Altitudinal Temperature Gradients Research Articles

Higher temperature accelerates carbon cycling in a temperate montane forest without decreasing soil carbon stocks

Global warming is expected to accelerate the cycling of soil organic carbon (SOC) and the assimilation of new carbon, but the net effect of those counteracting accelerations and their ultimate effects on SOC are still uncertain. This hinders the prediction of long-term changes in biospheric carbon stocks and SOC-climate feedbacks. Here, we studied the long-term effect of temperature on carbon cycling across a 3.2 °C altitudinal temperature gradient in a temperate forest ecosystem in New Zealand. Across the gradient, soil respiration rates increased with increasing temperature from 9.0 to 10.4 tC ha−1 yr−1, but SOC stocks down to 85 cm depth also tended to increase, from 154 to 176 tC ha−1, albeit non-significantly (P = 0.06). This system was able to maintain higher soil respiration rates at higher temperatures without reducing SOC because the higher respiration rates were sustained by higher litterfall rates. Aboveground litterfall increased from 1.8 to 2.4 tC ha−1 yr−1 and estimated belowground C inputs increased from 7.2 to 8.0 tC ha−1 yr−1 along the temperature gradient. These higher fluxes were associated with significantly (P < 0.05) increased biomass at higher temperatures. As a direct measure of the effect of temperature on carbon cycling processes, we also calculated the turnover rate of forest litter which increased about 1.4-fold across the temperature gradient. This study demonstrates that higher temperatures along the thermal gradient increased plant carbon inputs through enhanced gross primary production, which counteracted SOC losses through temperature-enhanced soil respiration. These results suggest that temperature sensitivities of both plant carbon inputs and SOC losses must be considered for predicting SOC-climate feedbacks.

Complementarity of gymnosperms and angiosperms along an altitudinal temperature gradient

In seasonal tropical forests, evergreen–deciduous mixtures are more productive than monocultures because they intercept more light throughout the year, reflecting complementary resource use by functional groups possessing different traits. This suggests that temperate and boreal forests may also exhibit overyielding, due to the difference in phenology between gymnosperms and angiosperms. However, complementarity could also arise from differences in morphology between needle leaves and broad leaves, or facilitation by N‐fixing species. Alternatively, mixtures may be more productive simply because interspecific competition is less intense than intraspecific competition. We used forest inventory data to assess the complementarity of the main functional groups in Switzerland. We employed a trait‐based analysis of competition to determine whether: 1) trait differences reduce the intensity of competition between complementary functional groups, 2) complementarity is observed along a broad altitudinal temperature gradient. N‐fixing species facilitated the growth of non‐fixing species, such that 50/50 mixtures were 50% more productive than monocultures, though half the overyielding was due to the alleviation of intraspecific competition. In contrast, we found no evidence of complementarity between evergreen and deciduous species. For example, in stands where larch was mixed with other gymnosperms, there was no reduction in heterospecific competition among evergreen species, even though evergreens cast shade on one another throughout the year. In cold montane forests, broadleaf species reduced the suppression of needleleaf species, and vice versa. Thus, 50/50 mixtures of needleleaf and broadleaf species were 15% more productive than needleleaf monocultures. However, in warm lowland forests, broadleaf species exacerbated the suppression of needleleaf species, completely offsetting the positive effect that needleleaf species had on broadleaf species. In summary, we found no evidence of complementarity between evergreen and deciduous species, but needleleaf–broadleaf mixtures exhibited overyielding in cold montane forests, which is consistent with the stress gradient hypothesis, though the underlying mechanisms remain uncertain.

Caterpillar biomass depends on temperature and precipitation, but does not affect bird reproduction

Complex changes in phenological events appear as temperatures are increasing: In deciduous forests bud burst, hatching of herbivorous caterpillars, egg laying and nestling time of birds when feeding chicks on caterpillars, may differentially shift into early season and alter synchronization. If timing of bird reproduction has to match with short periods of food availability, phenological mismatch could negatively affect reproductive success. Using a unique empirical approach along an altitudinal temperature gradient, we firstly asked whether besides temperature, also precipitation and leaf phenology interplay and affect caterpillar biomass, since impacts of rainfall on caterpillars have been largely neglected so far. Secondly, we asked whether abundance of caterpillars and thereby body mass of great tit nestlings, which are mainly fed with caterpillars, vary along the altitudinal temperature gradient. We demonstrated that next to temperature also precipitation and leaf phenology affected caterpillar biomass. In our beech forest, even along altitudes, caterpillars were available throughout the great tit breeding season but in highly variable amounts. Our findings revealed that although timing of leaf phenology and great tit breeding season were delayed with decreasing temperature, caterpillars occurred synchronously and were not delayed according to altitude. However, altitude negatively affected caterpillar biomass, but body mass of fledglings at high altitude sites was not affected by lower amounts of caterpillar biomass. This might be partially outweighed by larger territory sizes in great tits.

