Positive and negative effects of grass,cattle, and wild herbivores on <Emphasis Type="Italic">Acacia</Emphasis> saplings in an East African savanna

Plant–plant interactions can be a complex mixture of positive and negative interactions, with the net outcome depending on abiotic and community contexts. In savanna systems, the effects of large herbivores on tree–grass interactions have rarely been studied experimentally, though these herbivores are major players in these systems. In African savannas, trees often become more abundant under heavy cattle grazing but less abundant in wildlife preserves. Woody encroachment where cattle have replaced wild herbivores may be caused by a shift in the competitive balance between trees and grasses. Here we report the results of an experiment designed to quantify the positive, negative, and net effects of grasses, wild herbivores, and cattle on Acacia saplings in a Kenyan savanna. Acacia drepanolobium saplings under four long-term herbivore regimes (wild herbivores, cattle, cattle + wild herbivores, and no large herbivores) were cleared of surrounding grass or left with the surrounding grass intact. After two years, grass-removal saplings exhibited 86% more browse damage than control saplings, suggesting that grass benefited saplings by protecting them from herbivory. However, the negative effect of grass on saplings was far greater; grass-removal trees accrued more than twice the total stem length of control trees. Where wild herbivores were present, saplings were browsed more and produced more new stem growth. Thus, the net effect of wild herbivores was positive, possibly due to the indirect effects of lower competitor tree density in areas accessible to elephants. Additionally, colonization of saplings by symbiotic ants tracked growth patterns, and colonized saplings experienced lower rates of browse damage. These results suggest that savanna tree growth and woody encroachment cannot be predicted by grass cover or herbivore type alone. Rather, tree growth appears to depend on a variety of factors that may be acting together or antagonistically at different stages of the tree’s life cycle.     

The role of natural agents in the removal of coat-imposed dormancy in Dichrostachys cinerea (L.) Wight et Arn. seeds

Dichrostachys cinerea seeds are impermeable to water and do not germinate readily. The plant, however, contributes to a bush encroachment problem, minimising visibility in wildlife reserves and reducing the area available for grazing. Natural dormancy-breaking conditions must therefore exist, promoting the germinability of these normally dormant seeds. Diurnal temperature fluctuations (50°C/15°C), especially when combined with moisture were found to break dormancy. High temperatures, such as those generated by fire, destroyed seeds. Seeds ingested by herbivorous browsers such as nyala, became permeable. Seed burial for 48 weeks, was a more effective treatment than storage in promoting permeability. Distribution of D. cinerea will therefore be restricted to areas in which these conditions or factors occur. Seeds germinated optimally at 30C in the laboratory.     

Facilitation of seedling establishment by an endemic shrub in tropical coastal sand dunes

Facilitation is predicted to occur on coastal sand dunes as these ecosystems have the harsh physical conditions common during primary succession. In a tropical dune system on the Gulf of Mexico the spatial patterns of plant distribution were analyzed and the hypothesis that facilitation is the responsible mechanism was tested by monitoring changes in the micro-environment and performing a seedling planting experiment under natural conditions. Densities of seedlings and adults of late colonizer grasses were two to six times greater beneath the early colonizer shrub, Chamaecrista chamaecristoides, than in the exposed areas. Temperatures on the sand surface, wind speed and sand accretion were significantly reduced by the shade of Chamaecrista. Only phosphate contents during the dry season were significantly higher in sand in the shade than in exposed sand. In the field experiment, successful establishment of seedlings of two late colonizer grasses (Trachypogon plumosus and Schizachyrium scoparium) was low but was concentrated exclusively beneath the shrubs. The introduced plants were reproductive one year after onset of the experiment and remained vigorous after three years. The observed spatial aggregation among the target species may be a result of improved conditions in the shade of the shrub, which facilitated the survival and establishment of late colonizers. On top of the environmental amelioration, protection from recurrent disturbances such as substrate mobility is an additional beneficial effect of the shrub.     

