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.     

What drives elevational patterns of diversity? A test of geometric constraints,climate and species pool effects for pteridophytes on an elevational gradient in Costa Rica

Aim We studied pteridophyte species richness between 100 m and 3400 m along a Neotropical elevational gradient and tested competing hypotheses for patterns of species richness. Location Elevational transects were situated at Volcán Barva in the Braulio Carrillo National Park and La Selva Biological Station (100–2800 m) and Cerro de la Muerte (2700–3400 m), both on the Atlantic slope of Costa Rica, Central America. Method We analysed species richness on 156 plots of 20 × 20 m and measured temperature and humidity at four elevations (40, 650, 1800 and 2800 m). Species richness patterns were regressed against climatic variables (temperature, humidity, precipitation and actual evapotranspiration), regional species pool, area and predicted species number of a geometric null model (the mid‐domain effect, MDE). Results The species richness of the 484 recorded species showed a hump‐shaped pattern with elevation with a richness peak at mid‐elevations (c. 1700 m). The MDE was the single most powerful explanatory variable in linear regression models, but species richness was also associated strongly with climatic variables, especially humidity and temperature. Area and species pool were associated less strongly with observed richness patterns. Main conclusions Geometric models and climatic models exclusive of geometric constraints explained comparable amounts of the elevational variation in species richness. Discrimination between these two factor complexes is not possible based on model fits. While overall fits of geometric models were high, large‐ and small‐ranged species were explained by geometric models to different extents. Species with narrow elevational ranges clustered at both ends of the gradient to a greater extent than predicted by the MDE null models used here. While geometric models explained much of the pattern in species richness, we cannot rule out the role of climatic factors (or vice versa) because the predicted peak in richness from geometric models, the empirical peak in richness and the overlap in favourable environmental conditions all coincide at middle elevations. Mid‐elevations offer highest humidity and moderate temperatures, whereas at high elevations richness is reduced due to low temperatures, and at low elevations by reduced water availability due to high temperatures.     

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。     

Species Richness,Spatial Variation,and Abundance of the Soil Seed Bank of a Secondary Tropical Rain Forest1

Despite the importance of the soil seed bank in tropical forest regeneration, little is known about spatial variability in species composition and abundance of seeds stored in the soil. To develop sampling methods for comparative studies, we examined species richness, spatial variation, and abundance of germinants from the soil seed bank in a 16 year old secondary, tropical wet forest at La Selva Biological Station, Costa Rica. Surface soil (10 cm deep, 4.7 cm diameter) was collected at the intersection points of a gridded 1 ha plot (10 × 10-m grid, 121 samples) and in a nested 100 m² subplot (2 × 2-m grid, 36 samples). The 1 ha plot had a density of 4535 seeds/m² with 34 species observed. Based on a series of 100 randomized species accumulation curves, a Michaelis-Menten fit predicted a mean species richness of 36.3 species; the number of observed species was close to the predicted asymptote. A nonparametric, first-order jackknife species richness estimator predicted a species richness of 37.0 species. Eighty-five and 95 percent of the observed species richness is contained, on average, within 41 and 74 pooled samples, respectively. Within the 100 m² nested subplot, a density of 5476 seeds/m² was observed, comprising 26 species with an estimated species richness (Michaelis-Menten fit) of 29.1 species. The jackknife species richness estimator predicted a species richness of 36.7 species. For species richness and abundance of both plots, spatial autocorrelation statistics (Moran's I) were not significantly different from zero at lag distances from 2 to 100 m, indicating a random distribution at these spatial scales. For this site, accurate estimates of species composition depend upon the number of samples collected as well as the spatial distribution of sampling effort. Many small samples distributed over a large area provide greater accuracy and precision for estimating species richness of the soil seed bank.     

环境梯度下蒙古栎群落的物种多样性特征

通过样地法研究了东北地区处于不同经度、纬度和海拔的 13个地点蒙古栎群落的物种丰富度、Gini指数、PIE指数、Shannon指数和 Pielou指数 ,利用相关和回归的统计方法分析了不同地点物种的丰富度指数、Simpson多样性指数和 Shannon多样性指数与各地所处的经度、纬度和海拔的关系。结果发现 :不仅不同地点 (较大尺度 )的物种丰富度和多样性指数均有差异 ,即使在相同的地点 (较小尺度 ) ,物种丰富度及多样性指数也有差异 ,有时还具有很大的差异 ,呈现空间异质性分布的特征 ;因为影响这些多样性指数的环境因子更加复杂 ,不仅受经度、纬度和海拔的影响 ,也受地形、群落的年龄、干扰史等多种生态因子影响。不同地点的物种丰富度与海拔和纬度都具有明显的相关性 (p<0 .0 5 ) ,物种丰富度随海拔和纬度的升高而降低 ,依据显著度的大小可以推测物种丰富度与海拔的相关性比与纬度的相关性更密切 ;蒙古栎群落不同类群的植物种的丰富度具有不同的分布格局 ,木本植物的丰富度与当地纬度具有明显的相关性 (p<0 .0 5 ) ,而与所在地的海拔没有显著的关系 (p>0 .0 5 ) ,而草本植物受海拔的影响更显著 (p<0 .0 5 ) ,而与纬度之间没有显著的关系 (p>0 .0 5 )。群落的 Gini指数、PIE指数、Shannon指数和Pielou指数未发现与海     

Diversity,composition, and structure of tropical dry forests in Central America

Tropical dry forests have been reduced to less than 0.1% of their original expanse on the Pacific side of Central America and are considered by some to be the most endangered ecosystem in the lowland tropics. Plots 1000 m² were established in seven tropical dry forests in Costa Rica and Nicaragua in order to compare levels of species richness to other Neotropical dry forest sites and to identify environmental variables associated with species richness and abundance. A total of 204 species and 1484 individuals 2.5 cm were encountered. Santa Rosa National Park was the richest site with the highest family (33), genera (69), and species (75) diversity of all sites. Species richness and forest structure were significantly different between sites. Fabaceae was the dominant tree and shrub family at most sites, but no species was repeatably dominant based on number of stems in all fragments of tropical dry forest. Central American dry forests had similar species richness when compared to other Neotropical forests. There was no correlation between forest cover within reserves, or precipitation and plant species richness. There was a significant correlation between anthropogenic disturbance (intensity and frequency of fire, wood collection, grazing) and total species richness, tree and shrub species richness, and liana abundance. These results suggest controlling levels on anthropogenic disturbance within reserves should be a high priority for resource managers in Central America. Further research in forest fragments which examine individual and a combination of disturbance agents would help clarify the importance of anthropogenic disturbance on species richness and abundance.