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.     

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.     

Vegetative structure,microclimate, and leaf growth of a páramo tussock grass species,in undisturbed,burned and grazed conditions

In neotropical alpine grasslands (páramo), the natural tussock grass vegetation is extensively grazed and occasionally burned. The low productivity of the tussock grass seems to be the reason for the disappearance of this growth form in the most frequently intervened areas. The structure, microclimate and leaf elongation rates of new emerging leaves were studied for the dominant tussock grass species Calamagrostis effusa, at an undisturbed, a moderately grazed (7 year after fire) and a heavily grazed (3.5 years after fire) site. In absence of grazing and burning, the tussocks had a high standing crop (1.07±0.09 kg DW · m^-2) and leaf area per projected tussock cover (LAI: 9.6±1.4). Two thirds of the total mass was dead and more than half of the leaves were in horizontal position. The tussock growth form protects the meristems from severe climatic conditions. At midday, the temperature was higher at meristem level than in the rest of the tussock. At this level, photosynthetic irradiance (PI) was almost extinct at 2.9±0.74% of PI above the vegetation. The red/far red ratio (R/FR) was strongly decreased. Initial leaf elongation of new born leaves was 2.3 mm · day^-1, and constant during the year; estimated net annual production was 198±73.8 g m^-2. At the moderately grazed and the heavily grazed study sites, the tussocks were smaller, greener and more erect than those at the undisturbed site. More PI reached the meristems and R/FR was higher at the base of grazed tussocks. Leaf elongation rates were lower. Most of the litter disappeared during the fires. The lower elongation rate of leaves in the grazed areas might be a response to defoliation, resulting in increased tillering and a lack growth associated with poor temperature insulation and more UV-B damage.     

Does selective defoliation mediate competitive interactions in a semiarid savanna? A demographic evaluation

Abstract. Three patterns of target-neighbor plant defoliation were imposed on a late-seral, perennial, C₄-grass, Bouteloua curtipendula, in three long-term grazing regimes to determine the influence of selective defoliation on competitive interactions and species replacement in a semiarid savanna on the Edwards Plateau, Texas, USA. Short-term (3-yr) target plant defoliation did not significantly affect either tiller or plant responses in any of the three grazing regimes. Neighbor plant defoliation, either alone or in combination with target plants, produced a significant defoliation interaction with time for tiller number and basal area per plant, but not for tiller recruitment or mortality. The minimal effect of selective defoliation on the intensity of competitive interactions in this semiarid community indicates that selective grazing has a less definitive role in mediating herbivore-induced species replacement than it does in mesic grasslands and savannas. This interpretation is discussed within the context of long-term (45-yr) change in herbaceous vegetation associated with grazing in this community. Cumulative tiller recruitment in the intensively grazed regime was only 44% of that in the ungrazed regime because of greater plant mortality and fewer surviving plants that recruited tillers. Target plant mortality (50%) only occurred in the intensively grazed regime and the proportion of target plants that initiated tillers decreased by 70, 48 and 32% in the ungrazed, moderately and intensively grazed regimes, respectively, during the final two years of the investigation. The decrease in cumulative tiller recruitment in all grazing regimes was probably mediated by a drought-induced increase in median tiller age the second year of the study. However, tiller per tiller recruitment rate among plants that recruited at least one tiller remained relatively constant among grazing regimes and years. Intensive, long-term grazing has modified the population structure of this late-seral perennial grass to the extent that population responses to both herbivory and periodic drought have been altered in comparison with those of ungrazed and moderately grazed populations. Ecological consequences of a herbivore-induced transition in population structure may be to minimize the effect of selective herbivory on competitive interactions and to function as an avoidance mechanism to reduce the probability of localized population extinction in response to intensive long-term herbivory.     

