Distribution and Flowering Ecology of Bromeliads along Two Climatically Contrasting Elevational Transects in the Bolivian Andes1

We compared the diversity, taxonomic composition, and pollination syndromes of bromeliad assemblages and the diversity and abundance of hummingbirds along two climatically contrasting elevational gradients in Bolivia. Elevational patterns of bromeliad species richness differed noticeably between transects. Along the continuously wet Carrasco transect, species richness peaked at mid‐elevations, whereas at Masicurí most species were found in the hot, semiarid lowlands. Bromeliad assemblages were dominated by large epiphytic tank bromeliads at Carrasco and by small epiphytic, atmospheric tillandsias at Masicurí. In contrast to the epiphytic taxa, terrestrial bromeliads showed similar distributions across both transects. At Carrasco, hummingbird‐pollination was the most common pollination mode, whereas at Masicurí most species were entomophilous. The proportion of ornithophilous species increased with elevation on both transects, whereas entomophily showed the opposite pattern. At Carrasco, the percentage of ornithophilous bromeliad species was significantly correlated with hummingbird abundance but not with hummingbird species richness. Bat‐pollination was linked to humid, tropical conditions in accordance with the high species richness of bats in tropical lowlands. At Carrasco, mixed hummingbird/bat‐pollination was found especially at mid‐elevations, i.e., on the transition between preferential bat‐pollination in the lowlands and preferential hummingbird‐pollination in the highlands. In conclusion, both richness patterns and pollination syndromes of bromeliad assemblages varied in distinct and readily interpretable ways in relation to environmental humidity and temperature, and bromeliad pollination syndromes appear to follow the elevational gradients exhibited by their pollinators.     

Bird diversity along elevational gradients in the Andes of Colombia: area and mass effects

Aim The decrease in species richness with increasing elevation is a widely recognized pattern. However, recent work has shown that there is variation in the shape of the curve, such that both negative monotonic or unimodal patterns occur, influenced by a variety of factors at local and regional scales. Discerning the shape of the curve may provide clues to the underlying causes of the observed pattern. At regional scales, the area of the altitudinal belts and mass effects are important determinants of species richness. This paper explores the relationship between bird species richness, elevation, mass effects and area of altitudinal zones for birds in tropical mountains. Location The three Andean ranges of Colombia and the peripheral mountain ranges of La Macarena and Santa Marta. Methods Lists of bird species were compiled for altitudinal belts in eastern and western slopes of the three Andean Cordilleras and for La Macarena and Santa Marta. The area of the altitudinal belts was computed from digital elevation models. The effect of area was analysed by testing for differences among altitudinal belts in the slopes and intercepts of the species‐area relationships. Mass effects were explored by separately analysing two sets of species: broadly distributed species, i.e. lowland species whose distributions extend into the Andes, and tropical Andean species, i.e., species that evolved in the Andes. Results Plotting total number of species in each altitudinal belt revealed a decline in species richness with elevation. In slopes with a complete elevational gradient from lowlands to mountain peaks, the decrease was monotonic. In internal Andean slopes where the lower elevational belts are truncated, there was a peak at mid elevations. There was a linear relationship between number of species and area of the altitudinal belts. When controlling for area, there were no differences in the number of species among altitudinal belts (500–2600 m), except for the two upper‐elevation zones (2600–3200 and > 3200 m), which had lower species richness. Diversity of widely distributed species declined monotonically with elevation, whereas tropical Andean species exhibited a mid‐elevation peak. Main conclusions A large proportion of the variation in species richness with elevation was explained by area of the altitudinal belts. When controlling for area, species richness remained constant up to 2600 m and then decreased. This pattern contrasts with a previously reported hump‐shaped pattern for Andean birds. Diversity patterns of widely distributed species suggested that immigration of lowland species inflates diversity of lower elevational belts through mass effects. This influence was particularly evident in slopes with complete altitudinal gradients (i.e. connected to the lowlands). Tropical Andean species, in contrast, were more diverse in mid‐elevational belts, where speciation rates are expected to be higher. The influence of these species was more prevalent in internal Andean slopes with no connection to the lowlands. The decline of species richness at high elevations may be related to higher extinction rates and lower resource levels.     

