Performance of two Picea abies (L.) Karst. stands at different stages of decline

Summary The annual replacement of tillers of Agropyron desertorum (Fisch. ex Link) Schult., a grazing-tolerant, Eurasian tussock grass, was examined in the field following cattle grazing. Heavy grazing before internode (culm) elongation seldom affected tiller replacement. Heavy grazing during or after internode elongation, which elevates apical meristems, increased overwinter mortality of fall-produced tillers and reduced the number and heights of these replacement tillers. Unexpectedly, tussocks grazed twice within the spring growing season tended to have lower overwinter tiller mortality, greater tiller replacement, and larger replacement tillers than tussocks grazed only once in late spring. These responses of twice-grazed tussocks, however, were still less than those of ungrazed tussocks or tussocks grazed moderately in early spring. The presence of ungrazed tillers on partially grazed tussoks did not increase the replacement of associated grazed tillers relative to tillers on uniformly grazed plants. This result indicates that resource sharing among tillers, if present, is short-lived or ecologically unimportant in this species. Although A. desertorum is considered grazing-tolerant, tiller replacement on heavily grazed tussocks, particularly those grazed during or after internode elongation when apical meristems were removed, was usually inadequate for tussock maintenance. These observations at the tiller (ramet) level of organization in individual tussocks (genet) may explain the often noted reduction in stand (population) longevity with consistent heavy grazing.     

Effects of topography,soil moisture,wind and grazing on Festuca seedlings in a Patagonian grassland

Abstract. The effects of topography, soil moisture, wind and grazing on the emergence and survival of seedlings of Festuca spp. were examined in the steppe zone of Patagonia, Argentina. Ungrazed and grazed field treatment plots were established on a plain and a north-facing slope at the Media Luna Ranch (43° 36′S, 71° 25′W). On the leeward and windward sides of each of 15 Festuca plants, 0.1 m × 0.4 m quadrats were censused bimonthly for seedling emergence and survival over three growing seasons. Three categories were distinguished: recently germinated and up to the first leaf, two to four leaves, and from five leaves up to one tiller. Soil moisture content, litter cover and frost heaving effects were also determined for each treatment at each sampling date. Festuca spp. showed two emergence peaks, one in late fall and the other in early to mid-spring. Seedling emergence was significantly correlated with soil moisture content in the 0–5 cm of the soil during the three growing seasons. Seedlings that emerged in the fall had higher survivorship than those that emerged in spring. Seedling emergence and survival was significantly (p < 0.01) lower on slopes, in the grazing treatment, and on windward sides of adult plants. In this grassland, an increase in the availability of safe sites for seedling emergence and survival might be achieved by protecting vegetation from grazing, particularly on north-facing slopes.     

Spatial arrangement of tiller replacement in Agropyron desertorum following grazing

Summary The spatial arrangement of tiller replacement was assessed on grazed and ungrazed tussocks of Agropyron desertorum (Fisch. ex Link) Schult. for three annual cycles. Frequency distributions of the number of replacement tillers per single progenitor were also determined. Tiller replacement was usually greater on the perimeter of tussocks than within the core, with or without grazing. Replacement was inversely related to grazing intensity, both on the perimeter and within the core of tussocks. Heights of replacement tillers on the perimeter or within the core seldom differed. Furthermore, grazing seldom affected the number of replacement tillers per progenitor. Greater tillering on the perimeter than within the core indicates that the tussocks were expanding. Apparently, grazing neither enhances tussock expansion and subsequent disintegration, nor does it necessarily lead to patches of tillers (multiple tillering per progenitor) within tussocks of A. desertorum.     

GRAZING HISTORY,DEFOLIATION, AND FREQUENCY-DEPENDENT COMPETITION: EFFECTS ON TWO NORTH AMERICAN GRASSES

Agropyron smithii and Bouteloua gracilis plants from intensively grazed prairie dog colonies and from a grazing exclosure in Wind Cave National Park, South Dakota, were used to compare responses of conspecific populations with different histories of exposure to grazing and to competition for light. In separate experiments for each species, plants grown in monocultures and two-population replacement-series mixtures were used to examine effects of defoliation, frequency-dependent competition, and population on biomass and morphology. Colony and exclosure plants frequently responded differently. Defoliation more often adversely affected exclosure plants than colony plants, while interpopulation competition more often adversely affected colony plants. Defoliation frequently negated the competitive advantage of exclosure plants. Intrapopulation competition appeared to be greater among exclosure than colony plants. Our results indicate that conclusions based on studies of plants in long-term exclosures may not apply to plant populations having long histories of intensive grazing. While there were differences between species, in both, these experiments provide evidence of population differentiation, resulting in morphologically dissimilar populations which responded differently to defoliation and to inter- and intrapopulation competition.     

