Plant species richness responses to grazing protection and degradation history in a low productivity landscape

Questions: Does species richness and abundance accumulate with grazing protection in low productivity ecosystems with a short evolutionary history of grazing, as predicted by emerging theory? How do responses to grazing protection inform degradation history? Location: Mulga (Acacia aneura) dry forest, eastern Australia, generally considered chronically degraded by livestock grazing. Methods: Three paired exclosures (ungrazed, and macropod‐grazed) were compared with open‐grazed areas after 25 years using quadrats located on either side of the fences. Additionally, the regional flora for mulga dry forest was assessed to identify species that may have declined and could be threatened by grazing. Results: Low herbaceous biomass accumulation (<1.3 t ha^?1) with full grazing protection confirmed a low productivity environment. For most plant life forms the highest species richness was in macropod‐grazed exclosures, an intermediate grazing disturbance that best approximates the evolutionary history of the environment. This was the net outcome of species that both declined and increased in response to grazing. Regeneration and subsequent self‐thinning of mulga was promoted with grazing protection, but did not confound interpretation of species richness and abundance responses. At the regional scale only 11 native species out of 407 comprising the mulga dry forest flora were identified as rare and potentially threatened by grazing. Conclusions: Significant increases in richness or abundance of native plants with grazing protection, persistence of perennial grasses, regeneration of mulga and scant evidence of a major decline in the regional flora are not consistent with established assertions that long‐grazed mulga dry forest has crossed functional thresholds that limit recovery. Further, a peak in species richness under intermediate (macropod) grazing is counter to the shape of the response predicted by emerging theory for recovery of species richness in a low productivity environment. The finding prompts a more thorough understanding of the distinction between environments with inherently low productivity and those degraded by grazing.     

Land use and drought interactively affect interspecific competition and species diversity at the local scale in a semiarid steppe ecosystem

Few studies have considered interactive effects of grazing and drought on species composition and the relative contribution of species to total biomass, although it is important to understand the short-term dynamics and community succession in grazed ecosystems. We monitored species diversity and relative biomass contribution at one site protected from grazing since 1979 (UG79), and at winter grazing (WG) and heavily grazed (HG) sites. Continuous heavy grazing resulted in lower plant height and more but small individuals (tillers or stolons). Drought significantly reduced total plant density on all sites. Grazing affected species diversity more than drought. Species richness at site UG79 was significantly higher than at sites WG and HG, while drought only tended to reduce species diversity. Drought stress and grazing disturbance interactively controlled species competition and functional groups. Both perennial grasses and forbs had greater contribution to total biomass at site UG79, and perennial grasses contributed more than 97% of total biomass at site WG. The contribution to total biomass of annual forbs and semi-shrubs significantly increased at site HG after two dry years. The significant decrease in Potentilla acaulis and a substantial increase in annual species at this site indicate that the perennial vegetation of this ecosystem is in great danger of extinction under conditions of prolonged drought.     

Consequences of anthropogenic disturbances on soil seed bank diversity and nurse shrub effect in a semiarid rangeland

What is the effect of common land use histories on the diversity, richness, spatial distribution and abundance of the soil seed bank (SSB)? Does the effect change between different microsites under shrub canopy? To address these questions we selected seven sites with different grazing and fire histories in the Patagonian Monte desert. We took soil samples in seven microsites at different distances of the trunk along a windward/leeward transect through shrub patches to estimate the SSB of perennial grasses, shrubs and annuals. Shrubs SSB was scarce. The nurse shrub effect on perennial grass SSB was evident at sites with a low disturbance intensity (as sites without continuous grazing), and higher on the leeward side under the shrub canopy. High disturbance intensity (such as permanent grazing) promoted a decrease in perennial grass SSB and an increase in annuals SBB (especially non-native). Land use histories related to fire showed a moderated disturbance response (medium values for both groups). Differences between land use histories varied depending on the windward/leeward microsite from which the SSB was analysed. Our results suggest a nurse effect of the shrub patches on the spatial distribution and abundance of the SSB, but this effect decrease under continuous grazing. Fire seems to have a positive effect on perennial grass SSB. Nevertheless, fire characteristics must be controlled. Moreover, perennial grass SSB was almost depleted under continuous grazing, driving the grazed system towards a vulnerable state; and annual non-native species take advantage of this disturbance and dominate the SSB.     

