Montane Andean rain forests are a global diversity hotspot of geometrid moths

Aim Andean forests are known to be a major diversity hotspot for vascular plants and vertebrates, but virtually nothing is known about the diversity of arthropods. We examined whether montane rain forests in southern Ecuador are also a diversity hotspot for arthropods, and chose geometrid moths as a model group. Location The study area in southern Ecuador (Province Zamora‐Chinchipe, 79° W, 04° S) covers c. 40 km², with 39 collecting sites within an elevational range of 1040–2677 m a.s.l. Thirty‐five of the sites were situated in an area c. 2.5 km². Additional qualitative sampling was carried out in the same area and up to an elevation of 3100 m. Methods Nocturnal moths were collected quantitatively and qualitatively using portable light towers consisting of two 15 W fluorescent tubes, and diurnal moths were collected qualitatively using an insect net. Insects were sampled in six fieldwork periods in the years 1999–2003. As diversity measures, Fisher's alpha of the log‐series distribution as well as eight estimators of total species richness were applied. Results A total of 1266 species were recorded, 63% of which were identified to named species, whereas the remainder are likely to include many undescribed species. Quantitative samples at light towers collected 35,238 specimens representing 1223 species. The extrapolated species number for these data is 1420 (incidence coverage estimator). Twenty‐one additional nocturnal species and 22 exclusively diurnal species were sampled qualitatively at elevations between 1040 and 3100 m. The pooled value of Fisher's alpha for all quantitative samples is 246 ± 3. Main conclusions The diversity of Geometridae documented here is much higher than anywhere else in the world, even without the inclusion of additional species from adjacent lowland rain forests. The number of recorded species in this small area corresponds to more than 6% of the known world fauna of geometrid moths. Our study emphasizes the importance of protecting the remaining montane Andean rain forests. For setting priorities in conservation, more studies on insect diversity are urgently required in other regions of the Andes, since montane forests are being destroyed at an alarming rate.     

Phylogenetic diversity of geometrid moths decreases with elevation in the tropical Andes

Species diversity of geometrid moths (Lepidoptera, Geometridae) has previously been shown to be extremely and constantly high along a continuously forested elevational gradient in the Andes of southern Ecuador. We analysed samples taken from 32 sites between 1999 and 2011 in northern Podocarpus National Park and adjacent areas from 1020 to 2916 m a.s.l. We conjecture that high elevation habitats were historically mostly colonised by species from lower elevations, and that environmental filtering (e.g. through host plant specificity or temperature tolerance) constrained colonisation from lower elevations, which would yield a pattern of elevationally decreasing phylogenetic diversity. We analysed elevational phylogenetic patterns by means of: 1) the nearest‐taxon index (NTI), 2) DNA barcode‐based terminal branch lengths (TBLs) from maximum‐likelihood phylogeny, 3) the subfamily composition of the local assemblages, and 4), the rarefied number of morphologically defined genera per site. We counted a total of 1445 species. NTI values significantly increased with elevation, both in a conventional and a rarefaction approach. TBLs decreased significantly with elevation. Subfamily composition profoundly changed with elevation, particularly expressed as an increased proportion of the subfamily Larentiinae and decreased fractions of Sterrhinae and Geometrinae. The number of genera in equally rarefied species resamples significantly decreased with elevation. We conclude that environmental filtering indeed contributed to an altitudinal decrease in moth phylodiversity, but these constraints prevented only relatively few clades from colonising high elevation habitats.     

Determinants of diversity in afrotropical herbivorous insects (Lepidoptera: Geometridae): plant diversity,vegetation structure or abiotic factors?

