Diversity, distribution and biogeography of testate amoebae in China: implications for ecological studies in Asia

Testate amoebae are a group of shelled protozoa that occur in high density populations in wet environments. More than 1900 testate amoebae species or subspecies have been reported in published literature over the last 200 years, from many regions of the world. Testate amoebae are classified as Lobosea or Filosea respectively, according to the presence of lobose or filiform pseudopodia. Testate amoebae have proved an interesting group of indicator organisms in palaeoenvironmental studies and have also been used as bioindicators of human impact on ecosystems. Until recently, the testate amoebae of China were unknown to most western scientists, but our knowledge has improved greatly over the past 20 years. This paper summarizes the testate amoebae research in China along with relevant data from other countries in Asia, and provides the necessary context for future research.     

Diversity and biogeography of testate amoebae

Testate amoebae are amoeboid protists inhabiting a test (shell). They occur globally in soils, wetlands and freshwater, especially peats and mosses. They are of ancient origin, dating from at least the Mesozoic, with possible ancestors as old as the Neoproterozoic. Approximately 2,000 taxa have been described—a number which could easily rise to 4,000 with comprehensive recording. Whilst many protists appear to be cosmopolitan as morphospecies, some of the larger testate species (exceeding 100 μm) have long been considered, controversially, to be geographically restricted. Definitive conclusions have often been confounded by gaps in distributional data and misidentification. Recent increases in recording from previously little known regions, and the rise of molecular taxonomy, have started to resolve outstanding issues—processes still far from complete. Accordingly, biogeographical studies have concentrated on “flagship” species—those which can be identified with certainty and are sufficiently recorded to determine their ecological ranges. Apodera vas (Certes) has been proved to be largely restricted to the Gondwanaland continents and sub-Antarctic islands, but absent from the Holartic despite the availability of much suitable habitat. An early analysis postulated a Mesozoic origin of the species and a distribution influenced by continental drift. Recent molecular evidence could imply a later origin. Either way, its current distribution is clearly influenced by the pattern of global wind currents and lack of lowland tropical habitat. By contrast a “Gondwana-tropical” group of species appears to be restricted to latitudes unaffected by glaciation. Instances of local endemism, such as restriction to a single island, are also known, which await molecular evidence for substantiation. Special Issue: Protist diversity and geographic distribution. Guest editor: W. Foissner.     

The testate amoebae of New Zealand: A checklist,identification key and assessment of biogeographic patterns

New Zealand (NZ) is a well-known hotspot of biodiversity and endemism for macroscopic organisms, but its microbial diversity is comparatively poorly documented. We assembled all records on NZ testate amoebae published since the early 20th century and present a comprehensive taxonomic checklist for NZ. Testate amoebae are reported from six major habitat types across both the North and South Islands of NZ, but the sampling effort is ecologically and geographically biased in favour of wetlands and the South Island. As a result, 93% of all 128 morphotypes recorded in NZ occur in wetlands, 28% are restricted to the South Island, and diversity is greater at higher latitudes. Around 50% of morphotypes have a broad latitudinal distribution across the NZ mainland, whereas 15% have narrow latitudinal ranges. Future research should aim to broaden the geographical and ecological ranges. We predict that our list of NZ testate amoebae will expand substantially with future work, and that the latitudinal diversity gradient will be inverted. We also introduce an interactive, fully illustrated, online Lucid key for the rapid identification of NZ testate amoebae. As many morphospecies are cosmopolitan, this key provides a useful tool for testate amoebae identification in other parts of the world.     

Biogeography and comparative ecology of testate amoebae inhabiting Sphagnum-dominated peatlands in the Great Lakes and Rocky Mountain regions of North America

Testate amoebae (Protozoa: Rhizopoda) are common inhabitants of peatlands. Strong relationships between community composition and substrate moisture in Sphagnum‐dominated peatlands have made them particularly useful as hydrological proxies in environmental and palaeoenvironmental research. However, stability of these relationships in geographical space is important for widespread applicability. In this study, we compared testate amoeba communities inhabiting Sphagnum‐dominated peatlands of the Great Lakes and Rocky Mountain regions of North America. Our primary objectives were to describe patterns of community composition in the two regions, develop hypotheses to explain differences, and determine if taxa occupy similar ecological niches with respect to substrate moisture in both places. Our results indicated that testate amoeba communities are relatively different in the two regions, and these differences are probably caused by differences in climate and peatland trophic status, although other factors may also play a role. However, many taxa do occur in both regions and these taxa had comparable moisture preferences in each region, suggesting that the ecological niches of taxa with respect to substrate moisture are similar even within communities of relatively different composition.     