Effects of grazing on seedling establishment: the role of seed and safe-site availability

Abstract. The first objective of this paper was to assess the effects of grazing on seedling establishment of two species whose relative abundance at the adult stage is affected by grazing in a contrasting fashion. Second, we evaluated the relative importance of seed versus safe-site availability in explaining the effect of grazing on seedling establishment. We monitored seedling establishment on a grazed area, on two areas which had not been grazed for two and seven years, and on plots which had been experimentally defoliated. The species compared were Dan-thonia montevidensis, a native perennial grass which dominates both grazed and ungrazed communities, and Leontodón taraxacoides, an invading exotic rosette species from the Compositae family. Continuous grazing enhanced seedling establishment of both species through its effect on the availability of safe sites. Seed availability accounted for only one, but very important, grazing effect: the lack of response by L. taraxacoides to the defoliation in the seven-year old exclosure. Its seed supply was depleted by exclusion of grazing and, consequently, its short-term regeneration capacity after disturbance was lost.     

After grazing exclusion,is there any modification of strategy for two guerrilla species: <Emphasis Type="Italic">Elymus repens</Emphasis> (L.) Gould and <Emphasis Type="Italic">Agrostis stolonifera</Emphasis> (L.)?

Elymus repens (L.) Gould and Agrostis stolonifera (L.), are competitive grasses with guerrilla strategy that invade grasslands with a low stocking rate. In this work, we tested the hypotheses that grazing exclusion facilitates vegetative development of rhizomes and stolons of these clonal grasses and that such change is a key mechanism for their abundance in set-aside grasslands. The competitive capacities of these two guerrilla species were characterised by samples in plant community (species richness and biomass) and on the level of individual species (morphometric measurements on stolons and rhizomes) during a growing season. Compared to grasslands where grazing was excluded for three years, species richness was higher in grazed site and the plant community structure differed. Indeed, with grazing exclusion, a shift from annual species with a diversified growth-form to perennial species with a tall tussock and graminoid growth-form was monitored. In ungrazed situation, Elymus repens and Agrostis stolonifera were the dominant grasses, and the standing biomass for the lowland community showed a significant increase compared to the grazed site. Vegetative development increased competitive capacities of these two guerrilla species and led by phenomenon of competitive exclusion to the disappearance of annuals species. With grazing cessation, Elymus repens was found to increase the size of aerial traits (shoot length and the number of leaves per shoot) and this may both be propitious for achieving dominance within plant communities and also maintaining its competitive local advantage. By contrast, Agrostis stolonifera showed an increase in a root trait, i.e. rhizome length, in the fenced site, which provide good ability for spatial propagation and then to explore adjacent patches. We concluded that Elymus repens presented a morphological capacity to change its colonising strategy from a guerrilla strategy to a phalanx strategy, by morphological variability of aerial organs, when it was submitted to competitive stress and environment modifications. Agrostis stolonifera showed a capacity to escape aerial competition resulting from grazing cessation, than to increase underground propagation capacity. The present study highlighted the capacities of Elymus repens to respond in an adaptative way to competitive pressure.     

