Influence of altitude on local adaptation in upland tree species from central Argentina

• Steep climatic gradients boost morphological and physiological adjustments in plants, with consequences on performance. The three principal woody species of the Sierras Grandes Mountains of central Argentina have marked differences in sapling performance along their altitudinal distribution. We hypothesize that the steep gradient of climatic conditions across the species’ altitudinal distribution promotes trait differences between populations of different altitudes that are inherited by the following generation.
• Seeds from different altitudes were exposed to three temperature regimes to assess differential germination responses. Saplings were then transplanted to a greenhouse to assess possible variations in attributes and performance after 18 months.
• The three species showed differences in germination responses to temperature among altitudes and/or in sapling attributes and performance. In Maytenus boaria and Escallonia cordobensis, germination success was higher under high temperatures for the highest‐altitude, whereas lower temperatures boosted germination of the lowest altitudes. Polylepis australis showed no differences in germination among temperature treatments. In the greenhouse, saplings of the three species from intermediate altitudes showed high performance, whereas the upper and lower populations seemed to be adjusted to tolerating more stressful conditions (i.e., lower temperatures at the upper end and water stress at the lower end), showing lower performance toward both altitudinal limits.
• These patterns agree with those described for saplings growing under field conditions, suggesting adjustments in response to environmental changes undergone by populations along the altitudinal range. The marked adjustments of populations to the local environment suggest a potentially high impact of climatic change on species distribution.

     

Phenological responses of upland rice grown along an altitudinal gradient

High altitude upland rice (Oryza sativa L.) production systems are expected to benefit from climate change induced increase in temperatures. The potential yield of rice genotypes is governed by the thermal environment experienced during crop development phases when yield components are determined. Thus, knowledge on genotypic variability in phenotypic responses to variable temperature is required for assessing the adaptability of rice production to changing climate. Although, several crop models are available for this task, genotypic thermal constants used to simulate crop phenology vary strongly among the models and are under debate. Therefore, we conducted field trials with ten contrasting upland rice (O. sativa L.) genotypes on three locations along an altitudinal gradient with five monthly staggered sowing dates for two years in Madagascar with the aim to study phenological responses at different temperature regimes. We found that, crop duration is equally influenced by genotype selection, sowing date and year in the high altitude. In contrast, in mid altitudes genotype has no effect on crop duration. At low altitudes crop duration is more affected by sowing date. Grain yield is strongly affected by low temperatures at high altitudes and severly influenced by frequent tropical cyclones at low altitudes. In high altitude, genotype explained 68% of variation in spikelet sterility, whereas in mid and low altitudes environment explained more than 70% of the variation. The phenological responses determining crop duration and yield, the basic genotypic thermal constants, and the analyses of genotypic thermal responses with regard to spikelet sterility reported here, provide valuable information for the improvement of rice phenological models urgently needed to develop new genotypes and better adapted cropping calendars.     

Small-scale altitudinal effects on reproduction in bank voles

Elevation gradients and corresponding changes in environment have the potential to affect seasonal reproduction and mortality and hence the seasonal age structure of animal populations. This has been frequently shown in mammals and birds on larger geographic scales, but evidence for small-scale gradient effects is scanty. We studied such small scale altitudinal effects on the proportions of juveniles in a population of bank voles (Myodes glareolus) in a forested habitat along an elevation gradient of 355 m. This was done by regular live trapping in different altitudes over a two-year period. We also measured altitudinal differences in ambient temperatures and vegetational parameters in order to shed light on the underlying mechanisms of such purported differences in age structure. As could be expected for a seasonal breeder, the proportion of juveniles showed an initial increase in spring, an optimum in summer and a subsequent regression in early autumn, reflecting the seasonal pattern of reproduction. In addition, there was evidence for altitudinal effects on the occurrence of juveniles, which was decreased in higher altitudes, assumingly driven by altitudinal differences in onset and/or intensity of annual breeding activity. This altitudinal gradient in juvenile occurrence was partly explained by corresponding differences in ambient temperatures. Furthermore, there was good support that the higher abundance of herbs in lower altitudes played an important role in explaining the altitudinal differences in the proportion of juveniles. In conclusion, our study strongly supports the existence of altitudinal small-scale gradient effects on reproductive parameters in a small rodent, most probably due to the action of altitude-related factors such as microclimate and vegetational parameters.     

