Invasion of maritime chaparral by the introduced succulentCarpobrotus edulis

Invasion by the alien succulent,Carpobrotus edulis, has become a common occurrence after fire in maritime chaparral in coastal California, USA. We studied post-burnCarpobrotus establishment in chaparral that lackedCarpobrotus plants before the fire and compared seedbank and field populations in adjacent burned and unburned stands.Carpobrotus seeds were abundant in deer scat and in the soil before burning. Burning did not enhance germination: many seeds were apparently killed by fire and seed bank cores taken after fire revealed no germinable seeds. Laboratory tests showed that temperatures over 105°C for five minutes killedCarpobrotus seeds. In a field experiment involving use of herbivore exclosures, we found that herbivory was an important source of mortality for seedlings in both burned and unburned chaparral. All seedlings, however, died outside of the burn regardless of the presence of cages. Establishment there is apparently limited by factors affecting plant physiology. In the burned area, seedlings that escaped herbivory grew very rapidly. Overall, it appears that herbivory limited seedling establishment in both burned and unburned sites but that the post-burn soil environment supportedCarpobrotus growth in excess of herbivore use, thus promoting establishment.     

Plant functional group responses to fire frequency and tree canopy cover gradients in oak savannas and woodlands

Questions: How do fire frequency, tree canopy cover, and their interactions influence cover of grasses, forbs and understorey woody plants in oak savannas and woodlands? Location: Minnesota, USA. Methods: We measured plant functional group cover and tree canopy cover on permanent plots within a long‐term prescribed fire frequency experiment and used hierarchical linear modeling to assess plant functional group responses to fire frequency and tree canopy cover. Results: Understorey woody plant cover was highest in unburned woodlands and was negatively correlated with fire frequency. C4‐grass cover was positively correlated with fire frequency and negatively correlated with tree canopy cover. C3‐grass cover was highest at 40% tree canopy cover on unburned sites and at 60% tree canopy cover on frequently burned sites. Total forb cover was maximized at fire frequencies of 4–7 fires per decade, but was not significantly influenced by tree canopy cover. Cover of N‐fixing forbs was highest in shaded areas, particularly on frequently burned sites, while combined cover of all other forbs was negatively correlated with tree canopy cover. Conclusions: The relative influences of fire frequency and tree canopy cover on understorey plant functional group cover vary among plant functional groups, but both play a significant role in structuring savanna and woodland understorey vegetation. When restoring degraded savannas, direct manipulation of overstorey tree canopy cover should be considered to rapidly reduce shading from fire‐resistant overstorey trees. Prescribed fires can then be used to suppress understorey woody plants and promote establishment of light‐demanding grasses and forbs.     

Variation in insect herbivory across a salt marsh tidal gradient influences plant survival and distribution

Herbivore damage and impact on plants often varies spatially across environmental gradients. Although such variation has been hypothesized to influence plant distribution, few quantitative evaluations exist. In this study I evaluated patterns of insect herbivory on an annual forb, Atriplex patula var. hastata, across a salt marsh tidal gradient, and performed experiments to examine potential causes and consequences of variation in herbivory. Damage to plants was generally twice as great at mid-tidal elevations, which are more frequently inundated, than at higher, less stressful, elevations at five of six surveyed sites. Field herbivore assays and herbivore preference experiments eliminated the hypothesis that plant damage was mediated by herbivore response to differences in host plants across the gradient. Alternately, greater herbivore densities in the mid-marsh, where densities of an alternate host plant (Salicornia europaea) were high, were associated with greater levels of herbivory on Atriplex, suggesting spillover effects. The effect of insect herbivores on host plant performance varied between the two sites studied more intensively. Where overall herbivore damage to plants was low, herbivory had no detectable effect on plant survival or seed production, and plant performance did not significantly differ between zones. However, where herbivore damage was high, herbivores dramatically reduced both plant survival (>50%) and fruit production (40-70%), and their effects were stronger in the harsher mid-marsh than the high marsh. Thus herbivores likely play a role in maintaining lower Atriplex densities in mid-marsh. Overall, these results suggest that variation in herbivore pressure can be an important determinant of patterns of plant abundance across environmental gradients.     