Impact of Red Deer Cervus elaphus Grazing on Bilberry Vaccinium myrtillus and Composition of Ground Beetle (Coleoptera, Carabidae) Assemblage

We studied the role of red deer Cervus elaphus L. as ecosystem modifier in boreal forest (Tingvoll municipality, 62°52′ N, 8°20′ E, Norway), during early summer of 2001. The effect of grazing by red deer on ground beetles (Carabidae) abundance and diversity was investigated across a gradient of grazing pressures. We trapped ground beetles by pit-fall traps from three homogeneous winter grazing areas (ungrazed, medium grazed, heavily grazed). Bilberry Vaccinium myrtillus (the main winter food for red deer) was sampled and its dry weight was measured for the three locations. Gradient analyses showed that grazing by red deer affects carabid species composition. Grazing significantly affected the amount of bilberry, which correlated with species variation. According to our predictions, we found a higher abundance of carabids in the heavily grazed location, but the species richness and the diversity indices were similar for the three areas. This study shows that overall species composition is altered along a gradient as consequence of red deer winter grazing and that red deer act as ecosystem engineer, by reducing the bilberry heather which dominates the field layer in early summer.     

Does grazing reduce survival of indigenous perennial grasses of the semi-arid woodlands of western New South Wales?

Abstract Exclosures were used to examine the impact of grazing upon the mortality patterns of populations of six indigenous grass species. The experiment compared unfenced areas with areas from which either sheep only or sheep, rabbits and kangaroos were excluded. There were large interspecific differences in mortality patterns, with Stipa nitida and Aristida browniana having relatively high mortality rates and Eragrostis eriopoda having relatively low mortality rates. Grazing-induced mortality was observed in treatment areas that were grazed by sheep, rabbits and kangaroos and in areas grazed only by rabbits and kangaroos. The short-lived S. nitida appears less likely to suffer grazing-induced mortality than species of intermediate longevity such as Monachather paradoxa. These observations help explain the decline in endemic perennial grasses that has taken place in the vegetation of western New South Wales since European settlement. Management of these rangelands to encourage these grasses must consider total grazing pressure and not simply the impact of livestock.     

Plastic traits of an exotic grass contribute to its abundance but are not always favourable

In herbaceous ecosystems worldwide, biodiversity has been negatively impacted by changed grazing regimes and nutrient enrichment. Altered disturbance regimes are thought to favour invasive species that have a high phenotypic plasticity, although most studies measure plasticity under controlled conditions in the greenhouse and then assume plasticity is an advantage in the field. Here, we compare trait plasticity between three co-occurring, C(4) perennial grass species, an invader Eragrostis curvula, and natives Eragrostis sororia and Aristida personata to grazing and fertilizer in a three-year field trial. We measured abundances and several leaf traits known to correlate with strategies used by plants to fix carbon and acquire resources, i.e. specific leaf area (SLA), leaf dry matter content (LDMC), leaf nutrient concentrations (N, C:N, P), assimilation rates (Amax) and photosynthetic nitrogen use efficiency (PNUE). In the control treatment (grazed only), trait values for SLA, leaf C:N ratios, Amax and PNUE differed significantly between the three grass species. When trait values were compared across treatments, E. curvula showed higher trait plasticity than the native grasses, and this correlated with an increase in abundance across all but the grazed/fertilized treatment. The native grasses showed little trait plasticity in response to the treatments. Aristida personata decreased significantly in the treatments where E. curvula increased, and E. sororia abundance increased possibly due to increased rainfall and not in response to treatments or invader abundance. Overall, we found that plasticity did not favour an increase in abundance of E. curvula under the grazed/fertilized treatment likely because leaf nutrient contents increased and subsequently its' palatability to consumers. E. curvula also displayed a higher resource use efficiency than the native grasses. These findings suggest resource conditions and disturbance regimes can be manipulated to disadvantage the success of even plastic exotic species.     

Does grazing mediate soil carbon and nitrogen accumulation beneath C4, perennial grasses along an environmental gradient?