Dynamics in space and time of four testate amoebae (Difflugia spp.) co-existing in the zooplankton of a reservoir in southern China

We studied a long time series of the dynamics in space and time of four species of Difflugia (thecamoebae) that co-exist in the pelagic plankton of Liuxihe Reservoir, an oligo-mesotrophic impoundment in southern China, during 8–9 months (“summer” form March to November), and retreat to the benthos during the rest of the year (“winter”). We discuss the reasons for the winter retreat, and suggest that predator evasion may be involved, although temperature-linked physiological effects (like the rate of gas bubble production) appear more probable. Clear diel vertical migration of Difflugia was not observed, but patchiness was common. We found no evident lake edge-effects in the spatial pattern either, but the abundances were strongly influenced by trophic conditions and increased by up to one order of magnitude in the upstream, eutrophic sections of the reservoir.     

Comparative ecology of vascular plant,bryophyte and testate amoeba communities in four Sphagnum peatlands along an altitudinal gradient in Switzerland

Monitoring tools are needed to assess changes in peatland biotic communities and ecosystem functions in response to on-going climate and other environmental changes. Although the responses of soil organisms and plants to ecological gradients and perturbations do not always correlate, peatland monitoring is mainly based on vegetation surveys. Testate amoebae, a group of protists, are important contributors to carbon and nitrogen cycling in organic soils and are useful bioindicators in peatland ecology and paleoecology. There is however little comparative data on the value of testate amoebae, vascular plants and bryophytes as bioindicators of micro-environmental gradients in peatlands.We compared the relationships of testate amoebae, bryophytes, and vascular plants with soil temperature, water table depth, micro-habitats and the carbon and nitrogen content of Sphagnum mosses in four peatlands along a 1300 m altitudinal gradient in Switzerland. We used the full diversity of vascular plants and bryophyte but only a selection of ten easily identifiable testate amoeba morpho-taxa (i.e. species or species-complexes).Indirect and direct gradient ordinations, multiple factor analysis (MFA) and transfer function models for inferring water table depth showed that a selection of ten testate amoeba taxa are more powerful (% variance explained in RDA) and accurate (discrimination among habitats) indicators of local conditions (micro-habitat type, water table depth and Sphagnum C/N ratio) than the vegetation (vascular plants and bryophytes either individually or combined and considering the full diversity).Our study showed that a limited list of ten easily identifiable testate amoeba taxa have higher bioindication value than the full bryophytes and vascular plants. Furthermore, testate amoebae can be analyzed on samples collected at any season (accessibility allowing and if precise sampling sites are well marked) – a clear advantage for biomonitoring and can be used to infer past changes from the peat record at the same taxonomic resolution. This simple approach could therefore be very useful for biomonitoring of peatlands.     

Abrupt ecological changes in the last 800 years inferred from a mountainous bog using testate amoebae traits and multi-proxy data

Mountainous peatlands of Western Sudetes are considered a unique habitat in Central Europe. The region contains one of the largest raised bog complexes in temperate Europe and is thus of great importance for biodiversity conservation. In this first high-resolution study from this region we use long-term ecological data to assess how these mountain wetland ecosystems responded to anthropogenic impacts and climate change. We used testate amoebae morphological traits, micro- and macroscopic charcoal, pollen and plant macrofossils to reconstruct the history of peatland development over 800 years, illustrating shifts in its development and fire dynamics. Five hydrological stages of peatland development were recognized. Testate amoebae morphological traits reflected several abrupt ecological changes linked to anthropogenic modifications of landscape openness. A shift towards mixotrophic taxa, linked to hydrological change or shrubs expansion and shading, preceded aperture position change, which was associated to dust input through surrounding deforestation and simultaneous water-table increase. Fire reconstruction revealed increasing burning together with intensifying human activity, including the expansion of a nearby settlement. This study confirms the potential of testate amoeba communities and the use of morpho-functional traits as indicators of ecological effects of land-use change over long-temporal scales.     