Increase in the upper atmospheric temperature over tropospheric sources: Analysis of satellite measurements and numerical simulation

Variations in the temperature of the upper atmosphere caused by hurricanes are considered in this work on the basis of UARS satellite measurements. Analysis of the temperature variations shows that the temperature increases by 24–25 K in the mesopause over high-power tropospheric formations. Atmospheric gravity waves are considered a possible means of transferring disturbances from the Earth’s lower to the upper atmosphere. The maximal amplitude of atmospheric gravity waves was detected at altitudes of about 90 km during numerical simulation of propagation of the waves in a nonisothermal windless atmosphere with an accounting for the viscosity and thermal conductivity. A key factor of their attenuation and propagation is the altitudinal temperature gradient.

Do Great TitsParus majorNesting at High Altitudes Build Better Insulated Nests?

We investigated nest building behaviour of great tits Parus major in relation to altitudinal temperature gradients in Lower Austria. We hypothesised that colder environments at higher altitudes would select for a nest construction that increases thermal nest insulation. We collected data on nest architecture from 11 nests and on nest components from 33 nests along an altitudinal range (390 m) in 2010 to 2012, and experimentally measured specific insulation quality in 11 highly variable nests. Highly insulated nests included significantly more feathers than less well-insulated nests. Despite significantly lower temperatures at high altitude than at low altitude sites before clutch initiation, nest insulation quality did not increase with altitude. Moreover, we found that the date of clutch initiation was positively correlated with altitude. Birds may adjust to lower spring temperatures at high altitude by delaying clutch initiation, thus avoiding the need to invest in gathering highly insulating material.

Altitudinal variation in body size and age structure of the Sauter’s frog Rana sauteri in Taiwan

The tendency for organisms to be larger in cooler climates, the so-called Bergmann’s rule, has been widely observed in endotherms, but it is debatable in ectothermic anurans. Altitudinal variations in body size, age, and growth rate of the Sauter’s frog Rana sauteri were investigated with skeletochronology at six sites along the altitudinal gradients from 330 to 2,320 m in subtropical Taiwan. The aims of this study were to determine whether these life history traits vary with altitude and differ between males and females. The body size of male R. sauteri followed a converse Bergmann’s cline; the size decreased with increase in altitude, while no significant altitudinal change was found for females. The size dimorphism was female-biased and became greater at higher altitudes because of the altitudinal decrease in male size. Ages ranged between 1 and 5 years for males and 1 and 6 years for females. There was no significant difference in the age structures between the two sexes, but both sexes had higher average ages at higher-altitude sites. According to von Bertalanffy’s model, the growth coefficients decreased with increase in altitude for males but not for females. The annual growth rates were greater for all age classes of females than males, suggesting that growth rate was a major factor underlying body size patterns of both sexes. Growth rate was a major factor underlying body size patterns in both sexes of R. sauteri. It could be affected by not only altitudinal temperature gradient but also sex-biased life history traits and environmental factors.

Impact of Climate Change on the Distribution ofAedes albopictus(Diptera: Culicidae) in Northern Japan: Retrospective Analyses

The impact of climate change on the distribution of Aedes albopictus (Skuse) was analyzed in northern Japan, where chronological distribution records are incomplete. We analyzed local climate data using linear regression of the thermal suitability index (TSI) for the mosquito and mean annual temperature as functions of time. In northern Japan, thermal conditions since the early 20th century have become increasingly suitable for Ae. albopictus, more as a result of decreasing coldness in the overwintering season than increasing warmth in the reproductive season. Based on recent discovery records of Ae. albopictus in the northern border range, we determined thermal criteria for estimating when its persistent establishment became thermally possible. Retrospective analyses indicated that those criteria were reached in most coastal lowlands of northern Honshu before the accelerated temperature increase after the mid-1980s and the first records of this species after 1990; at some sites, temperature criteria were reached during or before the early 20th century. Expansion of the thermally suitable range after 1990 was supported only for inland areas and the northernmost Pacific coast. The estimated expansion rate was approximately 26 km per decade. Our analyses also demonstrated the importance of local climate heterogeneity (apart from north-south or altitudinal temperature gradients) in determining the expansion pattern.

