Resource partitioning between shrubs and grasses in the Patagonian steppe

Summary Experiments were conducted in the Patagonian steppe in southern South America to test the following hypotheses: (a) grasses take up most of the water from the upper layers of the soil and utilize frequent and short-duration pulses of water availability; (b) shrubs, on the contrary, take up most of the water from the lower layers of the soil and utilize infrequent and long-duration pulses of water availability. Grasses and shrubs were removed selectively and the performance of plants and the availability of soil resources were monitored. Results supported the overall hypothesis that grasses and shrubs in the Patagonian steppe use mainly different resources. Removal of shrubs did not alter grass production but removal of grasses resulted in a small increase in shrub production which was mediated by an increase in deep soil water and in shrub leaf water potential. The efficiency of utilization of resources freed by grass removal was approximately 25%. Shrubs used water exclusively from lower soil layers. Grasses took up most of the water from upper layers but they were also capable of absorbing water from deep layers. This pattern of water partitioning along with the lack of response in leaf nitrogen to the removal treatments suggested that shrubs may be at a disadvantage to grasses with respect to nutrient capture and led to questions about the role of nutrient recirculation, leaching, and nitrogen fixation in the steppe.     

不同刈牧强度对冷蒿生长与资源分配的影响

利用野外实验与盆栽实验,对不同刈牧强度下冷蒿生长与资源分配影响的研究结果表明,按比例刈割冷篙的再生生长大于留茬高度刈割,在生长季前期,不刈割冷蒿净生长高于刈割处理,而进入生长季中后期(8月中旬以后),轻度刈割净生长高于不刈割处理,冷篙种群生物量分配的总体格局是根>叶>茎,刈牧明显影响冷蒿生物量分配格局,尤其是叶和花的分配,3／4刈割或留茬4cm刈割叶生物量分配显著高于其它各处理,而花的生物量及其分配显著低于其它处理,根、茎生物量分配各处理间差异不显著．冷蒿有性生殖分配随刈牧强度的增加而降低,繁殖方式发生了改变,优先将光合产物分配给再生茎以及繁殖方式转向营养繁殖,通过克隆生长维持和扩大种群是冷蒿对强度放牧的生态适应对策。     

日本中部不同放牧条件下人工草地地上枯死量的时间变化*

日本国立草地研究所位于日本中部的西那须地区,为了提示草地生态系统的能流和碳循环与气象因素与人为干扰之间的关系的规律,在其所辖人工草地的放牧试验场内,自1974a至1994a间,进行了不同放牧条件的长期实验。对该人工草地在21a间的地上枯死量(包括立枯部和地面凋落物)随时间变化规律以及不同放牧处理(不同放牧强度和施肥量)对其影响进行了评价和分析。结果表明,地上立枯部分和凋落物的量随季节和年度变化很大,并与地上部现存生物量有显著的正相关关系;协方差分析结果表明放牧压力、季节和年度等变化因素对地上立枯部分和凋落物的量有着极强的影响(p<0.01),而施肥量的影响则无显著性差异。     

Herbivory resistance traits in populations of Poa ligularis subjected to historically different sheep grazing pressure in Patagonia

Seeding selected populations with high grazing resistance may foster recovery of plant populations threatened by overgrazing. Resistance to grazing depends on grazing avoidance (escape from grazers) and grazing tolerance (ability to growth after defoliation). Many studies of grazing tolerance defoliate plants at a fixed height instead of removing the same proportion of biomass and therefore confound tolerance with avoidance. For this reason, the information on evolution of tolerance to defoliation at the intraspecific level is remarkably scarce despite the abundance of papers published that evaluate responses to defoliation. The estimation of the cost of tolerance is also troublesome because current methods usually include spurious correlations due to correlation between variables that share common terms. The objectives of this paper were to assess the intraspecific variation in tolerance and in traits associated with avoidance and growth in populations with different sheep grazing histories. We also estimated the percentage of biomass removed when the defoliation treatment was imposed at fixed height in order to separate tolerance and avoidance. Finally, we estimated the cost of tolerance using a new method proposed for spurious correlations. Results of a greenhouse experiment indicated no difference in tolerance among the three compared populations. However, the populations from overgrazed fields had more prostrate growth form, higher specific leaf area, and higher tillering rate (when no defoliated) than populations from exclosures. We confirmed that fixed height defoliation would have removed a higher proportion of shoot biomass from taller than from shorter individual plants, confounding grazing tolerance and avoidance. Regarding the cost of tolerance, we found no differences from a null model of no cost, indicating that the evolution (or future breeding) of more tolerant genotypes would not be constrained by this cost.     

