Invasions by alien plant species of the agro-pastoral ecotone in northern China: Species-specific and environmental determinants

The establishment, reproduction, dispersal, and distribution of alien plants are affected by various factors during the transition from being newly introduced in a habitat to being invasive. In the agro-pastoral ecotone of northern China, comprising farmlands and natural grasslands, the biological characteristics of alien plant species were the key intrinsic factors (propagation characteristics and competitive ability), followed by such extrinsic factors as human interference and environmental heterogeneity. Among biological characteristics, the life form may be an important and useful indicator of the invasive ability of a species, and the risk of invasion is greater from alien species that are poisonous, inedible, and have traits that facilitate wide dispersal. Farmlands may serve as initial shelters for alien species, from which they spread into neighbouring habitats, whereas natural grassland may act as a barrier to plant invasions. Management practices detrimental to grasslands, including overgrazing, reclamation, and road construction, often facilitate the invasions; therefore, counter measures such as reseeding and a ban on grazing need special attention. Environmental factors including precipitation, nutrients, prevailing winds, fires, and topography may be other factors that promote or block the process of invasion. In studying ways of preventing or controlling such invasions, alien plants with short life cycle, prolific seed production, and strong competitiveness, deserve particular attention and so do human activities that may damage the environment and fragile habitats.     

放牧强度对华北农牧交错带典型草地土壤化学计量特征的短期影响

放牧是天然草地的主要利用方式之一,不同放牧强度可能通过影响家畜的选择性采食、凋落物输入和微生物的组成及结构等影响草地土壤化学计量特征.本研究通过在华北农牧交错带典型草地一个连续3年(2017-2019年)的生长季放牧试验,测定了土壤全碳(TC)、全氮(TN)、可溶性有机碳(DOC)和可溶性氮(DN)含量,以及土壤微生物...     

Patterns of species richness in dry grassland patches in an agricultural landscape

Eighty-five patches of semi-natural grassland of varying size scattered in a agricultural landscape were investigated for their flora of vascular plants. Relationships between species richness and patch area, spatial isolation and local habitat conditions including heterogeneity were examined. Differences between single species and among groups of species defined by life-history traits were also investigated.
Area was shown to be an important determinant of species richness irrespective of habitat heterogeneity. Isolation in space and habitat heterogeneity also play significant roles. These results are consistent with results from a multitude of studies on fragments of ancient deciduous woodland in northern Europe, They are, however, contradictory to results from previous studies in grasslands within the same region. Seed mass and dispersal syndrome were poor predictors of the degree to which the species were affected by isolation of grassland patches. Seed mass deviation from community median could explain a small percentage of the variation in regional abundance. Logistic regression on species occurrences showed that few species are associated with large patches, and less than half seem to avoid isolated patches.     

北方农牧交错带温性盐碱化草地土壤碳组分对模拟增温的响应机制

中国北方农牧交错带温性盐碱化草地土壤有机碳库对全球气候变暖的响应趋势存在较大不确定性。作为温性盐碱性草地的典型分布区,山西右玉农牧交错带是探索相关研究的理想生境。基于山西农业大学野外观测研究站开顶式气室模拟增温实验平台,通过采集生长旺季土壤样品,探索温性盐碱化草地不同土层有机碳、氮组分对模拟增温的响应与适应机制。结果表明：(1)不同增温处理对土壤有机碳(C)、总氮(N)、颗粒性有机碳(POM-C)和氮(POM-N)、矿物结合态有机碳(MAOM-C)和氮(MAOM-N)、可溶性有机碳(DOC)和氮(DON),以及微生物量碳(MBC)和氮(MBN)等组分无显著影响,但显著降低了MAOM-C/MBC的比值;(2)除土壤可溶性有机碳和微生物量碳外,土壤碳、氮各组分均随土层深度加深而呈现递减趋势,土壤碳、氮各组分之间的比值,除MAOM-N/N和MBC/C外,均随土层深度的增加而呈现显著上升趋势;(3)增温对POM-N/MBN和MAOM-N/MBN的影响与土层深度存在明显的交互效应;(4)不同土层氮组分比值对增温的响应与禾草丰度、杂类草丰度、凋落物量、土壤pH值及土壤含水量等因素有关。其中,凋落物...     