Using genetic variation to infer associations with climate in the common frog,Rana temporaria

Recent and historical species' associations with climate can be inferred using molecular markers. This knowledge of population and species-level responses to climatic variables can then be used to predict the potential consequences of ongoing climate change. The aim of this study was to predict responses of Rana temporaria to environmental change in Scotland by inferring historical and contemporary patterns of gene flow in relation to current variation in local thermal conditions. We first inferred colonization patterns within Europe following the last glacial maximum by combining new and previously published mitochondrial DNA sequences. We found that sequences from our Scottish samples were identical to (92%), or clustered with, the common haplotype previously identified from Western Europe. This clade showed very low mitochondrial variation, which did not allow inference of historical colonization routes but did allow interpretation of patterns of current fine-scale population structure without consideration of confounding historical variation. Second, we assessed fine-scale microsatellite-based patterns of genetic variation in relation to current altitudinal temperature gradients. No population structure was found within altitudinal gradients (average FST=0.02), despite a mean annual temperature difference of 4.5 °C between low- and high-altitude sites. Levels of genetic diversity were considerable and did not vary between sites. The panmictic population structure observed, even along temperature gradients, is a potentially positive sign for R. temporaria persistence in Scotland in the face of a changing climate. This study demonstrates that within taxonomic groups, thought to be at high risk from environmental change, levels of vulnerability can vary, even within species.

Of mast and mean: differential‐temperature cue makes mast seeding insensitive to climate change

Mast-seeding plants often produce high seed crops the year after a warm spring or summer, but the warm-temperature model has inconsistent predictive ability. Here, we show for 26 long-term data sets from five plant families that the temperature difference between the two previous summers (ΔT) better predicts seed crops. This discovery explains how masting species tailor their flowering patterns to sites across altitudinal temperature gradients; predicts that masting will be unaffected by increasing mean temperatures under climate change; improves prediction of impacts on seed consumers; demonstrates that strongly masting species are hypersensitive to climate; explains the rarity of consecutive high-seed years without invoking resource constraints; and generates hypotheses about physiological mechanisms in plants and insect seed predators. For plants, ΔT has many attributes of an ideal cue. This temperature-difference model clarifies our understanding of mast seeding under environmental change, and could also be applied to other cues, such as rainfall.

Exogenous selection shapes germination behaviour and seedling traits of populations at different altitudes in a Senecio hybrid zone

The Senecio hybrid zone on Mt Etna, Sicily, is characterized by steep altitudinal clines in quantitative traits and genetic variation. Such clines are thought to be maintained by a combination of 'endogenous' selection arising from genetic incompatibilities and environment-dependent 'exogenous' selection leading to local adaptation. Here, the hypothesis was tested that local adaptation to the altitudinal temperature gradient contributes to maintaining divergence between the parental species, S. chrysanthemifolius and S. aethnensis. Intra- and inter-population crosses were performed between five populations from across the hybrid zone and the germination and early seedling growth of the progeny were assessed. Seedlings from higher-altitude populations germinated better under low temperatures (9-13 °C) than those from lower altitude populations. Seedlings from higher-altitude populations had lower survival rates under warm conditions (25/15 °C) than those from lower altitude populations, but also attained greater biomass. There was no altitudinal variation in growth or survival under cold conditions (15/5 °C). Population-level plasticity increased with altitude. Germination, growth and survival of natural hybrids and experimentally generated F(1)s generally exceeded the worse-performing parent. Limited evidence was found for endogenous selection against hybrids but relatively clear evidence was found for divergence in seed and seedling traits, which is probably adaptive. The combination of low-temperature germination and faster growth in warm conditions might enable high-altitude S. aethnensis to maximize its growth during a shorter growing season, while the slower growth of S. chrysanthemifolius may be an adaptation to drought stress at low altitudes. This study indicates that temperature gradients are likely to be an important environmental factor generating and maintaining adaptive divergence across the Senecio hybrid zone on Mt Etna.

