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.     

Lack of intraspecific variation in resistance to defoliation in a grass that evolved under light grazing pressure

Grass species frequently show marked intraspecific variation inmorphology and tolerance to defoliation. Remarkably, most of this evidencecomesfrom grasslands with long evolutionary history of grazing. Here, we explore theintraspecific variation in grazing tolerance and morphometric traits associatedwith grazing avoidance of Paspalum dilatatum (Poir.), agrass from the Flooding Pampa (Argentina), where grazing is a novel disturbancein evolutionary time. We performed a clipping experiment in a greenhouse withtwo populations from sites with contrasting short term grazing regime:continuous grazing and 20 year-old grazing exclosure. The populations did notdiffer in their tolerance to clipping, and showed minor differences in the wayclipping affected plant height, a trait associated with grazing avoidance. Ourresults indicate that there are exceptions to the generalized findings of highlevels of intraspecific variation in grazing resistance among populations ofgrasses. These exceptions may be associated to evolutionary history of grazing.     

Acceptability of plant species along grazing gradients around watering points in Tunisian arid zone

Water points provide excellent sites for studying overgrazing effects on plant communities in dry areas. Distance from water can be considered like a surrogate of grazing pressure being high near the water and low away from it. The main aim of this study is to investigate overgrazing effects on acceptability of fodder plants along a grazing gradient around three natural watering points. To achieve this goal, we classified spontaneous plants according to their acceptability degree and we followed their cover, richness and density as well as the grazing value along a grazing gradient around these wells, using phyto-ecological studies during the spring 2004 and 2006. Main results show that very palatable plants (mainly constituted by annuals) are more dominant in both the closed and the more disturbed transect areas around wells. The unpalatable plants dominate sites with moderate disturbance around wells. Ligneous palatable species obviously have a lower degree of disturbance. During the studied seasons the grazing gradient around wells 1 and 2, the oldest ones, seemed to exert a feedback upon the grazing intensity.     

Determining suitable grazing time for Puccinella distans Parl. based on its phenology in West Azerbaijan Province of Iran

Studying the phenology, which is an important issue of plants, can be used for determining the best time of all kinds of utilizations, and also conservation and stability of plants’ growth and breeding will result. Therefore, this study was performed in Tezkharab rangelands, where Puccinella distans is an important vegetative element that is consumed by livestock, to determine the suitable grazing time for this plant based on its phenology, for about a 4-year period (2007–2010). For this purpose, plant phenology and changes in its growth characteristics were investigated weekly. Moreover, during this period, all phenological stages were adjusted to the monthly precipitation and temperature. The results indicated that the time of phenological stages had fluctuations which were related to environmental conditions. The best time to start grazing is early May and the best time for collecting the seeds of this species is late July to early August.     

Influence of rainfall and grazing on herbage dynamics in a seasonally dry tropical savanna

Species composition and herbage dynamics in relation to rainfall variability and cattle grazing were studied in permanently protected, grazed, and temporarily fenced treatments on three sites in a seasonally dry tropical savanna. Permanently protected sites, established between 1979 and 1984, were 55–79% similar with each other in species composition, and 14–25% similar with grazed sites during the period 1986–1988. Similarity among grazed sites was only 36–43%. Number of species was greater in the grazed treatment than in the permanently protected treatment. The percentages of annual grasses and non-leguminous forbs were greater in grazed savanna than in permanently protected savanna. Species diversity was higher in grazed savanna than in the corresponding permanently protected savanna. Species the two annual cycles studied, peak live shoot biomass was 614 g m^-2 in permanently protected savanna, 109 g m^-2 in grazed savanna, and 724 g m^-2 in temporarily fenced savanna. Live shoot biomass in temporarily fenced savanna was 18 to 44% greater than in permanently protected savanna. Peak canopy biomass ranged from 342 to 700 g m^-2 in permanently protected savanna. It was related with total rainy season rainfall, and was particularly sensitive to late rainy season rainfall. On the other hand, peak canopy biomass in grazed savanna ranged from 59 to 169 g m^-2 and was related to grazing intensity rather than either total rainy season rainfall or late rainy season rainfall. Coefficient of variation of green biomass in permanently protected savanna was related with rainfall variability indicating it to be a pulsed system which responds quickly to rainfall events. Biomass of woody species ranged from 2466 to 5298 g m^-2 in permanently protected savanna and from 744 to 1433 g m^–2 in the grazed savanna. Green foliage biomass was 3.7 to 6.4% of the woody biomass in permanently protected and 5.6 to 5.9% in grazed savanna, and supplements substantially the fodder resource during the dry periods of the year.     