Contrasting genetic metrics and patterns among naturalized rainbow trout (Oncorhynchus mykiss) in two Patagonian lakes differentially impacted by trout aquaculture

Different pathways of propagation and dispersal of non‐native species into new environments may have contrasting demographic and genetic impacts on established populations. Repeated introductions of rainbow trout (Oncorhynchus mykiss) to Chile in South America, initially through stocking and later through aquaculture escapes, provide a unique setting to contrast these two pathways. Using a panel of single nucleotide polymorphisms, we found contrasting genetic metrics and patterns among naturalized trout in Lake Llanquihue, Chile's largest producer of salmonid smolts for nearly 50 years, and Lake Todos Los Santos (TLS), a reference lake where aquaculture has been prohibited by law. Trout from Lake Llanquihue showed higher genetic diversity, weaker genetic structure, and larger estimates for the effective number of breeders (N_b) than trout from Lake TLS. Trout from Lake TLS were divergent from Lake Llanquihue and showed marked genetic structure and a significant isolation‐by‐distance pattern consistent with secondary contact between documented and undocumented stocking events in opposite shores of the lake. Multiple factors, including differences in propagule pressure, origin of donor populations, lake geomorphology, habitat quality or quantity, and life history, may help explain contrasting genetic metrics and patterns for trout between lakes. We contend that high propagule pressure from aquaculture may not only increase genetic diversity and N_b via demographic effects and admixture, but also may impact the evolution of genetic structure and increase gene flow, consistent with findings from artificially propagated salmonid populations in their native and naturalized ranges.     

不同放牧模式下高原鼠兔洞口的空间分布格局变化

2019年4-11月,在四川省若尔盖县选取全年禁牧、冬季放牧、全年放牧3种放牧模式样地,对高原鼠兔(Ochotona curzoniae)洞口的数量、空间分布格局变化及影响因子进行了研究.计算平均最近邻指数和S2/m比值来分析样方(10m×10m)和样地水平下的空间分布格局,通过非参数检验评估空间分布格局在不同放牧模式...     

高山草地绵羊放牧生态系统草畜营养供需动态

为了探讨放牧生态系统草畜间动态关系 ,缓减草畜营养供需的季节性不平衡 ,分别在高山草地春、夏、秋和冬季 ,进行了绵羊消化代谢试验 .结果表明 ,春季牧草消化能、粗蛋白质含量及粗蛋白质 消化能比值最高 ,分别为 9.76MJ、2 1.5 3%和 2 2 .0 6 g·MJ-1,有效能不足 ;高山草地牧草产量低 ,绵羊采食干物质量最低 (6 0 .5 1± 3.4 2 g·W0 .75kg-1·d-1) ;应补充饲料 ,尤其补饲能量饲料 .夏季牧草粗蛋白质 消化能比值为 17.6 2g·MJ-1,偏高 ,能量仍略显不足 ;补饲能量饲料 ,可提高绵羊对牧草N的利用率 .秋季与其它 3季牧地相比 ,草畜间的营养供需较平衡 ,放牧能满足绵羊的营养需要 ,其体内N存留率较高 .冬季牧草质差 ,消化能、粗蛋白质含量最低 ,分别为 4 .30MJ、4 .6 3% ,绵羊干物质采食量亦低 ,N代谢呈负平衡 ,应补饲干物质、有效能及粗蛋白质不足的部分 .为改善此现状 ,要实行季节畜牧业 ,建立植物生产与动物生产的耦合系统     

