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.     

Patterns of ant species richness along elevational gradients in an arid ecosystem

Aim In this study, we examine patterns of local and regional ant species richness along three elevational gradients in an arid ecosystem. In addition, we test the hypothesis that changes in ant species richness with elevation are related to elevation‐dependent changes in climate and available area. Location Spring Mountains, Nevada, U.S.A. Methods We used pitfall traps placed at each 100‐m elevational band in three canyons in the Spring Mountains. We compiled climate data from 68 nearby weather stations. We used multiple regression analysis to examine the effects of annual precipitation, average July precipitation, and maximum and minimum July temperature on ant species richness at each elevational band. Results We found that patterns of local ant species richness differed among the three gradients we sampled. Ant species richness increased linearly with elevation along two transects and peaked at mid‐elevation along a third transect. This suggests that patterns of species richness based on data from single transects may not generalize to larger spatial scales. Cluster analysis of community similarity revealed a high‐elevation species assemblage largely distinct from that of lower elevations. Major changes in the identity of ant species present along elevational gradients tended to coincide with changes in the dominant vegetation. Regional species richness, defined here as the total number of unique species within an elevational band in all three gradients combined, tended to increase with increasing elevation. Available area decreased with increasing elevation. Area was therefore correlated negatively with ant species richness and did not explain elevational patterns of ant species richness in the Spring Mountains. Mean July maximum and minimum temperature, July precipitation and annual precipitation combined to explain 80% of the variation in ant species richness. Main conclusions Our results suggest that in arid ecosystems, species richness for some taxa may be highest at high elevations, where lower temperatures and higher precipitation may support higher levels of primary production and cause lower levels of physiological stress.     

The influence of distance to perennial surface water on ant communities in Mopane woodlands,northern Botswana

Studies of biodiversity along environmental gradients provide information on how ecological communities change in response to biotic and abiotic factors. For instance, distance to water is associated with several factors that shape the structure and the functioning of ecosystems at a range of spatial scales. We investigated the influence of distance to a perennial water source on ant communities in a semi‐arid savanna in northern Botswana. Ant abundance, taxonomic richness, and both alpha and beta diversity were generally higher during the wet than the dry season. However, there were strong seasonal influences on the effects of distance to water, with more pronounced effects during the wet season. While both abundance and beta diversity declined with increasing distances to water during the wet season, there was a contrasting increase in alpha diversity. There was no major effect of distance to water on taxonomic richness during either season. Beta diversity was as high across as along gradients, and we found support for modular rather than nested community structures along gradients. Our study demonstrated that small‐scale gradients in distance to water can influence several aspects of ant communities in semi‐arid savannas. However, our results also point to strong effects of small‐scale environmental variation, for instance associated with vegetation characteristics, soil properties, and plant community structure that are not directly linked to water access.     

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.     

长白山不同海拔梯度裸肉足虫群落分布特征

裸肉足虫作为联结微生物和大中型土壤动物的重要环节, 在土壤生态系统物质循环和能量流动过程中起着重要作用。为探明裸肉足虫群落沿海拔梯度的分布特征及其主要驱动因子, 作者在长白山北坡选择不同海拔梯度(700 m、1,000 m、1,300 m、1,600 m、1,900 m和2,200 m), 采用最大可能数法对裸肉足虫进行了培养计数, 并采用平板培养、标记、分离再培养的方法进行了分类鉴定, 分析比较了不同海拔梯度裸肉足虫的群落组成和结构特征。结果表明: 长白山北坡裸肉足虫物种丰富, 不同海拔梯度裸肉足虫丰富度指数存在显著差异, 且与土壤酸碱度呈极显著正相关关系。其中林分较为单一的岳桦(Betula ermanii)林带(1,900 m)裸肉足虫丰富度最低, 位于植被交错带的针阔混交林带(1,000 m)裸肉足虫丰富度最大, Shannon-Wiener多样性指数和Pielou均匀度指数在不同海拔梯度间不存在显著性差异, 但变化趋势与丰富度一致。聚类分析结果显示, 1,300 m、1,600 m和1,900 m海拔带以及700 m和2,200 m海拔带裸肉足虫群落组成较为相似。典范对应分析(canonical correspondence analysis, CCA)显示, 裸肉足虫群落组成和结构主要受土壤酸碱度、铵态氮以及碳氮比的影响, 而海拔和土壤含水量对其没有显著影响。综上, 裸肉足虫群落多样性随海拔梯度的增加并未呈现递减或单峰的变化趋势, 土壤的基本理化性质是驱动裸肉足虫群落分布的主要因素; 此外, 地上植被也可能通过凋落物和根系分泌物间接影响裸肉足虫的群落组成和多样性。     