Plant morphology and grazing history:

Grazing-related, intraspecific, morphological variation was studied in four North American grasses (Bouteloua gracilis, Agropyron smithii, Schizachyrium scoparium, and Andropogon gerardii) from eight locales in Wind Cave National Park, South Dakota: three locales currently occupied and heavily grazed by prairie dogs (Cynomys ludovicianus), colonized (since settlement) for 2–100 years, where native ungulates concentrate grazing activities; an extinct colony locale from which prairie dogs were removed 30 years previously, moderately to lightly grazed by ungulates; two noncolony locales, moderately to lightly grazed by ungulates; and two locales from within a 50-year-old grazing exclosure, with no known history of grazing by prairie dogs nor any recent grazing by ungulates. Data were collected both in situ and in common environments.Active-colony plants were more frequently and more heavily grazed than those at other grazed locales. In situ, plants from heavily grazed populations were smaller and more prostrate than those from populations with little or no grazing (including the extinct colony) and interpopulation variation corresponded to current grazer use. After several growing seasons in common environments, there were still significant interpopulation differences; however, variation often corresponded with grazing history. Although differences between active-colony and noncolony plants were somewhat reduced (indicating some phenotypic plasticity), active-colony plants were still smaller and more prostrate. However, extinct-colony plants more closely resembled active-colony plants than noncolony plants. Morphological variation among these populations is the result of more than simple grazer use; historical factors and the dynamic nature of the grazing regimes are also contributing factors.Abbreviations A. gerardii Andropogon gerardii - A. smithii Agropyron smithii - BFC Bison Flats colony locale - BFN noncolony locale at Bison Flats - B. gracilis Bouteloua gracilis - EXT Upper Highland extinct-colony locale - GDN common garden - GH greenhouse - NEW new satellite colony locale - PVC Pringle Valley colony locale - PVN noncolony locale in Pringle Valley - S. scoparium Schizachyrium scoparium - WCNP Wind Cave National Park, South Dakota, USA - XFN exclosure locale just inside exclosure fence from BFN - XHQ exclosure locale near headquarters buildings     

Light field heterogeneity among tussock grasses: Theoretical considerations of light harvesting and seedling establishment in tussocks and uniform tiller distributions

Although the tussock growth form of caespitose graminoids is widespread, the effect of this growth form on light interception and carbon gain of tillers has received little attention. Daily incident photosynthetic photon flux density (PFD_inc) and carbon gain in monospecific stands of tussock grasses were compared with those of a hypothetical distribution with the equivalent tiller density per total ground area, but evenly distributed rather than clumped in tussocks. This was computed for two tussock grasses Pseudoroegneria spicata (Pursh) A. Löve (bluebunch wheatgrass) and Agropyron desertorum (Fisch, ex Link) Schult. (creasted wheatgrass) at different plant densities. Daily PFD_inc and net photosynthesis (A) were greater if tillers were distributed uniformly rather than clumped in tussocks, except when the density of tussocks was so great as to approach a uniform canopy. When tussock density per ground area was low, much of the difference between tussock and uniform tiller densities in PFD_inc and A was due to shading within the tussocks; up to 50–60% of the potential carbon gain was lost in A. desertorum due to shading within tussocks. In a matrix of tussocks, the light field for establishing seedlings was very heterogeneous; potential A ranged from 7 to 96% relative to an isolated seedling. The mean of daily PFD_inc and A for seedlings in a tussock stand were nearly identical to the values in corresponding stands of uniform tiller distributions. It is hypothesized that the loss of A resulting from clumping tillers into tussocks is offset by benefits of protecting sequestered belowground resources from invasion by seedlings of competitors.     