Community assembly, stability and signatures of dynamical constraints on food web structure

To understand the dynamics of natural species communities, a major challenge is to quantify the relationship between their assembly, stability, and underlying food web structure. To this end, two complementary aspects of food web structure can be related to community stability: sign structure, which refers to the distributions of trophic links irrespective of interaction strengths, and interaction strength structure, which refers to the distributions of interaction strengths with or without consideration of sign structure. In this paper, using data from a set of relatively well documented community food webs, I show that natural communities generally exhibit a sign structure that renders their stability sensitive to interaction strengths. Using a Lotka-Volterra type population dynamical model, I then show that in such communities, individual consumer species with high values of a measure of their total biomass acquisition rate, which I term “weighted generality”, tend to undermine community stability. Thus consumer species’ trophic modules (a species and all its resource links) should be “selected” through repeated immigrations and extinctions during assembly into configurations that increase the probability of stable coexistence within the constraints of the community's trophic sign structure. The presence of such constraints can be detected by the incidence and strength of certain non-random structural characteristics. These structural signatures of dynamical constraints are readily measurable, and can be used to gauge the importance of interaction-driven dynamical constraints on communities during and after assembly in natural communities.     

北京小型湖泊底栖无脊椎动物群落结构特征与营养状况的研究

1987年间逐月调查分析了北京6个小型湖泊底栖无脊椎动物群落结构特征。通过指示种、优势种、种类、数量、生物量、Shannon多样性指数及其与水质的关系分析,前3个湖相当于中营养湖泊,后3个湖相当于中-富营养湖泊。梨形环稜螺(Bellamya purificata)和粗腹摇蚊幼虫(Pelopia sp.)可作为北京6个小湖环境质量的指标生物,前者指示水质较好,后者指示水质较差。Shannon多样性指数值(H′)与湖水BOD、COD、TP、TN含量之间关系比较密切,用该指数并参考指示生物特征评价此类型浅水小湖营养状况是可行的。     

Testate amoebae and tintinnids as spatial and seasonal indicators in the intertidal margins of Guadiana Estuary (southeastern Portugal)

The present study gathers pioneering taxonomical and community data on testate amoebae and agglutinated tintinnids, analyzing their seasonal and spatial distribution patterns in the mesotidal system of the Guadiana Estuary, southeastern Portugal. To evaluate both groups’ potential as bioindicators in climate monitoring and paleoenvironmental reconstructions, their abundance, diversity and living proportions were compared to elevation in relation to mean sea-level, marine influence and to periods of elevated water levels. The distributions of testate amoebae and tintinnid total assemblages were also related to major physicochemical variables by means of multivariate analysis.From 49 surface sediment samples collected in winter and summer 2010, 17 species (25 strains) of testate amoebae were identified (in the 63 μm fraction). The Centropyxidae, mainly represented by Centropyxis aculeata, Centropyxis arcula and Centropyxis constricta, had the greatest density (71% of total individuals), while the Difflugidae represented the most species (82% of total species). Higher diversities and densities were observed in winter in the upper estuary, where salinity is negligible, and in the middle estuary where they are concentrated at the lower levels of the intertidal zone. Few or no individuals were observed in the highest marsh zones. In winter, dead testate amoebae were dominant, with empty tests accumulating in the sediments from post-mortem transport by high river discharge. In summer, an increase in the living fraction is observed, with living testate amoebae along the entire estuary, which could indicate that some species are able to live in the lower reaches of the estuary.Along with the testate amoebae, two species of agglutinated tintinnids (>63 μm), Stenosemella ventricosa and Tintinnopsis cf. lata, were abundant in the sediments. Their highest abundances were observed in summer. Living individuals were only recorded in summer, mainly in the vicinities of freshwater and sewage outflows, where elevated nutrient concentrations may be expected.The present study demonstrates that both testate amoebae and tintinnids have well defined patterns in their temporal and spatial distribution, offering high bioindicator potential in environmental/climate monitoring studies as well as in paleoenvironmental reconstructions.     

基于稳定同位素技术的海州湾海洋牧场食物网基础及营养结构的季节性变化

基于稳定同位素技术对2015年春季海州湾海洋牧场海域采集的中小型生物消费者,包括鱼类、虾类、蟹类、头足类、螺类和双壳类等与其潜在碳源样品进行分析,利用IsoSource模型计算该海域消费者碳源贡献率,并对2014年夏季生物学样品与2015年春季样品比较,分析食物网营养结构的季节性变化,根据稳定同位素测定结果绘制二维双标图,计算出6种营养结构的量化指标.结果表明: 2015年春季海州湾海洋牧场海域消费者的δ¹³C值范围为-18.9‰～-17.1‰,3种潜在碳源[浮游植物、悬浮颗粒有机物(POM)、沉积物(SOM)]的δ¹³C值范围为-18.1‰～-23.4‰,根据模型计算得出浮游植物对消费者的平均碳源贡献最大,为80.8%,其余依次为SOM和POM,分别为10.8%和8.4%.2014年夏季生物样品与2015年春季样品的δ¹³C值存在显著差异,而δ¹⁵N值无显著性差异;6种量化指标表明群落营养结构存在季节性差异, 2014年夏季的δ¹³C比值范围(CR)、总面积(TA)、平均最邻近距离(NND)和平均最邻近距离标准差(SDNND)均大于2015年春季,δ¹⁵N比值范围(NR)和平均离心距离(CD)无明显变化,夏季群落营养结构冗余度小于春季,且食源多样性水平高于春季,存在季节差异.     