Plasticity to simulated shade is associated with altitude in structured populations of Arabidopsis thaliana

Plants compete for photosynthesis light and induce a shade avoidance syndrome (SAS) that confers an important advantage in asymmetric competition for light at high canopy densities. Shade plasticity was studied in a greenhouse experiment cultivating Arabidopsis thaliana plants from 15 populations spread across an altitudinal gradient in the northeast area of Spain that contain a high genetic variation into a reduced geographical range. Plants were exposed to sunlight or simulated shade to identify the range of shade plasticity. Fourteen vegetative, flowering and reproductive traits were measured throughout the life cycle. Shade plasticity in flowering time and dry mass was significantly associated with the altitude of population origin. Plants from coastal populations showed higher shade plasticity indexes than those from mountains. The altitudinal variation in flowering leaf plasticity adjusted negatively with average and minimum temperatures, whereas dry mass plasticity was better explained by negative regressions with the average, maximum and minimum temperatures, and by a positive regression with average precipitation of the population origin. The lack of an altitudinal gradient for the widest number of traits suggests that shade light could be a driver explaining the distribution pattern of individuals in smaller geographical scales than those explored here.     

Altitudinal variation in life cycle syndromes of California populations of the grasshopper,Melanoplus sanguinipes (F.)

Summary Life cycles of California populations of the grasshopper, Melanoplus sanguinipes, varied along an altitudinal gradient. Temperature records indicate a longer season at low altitude on the coast, based on computation of degree days available for development, even though summer air temperatures are cooler than at high altitude; this is a result of warm soil temperatures. At high and low altitudes there was a high proportion of diapause eggs oviposited, while intermediate proportions of diapause eggs occurred at mid altitudes. The low altitude, and especially sea level, populations diapaused at all stages of embryonic development, while at high altitudes most diapause occurred in the late stages just before hatch. Diapause was more intense at high altitudes. One result of diapause differences was delayed hatching in the sea level population. Nymphal development and development of adults to age at first reproduction were both accelerated at high altitude relative to sea level. At lower temperatures (27° C) there was a tendency for short days to accelerate development of sea level nymphs, but not high altitude nymphs. In both sea level and high altitude grasshoppers, short days accelerated maturation of adults to onset of oviposition at warm temperature (33° C); there was little reproduction at 27° C. Population differences for all traits studied appear to be largely genetic with some maternal effects possible. We interpret diapause variation at low and mid altitudes to be responses to environmental uncertainty and variations in development rates to be adaptations to prevailing season lengths.     

Altitudinal variation of the thermal biology and running performance in the lizard Podarcis tiliguerta

Summary We studied, in the field and laboratory, aspects of the thermal biology in two populations of the lizard Podarcis tiliguerta along a 1450 m altitudinal gradient. Body temperatures (T_b) at high altitudes average lower, are more variable, but are more elevated above environmental temperatures than at sea level. Lizards partially reduced the impact of altitudinal changes in thermal loads through presumable subtle behavioural adjustments. A comparison of the thermal preferences in the laboratory, the maximal operative temperatures predicted from a biophysical model, and the activity T_b's at both sites, indicates that the main response to changing environmental conditions is an active shift in thermoregulatory set points. Integration of field T_b's and laboratory data on temperature specific sprint speeds, predicts that the mountainous lizards experience reduced running abilities that are especially acute in the early morning. Despite this impairment of running performance, the thermal sensitivity of running speed has not evolved to match the T_b's experienced by both populations. This result supports the view that the thermal physiology of this lizard is evolutionarily conservative, but the lack of information on the relation between running performance and fitness components impedes rejection of alternative hypotheses.     