Germination and population structure of spear thistle Cirsium vulgare in relation to experimentally controlled sheep grazing

Summary The size and number of rosettes of Cirsium vulgare were censused in a 4 ha sheep grazing trial on lowland calcareous grassland in paddocks receiving controlled winter, spring and summer grazing treatments. Spring grazing significantly increased thistle rosette numbers, and there were always fewer rosettes in size classes 250mm–300 mm under the heavier grazing treatments. The emergence of seeds of Cirsium vulgare sown by hand into each of the grazing treatments was monitored and showed a positive effect of spring grazing. When the percent emergence of sown seeds was used as a covariate in the analysis of rosette numbers in the experimental paddocks, it was found to account for 77% of the variance in thistle numbers. It was concluded that the main effect of grazing on thistle rosette numbers is an indirect one exercised through the effects of grazing on germination conditions. Seeds of Cirsium vulgare were sown into artificial gaps at another grassland site, to determine the effect of gap-size upon seedling emergence. Germination was poor, but significantly more seedlings emerged in gaps 10–20 cm diameter, than in 5 cm gaps or in control plots without a gap. A computer simulation model was used to explore the relationship between gap density and thistle population dynamics. A threshold density of gaps was found to exist, below which thistle populations went extinct, and above which the thistle population grew geometrically. The degree of aggregation of dispersed seeds did not alter the threshold gap-density for plant extinction, but did affect the rate of increase of the thistle population when the threshold gap-density was exceeded. It is concluded that strategies for controlling Cirsium vulgare populations by grazing manangement will be most effective if aimed at reducing suitable sites for establishment in spring. The success of such attempts will depend upon the soil fertility of the site, and sudden outbreaks of Cirsium vulgare infestation can be expected if a sward gradually deteriorates through over-grazing.     

Determining the growing season of land vegetation on the basis of plant phenology and satellite data in Northern China

The objectives of this study are to explore the relationships between plant phenology and satellite-sensor-derived measures of greenness, and to advance a new procedure for determining the growing season of land vegetation at the regional scale. Three phenological stations were selected as sample sites to represent different climatic zones and vegetation types in northern China. The mixed data set consists of occurrence dates of all observed phenophases for 50–70 kinds of trees and shrubs from 1983 to 1988. Using these data, we calculated the cumulative frequency of phenophases in every 5-day period (pentad) throughout each year, and also drew the cumulative frequency distribution curve for all station-years, in order to reveal the typical seasonal characteristics of these plant communities. The growing season was set as the time interval between 5% and 95% of the phenological cumulative frequency. Average lengths of the growing season varied between 188 days in the northern, to 259 days in the southern part of the research region. The beginning and end dates of the surface growing season were then applied each year as time thresholds, to determine the corresponding 10-day peak greenness values from normalized difference vegetation index curves for 8-km² pixels overlying the phenological stations. Our results show that, at the beginning of the growing season, the largest average greenness value occurs in the southern part, then in the northern, and finally the middle part of the research region. In contrast, at the end of the growing season, the largest average greenness value is measured in the northern part, next in the middle and lastly the southern part of the research region. In future studies, these derived NDVI thresholds can be applied to determine the growing season of similar plant communities at other sites, which lack surface phenological data. Received: 29 November 1999 / Revised: 14 March 2000 / Accepted: 15 March 2000     

The role of annual grasses in the phenology of Rhopalosiphum padi in the low rainfall belt of South Australia

Rhopalosiphum padi (L.) (Hemiptera: Aphididae) was the first species of cereal aphids to colonise annual grasses across the state each year. Numbers were higher in the Lower Murray Valley than in the Mount Lofty Ranges or Adelaide Plains. At all locations, numbers generally increased until grasses reached the boot stage after which they declined. Alatoid production commenced in late July in response to crowding, but thereafter plant age and photoperiod were more important determinants. Metopolophium dirhodum (Walker) (Hem-pitera: Aphididae) and Sitobion nr fragariae (Hemiptera: Aphididae) first appeared in annual grasses in late July.     