Soil and plant water relations in a crested wheatgrass pasture: response to spring grazing by cattle

Summary Few field studies have attempted to relate effects of actual livestock grazing on soil and plant water status. The present study was initiated to determine the effects of periodic defoliations by cattle during spring on soil moisture and plant water status in a crested wheatgrass (Agropyron cristatum (L.) Gaertn. and A. desertorum (Fisch. ex Link) Schult.) pasture in central Utah. Soil moisture in the top 130 cm of the soil profile was depleted more rapidly in ungrazed plots than in grazed plots during spring and early summer. Soil moisture depletion was more rapid in grazed plots in one paddock after 1 July due to differential regrowth, but there was no difference in soil water depletion between plots in another paddock during the same period. This difference in soil water depletion between paddocks was related to a difference in date of grazing. Although more water had been extracted from the 60 cm to 130 cm depths in ungrazed plots by late September, cumulative soil moisture depletion over the entire 193 cm profile was similar in grazed and ungrazed plots. Prior to 1 July, grazing had no effect on predawn leaf water potentials as estimated by a pressure chamber technique; however, after 1 July, predawn leaf water potentials were lower for ungrazed plants. Midday leaf water potentials were lower for grazed plants before 1 July, but did not differ between grazed and ungrazed plants after 1 July. A 4- to 8-day difference in date of defoliation did not affect either predawn or midday leaf water potentials. The observed differences in water use patterns during spring and early-summer may be important in influencing growth and competitive interactions in crested wheatgrass communities that are subject to grazing by domestic livestock.     

Effects of bison grazing on Andropogon gerardii and Panicum virgatum in burned and unburned tallgrass paririe

Summary Responses to clipping and bison grazing in different environmental contexts were examined in two perennial grass species, Andropogon gerardii and Panicum virgatum, on the Konza Prairie in northeastern Kansas. Grazed tillers had lower relative growth rates (RGR) than clipped tillers following defoliation but this difference was transient and final biomass was not affected by mode of defoliation. Grazed tillers of both species had higher RGR throughout the season than ungrazed tillers, resulting in exact compensation for tissue lost to defoliation. However, A. gerardii tillers which had been grazed repeatedly the previous year (1988) had reduced relative growth rates, tiller biomass and tiller survival in 1989. This suggests that the short-term increase in aboveground relative growth rates after defoliation had a cost to future plant growth and tiller survival.In general, the two species had similar responses to defoliation but their responses were altered differentially by fire. The increase in RGR following defoliation of A. gerardii was relatively greater on unburned than burned prairie, and was influenced by topographic position. P. virgatum responses to defoliation were similar in burned and unburned prairie. Thus grazing, fire, and topographical position all interact to influence tiller growth dynamics and these two species respond differently to the fire and grazing interaction. In addition, fire may interact with grazing pattern to influence a plants' grazing history and thus its long-term performance.     

Growth and tillering patterns within tussocks of Eriophorum vaginatum

Growth patterns were investigated for Eriophorum vaginatum tussocks from disturbed and undisturbed tussock tundra at two sites in Alaska. Total basal area of tussocks decreased with increased cryoturbation but mean basal area per tussock did not. Flowering was observed in tussocks of significantly smaller size on disturbed compared to undisturbed tundra. For tussocks with < 10% cover by shrubs and moss, number of tillers per tussock was linearly related to tussock diameter in most disturbed and undisturbed sites. Exceptions occurred in an area that had been bladed with a bulldozer 7 yr before our survey where tiller number increased as the square of diameter and in an area with much frost activity where tiller number was not emulated with diameter. The ratio of daughter tillers to adult tillers decreased with diameter in disturbed tundra, whereas the trend was less pronounced in undisturbed tundra.
Microsuccession in undisturbed tussock tundra was investigated by sampling tussocks with different amounts of shrub and moss cover for number of daughter tillers per adult tiller, weight per tiller, percent nitrogen, and percent phosphorus. Small tussocks without cover by other species and large, partially covered tussocks were not significantly different by any measures, but tussocks that were almost completely buried had significantly (p < 0.05) lower values of tillering index, weight per tiller, and percent phosphorus.     