Seed germination,seedling inoculation and establishment ofAlnus spp. in containers in greenhouse trials

Methods for production of containerized seedlings ofAlnus species were developed which permit nitrogen-fixing nodules to form on the root systems prior to outplanting, in order to provide an early nitrogen input during seedling establishment. The methods are based on procedures for inoculating root systems with suspensions ofFrankia (Actinomycetales), applied either directly in the container cell as a soil drench at the time of seeding, or as a root dip for seedlings transplanted into the containers. Germination of dried, stored seed was enhanced by light and by presoaking for 16 h in water. Pretreatments to overcome seed dormancy or to eliminate fungal pathogens did not further enhance germination. Some loss of seedlings was recorded in the early stages of growth shortly after germination, which is a factor in calculating projected seedling yield. Nodulation and seedling growth were evaluated in terms of growth media characteristics. Seedlings performed well in peat-vermiculite, at soil pH between 5.5 and 8.0.     

Selection on floral display in insect-pollinated Primula farinosa: effects of vegetation height and litter accumulation

Grazing reduces litter thickness and vegetation height and may thereby indirectly affect reproductive success and selection on floral characters in plants with prostrate growth. Reductions in litter thickness and vegetation height should influence both the resource status of plants with leaves positioned close to the ground and the significance of inflorescence height for interactions with pollinators and seed predators. We experimentally examined how simulated grazing of surrounding vegetation affected pollen limitation, fruit predation and fecundity of short-scaped and long-scaped Primula farinosa, which differ markedly in floral display and therefore in expected attractiveness to pollinators. Litter removal and pruning of surrounding vegetation increased fruit and seed production per plant in the year of the treatment and the probability of flowering in the following year. Pollen limitation of fruit and seed production was stronger in the short-scaped morph than in the long-scaped morph, but was not significantly affected by litter removal and simulated grazing of surrounding vegetation. Supplemental hand-pollination reduced seed size in the year of the treatment and flowering probability in the second year, and these effects did not differ among scape morphs or grazing treatments. The results suggest that grazing indirectly favours seed production in P. farinosa, mainly because it increases the resource status of plants that escape damage. Contrary to expectation, there was no strong evidence that litter accumulation and tall vegetation increase the severity of pollen limitation or reduce the relative performance of the short-scaped morph.     

Community changes following shrub invasion of grassland

Summary We studied the development of the shrub Baccharis pilularis ssp consanguinea and its effects on herbs of the annual grassland in Northern California. A series of shands of Baccharis was sampled of ages ranging from 1 yr to>9 yr, representing most of the life cycle of the shrub. In each stand we examined shrub biomass, structure and litterfall. We also determined cover and biomass of all herbaceous species and estimated seed production, seed rain and storage of seed in the soil. Abundances of all herbaceous species declined greatly after Baccharis formed a closed canopy at 2–3 yr, and little seed of herbaceous species was either dispersed into shrub stands or stored in the soil. Exclosures suggested that herbivory by small mammals in the closed shrub stands may be important in reducing the abundance of herbaceous species following shrub invasion of grassland.     

Irrigation and fertilization effects on seed number,size, germination and seedling growth: implications for desert shrub establishment