Effect of competition intensity on recruitment of palatable and unpalatable grasses

In this study, we made an attempt to reveal how competition intensity from established plants impacts on palatable and unpalatable grass seedlings recruitment, in a natural mesic grassland of central Argentina. Our objective was to assess the seedling recruitment of a palatable species (Chascolytrum subaristatum) and an unpalatable species (Nassella trichotoma) in microsites differing in competition intensity from established plants. Identity (C. subaristatum and N. trichotoma) and defoliation severity were used as surrogate for competition intensity. In March 2017, we permanently marked established individuals of N. trichotoma and C. subaristatum and placed two circular plots adjacent to each individual. In one plot we added seeds of N. trichotoma and in the other seeds of C. subaristatum. After seeding, established plants were randomly assigned to one of three level of defoliation: without defoliation, low defoliation severity and high defoliation severity. From April to November 2017 (i.e. over a complete annual growing cycle), we measured seedling density, recruitment and growth. Our results supported the hypothesis that seedlings of palatable grasses are more competitive than seedlings of unpalatable grasses. Seedling of the palatable grass C. subaristatum recruited successfully regardless the intensity of competition from established plants, whereas seedlings of the unpalatable grass N. trichotoma recruited better under low competitive pressure from established plants. Our results suggest that the availability of microsites with low competitive pressure from the established vegetation, created by selective grazing of palatable grasses, promotes the recruitment of unpalatable grass seedlings. This mechanism may contribute to the species replacement process commonly observed in heavy grazed grasslands.     

Continuous grazing disrupts desert grass-soil seed bank composition under variable rainfall

Long-term field studies help unveil mechanisms of grass soil seed bank (SSB) persistence and resilience. We wonder whether grazing is a disruptive force that changes grass SSB composition and size, while heavy rainfall is a positive bottom-up force that restores the grass SSB in grazed areas of the Monte desert, Argentina. We used core sampling to study the grass SSB in cattle-free (17 year) and grazed (10 year) grasslands. The SSB size in cattle-free sites usually responded in a continuous positive fashion to rainfall, and the proportion of palatable grasses in the SSB was similar among years. Most grass species formed a transient SSB, but the presence of a stand of perennial plants would prevent their local extinction. In contrast, the SSB size in the grazed habitats showed a threshold response to rainfall (i.e. it did not increase under low-moderate rainfall), the seeds of palatable grasses were always scarce, and the proportion of seeds of less palatable grasses increased under extreme rainy conditions. Seed production from grasses that grew tillers from axillary buds during a drought was zero, suggesting that the bank of plants could have a role in the replenishment of the grass SSB only in mesic years. Within the time span assessed here, continuous heavy grazing together with scarce as well as heavy rainfall were disruptive forces that reduced the number and proportion of seeds from palatable grasses in the SSB, which are also the species preferred by seed-eating animals.

     

Contrasting demography of two Patagonian shrubs under different conditions of sheep grazing and resource supply

Summary Two shrub species, dominant in western Patagonia, Argentina, exhibited different demographic behaviors under the effect of sheep grazing and of manipulated resource levels. Senecio filaginoides showed an increase in cover under increasing grazing pressures; the rise was explained by an increasing number of individuals and not by larger plants; this trend was reversed when no grazing took place. Mulinum spinosum showed a slight decrease in cover and density at increasing grazing pressures, and no change in plant size. In a field experiment in which water was added (30 mm in early summer) and the dominant grasses removed, both factors substantially augmented the number of 1-year-old seedlings of Senecio recruited; the only noticeable response of Mulinum was an increase in seedling emergence because of grass removal. Six years after this last treatment, Senecio exhibited a ten fold increase in density, and its population structure showed a peak in younger classes; Mulinum remained unaffected. In seedling surveys conducted within an exclsure and in its surroundings, the emergence of Senecio was similar in these areas in the two years of the study, whereas the emergence of Mulinum was higher inside the exclosure for one of the years. The emergence of Senecio in the wetter year was 3 times larger than in the drier one. Grazing did not affect first-year survival of any species; the major source of seedlign death in this case, as well as in the manipulative experiment, was desiccation during summer drought. We conclude that the contrasting responses of the two species to grazing are mostly due to the greater ability of Senecio to benefit from any increase in resources, including those not used by grazed plants. The difference might also be due to a reduction in the seed availability of Mulinum caused by sheep preference for its fruits. The dissimilar abilities of the two shrubs to employ the resources freed by grazing would derive more from their contrasting demographic potential (i.e. the ability to rapidly increase biomass/m²) than from a difference in their niche overlap with palatable species.     

