Small mammal species richness and turnover along elevational gradient in Yulong Mountain,Yunnan, Southwest China

Understanding the species diversity patterns along elevational gradients is critical for biodiversity conservation in mountainous regions. We examined the elevational patterns of species richness and turnover, and evaluated the effects of spatial and environmental factors on nonvolant small mammals (hereafter “small mammal”) predicted a priori by alternative hypotheses (mid‐domain effect [MDE], species–area relationship [SAR], energy, environmental stability, and habitat complexity]) proposed to explain the variation of diversity. We designed a standardized sampling scheme to trap small mammals at ten elevational bands across the entire elevational gradient on Yulong Mountain, southwest China. A total of 1,808 small mammals representing 23 species were trapped. We observed the hump‐shaped distribution pattern of the overall species richness along elevational gradient. Insectivores, rodents, large‐ranged species, and endemic species richness showed the general hump‐shaped pattern but peaked at different elevations, whereas the small‐ranged species and endemic species favored the decreasing richness pattern. The MDE and the energy hypothesis were supported, whereas little support was found for the SAR, the environmental stability hypothesis, and the habitat complexity. However, the primary driver(s) for richness patterns differed among the partitioning groups, with NDVI (the normalized difference vegetation index) and MDE being the most important variables for the total richness pattern. Species turnover for all small mammal groups increased with elevation, and it supported a decrease in community similarity with elevational distance. Our results emphasized for increased conservation efforts in the higher elevation regions of the Yulong Mountain.     

贡嘎山东坡非飞行小型兽类物种多样性的垂直分布格局

为了解贡嘎山地区物种多样性的垂直分布格局,2010年4—9月利用夹日法对贡嘎山东坡非飞行小型兽类的物种多样性进行了详细调查。调查在海拔1200—4000m之间按400m间隔设置了8个采集样地,累计布夹28800夹次,捕获非飞行小型兽类个体701个,观察记录到松鼠个体25个,共调查记录小兽个体726个,分属于3目6科16属25种。非参数估计的物种丰富度Chao2和Jackknife2指数以及物种累积曲线评估表明本次调查取样充分,能很好地反映该地区非飞行小型兽类物种多样性的垂直分布格局。结果表明:非飞行小型兽类物种多样性的垂直分布格局为单峰模型;物种丰富度和物种多度在中海拔地区最高,在低海拔和高海拔地区较低;相反,物种均匀度在中海拔地区较低,在低海拔和高海拔地区较高;而物种优势度则随着海拔的升高而逐渐增加;Shannon-Wiener、Fisher-α、Margalef三个综合性物种多样性指数均显示物种多样性在中海拔地区最高;与其它多样性指数相比,Simpson指数未能很好地反映出不同海拔段群落物种多样性的垂直分布差异;而与Shannon-Wiener和Simpson指数相比,Fisher-α和Margalef指数对不同物种组成的群落多样性区分较好。同时,基于不同海拔段物种组成的聚类分析结果也表明物种多样性在中海拔地区最高。物种多样性在中海拔地区最高的垂直分布模式提示我们在贡嘎山地区的生物多样性保护和生态管理中应特别重视中海拔地段,因为该地段中居于生态食物链中间位置的小兽物种最丰富,是山地生物多样性保护的关键。此外,规范统一的调查方法将有利于研究数据的整合和减少人为因素带来的误差。     

Ecological and evolutionary drivers of the elevational gradient of diversity

Ecological, evolutionary, spatial and neutral theories make distinct predictions and provide distinct explanations for the mechanisms that control the relationship between diversity and the environment. Here, we test predictions of the elevational diversity gradient focusing on Iberian bumblebees, grasshoppers and birds. Processes mediated by local abundance and regional diversity concur in explaining local diversity patterns along elevation. Effects expressed through variation in abundance were similar among taxa and point to the overriding role of a physical factor, temperature. This determines how energy is distributed among individuals and ultimately how the resulting pattern of abundance affects species incidence. Effects expressed through variation in regional species pools depended instead on taxon‐specific evolutionary history, and lead to diverging responses under similar environmental pressures. Local filters and regional variation also explain functional diversity gradients, in line with results from species richness that indicate an (local) ecological and (regional) historical unfolding of diversity–elevation relationships.     