Post‐fire succession and 20th century reduction in fire frequency on xeric southern Appalachian sites

Abstract. We document post‐fire succession on xeric sites in the southern Appalachian Mountains, USA and assess effects of 20th century reduction in fire frequency on vegetation structure and composition. Successional studies over 18 yr on permanent plots that had burned in 1976–1977 indicate that tree mortality and vegetation response varied with fuel load and fire season. In the first three years after fire, hardwood sprouts dominated tree regeneration. On sites where summer and autumn fires reduced litter depth to less than 1 cm, densities of shade‐intolerant Pinus seedlings increased steadily over this period. 4 to 8 yr after fire, large numbers of newly established seedlings and sprouts had grown to 1 – 10 cm DBH. By year 18 growth of these saplings led to canopy closure on most sites. Herbaceous cover and richness peaked in the first decade after fire, then declined. On similar sites that had not burned in more than 50 yr, regeneration of shade‐intolerant Pinus spp. and mean cover and richness of herbs were considerably lower than those observed on recently burned plots. Reconstructions of landscape conditions based on observed post‐fire succession and 20th century changes in fire regime suggest that reductions in fire frequency circa 1940 led to substantial changes in forest structure and decreases in cover and richness of herbaceous species.     

Effects of fire severity in a north Patagonian subalpine forest

Abstract. Question: What is the relative importance of fire‐induced canopy mortality, soil burning and post‐fire herbivory on tree seedling performance? Location: Subalpine Nothofagus pumilio forests at Challhuaco valley (41°13'S, 71°19'W), Nahuel Huapi National Park, Argentina. Methods: We fenced and transplanted soils of three burning severities along a fire severity gradient produced by a fire in 1996. Over two growing seasons we monitored soil water, direct incoming solar radiation, seedling survival, final seedling total biomass and root/shoot ratio. Additionally, we assessed severity‐related changes in soil properties. Results: Incoming radiation (an indicator of the amount of canopy cover left by the fire) was the primary factor influencing spring and summer top soil water availability, first and second‐year seedling survival and seedling growth. While seedling survival and soil water content were negatively affected by increased radiation, seedling final biomass was highest in very open microsites. Burned soils showed lower water holding capacity and soil carbon; however these changes did not affect topsoil water, and, contrary to expectation, there was a slight tendency toward higher seedling survival on more heavily burned soils. Herbivory significantly reduced seedling survival, but only under high‐radiation conditions. While the effect of radiation on final seedling biomass was not affected by herbivory, R/S ratios were significantly reduced by herbivory in high radiation micro sites. Conclusions: Despite inducing faster aerial growth, increased radiation and herbivory in severely burned sites may effectively prevent post‐fire regeneration in north Patagonian subalpine forest where seed sources are not limiting.     

Prefire grazing by cattle increases postfire resistance to exotic annual grass (Bromus tectorum) invasion and dominance for decades

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Introduced ungulate herbivore alters soil processes after fire

Ungulate herbivory can have profound effects on ecosystem processes by altering organic inputs of leaves and roots as well as changing soil physical and chemical properties. These effects may be especially important when the herbivore is an introduced species. Utilizing large mammal exclosures to prevent access by introduced elk at multiple sites along a fire chronosequence, we examined the effects of elk herbivory and fire on soil microbial activity and nutrient availability. Using time since fire as a co-variate and herbivore exclosures, paired with areas outside of the exclosures, we hypothesized that reductions in plant biomass due to herbivory would reduce organic inputs to soils and impact soil microbial activities and nutrient storage. We found three major patterns: (1) when elk were excluded, surface mineral soils had higher soil organic carbon (C), total nitrogen (N), microbial N pools, and increased extra-cellular enzyme activity of a C-acquiring enzyme across a gradient of time since fire. (2) When introduced elk are present, the activity of some extracellular enzymes as well as NO₃ ⁻ availability are enhanced in the soil but the post-fire patterns described above with respect to nutrient accrual over time are delayed. (3) Herbivory by an introduced ungulate upsets the trajectory of ecosystem “recovery” after wildfire and delays soil C and N dynamics by an estimated 14.5–21 years, respectively. These results suggest that introduced, browsing herbivores significantly decelerate ecosystem processes but herbivory by exotics may also result in unpredictability in specific soil responses.     