Rapid recovery of kohekohe (

Soil conditions, vegetation features and soil fauna were recorded in montane tall tussock grassland dominated by narrow- leaved snow tussock Chionochloa rigida ssp. rigida up to 30 months after a spring fire. Burning reduced the stature of tussocks and the size and density of tillers in the first growing season. After two growing seasons, tussock canopy development and tiller size remained below those found in the unburnt grassland nearby. New tillers and tussocks established following the prolific fire-induced flowering one year after burning. After the fire and sheep grazing, intertussock cover became progressively dominated by introduced grasses and herbs. While soil pH, moisture content, bulk density, surface litter and total nematodes showed significant treatment (burning) effects, these properties also showed significant year-to-year variation. The greatest increase in any nematode group was in Paratylenchus, a distinctive genus widespread in tussock grasslands and apparently responsive to environmental fluctuation and root development; its population was 100x and 29x greater in the burned area than in the control area 16 and 30 months after burning. Subject to detailed testing, populations of mites and collembola may provide relatively simple indicators of recovery of ecosystem function of such grasslands after burning.     

Growth and carbon allocation of Agropyron desertorum following autumn defoliation

Summary Growth and carbon allocation of a cool season tussock grass, Agropyron desertorum, following defoliation of newly initiated tillers in the autumn of 1988 and 1989 were investigated. Tiller density and mortality, reproductive shoot density, root density, biomass, individual tiller weight, carbon allocation, and soil water depletion were used to evaluate the response of A. desertorum to autumn grazing. Tiller recruitment was lower in the autumn-defoliated treatment in both years compared with the control because of the cessation of tiller development following autumn defoliation. Autumn defoliation also significantly reduced the movement of ¹³C to the roots in 1988 but not in 1989. Soils were cooler and drier in 1989. Other plant growth measurements and soil water depletion rates were not different between treatments. Autumn defoliation in 1988 did not influence tiller recruitment in the following autumn. Two consecutive years of autumn defoliation did not affect tiller overwinter mortality or peak standing crop in 1990.     

Demography of three dominant sedges under contrasting grazing regimes in the High Arctic

Abstract. Tiller demography of Carex aquatilis ssp. stans, Carex membranacea, and Eriophorum angustifolium ssp. triste was investigated in ungrazed and grazed high arctic vegetation on central Ellesmere Island, Canada. Tiller birth, growth, flowering and death were studied from excavated clonal fragments, and tiller density and biomass were studied from excavated turfs. Five life‐cycle stages were determined: dormant buds, juvenile, mature, flowering and dead tillers. A stage‐based transition matrix model was developed to estimate the long‐term dynamics of the sedge populations and to compare life‐history strategies between ungrazed and grazed populations. Short‐term and retrospective models, based on the growth during the sampling year and during the lifetime of the clonal fragments, respectively, were compared to see how well the short‐term model can describe demography of long‐lived plants. According to the short‐term model, tiller populations were decreasing (λ < 1 except for C. membranacea), whereas the retrospective model indicated that the tiller populations were increasing. Tiller population growth rates did not differ between ungrazed and grazed habitats. Nevertheless, the similar growth rates may be obtained by balanced differences in the vital rates between plants of the two habitats. The plants in the ungrazed habitat tended to remain in their current life‐cycle stage, whereas plants in the grazed habitat moved quickly to the next stage and died earlier. C. aquatilis ssp. stans appears to gain a competitive advantage over the other species under intensive grazing, as indicated by the higher tiller density and greater below‐ground biomass in grazed vegetation. The greater amount of below‐ground biomass apparently buffers C. aquatilis ssp. stans against grazing better than the other species.     

Grazing decisions of Soay sheep,Ovis aries,on St Kilda: a consequence of parasite distribution?