The mid-domain effect applied to elevational gradients: species richness of small mammals in Costa Rica

Aim The objective of this study was to comprehensively document and examine the alpha and gamma patterns of species richness in non-volant, small mammals (rodents, shrews and mouse opossums) along a tropical elevational gradient. These data were used to determine the support for existing hypotheses of species richness encompassing mid-domain null models, as well as climatic, and community overlap hypotheses. Location Field studies were conducted along a Caribbean slope of the Río Peñas Blancas watershed in the north-eastern region of Costa Rica between 750 and 1850 m at 10 sampling sites. Methods Species richness and abundances of small mammals were surveyed for four seasons including three temporal replicates at each of five elevational sites: late wet season (2000), early wet season (2001), and dry season (2002), and one spatial replicate at five different sites within the same elevations during the late wet season (2001). Species richness at elevations below 700 m was compiled from specimen records from 23 US national and international collections. Predictions of a null model based solely on geometric constraints were examined using a Monte Carlo simulation program, Mid-Domain Null. Results In 16,900 trap nights, 1561 individuals from 16 species were captured. Both alpha and gamma species richness peaked at mid-elevation between 1000 and 1300 m, with richness declining both at higher and lower elevations. Most of the empirical curves of species richness occur within 95% prediction curves of the mid-domain model, although deviations from the null model exist. Regression of the empirical richness on the null model predictions explained nearly half of the variation observed (r² = 0.45, P = 0.002). Main conclusions The geometric constraints of montane topography appear to influence the diversity pattern of small mammals, although climatic conditions including an intermediate rainfall and temperature regime, and distance from the persistent cloud cap also are correlated with the pattern of species richness. The predictions of productivity, and community overlap hypotheses are not supported with the empirical data.     

Community of small mammals along an elevational gradient in Biological Reserve of Serra do Japi,municipality of Jundiaí‐SP,Brazil

Studies of elevational gradients in forests are particularly interesting for the considerable differences that can be observed over short distances, such as in vegetation and temperature. Different taxonomic groups display varying types of distribution patterns along elevational gradients, with unimodal distribution being recognised as the most common pattern. The distribution of species can be affected by a range of factors that include, biotic, spatial, climatic, historic and energetic. Small mammals represent an ideal model for studies about distribution and habitat use as they can be highly abundant, tend to have different diets and use space differently. The aims of this study are to build a comprehensive understanding of the community of small mammals of the Biological Reserve of Serra do Japi and to explore its distribution pattern along elevational gradients. We investigated the influence of biomass of arthropods, fruits and seeds and percentage of ground cover, canopy cover and vertical vegetation at richness and abundance of small mammals at three different elevations. To accomplish this, we used seventy‐two pitfall traps of 63 L to capture small mammals and distributed them equally across three elevations defined as low (880–899 m), intermediate (1046–1089 m) and high (1170–1189 m). Each elevation had three lines or replicas of traps. Throughout the study, we captured one hundred and fourteen individuals belonging to eleven species of small mammals. The presence of rare and endemic species demonstrates the importance of conservation and maintenance of the Biological Reserve of Serra do Japi. In regard to the distribution of species, despite the short gradient range, we found a unimodal pattern and a positive correlation between ground cover (fallen twigs and branches up to 1 m high) and richness and abundance of small mammals. More ground cover can reduce the effects of competition and predation on small mammals’ communities. Abstract in Portuguese is available with online material.     

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.     