VARIATION IN LEAF DISSECTION AND LEAF ENERGY BUDGETS AMONG POPULATIONS OF ACHILLEA FROM AN ALTITUDINAL GRADIENT

The degree of leaf dissection differs dramatically among populations of the Achillea millefolium complex along an altitudinal gradient in the Sierra Nevada. The purpose of this study was to determine whether there was a genetic basis to differences in leaf shape among populations, and also to determine the importance of genetic variability within populations, plastic responses of leaf shape to the growth environment, and genetic differences among populations in plastic response to the environment. A second major goal of the research was to investigate the effects of differences in leaf dissection on the capacity for leaf temperatures to become uncoupled from air temperatures. Greenhouse experiments using clonal replicates of plants collected at different altitudes revealed that there were genetic differences among populations in the degree of dissection, and that other sources of phenotypic variation, such as plasticity, were also significant. Leaves from the highest altitude population had the most compact shape under all conditions, while those from lower altitude populations were always more open and highly dissected. In both simulations and actual measurements the dissected leaves of low altitude plants remained close to air temperatures, while the compact leaves of high altitude plants had the capacity to warm up substantially above air temperatures.     

Pattern of ant diversity in Korea: An empirical test of Rapoport's altitudinal rule

Two diversity patterns (hump-shaped and monotonic decrease) frequently occur along altitude or latitude gradients. We examined whether patterns of ant species richness along altitudes in South Korea can be described by these patterns and whether ranges of ant species follow Rapoport's altitudinal rule. Ants on 12 high mountains (> 1100 m) throughout South Korea (from 33° N to 38° N) were surveyed using pitfall traps at intervals of 200–300 m altitude. The temperatures at the sampling sites were determined from digital climate maps. Ant species richness decreased monotonically along the altitudinal gradient and increased along the temperature gradient. However, species richness of cold-adapted species (highland species) showed a hump-shaped pattern along altitude and temperature gradients. The altitude and temperature ranges of ant species followed Rapoport's rule. Sampling site temperature ranges were significantly correlated with coldness. Therefore, Rapoport's rule can be explained by high cold-tolerance of species inhabiting high altitudes or latitudes.     

Morphological and physiological responses to altitude in deer mice Peromyscus maniculatus.

Individuals within a species, living across a wide range of habitats, often display a great deal of phenotypic plasticity for organ mass and function. We investigated the extent to which changes in organ mass are variable, corresponding to environmental demand, across an altitudinal gradient. Are there changes in the mass of oxygen delivery organs (heart and lungs) and other central processing organs (gut, liver, kidney) associated with an increased sustainable metabolic rate that results from decreased ambient temperatures and decreased oxygen availability along an altitudinal gradient? We measured food intake, resting metabolic rate (RMR), and organ mass in captive deer mice (Peromyscus maniculatus bairdii) at three sites from 1,200 to 3,800 m above sea level to determine whether energy demand was correlated with organ mass. We found that food intake, gut mass, and cardiopulmonary organ mass increased in mice living at high altitudes. RMR was not correlated with organ mass differences along the altitudinal gradient. While the conditions in this study were by no means extreme, these results show that mice living at high altitudes have higher levels of energy demand and possess larger cardiopulmonary and digestive organs than mice living at lower altitudes.     

Altitudinal variation in egg retention and rates of embryonic development in oviparous Zootoca vivipara fits predictions from the cold-climate model on the evolution of viviparity

The evolution of reptilian viviparity is favoured, according to the cold‐climate hypothesis, at high latitudes or altitudes, where egg retention would entail thermal benefits for embryogenesis because of maternal thermoregulation. According to this hypothesis, and considering that viviparity would have evolved through a gradual increase in the extent of intrauterine egg retention, highland oviparous populations are expected to exhibit more advanced embryo development at oviposition than lowland populations. We tested for possible differences in the level of egg retention, embryo development time and thermal biology of oviparous Zootoca vivipara near the extreme altitudinal limits of the species distribution in the north of Spain (mean altitude for lowland populations, 235 m asl.; for highland populations, 1895 m asl.). Altitude influenced neither temperature of active lizards in the field nor temperature selected by lizards in a laboratory thermal gradient, and pregnant females selected lower temperatures in the thermal gradient than did males and nonpregnant females across altitudinal levels. Eggs from highland populations contained embryos more developed at the time of oviposition (Dufaure and Hubert's stages 33–35) than eggs of highland populations (stages 30–34) and partly because of this difference incubation time was shorter for highland embryos. When analysed for clutches from both altitudinal extremes at the same embryonic stage at oviposition (stage 33), again incubation time was shorter for highland populations, indicating genuine countergradient variation in developmental rate. Our results indicate that temperature is an environmental factor affecting the geographical distribution of different levels of egg retention in Z. vivipara, as predicted by the cold‐climate hypothesis on the evolution of viviparity.     