Effects of Prescribed Fire on Shinnery Oak ( Quercus havardii) Plant Communities in Western Oklahoma

Changes in structural and compositional attributes of shinnery oak (Quercus havardii Rydb.) plant communities have occurred in the twentieth century. These changes may in part relate to altered fire regimes. Our objective was to document effects of prescribed fire in fall (October), winter (February), and spring (April) on plant composition. Three study sites were located in western Oklahoma; each contained 12, 60 × 30‐m plots that were designated, within site, to be seasonally burned, annually burned, or left unburned. Growing season canopy cover for herbaceous and woody species was estimated in 1997–1998 (post‐treatment). At one year post‐fire, burning in any season reduced shrub cover, and spring burns reduced cover most. Winter and annual fires increased cover of rhizomatous tallgrasses, whereas burning in any season decreased little bluestem cover. Perennial forbs increased with fall and winter fire. Shrub stem density increased with fire in any season. Communities returned rapidly to pre‐burn composition with increasing time since fire. Fire effects on herbaceous vegetation appear to be manifested through increases in bare ground and reduction of overstory shrub dominance. Prescribed fire can be used as a tool in restoration efforts to increase or maintain within and between community plant diversity. Our data suggest that some plant species may require or benefit from fire in specific seasons. Additional research is needed to determine the long‐term effects of repeated fire over time.     

Increase of fast nutrient cycling in grassland microcosms through insect herbivory depends on plant functional composition and species diversity

Nutrient cycling in terrestrial ecosystems is affected by various factors such as plant diversity and insect herbivory. While several studies suggest insect herbivory to depend on plant diversity, their interacting effect on nutrient cycling is unclear. In a greenhouse experiment with grassland microcosms of one to six plant species of two functional groups (grasses and legumes), we tested the influence of plant species richness (diversity) and functional composition on plant community biomass production, insect foliar herbivory, soil microbial biomass, and nutrient concentrations in throughfall. To manipulate herbivory, zero, three or six generalist grasshoppers (Chorthippus parallelus) were added to the plant communities. Increasing plant species richness increased shoot biomass and grasshopper performance, without significantly affecting root biomass or insect herbivory. Plant functional composition affected all of these parameters, e.g. legume communities showed the highest shoot biomass, the lowest grasshopper performance and suffered the least herbivory. Nutrient concentrations (dissolved mineral N, PO₄‐P, SO₄‐S) and pH in throughfall increased with herbivory. PO₄‐P and pH increases were positively affected by plant diversity, especially under high herbivore pressure. Plant functional composition affected several throughfall variables, sometimes fully explaining diversity effects. Increasing plant diversity tended to increase soil microbial biomass, but only under high herbivore pressure. Faeces quantities strongly correlated with changes in pH and PO₄‐P; frass may therefore be an important driver of throughfall pH and a main source of PO₄‐P released from living plants. Our results indicate that insect herbivory may significantly influence fast nutrient cycling processes in natural communities, particularly so in managed grasslands.     

Impacts of feral horses and deer on an endangered woodland of Kosciuszko National Park

In recent years, the impacts of rapidly increasing populations of feral horses and deer on the vegetation and stability of soils have become highly visible and widespread in Kosciuszko National Park. We investigated these impacts in the White Cypress Pine (Callitris glaucophylla Joy Thomps. & L.A.S. Johnson) – White Box (Eucalyptus albens Benth) woodlands of the lower Snowy River valley. This woodland is a component of the White Box‐Yellow Box‐Blakely's Red Gum Grassy Woodland and Derived Native Grasslands complex that is nationally listed as a critically endangered ecological community. To investigate the severity of the impacts of feral horse (Equus caballus) and deer (Dama dama and Rusa unicolor) in the valley in 2013 and 2017/18, we surveyed fenced exclosures and paired grazed plots that were first established and surveyed in 1984 and re‐surveyed in 1987. Using LFA and VAST methodologies (not used in 1987), we compared the relative response of environmental variables in plots inside and outside the exclosures in an attempt to ascertain recent herbivore impacts. While there was no evidence of horses or deer from dung surveys in 1987, in 2018, 84% of the dung was from horses, 13% from deer, 1% from rabbits and 2% from macropods. Total herbivore dung density increased fourfold since the 1987 survey. On the understanding that all plots had the same starting condition in 1984 with respect to prior herbivory, we deduce that horses and deer are having significant ecological impacts. There was a far greater cover of understorey plants and the midstorey was denser and taller inside the exclosures. Outside the exclosures, the vegetation cover was far more sparse and soil erosion was active and extensive. The total number of invertebrates captured in small pitfall traps was nearly twice as many within the exclosures compared to the grazed plots. The dense even‐aged regrowth overstorey stands of White Cypress Pine, inside and outside the exclosures, have changed little in 34 years.     

