The missing Madagascan mid-domain effect

Species richness varies enormously across geographical gradients, a well-known phenomenon for which there are many hypothesized explanations. One recent hypothesis uses null models to demonstrate that random re-distribution of species' ranges within a given domain leads to a 'mid-domain effect' (MDE): increasing species richness towards the centre of the area. Madagascar is especially well-suited for empirical evaluation of mid-domain models by virtue of its large endemic fauna and its clearly defined boundaries. Lees et al. [ Biol. J. Linn. Soc. 67 (1999) 529] observed patterns of species richness consistent with MDEs in the Madagascan rainforest (a slim, north–south belt). In this study, we test one-dimensional and two-dimensional mid-domain model predictions for the birds and mammals of the entire island of Madagascar. When only latitudinal extents of species' distribution are considered, patterns of richness in Madagascar show an MDE. However, this pattern disappears for both taxa after accounting for the tendency of latitudinal bands nearer the middle of the country to be larger. Two-dimensional mid-domain model predictions of species richness are qualitatively opposite to observed patterns. Instead, island-wide spatial gradients of species richness in Madagascar relate strongly to patterns of primary productivity and amount of remaining natural habitat. Earlier work that showed a mid-domain peak within the rainforest biome (effectively after controlling for climate and natural habitat) seems likely to have reflected methodological artefacts. The classic case in which MDEs should occur is, in fact, inconsistent with the mid-domain hypothesis.     

An elevational shift in butterfly species richness and composition accompanying recent climate change

The geographic ranges of many species have shifted polewards and uphill in elevation associated with climate warming, leading to increases in species richness at high latitudes and elevations. However, few studies have addressed community‐level responses to climate change across the entire elevational gradients of mountain ranges, or at warm lower latitudes where ecological diversity is expected to decline. Here, we show uphill shifts in butterfly species richness and composition in the Sierra de Guadarrama (central Spain) between 1967–1973 and 2004–2005. Butterfly communities with comparable species compositions shifted uphill by 293 m (± SE 26), consistent with an upward shift of approximately 225 m in mean annual isotherms. Species richness had a humped relationship with elevation, but declined between surveys, particularly at low elevations. Changes to species richness and composition primarily reflect the loss from lower elevations of species whose regional distributions are restricted to the mountains. The few colonizations by specialist low‐elevation species failed to compensate for the loss of high‐elevation species, because there are few low‐elevation species in the region and the habitat requirements of some of these prevent them from colonizing the mountain range. As a result, we estimated a net decline in species richness in approximately 90% of the region, and increasing community domination by widespread species. The results suggest that climate warming, combined with habitat loss and other drivers of biological change, could lead to significant losses in ecological diversity in mountains and other regions where species encounter their lower latitudinal‐range margins.     

Latitudinal gradients in the species richness of Australian termites (Isoptera)

Abstract Using data on the geographic range of 260 described species in the Atlas of Australian Termites, seven ‘regions’ with more complete data, across a wide range of latitudes were selected for further analysis. For these regions, mean species richness (± SE) was calculated for (i) all species from all families, (ii) Termitidae (197 spp.), (iii) Amitermes spp. (Termitidae, 58 spp.), (iv) all families excluding Amitermes spp. (139 spp.), (v) Termopsidae (5 spp.), (vi) Kalotermitidae (32 spp.) and (vii) Rhinotermitidae (25 spp.). In addition, we compared the Atlas data with species richness for five regions, across a comparable range of latitudes, based on the pooled species richness of described and un-described species given in community studies. No group of termites showed a consistent decline in species richness from tropical to temperate latitudes for either data set. The Atlas data showed similar total species richness from the tropics to the mediterranean southwest, before declining to lowest species richness at the highest latitudes. Species richness of Amitermes spp. and Rhinotermitidae was highest in the southwest. Termopsidae and Kalotermitidae showed no latitudinal pattern in species richness. Community studies showed highest and lowest total species richness in the southwest and at the highest latitudes (south-coastal Western Australia), respectively, and similar species richness from the tropics to arid central Australia. Species richness of. Amitermes spp. was highest in the southwest (31 spp.). Kalotermitidae and Rhinotermitidae showed no clear latitudinal pattern. The latitudinal patterns of species richness for the Australian termites is consistent with that for the Australian vertebrates and ants in that they differ from patterns established for these taxa on other continents.     