Searching for Altitudinal Zonation: Species Distribution and Vegetation Composition in the Superpáramo of Volcán Iliniza, Ecuador

Altitudinal variation of the zonal superpáramo vegetation was studied between 4300 and 4630 m to test a possible occurrence of a fine altitudinal zonation within the superpáramo belt. A rectangular grid of 1 m² sample plots was established; 25 replicate plots separated by a 3 m space were located along a 100 m long transect parallel to the contours, and there were 17 such transects separated by 20 m of altitude. Species were scored using a 7-grade cover scale and basic environmental data were gathered for each sample. Major changes occur over a short altitudinal range, at around 4400 m, which corresponds to a transition between the lower and upper superpáramo. Species richness sharply declines but species turnover (per altitude) increases along the altitudinal gradient. The correlation between richness and bare ground or rock cover is negative, but the correlation to rocks becomes positive above 4500 m. Species from lower altitudes tend to have narrower altitudinal range, although a large number of species appear to be indifferent to altitude. Direct ordination analyses indicate that high-altitude species show stronger correlation to environmental variables, especially rock, than species from lower altitudes. TWINSPAN cluster analysis delimited 15 groups of samples. There is a change in the clustering pattern along the altitudinal gradient from a horizontal (i.e., within altitude) to vertical (i.e., across altitude) arrangement of the cluster groups, although this pattern is partly obscured at the highest altitudes due to a large number of empty samples. MANOVA tests for samples from adjacent altitudinal levels indicate two distinct altitudinal breaks at lower altitudes, corresponding to the Loricaria-belt in lower superpáramo and the transition between lower and upper superpáramo, while no indication of a zonation was found in upper superpáramo.     

Altitudinal zonation of human-disturbed vegetation on Mt. Tianmu,eastern China

Much of the primary vegetation at low altitudes has been greatly altered or destroyed by a long history of human activities. This is particularly true in eastern China, where low-altitude areas are now dominated by secondary forests or plantations. Altitudinal vegetation zonation of this region is often based on these secondary forests, resulting in seral vegetation with an obscure zonal sequence. Here, we deduced the potential climax vegetation according to the regeneration patterns of the dominant species of the secondary forests at low altitudes (below 1,000 m a.s.l.) on Mt. Tianmu (1,506 m a.s.l., 30°18′30″–30°21′37″N, 119°24′11″–119°27′11″E). Based on the potential climax vegetation combined with the floristic composition and community structure, three vegetation zones were identified, viz: (1) evergreen broad-leaved forest zone (400–950 m a.s.l.); (2) evergreen and deciduous broad-leaved mixed forest zone (950–1,100 m a.s.l.); (3) deciduous broad-leaved forest zone (1,100–1,506 m a.s.l.). The altitudinal vegetation zones identified in this study correspond with the thermal conditions on Mt. Tianmu. The distribution of vegetation on Mt. Tianmu was limited by lower temperatures in winter, and the altitudinal thermal vegetation zones on this mountain were more similar to the thermal vegetation of Japan than to that of China. The vertical distributions and roles of conifers were different between the eastern and the western regions along 30°N latitude in humid East Asia. Cryptomeria fortunei formed the emergent layer, towering above the broad-leaved canopy at middle altitudes as C. japonica on Yakushima, but disappeared at high altitudes with hydrothermal limitation on Mt. Tianmu.     