Extinction debt of high-mountain plants under twenty-first-century climate change

Quantitative estimates of the range loss of mountain plants under climate change have so far mostly relied on static geographical projections of species’ habitat shifts 1‐3 . Here, we use a hybrid model 4 that combines such projections with simulations of demography and seed dispersal to forecast the climate-driven spatio-temporal dynamics of 150 highmountain plant species across the European Alps. This model predicts average range size reductions of 44‐50% by the end of the twenty-first century, which is similar to projections from the most ‘optimistic’ static model (49%). However, the hybrid model also indicates that population dynamics will lag behind climatic trends and that an average of 40% of the range still occupied at the end of the twenty-first century will have become climatically unsuitable for the respective species, creating an extinction debt 5,6 . Alarmingly, species endemic to the Alps seem to face the highest range losses. These results caution against optimistic conclusions from moderate range size reductions observed during the twenty-first century as they are likely to belie more severe longer-term effects of climate warming on mountain plants. Many plant and animal species have already been shifting their ranges in response to the past century’s climatic trends 79 . In mountains, owing to the altitudinal temperature gradient, species should primarily move upslope under warming, as has indeed been frequently documented during the recent decades 10,11 as well as in the palaeorecord 12,13 . As mountains usually have conical shapes, upslope movement inevitably results in range loss and may even lead to ‘mountain-top extinctions’ 14 in extreme cases. However, previous predictions of the magnitude of such range and biodiversity losses during the twenty-first century have been criticized 4,15 for relying on static ‘niche-based’ modelling approaches 16 , which disregard several processes crucial to range

Digesta retention time in the Galápagos tortoise (Chelonoidis nigra)

The retention time of food in the digestive tract of animals has important implications for digestive physiology. Retention time impacts digestive efficiency and among herbivores affects plant–animal interactions including herbivory and seed dispersal. Poorly studied yet iconic Galápagos tortoises are large-bodied generalist herbivores and ecosystem engineers which migrate seasonally. Potentially variable digesta retention times due to strong seasonal and altitudinal temperature gradients may influence tortoise seed dispersal abilities and rates of herbivory. We fed captive adult tortoises living in semi-natural conditions on Galápagos with inert particles and seeds from locally available fruits to determine whether seed size and ambient temperature influenced retention time. Median retention time varied from 6 to 28days, with a mode of 12days. Seed size had no effect on any of our measures of retention time, but ambient temperature was inversely correlated with retention times. Long retention time facilitates long distance seed dispersal by Galápagos tortoises, which may improve effectiveness. The effect of temperature, which may double from hot lowlands to cold highlands through the seasonal cycle, on tortoise digesta retention time will strongly influence seed dispersal efficiency and may influence patterns of food selection and migration in this species.

Lateglacial and early Holocene vegetation and climate gradients in the Nordfjord-Ålesund area, western Norway

Birks, H. H. & van Dinter, M. 2010: Lateglacial and early Holocene vegetation and climate gradients in the Nordfjord–Alesund area, western Norway. Boreas, Vol. 39, pp. 783–798. 10.1111/j.1502-3885.2010.00161.x. ISSN 0300-9483. Modern climate in western Norway shows a strong west–east gradient in oceanicity–continentality (coast to inner fjord) and altitudinal temperature gradients that control the regional and altitudinal zonation of vegetation. To discover if similar gradients existed during the Lateglacial and early Holocene, plant-macrofossil analyses were made from five lacustrine sediment sequences in the Nordfjord–Alesund region selected to sample the present climatic gradients. The macrofossil assemblages could be interpreted as analogues of the present vegetation, thus allowing reconstruction of past vegetation and climates. When the five sites were compared, climatic gradients could be detected. During the Lateglacial interstadial, mid-alpine assemblages with Salix herbacea and S. polaris occurred at the lowland coast and upland inland sites, whereas the inland lowland site had low-alpine dwarf-shrub heath dominated by Betula nana, demonstrating a strong west–east gradient in temperature and precipitation and an altitudinal gradient inland. During the Younger Dryas stadial, assemblages at the lowland coast and upland inland sites resembled high-alpine vegetation, whereas the inland lowland site was warmer with mid-alpine vegetation, demonstrating west–east and altitudinal temperature gradients. Gradients became less pronounced in the Holocene. The early abundance of Betula nana in the inner fjord sites but its rarity at the coast is striking and reflects the oceanicity gradient. All sites became forested with Betula pubescens a few centuries into the Holocene. This forest was probably close to tree line at 370 m a.s.l. at the coast. Inland, there was no detectable altitudinal gradient, with the tree line well above 400 m a.s.l. reflecting the present pattern of tree-line elevation.