What drives the species richness patterns of non‐volant small mammals along a subtropical elevational gradient?

The biodiversity of non‐volant small mammals along an extensive subtropical elevational gradient was studied for the first time on Gongga Mountain, the highest mountain in Hengduan Mountain ranges in China, located in one of the 25 global biodiversity hotspots. Non‐volant small mammals were replicate sampled in two seasons at eight sampling sites between 1000 and 4200 m elevation on the eastern slope of Gongga Mountain. In all, 726 individual small mammals representing 25 species were documented in 28 800 trap nights. The species richness pattern for non‐volant small mammals along the elevational gradients was hump‐shaped with highest richness at mid‐elevations. However, different richness patterns emerged between endemic and non‐endemic species, between larger‐ranged and smaller‐ranged species and between rodents and insectivores. Temperature, precipitation, plant species richness and geometric constraints (mid‐ domain effect) were most significant in explaining species richness patterns. Based on the analysis of simple ordinary least squares (OLS) and stepwise multiple regressions, the overall richness pattern, as well as the pattern of insectivores, endemic species and larger‐ranged species showed strong correlation with geometric constraint predictions. However, non‐endemic species richness was more strongly correlated with temperature, while rodent richness was correlated with plant species richness. Our study shows that no single key factor can explain all richness patterns of non‐volant small mammals. We need to be cautious in summarizing a general richness pattern of large species groups (e.g. small mammals or mammals) from species in smaller groups having different ecological distributions and life histories. Elevational richness patterns and their driving factors for small mammals are more likely dependent on what kind of species we study.     

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.     

Testing,as opposed to supporting,the Mid‐domain Hypothesis: a response to Lees and Colwell (2007)

Both our analysis ( Kerr et al. 2006 ), and Lees and Colwell's (2007) reanalysis, of patterns of bird and mammal diversity on Madagascar show that the central peak of richness predicted by the Mid‐Domain Hypothesis (MDH) is not observed. Lees and Colwell emphasize an observation consistent with MDH predictions: a latitudinal mid‐domain richness peak in the rainforest biome. They find (but do not mention) that no analogous peak is observed in the other two main Madagascan biomes. MDH fails nearly all its tests in Madagascar.     

Can changes in ant diversity along elevational gradients in tropical and subtropical Australian rainforests be used to detect a signal of past lowland biotic attrition?

Increasing temperatures are predicted to have profound effects on montane ecosystems. In tropical forests, biotic attrition may reduce lowland diversity if losses of species due to upslope range shifts are not matched by influxes of warmer‐adapted species, either because there are none or their dispersal is impeded. Australian rainforests consist of a north–south chain of patches, broken by dry corridors that are barriers to the dispersal of rainforest species. These rainforests have repeatedly contracted and expanded during Quaternary glacial cycles. Many lowland rainforests are expansions since the Last Glacial Maximum and may, therefore, show a signal of historical biotic attrition. We surveyed ants from replicated sites along three rainforest elevational transects in eastern Australia spanning 200 to 1200 m a.s.l. and nearly 14° of latitude. We examined elevational patterns of ant diversity and if there was possible evidence of lowland biotic attrition. Each transect was in a different biogeographic region; the Australian Wet Tropics (16.3°S), the central Queensland coast (21.1°S) and subtropical south‐eastern Queensland (28.1°S). We calculated ant species density (mean species per site) and species richness (estimated number of species by incorporating site‐to‐site species turnover) within elevational bands. Ant species density showed no signal of lowland attrition and was high at low and mid‐elevations and declined only at high elevations at all transects. Similarly, estimated species richness showed no evidence of lowland attrition in the Wet Tropics and subtropical south‐east Queensland; species richness peaked at low elevations and declined monotonically with increasing elevation. Persistence of lowland rainforest refugia in the Wet Tropics during the Last Glacial Maximum and latitudinal range shifts of ants in subtropical rainforests during the Holocene climatic optimum may have counteracted lowland biotic attrition. In central Queensland, however, estimated richness was similar in the lowlands and mid‐elevations, and few ant species were indicative of lower elevations. This may reflect historical biotic attrition due perhaps to a lack of lowland glacial refugia and the isolation of this region by a dry forest barrier to the north.     

