Facilitation and interference in subalpine meadows of the central Caucasus

Abstract. We studied the effects of neighbours on the biomass of seven randomly chosen species in species‐rich sub‐alpine meadows in the central Caucasus Mountains by comparing the performance of plants with neighbours removed experimentally to that of paired plants with their neighbours left intact. In most cases the removal of neighbours led to significant increases in vegetative and total above‐ground biomass implying the species were limited by competition. However, the neighbour removal led also to an increased leaf wilting for target plants, as well as to strong decline in reproductive effort for some species. We hypothesise that competition may be the prevailing type of interaction in species‐rich sub‐alpine meadow communities, but competitive effects on vegetative production may be balanced, if not outweighed, by facilitation, at least for some species. Such a balance may enhance species coexistence in communities.     

Multiple resource limitation: nonequilibrium coexistence of species in a competition model using a synthesizing unit

During the last two decades, the simple view of resource limitation by a single resource has been changed due to the realization that co-limitation by multiple resources is often an important determinant of species growth. Hence, the multiple resource limitation hypothesis needs to be taken into account, when communities of species competing for resources are considered. We present a multiple species–multiple resource competition model which is based on the concept of synthesizing unit to formulate the growth rates of species competing for interactive essential resources. Using this model, we demonstrate that a more mechanistic explanation of interactive effects of co-limitation may lead to the known complex dynamics including nonequilibrium states as oscillations and chaos. We compare our findings with earlier investigations on biological mechanisms that can predict the outcome of multispecies competition. Moreover, we show that this model yields a periodic state where more species than limiting complementary resources can coexist (supersaturation) in a homogeneous environment. We identify two novel mechanisms, how such a state can emerge: a transcritical bifurcation of a limit cycle and a transition from a heteroclinic cycle. Furthermore, we demonstrate the robustness of the phenomenon of supersaturation when the environmental conditions are varied.     

Seasonal variation in exploitative competition between honeybees and bumblebees

Honeybees (Apis mellifera) and bumblebees (Bombus spp.) often undergo exploitative competition for shared floral resources, which can alter their foraging behaviour and flower choice, even causing competitive exclusion. This may be strongest in summer, when foraging conditions are most challenging for bees, compared to other times of the year. However, the seasonal dynamics of competition between these major pollinator groups are not well understood. Here, we investigate whether the strength of exploitative competition for nectar between honeybees and bumblebees varies seasonally, and whether competitive pressure is greatest in summer months. We carried out experimental bee exclusion trials from May to late September, using experimental patches of lavender, variety Grosso, in full bloom. In each trial, we compared the numbers of honeybees (HB) foraging on patches from which bumblebees had been manually excluded (bumblebee excluded, BBE) versus control (CON) patches, HB(BBE-CON). This measure of exploitative competition varied significantly with season. As expected, mean HB(BBE-CON) was significantly greater in summer trials than in spring or autumn trials. This was despite high nectar standing crop volumes in BBE patch flowers in spring and autumn trials. Mean HB(BBE-CON) was not different between spring and autumn trials. Our results show that nectar competition between honeybees and bumblebees varies seasonally and is stronger in summer than spring or autumn, adding to current understanding of the seasonality of resource demand and competition between bee species. This information may also help to inform conservation programs aiming to increase floral resources for bees by showing when these resources are most needed.     

The nutrient-load hypothesis: patterns of resource limitation and community structure driven by competition for nutrients and light

Resource competition theory predicts that the outcome of competition for two nutrients depends on the ratio at which these nutrients are supplied. Yet there is considerable debate whether nutrient ratios or absolute nutrient loads determine the species composition of phytoplankton and plant communities. Here we extend the classical resource competition model for two nutrients by including light as additional resource. Our results suggest the nutrient-load hypothesis, which predicts that nutrient ratios determine the species composition in oligotrophic environments, whereas nutrient loads are decisive in eutrophic environments. The underlying mechanism is that nutrient enrichment shifts the species interactions from competition for nutrients to competition for light, which favors the dominance of superior light competitors overshadowing all other species. Intermediate nutrient loads can generate high biodiversity through a fine-grained patchwork of two-species and three-species coexistence equilibria. Depending on the species traits, however, competition for nutrients and light may also produce multiple alternative stable states, suppressing the predictability of the species composition. The nutrient-load hypothesis offers a solution for several discrepancies between classical resource competition theory and field observations, explains why eutrophication often leads to diversity loss, and provides a simple conceptual framework for patterns of biodiversity and community structure observed in nature.     