Island biogeography and the species richness of introduced mammals on New Zealand offshore islands

Aim To investigate and establish the significance of various island biogeographic relationships (geographical, ecological and anthropological) with the species richness of introduced mammals on offshore islands. Location The 297 offshore islands of the New Zealand archipelago (latitude: 34–47°S; longitude: 166–179°E). Methods Data on New Zealand offshore islands and the introduced mammals on them were collated from published surveys and maps. The species richness of small and large introduced mammals were calculated for islands with complete censuses and regressed on island characteristics using a Poisson distributed error generalized linear model. To estimate the ‘z‐value’ for introduced mammals on New Zealand islands, least‐squares regression was used [log₁₀ S vs. log₁₀ A]. Results High collinearity was found between the area, habitat diversity and elevation of islands. The island characteristics related to the species richness of introduced mammals differed predictably between large and small mammals. The species richness of introduced large mammals was mostly related to human activities on islands, whereas species richness of introduced small mammals was mostly related to island biogeographical parameters. The ‘z‐value’ for total species richness is found to be expectedly low for introduced mammals. Main conclusions Distance appears to have become ecologically trivial as a filter for introduced mammal presence on New Zealand offshore islands. There is strong evidence of a ‘small island’ effect on New Zealand offshore islands. The species richness of both small and large introduced mammals on these islands appears to be most predominantly related to human use, although there is some evidence of natural dispersal for smaller species. The ecological complexity of some islands appears to make them less invasible to introduced mammals. Some human activities have an interactive effect on species richness. A small number of islands have outlying species richness values above what the models predict, suggesting that the presence of some species may be related to events not accounted for in the models.     

Determinants of species richness in patches of grassland and heathland in Himmerland (Denmark)

Island biogeographical theory predict species richness to increase with habitat area and to decrease with isolation from colonisation sources. This theory has been applied to habitat fragments, and the predictions tested and found valid in empirical studies on fragments of deciduous woodland in northern Europe. However, previous results on fragments of grassland have been ambiguous. In the present study the species richness of vascular plants was investigated in 63 patches of grassland and heathland scattered in an agricultural and forested landscape using multiple linear regression. The relationship between species richness and patch area, isolation and local habitat conditions including heterogeneity were examined. Area was an important determinant of species richness, both in the full data set and in a subset of small habitat patches. In contrast, spatial isolation and habitat heterogeneity were not important factors determining species richness. Differences in soil acidity were accounting for a large proportion of the variance in species richness. This result is probably due to differences in the size of the regional species pools of grasslands with different levels of soil pH.     