海岸不同坡向滨麦光合特性与风速异质环境的关系

在自然条件下,测定分析了夏季和秋季静风天及秋季大风天,海岸沙丘不同坡向环境因子及滨麦株高、叶片叶绿素含量、土壤含水量和光合日变化特性,以期明晰滨麦响应不同风速环境的生理调控机理。结果表明,不同坡向环境异质性明显,滨麦形态可塑性强。海岸迎风坡风大、温度低、湿度大,滨麦植株低矮、叶绿素含量较高;背风坡温度高、土壤干旱、空气流动差,滨麦植株高大,叶绿素含量低。在夏秋季静风天,迎风坡和背风坡滨麦叶片Pn出现光合"午休"现象,但迎风坡滨麦日均Gs、Tr、Pn均显著高于背风坡;而在秋季大风天,迎风坡和背风坡滨麦叶片Pn"午休"现象消失,并且背风坡滨麦日均Pn、Tr、Gs均显著高于迎风坡滨麦。同坡向相比,秋季大风天迎风坡滨麦日均Pn、Tr、Gs明显较秋季静风天低,而背风坡滨麦日均叶片Pn、Tr、Gs却较静风天分别增高126%、66.3%、134%。强海风吹袭引发迎风坡温度降低、滨麦叶片摇摆、气孔导度降低导致Pn下降,而强海风使背风坡空气流动加快、温度降低、气孔导度增大、"午休"消失使Pn增高。不同坡向滨麦对不同海风风速的适应表现出明显的光合生理可塑性,它在滨麦适应不同风力、提高其光合速率和增加物质积累上具有重要作用。而滨麦的形态和光合生理可塑性可能是其在不同海风强度下生存、生长、实现种群扩张的重要生理调控机理,这一特性在未来作物、牧草和树木抗风、抗盐育种中具有重要应用价值。     

Kobresia tibetica tussocks facilitate plant species inside them and increase diversity and reproduction

Many plant species grow inside tussocks of some graminoids, but the underlying mechanisms remain unclear. We address whether some species occur and flower mainly inside tussocks so that species diversity and sexual reproduction are higher inside than outside tussocks, and whether relieving biological and physical stress is the mechanism associated with the facilitative process. In a heavily grazed grassland on the eastern Tibetan Plateau, where both physical (due to high altitude) and biological conditions (due to heavy grazing) are extremely harsh, we investigated vegetation in paired plots inside and outside 150 Kobresia tibetica tussocks and measured tussock basal area (=plot area). We also measured temperatures at soil surface, 5 and 10 cm depth and the number of animals (yaks, sheep and horses) grazing inside and outside tussocks. Sixty-seven percent of the species occurred and 42% flowered more frequently inside than outside tussocks, but none less frequently. Inside tussocks 78% species flowered, but outside tussocks only 31% did. Consequently, number of species, number of flowering species and number of inflorescences were all markedly larger inside than outside tussocks. Differences in number of species, number of flowering species and number of inflorescences inside and outside tussocks increased with increasing tussock basal area. Soil temperatures were lower inside than outside tussocks, but grazing intensity was much larger outside tussocks. Therefore, tussocks of K. tibetica facilitated the species inside them likely by grazing prevention, but not by increasing warmth. This study provides evidence that plant species colonizing tussocks of graminoids can be facilitated by the tussock species, and facilitation by grazing prevention may be one mechanism causing the coexistence of the species inside tussocks.     

Impact of grazing regime on a Mongolian forest steppe

Question: What is the impact of grazing regime on plant species abundance, plant growth form, plant productivity and plant nutrient concentrations in a forest steppe? Location: Hustai National Park in the forest steppe region of Mongolia. Methods: On the Stipa steppe we applied three different grazing regimes by using; (1) one type of exclosure which excluded grazing by large mammalian herbivores, mainly takh (Przewalski horse), (2) another type of exclosure that excluded both large and small (Siberian marmots) mammalian herbivores, and (3) control plots which were freely grazed. We measured species frequencies, tiller densities, plant biomass and nitrogen concentrations of the vegetation. Results: Exclusion from grazing by takh and marmots significantly increased plant standing crop, but marmot grazing and full grazing did not show significant differences. Protection from grazing decreased forage quality, shown by a lower N-concentration of the standing crop. However, this was solely the result of the lower live-dead ratio of the vegetation. The frequency of the rhizomatous Leymus chinensis decreased under reduced grazing, as did the frequency of the total of rhizomatous species. The frequency of Stipa krylovii increased under reduced grazing, as did its basal areas, tiller density and tussock height. Conclusion: Reduced grazing leads to a lower abundance of rhizomatous species and an increase in tussock species.     