Aim This study was conducted to investigate the potential of predicting alpha diversity and turnover rates of a highly diverse herbivorous insect family (Geometridae) based on vascular plant species richness and vegetation structure. Location The study was carried out on the south‐western slopes of Mount Kilimanjaro within a wide range of habitats between 1200 and 3150 m elevation. Methods The floristic and structural composition of the vegetation was recorded at 48 plots of 400 m². Geometrid moths were sampled manually at light sources located at the plot centres. Principal components analysis, redundancy analysis and multiple linear regression were used to explore how alpha diversity and species turnover of geometrid moths are related to vegetation structure and plant species richness. Results Alpha diversity of geometrid moths was significantly correlated with species diversity patterns in the most common vascular plant families (R² = 0.49) and with plant structural parameters (R² = 0.22), but not with overall floristic diversity. Species turnover of geometrid moths was strongly linked to diversity changes in a range of plant families (40% explained variance), less strongly to changes in vegetation physiognomy (25%), and only weakly to overall floristic diversity (5%). Changes in elevation were a better predictor of both alpha diversity and species turnover of geometrid moths than any principal component extracted from the vegetation data. Main conclusions Vegetation composition, diversity and structure all showed significant correlations with the diversity and species composition of geometrid moth assemblages. Nevertheless, in most cases relationships were indirect, via environmental parameters such as temperature and humidity, which influenced both vegetation and moth fauna. Possible direct links between geometrid diversity and potential food plants were much weaker. The lack of a significant correlation between overall plant species richness and geometrid diversity indicates that tropical geometrid moths may not be very selective in their food plant choice. Accordingly, a clear correlation between floral diversity and herbivore species richness must be regarded as overly simplistic, and the diversity of vascular plants cannot universally be used as a suitable biodiversity indicator for diverse insect taxa at higher trophic levels.     

Bergmann's rule does not apply to geometrid moths along an elevational gradient in an Andean montane rain forest

Aim Bergmann's rule generally predicts larger animal body sizes with colder climates. We tested whether Bergmann's rule at the interspecific level applies to moths (Lepidoptera: Geometridae) along an extended elevational gradient in the Ecuadorian Andes. Location Moths were sampled at 22 sites in the province Zamora‐Chinchipe in southern Ecuador in forest habitats ranging from 1040 m to 2677 m above sea level. Methods Wingspans of 2282 male geometrid moths representing 953 species were measured and analysed at the level of the family Geometridae, as well as for the subfamily Ennominae with the tribes Boarmiini and Ourapterygini, and the subfamily Larentiinae with the genera Eois, Eupithecia and Psaliodes. Results Bergmann's rule was not supported since the average wingspan of geometrid moths was negatively correlated with altitude (r = ?0.59, P < 0.005). The relationship between body size and altitude in Geometridae appears to be spurious because species of the subfamily Larentiinae are significantly smaller than species of the subfamily Ennominae and simultaneously increase in their proportion along the gradient. A significant decrease of wingspan was also found in the ennomine tribe Ourapterygini, but no consistent body size patterns were found in the other six taxa studied. In most taxa, body size variation increases with altitude, suggesting that factors acting to constrain body size might be weaker at high elevations. Main conclusions The results are in accordance with previous studies that could not detect consistent body size patterns in insects at the interspecific level along climatic gradients.     

Arctiid moth ensembles along a successional gradient in the Ecuadorian montane rain forest zone: how different are subfamilies and tribes?