Herbivore‐induced changes in flower scent and morphology affect the structure of flower–visitor networks but not plant reproduction

Herbivory induces various responses in plants, thus altering the plants’ phenotype in chemical and morphological traits. Herbivore‐induced changes in vegetative plant parts, plant‐physiological mechanisms, and effects on plant‐animal interactions have been intensively studied from species to community level. In contrast, we are just beginning to examine herbivore‐induced effects on reproductive plant parts and flower–visitor interactions, especially in a community context. We investigated the effect of herbivory at different plant developmental stages on plant growth, floral and vegetative phenotype and reproduction in Sinapis arvensis (Brassicaceae). Additionally, we tested how herbivore‐induced plant responses affect flower–visitor interactions and plant reproduction in species‐rich communities. Our results indicate that the timing of herbivory affects the magnitude of changes in plant traits. Herbivory in early but not in late development accelerated the plant's flowering phenology, reduced vegetative growth, increased stem trichome density and altered floral morphology and scent. These findings suggest age‐dependent tradeoffs between growth, defense and reproduction. Herbivore‐induced changes in flower traits also affected flower–visitor interactions in a community context with effects on the structure of flower–visitor networks. However, changes in the network structure had neglectable effects on plant reproduction, i.e. plants were able to compensate altered flower visitor behavior. Thus, herbivory is a source of intraspecific variation in reproductive traits, which can be behaviorally relevant for potential pollinators. However, plants were capable to maintain reproductive success suggesting a tolerance against herbivory. We conclude that in our study system induced direct or indirect defenses that have often been shown to decrease negative effects of herbivores on vegetative plant parts come at no costs for plant reproduction.     

Loss of a large grazer impacts savanna grassland plant communities similarly in North America and South Africa

Large herbivore grazing is a widespread disturbance in mesic savanna grasslands which increases herbaceous plant community richness and diversity. However, humans are modifying the impacts of grazing on these ecosystems by removing grazers. A more general understanding of how grazer loss will impact these ecosystems is hampered by differences in the diversity of large herbivore assemblages among savanna grasslands, which can affect the way that grazing influences plant communities. To avoid this we used two unique enclosures each containing a single, functionally similar large herbivore species. Specifically, we studied a bison (Bos bison) enclosure at Konza Prairie Biological Station, USA and an African buffalo (Syncerus caffer) enclosure in Kruger National Park, South Africa. Within these enclosures we erected exclosures in annually burned and unburned sites to determine how grazer loss would impact herbaceous plant communities, while controlling for potential fire-grazing interactions. At both sites, removal of the only grazer decreased grass and forb richness, evenness and diversity, over time. However, in Kruger these changes only occurred with burning. At both sites, changes in plant communities were driven by increased dominance with herbivore exclusion. At Konza, this was caused by increased abundance of one grass species, Andropogon gerardii, while at Kruger, three grasses, Themeda triandra, Panicum coloratum, and Digitaria eriantha increased in abundance.     