Plants with limited resources adjust partitioning among growth, survival, and reproduction. We tested the effects of water and nutrient amendments on seed production, size, and quality in Sarcobatus vermiculatus (greasewood) to assess the magnitude and importance of changes in reproductive partitioning. In addition, we assessed interactions among the environment of seed-producing plants (adult plant scale), seed size, and seedling microenvironment (seedling scale) on successful seedling establishment. Interactions of these factors determine the scale of resource heterogeneity that affects seedling establishment in deserts. Both total number of seeds produced per plant and seed quality (weight and germination) increased significantly in the enriched treatment in a 3-year field experiment. Seedling length 3 days after germination and seed N concentration, other measures of seed quality, were higher for seed from both irrigated and enriched plants than for seed from control plants. Field S. vermiculatus seed production and quality can be substantially increased with irrigation and nutrient enrichment at the adult plant scale and this allows management of seed availability for restoration. However, based on a greenhouse study, seedling environment, not the environment of the seed-producing plant or seed size, was the most important factor affecting seedling germination, survival, and growth. Thus it appears that production of more seed may be more important than improved seed quality, because higher quality seed did not compensate for a low-resource seedling environment. For both natural establishment and restoration this suggests that heterogeneity at the scale of seedling microsites, perhaps combined with fertilization of adult shrubs (or multi-plant patches), would produce the greatest benefit for establishing seedlings in the field.     

Combined effects of two arthropod herbivores and water stress on growth of Hypericum species

The interaction between physiological stress and arthropod herbivory was studied using two perennial species of Hypericum. Seedlings of H. perforatum, a herb introduced to Australia and weedy in places, and H. gramineum, an indigenous species, were subjected to water stress and/or herbivory by a mite, Aculus hyperici and an aphid, Aphis chloris. Both arthropods have recently been released in Australia for biological control of H. perforatum. Individually, stresses reduced measures of plant growth. Combinations of the three stresses decreased plant growth by slightly more than the product of their separate effects, suggesting that there is a weak positive interaction exacerbating the damage caused by each stress. Seedlings of the target weed and the non-target indigenous species were equally affected by the arthropods. The implications for weed biological control are discussed.     

Leaf and root extracts of the invasive shrub, Lonicera maackii, inhibit seed germination of three herbs with no autotoxic effects

In addition to effects mediated by resource competition, some invasive plants may impact surrounding vegetation by secreting compounds that are directly inhibitory to growth. Lonicera maackii, an invasive Asian shrub of forests and open areas in eastern and midwestern North America, has devastating effects on understory vegetation, some of which persist even after this shrub is removed. In this study, we explored the potential of aqueous extracts of the leaves and roots of this plant to inhibit seed germination of Impatiens capensis, Alliaria petiolata, Arabidopsis thaliana, and L. maackii in Petri dish bioassays. Both L. maackii root and leaf extracts significantly decreased germination in the three herb species. This inhibitory effect generally increased with increasing extract concentration and was more pronounced with application of leaf extract than root extract. However, when the same extracts were applied to seeds of L. maackii itself, germination was delayed in some cases, but was not significantly reduced by the end of the experiment. Germination of L. maackii seeds even reached significantly higher levels in some extract treatments than in no-extract controls. This implies that L. maackii can successfully inhibit the germination of other plants with few autotoxic effects and may even promote the germination of its own seeds.     

Effects of introduction and exclusion of large herbivores on small rodent communities

In this study we analysed the effects of large herbivores on smallrodent communities in different habitats using large herbivore exclosures. Westudied the effects of three year grazing introduction by red deer(Cervus elaphus L.) in previously ungrazed pine and oakwoodland and the exclusion of grazing by red deer, roe deer(Capreoluscapreolus L.) and mouflon (Ovis ammon musiminL.) in formerly, heavily grazed pine woodland and heathland. At eight exclosuresites within each habitat type, small rodents were captured with live trapsusing trapping grids. At each trapping grid, seed plots of beechnuts(Fagus sylvatica L.) and acorns (Quercusrobur L.) were placed to measure seed predation by rodents.Exclusion of grazing by large herbivores in formerly, heavily grazedhabitats had a significant effect on small rodent communities. Insideexclosureshigher densities of mainly wood mice (Apodemus sylvaticusL.) and field voles (Microtus agrestis L.) were captured.Introduction of grazing by red deer appeared to have no significant negativeeffects on small rodent communities. The seed predation intensity of beechnutsand acorns by small rodents was significantly higher in ungrazed situations,particularly in habitats that were excluded from grazing. The differencesbetween grazing introduction and exclusion effects on small rodent communitiescan be explained by differences in vegetation structure development. Therecovery of heavily browsed understory vegetation after large herbivore grazingexclusion proceeded faster than the understory degradation due to grazingintroduction. Small rodents depend on structural rich vegetations mainly forshelter. We conclude that large herbivores can have significant effects onvegetation dynamics not only via direct plant consumption but also throughindirect effects by reducing the habitat quality of small rodent habitats.     