Effect of size and altitude on soil organic carbon stock in homegarden agroforestry system in Central Himalaya,India

Homegarden agroforestry system has potential to sequester carbon while simultaneously producing agriculture yield and tree products. This study analyzed relationship between homegarden size and soil organic carbon stock (SCS) along an altitudinal gradient in Central Himalaya, India. Homegardens were selected at four altitudes (very low, low, mid and high) and categorized into three size classes viz. small, medium and large. Soil samples from surface (0–15 cm) and sub surface soils (15–30 cm) were collected seasonally and analyzed. Results indicated that SCS ranged between 22.56 and 81.51 t C ha⁻¹ and increased with increasing altitude for small homegardens. For medium and large homegardens, it increased up to mid altitude and then decreased in high altitude homegardens. At each altitude, maximum SCS (38.56–66.16 t C ha⁻¹) was recorded for medium sized homegardens except at high altitude where it was maximum (66.01 t C ha⁻¹) for small homegardens. Seasonal variation in SCS showed inconsistent pattern along the altitudinal gradient. The differences in SCS due to size and altitude recorded in the present study are possibly caused by distinct quality of plant material and management practices.     

Plant species and growth form richness along altitudinal gradients in the southwest Ethiopian highlands

Questions: Do growth forms and vascular plant richness follow similar patterns along an altitudinal gradient? What are the driving mechanisms that structure richness patterns at the landscape scale? Location: Southwest Ethiopian highlands. Methods: Floristic and environmental data were collected from 74 plots, each covering 400 m². The plots were distributed along altitudinal gradients. Boosted regression trees were used to derive the patterns of richness distribution along altitudinal gradients. Results: Total vascular plant richness did not show any strong response to altitude. Contrasting patterns of richness were observed for several growth forms. Woody, graminoid and climber species richness showed a unimodal structure. However, each of these morphological groups had a peak of richness at different altitudes: graminoid species attained maximum importance at a lower elevations, followed by climbers and finally woody species at higher elevations. Fern species richness increased monotonically towards higher altitudes, but herbaceous richness had a dented structure at mid‐altitudes. Soil sand fraction, silt, slope and organic matter were found to contribute a considerable amount of the predicted variance of richness for total vascular plants and growth forms. Main Conclusions: Hump‐shaped species richness patterns were observed for several growth forms. A mid‐altitudinal richness peak was the result of a combination of climate‐related water–energy dynamics, species–area relationships and local environmental factors, which have direct effects on plant physiological performance. However, altitude represents the composite gradient of several environmental variables that were interrelated. Thus, considering multiple gradients would provide a better picture of richness and the potential mechanisms responsible for the distribution of biodiversity in high‐mountain regions of the tropics.     

Assessing the origin of Neotropical mountain dung beetle assemblages (Scarabaeidae: Scarabaeinae): the comparative influence of vertical and horizontal colonization

Aims The fauna of mountains and their surrounding regions are likely to be influenced principally by two biological processes: horizontal colonization along similar altitudinal levels by elements originating from lineages inhabiting higher latitudes; and vertical colonization by lineages from the same latitude, but at lower altitudes. We examine whether the expected patterns derived from the latter process can be observed in mountain dung beetle assemblages. Specifically, we study the variation in species composition and richness with altitude in five regions spanning elevation gradients, analysing whether the altitudinal rates of change in the number of species and genera differ, and whether beta‐diversity scores for adjacent sites in each altitudinal gradient are different for species and genera. Location Eastern Cordillera of the Colombian Andes. Methods Field work was carried out in 1997–99 at 27 sites in five regions with elevation gradients, with 10–32 pitfall traps placed in each site. For each altitudinal level the numbers of species and genera were analysed with respect to altitude, and the slope of the linear regression between these variables was calculated. The slope of the curve of the altitude against the cumulative number of species and genera was also calculated for each altitudinal gradient to describe the compositional change between adjacent sites (beta diversity). Species and generic slopes were compared using analysis of covariance. The turnover of species along each altitudinal gradient was measured using presence/absence data and Cody's beta‐diversity index between adjacent pairs of sites. A cluster analysis was used to detect faunistically homogeneous groups of localities. Results Species richness always decreased with altitude, although the slopes did not differ significantly from zero. The number of genera also decreased with increasing altitude, but generally at a significantly slower rate than for species. Variation in the species beta‐diversity scores between altitudinal levels did not follow a homogeneous pattern in the different regions. Two main altitudinal groups of sites with a boundary c. 1500–1750 m a.s.l. can be detected with respect to faunistic similarity. Low‐ and mid‐altitude sites are inhabited by all of the genera (19) and 80% of all species collected. Eight genera and 61 species (c. 60% of the total) are unable to inhabit high‐altitude sites, and only 20 species appear to be exclusive to these high‐altitude environments (> 2000 m a.s.l.). Main conclusions The dominant processes explaining dung beetle composition in the high north‐eastern Andean mountains are probably those of vertical colonization. The limited role of horizontal colonization processes, or colonization from northern or southern lineages, could be a consequence of the isolation and recent geological origin of these mountains.     