EFFECTS OF FIRE IN TALLGRASS PRAIRIE ON GROWTH AND REPRODUCTION OF PRAIRIE CONEFLOWER (RATIBIDA COLUMNIFERA: ASTERACEAE)

Effects of fire on growth and reproduction of the perennial forb Ratibida columnifera were studied on the Konza Prairie Research Natural Area in northeastern Kansas, USA. Populations were sampled in seven different tallgrass prairie watersheds that varied in fire frequency and in the number of years elapsed since the last fire. Plants from sites not burned for many years were 2.6 times larger and produced 50% more stems than plants from recently burned sites. Number of seeds per plant was also higher in long-unburned sites due to greater numbers of flower heads per plant and greater numbers of achenes produced per head. Reproductive effort (ratio of inflorescence biomass to total vegetative biomass) was 33% lower in annually burned prairie than in any of the other sites. Significant differences in the relationships of inflorescence biomass to vegetative plant biomass in burned vs. unburned sites indicated that burning causes direct changes in plant reproductive effort independent from its effects on plant size. There was no clear relationship between patterns of seed production among sites and patterns of R. columnifera abundance. Ratibida columnifera responses to fire are most likely a result of changes in the relative competitive abilities of forbs and the dominant perennial grasses due to post-fire changes in abiotic conditions rather than a result of direct effects of fire on the fate of buds and subsequent vegetative and floral development.     

Synergistic influences of introduced herbivores and fire on vegetation change in northern Patagonia,Argentina

Question: We investigated how cattle and European hares, the two most widespread exotic herbivores in Patagonia, affect species composition, life‐form composition and community structure during the first 6 years of vegetation recovery following severe burning of fire‐resistant subalpine forests and fire‐prone tall shrublands. We asked how the effects of introduced herbivores on post‐fire plant community attributes affect flammability of the vegetation. Location: Nahuel Huapi National Park, northwest Patagonia, Argentina Methods: We installed fenced plots to exclude livestock and European hares from severely burned subalpine forests of Nothofagus pumilio and adjacent tall shrublands of N. antarctica. The former is an obligate seed reproducer, whereas the latter and all other woody dominants of the shrubland vigorously resprout after burning. Results: Repeated measures ANOVA of annual measurements over the 2001‐2006 period indicate that cattle and hare exclusion had significant but complex effects on the cover of graminoids, forbs, climber species and woody species in the two burned community types. Significant interactions between the effects of cattle and hares varied by plant life forms between the two communities, which implies that their synergistic effects are community dependent. Conclusions: Following severe fires, the combined effects of cattle and hares inhibit forest recovery and favour transition to shrublands dominated by resprouting woody species. This herbivore‐induced trend in vegetation structure is consistent with the hypothesis that the effects of exotic herbivores at recently burned sites contribute to an increase in the overall flammability of the Patagonian landscape.     

Functional identity versus species richness: herbivory resistance in plant communities

The resistance of a plant community against herbivore attack may depend on plant species richness, with monocultures often much more severely affected than mixtures of plant species. Here, we used a plant–herbivore system to study the effects of selective herbivory on consumption resistance and recovery after herbivory in 81 experimental grassland plots. Communities were established from seed in 2002 and contained 1, 2, 4, 8, 16 or 60 plant species of 1, 2, 3 or 4 functional groups. In 2004, pairs of enclosure cages (1 m tall, 0.5 m diameter) were set up on all 81 plots. One randomly selected cage of each pair was stocked with 10 male and 10 female nymphs of the meadow grasshopper, Chorthippus parallelus. The grasshoppers fed for 2 months, and the vegetation was monitored over 1 year. Consumption resistance and recovery of vegetation were calculated as proportional changes in vegetation biomass. Overall, grasshopper herbivory averaged 6.8%. Herbivory resistance and recovery were influenced by plant functional group identity, but independent of plant species richness and number of functional groups. However, herbivory induced shifts in vegetation composition that depended on plant species richness. Grasshopper herbivory led to increases in herb cover at the expense of grasses. Herb cover increased more strongly in species-rich mixtures. We conclude that selective herbivory changes the functional composition of plant communities and that compositional changes due to selective herbivory depend on plant species richness.     

Complex effects of mammalian grazing on extramatrical mycelial biomass in the Scandes forest‐tundra ecotone