北方农牧交错带温性盐碱化草地土壤呼吸对不同形态氮添加和刈割的响应

农牧交错带草地生态系统兼受农业和牧业的影响, 属于脆弱生态系统, 尤其是养分贫瘠的盐碱化草地, 其生态系统结构和功能对外界干扰的响应更加强烈。位于晋西北地区的农牧交错带盐碱化草地, 地理位置独特, 区别于天然牧区草地生态系统。由于毗邻农田, 农业氮肥的过量使用促进了活性氮气体排放, 同时使得农牧交错带草地土壤碳氮循环发生改变。刈割是北方农牧交错草地生态系统的主要管理方式, 为了深入探究氮添加和刈割管理方式对农牧交错带草地碳循环的影响, 进一步厘清该区域草地生态系统的碳动态问题, 该研究设置了一个不同形态氮添加和刈割的裂区实验, 测定土壤呼吸对不同形态氮肥添加和刈割的响应, 为进一步科学管理该区域草地提供可靠的依据。实验样地位于山西省右玉县境内的“山西农业大学农牧交错带草地生态系统野外观测研究站”, 于2017年设置不同形态氮添加和刈割处理, 实验处理包括对照(不刈割和刈割)、尿素添加、缓释尿素添加、刈割+尿素添加、刈割+缓释尿素添加, 每种处理6个重复, 共36个小区。在不同处理条件下测定土壤呼吸速率、土壤温度、土壤水分、土壤微生物生物量、土壤无机氮含量、植物地上和地下生物量, 并计算土壤累积碳排放量及CO₂通量。研究结果表明: (1)短期(2017-2018年)尿素和缓释尿素的添加显著提高了该地区土壤呼吸速率和土壤累积碳排放量。与添加缓释尿素相比, 添加尿素处理下的土壤呼吸速率和累积碳排放量更高; (2)刈割显著降低土壤呼吸速率和累积碳排放量; (3)短期氮添加和刈割的交互作用对土壤呼吸速率没有显著影响。因此, 短期氮添加促进了北方农牧交错带盐碱化草地土壤碳释放, 刈割抑制土壤呼吸, 降低了累积碳排放量, 这可能是由于刈割移除地上植物, 减少了凋落物的输入, 底物减少导致土壤微生物活性降低。但是随着处理时间的延长, 氮添加和刈割对该农牧交错带盐碱化草地土壤碳动态的影响还有待进一步探究和发现。     

The emergy of metabolism in different ecosystems under the same environmental conditions in the agro-pastoral ecotone of northern China

The sustainability of ecosystem productivity and rules governing ecosystem development are important topics of scientific research. The emergy approach is an effective method for investigating these topics, especially when used to evaluate systems that have developed under the same environmental conditions, such as climate and soil. In this paper, emergy differences between terrestrial ecosystems were studied in Guyuan County, a region representative of the agro-pastoral ecotone in Hebei Province, China. A combination of field tests and a questionnaire survey were carried out between June and August 2015. The ecosystems studied included natural grassland, artificial grassland, field crops and commercial crops. These four ecosystems were further subdivided into a total of ten ecosystems. Natural grassland was divided into free-grazing and mowed ecosystems; artificial grassland consisted of oat, Chinese leymus and corn silage; field crops included naked oats, flax and wheat; and commercial crops consisted of cabbage and potatoes. The results showed that the rain input of 4.78 × 10¹⁴ seJ/ha/yr constituted the highest renewable natural resource emergy and that the purchased emergy inputs of the ten ecosystems ranged from 3.53 to 147.67 × 10¹⁴ seJ/ha/yr. Natural resource emergy input was the basic power to maintain the ecosystem, and purchased emergy input was the direct cause of the development of the ecosystems. Groundwater was the most important non-renewable purchased energy for the production of economic crops. The emergy investment ratios (EIR) for potatoes (27.81) and cabbage (19.03) were higher than those of the other ecosystems, but mowed and artificial Chinese leymus grassland had the higher emergy self-sufficiency rates (ESR). Natural grassland, artificial Chinese leymus grassland and traditional grain crops had a low environmental load and high sustainability, whereas potatoes and cabbage had a high environmental load and low sustainability. Overall, rain-fed artificial grassland has a high development potential from the perspective of environment and productivity.     

Earthworms promote greater richness and abundance in the emergence of plant species across a grassland-forest ecotone