Competitive interactions between palatable and unpalatable grasses native to a temperate semi-arid grassland of Argentina

The argument that selective grazing leads to competitive replacement of palatable grasses by unpalatable grasses is based upon the assumption that the competitive ability of the palatable species is higher than the one of unpalatable species in the absence of grazing. In order to test this hypothesis we have compared the competitive ability of Stipa clarazii (palatable) and S. trichotoma (unpalatable) under field conditions, and S. clarazii and S. gynerioides (unpalatable) under greenhouse conditions. The three species are native to a temperate semi-arid grassland of central Argentina. In the field experiment, plants of both species were grown either independently or in pairs (palatable + unpalatable), protected from grazing. Shoot and seed production were measured at the end of the growing seasons of 1993, 1994 and 1995. In the greenhouse experiment, plants of both species were grown in pots, either in monoculture or in mixture, under conditions of high and low water and mineral nutrient availability. Total biomass and seed production were measured at the end of the experimental period. In both experiments the presence of the unpalatable species did not affect (P < 0.05) the productive responses of the palatable species. On the contrary, the presence of the palatable species significantly reduced (P < 0.05) the productive responses of the unpalatable species. Our results support the assumption, on which most interpretations of floristic changes induced by grazing are based, that the competitive ability of palatable grasses is higher than the one of unpalatable grasses in the absence of grazing.     

Recovery of a dwarf birch (Betula nana) population following reduction in grazing by red deer (Cervus elaphus)

Summary

In Britain Betula nana is a nationally scarce plant, its distribution being limited by climate, heather burning and grazing by deer and sheep. The recovery of a population of B. nana was monitored for several years inside a Scots pine regeneration exclosure in N.E. Scotland. Browsing pressure was reduced in the exclosure but a few red deer (amounting to 3–4 km^?2) were deliberately retained. B. nana stem density increased after red-deer density was reduced. Over the same period there was a reduction in browsing inside the exclosure. B. nana stem density was negatively related to heather height, deer presence and tree canopy cover. Comparisons of growth were made with plants outside the exclosure between 1998 and 2000, the mean annual increment of stems, allowing for browsing loss, being respectively 1.25 and 0.45 cm for inside and outside the exclosure. Most stem basal diameters inside the exclosure were smaller than outside. Reducing red deer density to 3–4 km^?2 on similar upland sites with peaty soils could lead to regeneration of B. nana.     

The effects of feeding by the rice water weevil, Lissorhoptrus oryzophilus Kuschel, on the growth and yield components of rice, Oryza sativa

Abstract 1 The effects of feeding by larvae of the rice water weevil, Lissorhoptrus oryzophilus Kuschel, on the growth and yield components of rice, Oryza sativa, were evaluated using four varieties of rice, ‘Cocodrie’, ‘Cypress’, ‘Bengal’ and ‘XP1003’, over 2 years. 2 Both adults and larvae of L. oryzophilus feed on rice plants, but it is feeding by the larval stage that causes economic losses. Feeding by L. oryzophilus larvae resulted in extensive damage to root systems. Pruning of root systems resulted in a decrease in tiller number and shoot biomass of rice plants in the vegetative stage of growth. 3 Yield losses were due to a combination of decreases in panicle densities, numbers of grains per panicle, and grain weights. Decreases in panicle densities were a direct result of the reductions in tiller numbers. Reductions in numbers of grains per panicle and grain weights probably resulted from decreases in shoot biomass. 4 Injury by rice water weevil larvae is chronic. The tillering stage of rice suffered the majority of weevil damage, but the growth effects were not manifested until later.     

