabolism in birds

Summary The correlation of basal rate of metabolism with various factors is examined in birds. Chief among these is body mass. As in mammals, much of the remaining variation in basal rate among birds is associated with food habits. Birds other than passerines that feed on grass, nectar, flying insects, or vertebrates generally have basal rates that are similar to mammals of the same mass and food habits. In contrast, most invertebrate-eating birds that weigh over 100 g have higher basal rates than equally-sized, invertebrate-eating mammals. The high basal rates of small passerines equal those of small mammals that do not enter torpor and represent the minimal cost of continuous endothermy. Large passerines and small procellariiforms, charadriiforms, and psittaciforms generally have higher basal rates than mammals with the same mass and food habits. The high basal rates of passerines (in combination with altricial habits) may have significance in permitting high post-natal growth rates and the exploitation of seasonally abundant resources. These interrelations may contribute to the predominance of passerines in temperate land environments.     

Energetics of New Zealand's temperate parrots

Abstract

As in most parrots (Order: Psittaciformes) studied, New Zealand species have, independent of body mass, high basal rates of metabolism, low thermal conductances, and precisely regulated body temperatures. An analysis of covariance showed that basal rate in parrots correlates with body mass and thermal climate; temperate species have basal rates that are 21% higher than those of tropical species; and New Zealand's parrot have basal rates 32% higher than expected from body mass. Present information suggests that basal rate in parrots appears to be correlated neither with water availability in the environment nor with food habits. High basal rates in parrots are associated with large pectoral muscle masses. The very small pectoral muscle masses of the kakapo, Strigops habroptilus, predict that this flightless parrot has a low basal rate.     

Food habits, energetics, and the reproduction of marsupials

Basal rate of metabolism in marsupials and in eutherian mammals is principally correlated with body mass, food habits and activity. Feeding on fruit, the leaves of woody plants, or invertebrates is associated with low basal rates, especially at large masses, in both groups of mammals. These foods lead to low basal rates because they are seasonally unavailable, are indigestible, or need to be detoxified. The depression in basal rate associated with frugivory and folivory is increased when coupled with sedentary, arboreal habits in both marsupials and eutherians. In contrast, eutherians that feed on vertebrates or herbs generally have high basal rates, while marsupials that eat these foods do not have high basal rates. These foods permit high basal rates, which are exploited by eutherians because high basal rates in these mammals lead to high rates of reproduction. Marsupials have, at best, a limited correlation of reproduction with rate of metabolism, so that feeding on vertebrates or herbs does not lead to high basal rates in these mammals. This difference between marsupials and eutherians in the coupling of reproduction to energetics has at least two ecological consequences. 1) Marsupials generally do not tolerate cold-temperate environments because they do not accelerate growth and development to complete reproduction within a short spring and summer. 2) Marsupials coexist with ecologically similar eutherians as long as marsupials have food habits that are correlated with low rates of metabolism in eutherians (i.e. they feed on fruit, the leaves of woody plants, or invertebrates), but they tend to be displaced by eutherians when marsupials have food habits that are associated with high rates of metabolism in eutherians (i.e. when they feed on vertebrates and, probably, herbs).     

Ecological factors affect the level and scaling of avian BMR

The basal rate of metabolism (BMR) in 533 species of birds, when examined with ANCOVA, principally correlates with body mass, most of the residual variation correlating with food habits, climate, habitat, a volant or flightless condition, use or not of torpor, and a highland or lowland distribution. Avian BMR also correlates with migratory habits, if climate and a montane distribution is excluded from the analysis, and with an occurrence on small islands if a flightless condition and migration are excluded. Residual variation correlates with membership in avian orders and families principally because these groups are behaviorally and ecologically distinctive. However, the distinction between passerines and other birds remains a significant correlate of avian BMR, even after six ecological factors are included, with other birds having BMRs that averaged 74% of the passerine mean. This combination of factors accounts for 97.7% of the variation in avian BMR. Yet, migratory species that belong to Anseriformes, Charadriiformes, Pelecaniformes, and Procellariiformes and breed in temperate or polar environments have mass-independent basal rates equal to those found in passerines. In contrast, penguins belong to an order of polar, aquatic birds that have basal rates lower than passerines because their flightless condition depresses basal rate. Passerines dominate temperate, terrestrial environments and the four orders of aquatic birds dominate temperate and polar aquatic environments because their high BMRs facilitate reproduction and migration. The low BMRs of tropical passerines may reflect a sedentary lifestyle as much as a life in a tropical climate. Birds have BMRs that are 30-40% greater than mammals because of the commitment of birds to an expensive and expansive form of flight.     

