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.     

Contrasting patterns in local and zonal family richness of stream invertebrates along an Andean altitudinal gradient

1. Describing and understanding patterns in biological diversity along major geographical gradients is an important topic in ecology. Samples collected from a large number of physically and chemically comparable stream sites along a 4000 m gradient of altitude in the Andes of Ecuador served to characterise patterns of family richness of aquatic macroinvertebrates at the scale of the stream site (local) and at that of discrete altitudinal zones. 2. Both mean local and zonal family richness decreased by about 50% from sea level to 4000 m a.s.l. Local richness declined linearly, while zonal richness remained constant from sea level up to a threshold altitude of about 1800 m, whereafter it decreased. 3. From sea level to 1800 m few families were lost from zonal richness and few were gained. From 1800 to 3800 m the decrease in the number of families was accounted for by a loss of families present in lowland streams, with few new families gained. Hence, there was relatively little turnover of families along the entire gradient. 4. The diverging pattern of local and zonal richness was caused by sporadically occurring families inflating zonal richness at mid‐altitudes. If the sporadic families were represented by the same species found commonly in the lowlands, then the mid‐altitudinal zonal richness would be maintained by a ‘rescue effect’. More probably, however, the sporadically occurring families found at mid‐altitudes are each represented by new species replacing each other along the gradient, the families progressively diminishing in species richness and occurrence as the overall temperature tolerance of the family is approached. 5. This study demonstrates that spatial scale affects altitudinal patterns in the taxonomic richness of stream invertebrates. It also showed that family‐level identification can facilitate interpretation of sources and sinks of biodiversity along geographic gradients.     

Altitudinal patterns of seed plant richness in the Gaoligong Mountains, south-east Tibet, China

Elevational patterns of species richness and their underlying mechanisms have long been a controversial issue in biodiversity and biogeographical research, and several hypotheses have been proposed in the past decades. Local and regional studies have suggested that area and geometric constraint are two of major factors affecting the elevational pattern of species richness. In this study, using data of seed plants and their distribution ranges and a Digital Elevation Model data set, we explored altitudinal patterns of seed plant richness and quantified the effects of area and the mid-domain effect (MDE) on the richness patterns in a high mountain area, Gaoligong Mountains (ranging from 215 m to 5791 m a.s.l.) located in south-eastern Tibet, China. The results showed that richness and density (richness/log-transformed area) of seed plants at species, genus, and family levels all showed hump-shaped patterns along the altitudinal gradient. The altitudinal changes in richness of species with three different range sizes (< 500 m, 500–1500 m, and > 1500 m), species of different plant life-forms (trees, shrubs, and herbs), and endemic species further confirmed this finding. Analysis of Generalized Linear Model depicted that although the area of each elevational band was always in high correlation with the species richness, the MDE could explain 84.9%, 33.8%, 83.8%, and 84.5% of the total variation in richness for all species and the three species groups with different range sizes, respectively. This suggests that the MDE significantly influences the patterns of species richness and is likely be stronger for broad-ranged species than for narrow-ranged ones in the Gaoligong Mountains.     

Species Diversity and Altitudinal Gradient Patterns of Evergreen Broad leaved Forest in MeihuashanNational Natural Reserve,Fujian Province

In China, evergreen broad leaved forests (EBLFs) is one of the most important vegetation types which was widly distributed in subtropical area, and it plays a very important role in the global biological diversity and natural environment conservation also. In order to reveal species diversity and altitudinal gradient patterns of evergreen broad leaved forest in Meihuashan National Natural Reserve, Fujian Province. Five altitude transects were set up at a vertical interval of 200m between 375m and 1300m above sea level in the EBLFs distribution areas, and twenty four quadrats(14400m2) had been surveyed. Species richness(S), species richness index (dGl), Simpson index (D), Shannon Wiener index (H′), Pielou evenness index (J) had been used for analysis of species diversity and altitudinal gradient pattern of EBLFs. The average value of S, dGl, H′,J and D were 64.42, 10.75, 5.75, 3.50, 0.58 respectively. The difference of community species diversity index(S, dGl, D, H′, and J) was extremely significant between transects, and the altitudinal gradient patterns of species diversity presented the unimodal variable trend, with a peak in the mid altitude(700m-900m). The species richness and Shannon Wiener index of different layer were ranked as shrub layer (include young tree and the plants between layers)>arbor layer>herb layer. The species richness of tree and shrub layer, and Shannon Wiener index of tree layer were significantly different between at transects, and trends of altitude gradient was similar to community. The Shannon Wiener index of shrub layer and herb layer, and the species richness of herb layer did not change significantly along elevation gradient. Therefore, plant species diversity distribution pattern presented a unimodal variable trend along an elevation gradient, and supported “mid domain model” in EBLFs of Meihuashan National Nature Reserve.     