Diversity of geometrid moths (Lepidoptera: Geometridae) along an Afrotropical elevational rainforest transect

Geometrid moths were investigated at 26 sites on 9 elevational levels along an elevational transect at Mt. Kilimanjaro (Tanzania), stretching from the fine‐grained mosaic of small agroforest plots with combined cultivation of trees, shrubs and crops at 1650 m through mountain rainforest to heathland at 3300 m. We sampled moths manually at light between 19 : 00 and 22 : 00 in the rainy seasons of March to May and October to January in the years 2000, 2001 and 2002. Along the transect, the composition of moth communities changed from a domination by Sterrhinae and Ennominae to a dominance of Larentiinae with increasing elevation. Overall, alpha diversity was very low compared to other tropical mountain regions. Fisher's alpha showed a maximum of 30 in the agroforest mosaic at 1650 m and decreased to values around 12 in the mountain rainforest. Communities of geometrid moths within the forest belt were significantly dissimilar from communities outside the forest. The diversity patterns on Mt. Kilimanjaro can be related to the young age, island‐like position and history of the mountain. These factors have led to the formation of a homogeneous upper mountain rainforest habitat which in turn houses homogeneous moth communities with a low diversity compared to habitats at lower elevations. Here, a heterogeneous habitat mosaic allowing the intrusion of savannah species into this former forest habitat may account for an increased diversity. In the heath zone above the forest, climatic conditions are very harsh, permitting only few specialists to thrive in this ericaceous woodland. Edge effects were discernible at the forest–heathland boundary where some moth species from heathland invaded the closed forest. At the boundary between agroforest and a forest mosaic of exotic Acacia and Eucalyptus forest plantations and natural mountain forest, diversity values remained low as the dominant species Chiasmia fuscataria accounted for far higher proportions than other dominant species in any of the other habitats.     

Elevational zonation of afrotropical forest bird communities along a homogeneous forest gradient

Aim This study analyses the distribution and abundance of birds from a forested tropical gradient in order to determine whether elevationally distinct communities are detectable in this habitat. Location An avifaunal census was carried out on a single transect within the tropical forest of the Udzungwa Mountains in the Eastern Arc, Tanzania, covering a range in elevation from 300 to 1850 m. Methods Two complementary data sets on forest birds were analysed, encompassing (1) data derived from standardized 20‐ha spot‐mapping censuses performed at nine elevations over 175‐m intervals from 400 to 1800 m a.s.l., and (2) all observations of birds binned into 32 data points at 50‐m intervals, from 300 to 1850 m a.s.l. The degree of zonation in the avian community along the elevational gradient was examined using the chronological clustering method, an agglomerative hierarchical clustering method that can be carried out with a range of similarity indices. Results The chronological clustering analysis of the data set based on standardized spot‐mapping revealed a clearly defined boundary at c. 1200 m a.s.l., separating lowland from montane communities. Most bird species could be categorized as belonging to one of these two communities. The data set based on all observations revealed a number of potential secondary boundaries, although these boundaries delimited the entire elevational ranges of individual species in only relatively few cases. Main conclusions In contrast to previously published studies, we find evidence of an elevational zonation of distinct communities within a seemingly homogeneous habitat. Although similar boundaries have been assumed to arise as a result of vegetational ecotones, or because of interspecific competition, these mechanisms are poorly corroborated. We suggest that the causes of patterns of zonation are not well understood, and that the interplay among species distributions, species richness, and environmental factors needs more consideration. The chronological clustering method is proposed as an appropriate tool for studying these specific patterns.     