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.     

Assessing the vulnerability of species richness to anthropogenic climate change in a biodiversity hotspot

Aim To compare theoretical approaches towards estimating risks of plant species loss to anthropogenic climate change impacts in a biodiversity hotspot, and to develop a practical method to detect signs of climate change impacts on natural populations. Location The Fynbos biome of South Africa, within the Cape Floristic Kingdom. Methods Bioclimatic modelling was used to identify environmental limits for vegetation at both biome and species scale. For the biome as a whole, and for 330 species of the endemic family Proteaceae, tolerance limits were determined for five temperature and water availability‐related parameters assumed critical for plant survival. Climate scenarios for 2050 generated by the general circulation models HadCM2 and CSM were interpolated for the region. Geographic Information Systems‐based methods were used to map current and future modelled ranges of the biome and 330 selected species. In the biome‐based approach, predictions of biome areal loss were overlayed with species richness data for the family Proteaceae to estimate extinction risk. In the species‐based approach, predictions of range dislocation (no overlap between current range and future projected range) were used as an indicator of extinction risk. A method of identifying local populations imminently threatened by climate change‐induced mortality is also described. Results A loss of Fynbos biome area of between 51% and 65% is projected by 2050 (depending on the climate scenario used), and roughly 10% of the endemic Proteaceae have ranges restricted to the area lost. Species range projections suggest that a third could suffer complete range dislocation by 2050, and only 5% could retain more than two thirds of their range. Projected changes to individual species ranges could be sufficient to detect climate change impacts within ten years. Main conclusions The biome‐level approach appears to underestimate the risk of species diversity loss from climate change impacts in the Fynbos Biome because many narrow range endemics suffer range dislocation throughout the biome, and not only in areas identified as biome contractions. We suggest that targeted vulnerable species could be monitored both for early warning signs of climate change and as empirical tests of predictions.     

Species traits explain recent range shifts of Finnish butterflies

This study provides a novel systematic comparative analysis of the species characteristics affecting the range margin shifts in butterflies towards higher latitudes, while taking phylogenetic relatedness among species into account. We related observed changes in the northern range margins of 48 butterfly species in Finland between two time periods (1992–1996 and 2000–2004) to 11 species traits. Species with positive records in at least ten 10 km × 10 km grid squares (in the Finnish National Butterfly Recording Scheme, NAFI) in both periods were included in the study. When corrected for range size change, the 48 butterfly species had shifted their range margins northwards on average by 59.9 km between the study periods, with maximum shifts of over 300 km for three species. This rate of range shifts exceeds all previously reported records worldwide. Our findings may be explained by two factors: the study region is situated in higher latitudes than in most previous studies and it focuses on the period of most prominent warming during the last 10–15 years. Several species traits exhibited a significant univariate relationship with the range margin shift according to generalized estimation equations (GEE) taking into account the phylogenetic relatedness among species. Nonthreatened butterflies had on average expanded their ranges strongly northwards (84.5 km), whereas the distributions of threatened species were stationary (−2.1 km). Hierarchical partitioning (HP) analysis indicated that mobile butterflies living in forest edges and using woody plants as their larval hosts exhibited largest range shifts towards the north. Thus, habitat availability and dispersal capacity of butterfly species are likely to determine whether they will be successful in shifting their ranges in response to the warming climate.     

Assessing New Zealand fern diversity from spatial predictions of species assemblages

