Recovery and succession in a multi-species tropical seagrass meadow following experimental disturbance: the role of sexual and asexual reproduction

Recolonisation and succession in a multi-species tropical seagrass meadow was examined by creating gaps (50×50 cm) in the meadow and manipulating the supply of sexual and asexual propagules. Measurements of leaf shoot density and estimates of above-ground biomass were conducted monthly to measure recovery of gaps between September 1995 and November 1997. Measurements of the seeds stored in the sediment (seed bank) and horizontal rhizome growth of colonising species were also conducted to determine their role in the recovery process. Asexual colonisation through horizontal rhizome growth from the surrounding meadow was the main mechanism for colonisation of gaps created in the meadow. The seed bank played no role in recolonisation of cleared plots. Total shoot density and above-ground biomass (all species pooled) of cleared plots recovered asexually to the level of the undisturbed controls in 10 and 7 months, respectively. There was some sexual recruitment into cleared plots where asexual colonisation was prevented but seagrass abundance (shoot density and biomass) did not reach the level of unmanipulated controls. Seagrass species did not appear to form seed banks despite some species being capable of producing long-lived seeds. The species composition of cleared plots remained different to the undisturbed controls throughout the 26-month experiment. Syringodium isoetifolium was a rapid asexual coloniser of disturbed plots and remained at higher abundances than in the control treatments for the duration of the study. S. isoetifolium had the fastest horizontal rhizome growth of species asexually colonising cleared plots (6.9 mm day−1). Halophila ovalis was the most successful sexual coloniser but was displaced by asexually colonising species. H. ovalis was the only species observed to produce fruits during the study. Small disturbances in the meadow led to long-term (>2 years) changes in community composition. This study demonstrated that succession in tropical seagrass communities was not a deterministic process. Variations in recovery observed for different tropical seagrass communities highlighted the importance of understanding life history characteristics of species within individual communities to effectively predict their response to disturbance. A reproductive strategy involving clonal growth and production of long-lived, locally dispersed seeds is suggested which may provide an evolutionary advantage to plants growing in tropical environments subject to temporally unpredictable major disturbances such as cyclones

Improved models for estimating temporal changes in carbon sequestration in above-ground biomass of mixed-species environmental plantings

Plantings of mixed native species (termed 'environmental plantings') are increasingly being established for carbon sequestration whilst providing additional environmental benefits such as biodiversity and water quality. In Australia, they are currently one of the most common forms of reforestation. Investment in establishing and maintaining such plantings relies on having a cost-effective modelling approach to providing unbiased estimates of biomass production and carbon sequestration rates. In Australia, the Full Carbon Accounting Model (FullCAM) is used for both national greenhouse gas accounting and project-scale sequestration activities. Prior to undertaking the work presented here, the FullCAM tree growth curve was not calibrated specifically for environmental plantings and generally under-estimated their biomass. Here we collected and analysed above-ground biomass data from 605 mixed-species environmental plantings, and tested the effects of several planting characteristics on growth rates. Plantings were then categorised based on significant differences in growth rates. Growth of plantings differed between temperate and tropical regions. Tropical plantings were relatively uniform in terms of planting methods and their growth was largely related to stand age, consistent with the un-calibrated growth curve. However, in temperate regions where plantings were more variable, key factors influencing growth were planting width, stand density and species-mix (proportion of individuals that were trees). These categories provided the basis for FullCAM calibration. Although the overall model efficiency was only 39-46%, there was nonetheless no significant bias when the model was applied to the various planting categories. Thus, modelled estimates of biomass accumulation will be reliable on average, but estimates at any particular location will be uncertain, with either under- or over-prediction possible. When compared with the un-calibrated yield curves, predictions using the new calibrations show that early growth is likely to be more rapid and total above-ground biomass may be higher for many plantings at maturity. This study has considerably improved understanding of the patterns of growth in different types of environmental plantings, and in modelling biomass accumulation in young (<25. years old) plantings. However, significant challenges remain to understand longer-term stand dynamics, particularly with temporal changes in stand density and species composition. © 2014.