Forest aboveground biomass along an elevational transect in Sulawesi,Indonesia, and the role of Fagaceae in tropical montane rain forests

Aim This study investigates how estimated tree aboveground biomass (AGB) of tropical montane rain forests varies with elevation, and how this variation is related to elevational change in floristic composition, phylogenetic community structure and the biogeography of the dominant tree taxa. Location Lore Lindu National Park, Sulawesi, Indonesia. Methods Floristic inventories and stand structural analyses were conducted on 13 plots (each 0.24 ha) in four old‐growth forest stands at 1050, 1400, 1800 and 2400 m a.s.l. (submontane to upper montane elevations). Tree AGB estimates were based on d.b.h., height and wood specific gravity. Phylogenetic diversity and biogeographical patterns were analysed based on tree family composition weighted by AGB. Elevational trends in AGB were compared with other Southeast Asian and Neotropical transect studies (n = 7). Results AGB was invariant from sub‐ to mid‐montane elevation (309–301 Mg ha^?1) and increased slightly to 323 Mg ha^?1 at upper montane elevation. While tree and canopy height decreased, wood specific gravity increased. Magnoliids accounted for most of the AGB at submontane elevations, while eurosids I (including Fagaceae) contributed substantially to AGB at all elevations. Phylogenetic diversity was highest at upper montane elevations, with co‐dominance of tree ferns, Podocarpaceae, Trimeniaceae and asterids/euasterids II, and was lowest at lower/mid‐montane elevations, where Fagaceae contributed > 50% of AGB. Biogeographical patterns showed a progression from dominant tropical families at submontane to tropical Fagaceae (Castanopsis, Lithocarpus) at lower/mid‐montane, and to conifers and Australasian endemics at upper montane elevations. Cross‐continental comparisons revealed an elevational AGB decrease in transects with low/no presence of Fagaceae, but relatively high AGB in montane forests with moderate to high abundance of this family. Main conclusions AGB is determined by both changes in forest structure and shifts in species composition. In our study, these two factors traded off so that there was no net change in AGB, even though there were large changes in forest structure and composition along the elevational gradient. Southeast Asian montane rain forests dominated by Fagaceae constitute important carbon stocks. The importance of biogeography and species traits for biomass estimation should be considered by initiatives to reduce emissions from deforestation and forest degradation (REDD) and in taxon choice in reforestation for carbon offsetting.     

Vegetation patterns and environmental gradients in tropical dry forests of the northern Yucatan Peninsula

Patterns of plant species composition and their relationships to soil and topographic variables were investigated in tropical dry forests across the north central Yucatan, Mexico. Seven sites were studied in the oldest accessible forests along a 200–km transect oriented northwest to southeast; an eighth site was located in a little‐disturbed area located 75 km northeast of the transect. Two of the sites were on Mayan ruins. All sites were sampled using 9–24, 10m × 20m plots (<n= 132) for woody stems ≥ 3.0 cm diameter breast height. The important natural forest species were Bursera simaruba, Caesalpinia gaumeri, Gymnopodium floribundum, Piscidia piscipula, and Thouinia paucidentata. The two most important woody species in ruin woodlands were Brosimum alicastrum and Croton lundellii. Forest plots (n=108) had 17 species on average, ruin plots (n= 24) nine species. Mean basal area of stems at the forest plots (20.7 m².ha^‐1) was lower than in ruin plots (28.4 m².ha^‐1). Detrended Correspondence Analysis generally placed plots by site along the geographic transect. Natural forest plots and sites were separated from the plots on ruin sites. The five soil and topographic variables (slope, soil depth, percent surface rock, soil pH, total soil organic matter) differed significantly among sites. Plot values were correlated with DCA axe scores. Intersite floristic variation reflects an overall west to east environmental gradient affected by climate.     

Environmental determinants of leaf litter ant community composition along an elevational gradient

Ant communities are extremely diverse and provide a wide variety of ecological functions in tropical forests. Here, we investigated the abiotic factors driving ant composition turnover across an elevational gradient at Mont Itoupé, French Guiana. Mont Itoupé is an isolated mountain whose top is covered by cloud forests, a biogeographical rarity that is likely to be threatened according to climate change scenarios in the region. We examined the influence of six soil, climatic, and LiDAR‐derived vegetation structural variables on leaf litter ant assembly (267 species) across nine 0.12‐ha plots disposed at three elevations (ca. 400, 600, and 800m asl). We tested (a) whether species cooccurring within a same plot or a same elevation were more similar in terms of taxonomic, functional, and phylogenetic composition, than species from different plots/elevations, and (b) which environmental variables significantly explained compositional turnover among plots. We found that the distribution of species and traits of ant communities along the elevational gradient was significantly explained by a turnover of environmental conditions, particularly in soil phosphorus and sand content, canopy height, and mean annual relative humidity of soil. Our results shed light on the role exerted by environmental filtering in shaping ant community assembly in tropical forests. Identifying the environmental determinants of ant species distribution along tropical elevational gradients could help predicting the future impacts of global warming on biodiversity organization in vulnerable environments such as cloud forests.     