Effects of precipitation and soil water potential on drought deciduous phenology in the Kalahari

We utilized an ecosystem process model to investigate the influence of precipitation and soil water potential on vegetation phenology in the semi‐arid, drought‐deciduous ecosystems in the Kalahari region of South Africa. The timing of leaf flush was assumed to be the first day during which a rainfall event exceeded that day's estimate of potential evapotranspiration after a defined dry season. Leaf senescence was assumed to be a dynamic feedback between soil water potential and net plant carbon gain and was determined by dynamically modeling the effects of concomitant trends in soil water potential and net primary production on leaf area index (LAI). Model predictions of LAI were compared with satellite‐derived normalized difference vegetation indices (NDVI) for 3 years at two sites along the Kalahari transect. The mean absolute error for the prediction of modeled leaf flush date compared with leaf flush dates estimated from NDVI were 10.0 days for the Maun site and 39.3 days for the Tshane site. Correlations between model predicted 10‐day average LAI and 10‐day composite NDVI for both Maun and Tshane were high (ρ=0.67 and 0.74, respectively, P<0.001), suggesting that this method adequately predicts intra‐annual leaf area dynamics in these dry tropical ecosystems.     

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.     

The impact of grazing pressure in clearfelled,burned and undisturbed eucalypt forest

An area of dry eucalypt forest on dolerite in southeastern Tasmania with comparable sites that were a) unlogged, b) clearfelled, c) clearfelled and slash-burned, was monitored over a 24 month period. Detailed records were made of plant species composition and cover on floristically similar fenced and unfenced permanent plots in two major vegetation types. Mammal faecal pellets were cleared from each of the unfenced plots and subsequently counted and attributed to species of origin. For the six combinations of treatment and vegetation type pellet deposition rate was highly correlated with differences in the rate of change of species cover between fenced and unfenced plots and the lifeform composition of the vegetation. The greatest changes were observed in the clearfelled, burned area, while there was relatively little change in the undisturbed forest. Grass and herb species were generally disadvantaged by mammal exclusion while shrubs and graminoids generally grew faster in the fenced plots.Nomenclature of plant species follows Curtis & Morris (1975).This project was supported financially by the Forest Ecology Research Fund, Tasmania.     

Coexistence of multiple parasitoids on a single host due to differences in parasitoid phenology

There are many well-documented cases in which multiple parasitoids can coexist on a single host species. We examine a theoretical framework to assess whether parasitoid coexistence can be explained through differences in timing of parasitoid oviposition and parasitoid emergence. This study explicitly includes the phenology of host and parasitoid development and explores how this mechanism affects the population dynamics. Coexistence of the host with two parasitoids requires a balance between parasitoid fecundity and survival and occurs most readily if one parasitoid attacks earlier but emerges later than the other parasitoid. The host density can either be decreased or increased when a second coexisting parasitoid is introduced into the system. However, there always exists a single parasitoid type that is most effective at depressing the host density, although this type may not be successful due to parasitoid competition. The coexistence of multiple parasitoids also affects the population dynamics. For instance, population oscillations can be removed by the introduction of a second parasitoid. In general, subtle differences in parasitoid phenology can give rise to different outcomes in a host–multi-parasitoid system, and this may offer some insight into why establishing criteria for the ‘ideal’ biological control agent has been so challenging.     

The effect of time-varying mortality and carbon assimilation on models of carbon allocation in annual plants

Summary Models of optimal carbon allocation schedules have influenced the way plant ecologists think about life history evolution, particularly for annual plants. The present study asks (1) how, within the framework of these models, are their predictions affected by within-season variation in mortality and carbon assimilation rates?; and (2) what are the consequences of these prediction changes for empirical tests of the models? A companion paper examines the basic assumptions of the models themselves. I conducted a series of numerical experiments with a simple carbon allocation model. Results suggest that both qualitative and quantitative predictions can sometimes be sensitive to parameter values for net assimilation rate and mortality: for some parameter values, both the time and size at onset of reproduction, as well as the number of reproductive intervals, vary considerably as a result of small variations in these parameters. For other parameter values, small variations in the parameters result in only small changes in predicted phenotype, but these have very large fitness consequences. Satisfactory empirical tests are thus likely to require much accuracy in parameter estimates. The effort required for parameter estimation imposes a practical constraint on empirical tests, making large multipopulation comparisons impractical. It may be most practical to compare the predicted and observed fitness consequences of variation in the timing of onset of reproduction.     