福建梅花山常绿阔叶林植物物种多样性及其海拔梯度格局

常绿阔叶林是福建梅花山国家级自然保护区地带性植被。采用样带与典型群落调查法对区内的常绿阔叶林14400m2样地展开调查,并对植物多样性海拔梯度格局进行分析,结果表明：（1） 群落植物物种丰富度、Gleason丰富度指数、Simpson指数、Shannon Wiener指数和Pielou均匀度指数的均值分别为64.42、10.75、5.75、3.50、0.58,且这5种指数在各样带间差异极为显著,并随海拔的升高均呈单峰曲线变化,峰值出现在海拔700m~900m。（2） 群落各层次的植物物种丰富度、Shannon Wiener指数均呈现灌木层（包括幼树和层间植物）〉乔木层〉草本层的特征。乔木、灌木层物种丰富度与乔木层Shannon Wiener指数在海拔梯度上的样带间差异极显著,变化趋势与群落相似;灌木层与草本层Shannon Wiener指数以及草本层物种丰富度随海拔梯度变化不明显。因此,梅花山自然保护区常绿阔叶林植物物种多样性的海拔梯度格局呈现单峰分布,并支持中间高度膨胀模式（mid domain model）。     

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.     

Elevational gradients of species diversity, breeding system and floral traits of orchid species on Réunion Island

Aim This research investigates changes in orchid species composition and diversity, plant breeding system and floral traits along an elevational gradient spanning 2300 m (200–2500 m). Location The study was conducted on Réunion Island (Mascarene Islands, Indian Ocean). Methods Data on the distribution of 135 orchid species from 35 genera were gathered from 121 localities situated between 200 and 2500 m a.s.l. For each locality, 500‐m transects were walked and each orchid species was recorded. Measures of species diversity (species richness, a modified Shannon diversity index and the modified Shannon equitability index) were related to altitude using ordinary least‐squares regression. Species turnover and elevational gradients in species composition were determined by: (1) relating scores of detrended correspondence analysis to altitude using ordinary least‐squares regression, and (2) relating Sørensen similarity indices to differences in altitude using Mantel tests. Finally, the average proportion of species displaying similar floral traits or showing the same breeding system were compared among altitudinal zones. Results Species richness per transect ranged from 1 to 36 species (mean 14.3) and decreased significantly with increasing altitude. Similarly, species evenness decreased significantly with increasing altitude. Around 50% of all orchid species were rare (occurred in fewer than 5% of all localities), and only a few occurred in more than 50% of all localities. Orchid species composition changed continuously with altitude, indicating turnover of species with increasing altitude. Analogously, orchid breeding systems and floral traits also changed with altitude. Relatively more auto‐pollinating species were found at high altitudes compared with mid‐ and low‐altitude sites where animal‐pollinated species were most abundant. Species characterized by a cleistogamous pollination system were found almost exclusively in high‐altitude sites, whereas the proportion of species displaying floral traits related to pollination by long‐tongued moths (sphinx) and flies sharply decreased with increasing altitude. Main conclusions Environmental conditions associated with altitude exert a large influence on orchid species composition and the distribution of orchid breeding systems. Our results revealed a high proportion of auto‐pollinating species, and confirm earlier findings that auto‐pollinating species are more frequent in high‐altitude sites.     

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.     