Functional diversity and redundancy of subalpine meadows subjected to anthropogenic disturbances

人为干扰下的亚高山草甸功能多样性和功能冗余 人为干扰程度的加剧会引起植物群落物种多样性和功能多样性的改变,并降低生态系统的稳定性。较高水平的功能冗余对于物种损失引起的生态系统功能和稳定性的改变起到了重要的缓冲作用。然而,关于亚高山草甸植物群落功能冗余对干扰的响应的研究相对较少。本研究的目的是,确定人为干扰下不同退化程度的亚高山草甸,其植物群落功能多样性和功能冗余对干扰的响应特征,及不同物种对功能冗余的贡献程度及其在干扰梯度上的变化规律。研究区位于青藏高原东南缘玉龙雪山上的亚高山草甸。本研究识别出4类干扰强度(无干扰、弱干扰、中度干扰和重度干扰),测定了干扰梯度上的植物物种组成、土壤性质和5个关键的植物功能性状等指标,进一步计算了Simpson多样性指数、功能多样性指数(RaoQ)、功能冗余度、群落的植物功能性状加权平均值(CWM)以及物种水平的功能冗余等指数。研究结果表明,功能多样性和功能冗余在干扰梯度上呈现单峰变化规律,即二者分别在中度干扰、弱干扰下达到最大值。物种多样性在干扰梯度上整体呈下降趋势。随着干扰程度的增加,采取保守生长策略并具有相关性状特征(如较低比叶面积和较高的干物质含量)的物种的丰富度降低,采取资源获取策略并具有相关性状(如低矮植株、较高的比叶面积和较低的干物质含量)的物种增加。不同物种的功能冗余程度存在差异。值得注意的是,一些物种在群落中扮演着独特的功能角色,如无干扰梯度下的网脉橐吾(Ligularia dictyoneura)以及中度和重度干扰梯度下的滇西委陵菜(Potentilla delavayi)。     

Nutrient limitation along a productivity gradient in wet meadows

Conservation management in meadows often focuses on reducing soil fertility and consequently community productivity as to promote and sustain species-rich vegetations. The productivity level to which nutrients are limiting growth is, however, unclear, as well as the relationship between productivity and the type of nutrient limitation. We carried out a fertilization experiment with N, P and K in six annually mown meadows along an aerial phytomass gradient (200–650 g m^–2). All meadows were found to be growth-limited by nutrients. Low-productive meadows were N-limited, or N+P co-limited, whereas our higher productive meadows were co-limited by a combination of N, P and/or K. The results from our experiments were compared with the results from 45 other fertilization experiments with N, P and K in grasslands and wetlands (aerial phytomass range 50–1500 g m^–2). Our results were consistent in nitrogen being the most frequent (co)-limiting nutrient, and regarding the equal frequence of occurrence of P (co)-limitation and K (co)-limitation (both in ca. 25–30% of all sites). Co-limitation occurred more often in our sites than in the other experiments. There was no clear relationship between aerial phytomass and type of nutrient limitation, except that K (co)-limitation only occurred at sites with phytomass above 200 g m^–2, and P (co)-limitation below 600 g m^–2. A comparison of productivity and nutrient concentrations in aerial phytomass among two years indicated that the type of nutrient limitation is not a static site characteristic but may vary with dynamic environmental conditions such as soil wetness; drought seems to enhance N-availability which may induce P- and K-limitation.     