Continuum or zonation? Altitudinal gradients in the forest vegetation of Mt. Kilimanjaro

Based on the analysis of 600 vegetation plots using the method of Braun-Blanquet (1964) the composition of the whole vascular forest plant flora with about 1220 species was studied in the forests of Mt. Kilimanjaro. The altitudinal distribution of all strata (trees, shrubs, epiphytes, lianas and herbs) along a transect of 2400 m is discussed with respect to altitudinal zonation and ecological factors. With uni-dimensionally constraint clustering significant discontinuities were revealed that occurred simultaneously in the different strata. Thus even in structurally highly complex, multilayered tropical montane forests distinct community units exist that can be surveyed and classified by the Braun-Blanquet approach. This observed zonation was significantly correlated with altitude, temperature and soil acidity (pH); rainfall was of importance in particular for the zonation of epiphytes. Other key factors were humidity (influenced by stable cloud condensation belts) and minimum temperature (in particular the occurrence of frost at 2700 m altitude upslope). The contrary results of other transect studies in East Africa in respect to continuity of change in floristic composition appear to be caused by different sampling methods and intensities or mixing of data from areas with different climate conditions, whereas species richness did not influence the clarity of floristic discontinuities on Kilimanjaro and other parts of East Africa.     

Elevation‐dependent effects of forest fragmentation on plant–bird interaction networks in the tropical Andes

Tropical forests harbor diverse ecological communities of plants and animals that are organized in complex interaction networks. The diversity and structure of plant–animal interaction networks may change along elevational gradients and in response to human‐induced habitat fragmentation. While previous studies have analyzed the effects of elevation and forest fragmentation on species interaction networks in isolation, to our knowledge no study has investigated whether the effects of forest fragmentation on species interactions may differ along elevational gradients. In this study, we analyzed main and interaction effects of elevation and forest fragmentation on plant–frugivore interaction networks at plant and bird species level. Over a period spanning two years, we recorded plant–frugivore interactions at three elevations (1000, 2000 and 3000 m a.s.l.) and in two habitat types (continuous and fragmented forest) in tropical montane forests in southern Ecuador. We found a consistent effect of elevation on the structure of plant–frugivore networks. We observed a decrease in the number of effective bird partners of plants and, thus, a decline in the redundancy of bird species with increasing elevation. Furthermore, bird specialization on specific plant partners increased towards high elevations. Fragmentation had a relatively weak effect on the interaction networks for both plant and bird species, but resulted in a significant increase in bird specialization in fragmented forests at high elevations. Our results indicate that forest fragmentation may have stronger effects on plant–frugivore interaction networks at high compared to low elevations because bird species richness declined more steeply towards high elevations than plant species richness. We conclude that conservation efforts should prioritize the maintenance of consumer diversity, for instance by maintaining stretches of continuous forest. This applies in particular to species‐poor communities, such as those at high elevations, as the ecological processes in these communities seem most sensitive towards forest fragmentation.     

Changes in Species Richness,Abundance, and Composition of Arboreal Twig‐nesting Ants Along an Elevational Gradient in Coffee Landscapes