Mycorrhizal associations are widespread in high‐latitude ecosystems and are potentially of great importance for global carbon dynamics. Although large herbivores play a key part in shaping subarctic plant communities, their impact on mycorrhizal dynamics is largely unknown. We measured extramatrical mycelial (EMM) biomass during one growing season in 16‐year‐old herbivore exclosures and unenclosed control plots (ambient), at three mountain birch forests and two shrub heath sites, in the Scandes forest‐tundra ecotone. We also used high‐throughput amplicon sequencing for taxonomic identification to investigate differences in fungal species composition. At the birch forest sites, EMM biomass was significantly higher in exclosures (1.36 ± 0.43 g C/m²) than in ambient conditions (0.66 ± 0.17 g C/m²) and was positively influenced by soil thawing degree‐days. At the shrub heath sites, there was no significant effect on EMM biomass (exclosures: 0.72 ± 0.09 g C/m²; ambient plots: 1.43 ± 0.94). However, EMM biomass was negatively related to Betula nana abundance, which was greater in exclosures, suggesting that grazing affected EMM biomass positively. We found no significant treatment effects on fungal diversity but the most abundant ectomycorrhizal lineage/cortinarius, showed a near‐significant positive effect of herbivore exclusion (p = .08), indicating that herbivory also affects fungal community composition. These results suggest that herbivory can influence fungal biomass in highly context‐dependent ways in subarctic ecosystems. Considering the importance of root‐associated fungi for ecosystem carbon balance, these findings could have far‐reaching implications.     

Endophyte-grass-herbivore interactions: the case of Neotyphodium endophytes in Arizona fescue populations

Neotyphodium endophytes in introduced agronomic grasses are well known to increase resistance to herbivores, but little is known of interactions between Neotyphodium endophytes and herbivores in native grass populations. We investigated whether endophytes mediate plant-herbivore interactions in a native grass species, Festuca arizonica in the southwestern United States, in two ways. First, to test the prediction that the presence and frequency of endophyte-infected (E+) plants should increase with increasing herbivory, we determined endophyte frequencies over a 4-year period in six natural Arizona fescue populations. We compared Neotyphodium frequency among plants growing inside and outside long-term vertebrate grazing exclosures. Second, we experimentally tested the effects of Neotyphodium infection, plant clone, and soil nutrients on plant resistance to the native grasshopper Xanthippus corallipes. Contrary to predictions based upon the hypothesis that endophytes increase herbivore resistance, levels of infection did not increase in plants subjected to grazing outside of exclosures relative to ungrazed plants within exclosures. Instead, endophyte frequencies tended to be greater inside the exclosures, where long-term vertebrate grazing was reduced. The grasshopper bioassay experiment corroborated these long-term patterns. Survival of grasshoppers did not differ between infected (E+) and uninfected (E–) plants. Instead, mean relative growth rate of grasshoppers was higher on E+ grasses than on E– ones. Growth performance of newly hatched grasshopper nymphs varied among host plant clones, although two of six clones accounted for most of this variation. Our results suggest that Neotyphodium-grass-herbivore interactions may be much more variable in natural communities than predicted by studies of agronomically important Neotyphodium-grass associations, and herbivory is not always the driving selective force in endophyte-grass ecology and evolution. Thus, alternative hypotheses are necessary to explain the wide distribution and variable frequencies of endophytes in natural plant populations. Received: 15 February 1999 / Accepted: 19 July 1999     

Pyric Herbivory and the Nexus Between Forage,Fire and Native and Introduced Large Grazing Herbivores in Australian Tropical Savannas

Earth’s tropical savannas typically support high biomass of diverse grazing herbivores that depend on a highly fluctuating resource: high-quality forage. An annual wet–dry cycle, fire and herbivory combine to influence forage quality and availability throughout the year. In the savannas of northern Australia, a depauperate suite of large native (marsupial) herbivores (wallaroos [Osphranter spp.] and the agile wallaby [Notamacropus agilis]) compete for resources with non-native large herbivores introduced in the late nineteenth century, particularly bovines (feral and managed cattle [Bos spp.] and feral water buffalo [Bubalus bubalis]) that now dominate the landscape. Anecdotal reports of recent population declines of large macropods and negative impacts of bovines highlight the need to better understand the complex relationship between forage, fire and abundance of native and introduced large herbivores. The pyric herbivory conceptual model, which posits complex feedbacks between fire and herbivory and was developed outside Australia, predicts that native and introduced large herbivores will both respond positively to post-fire forage production in Australian savannas where they co-occur. We used grazing exclosures, forage biomass and nutrient analyses and motion-sensor camera-trapping to evaluate the overall robustness of the pyric herbivory model in the Australian context, specifically whether forage quantity and quality are impacted by herbivory, season and fire activity, and which forage attributes most influence large grazing herbivore abundance. Forage quantity, as measured by live, dead and total herbaceous biomass and proportion of biomass alive, was higher inside herbivore exclosures, even at relatively low densities of herbivores. Forage quality, as measured by fibre content, was not affected by herbivory, however, crude protein content of live herbaceous biomass was greater outside herbivore exclosures. Recent fire was an important predictor of all measures of forage quantity and quality. Recent fire occurrence decreased overall quantity (biomass) but increased quality (decreased fibre content and increased crude protein content); late dry season fires resulted in forage with the highest crude protein content. The predictions of the pyric herbivory conceptual model are consistent with observations of the feeding behaviour of introduced bovines and some large macropods in northern Australian savannas, lending support to the global generality of pyric herbivory in fire-prone grassy biomes.