Aims Chalk grasslands are subject to vegetation dynamics that range from species-rich open grasslands to tall and encroached grasslands, and woods and forests. In grasslands, earthworms impact plant communities and ecosystem functioning through the modification of soil physical, chemical and microbiological properties, but also through their selective ingestion and vertical transportation of seeds from the soil seed bank. Laboratory experiments showed that seed–earthworm interactions are species specific, but little is known on the impact of seed–earthworm interactions in the field. The overall aim of this study was to better understand seed–earthworm interactions and their impact on the plant community. First we analyzed the composition of seedlings emerging from casts after earthworm ingestion. Then we compared seedling composition in casts to the plant composition of emerging seedlings from the soil and of the aboveground vegetation along four stages of the secondary succession of chalk grasslands.Methods Four stages of the secondary succession of a chalk grassland—from open sward to woods—were sampled in Upper Normandy, France, in February 2010. Within each successional stage (×3 replicates), we sampled the standing vegetation, soil seed bank at three soil depths (0–2, 2–5 and 5–10cm) and earthworm surface casts along transects. Soil and cast samples were water sieved before samples were spread onto trays and placed into a greenhouse. Emerging seedlings were counted and identified. Effect of successional stage and origin of samples on mean and variability of abundance and species richness of seedlings emerging from casts and soil seed banks were analyzed. Plant compositions were compared between all sample types. We used generalized mixed-effect models and a distance-based redundancy multivariate analysis.Important findings Seedling abundance was always higher in earthworm casts than in the soil seed bank and increased up to 5-fold, 4-fold and 3.5-fold, respectively, in the tall grassland, woods and encroached grassland compared to the soil surface layer. Species richness was also higher in earthworm casts than in the soil seed bank in all successional stages, with a 4-fold increase in the encroached grassland. The plant composition of the standing vegetation was more similar to that of seedlings from casts than to that of seedlings from the soil seed bank. Seedlings diversity emerging from casts in the tall and encroached grasslands tended toward the diversity found in woods. Our results indicate that earthworms may promote the emergence of seedlings. We also suggest that the loss of some plant species in the seed bank and the tall grass vegetation in intermediary successional stages modify the local conditions and prevent the further establishment of early-successional plant species.     

The role of patch area and habitat diversity in explaining native plant species richness in disturbed suburban forest patches in northern Belgium

Relatively easy measurable patch characteristics (especially habitat diversity measures) have proven to be valuable indicators of forest plant species richness in forest fragments of relatively undisturbed areas. Urban and suburban forest patches, however, are characterized by a specific landscape ecological context implying that specific processes may influence ecosystem functioning and hence that other abiotic indicators for plant diversity are more appropriate. We studied the relation between functional ecological plant species groups and suburban forest patch characteristics such as patch area, habitat diversity and isolation. Some components of species richness were related to the isolation of the patches. In contrast to previous similar large-scale fragmentation studies in more rural areas, further results stressed the overwhelming importance of patch area relative to habitat variables in determining species richness. This suggests (1) the occurrence of density-dependent species extinction processes in small forest patches; or (2) the existence of external deterministic factors which put a major constraint on species richness in small patches. We tend to support the latter hypothesis and propose forest disturbance and associated black cherry (Prunus serotina Ehrh.) invasion as such a possible external factor. Small forest patches may be more sensitive to disturbance and biological invasion due to various reasons. Hence large forest patches are to be preferred for plant conservation in the suburban area.     

Flea (Siphonaptera) species richness in the Great Basin Desert and island biogeography theory

Numbers of flea (Siphonaptera) species (flea species richness) on individual mammals should be higher on large mammals, mammals with dense populations, and mammals with large geographic ranges, if mammals are islands for fleas. I tested the first two predictions with regressions of H. J. Egoscue's trapping data on flea species richness collected from individual mammals against mammal size and population density from the literature. Mammal size and population density did not correlate with flea species richness. Mammal geographic range did, in earlier studies. The intermediate‐sized (31 g), moderately dense (0.004 individuals/m²) Peromyscus truei (Shufeldt) had the highest richness with eight flea species on one individual. Overall, island biogeography theory does not describe the distribution of flea species on mammals in the Great Basin Desert, based on H. J. Egoscue's collections. Alternatively, epidemiological or metapopulation theories may explain flea species richness.     