Nonlinear response of dissolved organic carbon concentrations in boreal lakes to increasing temperatures

Recent increases in concentrations of dissolved organic carbon (DOC) in lakes and rivers over large regions have been related to both changes in the climate and in atmospheric deposition chemistry. Using a data set of 1041 boreal lakes along a 13° latitudinal gradient, sampled in 1995, 2000, and 2005, and an additional data set of 90 lakes along a 1000‐m altitudinal gradient at 68°N, we show that DOC concentrations increase in a nonlinear way along a latitudinal and altitudinal temperature gradient. The nonlinear relation of DOC to increasing temperatures was consistent over space and time. Out of 14 meteorological, catchment, morphometric, and atmospheric deposition variables tested, the variable best explaining this kind of nonlinear pattern was the number of days when air temperatures exceeded 0°C, i.e., the duration of the main growing and runoff season (DT&gt;0). Using DT&gt;0 as an input variable, we were able to predict the nonlinear temperature response of DOC concentrations, both spatially (R2 = 0.90, p &lt; 0.0001) and temporally (R2 = 0.90, p &lt; 0.0001). DT&gt;0 has an advantage over other variables because it includes the time factor, which is decisive for the duration that biogeochemical processes can take place. We suggest that DOC concentrations in lakes are influenced by climate change and that present temperature increases over Sweden result in an accelerated DOC increase toward warmer geographical regions.

Directional Evolution of the Slope of the Metabolic Rate–Temperature Relationship Is Correlated with Climate

Abstract The evolution of metabolic rate-temperature (MR-T) reaction norms is of fundamental importance to physiological ecology. Metabolic cold adaptation (MCA) predicts that populations or species from cooler environments will have either a higher metabolic rate at a common temperature or steeper MR-T relationships, indicating greater sensitivity of respiratory metabolism to temperature. Support for MCA has been found in some insect species by comparing species or populations differing in latitude. However, the generality of these findings are contentious, with most studies either unable to account for phenotypic plasticity or the evolutionary relatedness of species or populations. Hence, the importance of MCA is vigorously debated from both evolutionary and ecological perspectives. Furthermore, few species, particularly from tropical environments, have been shown to differ in MR-T sensitivity along altitudinal temperature gradients. Here, using four populations of tsetse flies (Glossina pallidipes, Diptera: Glossinidae) from thermally distinct geographic regions, we test the hypothesis that there is evolved variation in MR-T relationships to cold climates. We found that a high-altitude equatorial population from a cool habitat has a steeper MR-T reaction norm. By contrast, other populations from warmer environments in East Africa do not differ with respect to their MR-T reaction norms. Squared-change parsimony analyses, based on the combined mitochondrial 16S rDNA ribosomal subunit and cytochrome c oxidase subunit I (COI), support the hypothesis of adaptive differentiation of MR-T reaction norms in the cool-climate population. Seasonal adjustments or laboratory-temperature-induced phenotypic plasticity changed the intercept of the reaction norm rather than the slope, and thus the observed intraspecific variation in slopes of MR-T reaction norms could not be accounted for by phenotypic plasticity. These results therefore suggest evolutionary adaptation of MR-T reaction norms to cool climates (<22 degrees C) in tsetse and provide novel support for MCA within an insect species.

Computation of the space and time evolution of equilibrium-line altitudes on Andean glaciers (10°N–55°S)

A previous study of Fox [Fox, A.N. 1993. Snowline altitude and climate at present and during the Last Pleistocene Glacial Maximum in the Central Andes (5°–28°S). Ph.D. Thesis. Cornell University.] showed that for a fixed 0 °C isotherm altitude, the equilibrium-line altitude (ELA) of the Peruvian and Bolivian glaciers from 5 to 20°S can be expressed based on a log–normal expression of local mid-annual rainfall amount ( P). In order to extrapolate the function to the whole Andes (10°N to 55°S) a local 0 °C isotherm altitude is introduced. Two applications of this generalised function are presented. One concerns the space evolution of mean inter-annual ELA for three decades (1961–1990) over the whole South American continent. A high-resolution data set (grid data: 10′ for latitude/longitude) of mean monthly air surface temperature and precipitation is used. Mean annual values over the 1961–1990 period were calculated. On each grid element, the mean annual 0 °C isotherm altitude is determined from an altitudinal temperature gradient and mean annual temperature ( T) at ground level. The 0 °C isotherm altitude is then associated with the annual precipitation amount to compute the ELA. Using computed ELA and the digital terrain elevation model GTOPO30, we determine the extent of the glacierised area in Andean regions under modern climatic conditions. The other application concerns the ELA time evolution on Zongo Glacier (Bolivia), where inter-annual ELA variations are computed from 1995 to 1999. For both applications, the computed values of ELA are in good agreement with those derived from glacier mass balance measurements.