Indirect interactions between browsers and seed predators affect the seed bank dynamics of a chaparral shrub

Interactions between herbivores and seed predators may have long-term consequences for plant populations that rely on persistent seed banks for recovery after unpredictable fires. We assessed the effects of browsing by deer and seed predation by rodents, ants and birds on the densities of seeds entering the seed bank of Ceanothus cuneatus var. rigidus, a maritime chaparral shrub in coastal California. Ceanothus produced many more seeds when protected from browsers in long-term experimental exclosures than did browsed plants, but the seed densities in the soil beneath browsed and unbrowsed Ceanothus were the same at the start of an intensive one-year study. The density of seeds in the soil initially increased in both treatments following summer seed drop: while densities returned to pre-drop levels within a few weeks under browsed plants, soil seed densities remained high for 5–8 months beneath unbrowsed plants. Rodent abundance (especially deer mice) was higher near unbrowsed plants than >30 m away, and rodents removed Ceanothus seeds from dishes in the experimental plots. At least in the short term, rodent density and rates of seed removal were inversely related to the intensity of browsing. Our data have management implications for maintaining viable Ceanothus populations by regulating the intensity of browsing and the timing, intensity and frequency of fires.Electronic Supplementary Material Supplementary material is available to authorised users in the online version of this article at .     

Recovery patterns of three chaparral shrub species after wildfire

Summary In a mature, even aged stand of mixed chaparral, Rhus laurina (facultative resprouter) had consistently higher water potentials and deeper roots than Ceanothus spinosus (facultative resprouter) and Ceanothus megacarpus (obligate seeder). For two years following a wildfire, the same stand of chaparral had resprouts with higher survivorships, predawn water potentials, stomatal conductances, photosynthetic rates and shoot elongation rates than seedlings. Supplemental irrigation of seedlings during summer months removed differences between resprouts and seedlings suggesting that the cause of such differences was limited water availability to the shoot tissues of seedlings. After two years of postfire regrowth, mean seedling survivorship for the obligate seeder (C. megacarpus) was 42%, whereas seedling survivorship for facultative resprouters was only 18% (C. spinosus) and 0.01% (R. laurina). Our results are consistent with the hypothesis that lack of resprouting ability among obligate seeders is offset by an enhanced ability to establish seedlings after wildfire, allowing obligate seeders to maintain themselves in mixed populations through many fire cycles.     

Effects of two types of herbivores on the population dynamics of a perennial herb

The joint effects of multiple herbivores on their shared host plant have received increasing interest recently. The influence of herbivores on population dynamics of their host plants, especially the relative roles of different types of damage, is, however, still poorly understood. Here, we present a modelling approach, including both deterministic and stochastic matrix modelling, to be used in estimating fitness effects of multiple herbivores on perennial plants. We examined the effects and relative roles of two specialist herbivores, a pre-dispersal seed predator, Euphranta connexa, and a leaf-feeding moth, Abrostola asclepiadis, on the population dynamics and long-term fitness of their shared host plant, a long-lived perennial herb Vincetoxicum hirundinaria (Asclepiadaceae). We collected demographic data during 3 years and combined these data with the effects of natural levels of herbivory measured from the same individuals. We found that both seed predation and leaf herbivory reduced population growth of V. hirundinaria, but only very high damage levels changed the growth trend of the vigorously growing study populations from positive to negative. Demographic modelling indicated that seed predation had a greater impact on plant population growth than leaf herbivory. The effect of leaf herbivory was weaker and diminished with increasing level of seed predation. Evaluation of individual fitness components, however, suggested that leaf herbivory contributed more strongly to host plant fitness than seed predation. Our results emphasize that understanding the effects of a particular herbivore on plant population dynamics requires also knowledge on other herbivores present in the system, because the effect of a particular type of herbivory on plant population dynamics is likely to vary according to the intensity of other types of herbivory. Furthermore, evaluating herbivore impact from using individual fitness components does not necessarily reflect the long-term effects on total plant fitness.     