Butterfly diversity and historical land cover change along an altitudinal gradient

Land cover and climate change are both major threats for biodiversity. In mountain ecosystems species have to adapt to fragmented habitats and harsh environmental conditions but so far, altitudinal effects in combination with land cover change have been rarely studied. The objective of this study was to determine the effects of altitude and historical land cover change on butterfly diversity. We studied species richness patterns of butterflies occuring in wetlands and other open habitats along an altitudinal gradient in a low mountain region (340–750 m a.s.l., Bavaria, Germany) with drastic loss of open habitats within the last 40–60 years. We recorded in 27 sites a total of 4,523 individuals of 49 butterfly species and five species of burnet moths. Species richness peaked at mid elevation and increased with patch size. Land cover change was most pronounced at high altitudes, but neither current open habitats, nor the historical loss of open habitats affected the species richness of butterflies. Neither open land specialized butterflies nor generalist and forest species were significantly affected by the loss of open habitats. However, increasing forest area in high altitudes reduces possible refuge open habitats for butterflies at their thermal distribution limits. This could lead to extinction of such butterfly species when temperatures further rise due to global warming.     

Genetic differences and phenotypic plasticity in body size between high- and low-altitude populations of the ground beetle Carabus tosanus

The body size of a univoltine carabid beetle Carabus tosanus on Shikoku Island, Japan, was clearly smaller in higher‐altitude populations (subspecies), which possibly represents incipient speciation. To explore the determinants of altitudinal differences in body size in this species, we studied the degree of phenotypic plasticity by conducting rearing experiments at two constant temperatures and examined genetic differences through interpopulation crosses. At 15 °C, C. tosanus had a longer developmental period and a shorter adult body than at 20 °C. Nevertheless, variation in body size due to temperature effects (phenotypic plasticity) was small compared to the interpopulation differences, which suggests substantial genetic differences between populations (subspecies) at different altitudes. In F₁ offspring from crosses between a low‐altitude (subspecies tosanus) and a high‐altitude population (subspecies ishizuchianus), adult body length was affected by the genotypes of both parents, with an interaction effect of parental genotype and offspring sex. Further analyses revealed that adult body length was affected by sex‐linked factors in addition to autosomal factors. These genetic differences in body size may have resulted from adaptations to different altitudes and may be important for the process of incipient speciation because body size differences could contribute to premating reproductive isolation.     

Altitudinal trends in climate change result in radial growth variation of Pinus yunnanensis at an arid-hot valley of southwest China

Climate change has already had observable impact on the biophysical environment, and lead to the different sensitivity of vegetation to climate factors on spatio-temporal scale. Therefore, understanding how the radial growth respond to climate at different spatio-temporal scales is crucial to recognize forest growth dynamic and make scientific management decisions under the background of climatic change. In the present study, the tree ring of Pinus yunnanensis at six altitudes gradients between 1300 m and 2500 m from a typical arid-hot valley in Jinsha River, were collected. We analyzed the relationship between radial growth and climate at different altitudes, and the sensitivity of growth to climatic factors over time. The results showed that the mean width of tree rings decreased as the altitude increasing. The relationship between climatic factors and radial growth at low or high altitudes was different with that at mid altitudes. Radial growth was negatively correlated to the temperatures from February to July at both low altitudes (1300–1500 m) and at high altitudes (2200–2500 m), but positively correlated to the temperatures in October of the previous year to April at mid altitudes (1700–1900 m). Precipitation in October of the previous year, May, and June in growing year had a positive effect on radial growth at all altitudes. Temperature and precipitation in the previous year showed a time-lag effect on radial growth. A moving correlation analysis of the tree ring index and climate variables showed that the limiting factors for tree growth at different altitudes varied over time. The influence of drought on the tree growth increased gradually as the climate warming. In future research, evaluating the dynamic relationship between vegetation growth and climate warming at spatio--temporal scale will be particularly important to guide forest management.     

