Vertical stratification of vascular epiphytes in submontane and montane forest of the Bolivian Andes: the importance of the understory

We studied species richness, composition, and vertical distribution of vascular epiphytes at two sites in the Bolivian Andes. To account for the epiphyte flora on understory trees, epiphytes on shrubs and small trees were sampled in 20 × 20 m² subplots around each sampled canopy tree; this understory zone U is introduced as an addition to the well-established five vertical Johansson tree zones. More than 20% of about 500 species recorded were found only in the understory subplots, including ca. 40% of aroids, 35%–40% of piperoids, and 25%–30% of ferns. Habitat generalists (occurring in three or more zones) were most common, contributing about 50% of all species, specialists (occurring only in two zones, or in three continuous ones) 34%–42%, and hemiepiphytes 6%–16%. Canopy epiphytes (occurring > 90% in tree zones Z3–5) were mainly represented by orchids and ferns, many with special adaptations to drought stress such as pseudobulbs, succulence, and poikilohydry. Trunk epiphytes ( > 90% in understory and tree zones Z1–2) reached highest relative species numbers among piperoids and ferns. Most hemiepiphytes were also trunk epiphytes, due to their characteristic growth pattern, and included mainly aroids. The vertical distribution of epiphytes within a tree is determined by several microenvironmental gradients, with light intensity, wind speed, and air temperature increasing and air humidity decreasing from the ground level to the canopy.     

Diversity and composition of Arctiidae moth ensembles along a successional gradient in the Ecuadorian Andes

Andean montane rain forests are among the most species‐rich terrestrial habitats. Little is known about their insect communities and how these respond to anthropogenic habitat alteration. We investigated exceptionally speciose ensembles of nocturnal tiger moths (Arctiidae) at 15 anthropogenically disturbed sites, which together depict a gradient of forest recovery and six closed‐forest understorey sites in southern Ecuador. At weak light traps we sampled 9211 arctiids, representing 287 species. Arctiid abundance and diversity were highest at advanced succession sites, where secondary scrub or young forest had re‐established, followed by early succession sites, and were lowest, but still high, in mature forest understorey. The proportion of rare species showed the reverse pattern. We ordinated moth samples by non‐metric multidimensional scaling using the chord‐normalized expected species shared index (CNESS) index at various levels of the sample size parameter m. A distinct segregation of arctiid ensembles at succession sites from those in mature forest consistently emerged only at high m‐values. Segregation between ensembles of early vs. late succession stages was also clear at high m values only, and was rather weak. Rare species were responsible for much of the faunal difference along the succession gradient, whereas many common arctiid species occurred in all sites. Matrix correlation tests as well as exploration of relationships between ordination axes and environmental variables revealed the degree of habitat openness, and to a lesser extent, elevation, as best predictors of faunal dissimilarity. Faunal differences were not related to geographical distances between sampling sites. Our results suggest that many of the more common tiger moths of Neotropical montane forests have a substantial recolonization potential at the small spatial scale of our study and accordingly occur also in landscape mosaics surrounding nature reserves. These species contribute to the unexpectedly high diversity of arctiid ensembles at disturbed sites, whereas the proportion of rare species declines outside mature forest.     

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.     

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.     

Diversity of dry forest epiphytes along a gradient of human disturbance in the tropical Andes

Question: Disturbance effects on dry forest epiphytes are poorly known. How are epiphytic assemblages affected by different degrees of human disturbance, and what are the driving forces? Location: An inter‐Andean dry forest landscape at 2300 m elevation in northern Ecuador. Methods: We sampled epiphytic bryophytes and vascular plants on 100 trees of Acacia macracantha in five habitats: closed‐canopy mixed and pure acacia forest (old secondary), forest edge, young semi‐closed secondary woodland, and isolated trees in grassland. Results: Total species richness in forest edge habitats and on isolated trees was significantly lower than in closed forest types. Species density of vascular epiphytes (species per tree) did not differ significantly between habitat types. Species density of bryophytes, in contrast, was significantly lower in edge habitat and on isolated trees than in closed forest. Forest edge showed greater impoverishment than semi‐closed woodland and similar floristic affinity to isolated trees and to closed forest types. Assemblages were significantly nested; habitat types with major disturbance held only subsets of the closed forest assemblages, indicating a gradual reduction in niche availability. Distance to forest had no effect on species density of epiphytes on isolated trees, but species density was closely correlated with crown closure, a measure of canopy integrity. Main conclusions: Microclimatic changes but not dispersal constraints were key determinants of epiphyte assemblages following disturbance. Epiphytic cryptogams are sensitive indicators of microclimate and human disturbance in montane dry forests. The substantial impoverishment of edge habitat underlines the need for fragmentation studies on epiphytes elsewhere in the Tropics.     