The utility of explicit spatial predictions for biodiversity assessment is investigated with New Zealand fern flora. Distributions of 43 species were modelled from climatic and landform variables and predicted across New Zealand using generalised additive models (GAM). An original package of functions called generalised regression analysis and spatial prediction (GRASP) was developed to perform the analyses. On average, for the 43 models, the contributions of environmental variables indicate that mean annual temperature is the most important factor at this broad regional scale. Both annual solar radiation and its seasonality had higher correlations than temperature seasonality. Measures of water availability such as ratio of rainfall to potential evapotranspiration, air saturation deficit and soil water deficit presented significant contributions. Lithology was a better predictor than slope and drainage. These results are similar to those obtained from analyses of the distributions of New Zealand tree species and are consistent with the hypothesis that both tree and fern diversity are highest on sites conducive to high productivity. In order to identify hotspots of fern diversity, spatial predictions of individual species were summed up. The resulting map gave a very similar result to the direct prediction of their corresponding richness (number of species by plot out of 43 spp.). As a consequence, and where individual species models were not all available, the number of species within different species assemblages was directly modelled. Predicted richness hotspots of total species (out of 122 spp.), selected species (out of 43 and 21 spp.) and common species (out of 23 spp.) present very similar spatial patterns and are highly correlated. Richness of uncommon species (out of 39 spp.) was also accurately predicted, but presented a different spatial pattern. The number of rare species (out of 60 spp.) was not correctly modelled. Even though the lack of data for rare species clearly limits the application of this approach, fern community composition of more common species can be partially reconstructed from individual species predictions. This case study offers therefore a consistent approach not only for biodiversity hotspots identification, but also for setting targets to biodiversity assessment and restoration programs.     

Geographical patterns in species richness of the benthic polychaetes in the continental shelf of the Gulf of California, Mexican Pacific

The present study is the first attempt to describe meso-scale patterns in the species richness of polychaetes along the Gulf of California, which stretches from about 23°N to 31°N. We examine herein the spatial changes in species distribution and explore the overlapping of species’ ranges towards the centre of the Gulf, to test whether the mid-domain effect (MDE) could explain an expected mid-domain peak in species richness. The faunal composition and the latitudinal range of 244 species of polychaetes recorded along the continental shelf of the Gulf of California were analysed in latitude bands of 1°. The species composition changes around the Gulf’s archipelago (~29°N), and the highest values of species richness are found at the 25° (197 species) and 26° (193 species) of latitude. Although the species richness pattern could be described by a parabolic shape, the regional trend was not strongly consistent with the peak of diversity at 27°N (176–191 species) predicted by the mid-domain effect: the random sorting of species’ ranges within spatial domain does not explain satisfactorily the geographical patterns of diversity. Nevertheless, a partial contribution of MDE to these natural patterns of diversity could be detected, and the increase in species richness towards middle latitudes was basically determined by species with distribution ranges larger than 6°. The low level of significance between the empirical species richness pattern and the mid-domain model prediction for polychaetes in the Gulf does not restrict their use as a model for exploring the randomness of the diversity patterns.     

Distribution and species richness of woody dryland legumes in Baja California,Mexico

Abstract. We analysed the biogeographic patterns of woody legumes in the Baja California peninsula, NW Mexico. From the specimen labels of eight herbaria, we digitized 4205 records from 78 species, and projected them onto a grid of 205 cartographic cells (20’ longitude × 15’ latitude). Most species followed distribution patterns that coincide with floristic subdivisions of the peninsula. Endemism is high, reaching 60–70% in the centre of the peninsula, where the driest deserts are found and where significant floristic changes took place during Pleistocene glacial events. The number of cartographic cells (i.e. their geographic ranges) were log‐normally distributed, as has been reported for many other taxa. Floristic richness was found to be clumped around some cells where the observed richness is significantly higher than could be expected from chance variation. We tested the hypothesis that these ‘hotspots’ could be attributable to great collection efforts or to large land surfaces, but we still found 16 cells where richness is significantly high once these two factors are accounted for. Species richness and micro‐endemism increase towards the south, conforming to Rapoport's rule that predicts that species ranges become smaller towards the equator while richness increases. The floristic hotspots for woody legumes in Baja California occur in the Cape Region and along the Sierra de la Giganta in the southern Gulf Coast, where 77% of the total peninsular legume flora can be found. These hotspots are mostly unprotected, and should be considered priority areas for future conservation efforts.     