Reintroduction of large herbivores restored plant species richness in abandoned dry temperate grassland

Naturalistic grazing by large herbivores is an increasingly practiced way of managing habitats with conservational value. It has the potential to restore and enhance biodiversity, creating self-sustainable environments vital for organisms requiring regular disturbances to moderate and/or reverse successional changes. European bison, Exmoor pony, and Tauros cattle were introduced in 2015 to a former military training area in Milovice, Czech Republic. The prevailing vegetation type is a forest-steppe savanna with Bromus erectus-dominated xeric grasslands mixed with deciduous shrubs and trees. After the cessation of military use, the area was abandoned which led to successional changes, including the dominance of tall grasses, litter accumulation, and bush encroachment. In 2017–2021, we monitored grassland vegetation in 30 grazed permanent plots (2?×?2 m) and 5 control plots representative of ungrazed, abandoned vegetation adjacent to the grazed areas. Naturalistic grazing increased species richness and the cover of forbs, while the cover of grasses and legumes was minimally affected. Grazing increased functional diversity of plant community, promoted a compositional change to small statured species and an increased incidence of red-list species. Seven years of continuous grazing increased the conservation value of this forest-steppe vegetation, a habitat type rapidly declining in Europe.

     

Island theory, matrix effects and species richness patterns in habitat fragments

Island biogeography theory, created initially to study diversity patterns on islands, is often applied to habitat fragments. A key but largely untested assumption of this application of theory is that landscape matrix species composition is non‐overlapping with that of the islands. We tested this assumption in successional old field patches in a closely mowed matrix, and because our patches are appropriately viewed as sets of contiguous habitat units we studied patterns of species richness per unit area. Previous studies at our site did not find that diversity patterns on patch ‘islands’ conformed to predictions of island biogeography theory. Our results indicate that when matrix species are removed from the patch samples, diversity patterns conform better to theory. We suggest that classical island theory remains an appropriate tool to study diversity patterns in fragmented habitats, but that allowances should be made for spill‐over colonization of ‘islands’ from the ‘sea’.     

Changes in plant species richness in a calcareous grassland following changes in environmental conditions

This study presents results of a three and a half year experiment in a calcareous grassland on the Baltic island of Öland, south-eastern Sweden. Environmental conditions, viz. nutrient regime, water availability, light conditions and grazing pressure were changed in a randomised experimental design with ten different combinations of these factors, including a control. Weather conditions during the study period fluctuated considerably. Only the combination of reduced light and increased soil nutrient availability decreased species richness significantly, presumably by inducing asymmetric competition for light by competitively strong species. Physical space also became less available for weaker competitors. In cases where only nutrients or only water were supplied in sufficient amounts, competitive exclusion was inhibited by either low nutrient or low water availability. Under conditions which combined high levels of nutrient and water supply with regular disturbance (i.e. simulated grazing) high species richness was maintained. Although there was a significant negative correlation between species richness and amount of above-ground biomass, the number of species was not significantly reduced in the treatment which combined fertilisation and irrigation. One reason for this may be that the heterogeneous light regime in this treatment still allowed species persistence and coexistence.     

Soil carbon and nitrogen stores and storage potential as affected by land-use in an agro-pastoral ecotone of northern China

Equilibrium carbon stock is the result of a balance between inputs and outflows to the pool. Changes in land-use are likely to alter such balance, resulting in different carbon stores under different land-use types in addition to the impacts of global climate change. In an agro-pastoral ecotone of Inner Mongolia, northern China, we investigated productivity and belowground carbon and nitrogen stores under six different types of land-uses, namely free grazing (FG), grazing exclusion (GE), mowing (MW), corn plantation (CP), fallow (FL), and alfalfa pasture (AP), and their impacts on litter and fine roots in semiarid grassland ecosystems. We found that there were great variations in aboveground net primary production (ANPP) across the six land-use types, with CP having markedly high ANPP; the FG had significantly reduced soil organic carbon (SOC) and nitrogen stores (SON) to 100 cm depth compared with all other types of land uses, while very little litter accumulation was found on sites of the FG and CP. The top 20 cm of soils accounted for about 80% of the root carbon and nitrogen, with very little roots being found below 50 cm. About 60% of SOC and SON were stored in the top 30 cm layer. Land-use change altered the inputs of organic matters, thus affecting SOC and SON stores accordingly; the MW and GE sites had 59 and 56% more SOC and 61% more SON than the FG. Our estimation suggested that restoring severely degraded and overgrazed grasslands could potentially increase SOC and SON stores by more than 55%; conversion from the native grasses to alfalfa could potentially double the aboveground biomass production, and further increase SOC and SON stores by more than 20%. Our study demonstrated significant carbon and nitrogen storage potential of the agro-pastoral ecotone of northern China through land-use changes and improved management in the context of mitigating global climate change.     