Aim We examined changes in the species diversity and faunal composition of arctiid moths along a successional gradient at a fine spatial scale in one of the world's hot spots for moths, the Andean montane rain forest zone. We specifically aimed to discover whether moth groups with divergent life histories respond differentially to forest recovery. Location Southern Ecuador (province Zamora‐Chinchipe) along a gradient from early successional stages to mature forest understorey at elevations of 1800–2005 m a.s.l. Methods Moths were sampled with weak light traps at 21 sites representing three habitat categories (early and late succession, mature forest understorey), and were analysed at species level. Relative proportions were calculated from species numbers as well as from specimen numbers. Fisher's α was used as a measure of local diversity, and for ordination analyses non‐metric multidimensional scaling (NMDS) was carried out. Results Proportions of higher arctiid taxa changed distinctly along the successional gradient. Ctenuchini (wasp moths) contributed more strongly to ensembles in natural forest, whereas Lithosiinae (lichen moths) decreased numerically with forest recovery. Arctiid species diversity (measured as Fisher's α) was high in all habitats sampled. The three larger subordinated taxa contributed differentially to richness: Phaegopterini (tiger moths) were always the most diverse clade, followed by Ctenuchini and Lithosiinae. Local species diversity was higher in successional habitats than in forest understorey, and this was most pronounced for the Phaegopterini. Dominance of a few common species was higher, and the proportion of species represented as singletons was lower, than reported for many other tropical arthropod communities. NMDS revealed a significant segregation between ensembles from successional sites and from forest understorey for all larger subordinated taxa (Phaegopterini, Ctenuchini, Lithosiinae). Abandoned pastures held an impoverished, distinct fauna. Faunal segregation was more pronounced for rare species. Ordination axes reflected primarily the degree of habitat disturbance (openness of vegetation, distance of sites from mature forest) and, to a lesser extent, altitude, but not distance between sampling sites. Main conclusions Despite the geographical proximity of the 21 sites and the pronounced dispersal abilities of adult arctiid moths, local ecological processes were strong enough to allow differentiation between ensembles from mature forest and disturbed sites, even at the level of subfamilies and tribes. Differences in morphology and life‐history characteristics of higher arctiid taxa were reflected in their differential representation (proportions of species and individuals) at the sites, whereas patterns of alpha and beta diversity were concordant. However, concordance was too low to allow for reliable extrapolation, in terms of biodiversity indication, from one tribe or subfamily to the entire family Arctiidae. Phaegopterini (comprising more putative generalist feeders during the larval stages) benefited from habitat disturbance, whereas Ctenuchini (with host‐specialist larvae) were more strongly affiliated with forest habitats.     

The role of environment and mid-domain effect on moth species richness along a tropical elevational gradient

Aim The biodiversity of geometrid moths (Lepidoptera) along a complete tropical elevational gradient was studied for the first time. The patterns are described, and the role of geometric constraints and environmental factors is explored. Location The study was carried out along the Barva Transect (10° N, 84° W), a complete elevational gradient ranging from 40 to 2730 m a.s.l. in Braulio Carrillo National Park, Costa Rica, and adjacent areas. Methods Moths were sampled manually in 2003 and 2004 at 12 rain forest sites using light ‘towers’, each with two 15 W ultraviolet fluorescent tubes. We used abundance‐based rarefaction, statistical estimation of true richness (Chao 1), geographically interpolated observed richness and Fisher's alpha as measures of local diversity. Results A total of 13,765 specimens representing 739 species were analysed. All four measures showed a hump‐shaped pattern with maxima between 500 and 2100 m elevation. The two subfamilies showed richness and diversity maxima at either lower (Ennominae) or higher (Larentiinae) elevation than Geometridae as a whole. Among the four environmental factors tested, relative humidity yielded the highest correlation over the transect with the rarefaction‐based richness estimates as well as with estimated true species richness of Geometridae as a whole and of Larentiinae, while rainfall explained the greatest variation of Ennominae richness. The elevational pattern of moth richness was discordant with both temperature and with tree species richness. A combination of all environmental factors in a stepwise multiple regression produced high values of r² in Geometridae. The potential effects of geometric constraints (mid‐domain effect, MDE) were investigated by comparing them with observed, interpolated richness. Overall, models fitted very well for Geometridae as a whole and for Ennominae, but less well for Larentiinae. Small‐ranged species showed stronger deviations from model predictions than large‐ranged species, and differed strikingly between the two subfamilies, suggesting that environmental factors play a more pronounced role for small‐ranged species. We hypothesize that small‐ranged species (at least of the Ennominae) may tend to be host specialists, whereas large‐ranged species tend to be polyphagous. Based on interpolated ranges, mean elevational range for these moths was larger with increasing elevation, in accordance with Rapoport's elevational rule, although sampling effects may have exaggerated this pattern. The underlying mechanism remains unknown because Rapoport's ‘rescue’ hypothesis could not explain the observed pattern. Conclusions The results clearly show that moth diversity shows a hump‐shaped pattern. However, remarkable variation exists with regard to taxon and range size. Both environmental and geometric factors are likely to contribute to the observed patterns.     