Pasture production and composition response after killing Eucalypt trees with herbicides in central Queensland

Clearing woodlands is practised world-wide to increase crop and livestock production, but can result in unintended consequences including woody regrowth and land degradation. The pasture response of 2 eucalypt woodlands in the central Queensland rangelands to killing trees with herbicides, in the presence or absence of grazing and regular spring burning, was recorded over 7 or 8 years to determine the long-term sustainability of these common practices. Herbage mass and species composition plus tree dynamics were monitored in 2 replicated experiments at each site. For 8 years following herbicide application, killing Eucalyptus populnea F. Muell. (poplar box) trees resulted in a doubling of native pasture herbage mass from that of the pre-existing woodland, with a tree basal area of 8.7 m2 ha-1. Conversely, over 7 years with a similar range of seasons, killing E. melanophloia F. Muell. (silver-leaved ironbark) trees of a similar tree basal area had little impact on herbage mass grown or on pasture composition for the first 4 years before production then increased. Few consistent changes in pasture composition were recorded after killing the trees, although there was an increase in the desirable grasses Dichanthium sericeum (R. Br.) A. Camus (Queensland bluegrass) and Themeda triandra Forssk. (kangaroo grass) when grazed conservatively. Excluding grazing allowed more palatable species of the major grasses to enhance their prominence, but seasonal conditions still had a major influence on their production in particular years. Pasture crown basal area was significantly higher where trees had been killed, especially in the poplar box woodland. Removing tree competition did not have a major effect on pasture composition that was independent of other management impositions or seasons, and it did not result in a rapid increase in herbage mass in both eucalypt communities. The slow pasture response to tree removal at one site indicates that regional models and economic projections relating to tree clearing require community-specific inputs.

Productivity, carbon assimilation and intra-annual change in tropical reef platform seagrass communities of the Torres Strait, north-eastern Australia

Detailed data on seagrass distribution, abundance, growth rates and community structure information were collected at Orman Reefs in March 2004 to estimate the above-ground productivity and carbon assimilated by seagrass meadows. Seagrass meadows were re-examined in November 2004 for comparison at the seasonal extremes of seagrass abundance. Ten seagrass species were identified in the meadows on Orman Reefs. Extensive seagrass coverage was found in March (18,700 ha) and November (21,600 ha), with seagrass covering the majority of the intertidal reef-top areas and a large proportion of the subtidal areas examined. There were marked differences in seagrass above-ground biomass, distribution and species composition between the two surveys. Major changes between March and November included a substantial decline in biomass for intertidal meadows and an expansion in area of subtidal meadows. Changes were most likely a result of greater tidal exposure of intertidal meadows prior to November leading to desiccation and temperature-related stress. The Orman Reef seagrass meadows had a total above-ground productivity of 259.8 t DW day-1 and estimated carbon assimilation of 89.4 t C day-1 in March. The majority of this production came from the intertidal meadows which accounted for 81% of the total production. Intra-annual changes in seagrass species composition, shoot density and size of meadows measured in this study were likely to have a strong influence on the total above-ground production during the year. The net estimated above-ground productivity of Orman Reefs meadows in March 2004 (1.19 g C m-2 day-1) was high compared with other tropical seagrass areas that have been studied and also higher than many other marine, estuarine and terrestrial plant communities.

Perennial grassland dynamics on fertile plains: Is coexistence mediated by disturbance?

The response of grasslands to disturbance varies with the nature of the disturbance and the productivity of the landscape. In highly productive grasslands, competitive exclusion often results in decreased species richness and grazing may allow more species to coexist. Once widespread, grasslands dominated by Dichanthium sericeum (Queensland bluegrass) and Astrebla spp. (Mitchell grass) occur on fertile plains but have been reduced in extent by cultivation. We tested the effects of exclusion of livestock grazing on these grasslands by comparing the floristic composition of sites in a nature reserve with an adjacent stock reserve. In addition, sites that had been cultivated within the nature reserve were compared with those where grazing but no cultivation had occurred. To partition the effects of temporal variation from spatial variation we sampled sites in three different years (1998, 2002 and 2004). Some 194 taxa were recorded at the nature reserve and surrounding stock routes. Sampling time, the occurrence of past cultivation and livestock grazing all influenced species composition. Species richness varied greatly between sampling periods relating to highly variable rainfall and water availability on heavy clay soils. Native species richness was significantly lower at previously cultivated sites (13-22 years after cultivation), but was not significantly influenced by grazing exclusion. After 8 years it appears that reintroducing disturbance in the form of livestock grazing is not necessary to maintain plant species richness in the reserve. The highly variable climate (e.g. droughts) probably plays an important role in the coexistence of species by negating competitive exclusion and allowing interstitial species to persist.