Seed coat structure and dormancy

An understanding of dormancy mechanisms is of ecological and economic importance. Identification of the level at which dormancy is imposed appears to be species specific. The variation brought about by this therefore requires that developmental studies be included in seed coat dormancy experiments. In most cases, a site of permeability can be identified during the developmental process, and this information can be utilized later to remove dormancy. Under natural conditions, the removal of seed coat dormancy requires the interaction of a number of ecological and physiological dormancy-breaking cues.     

Acacia karroo invasion of grassland: environmental and biotic effects influencing seedling emergence and establishment

Acacia karroo Hayne is the most important woody invader of grassland in South Africa, and can greatly reduce the productivity of grassland. A field experiment was conducted to test the hypotheses that emergence, growth and the 1st year's survival of Acacia karroo would be enhanced by (1) defoliation of the grass sward, (2) increased irradiance, (3) increased moisture availability and (4) its germination within cattle dung pats. The study was conducted on one site above and one below the natural altitudinal treeline of this species in grassland of the eastern Cape, South Africa. Not one seedling emerged from dung pats. Neither location nor the other treatments affected the density of emerging seedlings, although only 40.4 seedlings m^–2 emerged of the 200 seeds m^–2 planted. Shading dramatically increased the density of surviving seedlings. In the open, only 3 and 1.5 seedlings m^–2 remained respectively at the end of the growing season or the beginning of the next, compared to 23.3 and 19.5 seedlings m^–2 under shading for these respective times. This was attributed to the effect of shade on moisture availability in a season which received only 54% of average rainfall. Seedling survival until the end of the growing season was enhanced (30%) by shade at both sites, but also by supplemental water (24%) and defoliation of the sward (7%) at the site above the treeline. Across sites and treatments, seedling survival was related to moisture availability, with no or poor survival for < 500 mm rainfall, indicating this species can only establish in certain rainfall years. Seedling survival over winter was not influenced by treatment, but was greater for larger seedlings. Treatments affected seedling size, in particular seedlings growing under shade and within a dense grass canopy were etiolated. A. karroo seedlings are capable of establishing and surviving within a dense grass sward for at least a year, tolerant of low irradiance and of interference, which, because most seeds do not persist for much longer than a year, suggests this species forms predominantly a seedling bank. This has implications for the invasion of grassland by woody species.     

The effects of neighbors on the growth and survival of shrub seedlings following fire

Where plant species vie for limited resources, disturbances might preclude competition by releasing a flush of nutrients, or by reducing biomass and thereby diminishing the consumption of resources. However, if new seedlings colonize in clumps, they may still deplete resources within the local aggregations, which may then reduce their growth and survivorship. We investigated competition among seedlings in a burned area by examining the relationship between the performance of newly established shrub seedlings of Ceanothus impressus and (1) the proximity and (2) the identity of their near neighbors. We also investigated the relationship between neighbor proximity and the availability of water. Both survivorship and growth of C. impressus were positively associated with increasing distance to near neighbors, in a manner consistent with resource competition. The availability of water (as determined by pre-dawn xylem pressure potentials) tended to be greater when neighbors were farther away, providing evidence that water was a resource for which plants were competing. This conclusion is reinforced by the finding that the effects of neighbors were stronger in drought years, suggesting that yearly variation in the availability of an important resource (water) can affect the strength of competitive interactions. This suggests that after disturbances, when some resources are apparently abundant on a large scale, competition may be important in determining the small scale patterns of seedling growth and survival.     