Resource use and food preferences in understory ant communities along a complete elevational gradient in Papua New Guinea

Elevational gradients provide an interesting opportunity for studying the effect of climatic drivers over short distances on the various facets of biodiversity. It is globally assumed that the decrease in species richness with increasing elevation follows mainly the decrease in ecosystem productivity, but studies on functional diversity still remain limited. Here, we investigated how resource use and food preferences by both individual ant species and communities foraging in the understory vary with elevation along a complete elevational gradient (200 to 3200 m asl). Five bait types reflecting some of the main ecosystem processes in which ants are involved were tested: mutualism (sucrose and melezitose), predation (live termites), and detritivory (crushed insects and chicken feces). The observed monotonic decrease in both species richness and occurrences with elevation increase was accompanied by changes in some of the tested ecosystem processes. Such variations can be explained by resource availability and/or resource limitation: Predation and bird feces removal decreased with increasing elevation possibly reflecting a decline in species able to use these resources, while insect detritivory and nectarivory were most probably driven by resource limitation (or absence of limitation), as their relative use did not change along the gradient. Consequently, resource attractiveness (i.e., food preferences at the species level) appears as an important factor in driving community structuring in ants together with the abiotic environmental conditions.     

Nutrient use by tropical ant communities varies among three extensive elevational gradients: A cross-continental comparison

Aim

Many studies demonstrate that climate limits invertebrates along tropical elevational gradients, but we have only a rudimentary understanding of the role of nutrient limitation and climatic seasonality. Here we examined the relationships between ant community structure, nutrient use and season along three undisturbed elevational gradients, each from a different continent.

Location

Ecuador (South America), Papua New Guinea (PNG: Oceania), Tanzania (Africa).

Time period

2011–2014.

Major taxa studied

Ants.

Methods

Along each of the three gradients, we placed six distinct nutrient types (amino acid, sucrose, sucrose + amino acid, lipid, NaCl, H₂O). In total, we distributed 2370 baits at 38 sites from 203 m to 3972 m. We used generalized linear models to test for the effects of elevation and season on ant species richness and activity and relative nutrient use. We also tested if changes in ant trophic guilds corresponded to changes in the use of particular nutrients.

Results

Both species richness and activity decreased with elevation along each gradient. However, there were significant interaction effects among elevation, region and season, as ant activity in the dry season was higher in Ecuador and Tanzania but lower in PNG. Relative nutrient use varied among regions: ant preference for some nutrients changed with increasing elevation in Ecuador (decrease in lipid use) and Tanzania (decrease in amino acid and H₂O use), while season affected nutrient use in PNG. There were common trends in trophic guilds along the three elevational gradients (e.g. proportional increase of predators), but these did not explain most of the nutrient use patterns.

Main conclusion

While the structure of ant communities changed similarly with elevation, both the seasonal and elevational effects on nutrient use by ants differed between continents. We argue that regional differences in climate and nutrient availability rather than ant functional composition shape nutrient use by ants.     

Biogeographic multi‐species occupancy models for large‐scale survey data

Ecologists often seek to infer patterns of species occurrence or community structure from survey data. Hierarchical models, including multi‐species occupancy models (MSOMs), can improve inference by pooling information across multiple species via random effects. Originally developed for local‐scale survey data, MSOMs are increasingly applied to larger spatial scales that transcend major abiotic gradients and dispersal barriers. At biogeographic scales, the benefits of partial pooling in MSOMs trade off against the difficulty of incorporating sufficiently complex spatial effects to account for biogeographic variation in occupancy across multiple species simultaneously. We show how this challenge can be overcome by incorporating preexisting range information into MSOMs, yielding a “biogeographic multi‐species occupancy model” (bMSOM). We illustrate the bMSOM using two published datasets: Parulid warblers in the United States Breeding Bird Survey and entire avian communities in forests and pastures of Colombia''s West Andes. Compared with traditional MSOMs, the bMSOM provides dramatically better predictive performance at lower computational cost. The bMSOM avoids severe spatial biases in predictions of the traditional MSOM and provides principled species‐specific inference even for never‐observed species. Incorporating preexisting range data enables principled partial pooling of information across species in large‐scale MSOMs. Our biogeographic framework for multi‐species modeling should be broadly applicable in hierarchical models that predict species occurrences, whether or not false absences are modeled in an occupancy framework.     