Scale-dependent effects of land use on plant species richness of mountain grassland in the European Alps

Traditionally managed mountain grasslands in the Alps are species‐rich ecosystems that developed during centuries of livestock grazing. However, changes in land use including fertilisation of well accessible pastures and gradual abandonment of remote sites are increasingly threatening this diversity. In five regions of the Swiss and French Alps we assessed the relationship between land use, soil resource availability, cover of the unpalatable species Veratrum album, species richness and vegetation composition of mountain grasslands across four spatial scales ranging from 1 to 1000 m². Mean species richness and the increase in the number of species with increasing area were lower in intensively grazed, fertilised pastures than in traditional pastures or in abandoned pastures. Species composition of abandoned pastures differed from that of the other management types. Plant species richness was influenced by different factors at different spatial scales. At the 1 m² scale, plant species richness was negatively related to soil nitrate and influenced by the cover of V. album, depending on land use: species richness and cover of V. album were negatively correlated in abandoned pastures, but positively correlated in fertilised grasslands. At the 1000 m² scale, a negative effect of fertilization on richness was evident. These results indicate that at small scales species richness in mountain grasslands is determined by competition for light, which should be more important if nutrient availability is high, and by positive and negative interactions with unpalatable plants. In contrast, species richness at the large scale appears to be mainly influenced by land use. This result emphasizes the importance of studying such inter‐relationships at multiple scales. Our study further suggests that the maintenance of the traditional land use scheme is crucial for the conservation of plant species richness of mountain pastures as both intensification and abandonment changed species composition and reduced plant species diversity.     

Soil nutrients and variation in biomass rather than native species richness influence introduced plant richness in a semi-arid grassland

Several different hypotheses account for the success of introduced species in new environments. Experimental studies show a negative native-exotic richness relationship (NERR), while observational studies suggest that this relationship is usually positive. Increased resource availability and environmental variation can also enable introduced species to establish in new environments. We conducted an observational study in a semi-arid grassland in the Thompson-Nicola District of British Columbia to examine the biotic and abiotic factors that account for variation in introduced and native species richness.In each of 12 sites, an 8 × 8 m area was set up, containing 64, 1-m² plots. We identified and categorized plant species in each site into introduced and native species. We tested the relationship between introduced species richness and native species richness at the 1-m² sampling grain and at sampling grains up to 64 m². We also analysed the relationship between native and introduced species, and within-plot biomass, and between native and introduced species and variation in biomass. For a representative subset of four sites, we tested the relationship between introduced and native species richness and nitrogen, phosphorus and potassium.We found no NERR at the 1 m² sampling grain, nor for the other sampling grains up to 64 m². Introduced species richness increased with phosphorus and nitrogen availability, and was also positively related to biomass heterogeneity.Our results indicate that introduced species richness in these grasslands is likely influenced by phosphorus and nitrogen, and by variation in vegetation biomass, but not by native species. More non-native plants are likely to occupy nutrient-rich plots compared to nutrient-poor plots in these grasslands. Variation in biomass can leave gaps for the establishment of introduced species. These results should inform management considerations for the control of invasive species to optimize preservation of grasslands.