Relationships between the height of grass swards and herbage intake have been established for a number of large herbivores and sward height plays a major role in determining the selection of herbivore diets. However, tall grass swards also represent a more stable damp microclimate for gastrointestinal parasite larvae and tall swards contain generally greater numbers of parasites than short swards. Herbivores may then trade-off the need to maximise nutrient intake through the selection of tall swards with the need to avoid parasite ingestion. Stratified sward sampling techniques were used to determine the distribution of nutrients and parasites in a heterogeneous sward structure on the island of Hirta, St Kilda, which is grazed by a population of feral Soay sheep. The disparity between the short gap vegetation and the tall tussock vegetation was greatest in spring, when gap vegetation was some three-fold shorter than tussock vegetation; this led to tussocks offering greater nutrient and energy intake rate when compared to gap vegetation in spring. Parasites were concentrated in tussocks in spring, thus creating trade-offs. Such trade-offs were not present in summer when parasites were more evenly distributed across the gap/tussock sward structure and the nutritional advantages associated with grazing tussocks were diminished. The diet selection of the resident population of sheep was determined by recording the number of bites taken from gap and tussock vegetation by randomly selected focal animals over repeated 5-min grazing periods. Overall, all animals avoided grazing tussocks, which were most strongly avoided in the spring, and older animals avoided grazing tussocks to a greater extent. Overall, females with lambs and males avoided tussocks to a similar degree and both avoided them less than barren females. Faecal egg counts (FEC) of female sheep were negatively correlated with tussock selectivity and vice versa for male sheep. The interaction between the grazing behaviour of each sex and FEC is discussed in relation to the immunocompetence and nutrient requirements of the different sexes. The maximum disparity between the costs and benefits of the trade-off studied occurred in late winter/early spring which also represents the time of greatest nutrient demand in the Soay sheep, since many are close to starvation and/or are in the advanced stages of pregnancy. Grazing decisions of individuals at this time, determining relative nutrient and parasite intake, may then be related to subsequent fitness and survival.     

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     

Effects of livestock exclusion on density,survival and biomass of the perennial sagebrush grass Hymenachne pernambucense (Poaceae) from a temperate fluvial wetland

In Argentina, the intensification of soybean production has displaced a substantial proportion of cattle ranching to fluvial wetlands such as those in the Delta of the Paraná River. Cattle grazing affects structure and dynamics of native forage plants but there is little information on this impact in populations from fluvial wetlands. This study addresses the effect of cattle ranching on density, survival, mean life-span and aerial biomass of Hymenachne pernambucense (Poaceae), an important forage species in the region. The study was carried out monthly for one year in permanents plots subject to continuous grazing and plots excluded from grazing in the Middle Delta of the Paraná River. In plots excluded from grazing, tillers showed significantly higher population density and survival, and a two-fold increase in mean life-span, while continuous grazing decreased survival of cohorts. The largest contribution to tiller density in ungrazed and grazed populations was made by spring and summer cohorts, respectively. Total and green biomass were significantly higher in the ungrazed population, with highest differences in late spring-early summer. Cattle grazing affected the relationship between tiller density and green biomass suggesting that cattle prefer sprouts because they are more palatable and nutritious than older tissue.     

Morphological Analysis of Tillering in Agropyron spicatum and Agropyron desertorum

The caespitose grasses Agropyron spicatum and Agropyron desertorumexhibit a striking difference in tillering response followingexperimental clipping treatment, with plants of A. desertorumproducing up to 18 times more tillers. The two species are similarin many aspects of their phenology and physiology. Previousexamination of current photosynthate production and levels ofstored carbohydrates indicate only slight differences betweenthe species. The possible role of three anatomical/morphologicalconstraints in controlling tillering was examined. No evidencefor such constraints was found. A basal cluster of buds is presenton the parent tillers. The mean bud number per tiller was similarfor both species and the range (3–9) was identical. Nearlyall of the bud apical meristems appeared anatomically viablethroughout the growing season and vascular development occurredto within 250 to 490 µm of the various bud apices of bothspecies. Both normal fall tillers and summer tillers producedunder clipping treatment originated from the largest, most distalbuds of the basal cluster of buds. However, precocious, morphologicallydistinctive, second-order tillers occasionally grew out fromthe smaller, most basal buds of some elongating fall tillers. Agropyron spicatum, Agropyron desertorum, bluebunch wheatgrass, crested wheatgrass, bud, tiller, tillering ability, meristematic potential, vascular development, regrowth     