The distribution, diversity, and assembly of tropical insects have long intrigued ecologists, and for tropical ants, can be affected by competitive interactions, microhabitat requirements, dispersal, and availability and diversity of nesting sites. Arboreal twig‐nesting ants are limited by the number of hollow twigs available, especially in intensive agricultural systems. Ant diversity and abundance may shift along elevation gradients, but no studies have examined if the proportion of occupied twigs or richness of arboreal twig‐nesting ants vary with elevation. In coffee agroecosystems, there are over 40 species of arboreal twig‐nesting ants. We examined communities of twig‐nesting ants in coffee plants along an elevational gradient to answer the following questions: (1) Do species richness and colony abundance decline with elevation or show a mid‐elevation peak? (2) Does community composition change with elevation? (3) Is elevation an important predictor of change in ant abundance, richness, and relative abundance of common species? We surveyed 42 10 × 10 m plots in 2013 from 450 to1550 m elevation across a coffee landscape in Chiapas, Mexico. We sampled a total of 2211 hollow coffee twigs, 77.1 percent of which were occupied by one of 28 species of ants. Pseudomyrmex simplex was more abundant in lower elevations, whereas Pseudomyrmex ejectus dominated in high elevations. Species richness and the percent of occupied hollow twigs both peaked at mid‐elevations (800–1050 m). In sum, we found that species richness, abundance, and composition of arboreal twig‐nesting ants shift with elevation. These findings may provide important insights for understanding ant communities in coffee agroecosystems.     

Determining woodpecker diversity in the sub‐Himalayan forests of northern India using call playbacks

ABSTRACT Tropical forests have exceptional woodpecker diversity, but little is known about the abundance and diversity of woodpeckers in the Indian subcontinent, particularly for the Shorea robusta‐dominated moist deciduous forests of northern India. Our objective was to compare the number of woodpecker species and number of individuals detected using playback surveys and visual/aural transect surveys at five sites. Each site was surveyed 5–6 times along a 2000‐m transect, with woodpeckers detected using two methods: (1) visual and aural cues, and (2) playing back calls of 13 species at 400‐m intervals. Both methods involved similar effort per survey (100–110 min). During surveys, we detected 11 species of woodpeckers. More species and more than twice as many individuals were detected during playback surveys than during visual/aural surveys. In addition, species accumulation curves showed that we detected the species known to be present based on previous work faster with playback surveys than with visual/aural surveys at four of the five sites. During field trials, 97% of targeted individuals (N= 269) of 12 species responded to playback, and 83% of the responses occurred within 1 min of broadcast. The number of species of woodpeckers in our study area (11 species) was typical for a structurally diverse, tropical/subtropical moist broad‐leaved forest. Our results demonstrate that playback surveys are more efficient and accurate than visual/aural surveys, and that playback surveys can be useful for assessing and monitoring woodpecker diversity in tropical forests.     

Diversity patterns of vascular epiphytes along an elevational gradient in the Andes

Aim To document the elevational pattern of epiphyte species richness at the local scale in the tropical Andes with a consistent methodology. Location The northern Bolivian Andes at 350–4000 m above sea level. Methods We surveyed epiphytic vascular plant assemblages in humid forests in (a) single trees located in (b) 90 subplots of 400 m² each located in (c) 14 plots of 1 ha each. The plots were separated by 100–800 m along the elevational gradient. Results We recorded about 800 epiphyte species in total, with up to 83 species found on a single tree. Species richness peaked at c. 1500 m and declined by c. 65% to 350 m and by c. 99% to 4000 m, while forests on mountain ridges had richness values lowered by c. 30% relative to slope forests at the same elevations. The hump‐shaped richness pattern differed from a null‐model of random species distribution within a bounded domain (the mid‐domain effect) as well as from the pattern of mean annual precipitation by a shift of the diversity peak to lower elevations and by a more pronounced decline of species richness at higher elevations. With the exception of Araceae, which declined almost monotonically, all epiphyte taxa showed hump‐shaped curves, albeit with slightly differing shapes. Orchids and pteridophytes were the most species‐rich epiphytic taxa, but their relative contributions shifted with elevation from a predominance of orchids at low elevations to purely fern‐dominated epiphyte assemblages at 4000 m. Within the pteridophytes, the polygrammoid clade was conspicuously overrepresented in dry or cold environments. Orchids, various small groups (Cyclanthaceae, Ericaceae, Melastomataceae, etc.), and Bromeliaceae (below 1000 m) were mostly restricted to the forest canopy, while Araceae and Pteridophyta were well represented in the forest understorey. Main conclusions Our study confirms the hump‐shaped elevational pattern of vascular epiphyte richness, but the causes of this are still poorly understood. We hypothesize that the decline of richness at high elevations is a result of low temperatures, but the mechanism involved is unknown. The taxon‐specific patterns suggest that some taxa have a phylogenetically determined propensity for survival under extreme conditions (low temperatures, low humidity, and low light levels in the forest interior). The three spatial sampling scales show some different patterns, highlighting the influence of the sampling methodology.     