Altitudinal variation of dung beetle (Scarabaeidae: Scarabaeinae) assemblages in the Colombian Andes

Aim We describe the changes in species richness, rarity and composition with altitude, and explore whether the differences in Scarabaeinae dung beetle composition along five altitudinal transects of the same mountain range are related to altitude or if there are interregional differences in these altitudinal gradients. Location Field work was carried out on the eastern slope of the eastern Cordillera, Colombian Andes, between Tamá Peak to the north, in the Tamá National Park (07°23′ N, 72°23′ W) and the San Miguel River (00°28′ N, 77°17′ W) to the south. Methods Sampling was carried out between February 1997 and November 1999 in five regions spanning elevation gradients. In each gradient, six sites were chosen at 250 m intervals between 1000 and 2250 m a.s.l. Results We found a curvilinear relationship between altitude and mean species richness, with a peak in richness at middle elevations. However, the diversity of dung beetle assemblages does not seem to be related to the interregional differences in environmental conditions. The number of geographically restricted species is negatively and significantly related to altitude, with geographically restricted species more frequent at low altitude sites. Ordination delimited the two main groups according to altitude: one with all the highest sites (1750–2250 m a.s.l.) and a second group with the remaining sites (< 1750 m a.s.l.). Analysis of species co‐occurrence shows that these dung beetle assemblages seem to be spatially structured when all sites have the same probability of being chosen. In contrast, the spatial structure of species assemblages seems to be random when the probability of choosing any site is proportional to its altitude. Main conclusions The altitude of sites is the main factor that influences the diversity of these dung beetle assemblages. The peak in species richness at middle elevations, the higher number of geographically restricted species at lower altitudinal levels, and the compositional differences along these mountain gradients seem to result from the mixing at these altitudes of dung beetle assemblages that have different environmental adaptations and, probably, different origins. The relevance of altitude in these assemblages is related to the limited role of these Neotropical high altitude environments as centres of refuge and vicariance for a monophyletic group of warm‐adapted species, for which the vertical colonization of these high mountain environments by lineages distributed at lower altitudes would have been very difficult.     

Terrestrial isopod diversity in the wadi Moula‐Bouterfess catchment area (Kroumirie,north‐west of Tunisia)

We assessed the diversity of terrestrial isopods according to habitat and altitude in the wadi of Moula‐Bouterfess catchment area. The most representative habitat types for the area were selected within altitudinal range from 6 to 550 m a.s.l. In total nine sites were sampled: four with maquis and garrigue vegetation, one at a meadow and four sites with forest vegetation. In each sampled habitat, individuals were collected by hand in April 2005 using 30 replicates of a quadrate of 0.5 × 0.5 m. During the study, 582 individuals belonging to 11 terrestrial isopod species from five families were collected. Among these, three species were newly mentioned in Tunisia. The genus Armadillidium was the most abundant genus (60% of the total number of collected specimens) and P. variabilis was the most common species as it was sampled in eight of the nine studied habitats. Terrestrial isopod community structure differs among the nine sampling habitats. Abundance and species richness values are low in the different studied habitats. The Shannon–Wiener H′ varied from 0.67 to 2.06. The Canonical Correspondence Analysis (CCA) applied to our data showed that the majority of terrestrial isopod species seem to be more sensitive to vegetal associations than to altitude. No relationship was found between species richness/diversity (Shannon–Wiener index H’) and altitude. The studied sites were separated into open and closed areas based on the Bray–Curtis index for similarity.     

From lowlands to highlands: searching for elevational patterns of species richness and distribution of scarab beetles in Costa Rica

Aim Understanding the heterogeneous distribution of species on mountains is an important aim in ecology. Altitudinal gradients have enormous potential for improving our knowledge of trends in biodiversity and conservation. In this study, we investigated the variation in scarab beetle diversity (Dynastinae, Rutelinae and Melolonthinae) along an elevational tropical forest gradient. Location The Atlantic slope of the Guanacaste mountain range in Costa Rica. Methods Ultraviolet light traps placed in six forests situated from 100 to 1510 m were used. Changes in species composition and richness among elevations were investigated. Differences in the altitudinal patterns using different groups of species were examined: the whole assemblage, each separate subfamily and two different trophic habits (phytophagous or saproxylic). The effects of temperature, humidity and elevation on scarab distribution were tested using canonical correspondence analyses. The relationship between the community similarity of the studied forests and the altitudinal distance among them was also analysed. Results Species composition and richness changed along the gradient. The peak in species richness varied depending on the species group considered and in all cases occurred 500 or 800 m. Forests at these altitudes were also the richest in exclusive species. Species composition turnover among elevations appeared with a clear separation between lowland and highland fauna. The latter was lower in richness but also had exclusive species. Temperature, humidity and altitude affected species distribution, with altitude being the most important factor for all the subfamilies studied. Main conclusions Our results showed that species distribution fits a hump‐shaped pattern. The peak of this pattern varied depending on the taxonomic group and mountain analysed, highlighting the importance of evolutionary processes as species distribution drivers. The fact that species richness peaked at elevations where human impact is currently important underlines the value of the development of conservation strategies for these areas.     