An experiment was conducted to evaluate the influence of long-term (>25 yrs) grazing on soil organic carbon (SOC) and total soil nitrogen (N) accumulation beneath individual plants of three perennial grasses along an environmental gradient in the North American Great Plains. The zone of maximum SOC and N accumulation was restricted vertically to the upper soil depth (0-5 cm) and horizontally within the basal area occupied by individual caespitose grasses, which contributed to fine-scale patterning of soil heterogeneity. Long-term grazing mediated SOC and N accumulation in the tall-, mid- and shortgrass communities, but the responses were community specific. SOC and N were lower beneath Schizachyrium scoparium plants in long-term grazed sites of the tall- and midgrass communities, but higher beneath Bouteloua gracilis plants in the long-term grazed site of the shortgrass community. SOC, but not N, was greater in soils beneath compared to between S. scoparium plants in an abandoned field seeded in 1941, indicating that this caespitose grass accumulated SOC more rapidly than N. SOC and N were greater in the 0-5 cm soil depth beneath a caespitose grass (S. scoparium) compared to a rhizomatous grass (Panicum virgatum) in the tallgrass community, with no significant accumulation of either SOC or N beneath P. virgatum plants. Grazing appears to indirectly mediate nutrient accumulation beneath caespitose grasses along the environmental gradient by modifying the size class distribution of plants. Populations with a greater proportion of large plants have a greater potential for biomass incorporation into soils and may more effectively capture redistributed organic matter from between plant locations. Contrasting plant responses to grazing at various locations along the environmental gradient conform to the predictions of the generalized grazing model, as the selection pressures of grazing and aridity may have also influenced the ability of caespitose grasses to accumulate nutrients in soils beneath them by mediating grazing resistance, competitive ability and population structure.     

Patch dynamics in grazed subtropical native pastures in south‐east Queensland

Abstract Patch formation is common in grazed grasslands but the mechanisms involved in the formation and maintenance of patches are not clear. To increase our knowledge on this subject we examined possible reasons for patch formation and the influence of management on changes between patch states in three experiments in native pasture communities in the Crows Nest district, south‐east Queensland. In these communities, small‐scale patches (tall grassland (dominated by large and medium tussock grasses), short swards (dominated by short tussock grasses and sedges), and lawns (dominated by stoloniferous and/or rhizomatous grasses)) are readily apparent. We hypothesized that the formation of short sward and lawn patches in areas of tall grassland was due to combinations of grazing and soil fertility effects. This was tested in Experiment 1 by applying a factorial combination of defoliation, nutrient application and transplants of short tussock and stoloniferous species to a uniform area of tall grassland. Total species density declined during the experiment, was lower with high nutrient applications, but was not affected by defoliation. There were significant changes in abundance of species that provided support for our hypotheses. With light defoliation and low nutrients, the tall grassland remained dominated by large tussock grasses and contained considerable amounts of forbs. With heavy defoliation, the pastures were dominated by medium tussock grasses and there were significant decreases in forbs and increases in sedges (mainly with low nutrients) and stoloniferous grasses (mainly with high nutrients). Total germinable seed densities and those of most species groups were significantly lower in the heavy defoliation than the light defoliation plots. Total soil seed numbers were not affected by nutrient application but there were fewer seeds of the erect forbs and more sedge seeds in plots with high nutrients. The use of resting from grazing and fire to manage transitions between patches was tested. In Experiment 2 , changes in species density and abundance were measured for 5 years in the three patch types with and without grazing. Experiment 3 examined the effects of fire, grazing and resting on short sward patches over 4 years. In Experiment 2 , total species density was lower in lawn than short sward or tall grassland patches, and there were more species of erect forbs than other plant groups in all patch types. The lawn patches were originally dominated by Cynodon spp. This dominance continued with grazing but in ungrazed patches the abundance of Cynodon spp. declined and that of forbs increased. In the short sward patches, dominance of short tussock grasses continued with grazing but in ungrazed plots their abundance declined while that of large tussock grasses increased. The tall grassland patches remained dominated by large and medium tussock species. In Experiment 3 , fire had no effect on species abundance. On the grazed plots the short tussock grasses remained dominant but where the plots were rested from grazing the small tussock grasses declined and the large tussock grasses increased in abundance. The slow and relatively small changes in these experiments over 4 or 5 years showed how stable the composition of these pastures is, and that rapid changes between patch types are unlikely.     