Diverging diversity patterns of vascular plants and geometrid moths during forest regeneration on Mt Kilimanjaro, Tanzania

Aim This study investigates diversity patterns of vascular plants and plant‐feeding geometrid moths during montane rain forest regeneration in relation to the biogeographical and historical conditions of Mt Kilimanjaro. Location Investigations were undertaken on the south‐western slopes of Mt Kilimanjaro at altitudes between 2075 and 2265 m. Methods Thirteen plots were selected for this study. Four of these were situated in the middle of large clearings (> 1000 m²), three in secondary forest, two in mature forest remnants surrounded by secondary forest and four plots within continuous closed mature forest. Vascular plant species were recorded in an area of 20 × 20 m². Geometrid moths were attracted using lamps placed inside reflective gauze cylinders. Results Ninety‐three species of vascular plants were recorded on the plots. Plant diversity increased in the course of forest regeneration from clearings and secondary forest to mature forest remnants and mature forest. This increase was visible in all vegetation strata as well as in the species number of Dicotyledoneae. The diversity of geometrid moths conversely decreased from early to late successional stages. A total of 2276 Geometridae representing 114 morphospecies were included in the study. Local values of Fisher's α varied from 10.3 to 18.3 on clearings and in secondary forest, whereas they remained below 8.0 in mature forest and mature forest remnants. There was a significant negative correlation between the diversity of Geometridae and the number of dicots, and of plant species in the shrub layer. Main conclusions Contrary to an expected positive correlation between the diversity of vascular plants and herbivorous geometrid moths, diversity patterns of these two groups are strongly diverging due to biogeographical and ecological factors differently affecting the two groups. The increase in plant diversity can chiefly be explained with an increase in epiphyte diversity which is related to the occurrence of suitable habitats in extensive moss layers on huge Ocotea usambarensis (Engl.) trees in the mature forest. The low diversity of geometrid moths in these forests may be connected to the isolation and relatively young age of the montane rain forests on Mt Kilimanjaro. Hence only a small number of moth species adapted to the cool and perhumid conditions within moist mature forest have so far immigrated into these habitats, and time was insufficient for the evolution of many new species.     

Turning Up the Heat on a Hotspot: DNA Barcodes Reveal 80% More Species of Geometrid Moths along an Andean Elevational Gradient

We sampled 14,603 geometrid moths along a forested elevational gradient from 1020–3021 m in the southern Ecuadorian Andes, and then employed DNA barcoding to refine decisions on species boundaries initially made by morphology. We compared the results with those from an earlier study on the same but slightly shorter gradient that relied solely on morphological criteria to discriminate species. The present analysis revealed 1857 putative species, an 80% increase in species richness from the earlier study that detected only 1010 species. Measures of species richness and diversity that are less dependent on sample size were more than twice as high as in the earlier study, even when analysis was restricted to an identical elevational range. The estimated total number of geometrid species (new dataset) in the sampled area is 2350. Species richness at single sites was 32–43% higher, and the beta diversity component rose by 43–51%. These impacts of DNA barcoding on measures of richness reflect its capacity to reveal cryptic species that were overlooked in the first study. The overall results confirmed unique diversity patterns reported in the first investigation. Species diversity was uniformly high along the gradient, declining only slightly above 2800 m. Species turnover also showed little variation along the gradient, reinforcing the lack of evidence for discrete faunal zones. By confirming these major biodiversity patterns, the present study establishes that incomplete species delineation does not necessarily conceal trends of biodiversity along ecological gradients, but it impedes determination of the true magnitude of diversity and species turnover.     

Species richness and diversity across rocky intertidal elevation gradients in Helgoland: testing predictions from an environmental stress model

Environmental stress affects species richness and diversity in communities, but the precise form of the relationship is unclear. We tested an environmental stress model (ESM) that predicts a unimodal pattern for total richness and diversity in local communities across the full stress gradient where a regional biota can occur. In 2008, we measured richness and diversity (considering all macrobenthic species) across the entire intertidal range on three rocky shores on Helgoland Island, Germany. Intertidal elevation is known to be positively related to abiotic stress. Since Helgoland is between the northern and southern biogeographic boundaries for the cold-temperate NE Atlantic intertidal biota, it exhibits low stress levels for this biota at low elevations and high stress at high elevations because of long (>6 h) emersion times. Thus, we predicted a unimodal trend for richness and diversity across elevation. On all three shores, richness increased from high to middle elevations, but remained similar between middle and low elevations. Diversity followed the same trend on one shore and different trends (although also non-unimodal) on the other two. Evenness explained the trend differences between richness and diversity. Overall, our study yielded little support for the ESM. Reasons for richness and diversity not decreasing at low elevations may be related to influences of mostly subtidal species, Helgoland’s intertidal range, or sampling resolution. Our study also suggests that the ESM must be developed further to differentiate between richness and diversity. We offer recommendations to improve future ESM research using intertidal systems.     