Structure and dynamics of subalpine forests in the Wang Lang Natural Reserve, Sichuan, China

Population structure (size, age, spatial patterns) and radial growth patterns are used to analyze regeneration patterns of Abies faxoniana, Betula albosinensis, Betula utilis, Larix potaninii, Picea purpurea, and Sabina saltuaria and reconstruct disturbance history in 8 subalpine forest stands in Wang Lang Natural Reserve, Sichuan, China. In old-growth stands tree regeneration occurs in tree-fall gaps whereby A. faxoniana, Betula sp., P. purpurea, and S. saltuaria persist at stand scales by gap-phase regeneration. Clump sizes of young populations are similar to canopy gap sizes but clumps sizes vary among species. Young Betula patches are larger than those of A. faxoniana suggesting that gap-partitioning by size contributes to species coexistence in mixed stands. Picea purpurea and S. saltuaria are longer lived than A. faxoniana which may compensate for lower recruitment and prevent their replacement by A. faxoniana. Tree regeneration and community structure are also influenced by the understory bamboo Fargesia denudata. Seedlings, saplings, and shrub density all decline with an increase in bamboo cover. Species that regenerate in old-growth forest also regenerate after flooding as do species that establish only on bare substrates (i.e. Larix potaninii, Prunus sp.). Structural and compositional patterns in Wang Lang forests are a reflection of disturbance history, canopy species life history attributes such as dispersal ability, shade tolerance, growth rates, and longevity, and competition of trees and shrubs with understory bamboos.     

杭州湾南岸大桥建设区域潮间带大型底栖动物功能群及营养等级的季节动态

2006年5月至2007年1月在杭州湾南岸跨海大桥附近潮滩共采集到32种大型底栖动物,根据其食性类型划分为5种功能群。用功能群方法对这些物种生境变化的关系进行了分析,结果表明:大型底栖动物各多样性指数在潮带间差异显著,季节间则只有Margalef种类丰度(S)和Shannon-Wiener指数(H′)存在显著性差异;各功能群密度在季节间均不存在显著性差异,而在潮带间除了浮游生物食者外,其它类群均存在显著性差异;肉食者的种类数在潮带间不存在显著性差异,而在季节间存在显著性差异,其它功能群刚好相反。滩涂大型底栖动物群落主要包括4个营养级别,各营养级的构成密度在季节间差异不显著,在潮带间则差异显著;各营养级种类数是处于第四级的肉食者季节间差异显著,潮带间变化不显著,处于第二、三营养等级的功能群则正好相反。底栖动物功能群多样性综合反映了杭州湾南岸大桥建设区域潮间带大型底栖动物群落变化情况。     

三江平原沟渠网络结构对区域景观格局的影响

农业开发进程中产生的沟渠作为农业景观中廊道内部的动因主体,会对周围自然生态系统产生不同程度的影响;同时沟渠网络的形成,又使得这种影响作用更加深入和复杂.以根据三江平原建三江管局辖区2004年的Landsat-TM卫星影像解译的土地利用现状图以及沟渠分布水利图为基础资料,在地理信息系统(GIS)支持下,选择3个30×30km2的取样区为研究对象,分析了各样区的景观结构特征和沟渠网络结构.在此基础上,用典范相关分析(CCA)定量研究了沟渠网络结构对于区域景观格局的影响.结果表明:沟渠网络的结构对区域景观格局有显著影响,沟渠密度越大,网络结构越复杂,则区域景观中斑块数目越多,平均斑块面积越小;同时农业用地所占比例越大,自然湿地所占比例越小.通过各沟渠网络结构指标与CCA排序轴的相关性分析发现,对区域景观格局影响最大的指标是沟渠密度,其次是线点率和环度,而反映沟渠网络连通性的连通度指标影响相对较小.     