Initial effects of experimental warming on carbon exchange rates, plant growth and microbial dynamics of a lichen-rich dwarf shrub tundra in Siberia

Biological soil crusts can affect seed germination and seedling establishment. We have investigated the effect of biological soil crusts on seed water status as a potential mechanism affecting seed germination. The seed water potential of two annual grasses, one exotic Bromus tectorum L. and another native Vulpia microstachys Nutt., were analyzed after placing the seeds on bare soil, on a crust that contains various lichens and mosses (mixed crust), or on a crust dominated by the crustose lichen Diploschistes muscorum (Scop.) R. Sant. (Diploschistes crust). Seed water potential and germination were similar on the bare soil and the mixed crust, except for the initial germination of V. microstachys, which was higher on the mixed crust than on the bare soil. For the two grasses studied, seed water potential was significantly higher on the bare soil and mixed crust than on the Diploschistes crust. These differences in water potential correlated with differences in germination, which was much lower on the lichen crust. Experiments were conducted under two watering regimens. Increasing the frequency of watering amplified the differences in seed water potential and germination between the Diploschistes crust and the other two surfaces. For a particular watering regimen, the bare soil, mixed crust, and Diploschistes crust received the same amount of water, but they reached significantly different water potentials. Throughout the experiments, the water potential of the soil and mixed crust remained above −0.6 MPa, while there was a marked decline in the water potential of the Diploschistes surface to about −4 MPa. To ascertain that water was the major factor limiting germination on the Diploschistes crust, we conducted germination tests in an environment with 100% relative humidity. Under these conditions, germination on the Diploschistes crust was similar to that on the bare soil. However, the seeds that germinated on the Diploschistes crust did not penetrate this surface and approximately 60% of their root tips became necrotic. Our results indicate that the presence of D. muscorum can inhibit seedling establishment by two mechanisms: a reduction in seed water absorption and an increase in root tip mortality.     

Effects of herbivory and herb interference on oak establishment in a semi-arid temperate savanna

Abstract. Early establishment of Quercus emoryi seedlings in the grassland phase of a semi-arid temperate savanna was evaluated in southern Arizona, USA. Controlled-environment experiments indicated that Quercus seedlings are tolerant of frequent clipping, but undipped seedlings produce more biomass than clipped seedlings. Seedlings were much less tolerant of clipping or defoliation in the field than in the growth chamber, and most seedlings died after being clipped or defoliated by herbivores. Invertebrate herbivory, which is rarely mentioned as an important factor limiting seedling establishment, was the leading cause of seedling mortality in the field study. Below-ground interference from herbs had a greater impact on seedling survival and growth than above-ground interference.     

Resistance to herbicide and susceptibility to herbivores: environmental variation in the magnitude of an ecological trade-off

Trade-offs can maintain genetic diversity and constrain adaptation; however, their magnitude may depend on ecological factors. I considered whether resistance to the herbicide triazine in Amaranthus hybridus (Amaranthaceae) imposed the trade-off of increasing susceptibility to herbivorous insects. I grew triazine-resistant and triazine-susceptible plants under contrasting levels of light and fertilization, and quantified susceptibility to herbivores using the specialist Disonycha glabrata (Coleoptera: Chrysomelidae) and the generalist Trichoplusia ni (Lepidoptera: Noctuidae). Resistance to triazine increased susceptibility to both species of herbivorous insects, as manifested by greater feeding preference, growth, and survival of herbivores. However, these effects were more pronounced with T. ni and for plants grown under high light. My results demonstrate the presence of a trade-off between resistance to triazine and susceptibility to herbivorous insects that may in turn impose an ecologically based fitness cost, and illustrate the potential for this cost to vary across environments.