The effects of sheep grazing on seedling establishment and survival in grassland

This paper examines the effects of sheep and insect grazing on the generation of natural gaps in turf and the establishment of seedlings therein. The site is on an old field on limestone. There were five sheep grazing treatments: ungrazed, and grazed briefly in spring, briefly in autumn, in autumn or in summer and autumn. The first three of these treatments also contained plots in which insecticide was applied regularly during the growing season. Pins of increasing size (1.6 mm–12.8 mm diameter) were lowered at grid intersections in permanent quadrats to record gap type and the occurrence of seedlings on seven dates over 15 months.Heavily-grazed plots had more bare ground in spring and more herb seedlings established in them. Seedlings of annual species of Bromus were able to establish in gaps (bare ground and litter) of smaller size than was required by herb seedlings (which had much smaller seeds). The use of an insecticide increased seedling establishment in October in ungrazed and in spring-grazed paddocks but decreased it in autumn-grazed paddocks. This may indicate the greater importance of insect herbivory in making gaps in the matted sward of autumn-grazed plots.Seedlings which established in October 1986 and March 1987 were more likely to die than those establishing earlier in 1986. This may be because of winter-kill and especially vigorous spring vegetation growth respectively. Seedling mortality was not affected by gap size or grazing treatment (unless such effects operated before we detected the seedlings).     

Interactions between herbivory and resource availability on grazing tolerance of Leymus chinensis

Herbivory and resource interact to influence plant regrowth following grazing, but few detailed investigations on grazing tolerance at population levels are available. We conducted two pot experiments along a simulated grazing gradient (0%, 25%, 50% and 75% of shoot removal) at three water or nutrient levels to determine the interaction of resource and herbivory on Leymus chinensis, a perennial, dominant species in the eastern Eurasian steppes. Interactions between water availability and clipping intensity on the relative height growth rate (RHGR) and bud number were significant. Significant interactions between nutrient and clipping on RHGR, total biomass and specific leaf area (SLA) were also found. Total biomass and bud number, showing a unimodal curve along the clipping gradient in resource-rich environments, were highest at light clipping level, suggesting that this species has the plastic compensatory responses from under- to overcompensation. Interactions between herbivory and water or nutrient were opposite to each other. The “cooperative” interactions between water and herbivory magnified the difference in grazing tolerance of L. chinensis between high and low water treatments. The “antagonistic” interactions between nutrient and herbivory, on the other hand, were reflected in the lower tolerance to heavy clipping in the high nutrient than low nutrient treatments. Results partly support the limiting resource model (LRM). A modified and simplified graphic model of the LRM was proposed based on our results. The new LRM clearly demonstrated that “cooperative” interactions between varying water levels and clipping intensities aggravate the detrimental impacts of herbivores on plant growth and reproduction, whereas “antagonistic” interactions between nutrient and grazing alleviate the negative effects of herbivores. Biomass compensation and density compensation were identified as main mechanisms of herbivory tolerance in this clonal species.     

华南丘陵人工马占相思林下灌木湿润季节蒸腾与小气候的关系

在湿润季节测定了广东丘陵常见的人工马占相思林内６种灌木的蒸腾日变化, 描述其蒸腾日变化的规律．并分析叶片蒸腾与林内的光、气温、湿度等小气候因子及叶温、 叶片气孔扩散阻力的回归关系．所有被观测灌木的蒸腾耗水日变化格局相似,阳性种类桃 金娘在人工林的隐蔽条件下生长良好．其蒸腾速率最高,山苍子的蒸腾速率次之,其它种 类的蒸腾表现较为适中．回归分析结果表明,灌木蒸腾变化与空气相对湿度呈负相关,与 林内气温、全光照、光合有效辐射呈正相关．叶温和叶片气孔扩散阻力与灌木蒸腾变化分 别呈正、负相关．     

