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.     

Support for the elevational Rapoport's rule among seed plants in Nepal depends on biogeographical affinities and boundary effects

As one of the most important hypotheses on biogeographical distribution, Rapoport's rule has attracted attention around the world. However, it is unclear whether the applicability of the elevational Rapoport's Rule differs between organisms from different biogeographical regions. We used Stevens’ method, which uses species diversity and the averaged range sizes of all species within each (100 m) elevational band to explore diversity‐elevation, range‐elevation, and diversity‐range relationships. We compared support for the elevational Rapoport's rule between tropical and temperate species of seed plants in Nepal. Neither tropical nor temperate species supported the predictions of the elevational Rapoport's rule along the elevation gradient of 100–6,000 m a.s.l. for any of the studied relationships. However, along the smaller 1,000–5,000 m a.s.l. gradient (4,300 m a.s.l. for range‐elevation relationships) which is thought to be less influenced by boundary effects, we observed consistent support for the rule by tropical species, although temperate species did not show consistent support. The degree of support for the elevational Rapoport's rule may not only be influenced by hard boundary effects, but also by the biogeographical affinities of the focal taxa. With ongoing global warming and increasing variability of temperature in high‐elevation regions, tropical taxa may shift upward into higher elevations and expand their elevational ranges, causing the loss of temperate taxa diversity. Relevant studies on the elevational Rapoport's rule with regard to biogeographical affinities may be a promising avenue to further our understanding of this rule.     

Contrasting patterns of intraspecific trait variability in native and non-native plant species along an elevational gradient on Tenerife,Canary Islands

Background and AimsNon-native plant species are not restricted to lowlands, but increasingly are invading high elevations. While for both native and non-native species we expected variability of plant functional traits due to the changing environmental conditions along elevational gradients, we additionally assumed that non-native species are characterized by a more acquisitive growth strategy, as traits reflecting such a strategy have been found to correlate with invasion success. Furthermore, the typical lowland introduction of non-native species coming from multiple origins should lead to higher trait variability within populations of non-native species specifically at low elevations, and they might therefore occupy a larger total trait space.MethodsAlong an elevational gradient ranging from 55 to 1925 m a.s.l. on Tenerife, we collected leaves from eight replicate individuals in eight evenly distributed populations of five native and six non-native forb species. In each population, we measured ten eco-morphological and leaf biochemical traits and calculated trait variability within each population and the total trait space occupied by native and non-native species.Key ResultsWe found both positive (e.g. leaf dry matter content) and negative (e.g. leaf N) correlations with elevation for native species, but only few responses for non-native species. For non-native species, within-population variability of leaf dry matter content and specific leaf area decreased with elevation, but increased for native species. The total trait space occupied by all non-native species was smaller than and a subset of that of native species.ConclusionsWe found little evidence that intraspecific trait variability is associated with the success of non-native species to spread towards higher elevations. Instead, for non-native species, our results indicate that intermediate trait values that meet the requirements of various conditions are favourable across the changing environmental conditions along elevational gradients. As a consequence, this might prevent non-native species from overcoming abruptly changing environmental conditions, such as when crossing the treeline.     

Home Range Use and Movement Patterns of Non-Native Feral Goats in a Tropical Island Montane Dry Landscape

Advances in wildlife telemetry and remote sensing technology facilitate studies of broad-scale movements of ungulates in relation to phenological shifts in vegetation. In tropical island dry landscapes, home range use and movements of non-native feral goats (Capra hircus) are largely unknown, yet this information is important to help guide the conservation and restoration of some of the world’s most critically endangered ecosystems. We hypothesized that feral goats would respond to resource pulses in vegetation by traveling to areas of recent green-up. To address this hypothesis, we fitted six male and seven female feral goats with Global Positioning System (GPS) collars equipped with an Argos satellite upload link to examine goat movements in relation to the plant phenology using the Normalized Difference Vegetation Index (NDVI). Movement patterns of 50% of males and 40% of females suggested conditional movement between non-overlapping home ranges throughout the year. A shift in NDVI values corresponded with movement between primary and secondary ranges of goats that exhibited long-distance movement, suggesting that vegetation phenology as captured by NDVI is a good indicator of the habitat and movement patterns of feral goats in tropical island dry landscapes. In the context of conservation and restoration of tropical island landscapes, the results of our study identify how non-native feral goats use resources across a broad landscape to sustain their populations and facilitate invasion of native plant communities.     