Diversity of terrestrial isopod species in the Chambi National Park (Kasserine,Tunisia)

Diversity of terrestrial isopods across habitats and altitude was studied in the National Park of Chambi (central Tunisia). Samples were collected over five years in seven sites (S1–S7), within an altitudinal range from 750 to 1,500 m a.s.l. Twelve species belonging to five families were identified, with one endemic species (Porcellio djahizi) to this area, and three species were present in all sites (P. djahizi, Leptotrichus panzerii and Armadillidium tunisiense). Agnaridae represented by Hemilepistus reaumurii species was the most abundant family (58.77%) and found in only one site. Our results showed differences in species richness between sites. Indeed, S2 and S3 (900 and 1,050 m) showed the highest species richness with 11 species each. This result validates the hypothesis of mid‐elevational richness peak and contradicts the theory of large sampling regimes in determining the relationship between species richness and elevation. Terrestrial isopod species richness in the Chambi Mountain tends to decrease with altitude: 11 species in S2 and S3 to four species in S7. The highest Shannon–Wiener diversity index value was observed in S2 (H′ = 3.21 bits). Canonical correspondence analysis revealed that H. reaumurii was positively correlated with temperature whereas A. tunisiense and P. djahizi were the only species positively correlated with altitude.     

The distribution of ground spiders (Araneae, Gnaphosidae) along the altitudinal gradient of Crete, Greece: species richness, activity and altitudinal range

Aim To study the altitudinal variation of ground spiders (Araneae, Gnaphosidae) of Crete, Greece, as far as species composition, species richness, activity and range of distribution are concerned. Location Altitudinal zones (0–2400 m) along the three main mountain massifs of the island of Crete. Methods Thirty‐three sampling sites were located from 0 to 2400 m a.s.l. on Crete, and sampled using pitfall traps. Material from the high‐activity period of Gnaphosidae (mid‐spring to mid‐autumn) was analysed. Sampling sites were divided into five altitudinal zones of 500 m each. Statistical analysis involved univariate statistics (anova ) and multivariate statistics, such as multidimensional scaling (MDS) and cluster analysis (UPGMA) using binomial data of species presence or absence. Results Species richness declines with altitude and follows a hump‐shaped pattern. The activity pattern of the family, as a whole, is not correlated with altitude and is highly species‐specific. In the highest zone, both species richness and activity decline dramatically. The altitudinal range of species distribution increases with altitude. On the Cretan summits live highly tolerant lowland species and isolated residents of the high mountains of Crete. Two different patterns of community structure are recorded. Main conclusions Communities of Gnaphosidae on Crete present two distinct structures following the altitudinal gradient, these being separated by a transitional zone between 1600 and 2000 m. This study supports previous results which show a hump‐shaped decline in species richness of Gnaphosidae along altitudinal gradients, leading to a peak at 400–700 m, where an optimum of environmental factors exists. This makes this zone the meeting point of the often opportunistic lowland species with the older and most permanent residents of the island. Rapoport's rule on the positive correlation of the altitudinal range of species distributions with altitude is also supported. The high activity recorded for the species that persist on the high mountains of Crete is indicative of a tolerant arachnofauna, and is considered to result from relaxation of competitive interactions with other species. This is related to a reduction in species numbers, shortening of the activity period on high mountains and the unique presence of high mountain species that thrive only there. As shown in our study, strategies to cope with altitude are species‐specific. Therefore, there cannot exist one single model to describe how animals react to the change in altitude, even under the same environmental conditions.     