Disturbance: a double-edged sword for restoration in a changing climate

Ecological restoration often relies on disturbance as a tool for establishing target plant communities, but disturbance can be a double-edged sword, at times initiating invasion and unintended outcomes. Here we test how fire disturbance, designed to enhance restoration seeding success, combines with climate and initial vegetation conditions to shift perennial versus annual grass dominance and overall community diversity in Pacific Northwest grasslands. We seeded both native and introduced perennial grasses and native forbs in paired, replicated burned-unburned plots in three sites along a latitudinal climate gradient from southern Oregon to central-western Washington. Past restoration and climate manipulations at each site had increased the variation of starting conditions between plots. Burning promoted the expansion of extant forbs and perennial grasses across all sites. Burning also enhanced the seeding success of native perennial grass and native forbs at the northern and central site, and the success of introduced perennial grasses across all three sites. Annual grass dominance was driven more by latitude than burning, with annuals maintaining their dominance in the south and perennials in the north. At the same time, unrestored grasslands surrounding all sites remained dominated by perennial grasses, suggesting that initial plot clearing may have allowed for annual grass invasion in the southern site. When paired with disturbance, further warming may increase the risk of annual grass dominance, a potentially persistent state.     

Grazing history effects on above- and below-ground litter decomposition and nutrient cycling in two co-occurring grasses

Large herbivores may alter carbon and nutrient cycling in soil by changing above- and below-ground litter decomposition dynamics. Grazing effects may reflect changes in plant allocation patterns, and thus litter quality, or the site conditions for decomposition, but the relative roles of these broad mechanisms have rarely been tested. We examined plant and soil mediated effects of grazing history on litter mass loss and nutrient release in two grazing-tolerant grasses, Lolium multiflorum and Paspalum dilatatum, in a humid pampa grassland, Argentina. Shoot and root litters produced in a common garden by conspecific plants collected from grazed and ungrazed sites were incubated under both grazing conditions. We found that grazing history effects on litter decomposition were stronger for shoot than for root material. Root mass loss was neither affected by litter origin nor incubation site, although roots from the grazed origin immobilised more nutrients. Plants from the grazed site produced shoots with higher cell soluble contents and lower lignin:N ratios. Grazing effects mediated by shoot litter origin depended on the species, and were less apparent than incubation site effects. Lolium shoots from the grazed site decomposed and released nutrients faster, whereas Paspalum shoots from the grazed site retained more nutrient than their respective counterparts from the ungrazed site. Such divergent, species-specific dynamics did not translate into consistent differences in soil mineral N beneath decomposing litters. Indeed, shoot mass loss and nutrient release were generally faster in the grazed grassland, where soil N availability was higher. Our results show that grazing influenced nutrient cycling by modifying litter breakdown within species as well as the soil environment for decomposition. They also indicate that grazing effects on decomposition are likely to involve aerial litter pools rather than the more recalcitrant root compartment.     

Influence of Grazing on Soil Seed Banks Determines the Restoration Potential of Aboveground Vegetation in a Semi‐arid Savanna of Ethiopia

Species composition, number of emerging seedlings, species diversity and functional group of the soil seed banks, and the influence of grazing on the similarity between the soil seed banks and aboveground vegetation, were studied in 2008 and 2009 in a semi‐arid savanna of Ethiopia. We tested whether the availability of persistent seeds in the soil could drive the transition from a degraded system under heavy grazing to healthy vegetation with ample perennial grasses. A total of 77 species emerged from the soil seed bank samples: 21 annual grasses, 12 perennial grasses, 4 herbaceous legumes, 39 forbs, and 1 woody species. Perennial grass species dominated the lightly grazed sites, whereas the heavily grazed sites were dominated by annual forbs. Heavy grazing reduced the number of seeds that can germinate in the seed bank. Species richness in the seed bank was, however, not affected by grazing. With increasing soil depth, the seed density and its species richness declined. There was a higher similarity in species composition between the soil seed bank and aboveground vegetation at the lightly grazed sites compared with the heavily grazed sites. The mean similarity between the seed banks and aboveground vegetation was relatively low, indicating the effect of heavy grazing. Moreover, seeds of perennial grasses were less abundant in the soil seed banks under heavy grazing. We concluded that restoration of grass and woody species from the soil seed banks in the heavily grazed areas could not be successful in semi‐arid savannas of Ethiopia.     