Growth of the tropical Pinus kesiya as influenced by climate and nutrient availability along an elevational gradient

与温带地区相比,人们对在热带山脉中,尤其是热带针叶林中,沿海拔梯度的个体树种生长减少的原因知之甚少。我们的研究目的是检验气候或土壤条件是否会导致在高海拔分布的卡西亚松(Pinus kesiya,一种在南亚和东南亚广泛分布的松树)的生长减少。我们对越南中南部山脉中沿海拔梯度约在900~2000米之间的卡西亚松的树干直径增量和其年轮中13C(△13C)的同位素判别进行分析。同时确定其生长与△13C、气候和土壤变量的关系。研究发现,树木的断面积生长量(BAI)与温度或降水之间没有一致的相关关系。相反,沿海拔梯度,我们发现BAI与△13C以及与上层矿质土壤的C/N比和δ15N特征呈显著负相关。BAI与土壤中有效磷(Pa)和“碱性”阳离子(钙、镁、钾)的浓度呈正相关。我们的结论是,较高海拔下的较低温度会对树木生长产生间接影响,主要通过产生较高的C/N比并降低氮(N)和P矿化率而产生作用,而这些影响因其他负面反馈而进一步受到阻碍。主要包括：降水增加的淋溶作用而导致较低浓度的“碱性”阳离子以及较高海拔土壤矿化的N和P的有效性降低。本研究结果可以进一步解释卡西亚松的最高生长极限以及该物种在其他山区出现的情况     

The inability of tropical cloud forest species to invade grasslands above treeline during climate change: potential explanations and consequences

The upper elevational range edges of most tropical cloud forest tree species and hence the ‘treeline’ are thought to be determined primarily by temperatures. For this reason, the treeline ecotone between cloud forests and the overlying grasslands is generally predicted to shift upslope as species migrate to higher elevations in response to global warming. Here, we propose that other factors are preventing tropical trees from shifting or expanding their ranges to include high elevation areas currently under grassland, resulting in stationary treelines despite rising mean temperatures. The inability of cloud forest species to invade the grasslands, a phenomenon which we refer to as the ‘grass ceiling’ effect, poses a major threat to tropical biodiversity as it will greatly increase risk of extinctions and biotic attrition in diverse tropical cloud forests. In this review, we discuss some of the natural factors, as well as anthropogenic influences, that may prevent cloud forest tree species from expanding their ranges to higher elevations. In the absence of human disturbances, tropical treelines have historically shifted up‐ and down‐slope with changes in temperature. Over time, increased human activity has limited forests to lower elevations (i.e. has depressed treelines), and often broken the equilibrium between species range limits and climate. Yet even in areas where anthropogenic influences are halted, cloud forests have not expanded to higher elevations. Despite the critical importance of understanding the distributional responses of tropical species to climate change, few studies have addressed the factors that influence treeline location and dynamics, severely hindering our ability to predict the fate of these diverse and important ecosystems.     

Influence of past land use and current human disturbance on non-native plant species on small Italian islands

Biological invasions are regarded as one of the main drivers of habitat degradation in island ecosystems. Mediterranean islands have been subjected to a high degree of land conversion over the past 60 years, resulting in a massive reduction in the amount of rural land and the sprawl of tourist activities. The aims of this paper are to evaluate the current level of invasion of alien plant species in semi-natural vegetation types that have developed after the abandonment of agriculture and to analyze the relationships between non-native species, native flora, and environmental characteristics. Two Italian islands (Ponza and Ventotene) were surveyed using a random-stratified sampling. The occurrence and relative cover of alien plant species were compared and separate analyses were performed for the native flora. Abundance patterns of both native and alien species were then studied in the light of the environmental and anthropogenic features. Although we found that some non-native species are extremely widespread, their relative cover at the plot level is low. Permutational Multivariate Analysis of Variance and Indicator Species Analysis revealed dissimilarities in the native species composition, while Mann–Whitney and Kruskal–Wallis tests showed differences in the ecological requirements (moisture, soil reaction, and nitrogen) of the native species pool. Canonical Correspondence Analysis pointed to the importance of the proximity to agricultural areas, human disturbance, and past land management, particularly residual terraces, in determining the difference between plant communities on the two islands. The results of our study suggest that traditional forms of agriculture may represent a key element for countering the establishment and spread of non-native plants in Mediterranean areas.     

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.     