Standard energetics of leaf-nosed bats (Hipposideridae): its relationship to intermittent- and protracted-foraging tactics in bats and birds

Basal rates of metabolism within the insectivorous genera Hipposideros and Ascelliscus, Old World leaf-nosed bats (Hipposideridae), ranged from 58% to 77% of the mammalian standard. The larger species, Hipposideros diadema and Hipposideros maggietaylori, effectively thermoregulated at ambient temperatures down to 9 degrees C, whereas two smaller species, Hipposideros galeritus and Hipposideros cervinus, occasionally permitted body temperatures to fall below 32 degrees C. The low basal rates of metabolism in hipposiderids correlated with a predatory life-style characterized by intermittent flight from a perch to capture insects, a correlation similar to that found in nonpasserine birds. Intermittent-foraging bats and nonpasserines collectively had basal rates of metabolism that averaged 75% of those that pursue insects during protracted flight. However, no difference in basal rate was found between protracted- and intermittent-foraging passerines, which had basal rates 1.8- and 2.4-times those of protracted-foraging and intermittent-foraging bats and nonpasserines, respectively. Bats, swifts, and caprimulgids that enter torpor have basal rates that are 85% of those of similar species that do not enter torpor. Body mass, order affiliation, foraging mode, and propensity to enter into torpor collectively account for 97% of the variation in basal rate of metabolism in insectivorous bats and birds. Foraging style therefore appears to be a factor contributing to the diversity in endotherm energetics. Minimal thermal conductance in the genus Hipposideros ranged from 75% to 102% of the mammalian standard. Birds have minimal thermal conductances that are 75% of mammals and intermittent foragers have minimal conductances that are 78% of protracted foragers.     

Altitudinal migration by birds: a review of the literature and a comprehensive list of species

Altitudinal migration is the seasonal altitudinal movement of birds from breeding areas to non‐breeding or wintering areas at different elevations. Although this type of migration is widely reported, questions remain concerning the number of species that perform altitudinal migration, possible variation among different taxa and geographic locations in the extent of altitudinal migration, and the foraging guilds of altitudinal migrants. We conducted an extensive bibliographic survey and compiled a list of altitudinal migrant birds worldwide. We characterized species in terms of their foraging guilds because the spatial distribution of food resources along altitudinal gradients is often evoked as a driver of bird altitudinal migration. We identified 1238 species of altitudinal migrants, ~10% of the ~10,000 extant species of birds. We found a strong geographic bias in publications focusing on avian altitudinal migration toward the United States and Costa Rica, and a paucity of studies in megadiverse regions such as the Afrotropical and Indomalayan realms, and areas in the Neotropics other than Costa Rica. We also found that most species of altitudinal migrants were invertivores rather than frugivores or nectarivores. This general pattern held true for all zoogeographic realms except the Neotropics, where nectarivores and frugivores predominated among altitudinal migrants. The prevalence of invertivore birds among altitudinal migrants is not unexpected because this is the most common foraging guild among birds worldwide. Overall, we found no prevalence of any specific foraging guild among altitudinal migrants across zoogeographic regions. The results of studies to date suggest that altitudinal migration by birds may be driven by a number of factors, including access to increased food resources for breeding or molting, weather conditions, and mating and nesting opportunities. However, to better understand the mechanisms underlying altitudinal migration, broadening the geographic scope of studies is paramount, with additional study of altitudinal migration especially needed in the megadiverse tropical regions of sub‐Saharan Africa, Southeast Asia, and South America.     

Seasonal energetics of northern phocid seals

The metabolic rate of harp (Pagophilus groenlandicus), harbor (Phoca vitulina), and ringed seals (Pusa hispida) was measured at various temperatures in air and water to estimate basal metabolic rates (BMRs) in these species. The basal rate and body composition of three harp seals were also measured throughout the year to examine the extent to which they vary seasonally. Marine mammalian carnivores generally have BMRs that are over three times the rates expected from body mass in mammals generally, both as a response to a cold-water distribution and to carnivorous food habits with the basal rates of terrestrial carnivores averaging about 1.8 times the mean of mammals. Phocid seals, however, have basal rates of metabolism that are 30% lower than other marine carnivores. Captive seals undergo profound changes in body mass and food consumption throughout the year, and after accounting for changes in body mass, the lowest rate of food intake occurs in summer. Contrary to earlier observations, harp seals also have lower basal rates during summer than during winter, but the variation in BMR, relative to mass expectations, was not associated with changes in the size of fat deposits. The summer reduction in energy expenditure and food consumption correlated with a reduction in BMR. That is, changes in BMR account for a significant portion of the seasonal variation in energy expenditure in the harp seal. Changes in body mass of harp seals throughout the year were due not only to changes in the size of body fat deposits, but also to changes in lean body mass. These results suggest that bioenergetics models used to predict prey consumption by seals should include time-variant energy requirements.     