Natural enemies of Paropsis atomaria Olivier (Coleoptera: Chrysomelidae) in south-eastern Queensland eucalypt plantations

A field survey for natural enemies of Paropsis atomaria was conducted at two south-eastern Queensland Eucalyptus cloeziana plantation sites during 2004-2005. Primary egg and larval parasitoids and associated hyperparasitoids were identified to genus or species, and parasitism rates were determined throughout the season. Predators were identified to family level but their impact was not quantified. P. atomaria adults were also examined as potential hosts for parasitic mites and nematodes. An undescribed species of Neopolycystus (Pteromalidae) was the major primary egg parasitoid species reared from egg batches, parasitising half of all egg batches collected. Three hyperparasitoid species (Baeoanusia albifunicle (Encyrtidae), Neblatticida sp. (Encyrtidae) and Aphaneromella sp. (Platygasteridae) were present, representing around one-quarter to one-third of all emergent wasps; this is the first host association record for Neopolycystus-B. albifunicle. In contrast to populations of P. atomaria from the Australian Capital Territory, primary larval parasitism was very low, around 1%, and attributable only to the tachinid flies Anagonia sp. and Paropsivora sp. However, the presence of the sit-and-wait larval hyperparasitoid, Perilampus sp. (Perilampidae) was high, emerging from around 17% of tachinid pupae, with planidia infesting a further 40% of unparasitised hosts. Three species of podapolipid mites parasitised sexually mature P. atomaria adults, while no nematodes were found in this study. Spiders were the most common predators and their abundance was positively correlated with P. atomaria adult and egg numbers. Although natural enemy species composition was identical between our two study sites, significant differences in abundance and frequency were found between sites.

Evidence of a latitudinal gradient in spider diversity in Australian cotton

The most common explanation for species diversity increasing towards the tropics is the corresponding increase in habitats (spatial heterogeneity). Consequently, a monoculture (like cotton in Australia) which is grown along a latitudinal gradient, should have the same degree of species diversity throughout its range. We tested to see if diversity in a dominant cotton community (spiders) changed with latitude, and if the community was structurally identical in different parts of Australia. We sampled seven sites extending over 20 degrees of latitude. At each site we sampled 1-3 fields 3-5 times during the cotton growing season using pitfall traps and beatsheets, recording all the spiders collected to family. We found that spider communities in cotton are diverse, including a large range of foraging guilds, making them suitable for a conservation biological control programme. We also found that spider diversity increased from high to low latitudes, and the communities were different, even though the spiders were in the same monocultural habitat. Spider beatsheet communities around Australia were dominated by different families, and responded differently to seasonal changes, indicating that different pest groups would be targeted at different locations. These results show that diversity can increase from high to low latitudes, even if spatial heterogeneity is held constant, and that other factors external to the cotton crop are influencing spider species composition. Other models which may account for the latitudinal gradient, such as non-equilibrium regional processes, are discussed.

Action learning in partnership with Landcare and catchment management groups to support increased pasture sowings in southern inland Queensland