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.     

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 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 variation in diversity and composition of land-snail faunas in a Tanzanian forest

We sampled terrestrial molluscs at fifteen elevations between 400 and 2000 m in Mwanihana Forest Reserve, Tanzania and recorded 84 taxa. Four diversity measures – species density (D_P), species richness (D₂₀) and observed (S_O) and interpolated (S_I) alpha diversity – were measured; beta diversity and abundance were also investigated. Mean elevational range was 470 m. D_P, D₂₀, alpha diversity and mollusc abundance increased with elevation, but alpha diversity peaked at 1695 m (S_O) or 1500 m (S_I) and declined at higher elevations. Maximum beta diversity was at 1000 m. Soil pH was negatively correlated with elevation and D_P. Cluster analysis divided the sites into lowland (400–900 m) and highland (>1000 m) groups. Axis 1 of a canonical correspondence analysis was associated with altitude and suggested a faunal discontinuity at 1000 m. Variation within the highland (>1000 m) and lowland faunal sets was related to elevation and forest structure respectively. The findings indicate that mollusc diversity peaks at intermediate elevations. This may be related to the combined effects of low rainfall conditions at low elevations and increasing effects of soil leaching at high elevations, both of which may limit mollusc diversity and abundance. Diversity at intermediate altitudes may be further elevated because of faunal mixing of lowland and highland groups.     

Faunal composition of geometrid moths changes with altitude in an Andean montane rain forest

Aim The objective of this study was to describe and interpret the changes in faunal composition in the moth family Geometridae (Lepidoptera) along a small‐scale elevational gradient in a tropical montane rain forest. This gradient was compared with a large‐scale latitudinal gradient in Europe. Location Investigations were carried out in the province Zamora‐Chinchipe in southern Ecuador along a gradient ranging from 1040 to 2677 m above sea level at twenty‐two sites. Methods Moths were sampled with light‐traps in three field periods in 1999 and 2000 and subsequently sorted and determined to species or morphospecies. Results We analysed 13,938 specimens representing 1010 species of geometrid moths. The proportional contribution of subtaxa to the local geometrid fauna changes along the elevational gradient at all systematic levels considered. While proportions of species of the subfamilies Ennominae, Sterrhinae and Geometrinae significantly decrease, the proportion of Larentiinae increases with increasing altitude. Changes also occur within the subfamilies Ennominae and Larentiinae. The host–plant specialist ennomine tribes Cassymini, Macariini and Palyadini completely vanish, and the proportion of the tribe Boarmiini decreases at high altitudes. In contrast, the remaining tribes (mostly comprising polyphagous species) either do not show proportional changes (Azelinini, Nacophorini, Nephodiini, Ourapterygini) or even increase (Caberini, ‘Cratoptera group’). Within Larentiinae, the species proportion of the genus Eois decreases, whereas concomitantly the proportion of Eupithecia increases. There is a remarkable similarity between the altitudinal patterns in Ecuador and those found along the latitudinal gradient in Europe. Main conclusions Species of the subfamily Larentiinae seem to be particularly well‐adapted to harsh environmental conditions, towards both high altitudes and latitudes. They might disproportionately profit from lower predation at higher altitudes. Many changes in the faunal composition can be explained by expected host–plant requirements of the species involved. Our results show that diversity estimates based on taxon ratios which are assumed to be constant must be regarded with caution because such ratios can change rapidly along environmental gradients.     