Determinants of regional species richness: an empirical analysis of the number of hawkmoth species (Lepidoptera: Sphingidae) on the Malesian archipelago

Aims Major patterns and determinants of the species richness of Sphingidae in the Malesian archipelago were investigated, including a distinction of richness patterns between subfamilies and range‐size classes. Location Southeast Asia, Malesia. Methods Using a compilation of specimen‐label data bases, geographic information system (GIS)‐supported estimates of distributional ranges for all Sphingidae species of Southeast Asia were used to assess the species richness of islands. Range maps for all species and checklists for 114 islands can be found at http://www.sphingidae‐sea.biozentrum.uni‐wuerzburg.de . Potential determinants of the species richness of islands were tested with general linear models. Results The estimated species richness of islands in the region is determined by biogeographical association, seasonality, availability of rain forest and island size. Species–area relationships are linear on a semi‐logarithmic representation, but not on a double‐logarithmic scale. Species richness of all sphingid subfamilies is influenced by biogeography. The presence of large rain‐forest areas affects mainly Smerinthinae, whereas distance from continental Asia is conspicuously irrelevant for this group. Widespread rather than geographically restricted species shape the overall distribution patterns of species richness. The altitudinal range of islands does not significantly affect species‐richness patterns, but its potential effects on geographically restricted species are discussed. Main conclusions As well as being affected by climatic and vegetation parameters, sphingid species richness is strongly influenced by a historical, directional dispersal process from continental Southeast Asia to the Pacific islands. This process did not apply equally to species of different taxonomic groups or range sizes. Widespread species decline in species richness towards the south‐east, whereas geographically restricted species exhibit an inverse pattern of species richness, probably because speciation becomes more important in this group within the more isolated island groups.     

The mid‐domain effect matters: simulation analyses of range‐size distribution data from Mount Kinabalu,Borneo

Aim In simulation exercises, mid‐domain peaks in species richness arise as a result of the random placement of modelled species ranges within simulated geometric constraints. This has been called the mid‐domain effect (MDE). Where close correspondence is found between such simulations and empirical data, it is not possible to reject the hypothesis that empirical species richness patterns result from the MDE rather than being the outcome (wholly or largely) of other factors. To separate the influence of the MDE from other factors we therefore need to evaluate variables other than species richness. The distribution of range sizes gives different predictions between models including the MDE or not. Here, we produce predictions for species richness and distribution of range sizes from one model without the MDE and from two MDE models: a classical MDE model encompassing only species with their entire range within the domain (range‐restricted MDE), and a model encompassing all species with the theoretical midpoint within the domain (midpoint‐restricted MDE). These predictions are compared with observations from the elevational pattern of range‐size distributions and species richness of vascular plants. Location Mount Kinabalu, Borneo. Methods The data set analysed comprises more than 28,000 plant specimens with information on elevation. Species ranges are simulated with various assumptions for the three models, and the species simulated are subsequently subjected to a sampling that simulates the actual collection of species on Mount Kinabalu. The resulting pattern of species richness and species range‐size distributions are compared with the observed pattern. Results The comparison of simulated and observed patterns indicates that an underlying monotonically decreasing trend in species richness with elevation is essential to explain fully the observed pattern of richness and range size. When the underlying trend is accounted for, the MDE model that restricts the distributions of theoretical midpoints performs better than both the classical MDE model and the model that does not incorporate geometric constraints. Main conclusions Of the three models evaluated here, the midpoint‐restricted MDE model is found to be the best for explaining species richness and species range‐size distributions on Mount Kinabalu.     

Harvestman (Arachnida: Opiliones) species distribution along three Neotropical elevational gradients: an alternative rescue effect to explain Rapoport's rule?