Water use efficiency of twenty-five co-existing Patagonian species growing under different soil water availability

The variation of plant water use efficiency (WUE) with water availability has two interacting components: a plastic response, evident when individuals of the same genotype are compared (e.g. wet versus dry years), and an interspecific response, evident when different species living in habitats with different water availability are compared. We analysed the WUE of 25 Patagonian species that belong to four life forms (grasses, shrubs, annual herbs and perennial herbs) in relation to the climatic conditions of 2 years and the mean historic water availability experienced by each species. To estimate water availability, we calculated the effective soil water potential (EWP) of each species, based on available information about soil water dynamics, phenology and root system structure. To estimate WUE, we used isotopic discrimination of leaf C (Δ¹³C) and mean annual water vapour difference between leaves and atmosphere (Δe) measured in situ. For the plastic response, for every species and life form, WUE increased from the dry to the wet year. We hypothesize that photosynthesis was less nutrient limited in the wet than in the dry year, facilitating higher net photosynthesis rates per unit of stomatal conductance in the wet year. For the interspecific response, WUE was lower in species native to drier habitats than in species native to wetter habitats. This response was mostly accounted for by a decrease in Δe with EWP. Annual herbs, which avoid drought in time (they have the earliest growth cycle), and shrubs, which avoid drought in space (they have the deepest roots), showed the highest EWP and WUE. We conclude that the conventional wisdom which states that the highest WUE occurs within a species during the driest years, and among species in the driest habitats, does not always hold true, and that co-existing life forms drastically differ in water availability and water economy.     

The effect of grazing and nutrient supply on periphyton associated bacteria

The effects of nutrient additions and grazing by macro-invertebrates on periphyton-associated algae and bacteria were studied by performing an enclosure experiment on three occasions from early spring to summer at mesotrophic Lake Erken and V?dd?, at the Swedish Baltic coast. There were significant interactions between nutrient additions and grazing on bacterial biomass and specific activity in Lake Erken. Thus, the importance of either bottom-up or top-down effects could not be singled out. Bacterial biomass increased with enrichment only in the absence of grazers. Grazer presence tended to increase bacterial biomass in ambient nutrient conditions, but to decrease bacterial biomass under enrichment. For specific activity the positive response to enrichment was restricted to grazer presence. Hence, grazing by macro-invertebrates may have an indirect positive effect on bacterial activity by enhancing nutrient conditions through their feeding activities and/or fecal pellets production. In addition, we found a significant relationship between bacterial production and chlorophyll a at both sites. This relationship weakened in the presence of macro-invertebrates. Thus, the importance of internal nutrient regeneration by bacteria and algae decreased, possibly due to increased nutrient availability, in the presence of macro-invertebrate grazers.     

Interspecific competition and coexistence of the two chrysomelids,Gastrophysa atrocyanea motschulsky andGalerucella vittaticollis baly (Coleoptera: Chrysomelidae), under limited food resource conditions

Field experiments were conducted in order to investigate the mode of exploitation of food resources and the mechanism of coexistence of mixed larval populations of the two chrysomelids,Gastrophysa atrocyanea andGalerucella vittaticollis, under limited food resource conditions. The larval survival rates seemed high enough to assure coexistence when hatchlings of the two species were released in 1∶1 and 1∶3 ratios on a host plant. However, the survival rate became almost nil for both species when a 3∶1 ratio was employed, suggesting asymmetrical interspecific competition. Wasted food consumption was much higher inG. atrocyanea larvae. The population ofG. atrocyanea seemed to be regulated more by intraspecific competition, while on the other hand, the population ofG. vittaticollis was considered to be more likely affected by the interspecific competition withG. atrocyanea, depending on the initial ratio and density of the two species.     