Diversity and biogeography of vascular epiphytes in Western Amazonia, Yasuní, Ecuador

Aim Although vascular epiphytes are important components of species richness and complexity of Neotropical forests, vascular epiphytes are under‐represented in large scale biogeographical analyses. We studied the diversity, biogeography and floristic relationships of the epiphytic flora of the Yasuní region (Western Amazonia) in a Neotropical context, with special emphasis on the influence of the Andean flora on floristic composition and diversity of surrounding lowland forests. Location Western Amazonian lowland rainforest, Tiputini Biodiversity Station (0°38′ S 76°09′ W, 230 m a.s.l., 650 ha), Yasuní National Park, Ecuador. Methods We compared the vascular epiphyte flora of Yasuní with 16 published Neotropical epiphyte inventories. Secondly, based on a floristic database with records of more than 70,000 specimens of vascular epiphytes from the Neotropics the elevational composition of eight selected inventories was analysed in detail. Results The vascular epiphyte flora of Yasuní is characterized by a very high species richness (313 spp.). A moderate portion of species is endemic to the Upper Napo region (c. 10%). However, this figure is much higher than previous analyses primarily based on woody species suggested. Geographical ranges of these species match with a proposed Pleistocene forest refuge. Compared with Northern and Central Amazonian sites, Western Amazonian epiphyte communities are characterized by a higher portion of montane and submontane species. Species richness of vascular epiphytes at the sites was correlated with the amount of rainfall, which is negatively correlated with the number of dry months. Main conclusion Recent and historic patterns of rainfall are the driving forces behind diversity and floristic composition of vascular epiphytes in Western Amazonia: high annual rainfall in combination with low seasonality provides suitable conditions to harbour high species richness. The proximity to the Andes, the most important centre of speciation for most Neotropical epiphytic taxa, in combination with the climatic setting has allowed a continuous supply of species richness to the region. At least for epiphytes, the borderline between the Andean and Amazonian flora is much hazier than previously thought. Moreover, the comparatively moist climate in Western Amazonia during the Pleistocene has probably led to fewer extinctions and/or more speciation than in more affected surrounding lowlands.     

Altitudinal biodiversity patterns of seed plants along Gongga Mountain in the southeastern Qinghai–Tibetan Plateau

The mechanisms underlying elevation patterns in species and phylogenetic diversity remain a central issue in ecology and are vital for effective biodiversity conservation in the mountains. Gongga Mountain, located in the southeastern Qinghai–Tibetan Plateau, represents one of the longest elevational gradients (ca. 6,500 m, from ca. 1,000 to 7,556 m) in the world for studying species diversity patterns. However, the elevational gradient and conservation of plant species diversity and phylogenetic diversity in this mountain remain poorly studied. Here, we compiled the elevational distributions of 2,667 native seed plant species occurring in Gongga Mountain, and estimated the species diversity, phylogenetic diversity, species density, and phylogenetic relatedness across ten elevation belts and five vegetation zones. The results indicated that species diversity and phylogenetic diversity of all seed plants showed a hump‐shaped pattern, peaking at 1,800–2,200 m. Species diversity was significantly correlated with phylogenetic diversity and species density. The floras in temperate coniferous broad‐leaved mixed forests, subalpine coniferous forests, and alpine shrublands and meadows were significantly phylogenetically clustered, whereas the floras in evergreen broad‐leaved forests had phylogenetically random structure. Both climate and human pressure had strong correlation with species diversity, phylogenetic diversity, and phylogenetic structure of seed plants. Our results suggest that the evergreen broad‐leaved forests and coniferous broad‐leaved mixed forests at low to mid elevations deserve more conservation efforts. This study improves our understanding on the elevational gradients of species and phylogenetic diversity and their determinants and provides support for improvement of seed plant conservation in Gongga Mountain.     