Influence of local and landscape factors on bumblebees in semi-natural meadows: a multiple-scale study in a forested landscape

Understanding the effects of local and landscape factors on bumblebees is relevant for the conservation of this group of pollinators. Bumblebees have been well-studied in agricultural landscapes of Western Europe, Asia and North America, but few studies have been developed on bumblebees in forest-dominated landscapes of Eastern Europe. We developed this study in 22 semi-natural meadows located in a patchy forested landscape of Estonia. We investigated the influence of habitat characteristics and landscape factors (calculated at four spatial scales: 250, 500, 1,000 and 2,000 m radius) on the total species richness and abundance of bumblebees. Correlation analysis, partial least squares (PLS) and stepwise forward-selection multiple regression analysis were applied in this study. The presence of a high diversity of flowering plants in semi-natural meadows may benefit the abundance of bumblebees. At the local level, patch area and shape seem to have positive and negative influences, respectively, on bumblebee species richness. At the landscape level, human settlements with the presence of gardens may favour bumblebee richness and abundance. Also, bumblebee species may increase with a high presence of meadows in the landscape, and may decrease with high percentages of forest and young forest. Overall, forested landscapes with a strong presence of edges and a diverse matrix may support a higher species richness and abundance of bumblebees. Both local and landscape factors should be considered when designing conservation strategies and agri-environmental measures.     

The influence of 3-years of warming and N-deposition on ecosystem dynamics is small compared to past land use in subalpine meadows

Background and aims

Drivers of ecosystem dynamics that are under human influence range from local, land-management decisions to global processes such as warming temperatures and N deposition. The goal of this study was to understand how multiple, potentially interacting factors influence net primary production, N mineralization, and water and soil CO₂ fluxes.

Methods

Here I report on a three-year experiment that manipulated air temperature using ITEX passive warming cones and N deposition in a mountain meadow ecosystems that were historically grazed or protected from grazing.

Results

The strongest and most consistent effect was due to the legacy of grazing, with previously grazed sites having lower primary production, lower soil respiration rates, lower soil moisture, and lower soil C and N stocks than historically ungrazed sites. Warming increased soil respiration, but the effect was transient, and decreased over the 3-year study. Nitrogen addition increased primary production in the second and third year of the experiment but had no significant effect on soil respiration. The effect of historical grazing on primary production was approximately double the effect of N addition. Temperature and N deposition rarely interacted except for increasing N availability during the warm, wet growing season of 2004.

Conclusions

These findings indicate that the legacies of land use, with their influence on plant community composition and hydrologic processes, are locally more important than short-term step changes in temperature and nutrient availability.     

Teasing apart plant community responses to N enrichment: the roles of resource limitation,competition and soil microbes

Although ecologists have documented the effects of nitrogen enrichment on productivity, diversity and species composition, we know little about the relative importance of the mechanisms driving these effects. We propose that distinct aspects of environmental change associated with N enrichment (resource limitation, asymmetric competition, and interactions with soil microbes) drive different aspects of plant response. We test this in greenhouse mesocosms, experimentally manipulating each factor across three ecosystems: tallgrass prairie, alpine tundra and desert grassland. We found that resource limitation controlled productivity responses to N enrichment in all systems. Asymmetric competition was responsible for diversity declines in two systems. Plant community composition was impacted by both asymmetric competition and altered soil microbes, with some contributions from resource limitation. Results suggest there may be generality in the mechanisms of plant community change with N enrichment. Understanding these links can help us better predict N response across a wide range of ecosystems.     

云南香格里拉地区亚高山草甸不同放牧管理方式下的碳排放

青藏高原由于高寒低温限制了有机碳的分解,大量的碳积累在土壤碳库中,对全球升温的反应很敏感.放牧可能会对该区草甸的碳排放产生显著影响.采用闭合箱动态法测定了云南香格里拉地区不同放牧管理方式下的亚高山草甸生态系统呼吸与土壤呼吸.常年放牧草甸与季节性放牧草甸的生态系统呼吸和土壤呼吸均呈现相似且明显的单峰季节变化特征,7月份达最大值,生态系统呼吸分别为9.77、8.03 μmol CO2 m-2 s-1,土壤呼吸分别为8.05、7.74 μmol CO2 m-2 s-1;1月份达最低值,生态系统呼吸分别为0.21、0.48 μmol CO2 m-2 s-1,土壤呼吸分别为0.16、0.49 μmol CO2 m-2 s-1.受一天中气温和土温的影响,常年放牧草甸的生态系统呼吸与土壤呼吸的日变化在夏季与冬季均呈现明显的单峰曲线变化,最高值都出现在14:00左右,最低值出现在凌晨.在夏季6～10月份,常年放牧草甸的呼吸显著大于季节性放牧草甸,表明较高的放牧强度增加了亚高山草甸的碳排放.土壤温度的指数模型F = aebT比土壤水分能更好地解释呼吸的变异性(R2 = 0.50～0.78,P < 0.0001).二元回归模型F=aebTWc比单因子模型的效果更好(R2=0.56～0.89,P < 0.0001).土壤呼吸在整个亚高山草甸生态系统呼吸中占主导地位,在常年放牧草甸与季节性放牧草甸分别为63.0%～92.7%和47.5%～96.4%,地上植物呼吸随生长季的变化而变化,在生长旺季占有较大的比例.生态系统呼吸和土壤呼吸的长期Q10(1a)是短期Q10(1d)的2倍左右.季节性放牧草甸的长期Q10小于常年放牧草甸,表明在温度上升的背景下,放牧压力较小的草甸碳库较为稳定,具有较好的碳截存能力.     