     

The Role of Seed and Vegetative Reproduction in Plant Recruitment and Demography in Tallgrass Prairie

Recruitment, establishment and survivorship of seed- and vegetatively-derived shoots were quantified biweekly in annually burned and infrequently burned tallgrass prairie to investigate the contributions of seed and vegetative reproduction to the maintenance and dynamics of tallgrass prairie plant populations, the demography of seedlings and ramets, and the influence of fire on the demography of grasses and forbs. Clonally produced grass and forb ramets comprised >99%of all established shoots present at the end of the growing season, whereas established seedlings accounted for <1%,emphasizing the rarity of successful seedling establishment and the importance of vegetative reproduction in driving the annual regeneration and dynamics of aboveground plant populations in tallgrass prairie. Most recruitment from vegetative reproduction occurred early in the growing season and was higher in annually burned than infrequently burned sites, although low levels of new stem recruitment occurred continuously throughout the growing season. Peak recruitment on annually burned prairie coincided with peak recruitment of the dominant C₄ grasses Andropogon gerardii and Sorghastrum nutans prior to prescribed spring fire, with a second peak in recruitment occurring following fire. On infrequently burned prairie, grass and forb recruitment was highest in early April and declined steadily through May. The naturalized C₃ grass, Poa pratensis, was responsible for most of the early recruitment on unburned sites, whereas A. gerardii contributed most to recruitment later in May. Infrequently burned prairie was dominated by these two grasses and contained a larger forb component than annually burned prairie. The principal demographic effect of fire was on ramet natality rather than mortality. Fire regime, plant functional group, or timing of cohort emergence before or after fire did not affect ramet survivorship. C₄ grass shoots that emerged early and were damaged by fire showed similar survivorship patterns to tillers that emerged after fire. Differences in species composition between annually burned and infrequently burned prairie are driven by fire effects on vegetative reproduction and appear to be related principally to the effect of fire and detritus accumulation on the development of belowground vegetative meristems of C₄ grasses and their emergence dynamics.     

Interactive response of herbivores,soils and vegetation to annual burning in a South African savanna

Abstract Despite decades of research, the primary factors determining savanna structure remain elusive – a conundrum termed ‘the savanna problem’. After 47 years of annual burning in Terminalia woodland and Acacia/Combretum savanna on sandy, granite‐derived soils in the southern Kruger National Park, South Africa, a dense cover of trees and shrubs persists on some burnt plots and is largely absent from others. We postulated that intense browsing pressure by antelope and other herbivores prevents recruitment of trees in burnt plots and that herbivores concentrate on plots that are richest in nutrients. Herbivore abundance did not show a relationship with soil macronutrients and we consequently investigated micronutrient status. The reduction in tree cover as a result of annual burning was positively correlated with mass of herbivores (15–1500 kg) (r ² = 0.61, n = 8). This index of herbivore abundance was in turn positively correlated with total Zn (r ² = 0.64, n = 8). Other indices of herbivore abundance showed significant relationships with total clay content and total Mn. We suggest that herbivores concentrate on sites with greater clay content (possibly due to a greater availability of micronutrients), and that tree cover can remain relatively dense under a regime of annual burning if browsing pressure is not intense. The long‐term burn experiments in the Kruger National Park savanna provide a platform for unravelling the savanna problem. Determining possible interactions between soil properties, herbivory and fire is a step in this direction.