Beaver herbivory on aquatic plants

Herbivores have strong impacts on marine and terrestrial plant communities, but their impact is less well studied in benthic freshwater systems. For example, North American beavers (Castor canadensis) eat both woody and non-woody plants and focus almost exclusively on the latter in summer months, yet their impacts on non-woody plants are generally attributed to ecosystem engineering rather than herbivory. Here, we excluded beavers from areas of two beaver wetlands for over 2 years and demonstrated that beaver herbivory reduced aquatic plant biomass by 60%, plant litter by 75%, and dramatically shifted plant species composition. The perennial forb lizard’s tail (Saururus cernuus) comprised less than 5% of plant biomass in areas open to beaver grazing but greater than 50% of plant biomass in beaver exclusions. This shift was likely due to direct herbivory, as beavers preferentially consumed lizard’s tail over other plants in a field feeding assay. Beaver herbivory also reduced the abundance of the invasive aquatic plant Myriophyllum aquaticum by nearly 90%, consistent with recent evidence that native generalist herbivores provide biotic resistance against exotic plant invasions. Beaver herbivory also had indirect effects on plant interactions in this community. The palatable plant lizard’s tail was 3 times more frequent and 10 times more abundant inside woolgrass (Scirpus cyperinus) tussocks than in spatially paired locations lacking tussocks. When the protective foliage of the woolgrass was removed without exclusion cages, beavers consumed nearly half of the lizard’s tail leaves within 2 weeks. In contrast, leaf abundance increased by 73–93% in the treatments retaining woolgrass or protected by a cage. Thus, woolgrass tussocks were as effective as cages at excluding beaver foraging and provided lizard’s tail plants an associational refuge from beaver herbivory. These results suggest that beaver herbivory has strong direct and indirect impacts on populations and communities of herbaceous aquatic plants and extends the consequences of beaver activities beyond ecosystem engineering.     

Axillary bud banks of two semiarid perennial grasses: occurrence, longevity, and contribution to population persistence

The occurrence, longevity, and contribution of axillary bud banks to population maintenance were investigated in a late-seral perennial grass, Bouteloua curtipendula, and a mid-seral perennial grass, Hilaria belangeri, in a semiarid oak-juniper savanna. Axillary buds of both species were evaluated over a 2-year period in communities with contrasting histories of grazing by domestic herbivores. A double staining procedure utilizing triphenyl tetrazolium chloride and Evan's blue indicated that both viable and dormant axillary buds remained attached to the base of reproductive parental tillers for 18–24 months which exceeded parental tiller longevity by approximately 12 months. Bud longevity of the late-seral species, B. curtipendula, exceeded bud longevity of the mid-seral species, H. belangeri, by approximately 6 months. Younger buds located on the distal portion of the tiller base were 3.2 and 1.4 times more likely to grow out than older proximal buds of B. curtipendula and H. belangeri, respectively. The percentage of older proximal buds, which included comparable portions of viable and dormant buds, that grew out to produce tillers following mortality of parental tillers was 6.0% for B. curtipendula and 8.4% for H. belangeri. In spite of the occurrence of relative large axillary bud banks for both species, the magnitude of proximal bud growth did not appear sufficient to maintain viable tiller populations. We found no evidence to support the hypothesis of compensatory bud growth on an individual tiller basis for either species. Grazing history of the communities from which the buds were collected did not substantially affect the number, status, longevity, or outgrowth of axillary buds on an individual tiller basis for either species. However, long-term grazing by domestic herbivores influenced axillary bud availability by modifying population structure of these two species. Bud number per square meter for B. curtipendula was 25% lower in the long-term grazed compared to the long-term ungrazed community based on a reduction in both tiller number per plant and plant number per square meter. In contrast, bud number per square meter for H. belangeri was 190% greater in the long-term grazed than in the long-term ungrazed community based on a large increase in plant density per square meter. Minimal contributions of axillary bud banks to annual maintenance of tiller populations in this mid- and late-seral species underscores the ecological importance of consistent tiller recruitment from recently developed axillary buds. Consistent tiller recruitment in grasslands and savannas characterized by intensive grazing and periodic drought implies that (1) bud differentiation and maturation must be remarkably tolerant of adverse environmental conditions and/or (2) tiller recruitment may resume from buds that mature following the cessation of severe drought and/or grazing, rather than from mature buds that survive these disturbances. These scenarios warrant additional research emphasis given the critical importance of this demographic process to tiller replacement in species populations and the maintenance of relative species abundance in grasslands and savannas. Received: 12 August 1996 / Accepted: 30 December 1996     

Assessment of interaction between sheep and poorly palatable grass: a key tool for grassland management and restoration