Aim Relationships between elevation and litter‐dweller harvestman (Arachnida: Opiliones) species richness along three elevational gradients in the Brazilian Atlantic Forest were evaluated. Specifically, three candidate explanatory factors for the observed patterns were tested: (1) the mid‐domain effect, (2) the Rapoport effect, and (3) the influence of environmental variables on species density and specimen abundance. Location Cuscuzeiro, Corcovado and Capricórnio mountains, in Ubatuba (23°26′ S, 45°04′ W), a coastal municipality in São Paulo state, south‐eastern Brazil. Methods We recorded harvestman species and abundance through active sampling using 8 × 8‐m plots in both summer and winter. At each plot we measured the temperature, humidity and mean litter depth. Harvestman species richness per elevational band was the sum of all species recorded in each band, plus the species supposed to occur due to the interpolation of the upper and lower elevational records. Differences between observed and expected species richness per elevational band, based on the mid‐domain effect, were examined through a Monte Carlo simulation. The Rapoport effect was evaluated using both the midpoint method and a new procedure proposed here, the ‘specimen method’. We applied multiple regression analysis to evaluate the contribution of each environmental variable (elevation, temperature, humidity and litter depth) on species density and specimen abundance per plot. Results Harvestman abundance and species richness decreased at higher elevations in the three mountains. The decrease in species richness was not monotonic and showed a plateau of high species richness at lower elevations. The number of harvestman species per elevational band does not fit that predicted by the mid‐domain effect based solely on geometric constraints assuming hard boundaries. Species with their midpoints at higher elevations tended to cover broader elevational range sizes. Both the midpoint method and the specimen method detected evidence of the Rapoport effect in the data. At fine spatial scales, temperature and humidity had positive effects on species density and specimen abundance, while mean litter depth had no clear effect. These relationships, however, were not constant between seasons. Main conclusions Our results suggest that harvestman species density declines at higher elevations due to restrictions imposed by temperature and humidity. We found a pattern in species range distribution as predicted by the elevational Rapoport effect. However, the usual rescue effect proposed to explain the Rapoport effect does not apply in our study. Since the majority of harvestman species covering broader elevational ranges do not exhibit reduced abundance at low elevations, an alternative rescue effect is proposed here. According to this alternative rescue effect, the decrease in species richness at higher elevations occurs due to differential upper limits of species with source populations below mid‐elevations. The seasonal differences in the relationships between environmental variables and species richness/specimen abundance per plot is an indication that species occurrence on elevational gradients is seasonally dependent. Thus relationships and hypotheses based on data recorded over short time periods, or in a single season, should be viewed cautiously.     

Beyond Rapoport's rule: evaluating range size patterns of New World birds in a two-dimensional framework

Aim To evaluate Rapoport's rule for New World birds in two‐dimensional geographical space. We specifically test for a topography × climate interaction that predicts little difference in range sizes between lowlands and mountains in cold climates, whereas in the tropics, montane species have narrow ranges and lowland species have broad ranges. Location The western hemisphere. Methods We used digitized range maps of breeding birds to generate mean range sizes in grids of 27.5 × 27.5 km and 110 × 110 km across North and South America. We examined the geographical pattern with respect to range in elevation, mean temperature in the coldest month, their interaction, biome size and continental width, using model II analysis of variance, multiple regression and simple correlation. Results In northern latitudes species have broad ranges in both mountainous and flat areas. However, range sizes in the mountains and lowlands diverge southwards, with the most extreme differences in the tropics. Further, there are minimal differences in range sizes across latitudes in lowlands. The smallest mean ranges occur in the tropical Andes. Mean range sizes in north‐central Canada, Central America and Argentina/Chile are also small, reflecting the narrowing of the continents in these areas. The best regression model explained 51% of the variation in mean range size. Main conclusions The two‐dimensional range size pattern indicates that neither winter temperature nor annual variability in temperature strongly influences the distribution of range sizes directly; rather, climate influences bird range sizes indirectly via effects on habitat size. Also, macroclimate interacts with topographic relief across latitudes, generating sharp mesoscale habitat gradients in tropical mountains but not in high latitude mountains or in lowlands at any latitude. Birds respond to these habitat gradients, resulting in ‘latitudinal’ range size gradients in topographically complex landscapes but not in simple landscapes.     

Biogeographic anomalies in the species richness of Chilean forests: Incorporating evolution into a climatic – historic scenario