Interactive effects of grazing and shrubs on the annual plant community in semi‐arid Mediterranean shrublands

Question: We studied the interactive effects of grazing and dwarf shrub cover on the structure of a highly diverse annual plant community. Location: Mediterranean, semi‐arid shrubland in the Northern Negev desert, Israel. Methods: Variation in the biomass and plant density of annual species in the shrub and open patches was monitored during four years, inside and outside exclosures protected from sheep grazing, in two contrasting topographic sites: north and south‐facing slopes that differed in their dominant dwarf shrubs species: Sarcopoterium spinosus and Corydothymus capitatus, respectively. Results: Above‐ground biomass, density and richness of annual species were lower under the canopy of both shrub species compared to the adjacent open patches in the absence of grazing. Grazing reduced the biomass of annuals in open patches of both topographic sites, but not in the shrub patches. On the north‐facing slope, grazing also reduced plant density and richness in the open patches, but increased plant density in the shrub patches. At the species level, various response patterns to the combined effects of grazing and patch type were exhibited by different annuals. Protection against the direct impacts of grazing by shrub cover as well as species‐specific interactions between shrubs and annuals were observed. A conceptual mechanistic model explaining these interactions is proposed. Conclusion: In semi‐arid Mediterranean shrublands grazing and dwarf shrub cover interact in shaping the structure of the annual plant community through (1) direct impacts of grazing restricted to the open patches, (2) species‐specific facilitation/ interference occurring in the shrub patches and (3) subsequent further processes occurring among the interconnected shrub and open patches mediated through variation in seed flows between patches.     

The effects of fire frequency and grazing on tallgrass prairie productivity and plant composition are mediated through bud bank demography

Periodic fire, grazing, and a variable climate are considered the most important drivers of tallgrass prairie ecosystems, having large impacts on the component species and on ecosystem structure and function. We used long-term experiments at Konza Prairie Biological Station to explore the underlying demographic mechanisms responsible for tallgrass prairie responses to two key ecological drivers: fire and grazing. Our data indicate that belowground bud banks (populations of meristems associated with rhizomes or other perennating organs) mediate tallgrass prairie plant response. Fire and grazing altered rates of belowground bud natality, tiller emergence from the bud bank, and both short-term (fire cycle) and long-term (>15 year) changes in bud bank density. Annual burning increased grass bud banks by 25% and decreased forb bud banks by 125% compared to burning every 4 years. Grazing increased the rate of emergence from the grass bud bank resulting in increased grass stem densities while decreasing grass bud banks compared to ungrazed prairie. By contrast, grazing increased both bud and stem density of forbs in annually burned prairie but grazing had no effect on forb bud or stem density in the 4-year burn frequency treatment. Lastly, the size of the reserve grass bud bank is an excellent predictor of long-term ANPP in tallgrass prairie and also of short-term interannual variation in ANPP associated with fire cycles, supporting our hypothesis that ANPP is strongly regulated by belowground demographic processes. Meristem limitation due to management practices such as different fire frequencies or grazing regimes may constrain tallgrass prairie responses to interannual changes in resource availability. An important consequence is that grasslands with a large bud bank may be the most responsive to future climatic change or other global change phenomena such as nutrient enrichment, and may be most resistant to exotic species invasions.     

不同供氮水平下反枝苋的化感作用与资源竞争能力

为探讨反枝苋(Amaranthus retroflexus)在入侵农田生态系统过程中的化感作用与资源竞争能力之间的关系,采用添加活性炭的方法,比较了在有无活性炭条件下,反枝苋与大豆(Glycine max)的生物量积累、生物量分配及不同器官含氮量随供氮水平的变化情况。结果表明,在低供氮水平时,化感作用在竞争中占有的比重较大,在高供氮水平时,化感作用所占的比重相对降低;在高供氮条件下,反枝苋增加了对根和叶的生物量分配,而在低供氮条件下,则增加对繁殖器官的生物量分配;在氮素营养贫瘠的条件下,添加活性炭的大豆含氮量要明显小于不添加活性炭时,随着供氮水平的上升,差异逐渐减小。说明在入侵过程中,反枝苋可以通过调节自身的化感作用和生物量分配来适应不同的氮素营养条件。     