Breeding systems and phylogenetic diversity of seed plants along a large‐scale elevational gradient

Aim The factors affecting the distribution of dioecious species have received surprisingly little attention and, as a consequence, are poorly understood. For example, there is a well‐documented negative association between dioecy and latitude, for which we have no candidate mechanisms. Conversely, it has been suggested that the relative proportion of dioecious species should be positively correlated with changes in elevation. Location Costa Rica, Central America. Methods We made use of data on the distribution of 175 seed plant species from a series of plots along a transect in Costa Rica that ranged from 30 to 2600 m a.s.l. to test the prediction that dioecy is correlated with elevation. Specifically, we examined relationships between the proportion of dioecy, at the species and individual levels, and the elevation, species richness, number of individuals, and phylogenetic diversity (PD) of plots. For comparison, we repeated all analyses with monoecious species, which also have unisexual flowers but do not suffer from reduced mate assurance and the seed shadow effect that may be the outcomes of having spatially separated sexes. Results The relative proportions of dioecious species and individuals displayed a unimodal relationship with elevation, both peaking at 750 m a.s.l. In contrast, the relative proportions of monoecious species and individuals displayed negative associations with elevation. In addition, the proportion of dioecious species was significantly positively associated with species richness and with the PD of plots. The proportion of monoecious species was not associated with species richness and was weakly positively associated with the PD of plots. Main conclusions Our results suggest that the selection pressure of elevation on the pollinators and life‐history strategy of dioecious species is much less than expected, and is overshadowed by the as‐yet unexplained correlation of dioecy with species richness. Additional studies of the ecology of cosexual and unisexual (i.e. dioecious and monoecious) species along large environmental gradients are needed.     

Contrasting patterns in elevational diversity between microorganisms and macroorganisms

Aim Data and analyses of elevational gradients in diversity have been central to the development and evaluation of a range of general theories of biodiversity. Elevational diversity patterns have, however, been severely understudied for microbes, which often represent decomposer subsystems. Consequently, generalities in the patterns of elevational diversity across different trophic levels remain poorly understood. Our aim was to examine elevational gradients in the diversity of macroinvertebrates, diatoms and bacteria along a stony stream that covered a large elevational gradient. Location Laojun Mountain, Yunnan province, China. Methods The sampling scheme included 26 sites spaced at elevational intervals of 89 m from 1820 to 4050 m elevation along a stony stream. Macroinvertebrate and diatom richness were determined based on the morphology of the specimens. Taxonomic richness for bacteria was quantified using a molecular fingerprinting method. Over 50 environmental variables were measured at each site to quantify environmental variables that could correlate with the patterns of diversity. We used eigenvector‐based spatial filters with multiple regressions to account for spatial autocorrelation. Results The bacterial richness followed an unexpected monotonic increase with elevation. Diatoms decreased monotonically, and macroinvertebrate richness showed a clear unimodal pattern with elevation. The unimodal richness pattern for macroinvertebrates was best explained by the mid‐domain effect (r² = 0.72). The diatom richness was best explained by the variation in nutrient supply, and the increase in bacterial richness with elevation may be related to an increased carbon supply. Main conclusions We found contrasting patterns in elevational diversity among the three studied multi‐trophic groups comprising unicellular and multicellular aquatic taxa. We also found that there may be fundamental differences in the mechanisms underlying these species diversity patterns.     

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.     

Phylogenetic diversity,structure and diversification patterns of endemic plants along the elevational gradient in the Eastern Himalaya

ABSTRACT

Background: Several studies have documented the variation in species diversity patterns along elevational gradients in the Himalaya, but few have reported the evolutionary and biogeographic processes behind these patterns.

Aims: To understand whether evolutionary history and phylogeny have any role in structuring plant species communities along an elevational gradient in the Sikkim Himalaya.

Methods: We used data on endemic plant species occurrence from primary and secondary sources to construct family-level phylogenetic supertrees for different growth forms with the help of Phylomatic tool of Phylocom. These phylogenetic supertrees were used as a base for testing phylogenetic diversity (PD), niche conservatism, diversification time patterns and phylogenetic structure of various plant growth forms along an elevational gradient.

Results: PD was the highest at mid-elevations for all growth forms and PD had a significant positive correlation with endemic species richness. Species at mid-elevations were dominated by the ancestral/primitive taxa. There was phylogenetic clustering at higher elevations and phylogenetic overdispersion at lower and mid-elevations for the majority of the growth forms.

Conclusions: Time-for-speciation effect and niche conservatism along elevation (retention of niche-related ancestral elevational distribution over evolutionary time scale by species) together determine plant species diversity patterns in the Himalaya.     