Variation in the phenology and abundance of flowering by native and exotic plants in subalpine meadows

The timing and abundance of flower production is important to the reproductive success of angiosperms as well as pollinators and floral and seed herbivores. Exotic plants often compete with native plants for space and limiting resources, potentially altering community floral dynamics. We used observations and a biomass-removal experiment to explore the effects of an invasive exotic flowering plant, Linaria vulgaris, on community and individual species flowering phenology and abundance in subalpine meadows in Colorado, USA. Invasion by L. vulgaris was associated with a shift in both the timing and abundance of community flowering. Invaded plant communities exhibited depressed flowering by 67% early in the season relative to uninvaded communities, but invaded sites produced 7.6 times more flowers than uninvaded sites once L. vulgaris began flowering. This increase in flowers at the end of the season was driven primarily by prolific flowering of L. vulgaris. We also found lower richness and evenness of resident flowering species in invaded plots during the period of L. vulgaris flowering. At the species level, a common native species (Potentilla pulcherrima) produced 71% fewer flowers in invaded relative to uninvaded plots, and the species had reduced duration of flowering in invaded relative to uninvaded sites. This result suggests that L. vulgaris does not simply alter the flowering of subordinate species but also the flowering of an individual common species in the plant community. We then used observational data to explore the relationship between L. vulgaris density and resident floral production but found only partial evidence that higher densities of L. vulgaris were associated with stronger effects on resident floral production. Taken together, results suggest that a dominant invasive plant can affect community and individual-species flowering.     

取样尺度对亚高寒草甸物种多样性与生产力关系的影响

植物物种多样性与生产力之间的关系是群落生态学的一个热点问题, 目前仍存在着很多争议。为探究自然群落中二者之间的关系, 对青藏高原亚高寒草甸3个样地的自然植物群落分别进行了不同取样面积的抽样调查。结果显示, 取样样地和取样尺度均对物种丰富度有显著影响, 取样样地而非取样尺度对群落地上生物量有显著性影响。在某一时刻对某一样地进行取样, 其单位面积生产力并不因取样面积的增加而提高, 而是保持恒定的, 尽管物种数随取样面积的增加而有明显增多。物种多样性与生产力之间的回归关系因样地与取样尺度不同而不同, 有U型、单峰型、正线性相关和无相关性, 其中无相关性出现的最多。据此推测, 亚高寒草甸群落物种多样性与生产力之间不存在某种确定性关系, 或者说, 亚高寒草甸物种多样性和生产力之间不存在必然的因果联系。     