Natives and non‐natives plants show different responses to elevation and disturbance on the tropical high Andes of Ecuador

The aim was to assess patterns of plant diversity in response to elevation and disturbance in a tropical mountain. The study area was located in north‐central portion of the Eastern Cordillera of the Ecuadorian Andes, on a road from 1,150 m a.s.l. (Osayacu) to 4,000 (Papallacta). Along a mountain road spanning a wide altitudinal gradient, at 20 elevations we sampled three plots: one at the roadside and two perpendicular to the roadside. The relationship between elevation and species richness was assessed using linear and quadratic regressions, the effect of disturbance on species richness was determined by ANCOVA and a t test with parameters obtained from quadratic equations. Similarity of species composition among the roadside and sites distant was evaluated with the Chao‐Jaccard and classic Jaccard similarity indices, the distribution of non‐native species according to their origin were analyzed with linear and quadratic regression. The native species showed a linearly monotonic decrease with elevation, whereas non‐natives showed a quadratic distribution. Disturbed areas had the greatest number of non‐native species and lower native species richness, showing also a high floristic similarity; less disturbed areas showed the opposite. The non‐native species of temperate origin were more numerous and showed unimodal elevational distribution, while species of tropical origin were few and decreased linearly with elevation. We conclude that in a tropical highland mountain range, native and non‐native plant species respond differently to elevation: native species exhibit a monotonically linear decrease, and non‐native species show a unimodal trend. Disturbance positively affects non‐native species showing higher richness and fewer species turnover. In addition, the non‐native species are located along of the elevational gradient in relation to their biogeographic origin.     

Elevational Ranges of Montane Birds and Deforestation in the Western Andes of Colombia

Deforestation causes habitat loss, fragmentation, degradation, and can ultimately cause extinction of the remnant species. Tropical montane birds face these threats with the added natural vulnerability of narrower elevational ranges and higher specialization than lowland species. Recent studies assess the impact of present and future global climate change on species’ ranges, but only a few of these evaluate the potentially confounding effect of lowland deforestation on species elevational distributions. In the Western Andes of Colombia, an important biodiversity hotspot, we evaluated the effects of deforestation on the elevational ranges of montane birds along altitudinal transects. Using point counts and mist-nets, we surveyed six altitudinal transects spanning 2200 to 2800m. Three transects were forested from 2200 to 2800m, and three were partially deforested with forest cover only above 2400m. We compared abundance-weighted mean elevation, minimum elevation, and elevational range width. In addition to analysing the effect of deforestation on 134 species, we tested its impact within trophic guilds and habitat preference groups. Abundance-weighted mean and minimum elevations were not significantly different between forested and partially deforested transects. Range width was marginally different: as expected, ranges were larger in forested transects. Species in different trophic guilds and habitat preference categories showed different trends. These results suggest that deforestation may affect species’ elevational ranges, even within the forest that remains. Climate change will likely exacerbate harmful impacts of deforestation on species’ elevational distributions. Future conservation strategies need to account for this by protecting connected forest tracts across a wide range of elevations.     

Latitudinal shifts of introduced species: possible causes and implications

This study aims to document shifts in the latitudinal distributions of non-native species relative to their own native distributions and to discuss possible causes and implications of these shifts. We used published and newly compiled data on inter-continentally introduced birds, mammals and plants. We found strong correlations between the latitudinal distributions occupied by species in their native and exotic ranges. However, relatively more non-native species occur at latitudes higher than those in their native ranges, and fewer occur at latitudes lower than those in their native ranges. Only a small fraction of species examined (i.e. <20% of animals and <10% of plants) have expanded their distributions in their exotic range beyond both high- and low-limits of their native latitudes. Birds, mammals and plants tended to shift their exotic ranges in similar ways. In addition, most non-native species (65–85% in different groups) have not reached the distributional extent observed in their native ranges. The possible drivers of latitudinal shifts in the exotic range may include climate change, greater biotic resistance at lower latitudes, historical limitations on ranges in native regions, and the impacts of humans on species distributions. The relatively restricted distribution of most species in their exotic range highlights the great potential of future spread of most introduced species and calls for closely monitoring their directional spread under climate change.     

Processes at multiple scales affect richness and similarity of non‐native plant species in mountains around the world