高黎贡山南段海拔梯度森林乔木层时空动态

研究群落物种组成和多样性的时空动态对揭示生物多样性的分布规律以及预测全球变化情景下生物多样性的变化趋势具有重要意义。然而,在山地生态系统中,不同海拔梯度的森林群落物种多样性和系统发育多样性如何随着时间尺度的变化仍不清楚。该研究以高黎贡山南段东、西坡海拔梯度(960～2 878 m)森林群落固定监测样带的17个样方为研究对象,基于2004、2008和2013年乔木层(DBH ≥ 5 cm)重调查数据,分析样方内物种组成、物种多样性和系统发育多样性的时空动态变化。结果表明:(1)沿着海拔梯度,物种多样性呈现单峰分布格局,系统发育多样性呈现上升的趋势,系统发育结构呈现聚集到离散或者随机的结构。(2)在时间尺度上,森林乔木层在物种多样性和系统发育多样性上并未发生显著性变化。然而,系统发育结构随着时间的推移呈现更加聚集的趋势。(3)在海拔梯度上,东坡低海拔区域(960～1 381 m)的森林群落样方呈现显著的物种丧失,其植被完全被耕地所替代。其中,诃子(Terminalia chebula)、麻栎(Quercus acutissima)、清香木(Pistacia weinmanniifolia)、枳椇(Hovenia acerba)和假香冬青(Ilex wattii)等为主要的丧失物种。相反,物种获得主要集中在西坡低海拔的样方,群落中丰富度显著增加的物种主要为曼青冈(Cyclobalanopsis oxyodon)、华山矾(Symplocos chinensis)和台湾杉(Taiwania cryptomerioides)等。据此,我们推测高黎贡山海拔梯度森林乔木层的群落结构和多样性的动态变化在中高海拔受群落演替和气候变化的制约,而在低海拔主要受人类活动的影响。该研究结果加深了对高黎贡山亚热带常绿阔叶林植物群落动态变化的认识,也有助于该地区精准保护策略的制定。     

Predators and priority effects suggested as potential drivers of microfauna communities in a community transplantation experiment along an elevational gradient

Transplantation experiments are a useful method to identify responses of organisms to environmental change. However, they are typically restricted to single or few species. Our experiment was carried out using entire bromeliad‐inhabiting microfauna communities which were transplanted along an elevational gradient, simulating environmental change acting on the communities. Additionally, we manipulated trophic interactions, i.e. resource availability and predator presence, thus combining abiotic and biotic effects in a full‐factorial experimental design. Using this experiment, we found a strong signal of original elevation in microfauna community structure (abundance, evenness, functional composition) with a shift from amoeba‐dominated to flagellate‐dominated communities with increasing original elevation. Surprisingly, the transplantation of communities along the elevational gradient did not affect community structure, indicating strong priority effects. Predation decreased microfauna abundance and increased microfauna evenness, specifically in higher original elevation and high resource levels. In summary, our results suggest that microfauna communities in bromeliads might be primarily shaped by priority effects and predator presence. However, interacting effects (between predator presence and resource availability, as well as between predator presence and original elevation) highlight the usefulness of studies with full‐factorial experimental designs to understand community‐structuring processes. Bromeliads and other micro‐ecosystems provide convenient study systems for community level approaches that could be used in future studies concerning the effects of environmental change, for example climate change on community structure.     

A dynamic multi‐scale occupancy model to estimate temporal dynamics and hierarchical habitat use for nomadic species

Distribution models are increasingly being used to understand how landscape and climatic changes are affecting the processes driving spatial and temporal distributions of plants and animals. However, many modeling efforts ignore the dynamic processes that drive distributional patterns at different scales, which may result in misleading inference about the factors influencing species distributions. Current occupancy models allow estimation of occupancy at different scales and, separately, estimation of immigration and emigration. However, joint estimation of local extinction, colonization, and occupancy within a multi‐scale model is currently unpublished. We extended multi‐scale models to account for the dynamic processes governing species distributions, while concurrently modeling local‐scale availability. We fit the model to data for lark buntings and chestnut‐collared longspurs in the Great Plains, USA, collected under the Integrated Monitoring in Bird Conservation Regions program. We investigate how the amount of grassland and shrubland and annual vegetation conditions affect bird occupancy dynamics and local vegetation structure affects fine‐scale occupancy. Buntings were prevalent and longspurs rare in our study area, but both species were locally prevalent when present. Buntings colonized sites with preferred habitat configurations, longspurs colonized a wider range of landscape conditions, and site persistence of both was higher at sites with greener vegetation. Turnover rates were high for both species, quantifying the nomadic behavior of the species. Our model allows researchers to jointly investigate temporal dynamics of species distributions and hierarchical habitat use. Our results indicate that grassland birds respond to different covariates at landscape and local scales suggesting different conservation goals at each scale. High turnover rates of these species highlight the need to account for the dynamics of nomadic species, and our model can help inform how to coordinate management efforts to provide appropriate habitat configurations at the landscape scale and provide habitat targets for local managers.     