The significance of waterfowl grazing in the primary vegetation succession on embanked sandflats

In the annual stage of a primary plant succession on desalinating sandflats of the Lauwerszeepolder, N Netherlands, grazing by waterfowl is a dominant factor, as is illustrated by exclosures. It is concluded from population dynamical data, that both the removal of biomass and the predation of seeds sustain the annual species and retard succession to wards the next stage of a perennial grassland by several years. Yearly mowing in September slightly enhances succession by selectively affecting the dominant annual Salicornia species. The recurrence of high salinities in the upper soil layer of bare, c.q. grazed surfaces is expected to limit the invasion of grass species and other perennials.Nomenclature of plant taxa follows Heukels/van der Meijden (1983), Flora van Nederland, 20th ed. Wolters-Noordhoff, Groningen.This article would not have been written without the contributions of many students and colleagues, equally aware of the rare opportunity we had to study a genuine primary succession. I wish to thank them all. The help in the moving experiment of J. Franke, S. Nijdam, the late M. Robertus, H. Vrieling and J. D. D. Hofman is gratefully acknowledged. Thanks are due to the State Authority of the IJssellake Polders, for permitting access to and use of the experimental areas.     

Adaptive phenology of desert and Mediterranean populations of annual plants grown with and without water stress

Summary The dynamics of vegetative and reproductive growth were compared in matched pairs of Mediteranean and desert populations of three unrelated annual species, Erucaria hispanica (L.) Druce, Brachypodium distachyon (L.) Beauv. and Bromus fasciculatus C. Presl., under high and low levels of water availability in a common-environment experiment. Plants of all desert populations showed earlier switches to reproductive development and to subsequent phenophases, and the transition to flowering occurred at smaller plant sizes. Water stress had no effect (E. hispanica) or slightly accelerated the transition to flowering in B. fasciculatus (by 1–2 days) and in B. distachyon (by 4–6 days). Plant senescence was strongly enhanced by water stress, and this enhancement was greater in desert populations than in corresponding Mediterranean ones. Duration of life cycle was greatly shortened by water stress in all three species. Desert and Mediterranean populations of the three species exhibited small differences in their relative response, i.e. phenotypic plasticity, to water stress for phenological and plant size parameters. In E. hispanica and B. fasciculatus the population x water regime interaction amounted to less than 3% of total variance. By contrast, the Mediterranean population of B. distachyon was much more plastic in its response to water stress than the desert population in its transition to plant senescence. Plants from the desert populations appeared to be adapted to shorter, more compact growth cycles, culminating in earlier dates of seed maturation and plant senescence. In addition, they showed larger phenotypic plasticity in the transition to plant senescence, which trait was enhanced or magnified by sustained or repeated lack of water. By contrast, plants from Mediterranean populations delayed switchover from one phenophase to the next, seeming thus to bet on more water being forthcoming.     

Optimal invasive species management under multiple uncertainties

The management programs for invasive species have been proposed and implemented in many regions of the world. However, practitioners and scientists have not reached a consensus on how to control them yet. One reason is the presence of various uncertainties associated with the management. To give some guidance on this issue, we characterize the optimal strategy by developing a dynamic model of invasive species management under uncertainties. In particular, focusing on (i) growth uncertainty and (ii) measurement uncertainty, we identify how these uncertainties affect optimal strategies and value functions. Our results suggest that a rise in growth uncertainty causes the optimal strategy to involve more restrained removals and the corresponding value function to shift up. Furthermore, we also find that a rise in measurement uncertainty affects optimal policies in a highly complex manner, but their corresponding value functions generally shift down as measurement uncertainty rises. Overall, a rise in growth uncertainty can be beneficial, while a rise in measurement uncertainty brings about an adverse effect, which implies the potential gain of precisely identifying the current stock size of invasive species.