Functional morphology and microclimate of Festuca orthophylla, the dominant tall tussock grass in the Andean Altiplano

Plant growth is driven by the rate of photosynthetic uptake of carbon, the loss of carbon and by allocation of photoassimilates to certain plant compartments, which leads to particular morphologies. Performance, vitality and persistence of a plant are affected by this partitioning process and vice versa. Under harsh climatic conditions such as cold temperature and seasonal drought, perennial plants often invest more in below-ground than above-ground structures. Festuca orthophylla in the subtropical Bolivian Altiplano does not match this ‘rule’. This species produces tall, evergreen tussocks, persisting decades and dominating the semi-arid, Andean landscape over thousands of square kilometers at elevation between 3600 and 4600 m a.s.l. The shallow rooting system represents only 21% of total biomass. The tussock base (root-stocks composed of the network of branching below-ground shoots and tiller meristems) comprises 28% of the total biomass. Although located partly below the soil surface, much of this biomass compartment is functionally above-ground (the basis of shoots). With their below-ground position, tiller meristems are protected against grazing and trampling by camelids as well as, to some degree, against fire and freezing. Fifty one percent of the biomass is above-ground (live leaves and inflorescences). In terms of phytomass (including attached necromass), 75% is above-ground. On average, a tussock consists of 3200 tightly packed total tillers (56% are live). Tillers emerge regularly intravaginally (i.e. within the leaf sheath of an existing mother tiller), resulting in dense canopies with strong self-shading: eighty percent of green foliage experience less than 50% of the incident light. The most important Altiplano plant species thus has morphological traits in favour of protection and survival rather than productivity.     

Negative Plant–Soil Feedback and Positive Species Interaction in a Herbaceous Plant Community

Increasing evidence shows that facilitative interaction and negative plant–soil feedback are driving factors of plant population dynamics and community processes. We studied the intensity and the relative impact of negative feedback on clonal growth and seed germination of Scirpus holoschoenus, a ‘ring’ forming sedge dominant in grazed grassland, and the consequences for species coexistence. The structure of aboveground tussocks was described. A Lithium tracer assessed belowground distribution of functional roots. Seed rain and seedling emergence were compared for different positions in relation to Scirpus tussocks. Soil bioassays were used to compare growth on soil taken from inside and outside Scirpus tussocks of four coexisting species (Mentha acquatica, Pulicaria dysenterica, Scirpus holoschoenus and Dittrichia viscosa). We also compared plant performance of dominant plant species inside and outside Scirpus tussocks in the field. The ‘ring’ shaped tussocks of S. holoschoenus were generated by centrifugal rhizome development. Roots were functional and abundant under the tillers and extending outside the tussocks. The large roots mats that were present in the inner tussock zone were almost all dead. Seedling emergence and growth both showed a strong negative feedback of Scirpus in the inner tussock zone. Scirpus clonal development strongly reduced grass biomass. In the degenerated tussock zone, Pulicaria and Mentha mortality was lower, and biomass of individual plants and seed production were higher. This positive indirect interaction could be related to species-specific affinity to soil conditions generated by Scirpus, and interspecific competitive release in the degenerated tussock zone. We conclude that Scirpus negative feedback affects its seedling emergence and growth contributing to the development of the degenerated inner tussock zone. Moreover, this enhances species coexistence through facilitative interaction because the colonization of the inner tussock zone is highly species-specific.     