积雪对祁连山亚高山草甸土壤呼吸速率的影响

草地作为陆地生态系统的重要组成部分,是分布最广的植被类型之一,全球草地面积约占陆地面积的1/4,土壤碳储量约占全球总碳储量的1/5,在气候变化和陆地生态系统的碳循环方面起着重要作用。我国各类天然草原面积近4亿hm~2,约占国土面积的41.7%,草地的总碳储量约占陆地生态系统总碳储量的16.7%,其中土壤层占93.1%,在碳储量中占有重要的地位。土壤呼吸是陆地生态系统碳循环的主要环节,其中草地土壤呼吸占陆地土壤呼吸量的5.6%,因此草地土壤碳库正成为草地生态系统研究的热点。祁连山草地资源丰富,亚高山草甸是祁连山自然保护区的基质景观。积雪作为冰冻圈的重要组成部分,对土壤温度和土壤水分具有调控作用,是影响土壤呼吸的重要因素,但是目前关于积雪对土壤呼吸的影响研究少见。研究区选择祁连山中部天涝池流域,利用LI-8100开路式土壤碳通量测定系统,探讨了土壤不同融化状态下积雪对祁连山亚高山草甸生态系统呼吸速率和土壤呼吸速率的影响,并分析了地表温度和土壤温度、地表空气相对湿度和土壤体积含水量对土壤呼吸的影响。结果表明:(1)雪覆盖降低了生态系统呼吸和土壤呼吸速率,但当日融化的积雪对次日生态系统呼吸有促进作用;(2)呼吸速率与土壤温度和地表温度呈显著指数关系(P0.01),与地表空气相对湿度呈线性相关关系。降雪后,全融土壤中呼吸速率与温度和地表空气相对湿度间的相关性好;(3)温度较高时,温度对呼吸速率的作用强;温度较低时,温度对呼吸速率的影响不大。     

Foraging trip duration of bumblebees in relation to landscape-wide resource availability

Abstract.  1. The study tested the hypotheses that bumblebees have shorter foraging trips in environments that provide abundant resources than in environments that provide sparse resources, and that shorter foraging trips translate into greater colony growth.
2. Six even-aged Bombus terrestris colonies were established in contrasting resource environments. Three colonies had access to abundant resources ( Phacelia tanacetifolia fields with high flower densities), and three colonies were placed in an environment with sparse resources (scattered semi-natural habitats with food plants at lower densities).
3. A total of 870 foraging trips of 220 marked B. terrestris foragers were observed using automated camcorder recordings.
4. The duration of foraging trips was shorter in environments with abundant resources (66 ± 4.6 min) than in environments with sparse resources (82 ± 3.7 min). Within 34 days colonies that had access to abundant resources gained significantly more weight (129 ± 40 g) than colonies foraging on sparse resources (19 ± 7 g).
5. Thus, the spatial distribution and quality of resources at landscape level affected the duration of foraging trips and the colony growth. It was concluded that future conservation schemes need to improve the spatial and temporal availability of resources in agricultural landscapes to counteract the ongoing decline of bumblebees.     

Pollen and resource limitation in a gynodioecious species

Differences between plant sex morphs in pollen or resource availability may affect their relative fitness and thereby the sex ratio of dimorphic species. In gynodioecious species, in which hermaphroditic and female plants coexist, a variety of factors (e.g., hermaphrodite self-fertility or rarity or pollinator discrimination against females) might be expected to lead to stronger pollen limitation in females than in hermaphrodites. On the other hand, females have been found to be superior compared to hermaphrodites in low-nutrient conditions. The effects of supplemental hand-pollination and resource addition on the reproductive output of the self-fertile gynodioecious perennial Geranium sylvaticum (Geraniaceae) were tested for several populations that differ in their female frequency (4.4-23.0%). Both pollen and resource availability limited fruit set and the number of seeds produced per plant; however, seed set (i.e., the number of seeds produced per fruit) was limited only by resources. Because pollen limitation in females did not correlate with female frequency, our results suggest that pollen limitation in females does not depend on the frequency of the pollen-producing hermaphrodites. Furthermore, because pollen and resource availability limited reproductive output of both sex morphs, these factors may not contribute significantly to maintenance and evolution of gynodioecy in G. sylvaticum.     

Photosynthetic limitation of several representative subalpine species in the Catalan Pyrenees in summer

Information on the photosynthetic process and its limitations is essential in order to predict both the capacity of species to adapt to conditions associated with climate change and the likely changes in plant communities. Considering that high‐mountain species are especially sensitive, three species representative of subalpine forests of the Central Catalan Pyrenees: mountain pine (Pinus uncinata Mill.), birch (Betula pendula Roth) and rhododendron (Rhododendron ferrugineum L.) were studied under conditions associated with climate change, such as low precipitation, elevated atmospheric [CO₂] and high solar irradiation incident at Earth's surface, in order to detect any photosynthetic limitations. Short‐term high [CO₂] increased photosynthesis rates (A) and water use efficiency (WUE), especially in birch and mountain pine, whereas stomatal conductance (g_s) was not altered in either species. Birch showed photosynthesis limitation through stomatal closure related to low rainfall, which induced photoinhibition and early foliar senescence. Rhododendron was especially affected by high irradiance, showing early photosynthetic saturation in low light, highest chlorophyll content, lowest gas exchange rates and least photoprotection. Mountain pine had the highest A, photosynthetic capacity (A_max) and light‐saturated rates of net CO₂ assimilation (A_sat), which were maintained under reduced precipitation. Furthermore, maximum quantum yield (F_v/F_m), thermal energy dissipation, PRI and SIPI radiometric index, and ascorbate content indicated improved photoprotection with respect to the other two species. However, maximum velocity of carboxylation of RuBisco (V_cmax) indicated that N availability would be the main photosynthetic limitation in this species.     