Photosynthetic Pathway Types of Forage Species Along Grazing Gradient from the Songnen Grassland,Northeastern China

Photosynthetic pathway types (C₃ and C₄ species) and their dynamics along grazing gradient were determined for 42 plant species in 30 genera and 13 families from the Songnen grassland, Northeastern China. Of the total, 10 species in 9 genera and 4 families had C₄ photosynthesis; 32 species in 21 genera and 12 families had C₃ photosynthesis. The proportion of C₄ species in total plants and C₄/C₃ increased with grazing intensity, and peaked in overgrazed plot. Most of the increased C₄ species (6 of 10) along the grazed gradient were annual grasses and halophytes. This indicated that the C₄ species had greater capacity to tolerate environmental stresses (e.g. drought and saline) caused by animal grazing in the Songnen grassland, Northeastern China.     

Effect of grazing on plant patterns in arid ecosystems of Patagonian Monte

Our objective was to assess the relationship between the spatial patterning of perennial grasses (total, grazed, and non‐grazed) and shrub patches in rangelands under different grazing pressures of the Patagonian Monte. We selected three grazed paddocks with the usual stocking rate for the area, where previous studies showed that a piosphere formation is common. At each paddock, we analysed the grain of heterogeneity at sites located at two distances from the single watering point (near, far), using high‐resolution aerial photographs. At these sites, we also assessed in the field the density, size, cover, and spatial patterning of grazed and non‐grazed perennial grasses and shrub patches. The grain of heterogeneity of shrub patches was coarser in sites near the watering point than in those distant from it, as a consequence of the increase in size of both, bare soil and shrub patches. Field sampling showed that a coarse grain of heterogeneity relative to fine‐grained sites resulted from changes in species composition, increased bare soil areas and reduced perennial grass cover. In coarse‐grained sites, lower perennial grass cover resulted from lower density and/or smaller size of grass bunches than in fine‐grained sites. We did not find significant differences among sites in the proportion of perennial grazed grasses. Since the density and cover of perennial grasses was higher in fine‐ than in coarse‐grained sites, we suggested that fine‐grained sites are more important as feeding stations than coarse‐grained sites. The consequences of this differential use could lead to degradation of fine‐grained sites and to higher homogeneity in spatial plant structure and floristic composition within paddocks with respect to the condition observed at present, increasing the size of the highly degraded zone within the piosphere. At the patch level, we found that at about one third of the sampled transects, both total and non‐grazed perennial grasses were spatially aggregated with shrub patches. However, in most transects grazed perennial grasses were indifferently distributed in relation with shrub patches, showing that grazers display high selectivity of foraging sites at macro level (i.e. high and low grazing pressure sites at the paddock level), but random occupancy of vegetation units (randomness in the distribution of grazed perennial grasses at the patch level). The intensity of the positive association between non‐grazed grasses and shrub patches was higher in fine‐grained than in coarse‐grained sites and may be attributed to higher protection against herbivores associated to denser shrub patches in fine‐ relative to coarse‐grained sites. We concluded that a feedback exists between the spatial distribution of species preferred by grazers and the spatial patterning of use of these species.     

Long‐term vegetation changes in experimentally grazed and ungrazed back‐barrier marshes in the Wadden Sea