Invasion of poorly palatable grasses due to abandonment or improper grazing management decreases pastures feeding value and biodiversity. With the aim to control their spread, we assessed the relationship between sheep foraging behavior and changes in aboveground phytomass, leaf traits, and chemical features of the tall grass Brachypodium rupestre and evaluated the effects of a B. rupestre-based diet on epithelium keratinization of rumen. Our results demonstrated that sheep became less selective throughout the experimental trials and B. rupestre decreased its aboveground phytomass. Some leaf traits showed significant changes (LDMC, LA, and ADL were higher in ungrazed areas; leaf nitrogen content was higher in the grazed ones). In addition, we detected an increase of the degree of epithelium keratinization of sheep. Thus high grazing pressure can be used to control the spread of B. rupestre, but negative effects on animal welfare due to the increase of rumen keratinization might be expected.     

Differential responses to defoliation frequency in little bluestem (<Emphasis Type="Italic">Schizachyrium scoparium</Emphasis>) in tallgrass prairie: implications for herbivory tolerance and avoidance

Plant responses to herbivory are complex. In grasses, relative growth rate (RGR), seed, and vegetative reproduction, resource allocation, and architecture vary differentially and often nonlinearly with grazing intensity. High grazing tolerance may be achieved through compensatory photosynthesis and leaf growth, or through demographic mechanisms such as activation of a belowground dormant bud bank. This study assessed the relationship between grazing frequency and responses of Schizachyrium scoparium (little bluestem) in a tallgrass prairie, and examined the roles of tiller growth, reproduction, and bud (meristem) populations in its persistence under grazing. Genets were subjected to varying simulated grazing frequencies for a period of 2 years. Strong differential responses were observed among plant traits. RGR, biomass, and flowering showed strong nonlinear reductions in response to increasing clipping frequency, with no evidence of threshold effects. However, meristem density was unaffected, and plants maintained a large bud bank across all clipping treatments. Tiller natality decreased initially, but increased with >4 clippings, suggesting that declines in tiller RGR are partially offset by increasing tiller natality, and that variation in genet size is driven more by demography than by variation in individual tiller growth. Increased grazing frequency also resulted in differential activation of buds at different positions (emerging within vs. outside the subtending leaf sheath), explaining the shift to a more prostrate growth form observed in many caespitose grasses under persistent grazing. Thus, although this grass species lacks the capacity for compensatory foliage re-growth, the maintenance of a large dormant bud bank and the differential activation of buds in different positions contribute to its grazing tolerance and avoidance, respectively, and its long-term persistence in grazed grasslands.     

Grass-to-tree facilitation in an arid grazed environment (Aïr Mountains, Sahara)

The scarcity of tree regeneration is a major concern for the long-term conservation of tree cover in the Aïr-Ténéré Nature Reserve (Sahara, Niger), and the causes of this shortage are poorly understood. We raised the hypothesis that livestock pressure is a key-factor in the mortality of tree seedlings and that facilitation processes among plants may enhance tree regeneration. We tested whether the tussock grass Panicum turgidum could facilitate the regeneration of a keystone tree species, Acacia tortilis var. raddiana, under a grazing regime. We transplanted 3-month-old seedlings of Acacia to plots representing different neighbourhood conditions (in areas without vegetation, amongst intact tussocks, among cropped tussocks, and under branches), which were either fenced or exposed to grazing by large herbivores. Survival and height of 160 transplants were recorded during one year. Acacia performance was much lower in the grazed plots, indicating that large herbivores had a strong negative impact on tree regeneration. Presence of Panicum tussocks had a positive effect on the height of seedlings, likely because they improved soil properties and water availability. No aboveground positive effect on Acacia was detected though the grass did reduce atmospheric aridity. We detected an indirect positive effect of Panicum on Acacia survival in the presence of herbivores when aridity and grazing pressure reached their maximum levels. We obtained evidence that indirect facilitation may occur in arid grazed areas, although it has been predicted to mainly exist in mesic competitive environments. Additionally, our results support the stress-gradient hypothesis against assumptions that competitive effects may dominate in extremely arid environments. Since most of the positive effect on Acacia survival is indirect (biotic grazing refuge) due to the presence of herbivores, transplantation of tree seedlings amongst Panicum may be a simple and low-cost technique for the restoration of Saharan ecosystems degraded by herbivore grazing.