Food selection by the guanaco (Lama guanicoe) along an altitudinal gradient in the Southern Andean Precordillera (Argentina)

Wild ungulates like the guanaco are exposed to important changes in climate and plant diversity along altitudinal gradients in the Andes Mountains, such as in the Southern Andean Precordillera where three phytogeographic provinces are present in altitudinal belts. The guanaco’s diet and food availability were seasonally analyzed using microhistological analysis and point-quadrat transects at six sampling sites, representative of the phytogeographic belts along the altitudinal gradient. Plant cover and diversity decreased with growing altitude. Richness of plant species was poorer at the summit than in the lower altitudes, whereas the proportion of species eaten by guanacos increased with altitude. The diet included 77 species. Grasses were preferred and shrubs were avoided all year round. The grass Poa spp. occupied more than 50 % of the diet at all altitudes. Grasses were the main dietary item even at low altitudes, where shrubs constituted the main food available. Decreasing generalism with descending phytogeographic belts agrees with the prediction for altitudinal gradients. The increase of diversity in the diet during the winter decline of plant cover at high and middle altitudes follows that expected from the optimal foraging theory. The winter decline of vegetation and the dietary shift from grazing to browsing proved to be stronger as altitude increases and the climate become more rigorous. Plant species richness, food scarcity, and climate severity are relevant variables to explain altitudinal and seasonal changes in the diet of adaptive ungulates in mountain environments, such as the guanaco in the Southern Andean Precordillera.     

Ontogeny of metabolism,thermoregulation and torpor in the house martin Delichon u. urbica (L.) and its ecological significance

Summary Special energetic adaptations are of great evolutionary significance for birds that encounter transient problems in finding food during the breeding season. House martins, as aerial insectivores, encounter such problems during spells of bad weather, when they must survive on body reserves. This species employs the following behavioural and physiological adaptations to save energy: Low basal metabolic rate (only 43% of the values predicted by allometric equations); low thermal conductance 51% (day) and 67% (night) of the predicted values; clustering behaviour; high tolerance of the young to periods of low food supply; and the ability to become torpid, found in adults and young from the age of 11 days on. House martins are the first passerine birds in which torpor has been found. These adaptations might have played a role in the great success of the house martin, one of the 10–15 most abundant bird species in Europe.Abbreviations BMR Basal metabolic rate, J/g·h - C Thermal conductance, J/g·h°C - M Energy metabolism, J/g·h - Ta Ambient temperature, °C - Tb Body temperature, °C - W Body mass, g A great part of these investigations were done in the laboratory of Prof. Dr. E. Kulzer, Physiologische Ökologie, Auf der Morgenstelle 28, D-7400 Tübingen, FRG     

Cellular Metabolic Rate Is Influenced by Life-History Traits in Tropical and Temperate Birds

In general, tropical birds have a “slow pace of life,” lower rates of whole-animal metabolism and higher survival rates, than temperate species. A fundamental challenge facing physiological ecologists is the understanding of how variation in life-history at the whole-organism level might be linked to cellular function. Because tropical birds have lower rates of whole-animal metabolism, we hypothesized that cells from tropical species would also have lower rates of cellular metabolism than cells from temperate species of similar body size and common phylogenetic history. We cultured primary dermal fibroblasts from 17 tropical and 17 temperate phylogenetically-paired species of birds in a common nutritive and thermal environment and then examined basal, uncoupled, and non-mitochondrial cellular O₂ consumption (OCR), proton leak, and anaerobic glycolysis (extracellular acidification rates [ECAR]), using an XF24 Seahorse Analyzer. We found that multiple measures of metabolism in cells from tropical birds were significantly lower than their temperate counterparts. Basal and uncoupled cellular metabolism were 29% and 35% lower in cells from tropical birds, respectively, a decrease closely aligned with differences in whole-animal metabolism between tropical and temperate birds. Proton leak was significantly lower in cells from tropical birds compared with cells from temperate birds. Our results offer compelling evidence that whole-animal metabolism is linked to cellular respiration as a function of an animal’s life-history evolution. These findings are consistent with the idea that natural selection has uniquely fashioned cells of long-lived tropical bird species to have lower rates of metabolism than cells from shorter-lived temperate species.     

Upward migration of vascular plants following a climate warming trend in the Alps

The aim of this study was to understand (1) whether warmer climatic conditions affected the vascular plant species composition, (2) the magnitude and rate of altitudinal changes in species distributions, and (3) whether an upward migration of alpine plants is connected to wind dispersal of diaspores. We compared historical records (1954–1958) with results from recent plant surveys (2003–2005) from alpine to nival ecosystems in the Rhaetian Alps, N-Italy. The presence of all vascular plant species and their maximum altitude were recorded along a continuous altitudinal transect of 730 m. An increase in species richness from 153 to 166 species was observed. Moreover, 52 species were recorded from altitudes 30–430 m higher than their 1950s limits, which corresponds to a median migration rate of 23.9 m/decade. In order to explain the observed migrations, the species wind-dispersal ability (diaspore weight and morphology) and the air temperature variation from 1926 to 2003 were considered. Species with more pronounced altitudinal shifts possess lighter diaspores. The highest increase in species richness was found between 2800 and 3100 m a.s.l.; this appears to be related to an estimated shift of the permafrost limit by +240 m during the last 50 years. The mean air temperature in the region rose by +1.6 °C in summer and by +1.1 °C in winter within this period. Climate warming is therefore considered as a primary cause of the observed upward migration of high mountain plants. Calculated altitudinal migration rates, however, varied remarkably among species. This would imply differential abilities of species to persist in an increasingly warmer climate. Species-specific conservation measures, including ex situ conservation, may therefore be required.