The incorporation of sown pastures as short-term rotations into the cropping systems of northern Australia has been slow. The inherent chemical fertility and physical stability of the predominant vertisol soils across the region enabled farmers to grow crops for decades without nitrogen fertiliser, and precluded the evolution of a crop–pasture rotation culture. However, as less fertile and less physically stable soils were cropped for extended periods, farmers began to use contemporary farming and sown pasture technologies to rebuild and maintain their soils. This has typically involved sowing long-term grass and grass–legume pastures on the more marginal cropping soils of the region. In partnership with the catchment management authority, the Queensland Murray–Darling Committee (QMDC) and Landcare, a pasture extension process using the LeyGrain™ package was implemented in 2006 within two Grain & Graze projects in the Maranoa-Balonne and Border Rivers catchments in southern inland Queensland. The specific objectives were to increase the area sown to high quality pasture and to gain production and environmental benefits (particularly groundcover) through improving the skills of producers in pasture species selection, their understanding and management of risk during pasture establishment, and in managing pastures and the feed base better. The catalyst for increasing pasture sowings was a QMDC subsidy scheme for increasing groundcover on old cropping land. In recognising a need to enhance pasture knowledge and skills to implement this scheme, the QMDC and Landcare producer groups sought the involvement of, and set specific targets for, the LeyGrain workshop process. This is a highly interactive action learning process that built on the existing knowledge and skills of the producers. Thirty-four workshops were held with more than 200 producers in 26 existing groups and with private agronomists. An evaluation process assessed the impact of the workshops on the learning and skill development by participants, their commitment to practice change, and their future intent to sow pastures. The results across both project catchments were highly correlated. There was strong agreement by producers (>90%) that the workshops had improved knowledge and skills regarding the adaptation of pasture species to soils and climates, enabling a better selection at the paddock level. Additional strong impacts were in changing the attitudes of producers to all aspects of pasture establishment, and the relative species composition of mixtures. Producers made a strong commitment to practice change, particularly in managing pasture as a specialist crop at establishment to minimise risk, and in the better selection and management of improved pasture species (particularly legumes and the use of fertiliser). Producers have made a commitment to increase pasture sowings by 80% in the next 5 years, with fourteen producers in one group alone having committed to sow an additional 4893 ha of pasture in 2007–08 under the QMDC subsidy scheme. The success of the project was attributed to the partnership between QMDC and Landcare groups who set individual workshop targets with LeyGrain presenters, the interactive engagement processes within the workshops themselves, and the follow-up provided by the LeyGrain team for on-farm activities.

What drives plant biodiversity in the clay floodplain grasslands of NSW?

An assessment of the relative influences of management and environment on the composition of floodplain grasslands of north-western New South Wales was made using a regional vegetation survey sampling a range of land tenures (e. g. private property, travelling stock routes and nature reserves). A total of 364 taxa belonging to 55 different plant families was recorded. Partitioning of variance with redundancy analysis determined that environmental variables accounted for a greater proportion (61.3%) of the explained variance in species composition than disturbance-related variables (37.6%). Soil type (and fertility), sampling time and rainfall had a strong influence on species composition and there were also east-west variations in composition across the region. Of the disturbance-related variables, cultivation, stocking rate and flooding frequency were all influential. Total, native, forb, shrub and subshrub richness were positively correlated with increasing time since cultivation. Flood frequency was positively correlated with graminoid species richness and was negatively correlated with total and forb species richness. Site species richness was also influenced by environmental variables (e. g. soil type and rainfall). Despite the resilience of these grasslands, some forms of severe disturbance (e. g. several years of cultivation) can result in removal of some dominant perennial grasses (e. g. Astrebla spp.) and an increase in disturbance specialists. A simple heuristic transitional model is proposed that has conceptual thresholds for plant biodiversity status. This knowledge representation may be used to assist in the management of these grasslands by defining four broad levels of community richness and the drivers that change this status.

Climate change effects on pasture systems in south-eastern Australia.

Climate change projections for Australia predict increasing temperatures, changes to rainfall patterns, and elevated atmospheric carbon dioxide (CO2) concentrations. The aims of this study were to predict plant production responses to elevated CO2 concentrations using the SGS Pasture Model and DairyMod, and then to quantify the effects of climate change scenarios for 2030 and 2070 on predicted pasture growth, species composition, and soil moisture conditions of 5 existing pasture systems in climates ranging from cool temperate to subtropical, relative to a historical baseline. Three future climate scenarios were created for each site by adjusting historical climate data according to temperature and rainfall change projections for 2030, 2070 mid-and 2070 high-emission scenarios, using output from the CSIRO Mark 3 global climate model. In the absence of other climate changes, mean annual pasture production at an elevated CO2 concentration of 550 ppm was predicted to be 24-29% higher than at 380 ppm CO2 in temperate (C-3) species-dominant pastures in southern Australia, with lower mean responses in a mixed C-3/C-4 pasture at Barraba in northern New South Wales (17%) and in a C-4 pasture at Mutdapilly in south-eastern Queensland (9%). In the future climate scenarios at the Barraba and Mutdapilly sites in subtropical and subhumid climates, respectively, where climate projections indicated warming of up to 4.4 degrees C, with little change in annual rainfall, modelling predicted increased pasture production and a shift towards C-4 species dominance. In Mediterranean, temperate, and cool temperate climates, climate change projections indicated warming of up to 3.3 degrees C, with annual rainfall reduced by up to 28%. Under future climate scenarios at Wagga Wagga, NSW, and Ellinbank, Victoria, our study predicted increased winter and early spring pasture growth rates, but this was counteracted by a predicted shorter spring growing season, with annual pasture production higher than the baseline under the 2030 climate scenario, but reduced by up to 19% under the 2070 high scenario. In a cool temperate environment at Elliott, Tasmania, annual production was higher than the baseline in all 3 future climate scenarios, but highest in the 2070 mid scenario. At the Wagga Wagga, Ellinbank, and Elliott sites the effect of rainfall declines on pasture production was moderated by a predicted reduction in drainage below the root zone and, at Ellinbank, the use of deeper rooted plant systems was shown to be an effective adaptation to mitigate some of the effect of lower rainfall.