The vertical distribution of pronophiline butterflies (Nymphalidae, Satyrinae) along an elevational transect in Monte Zerpa (Cordillera de Mérida, Venezuela) with remarks on their diversity and parapatric distribution

Aim The distribution of neotropical butterflies of the tribe Pronophilini (Lepidoptera, Satyridae) was analysed with the aim of determining (i) the elevational ranges of distribution of each species (ii) the elevational gradient in diversity, and (iii) the existence of parapatric distributions of some closely related species pairs. Location The field study was carried out in the middle and upper section of the valley of the Río Albarregas, a tributary of the Río Chama on the southern slopes of the Serranía de la Culata, in the central part of the Cordillera de Mérida, Venezuela. Methods The material was collected along an elevational transect set on a trail leading from Merida to the Páramo de Los Conejos in the area known as Monte Zerpa. It consisted of a series of 32 collection sites set at every 25 m of altitude and covered an elevation from 2250 m to 3025 m. Results The ranges of Lymanopoda obsoleta, L. albocincta, L. zapatoza, Corades chelonis, C. pannonia, C. medeba, Mygona irmina, Pedaliodes epidipnis, P. japhleta, P. montagna and P. panyasis were all restricted to the lower half of the cloud forest zone and L. diezti, C. pax, P. polla, P. ornata and P. ferratilis were all restricted to the upper part. Two species, Erethris porphyria and Steroma bega, crossed the entire elevational range of the cloud forest zone.The biodiversity, as measured by Shannon’s index, attained a maximum at 2700 m, beyond which it gradually decreased. A parapatric distribution was confirmed for three pairs of closely related species: L. obsoleta and L. diezti, C. chelonis and C. pax and P. montagna and P. ferratilis. Main conclusions The peak in diversity at middle elevations seems to result from the overlapping ranges of species distributed over lower and upper parts of the cloud forest. Correlation of species composition at particular transect sites with elevation showed that increase in elevation was associated with an increase in species composition difference. Post‐mating isolation was suggested as a primary factor responsible for maintaining the parapatric distributions of species occupying the upper and lower zones and preventing their respective distributions from expanding.     

A simulation approach to determine statistical significance of species turnover peaks in a species-rich tropical cloud forest

Use of β-diversity indices in the study of spatial distribution of species diversity is hampered by the difficulty of applying significance tests. To overcome this problem we used a simulation approach in a study of species turnover of ferns, aroids, bromeliads, and melastomes along an elevational gradient from 1700 m to 3400 m in a species-rich tropical cloud forest of Bolivia. Three parameters of species turnover (number of upper/lower elevational species limits per elevational step, Wilson–Shmida similarity index between adjacent steps) were analysed. Significant species turnover limits were detected at 2000 (± 50) m and 3050 m, which roughly coincided with the elevational limits of the main vegetation types recognized in the study area. The taxon specificity of elevational distributions implies that no single plant group can be used as a reliable surrogate for overall plant diversity and that the response to future climate change will be taxon-specific, potentially leading to the formation of plant communities lacking modern analogues. Mean elevational range size of plant species was 490 m (± 369). Elevational range sizes of terrestrial species were shorter than those of epiphytes. We conclude that our simulation approach provides an alternative approach for assessing the statistical significance of levels of species turnover along ecological gradient without the limitations imposed by traditional statistical approaches.     

Elevational gradients in species richness and endemism of selected plant groups in the central Bolivian Andes