Aim To investigate how species richness and similarity of non‐native plants varies along gradients of elevation and human disturbance. Location Eight mountain regions on four continents and two oceanic islands. Methods We compared the distribution of non‐native plant species along roads in eight mountainous regions. Within each region, abundance of plant species was recorded at 41–84 sites along elevational gradients using 100‐m² plots located 0, 25 and 75 m from roadsides. We used mixed‐effects models to examine how local variation in species richness and similarity were affected by processes at three scales: among regions (global), along elevational gradients (regional) and with distance from the road (local). We used model selection and information criteria to choose best‐fit models of species richness along elevational gradients. We performed a hierarchical clustering of similarity to investigate human‐related factors and environmental filtering as potential drivers at the global scale. Results Species richness and similarity of non‐native plant species along elevational gradients were strongly influenced by factors operating at scales ranging from 100 m to 1000s of km. Non‐native species richness was highest in the New World regions, reflecting the effects of colonization from Europe. Similarity among regions was low and due mainly to certain Eurasian species, mostly native to temperate Europe, occurring in all New World regions. Elevation and distance from the road explained little of the variation in similarity. The elevational distribution of non‐native species richness varied, but was always greatest in the lower third of the range. In all regions, non‐native species richness declined away from roadsides. In three regions, this decline was steeper at higher elevations, and there was an interaction between distance and elevation. Main conclusions Because non‐native plant species are affected by processes operating at global, regional and local scales, a multi‐scale perspective is needed to understand their patterns of distribution. The processes involved include global dispersal, filtering along elevational gradients and differential establishment with distance from roadsides.     

Geographic variation in field body temperature of sceloporus lizards

1. Using data from the literature, I assessed how broad climatic patterns affected field body temperatures (T_b’s) of lizards in the genus Sceloporus.

2. Sceloporus at temperate latitudes had mean T_b’s of 35°C throughout their elevational range. This pattern is associated with “tropical” temperatures that extend into high north latitudes during the summer and the relatively low elevations occupied by the lizards.

3. At tropical latitudes, mean T_b declined from 35°C at low elevations to 31°C at high elevations. This pattern is associated with low seasonal variation in temperature at tropical latitudes and the relatively high elevations occupied by the lizards.     

Niche shift in four non‐native estrildid finches and implications for species distribution models

Non‐native species can have severe impacts on ecosystems. Therefore, predictions of potentially suitable areas that are at risk of the establishment of non‐native populations are desirable. In recent years, species distribution models (SDMs) have been widely applied for this purpose. However, the appropriate selection of species records, whether from the native area alone or also from the introduced range, is still a matter of debate. We combined analyses of native and non‐native realized climate niches to understand differences between models based on all locations, as well as on locations from the native range only. Our approach was applied to four estrildid finch species that have been introduced to many regions around the world. Our results showed that SDMs based on location data from native areas alone may underestimate the potential distribution of a given species. The climatic niches of species in their native ranges differed from those of their non‐native ranges. Niche comparisons resulted in low overlap values, indicating considerable niche shifts, at least in the realized niches of these species. All four species have high potential to spread over many tropical and subtropical areas. However, transferring these results to temperate areas has a high degree of uncertainty, and we urge caution when assessing the potential spread of tropical species that have been introduced to higher latitudes.     

The biogeography of land snails in the islands of the Gulf of Guinea

Data on land snail diversity in the Gulf of Guinea islands is presented and the biogeography of the Gulf of Guinea fauna is discussed with reference to island snail faunas in the other areas of the world. Although the land snail faunas of the four islands clearly have west and central African affinities, speciation events following rare colonizations have led to high rates of endemism at the species and generic levels. The influence of island size, altitude, isolation and other factors on the development of the land snail faunas is discussed. The effect of land use changes on the land snail fauna over the last 150 years is evaluated. Current threats include recent changes in agricultural practice leading to the destruction of snail habitats in both forest and plantation areas. The potential threat posed by introduced species is also discussed. The Gulf of Guinea snail faunas appear to be in relatively good health in comparison to many other island snail faunas. However, the uniqueness of the faunas makes it imperative that further ecological and taxonomic research is undertaken both to understand the processes by which they developed and to evaluate the current status of many species in terms of distribution and threats.     

Colonization and coexistence of non-native ants on a model Atlantic island

Aims

Colonization by non-native ants represents one of the gravest potential threats to island ecosystems. It is necessary to identify general mechanisms by which non-native species are able to colonize and persist in order to inform future prevention and management. We studied a model-island assemblage of 17 non-native ant species with aim of identifying the spatial source of introductions and assessing how such a diversity of species are able to coexist.

Location

Data were collected on Ascension Island: an ideal study system for its intermediate area, compact shape, spatial heterogeneity, lack of native ant species, and availability of non-native ant records dating back to the 1800s.

Methods

We collected over 47,000 individual ants from 73 sites using a range of baited traps and survey techniques. We combined this novel data with past occurrence records in order to determine whether human settlements have historically been the source of ant introductions and to quantify the mean rate at which species have dispersed across the island. Analysis of standardized field data revealed the extent to which ants were partitioning ecological niche space via (1) habitat separation, (2) fine-scale resource partitioning and (3) climatic heterogeneity.