Resilience of a high-conservation-value, semi-arid grassland on fertile clay soils to burning, mowing and ploughing.

In grassland reserves, managed disturbance is often necessary to maintain plant species diversity. We carried out experiments to determine the impact of fire, kangaroo grazing, mowing and disc ploughing on grassland species richness and composition in a nature reserve in semi-arid eastern Australia. Vegetation response was influenced by winter-spring drought after establishment of the experiments, but moderate rainfall followed in late summer-autumn. Species composition varied greatly between sampling times, and the variability due to rainfall differences between seasons and years was greater than the effects of fire, kangaroo grazing, mowing or disc ploughing. In the fire experiment, species richness and composition recovered more rapidly after spring than autumn burning. Species richness and composition were similar to control sites within 12 months of burning and mowing, suggesting that removal of the dominant grass canopy is unnecessary to enhance plant diversity. Two fires (separated by 3 years) and post-fire kangaroo grazing had only minor influence on species richness and composition. Even disc ploughing caused only a small reduction in native richness. The minor impact of ploughing was explained by the small areas that were ploughed, the once-off nature of the treatment, and the high degree of natural movement and cracking in these shrink-swell soils. Recovery of the composition and richness of these grasslands was rapid because of the high proportion of perennial species that resprout vegetatively after fire and mowing. There appears to be little conservation benefit from fire, mowing or ploughing ungrazed areas, as we could identify no native plant species dependent on frequent disturbance for persistence in this grassland community. However, the ability of the Astrebla- and Dichanthium-dominated grasslands to recover quickly after disturbance, given favourable seasonal conditions, suggests that they are well adapted to natural disturbances (e.g. droughts, fire, flooding and native grazing).

Invasion impacts on biodiversity: responses of ant communities to infestation by cat’s claw creeper, Macfadyena unguis-cati (Bignoniaceae) in subtropical Australia

Ants are the dominant soil faunal group in many if not most terrestrial ecosystems, and play a key role in soil structure and function. This study documents the impacts of invasion by the exotic cat’s claw creeper vine, Macfadyena unguis-cati (L.) Gentry (Bignoniaceae) on surface-situated (epigaeic) and subterranean (hypogaeic) ant communities in subtropical SE Queensland Australia where it is a major environmental weed of riparian areas, rainforest communities and remnant natural vegetation, smothering standing vegetation and causing canopy collapse. Soil ants were sampled in infested and uninfested areas at eight sites spanning both riparian and non-riparian habitats in subtropical SE Queensland. Patterns of ant species composition and functional grouping in response to patch invasion status, landscape type and habitat stratum were investigated using ANOVA and non-metric multidimensional scaling ordination. The epigaeic and subterranean strata supported markedly different ant assemblages, and ant communities also differed between riparian and non-riparian habitats. However, M. unguis-cati invasion had a surprisingly limited impact. There was a tendency for ant abundance and species richness to be lower in infested patches, and overall species composition was different between infested and uninfested patches, but these differences were relatively small, and did not occur consistently across sites. There were changes in functional group composition that conformed to known functional group responses to environmental change, but these were similarly limited and inconsistent across sites. Our study has shown that ant communities are surprisingly resilient to invasion by M. unguis-cati, and serves as a warning against making assumptions about invasion impacts based on visual appearances.

Management of the invasive hill raspberry (Rubus niveus) on Santiago Island, Galapagos: Eradication or indefinite control?