Net primary productivity allocation and cycling of carbon along a tropical forest elevational transect in the Peruvian Andes

The net primary productivity, carbon (C) stocks and turnover rates (i.e. C dynamics) of tropical forests are an important aspect of the global C cycle. These variables have been investigated in lowland tropical forests, but they have rarely been studied in tropical montane forests (TMFs). This study examines spatial patterns of above‐ and belowground C dynamics along a transect ranging from lowland Amazonia to the high Andes in SE Peru. Fine root biomass values increased from 1.50 Mg C ha^?1 at 194 m to 4.95 ± 0.62 Mg C ha^?1 at 3020 m, reaching a maximum of 6.83 ± 1.13 Mg C ha^?1 at the 2020 m elevation site. Aboveground biomass values decreased from 123.50 Mg C ha^?1 at 194 m to 47.03 Mg C ha^?1 at 3020 m. Mean annual belowground productivity was highest in the most fertile lowland plots (7.40 ± 1.00 Mg C ha^?1 yr^?1) and ranged between 3.43 ± 0.73 and 1.48 ± 0.40 Mg C ha^?1 yr^?1 in the premontane and montane plots. Mean annual aboveground productivity was estimated to vary between 9.50 ± 1.08 Mg C ha^?1 yr^?1 (210 m) and 2.59 ± 0.40 Mg C ha^?1 yr^?1 (2020 m), with consistently lower values observed in the cloud immersion zone of the montane forest. Fine root C residence time increased from 0.31 years in lowland Amazonia to 3.78 ± 0.81 years at 3020 m and stem C residence time remained constant along the elevational transect, with a mean of 54 ± 4 years. The ratio of fine root biomass to stem biomass increased significantly with increasing elevation, whereas the allocation of net primary productivity above‐ and belowground remained approximately constant at all elevations. Although net primary productivity declined in the TMF, the partitioning of productivity between the ecosystem subcomponents remained the same in lowland, premontane and montane forests.     

Distribution and Flowering Ecology of Bromeliads along Two Climatically Contrasting Elevational Transects in the Bolivian Andes1

We compared the diversity, taxonomic composition, and pollination syndromes of bromeliad assemblages and the diversity and abundance of hummingbirds along two climatically contrasting elevational gradients in Bolivia. Elevational patterns of bromeliad species richness differed noticeably between transects. Along the continuously wet Carrasco transect, species richness peaked at mid‐elevations, whereas at Masicurí most species were found in the hot, semiarid lowlands. Bromeliad assemblages were dominated by large epiphytic tank bromeliads at Carrasco and by small epiphytic, atmospheric tillandsias at Masicurí. In contrast to the epiphytic taxa, terrestrial bromeliads showed similar distributions across both transects. At Carrasco, hummingbird‐pollination was the most common pollination mode, whereas at Masicurí most species were entomophilous. The proportion of ornithophilous species increased with elevation on both transects, whereas entomophily showed the opposite pattern. At Carrasco, the percentage of ornithophilous bromeliad species was significantly correlated with hummingbird abundance but not with hummingbird species richness. Bat‐pollination was linked to humid, tropical conditions in accordance with the high species richness of bats in tropical lowlands. At Carrasco, mixed hummingbird/bat‐pollination was found especially at mid‐elevations, i.e., on the transition between preferential bat‐pollination in the lowlands and preferential hummingbird‐pollination in the highlands. In conclusion, both richness patterns and pollination syndromes of bromeliad assemblages varied in distinct and readily interpretable ways in relation to environmental humidity and temperature, and bromeliad pollination syndromes appear to follow the elevational gradients exhibited by their pollinators.     

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。     

Stem respiration in tropical forests along an elevation gradient in the Amazon and Andes

Autotrophic respiration involves the use of fixed carbon by plants for their own metabolism, resulting in the release of carbon dioxide as a by‐product. Little is known of how autotrophic respiration components vary across environmental gradients, particularly in tropical ecosystems. Here, we present stem CO₂ efflux data measured across an elevation transect spanning ca. 2800 m in the Peruvian Amazon and Andes. Forest plots from five elevations were studied: 194, 210, 1000, 1500, and 3025 m asl Stem CO₂ efflux (R_s) values from each plot were extrapolated to the 1‐ha plot level. Mean R_s per unit stem surface area declined significantly with elevation, from 1.14±0.12 at 210 m elevation to 0.62±0.09 μmol C m⁻² s⁻¹ at 3025 m elevation. When adjusted for changing forest structure with elevation, this is equivalent to 6.45±1.12 Mg C ha⁻¹ yr⁻¹ at 210 m elevation to 2.94±0.19 Mg C ha⁻¹ yr⁻¹ at 3025 m elevation. We attempted to partition stem respiration into growth and maintenance respiration components for each site. Both growth and maintenance respiration rates per unit stem showed similar, moderately significant absolute declines with elevation, but the proportional decline in growth respiration rates was much greater. Stem area index (SAI) showed little trend along the transect, with declining tree stature at higher elevations being offset by an increased number of small trees. This trend in SAI is sensitive to changes in forest stature or size structure. In the context of rapid regional warming over the 21st century, such indirect, ecosystem‐level temperature responses are likely to be as important as the direct effects of temperature on maintenance respiration rates.     