不同温度与寄主条件下桃蚜生命表的研究

在寄主、温度对桃蚜生物学特性影响研究的基础上,研究了不同温度和寄主对桃蚜生命表的影响．结果表明,桃蚜生长发育最适温度均为２８℃,致死温度为３３℃．寄主对桃蚜最适内禀增长力（ｒ＇ｍ）有显著影响,不同寄主间２组合的蚜虫最适温度分别为２０．６６和２６．４４℃,同一寄主间为２４．４３和２８．９９℃;随着温度的升高,种群加倍时间呈负指数曲线下降．改换寄主后的蚜虫对低温条件更敏感．     

The resource economics of chemical and structural defenses across nitrogen supply gradients

In order to better understand the role of nutrient supplies in determining the prevalence of plant defense types, we investigated the theoretical relationships between ecosystem N supply and the net C gain of shoots that were undefended or defended in one of three ways: (1) by N-free chemical compounds, (2) by N-containing chemical compounds, or (3) by structural defenses. By extending economic models of shoot resource balance to include the relative value of C and N, depreciation, and amortization, we were able to show that the relative net C gain of the three defense types were similar to changes in their generally understood abundance along an N supply gradient. At low N supply, the additional C acquired when investing C in defense is much higher than investing N in defenses. Only at high N supply is it better to invest large quantities of N in defense rather than additional photosynthesis. In a sensitivity analysis, net C gain of shoots was most sensitive to factors that affect the relative value of C and N and the rate of herbivory. Although there is support for the relative value of C and N influencing defense strategies, more research is necessary to understand why tannins are not more prevalent at high N supply and why moderate amounts of N-based defenses are not used at low N supply.Due to an error in the citation line, this revised PDF (published in December 2003) deviates from the printed version, and is the correct and authoritative version of the paper.     

The life histories and population dynamics of two carabid species on a Dutch heathland

Summary We deal with the causes of the synchronously fluctuating numbers of subpopulations of the carabid species Calathus melanocephalus as compared with the asynchronously fluctuating numbers of subpopulations of the carabid Pterostichus versicolor. Both species continuously occupy a large heath area, Dwingelder Veld (1600 ha), in The Netherlands, and are studied there in the same localities with the same methods. Of the adults of C. melanocephalus, 90% do not cover more than 2 ha during the entire reproductive season, while 90% of adults of P. versicolor cover no more than 12 ha. In C. melanocephalus egg production in the field is usually similar to that under optimal feeding conditions in the laboratory, but in P. versicolor egg production seems to be much lower in the field. In the field 70–80% of the eggs most probably are killed by eelworms, followed by more than 90% mortality among the remaining larvae. Comparing mortality of developmental stages in laboratory experiments with that in field experiments in enclosures, it appears that mortality of larvae is not density-dependent, even when density in the experiments is much higher than it ever is in the field. Larval mortality mainly results from the poor ability of the larvae to find prey, even when in field experiments prey density is increased far above natural densities. We discuss why these poor prey-finding abilities are not improved by natural selection. In the spring breeder P. versicolor differences between localities both in abiotic factors, soil moisture and surface temperature, and biotic factors, reactions of prey species to abiotic factors, in spring and summer when the larvae are maturing contribute to the asynchronous fluctuations of numbers between subpopulations. In the autumn breeder C. melanocephalus possible differences in biotic factors between sites are outnumbered by the effects of winters with a higher or lower than normal amount of precipitation respectively. During a wet winter mortality among the larvae is much higher than during a dry winter. As these winter conditions are similar over large areas (many km²) the fluctuations of numbers between subpopulations are synchronous.Communication No. 443 of The Biological Station, Wijster