The eradication of an invasive plant species can provide substantial ecological and economic benefits by eliminating completely the negative effects of the weed and reducing the high cost of continuing control. A 5-yr program toward the eradication of hill raspberry (Rubus niveus Thunb.) in Santiago Island is evaluated using delimitation and extirpation criteria, as well as assessment of the ecological community response to management techniques. Currently, hill raspberry is located in the humid zone of Santiago island. It is distributed over three main infestations, small patches, and many scattered individuals within an area of approximately 1,000 ha. New infestations are constantly being found; every year, new detections add an area of approximately 175 ha. Adult and juvenile individuals are still found, both beyond and within known infestations. Both plant and seed bank density of hill raspberry decreased over time where infestations were controlled. Species composition in the seed bank and existing vegetation were significantly different between areas under intensive control and adjacent uninvaded forest. After 5 yr of intensive management, delimitation of hill raspberry has not been achieved; new populations are found every year, increasing the infested area that requires management. Off-target effects on native species resulting from control efforts seem to be substantial. Although a vast increase in economic investment would allow intensive searching that might enable all individuals to be found and controlled, the resultant disturbance and off-targets effects could outweigh the conservation benefits of eradication.

Diversity and abundance of fungivorous thrips (Thysanoptera) associated with leaf-litter and bark across forest types and two tree genera in subtropical Australia

Leaf-litter thrips were much more common and diverse in dry sclerophyll forest than in wetter forest types in subtropical southeast Queensland, Australia. In dry sclerophyll forest, the species composition of thrips in leaf-litter was strongly differentiated from the thrips fauna associated with bark of the trees Eucalyptus major and Acacia melanoxylon (4 of 34 species in common). The species composition of bark-dwelling thrips was similar across the two tree species and also across two eucalypts with different bark types, Eucalyptus major (flaky) and Eucalyptus siderophloia (rough). The diversity of thrips from the leaf-litter was not differentiated across all of these tree species. Virtually all thrips collected were Phlaeothripidae, subfamilies Idolothripinae and Phlaeothripinae. Idolothripinae were associated almost exclusively with leaf-litter, but Phlaeothripinae were in leaf-litter and bark. The association of fungal-feeding thrips with dry sclerophyll forest raises questions about their ecological requirements and the role they play in nutrient cycling. © 2012 Copyright Taylor and Francis Group, LLC.

A comparison of stocking methods for beef production in northern Australia: pasture and soil surface condition responses

Historical stocking methods of continuous, season-long grazing of pastures with little account of growing conditions have caused some degradation within grazed landscapes in northern Australia. Alternative stocking methods have been implemented to address this degradation and raise the productivity and profitability of the principal livestock, cattle. Because information comparing stocking methods is limited, an evaluation was undertaken to quantify the effects of stocking methods on pastures, soils and grazing capacity. The approach was to monitor existing stocking methods on nine commercial beef properties in north and south Queensland. Environments included native and exotic pastures and eucalypt (lighter soil) and brigalow (heavier soil) land types. Breeding and growing cattle were grazed under each method. The owners/managers, formally trained in pasture and grazing management, made all management decisions affecting the study sites. Three stocking methods were compared: continuous (with rest), extensive rotation and intensive rotation (commonly referred to as 'cell grazing'). There were two or three stocking methods examined on each property: in total 21 methods (seven continuous, six extensive rotations and eight intensive rotations) were monitored over 74 paddocks, between 2006 and 2009. Pasture and soil surface measurements were made in the autumns of 2006, 2007 and 2009, while the paddock grazing was analysed from property records for the period from 2006 to 2009. The first 2 years had drought conditions (rainfall average 3.4 decile) but were followed by 2 years of above-average rainfall. There were no consistent differences between stocking methods across all sites over the 4 years for herbage mass, plant species composition, total and litter cover, or landscape function analysis (LFA) indices. There were large responses to rainfall in the last 2 years with mean herbage mass in the autumn increasing from 1970 kg DM ha(-1) in 2006-07 to 3830 kg DM ha(-1) in 2009. Over the same period, ground and litter cover and LFA indices increased. Across all sites and 4 years, mean grazing capacity was similar for the three stocking methods. There were, however, significant differences in grazing capacity between stocking methods at four sites but these differences were not consistent between stocking methods or sites. Both the continuous and intensive rotation methods supported the highest average annual grazing capacity at different sites. The results suggest that cattle producers can obtain similar ecological responses and carry similar numbers of livestock under any of the three stocking methods.