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

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

A human-induced downward-skewed elevational abundance distribution of pteridophytes in the Bolivian Andes

Aim  To search for differences in the spatial variability of upper and lower elevational distribution limits of tropical ferns, based on the assumption that these are determined to different degrees by biotic and abiotic factors.
Location  The Yungas biogeographical region, in the Bolivian Andes.
Methods  From a data base of > 25,000 herbarium records, we analysed the skewness of the elevational distribution of 220 montane pteridophyte species, each with  15 records. Additionally, we compared the spatial variability of upper and lower elevational range limits of ferns in 351 plots of 400 m² each along four elevational transects separated by 15–450 km.
Results  Individual species showed variable elevational distribution patterns, ranging from symmetric to asymmetric, i.e. downward and upward skewed, but overall there was a statistically significant trend towards asymmetric distributions with abrupt upper limits and diffuse lower limits. This trend, however, was almost exclusively due to terrestrial species occurring at and above the current timberline. The analysis of the elevational transects revealed no significant trends.
Main conclusions  The downward-skewed distributional abundance of terrestrial, open-country ferns near the timberline appears to be a result of the extensive forest destruction that has lowered the timberline in the high Andes by 500–800 m, opening up habitats for a restricted suite of species. Our study shows that a limited number of species can cause a general trend in the overall data set, and that failure to extract these data may result in unsupported conclusions, in our case to assign a greater importance to biotic and abiotic factors in the elevational limitation of plants at lower and upper elevations, respectively.     

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。     

Differences in carbon stocks along an elevational gradient in tropical mountain forests of Colombia

Tropical mountain forests provide an exceptional opportunity to evaluate the patterns of variation in carbon stocks along elevational gradients that correspond to well‐defined temperature gradients. We predicted that carbon stored in live aboveground biomass, aboveground necromass, and soil components of forests on the eastern flank of the Colombian Andes would change with elevation along this gradient extending from 750 to 2,800 m above sea level. The rationale was that the corresponding change in temperature (14–26°C) would influence tree growth and decomposition of organic matter. To address this hypothesis, we examined the carbon stored in these three components using data from 20 0.25‐ha plots located along this elevational gradient. The mean total carbon stock found in the study region was 241.3 ± 37.5 Mg C/ha. Aboveground carbon stocks decreased with elevation (p = 0.001), as did necromass carbon stocks (p = 0.016). Although soil organic carbon stocks did not differ significantly along the gradient (p = 0.153), they contributed proportionately more at higher than at lower elevations, counterbalancing the opposite trends in aboveground carbon and necromass carbon stocks. As such, total carbon stocks did not vary significantly along the elevational gradient (p = 0.576).     

中国森林附生维管植物多样性数据集

附生维管植物是热带、亚热带湿性山地森林植物群落中物种最为丰富的类群之一, 在维系森林生物多样性及生态系统功能与服务中发挥着重要作用, 然而附生维管植物调查相对困难, 在多样性调查和统计中往往被忽视, 迄今为止我国尚缺乏全国范围的森林附生维管植物名录。本项研究基于已公开发布的数据库, 搜集并整理了1982-2022年间的关于中国森林附生维管植物研究的文献书籍。首先, 提取文献资料中的有效信息, 依据物种2000中国节点的数据进行标准化校正, 整理出中国森林附生维管植物共计49科246属1,739种, 然后据此编写了相对准确、完整的中国森林附生维管植物物种名录。本名录作为我国第一个全国性森林附生维管植物名录, 可为森林生物多样性保护工作提供重要的数据支持, 并能为深入研究附生维管植物的进化生物学、生物地理学及宏观生态学等领域提供重要的基础信息。     

Replicated hybrid zones of Xiphophorus swordtails along an elevational gradient

Natural hybrid zones provide opportunities to study a range of evolutionary phenomena from speciation to the genetic basis of fitness-related traits. We show that widespread hybridization has occurred between two neo-tropical stream fishes with partial reproductive isolation. Phylogenetic analyses of mitochondrial sequence data showed that the swordtail fish Xiphophorus birchmanni is monophyletic and that X. malinche is part of an independent monophyletic clade with other species. Using informative single nucleotide polymorphisms in one mitochondrial and three nuclear intron loci, we genotyped 776 specimens collected from twenty-three sites along seven separate stream reaches. Hybrid zones occurred in replicated fashion in all stream reaches along a gradient from high to low elevation. Genotyping revealed substantial variation in parental and hybrid frequencies among localities. Tests of F(IS) and linkage disequilibrium (LD) revealed generally low F(IS) and LD except in five populations where both parental species and hybrids were found suggesting incomplete reproductive isolation. In these locations, heterozygote deficiency and LD were high, which suggests either selection against early generation hybrids or assortative mating. These data lay the foundation to study the adaptive basis of the replicated hybrid zone structure and for future integration of behaviour and genetics to determine the processes that lead to the population genetic patterns observed in these hybrid zones.