Spatial gradients in species diversity of microscopic animals: the case of bdelloid rotifers at high altitude

Aim Organisms smaller than 2 mm appear not to follow the spatial patterns in richness and diversity commonly observed in macroscopic organisms. We describe spatial patterns in species diversity in a group of microscopic organisms, bdelloid rotifers, living in moss and lichen patches, in order to test the hypotheses of no relationship between species richness and composition and spatial gradients, suggested by previously published patterns in microscopic organisms. Location Moss and lichen patches as habitats for bdelloids, on high‐elevation peaks at altitudes between 2984 and 4527 m a.s.l. across the Italian, French and Swiss Alps, with distances among sample sites ranging from 1 m to 420 km, in comparison with lower‐elevation samples at altitudes from 850 to 1810 m a.s.l. Methods We sampled species assemblages of bdelloid rotifers living in isolated moss and lichen patches in 47 sites. We described the observed α, β and γ diversities; the heterogeneity of species assemblages; and the estimated number of species (incidence‐based coverage estimator). Patterns in species distribution were analysed at three different levels: (1) habitat, comparing species richness on moss and lichen substrates, testing differences in α diversity and heterogeneity (anova ), species composition (analysis of similarities test), and γ diversity (rarefaction curves); (2) altitude, comparing the observed richness with previously published data from locations well below 2000 m; and (3) distances between sites, correlating the matrix of Jaccard dissimilarities and the matrix of geographical distances with a Mantel test. Results Both species richness and species composition of bdelloid rotifers differed significantly between mosses and lichens at high elevations, but no difference was found in the heterogeneity of species assemblages. Alpha diversity was significantly lower at high‐elevation than at low‐elevation sites, but the estimated number of species was not reduced when compared with sites at low elevations. Geographical distance between sites had no effect on species composition in either mosses or lichens. The distribution of species was highly heterogeneous, with a low similarity among assemblages. Main conclusions As expected, bdelloids appear to occupy habitats selectively. The altitudinal gradient in species richness for bdelloid rotifers is limited to a decrease in α diversity only; such a decrease is not caused by a lower number of species (low γ diversity) being able to tolerate harsh conditions, and high‐altitude species are not a subset of species living at lower elevations. The observed values of α, β and γ diversity at high altitudes in the Alps are compatible with the scenario of a very low number of available propagules because of the low density of patches of favourable habitat. Our results suggest that the geographical distribution of animals, and therefore biodiversity patterns, may be strongly influenced by animal size, as small organisms such as bdelloids appear to show spatial patterns that differ from those known in larger animals. Differences in body size should be taken into account carefully in future studies of biodiversity patterns.     

Variation in invertebrate–bryophyte community structure at different spatial scales along altitudinal gradients

Aim This study assessed changes in diversity and assemblage composition in bryophytes and their associated invertebrates along altitudinal gradients in Australia and New Zealand. The importance of altitude in shaping these communities and for the diversity of both invertebrates and bryophytes was examined at different spatial scales, including local, altitudinal, regional and biogeographical. Location Samples were taken from four Australasian mountain ranges between 42° and 43°S: Mt Field and Mt Rufus, Tasmania, Australia, and Otira Valley and Seaward Kaikoura Mountains, South Island, New Zealand. Methods On both Tasmanian mountains, five altitudes were assessed (250, 500, 750, 1000 and 1250 m). At each location (mountain/altitude combination) two sites were chosen and six samples were taken. Six altitudes were assessed on New Zealand mountains (Otira: 250, 500, 750, 1000, 1250 and 1500 m; Kaikoura: 1130, 1225, 1325, 1425, 1525 and 2000 m). Bryophyte substrate was collected, and all samples were stored in 70% ethanol. Invertebrates were extracted from bryophytes using kerosene‐phase separation and all invertebrates were identified to family. At each location in Tasmania, all bryophyte species within six 25‐cm² grids per site were collected and identified to species. Bryophytes from New Zealand were identified to species from the invertebrate sample substrate because of sampling constraints. Results Altitude did have a significant effect on diversity, however, no general trend was found along the altitudinal gradient on the four mountains. There were distinct differences in diversity between biogeographical regions, mountains, altitudes and sites. In Tasmania, Mt Field had the highest diversity in invertebrates and bryophytes at 750 m. In contrast, Mt Rufus had consistent low invertebrate and bryophyte diversity along the entire altitudinal gradient. There were also distinctive differences between locations in the composition of invertebrate and bryophyte communities in Tasmania. Along the two altitudinal gradients in New Zealand, Otira had highest diversity for both invertebrates and bryophytes at low altitudes, whereas Kaikoura had highest invertebrate and lowest bryophyte diversity at the highest altitude. Main conclusions There was an effect of altitude, however, there were no consistent changes in diversity or composition on the four different mountains. There was considerable local and regional variation, and, despite a strong sampling design, no underlying altitudinal trends were detectable. This study demonstrates the importance of examining a range of spatial scales if patterns in community structure along altitudinal gradients are to be studied. The implications of this study are discussed with reference to survey design, taxonomic resolution, climate change and conservation of habitat.     