Habitat heterogeneity,temperature, and primary productivity drive elevational gradients in avian species diversity

AimAnticipating and mitigating the impacts of climate change on species diversity in montane ecosystems requires a mechanistic understanding of drivers of current patterns of diversity. We documented the shape of elevational gradients in avian species richness in North America and tested a suite of a priori predictions for each of five mechanistic hypotheses to explain those patterns.LocationUnited StatesMethodsWe used predicted occupancy maps generated from species distribution models for each of 646 breeding birds to document elevational patterns in avian species richness across the six largest U.S. mountain ranges. We used spatially explicit biotic and abiotic data to test five mechanistic hypotheses proposed to explain geographic variation in species richness.ResultsElevational gradients in avian species richness followed a consistent pattern of low elevation plateau‐mid‐elevation peak (as per McCain, 2009). We found support for three of the five hypotheses to explain the underlying cause of this pattern: the habitat heterogeneity, temperature, and primary productivity hypotheses.Main ConclusionsSpecies richness typically decreases with elevation, but the primary cause and precise shape of the relationship remain topics of debate. We used a novel approach to study the richness‐elevation relationship and our results are unique in that they show a consistent relationship between species richness and elevation among 6 mountain ranges, and universal support for three hypotheses proposed to explain the underlying cause of the observed relationship. Taken together, these results suggest that elevational variation in food availability may be the ecological process that best explains elevational gradients in avian species richness in North America. Although much attention has focused on the role of abiotic factors, particularly temperature, in limiting species’ ranges, our results offer compelling evidence that other processes also influence (and may better explain) elevational gradients in species richness.     

The mid‐domain effect and habitat complexity applied to elevational gradients: Moss species richness in a temperate semihumid monsoon climate mountain of China

The utility of elevational gradients as tools to test either ecological hypotheses and delineate elevation‐associated environmental factors that explain the species diversity patterns is critical for moss species conservation. We examined the elevational patterns of species richness and evaluated the effects of spatial and environmental factors on moss species predicted a priori by alternative hypotheses, including mid‐domain effect (MDE), habitat complexity, energy, and environment proposed to explain the variation of diversity. Last, we assessed the contribution of elevation toward explaining the heterogeneity among sampling sites. We observed the hump‐shaped distribution pattern of species richness along elevational gradient. The MDE and the habitat complexity hypothesis were supported with MDE being the primary driver for richness patterns, whereas little support was found for the energy and the environmental factors.     

Species richness and assemblages of bats along a forest elevational transect in Papua New Guinea

Over the past decades, elevational gradients have become a powerful tool with which to understand the underlying cause(s) of biodiversity. The Mt. Wilhelm elevational transect is one such example, having been used to study the birds, insects, and plants of Papua New Guinea (PNG). However, a survey of mammals from this forest elevational transect was lacking. We thus aimed to investigate patterns in the community structure and species richness of bats (Chiroptera) along the transect, link the species to available regional data, and explain the observed patterns by including environmental characteristics. Bat assemblages were surveyed between 200 m and a timberline at 3700 m a.s.l. at eight study sites separated by 500 m in elevation. We conducted mist-netting and acoustic surveys to detect and identify species at each site. Regional data were compiled to compare local with regional diversity. Finally, biotic (i.e., food availability, habitat features) and abiotic (i.e., mean daily temperature) factors were included in our analyses to disentangle the ecological drivers underlying bat diversity. Results revealed that species richness decreases with ascending elevation and was best explained by a corresponding decrease in temperature. We observed both turnover and nestedness of the species composition at regional scale whereas turnover was dominant at local scale. Extensions and shifts of bat elevational ranges were also found in Mt. Wilhelm. Consequently, despite that the study was restricted to one mountain in PNG, it demonstrates how basic inventory surveys can be used to address ecological questions in other similar and undisturbed tropical mountains.     