山西吕梁山亚高山草甸物种多样性的时空变化格局

在山西吕梁山系按照纬度从北向南依次选取马伦草原、荷叶坪、云顶山作为研究区,于2015、2016、2017年7月下旬进行亚高山草甸植被群落多样性调查,研究吕梁山亚高山草甸物种多样性不同年份、不同纬度的时空变化格局。结果表明:(1)吕梁山亚高山草甸群落中矮嵩草(Kobresia humilis)和珠芽蓼(Polygonum viviparum)、线叶嵩草(Kobresia capillifolia)和珠芽蓼、车前草(Plantago asiatica)和披碱草(Elymus dahuricus)分别为马伦草原、荷叶坪、云顶山的优势种,不同山地植物优势种和次优势种均以菊科(Asteraceae)、蔷薇科(Rosaceae)、莎草科(Cyperaceae)、禾本科(Gramineae)植物为主。(2)吕梁山亚高山草甸物种多样性在不同年份分布较为稳定,Pielou指数在连续3年内均无较大变化; Shannon-Wiener指数、Simpson指数和Patrick指数在2016年最高,同时该年份的降水天数也是最多,表明吕梁山亚高山草甸物种多样性在时间上呈现出受降水条件影响较大的变化格局。(3)吕梁山偏南部的云顶山多样性指数高于偏北部的马伦草原和荷叶坪,表明吕梁山亚高山草甸物种多样性在空间上表现出由北向南逐渐升高的变化格局。因此,吕梁山亚高山草甸物种多样性在时空上呈现出受降水条件影响较大,且由北向南逐渐升高的变化格局,降水条件和纬度梯度对该山地亚高山草甸物种多样性的时空变化格局产生了重要影响。     

The effects of snowpack properties and plant strategies on litter decomposition during winter in subalpine meadows

Background and aims

Approximately 50 % of belowground organic carbon is present in the northern permafrost region and due to changes in climate there are concerns that this carbon will be rapidly released to the atmosphere. The release of carbon in arctic soils is thought to be intimately linked to the N cycle through the N cycle’s influence on microbial activity. The majority of new N input into arctic systems occurs through N₂-fixation; therefore, N₂-fixation may be the key driver of greenhouse gases from these ecosystems.

Methods

At Alexandra Fjord lowland, Ellesmere Island, Canada concurrent measurements of N₂-fixation, N mineralization and nitrification rates, dissolved organic soil N (DON) and C, inorganic soil N and surface greenhouse gas fluxes (CO₂, N₂O and CH₄) were taken in two ecosystem types (Wet Sedge Meadow and Dryas Heath) over the 2009 growing season (June-August). Using Structural Equation Modelling we evaluated the hypothesis that CO₂, CH₄ and N₂O flux are linked to N₂-fixation via the N cycle.

Results

The soil N cycle was linked to CO₂ flux in the Dryas Heath ecosystem via DON concentrations, but there was no link between the soil N cycle and CO₂ flux in the Wet Sedge Meadow. Methane flux was also not linked to the soil N cycle, nor surface soil temperature or moisture in either ecosystem. The soil N cycle was closely linked to N₂O emissions but via nitrification in the Wet Sedge Meadow and inorganic N in the Dryas Heath, indicating the important role of nitrification in net N₂O flux from arctic ecosystems.

Conclusions

Our results should be interpreted with caution given the high variability in both the rates of the N cycling processes and greenhouse gas flux found in both ecosystems over the growing season. However, while N₂-fixation and other N cycling processes may play a more limited role in instantaneous CO₂ emissions, these processes clearly play an important role in controlling N₂O emissions.