Spatial diversity patterns of Pristimantis frogs in the Tropical Andes

Although biodiversity gradients have been widely documented, the factors governing broad‐scale patterns in species richness are still a source of intense debate and interest in ecology, evolution, and conservation biology. Here, we tested whether spatial hypotheses (species–area effect, topographic heterogeneity, mid‐domain null model, and latitudinal effect) explain the pattern of diversity observed along the altitudinal gradient of Andean rain frogs of the genus Pristimantis. We compiled a gamma‐diversity database of 378 species of Pristimantis from the tropical Andes, specifically from Colombia to Bolivia, using records collected above 500 m.a.s.l. Analyses were performed at three spatial levels: Tropical Andes as a whole, split in its two main domains (Northern and Central Andes), and split in its 11 main mountain ranges. Species richness, area, and topographic heterogeneity were calculated for each 500‐m‐width elevational band. Spatial hypotheses were tested using linear regression models. We examined the fit of the observed diversity to the mid‐domain hypothesis using randomizations. The species richness of Pristimantis showed a hump‐shaped pattern across most of the altitudinal gradients of the Tropical Andes. There was high variability in the relationship between area and species richness along the Tropical Andes. Correcting for area effects had little impact in the shape of the empirical pattern of biodiversity curves. Mid‐domain models produced similar gradients in species richness relative to empirical gradients, but the fit varied among mountain ranges. The effect of topographic heterogeneity on species richness varied among mountain ranges. There was a significant negative relationship between latitude and species richness. Our findings suggest that spatial processes partially explain the richness patterns of Pristimantis frogs along the Tropical Andes. Explaining the current patterns of biodiversity in this hot spot may require further studies on other possible underlying mechanisms (e.g., historical, biotic, or climatic hypotheses) to elucidate the factors that limit the ranges of species along this elevational gradient.     

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.     

三江并流地区干旱河谷植物物种多样性海拔梯度格局比较

在滇西北三江并流地区典型干旱河谷段, 在怒江、澜沧江和金沙江的东、西坡共设置了6条海拔梯度样带, 通过标准样地的植物群落调查, 分析各条样带植物的物种丰富度、物种更替率的海拔梯度格局, 并比较了地理和植被变量对分布格局的解释。干旱河谷植被带位于海拔3,000 m以下, 以灌丛和灌草丛为主, 其在各河谷的分布上限自西向东依次升高。植物物种丰富度的分布主要与海拔、流域、经纬度和植被带有关, 沿纬度和海拔梯度升高而显著增加的格局主要表现在草本层和灌木层, 灌木物种丰富度还呈现自西向东显著增加的趋势。怒江的灌木和草本种物种丰富度显著高于金沙江和澜沧江, 三条江的乔木种丰富度差异则不显著。森林带的样方草本物种丰富度显著低于灌草丛带样方, 并且还拥有后者没有的乔木种。不同样带的植物物种更替速率呈现了不一致的海拔梯度格局, 但均在样带海拔下部的灌草丛群落与海拔上部森林群落之间的交错带出现峰值。森林-灌草丛植被交错带在怒江样带处于海拔1,900-2,100 m处, 在澜沧江河谷位于海拔2,300-2,400 m, 在金沙江河谷位于海拔2,700-2,900 m。所有海拔样带的森林段或灌草丛段相对于同一样带不同植被段之间的物种更替程度为最小, 不仅小于同一流域不同样带相同植被段之间物种更替率的均值, 更小于所有样带相同植被段之间的更替率均值。在三条河流6条海拔样带的12个植被带段之间的物种更替变化中, 空间隔离因素可以解释34.2%, 而植被类型差异仅能解释不到0.5%。本研究结果显示了环境差异对不同植被类型物种丰富度的首要影响, 和各河流之间的空间隔离对植物群落构建和物种构成的主要作用。     