Patterns in diversity of anurans along an elevational gradient in the Western Ghats, South India

Aim To examine patterns in anuran species richness along an elevation gradient and identify factors that govern anuran species richness on a tropical elevational gradient. Location Sampling for anurans was carried out in Kalakad Mundanthurai Tiger Reserve (KMTR) in the southern Western Ghats, India. Methods Night‐time sampling for anuran species richness was carried out from 20 November 2004 to 20 April 2005, during the north‐east monsoon and dry seasons, using transects (50 × 2 m) and visual encounter surveys along the streams. The entire gradient was classified into thirteen 100‐m elevation zones. Sampling at the alpha (single drainage basin) level was carried out in the Chinnapul River drainage basin (40–1260 m a.s.l.) and at the gamma (landscape) level in four drainage basins. Additionally, published records were used to arrive at an empirical species richness (S) for the entire landscape. Mid‐Domain Null software was used to test for the possible influence of geometric constraints on anuran species at both the alpha and gamma levels. The influence of area under each elevation zone on empirical S was tested. The pattern in anuran species richness along the elevational gradient was investigated using: (1) species boundaries in each elevation zone and their habitat correlates, (2) abiotic factors as predictor variables, (3) mean snout vent lengths of anurans, and (4) correlation between the matrices of distance in the elevation zones based on microhabitat parameters and species composition. Cluster analysis on species presence–absence in the elevation zones was used to categorize the entire gradient into high, middle and low elevations. In these three elevation categories, pattern in composition of species was examined for endemism in Western Ghats–Sri Lanka biodiversity hotspot, uniqueness to an elevation zone, adaptations of adults and modes of breeding. Results Species richness at the alpha level increased linearly with elevation, while at the gamma level there were three peaks. Maximum species richness was observed at the highest elevation (1200 m) at both the alpha and the gamma levels. The observed patterns differed significantly from mid‐domain null predictions. The multi‐modal pattern in species richness was a consequence of overlapping species range boundaries. Soil temperature was the best single measure in explaining the majority of variation in species richness at the alpha level (r² = 0.846, P < 0.01). However, soil moisture was the best predictor when both the alpha and the gamma sites were pooled (r² = 0.774, P < 0.01). Anuran body size decreased with an increase in elevation. The highest proportions of endemic and unique species were found at high elevations (> 700 m). The proportion of arboreal anurans increased from low to high elevation. Anurans exhibiting direct development were predominantly found at high elevations. Main conclusions Geometric constraints did not influence anuran species richness along the elevational gradient. Overlapping range boundaries influenced species richness at the gamma level. Abiotic factors such as soil temperature and moisture influenced anuran species richness in the mountain range. The ‘Massenerhebung effect’ could be responsible for range restriction and endemism of anurans, differences in guilds and mode of reproduction. These findings highlight the importance of cloud forests for endemic anurans.     

Elevational gradients of small mammal diversity on the northern slopes of Mt. Qilian, China

Aim Small mammal species richness and relative abundance vary along elevational gradients, but there are different patterns that exist. This study reports the patterns of distribution and abundance of small mammals along the broader elevational gradient of Mt. Qilian range. Location The study was conducted in the Mt. Qilian range, north‐western China, from June to August 2001. Methods Removal trapping was conducted using a standardized technique at 7 sites ranging between 1600 and 3900 m elevation within three transects. Correlation, regression and graphical analyses were used to evaluate the diversity patterns along this elevational gradient. Results In total, 586 individuals representing 18 nonvolant small mammal species were collected during 20 160 trap nights. Species composition was different among the three transects with 6 (33%) of the species found only within one transect. Elevational distribution and relative abundance of small rodents showed substantial spatial variation, with only 2 species showing nonsignificant capture frequencies across elevations. Despite these variations, some general patterns of elevational distribution emerged: humped‐shape relationships between species diversity and elevation were noted in all three transects with diversity peaks at middle elevations. In addition, relative abundance was negatively correlated with elevation. Conclusions Results indicate that maximum richness and diversity of nonvolant small mammals occurred at mid‐elevations where several types of plants reached their maximum diversity and primary productivity, and where rainfall and humidity reached a maximum. It is demonstrated that the mid‐elevation bulge is a general feature of at least a large portion of the biota on the Mt. Qilian range.