Altitudinal Differences in Organic Matter Mass Loss and Fungal Biomass in a Subalpine Coniferous Forest, Mt. Fuji, Japan

In order to study the long-term effect of climatic conditions on organic matter decomposition, the difference in the mass loss rate of organic substrates was examined along the altitudinal temperature gradient (1500–2400 m) of Mt. Fuji, Japan. Two standard substrates, cellulose filter paper and wood chips of beech (Fagus crenata), were placed in the L and FH layers at each altitude. The mass loss rates tended to be smaller with increasing altitude and were significantly correlated with the annual mean air temperature in a log-linear fashion. The temperature dependence of the mass loss rates was much larger than that of microbial respiration. The initial increase in ergosterol content (an indicator of fungal biomass) of organic substrates was more rapid at low altitudes than at higher altitudes. The effects of fungal biomass and freezing temperatures on the temperature dependence of decomposition were examined using a simple simulation model. The results suggest that the effects of temperature increase on decomposition could be much larger than those estimated from the temperature-microbial respiration curves.

A fine‐scaled predictive model for changes in species distribution patterns of high mountain plants induced by climate warming

SummaryInduced by global warming, mountain plant species are migrating upwards. Species inhabiting the nival zone of today are threatened by competitors which move from the alpine zone towards the summits. The manner in which species move depends on their abilities to cope with microtopographical situations. We present a spatially explicit predictive model which draws scenarios of future species distribution patterns at a typical high mountain of the European Alps. The altitudinal temperature gradient is examined. Based on the lapse rate and on definitions of topographical niches of species, a +1 °C‐ and a +2 °C‐warming scenario are modelled using a fine‐scaled digital elevation model. Nival species will lose area and become restricted to specific topographical situations. Alpine and subnival grassland species are predicted to expand their area, mainly along stable surface situations. Whether the migration will take place as a filling or a moving process is specific to the particular species. Overall, biodiversity is apparently not threatened on the decadal scale. In special cases, however, genetic losses are likely both on a local and on a regional scale.

Climatic Shift of the Equilibrium Line: Kuhn's Concept Applied to the Greenland Ice Cap

The shift of the equilibrium line in response to climatic disturbances is calculated for the International Glaciological Greenland Expedition (EGIG) profile of the Greenland Ice Cap. Data of heat balance studies carried out during EGIG 1959 and EGIG 1967 are used according to Kuhn’s algorithm. The following parameters are applied; the heat transfer coefficient of the sensible heat flux; the effective emissivity of the atmosphere for long wave downward radiation; a factor relating net radiation to cloudiness; the duration of the ablation season; the altitudinal temperature gradient; the altitudinal accumulation gradient.The altitudinal shift of the equilibrium line results for the EGIG profile + 77m/°C and -4 m per 1/10 cloudiness with 35d of ablation. Changes in heat balance by formation of superimposed ice are taken into account.

Climatic Shift of the Equilibrium Line: Kuhn's Concept Applied to the Greenland Ice Cap

The shift of the equilibrium line in response to climatic disturbances is calculated for the International Glaciological Greenland Expedition (EGIG) profile of the Greenland Ice Cap. Data of heat balance studies carried out during EGIG 1959 and EGIG 1967 are used according to Kuhn’s algorithm. The following parameters are applied; the heat transfer coefficient of the sensible heat flux; the effective emissivity of the atmosphere for long wave downward radiation; a factor relating net radiation to cloudiness; the duration of the ablation season; the altitudinal temperature gradient; the altitudinal accumulation gradient. The altitudinal shift of the equilibrium line results for the EGIG profile + 77m/°C and -4 m per 1/10 cloudiness with 35d of ablation. Changes in heat balance by formation of superimposed ice are taken into account.