Broad‐scale richness gradients are closely associated with temperature and water availability. However, historical and evolutionary processes have also contributed to shape current diversity patterns. In this paper we focus on the potential influences of Pleistocene glaciation and phylogenetic niche conservatism (the tendency for traits to be maintained during diversification) on the tree diversity gradient in Chile, and we quantify its primary climatic correlates. Tree species richness is greatest at mid latitudes, particularly in the Andes and Coastal ranges, and decreases abruptly to the south and north. Regression tree analysis identified annual precipitation and annual temperature as the primary probable drivers of this gradient. Ice cover during the Last Glacial Maximum was also identified as an ‘important’ variable, but the contemporary and historical predictors are strongly collinear. Geographically weighted regression indicated that the relationships between richness and environmental variables vary regionally: the relationship between tree richness and precipitation is stronger in north‐central Chile, whereas tree richness and temperature are most strongly associated in south‐central Chile. By assigning each species the age of the family to which it belongs and averaging all species in each geographical unit, we also found that species from the oldest families are distributed mainly in mid to high latitudes and species from younger families are distributed mainly at lower latitudes. This pattern is closely associated with annual precipitation. Thus, the ecological component of tree richness follows contemporary climatic gradients of both energy and water, but the aridification of the Atacama Desert was an important driver over evolutionary time. The influence of recent Pleistocene glaciation remains unresolved but it cannot be discounted.     

欧亚大陆东部毛茛科植物多样性格局及主导因子

毛茛科是真双子叶植物的基部类群之一, 包含多种药用植物, 具有较高的保护价值, 但关于毛茛科物种多样性和谱系多样性大尺度格局及其影响因子的研究还比较匮乏, 特别是以较高分辨率分布数据为基础的物种多样性格局研究尚未见报道。本文旨在: (1)建立欧亚大陆东部毛茛科植物分布数据库, 估算不同生活型物种多样性和谱系多样性格局, 并探究格局的形成机制。(2)分析毛茛科物种多样性和谱系多样性的相关关系, 确定多样性热点地区, 为毛茛科保护规划提供依据。根据中国、哈萨克斯坦、吉尔吉斯斯坦、塔吉克斯坦、土库曼斯坦、乌兹别克斯坦、蒙古和俄罗斯等国家的区域和地方植物志, 建立了“欧亚大陆东部地区毛茛科物种分布数据库”。该数据库包含了欧亚大陆东部地区1,688种毛茛科物种的分布数据, 空间分辨率为100 km × 100 km。在此基础上, 估算了毛茛科全部及不同生活型的物种多样性和谱系多样性格局, 并利用广义线性模型和等级方差分离方法分析了毛茛科物种和谱系多样性格局与环境因子的关系。最后比较了物种多样性和谱系多样性的相关关系, 确定了毛茛科的古热点地区。结果显示: (1)欧亚大陆东部毛茛科植物物种和谱系多样性均呈明显的纬度格局, 且在山区具有较高的多样性。(2)毛茛科植物物种和谱系多样性受现代气候、地形异质性和末次冰期以来的气候变化的共同影响, 但不同影响因子的相对贡献率在物种和谱系多样性及不同生活型之间差异显著。(3)中高纬度地区的谱系多样性高于给定物种数的预期, 是毛茛科的古热点地区, 在毛茛科保护规划中应受到重视。     

The determinants of land snail diversity along a tropical elevational gradient: insularity,geometry and niches

Aim We investigated the patterns of species richness in land snails and slugs along a tropical elevational gradient and whether these patterns correlate with area, elevation, geographic constraints, and productivity. We did so both at the scale at which land snail population processes take place and at the coarser scale of elevational zones. Location Mount Kinabalu (4096 m) and the adjacent Mount Tambuyukon (2588 m) in Kinabalu Park, Sabah, Malaysian Borneo. Methods We used an effort‐controlled sampling protocol to determine land snail and slug species richness in 142 plots of 0.04 ha at elevations ranging from 570 to 4096 m. Extents of elevational ranges were determined by interpolation, extended where appropriate at the lower end with data from lowlands outside the study area. We used regression analysis to study the relationships between species density and richness on the one hand and elevation and area on the other. This was done for point data as well as for data combined into 300‐m elevational intervals. Results Species density (based on the individual samples) showed a decline with elevation. Elevational range length profiles revealed that range lengths are reduced at greater elevations and that a Rapoport effect is absent. Diversity showed a mild mid‐domain effect on Kinabalu, but not on Tambuyukon. When the data were combined into 300‐m elevational intervals, richness correlated more strongly with elevation than with area. Ecomorphospace was seen to shrink with increasing elevation. Main conclusions The elevational species richness patterns show the combined effects of (1) reduced niche diversity at elevations with lower productivity and (2) historical events in which the upward migration of lowland species as well as the speciation of highland endemics took place.