Disturbance from traditional fire management in subalpine heathlands increases Afro‐alpine plant resilience to climate change

Species are often controlled by biotic factors such as competition at the warm edge of their distribution range. Disturbances at the treeline, disrupting competitive dominance, may thus enable alpine species to utilize lower altitudes. We searched for evidence for range expansion in grazed, fire‐managed Ethiopian subalpine Erica heathlands across a 25‐year chronosequence. We examined vascular plant composition in 48 plots (5 × 5 m) across an altitudinal range of 3,465–3,711 m.a.s.l. and analyzed how community composition changed in relation to increasing competition over time (using a Shade index based on Erica shrub height and cover) and altitude. Species‘ habitats and altitudinal ranges were derived from literature. Time since fire explained more variation (r² = .41) in species composition than altitude did (r² = .32) in an NMDS analysis. Community‐weighted altitudinal optima for species in a plot decreased strongly with increasing shade (GLM, Standardized Regression Coefficient SRC = ?.41, p = .003), but increased only weakly with altitude (SRC = .26, p = .054). In other words, young stands were dominated by species with higher altitudinal optima than old stands. Forest species richness increased with Log Shade index (SRC = .12, p = .008), but was unaffected by altitude (SRC = ?.07, p = .13). However, richness of alpine and heathland species was not highest in plots with lowest Shade index, but displayed a unimodal pattern with an initial increase, followed by a decrease when shading increased (altitude was not significant). Our results indicate that disturbance from the traditional patch burning increases the available habitat for less competitive high‐altitude plants and prevents tree line ascent. Therefore, maintaining, but regulating, the traditional land use increases the Afro‐alpine flora's resilience to global warming. However, this system is threatened by a new REDD+ program attempting to increase carbon storage via fire suppression. This study highlights the importance of understanding traditional management regimes for biodiversity conservation in cultural landscapes in an era of global change.     

An altitudinal comparison of caterpillar (Lepidoptera) assemblages on Ficus trees in Papua New Guinea

Aim This analysis of caterpillar (Lepidoptera) beta‐diversity between tropical lowlands and highlands attempts to separate the effects of between‐site (1) turnover of herbivore species on particular host plants, (2) changes in host use by herbivores, and (3) turnover of plant species on changes in herbivore assemblages. Location Two rain forest areas 130 km and 1700 altitudinal metres apart were studied in Papua New Guinea: one in the lowlands (100 m a.s.l.) on the northern coast of the island and one in the central New Guinean cordillera at 1800 m a.s.l. Methods The analysis is based on caterpillar feeding records obtained by quantitative sampling and rearing of caterpillars from four Ficus species studied in the mountains and 21 Ficus species and 62 plant species from other genera and families studied in the lowlands, including three Ficus species studied in both areas. Results Only 17% of species feeding on Ficus in the highlands also occurred in the lowlands. These species represented 1–46% of individuals in caterpillar assemblages on particular Ficus hosts. Widespread species included both Ficus specialists and generalists feeding on numerous plant families. Some of the Ficus specialists changed their preferred host species with altitude. High species turnover was not explained by changes in the species composition of host plants with altitude as lowland and montane assemblages feeding on the same Ficus species showed high turnover. Despite the rarity of widespread caterpillars, the lowland and montane Ficus assemblages were remarkably similar in their dominance structure, species richness, host specificity, generic composition and familial composition. Main conclusions Ficus‐feeding Lepidoptera assemblages between tropical lowlands and highlands are characterized by substantial species turnover not explained by altitudinal changes in the composition of the vegetation. Further, species‐rich plant genera can support caterpillar assemblages with relatively low beta‐diversity compared with species‐poor genera as caterpillars can switch their host preferences from one congeneric host species to another along an altitudinal gradient. Closely related plant species can thus represent a broad, continuously distributed resource along such gradients.