Intra-habitat heterogeneity of microbial food web structure under the regime of eutrophication and sediment resuspension in the large subtropical shallow Lake Taihu,China

Planktonic microbial community structure and classical food web were investigated in the large shallow eutrophic Lake Taihu (2338 km², mean depth 1.9 m) located in subtropical Southeast China. The water column of the lake was sampled biweekly at two sites located 22 km apart over a period of twelve month. Site 1 is under the regime of heavy eutrophication while Site 2 is governed by wind-driven sediment resuspension. Within-lake comparison indicates that phosphorus enrichment resulted in increased abundance of microbial components. However, the coupling between total phosphorus and abundance of microbial components was different between the two sites. Much stronger coupling was observed at Site 1 than at Site 2. The weak coupling at Site 2 was mainly caused by strong sediment resuspension, which limited growth of phytoplankton and, consequently, growth of bacterioplankton and other microbial components. High percentages of attached bacteria, which were strongly correlated with the biomass of phytoplankton, especially Microcystis spp., were found at Site 1 during summer and early autumn, but no such correlation was observed at Site 2. This potentially leads to differences in carbon flow through microbial food web at different locations. Overall, significant heterogeneity of microbial food web structure between the two sites was observed. Site-specific differences in nutrient enrichment (i.e. nitrogen and phosphorus) and sediment resuspension were identified as driving forces of the observed intra-habitat differences in food web structure.     

Stand structure and vegetation dynamics of a subalpine wooded fen in Rocky Mountain National Park,Colorado

Abstract. The tree population size structure and relationship between tree diameter and age were examined in a subalpine fen and surrounding Picea-Abies forest in northern Colorado. The fen grades from a sedge fen, through an ecotone, to a treed fen. Tree growth rate varies across the vegetational gradient, with the sedge fen having the slowest growth, and the upland forest having the fastest growth. Differences in growth rate are related to the average size of peat hummocks, with areas containing tall hummocks exhibiting the highest tree growth rates. Size structures display the characteristic reverse-J distribution generally indicative of stable populations, but forest vegetation is expanding into the open regions of the fen, and within the treed fen an increase in Abies lasiocarpa is occurring. These changes are primarily attributed to a positive feedback situation wherein the fen's surface is built up by peat accumulation. Distinct hummocks form first on the open fen but then coalesce to form raised peat islands in the treed fen. This new substrate provides habitat with a comparatively low water table and allows the growth of mesophytic forest vegetation. A pathway for this vegetational development is proposed.     

Decomposition and nutrient dynamics of marsh litter in the Sanjiang Plain, Northeast China

The litterbag technique was used to study the decomposition and nutrient dynamics of marsh litter in the four communities, Carex pseudocuraica (C.pa), C. lasiocarpa (C.la), Deyeuxia angustifolia (D.aa), and D. angustifolia-Shrub (D.aa-Srb), in Sanjiang Plain, Northeast China. Decomposition was divided into two periods in the first year, with the mass loss ranging from 11.7% to 31.4% of the initial mass during summer and autumn, accounting for more than 75% of the annual loss. The decomposition rates ranged from 0.000 612 to 0.000 945 d^?1 depending on the depth of the flooding and its duration, and differed significantly in each community. The litter decomposed faster in communities with deeper and perennial flooding than in those with shallow and seasonal flooding. The initial ratios of C:N and C:P were also different among the four litter types, but these differences had no impact on the decomposition rates, suggesting that the main factor influencing the decomposition rates of marsh litter was the flooding status rather than the litter quality. The N concentrations in C.pa and C.la almost continuously increased over time, with their final values being 2.8 and 2.4 times higher than the initial ones, respectively. However, the nutrient dynamics in D.aa and D.aa-Srb offered another pattern, sharply falling in the first month and then gradually rising, with the values at the end of the experiment being close to those at the beginning. The litter accumulated substantial amounts of N in C.pa and C.la, while net N release from the litter was observed in both D.aa and D.aa-Srb. The difference may be caused by microorganisms' demand for nutrition, and then limited by the C:N ratios of litter and the availability of nitrogen from the soil and marsh water. In contrast with N dynamics, P concentrations of all the litter types apparently decreased during the first month, and then continued to decline in C.pa, remained constant in C.la and D.aa and increased slightly in D.aa-Srb. At the end of the experiment, the P concentrations decreased, respectively, by 56%, ?5%, 47% and 24% of the initial values of C.la, C. pa, D.aa and D.aa-Srb. The net P release was observed in all marsh litter over 480 days of decomposition and the intensity of the P release was different amongst communities, which may be regulated by ratios of initial C:P. The results suggested that in the marsh with the N limitation, litter tended to accumulate N and release P during decomposition and the intensity of accumulation or release was closely related to the initial C:N and C:P ratios.