Multi-scale altitudinal patterns in species richness of land snail communities in south-eastern France

Aim Species richness is an important feature of communities that varies along elevational gradients. Different patterns of distribution have been described in the literature for various taxonomic groups. This study aims to distinguish between species density and species richness and to describe, for land snails in south‐eastern France, the altitudinal patterns of both at different spatial scales. Location The study was conducted on five calcareous mountains in south‐eastern France (Etoile, Sainte Baume, Sainte Victoire, Ventoux and Queyras). Methods Stratified sampling according to vegetation and altitude was undertaken on five mountains, forming a composite altitudinal gradient ranging from 100 to 3100 m. Visual searching and analysis of turf samples were undertaken to collect land snail species. Species density is defined as the number of species found within quadrats of 25 m². Species richness is defined as the number of species found within an elevation zone. Different methods involving accumulation curves are used to describe the patterns in species richness. Elevation zones of different sizes are studied. Results Eighty‐seven species of land snails were recovered from 209 samples analysed during this study. Land snail species density, which can vary between 29 and 1 species per 25 m², decreases logarithmically with increasing altitude along the full gradient. However, on each mountain separately, only a linear decrease is observable. The climatic altitudinal gradient can explain a large part of this pattern, but the great variability suggests that other factors, such as heterogeneity of ground cover, also exert an influence on species density. The altitudinal pattern of species richness varies depending on the spatial resolution of the study. At fine resolution (altitudinal zones of 100 m) land snail species richness forms a plateau at altitudes below 1000 m, before decreasing with increasing altitude. At coarse resolution (altitudinal zones of 500 and 1000 m) the relationship becomes linear. Main conclusions This study reveals that land snail species density and land snail species richness form two different altitudinal patterns. Species density exhibits strong variability between sites of comparable altitude. A large number of samples seem necessary to study altitudinal patterns of species density. Species density decreases logarithmically with increasing altitude. Above a critical altitudinal threshold, this decrease lessens below the rate seen in the first 1500 m. Different methods exist to scale‐up species density to species richness but these often produce different patterns. In this study, the use of accumulation curves has yielded a pattern of species richness showing a plateau at low altitude, whereas simple plotting of known altitudinal ranges from single mountains would have produced stronger mid‐altitudinal peaks. This study shows that not only factors such as temperatures and habitat heterogeneity, but also an ecotone effect, are responsible for the observed patterns.     

Altitudinal divergence in maternal thermoregulatory behaviour may be driven by differences in selection on offspring survival in a viviparous lizard

Plastic responses to temperature during embryonic development are common in ectotherms, but their evolutionary relevance is poorly understood. Using a combination of field and laboratory approaches, we demonstrate altitudinal divergence in the strength of effects of maternal thermal opportunity on offspring birth date and body mass in a live-bearing lizard (Niveoscincus ocellatus). Poor thermal opportunity decreased birth weight at low altitudes where selection on body mass was negligible. In contrast, there was no effect of maternal thermal opportunity on body mass at high altitudes where natural selection favored heavy offspring. The weaker effect of poor maternal thermal opportunity on offspring development at high altitude was accompanied by a more active thermoregulation and higher body temperature in highland females. This may suggest that passive effects of temperature on embryonic development have resulted in evolution of adaptive behavioral compensation for poor thermal opportunity at high altitudes, but that direct effects of maternal thermal environment are maintained at low altitudes because they are not selected against. More generally, we suggest that phenotypic effects of maternal thermal opportunity or incubation temperature in reptiles will most commonly reflect weak selection for canalization or selection on maternal strategies rather than adaptive plasticity to match postnatal environments.