Characteristics of successful competitors: an evaluation of potential growth rate in two cold desert tussock grasses

Summary Within the first few weeks after seedling emergence, Agropyron desertorum, a more competitive tussock grass, had a much higher mean relative growth rate (RGR) than Agropyron spicatum, a very similar, but less competitive species. However, beyond the early seedling stage, the two grasses had a remarkably similar whole-plant RGR in hydroponic culture and aboveground RGR in glasshouse soil, if root temperatures were above approximately 12°C. At soil temperatures between 5 and 12°C, A. desertorum exhibited a 66% greater aboveground RGR than A. spicatum (P<0.05). Both species responded similarly to warming soil temperatures. In the field, however, tiller growth rates were generally similar. Neither species showed marked tiller elongation until a couple of weeks after snowmelt, by which time soil temperatures, at least to a depth of 10 cm, were above 12°C for a significant portion of the day. Aboveground biomass accumulation over a three-year period indicated that both grasses had similar potential growth rates whereas Artemisia tridentata ssp. vaseyana, a common neighbor planted in the same plots, had a much greater potential growth rate. The greater competitive ability of adult A. desertorum, as compared to A. spicatum, cannot be attributed to appreciable differences in potential growth rates.     

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.     

Compensatory growth of the kelp Macrocystis integrifolia (Phaeophyceae, Laminariales) against grazing of Peramphithoe femorata (Amphipoda, Ampithoidae) in northern-central Chile

Compensation of tissue loss has been considered an alternative strategy for seaweeds that have no or only minor chemical or structural defense against herbivory. Compensatory responses are facilitated by resource transfer among different tissues and have been suggested for large kelps. Macrocystis integrifolia (Bory) is a common kelp species from northern-central Chile, which is characterized by high growth rates and the absence of lipophilic chemical defenses against herbivore grazing. Herein, we used the giant kelp M. integrifolia to test for compensatory growth in response to grazing by the nest-dwelling amphipod Peramphithoe femorata (Krøyer). Amphipods were allowed to graze inside nests on subapical blades of M. integrifolia sporophytes for 14 days. We measured growth and chemical composition (C, N, laminaran and mannitol) of apical and subapical blades of grazed and ungrazed (control) sporophytes. Our results revealed the capability of M. integrifolia to maintain elongation rates in grazed subapical blades, which were similar to those of subapical blades from ungrazed sporophytes. Apical blades grew slower in grazed than in ungrazed sporophytes indicating a trade-off between apical and subapical blades when herbivores were present. Thus, compensation occurs in blades directly attacked by grazers and is probably mediated by vertical resource allocation within sporophytes to subapical blades, a suggestion supported by the fact that stipe internodes in these regions grew more on grazed sporophytes. In general, our study indicates that M. integrifolia exhibits compensatory growth against the herbivore amphipod P. femorata, and we suggest that this could be an important strategy of large kelp species to tolerate moderate grazing intensities.     

A numerical study of successions in an abandoned,damp calcareous meadow in S Sweden

Summary Two sites at Örup, SE skåne, Sweden, have been investigated, viz. a grazed, unimproved, tussocky pasture on a calcareous moraine clay; and an originally similar, adjacent area that was abandoned about 1960. On the grazed site the vegetation is extremely rich in species. This vegetation type was formerly widespread, but nowadays it is rare and therefore it is important to try to conserve examples of it. In total, 80 0.1 m² quadrates distributed at random within 4 homogeneous plots were investigated. The species composition and the cover, the height of the vegetation, the position on a tussock or in a depression and the amount of litter were recorded. The cover data were ordinated (PCA). The differentiation between tussocks and depressions and the effects of ceasing grazing were clearly separated. The connections between the vegetation and the position of the quadrat, grazing intensity etc. were investigated. The vegetation of the ungrazed parts had become dominated by Filipendula ulmaria and Carex disticha; on an average there were 5.5 species/0.1 m². The grazed quadrats contained three times as many species and showed much higher spatial variation, important species being Carex panicea, C. flacca, Molinia caerulea, Festuca ovina, Potentilla erecta, Centaurea jacea and Serratula tinctoria.Nomenclature of vascular plants follows Lid (1974), for names of bryophytes see Nyholm (1954–69) and Arnell (1956).This paper is mainly based on a more detailed paper in Swedish (Regnéll 1979).I thank Dr. E. van der Maarel, Prof. Nils Malmer, Fil. lic. Anders Larsson, Fil. Dr. Eva Waldemarsson-Jensén and Fil. kand. Stefan Persson for valuable discussions and encouragement.