Abundance variations within feeding guilds reveal ecological mechanisms behind avian species richness pattern along the elevational gradient of Mount Cameroon

Two distinct diversity patterns are observed along tropical elevations: (a) decreasing number of species toward high elevations and (b) a hump-shaped pattern with the peak at mid-elevations. As diversity is likely supported by ecological capacity of the environment, decomposition of the overall richness into ecological facets and considering number of individuals within them is crucial for the proper understanding of richness patterns. We examined abundances of different avian guilds along the forested part of the elevational gradient on Mt. Cameroon. We (a) compared richness and abundance elevational patterns, (b) assessed the effective contribution of multiple guilds to richness and abundance patterns, and (c) assessed to what extent observed abundances of guilds differed from those expected by chance. We sampled birds in 2011–2015 during the dry season at seven elevations (30 m, 350 m, 650 m, 1100 m, 1500 m, 1850 m, 2200 m a.s.l.). For each assemblage, we estimated proportions of species and individuals that use particular diets, foraging modes, and feeding strata. We found that a rather decreasing pattern of species richness turns into a hump-shaped one if we look at the total abundances, implying different mechanisms behind these patterns. The number of species and individuals thus do not seem to be directly related, contrary to “the more-individuals hypothesis.” Abundances of foliage gleaners at mid-elevations, nectarivores at high elevations, and frugivores at low elevations deviated from random expectations. Our results imply that parts of ecological space are filled separately by bird species and individuals along elevation of Mt. Cameroon.     

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.     

Explaining the elevational diversity pattern of geometrid moths from Borneo: a test of five hypotheses

Aim The aim of this study was to document quantitatively a hump‐shaped elevational distribution of diversity for the geometrid moths of northern Borneo. Furthermore, we aimed to test rigorously a number of hypotheses that were proposed as ‘general explanations’ for such patterns, namely (1) the mid‐domain effect, (2) overlap between lowland and montane (or endemic) fauna, (3) water and energy availability (i.e. temperature‐limitation at high elevations, water‐limitation at low elevations), (4) area size of elevational bands, and (5) determination by (host) plant diversity. Location Northern Borneo, Malaysia. Methods We used quantitative light‐trapping data across northern Borneo (from our own field work and published literature) for 52 sites (covering > 800 species) to quantify local diversity (as Fisher’s α). We used these data to test predictions of the five hypotheses in spatially explicit correlation analyses, using published data on climate, plant diversity and topography (Digital Elevation Model) as predictor variables. Results We documented a pronounced diversity peak at c. 650 m a.s.l. (breakpoint regression, r² = 0.55). We observed additional effects of other habitat parameters, sampling methodology and time on diversity only for data below 655 m a.s.l. Furthermore, we did not find unequivocal support for any of the above‐stated hypotheses. The mid‐domain effect and faunal overlap can be ruled out as single explanations of observed patterns, whereas the data equally fit predictions from water–energy dynamics, area and plant diversity only at higher elevations, but not in the lower part of the gradient. Of those, only effects of energy (i.e. temperature) were expected a priori to be restricted to elevations above the breakpoint. Main conclusions We tentatively conclude that temperature (i.e. energy limitation) is the best explanation for decreasing diversity towards high elevations, but we did not find a compelling explanation of lowered diversity near sea level. We discuss and present some preliminary data on further possible explanations, i.e. niche conservatism of higher taxonomic groups.     

What drives elevational patterns of diversity? A test of geometric constraints,climate and species pool effects for pteridophytes on an elevational gradient in Costa Rica