The Argentine ant, Linepithema humile, in Japan: Assessment of impact on species diversity of ant communities in urban environments

The Argentine ant Linepithema humile (Mayr) invaded the Hiroshima Prefecture in south‐west Japan some time before 1990. In this report, we describe the distribution of this exotic ant species and assess its impact on indigenous ant communities in urban areas. L. humile is now widely distributed mainly in urban areas and surrounding secondary vegetation of the cities Hatsukaichi and Hiroshima. The impact assessment suggested that L. humile reduced species diversity of local, indigenous ant communities. There was differential sensitivity of indigenous ant species to the invasion of L. humile. Some ant species disappeared in parks infested with L. humile; for example, Pheidole noda, Pheidole indica and Lasius japonicus. L. humile seemed to be superior to these ant species in certain traits and habits, such as mobility, recruitment ability, aggressiveness and omnivory. In contrast, Paratrechina sakurae and Camponotus vitiosus were less affected by L. humile infestation. The mechanisms allowing such coexistence seemed to be small body size (P. sakurae) and arboreal nesting habits (C. vitiosus).     

Patterns of elevational beta diversity in micro‐ and macroorganisms

Aim While ecologists have long been interested in diversity in mountain regions, elevational patterns in beta diversity are still rarely studied across different life forms ranging from micro‐ to macroorganisms. Also, it is not known whether the patterns in turnover among organism groups are affected by the degree to which the environment is modified by human activities. Location Laojun Mountain, Yunnan Province, China. Methods The beta diversity patterns of benthic microorganisms (i.e. diatoms and bacteria) and macroorganisms (i.e. macroinvertebrates) in a stony stream were simultaneously investigated between elevations of 1820 and 4050 m. Data were analysed by using a distance‐based approach and variation partitioning based on canonical redundancy analysis. Results Analyses of community dissimilarities between adjacent sampling sites showed comparable small‐scale beta diversity along the elevational gradient for the organism groups. However, bacteria clearly showed the lowest elevational turnover when analyses were conducted simultaneously for all pairwise sites. Variation partitioning indicated that species turnover was mostly related to environmental heterogeneity and spatial gradients including horizontal distance and elevation, while purely human impacts were shown to be less important. Main conclusions The elevational beta diversity at large scales was lower for bacteria than for eukaryotic microorganisms or macroorganisms, perhaps indicative of high dispersal ability and good adaptability of bacteria to harsh environmental conditions. However, the small‐scale beta diversity did not differ among the groups. Elevation was the major driver for the turnover of eukaryotic organisms, while the turnover of bacteria was correlated more with environmental variation.     

Multi‐scale ant diversity in savanna woodlands: an intercontinental comparison

Ecological patterns and processes are highly scale‐dependent, but few studies have used standardized methodology to examine how scale dependency varies across continents. This paper examines scale dependency in comparative ant species richness and turnover in savannas of Australia and Brazil, which are well‐matched climatically but whose ant faunas have contrasting biogeographic origins. The study was conducted in savanna woodland near Darwin in northern Australia and Uberlândia in central Brazil. The sampling design consisted of eight 400‐m line transects, four in each continent, with eight pitfall traps located on and around each of 20 trees evenly spaced along each transect. Ant richness and species turnover were compared at three spatial scales: pitfalls associated with a tree, trees within a transect and transects within a savanna. The composition of the Australian and Brazilian savanna ant faunas was broadly similar at the subfamily level, despite the very low proportion of shared genera and species. The ground and arboreal ant faunas were very distinct from each other in both savannas, but especially in Brazil. Overall ant abundance was almost three times higher in Australia than in Brazil, both on the ground and on vegetation, but overall species richness was higher in Brazil (150 species) than in Australia (93). There was no significant difference in the mean number of species per pitfall trap, but the mean species richness was significantly higher in Brazil than in Australia at both the tree and transect scales. We attribute these scale‐dependent intercontinental differences to biogeographical and historical factors in Brazil that have led to a large regional pool of arboreal species of rainforest origin. Our study underlines the importance of biogeographical context when conducting comparative analyses of community structure across biogeographical scales, and highlights the importance of process acting at regional scales in determining species richness in ant communities.     

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.