I analyzed the distribution of Acanthaceae, Araceae, Bromeliaceae, Cactaceae, Melastomataceae, and Pteridophyta in 62 vegetation plots of 400 m² along an elevational transect between 500 m and 2450 m, and at a nearby lowland site in western Santa Cruz department, Bolivia. These groups were selected because they are physiognomically distinctive, have high species numbers, are comparatively easy to identify, adequately reflect overall floristic relationships, include a wide range of life forms, and are small. The transect was located in the Tucumano-Boliviano biogeographic zone and included drought-deciduous (<850–1000 m), mixed evergreen (850–1000 m to 1800 m), and evergreen Podocarpus-dominated (>1800 m) forests. Elevational patterns of species richness were group-specific and probably related to the ecophysiological properties of each group. Species richness in Pteridophyta and Melastomataceae was correlated with moss cover (i.e., humidity), with elevation (i.e., temperatures) in Acanthaceae and epiphytic Bromeliaceae, with potential evapotranspiration (i.e., ecosystem productivity) in Araceae, and with light availability at ground level in terrestrial Bromeliaceae and Cactaceae. Community endemism generally increased with elevation, but showed a maximum at 1700 m for terrestrial Pteridophyta, and a nonsignificant decline for epiphytic Bromeliaceae and Cactaceae. Endemism was higher for terrestrial than for epiphytic taxa, and was lower among Pteridophyta compared to all other groups, reflecting different dispersal ability among taxonomic and ecological groups. Elevational zonation, tested against a null-model of random distribution of elevational limits, revealed a significant accumulation of upper and lower elevational range boundaries at 900–1050 m and at 1500–1850 m, corresponding to the elevational limits of the main physiognomic vegetation types.     

Elevational diversity patterns of small mammals on Mount Kinabalu, Sabah, Malaysia

1 Diversity patterns of small mammals were studied along an elevational transect on Mount Kinabalu, the highest mountain in South‐east Asia, utilizing data from previously existing sources and a new field study. A mark‐and‐release study (conducted during wet and dry seasons between November 1994 and April 1995) resulted in captures of 12 small mammal species, including two species of squirrels, two tree shrews, seven murid rodents and one gymnure. 2 Based on data compiled from this survey, museum specimens, and published and unpublished literature (analysed by locally weighted sums of squares and quadratic polynomial regressions), species richness of small mammals formed a middle elevation bulge, highest at about 1200–1400 m and declining at lower and higher elevations. Trapping during two seasons did not change the assessment of the pattern. 3 A cluster analysis of these data indicated that there are two elevationally associated faunas, one in the highlands and another in the lowlands. The transition between these two assemblages is at 1700–1800 m elevation. The lowland faunal assemblage has the highest number of species, with maximum species richness at about 1300 m for total small mammal species, about 1200 m for arboreal species and about 1400 m for terrestrial species. 4 The areas where much overlapping of species occurs are the elevations where climate and vegetation change rapidly from lowland to montane types. Tree species, gymnosperms, orchids and ferns showed a similar curvilinear pattern along the same elevational gradient, with maximum species richness at about 1400–1500 m. Temperature declined progressively with increasing elevation, but rainfall and humidity reached their highest levels at about 1700 m. 5 Maximum diversity of small mammals thus occurred at the elevation where a highland and a lowland assemblage overlapped, where several types of plants reached their maximum diversity, and where rainfall and humidity reached their maxima. Similar patterns have been documented for small mammals, plants, and climate at sites scattered in Indo‐Australia from Taiwan to New Guinea.     

Elevational distribution and zonation of tropical pteridophyte assemblages in Costa Rica

The use of species turnover rates to identify discontinuities in vegetation assemblages along environmental gradients is hampered by the difficulty of the application of significance tests. We used a simulation procedure to detect abrupt transitions in fern species assemblages along a tropical elevational gradient to detect elevational zonations between 100 and 3400 m in a species-rich tropical forest in Costa Rica. Significant species turnover values were only observed at 300, 1000, and 3200 m. These boundaries in vegetation assemblages were caused by different lifeforms within the pteridophyte species set: epiphytes attained high turnover values at low, terrestrials at high elevations. Accordingly, the mean elevational ranges of epiphytes and terrestrials differed clearly at high elevations, where those of terrestrial species showed a sharp decline above ca. 2500 m, but those of epiphytes kept more or less constant. The boundaries of significant range limit accumulations coincide with shifts in the environmental settings, namely humidity conditions and forest structure. The location of the observed boundaries in turn influences the elevational amplitudes of species restricted to the resulting elevational zones, determining the elevational distribution of range amplitudes. Between 1000 and 3200 m no significant boundaries were observed, although it has been argued that the lower cloud condensation layer and the frost line are the prime drivers of elevational zonations.