Abstract. Vegetation succession in three back‐barrier salt marshes in the Wadden Sea was studied using a data set comprising 25 years of vegetation development recorded at permanent quadrats. The effect of livestock grazing on succession was assessed by comparing quadrats where grazing was experimentally prevented or imposed. We studied changes at the species level as well as at the level of the plant community. Special attention is given to effects on plant species richness and community characteristics that are relevant for lagomorphs (hares and rabbits) and geese. Inundation frequency and grazing were most important in explaining the variation in species abundance data. The three marshes studied overlap in the occurrence of different plant communities and the observed patterns were consistent between them. Clear differences in frequency and abundance of plant species were observed related to grazing. Most plant species had a greater incidence in grazed treatments. Species richness increased with elevation, and was 1.5 to 2 × higher in the grazed salt marsh. Grazing negatively influenced Atriplex portulacoides and Elymus athericus, whereas Puccinellia maritima and Festuca rubra showed a positive response. The communities dominated by Elymus athericus, Artemisia maritima and Atriplex portulacoides were restricted to the ungrazed marsh. Communities dominated by Puccinellia maritima, Juncus gerardi and Festuca rubra predominantly occurred at grazed sites. As small vertebrate herbivores prefer these plants and communities for foraging, livestock grazing thus facilitates for them.     

Abiotic effects predominate under prolonged livestock‐induced disturbance

Despite the widespread recognition that disturbance by livestock affects multiple indices of landscape health, few studies have examined their effects on both biotic and abiotic processes. We examined the effects of livestock disturbance on soil, vascular plants and reptiles across a disturbance gradient in a semi‐arid Australian woodland. Our gradient ranged from long‐ungrazed water remote sites, through intermediately grazed recovering sites, to currently grazed sites close to water. Our aim was to examine the nature of the effects of grazing‐induced disturbance on biotic and abiotic processes along the gradient. We detected small biotic effects, but no abiotic effects, at low levels of disturbance (intermediate sites). We could not detect a consistent biotic effect on plants or reptiles along the gradient, except between the extreme disturbances. In contrast, we recorded substantial reductions in abiotic structure and function at the most disturbed sites. Structural changes included reductions in the cover of shrub hummocks and increases in bare soil, and reductions in cryptogamic soil crusts. Structural changes were associated with declines in function (soil stability and nutrient indices). Our data are consistent with the notion that abiotic effects predominate at high levels of disturbance in rangelands. Given the extent of abiotic modification at currently grazed sites, the cover of abiotic elements such as hummocks and soil surfaces would seem a better indicator of the long‐term effect of grazing‐induced disturbance than biotic components. The extent of disturbance at currently grazed sites across large areas of rangeland suggests that autogenic recovery will be protracted.     

C4 plants in the vegetation of Mongolia: their natural occurrence and geographical distribution in relation to climate

The natural geographical occurrence, carbon assimilation, and structural and biochemical diversity of species with C₄ photosynthesis in the vegetation of Mongolia was studied. The Mongolian flora was screened for C₄ plants by using ¹³C/¹²C isotope fractionation, determining the early products of ¹⁴CO₂ fixation, microscopy of leaf mesophyll cell anatomy, and from reported literature data. Eighty C₄ species were found among eight families: Amaranthaceae, Chenopodiaceae, Euphorbiaceae, Molluginaceae, Poaceae, Polygonaceae, Portulacaceae and Zygophyllaceae. Most of the C4 species were in three families: Chenopodiceae (41 species), Poaceae (25 species) and Polygonaceae, genus Calligonum (6 species). Some new C₄ species in Chenopodiaceae, Poaceae and Polygonaceae were detected. C₄ Chenopodiaceae species make up 45% of the total chenopods and are very important ecologically in saline areas and in cold arid deserts. C₄ grasses make up about 10% of the total Poaceae species and these species naturally concentrate in steppe zones. Naturalized grasses with Kranz anatomy,of genera such as Setaria, Echinochloa, Eragrostis, Panicum and Chloris, were found in almost all the botanical-geographical regions of Mongolia, where they commonly occur in annually disturbed areas and desert oases. We analyzed the relationships between the occurrence of C₄ plants in 16 natural botanical-geographical regions of Mongolia and their major climatic influences. The proportion of C₄ species increases with decreasing geographical latitude and along the north-to-south temperature gradient; however grasses and chenopods differ in their responses to climate. The abundance of Chenopodiaceae species was closely correlated with aridity, but the distribution of the C₄ grasses was more dependent on temperature. Also, we found a unique distribution of different C₄ Chenopodiaceae structural and biochemical subtypes along the aridity gradient. NADP-malic enzyme (NADP-ME) tree-like species with a salsoloid type of Kranz anatomy, such as Haloxylon ammodendron and Iljinia regelii, plus shrubby Salsola and Anabasis species, were the plants most resistant to ecological stress and conditions in highly arid Gobian deserts with less than 100 mm of annual precipitation. Most of the annual C₄ chenopod species were halophytes, succulent, and occurred in saline and arid environments in steppe and desert regions. The relative abundance of C₃ succulent chenopod species also increased along the aridity gradient. Native C₄ grasses were mainly annual and perennial species from the Cynodonteae tribe with NAD-ME and PEP-carboxykinase (PEP-CK) photosynthetic types. They occurred across much of Mongolia, but were most common in steppe zones where they are often dominant in grazing ecosystems. Received: 17 March 1999 / Accepted: 1 November 1999     