Aim We studied pteridophyte species richness between 100 m and 3400 m along a Neotropical elevational gradient and tested competing hypotheses for patterns of species richness. Location Elevational transects were situated at Volcán Barva in the Braulio Carrillo National Park and La Selva Biological Station (100–2800 m) and Cerro de la Muerte (2700–3400 m), both on the Atlantic slope of Costa Rica, Central America. Method We analysed species richness on 156 plots of 20 × 20 m and measured temperature and humidity at four elevations (40, 650, 1800 and 2800 m). Species richness patterns were regressed against climatic variables (temperature, humidity, precipitation and actual evapotranspiration), regional species pool, area and predicted species number of a geometric null model (the mid‐domain effect, MDE). Results The species richness of the 484 recorded species showed a hump‐shaped pattern with elevation with a richness peak at mid‐elevations (c. 1700 m). The MDE was the single most powerful explanatory variable in linear regression models, but species richness was also associated strongly with climatic variables, especially humidity and temperature. Area and species pool were associated less strongly with observed richness patterns. Main conclusions Geometric models and climatic models exclusive of geometric constraints explained comparable amounts of the elevational variation in species richness. Discrimination between these two factor complexes is not possible based on model fits. While overall fits of geometric models were high, large‐ and small‐ranged species were explained by geometric models to different extents. Species with narrow elevational ranges clustered at both ends of the gradient to a greater extent than predicted by the MDE null models used here. While geometric models explained much of the pattern in species richness, we cannot rule out the role of climatic factors (or vice versa) because the predicted peak in richness from geometric models, the empirical peak in richness and the overlap in favourable environmental conditions all coincide at middle elevations. Mid‐elevations offer highest humidity and moderate temperatures, whereas at high elevations richness is reduced due to low temperatures, and at low elevations by reduced water availability due to high temperatures.     

Alpha diversity of communities and their variety along altitude gradient on northern slope of Changbai Mountain]

From 700 to 2600 m on northern slope of Changbai Mountain, twenty plots were investigated with an interval of 100 m in altitude, and alpha diversity of communities and their variety along altitude gradient were studied. The results showed that in different successional layers of trees, or shrub and herb layers, richness and diversity decreased linearly with the increase of altitude. Tree species decreased from 19 species at 700 m to only 1 species at timberline. Shrub and herb species decreased from 20 and 61 species at 700 m to 2 and 15 species at 2600 m respectively. The peak values of richness of trees and herbs along altitude gradient were appeared just in the ecotones between different types of vegetation, and it reflected the characteristics of high species richness in ecotones.     

新疆农田系统瓢虫资源调查与多样性研究

为全面了解新疆农田系统瓢虫资源组成以及多样性结构差异,本研究采用网捕法于2018年、2019年连续两年对新疆11个地州50个县/市/区15种农作物生境的406个采集点开展了瓢虫资源系统调查.结果表明:共获7001号瓢虫标本,隶属3亚科13属26种,其中多异瓢虫Hippodamia variegata为绝对优势种类,其个体数量占比达55.11％,深点食螨瓢虫Stethorus punctillum、龟纹瓢虫Propylea japonica等10种为常见种;不同生境瓢虫群落多样性分析显示,苜蓿田和麦田的丰富度指数(5.598150、7.546581)、多样性指数(0.874884、0.708508)、均匀度指数(0.661678、0.544575)都较高,但优势度指数则分别仅为0.224261、0.342884;大豆田和西瓜田的多样性指数、丰富度指数以及均匀度指数均显著低于其他生境类型(P<0.05),但优势度指数远高于其他生境,为0.663091、0.658502,可见苜蓿田、麦田等生境瓢虫物种数和个体数量丰富,且分布均匀,而大豆田和西瓜田种类相对贫乏,且分布不均匀;不同地州的瓢虫群落多样性分析显示,伊犁州的丰富度指数(7.363638)、多样性指数(0.970345)、均匀度指数(0.712583)都较高,但优势度指数仅为0.135821;阿克苏地区的多样性指数、丰富度指数以及均匀度指数都显著低于其他地州,但优势度指数远高于其他地州,为0.663477,可见伊犁州的瓢虫物种数和个体数量丰富,且分布均匀,而阿克苏地区种类相对贫乏,且分布不均匀;不同海拔带瓢虫群落多样性分析则显示,300 m～1300 m海拔区间农田瓢虫群落多样性指数和丰富度指数显著高于其它海拔区间,且低海拔(-130 m～300 m)或高海拔(1600 m～2800 m)区间的瓢虫群落丰富度和多样性指数较低,但优势度较高,这表明300 m～1300 m海拔区间内瓢虫种类比较丰富,而在低海拔或高海拔区间农田生境内的瓢虫物种贫乏,个体数量也比较少.本研究基本明确了新疆农田系统瓢虫资源组成及多样性,揭示了其生态系统和生物多样性保护中的重要价值,同时为生物防治提供了丰富捕食性天敌瓢虫资源.