Results

Ants were radiating at a linear rate of approximately 0.5 km² per year from human settlements on this island, with the most widespread species having been introduced earliest. After accounting for incomplete colonization, we found no evidence to suggest habitat separation between species. Instead, we found significant niche separation through resource partitioning and weather-dependent activity patterns.

Main Conclusions

Our results indicate that non-native ants can coexist in very close proximity and are therefore capable of existing at great diversity on even small islands. It is inevitable that ant colonization will continue without increased biosecurity measures, habitat restoration around settlements and conservation of native species populations.     

Range dynamics of small mammals along an elevational gradient over an 80‐year interval

One expected response to observed global warming is an upslope shift of species elevational ranges. Here, we document changes in the elevational distributions of the small mammals within the Ruby Mountains in northeastern Nevada over an 80‐year interval. We quantified range shifts by comparing distributional records from recent comprehensive field surveys (2006–2008) to earlier surveys (1927–1929) conducted at identical and nearby locations. Collector field notes from the historical surveys provided detailed trapping records and locality information, and museum specimens enabled confirmation of species' identifications. To ensure that observed shifts in range did not result from sampling bias, we employed a binomial likelihood model (introduced here) using likelihood ratios to calculate confidence intervals around observed range limits. Climate data indicate increases in both precipitation and summer maximum temperature between sampling periods. Increases in winter minimum temperatures were only evident at mid to high elevations. Consistent with predictions of change associated with climate warming, we document upslope range shifts for only two mesic‐adapted species. In contrast, no xeric‐adapted species expanded their ranges upslope. Rather, they showed either static distributions over time or downslope contraction or expansion. We attribute these unexpected findings to widespread land‐use driven habitat change at lower elevations. Failure to account for land‐use induced changes in both baseline assessments and in predicting shifts in species distributions may provide misleading objectives for conservation policies and management practices.     

Elevational gradients in bird diversity in the Eastern Himalaya: an evaluation of distribution patterns and their underlying mechanisms

Background

Understanding diversity patterns and the mechanisms underlying those patterns along elevational gradients is critically important for conservation efforts in montane ecosystems, especially those that are biodiversity hotspots. Despite recent advances, consensus on the underlying causes, or even the relative influence of a suite of factors on elevational diversity patterns has remained elusive.

Methods and Principal Findings

We examined patterns of species richness, density and range size distribution of birds, and the suite of biotic and abiotic factors (primary productivity, habitat variables, climatic factors and geometric constraints) that governs diversity along a 4500-m elevational gradient in the Eastern Himalayan region, a biodiversity hotspot within the world''s tallest mountains. We used point count methods for sampling birds and quadrats for estimating vegetation at 22 sites along the elevational gradient. We found that species richness increased to approximately 2000 m, then declined. We found no evidence that geometric constraints influenced this pattern, whereas actual evapotranspiration (a surrogate for primary productivity) and various habitat variables (plant species richness, shrub density and basal area of trees) accounted for most of the variation in bird species richness. We also observed that ranges of most bird species were narrow along the elevation gradient. We find little evidence to support Rapoport''s rule for the birds of Sikkim region of the Himalaya.

Conclusions and Significance

This study in the Eastern Himalaya indicates that species richness of birds is highest at intermediate elevations along one of the most extensive elevational gradients ever examined. Additionally, primary productivity and factors associated with habitat accounted for most of the variation in avian species richness. The diversity peak at intermediate elevations and the narrow elevational ranges of most species suggest important conservation implications: not only should mid-elevation areas be conserved, but the entire gradient requires equal conservation attention.     

Can Rapoport's rule explain tree species richness along the Himalayan elevation gradient,Nepal?

Rapoport's rule applied to an elevation gradient predicts a positive correlation between elevation ranges and elevation. This is supposed to be caused by the increasing magnitude of the climatic extremes at higher elevations, and thus, it is deduced that species richness should decrease with increasing elevation. The distribution of 614 tree species was used to test Rapoport's elevational rule along a gradient from 100 to 4300 m a.s.l., in the Nepalese Himalaya. The relationship between species richness and elevation was analysed by using generalized linear models (GLM). Generalized additive models (GAM) were used to examine the relationship between elevational range and the elevational mid-point of a species along the gradient. The widest elevation ranges are observed at mid-elevations, and narrow elevation ranges are observed at both ends of the gradient. This does not support Rapoport's elevation rule, as proposed by Stevens. There is a peak in species richness between 900 and 1000 m, and not in the tropical lowland as projected by Rapoport's elevation rule.