Influence of livestock grazing on vegetation in a saline area in Northeast Thailand

The influence of grazing by water buffalo (Bubalas bufalis) and cattle (Bos taurus) was estimated for vegetation inside and outside cages in a saline area at Khon Kaen, Northeast Thailand. The home range of water buffalo and cattle shifted in response to the period of rice cultivation: during the rice-growing season these animals grazed on roadsides and abandoned places such as the study area; after rice harvest they grazed mainly rice stubble on the paddy. The vegetation in the study area was divided into three types: 1) dominated by the annual grassesChloris barbata andIschaemum rugosum; vegetational cover and plant height in the cage increase due to the increase of these grasses; 2) thorny shrub patch ofMaytenus mekongensis; other species in this patch almost died a year after experimental elimination of this shrub; this salt-tolerant shrub not only protected the co-existing species from grazing, but also suppressed salt accumulation; 3) almost pure stand ofPanicum repens; livestock preferred this perennial grass over others. Electrical conductivity (EC) of surface soil did not increase under the vegetation protected from grazing. Litter and other organic matter in the soil suppressed the upward movement of brine. Grazing led to an increase of bare ground where NaCl accumulated, and modified the heterogeneity of vegetation, which was reflected in the degree of salt accumulation.     

Woodlands on farms in southern New South Wales: A longer-term assessment of vegetation changes after fencing

Summary Fencing incentive programmes have been widely used throughout Australia to assist landholders to fence remnant woodland vegetation, to control grazing and improve native vegetation condition. This study investigated vegetation and soil condition in remnant woodlands fenced for 7–9 years in the Murray catchment area in southern New South Wales. Surveys were undertaken at 42 sites, where vegetation condition was assessed in paired fenced and unfenced sites. Semi‐structured interviews were also conducted with landholders to gather management information. Woodlands surveyed were Yellow Box/Blakely's Red Gum (Eucalyptus melliodora/E. blakelyi, 15 sites), Grey Box (E. microcarpa, 13 sites) and White Cypress Pine (Callitris glaucophylla, 14 sites). Fencing resulted in a range of responses which were highly variable between sites and vegetation types. In general, fenced sites had greater tree regeneration, cover of native perennial grasses, less cover of exotic annual grasses and weeds, and less soil compaction than unfenced sites. However, there was greater tree recruitment in remnants to the west of the study area, and tree recruitment was positively correlated with time since fencing. Within sites, tree recruitment tended to occur in more open areas with a good cover of native perennial grasses, as compared to sites with a dense tree canopy, or dominated by exotic annuals grasses or weeds. Forty‐eight per cent of fenced sites had no tree regeneration. There was a significant decline in native perennial grasses, and increase of several unpalatable weeds in many fenced areas, suggesting certain ecological barriers may be preventing further recovery. However, drought conditions and associated grazing are the most likely cause of this trend. A range of grazing strategies was implemented in fenced sites which require further research as a conservation management tool. Continued long‐term monitoring is essential to detect key threats to endangered woodland remnants.