Effect of nutrient availability, grazer assemblage and seagrass source population on the interaction between Thalassia testudinum (turtle grass) and its algal epiphytes

Seagrass leaves are often densely covered by epiphytic algae which can suppress seagrass productivity and has been implicated in declines of seagrass meadows worldwide. The net effect of epiphytes on seagrass growth and morphology depends on the independent and interactive effects of a variety of factors, including nutrient availability and the intensity of grazing on epiphytes. Here I report the results of a mesocosm experiment designed to test the effects of nutrient addition and within-functional group variation (grazer species composition and the source population of seagrass) on the strength of the interactions among grazers, epiphytes, and turtle grass (Thalassia testudinum). Turtle grass ramets from two sites in the northern Gulf of Mexico were cleared of epiphytes and transplanted into common-garden mesocosms. Replicate ramets were grown in a split-split plot design with two levels of dissolved nutrients and four different grazer species combinations (Tozeuma carolinense alone, Pagurus maclaughlinae alone, both species together, and no grazers present). As expected, grazers had a significant negative effect on epiphyte biomass/leaf area and a significant positive effect on turtle grass growth in the mesocosms. The two species were more similar in their direct effects on epiphyte biomass than in their indirect effects on turtle grass growth; this may reflect differences in epiphyte community composition under different grazer treatments. The effect of nutrient addition on turtle grass growth depended critically on the intensity of grazing: in the presence of grazers, turtle grass tended to produce a greater biomass of new leaf tissue in the tanks with nutrients added than in the control tanks. However, when grazers were absent, the direction of the effect was reversed, and plants with nutrients added grew less than the control plants. The two source populations of turtle grass differed significantly in epiphyte biomass/leaf area accrued in the mesocosms as well as in the strength of the effect of grazers on turtle grass growth. This suggests that population differentiation in seagrass interactions with epiphytes, as well as spatial and temporal variation in resources and grazer community composition, can greatly effect the role of epiphytes in limiting seagrass productivity.     

Ecological Engineering by a Mega-Grazer: White Rhino Impacts on a South African Savanna

Herbivory can change the structure and spatial heterogeneity of vegetation. We ask whether all species of grazers in a savanna ecosystem can have this effect or whether megaherbivores (>1000 kg) have a ‘special’ role that cannot be replicated by other species of grazers. We performed a replicated landscape scale experiment that examined the effects of White Rhino on the grass sward, on other species of grazing mammals and on the movement of fire through the landscape. White Rhino maintained short grass (‘lawn’) patches in mesic areas (∼750 mm pa) with increases in grass sward height when they were removed. Other species of grazers were unable to maintain short grass communities when White Rhino were removed. In semi-arid areas (∼600 mm pa) other, smaller grazers were able to maintain short grass communities in the absence of White Rhino and sward height did not increase. White Rhino removals affected fire by increasing fuel loads and fuel continuity. This resulted in larger, less patchy fires. We propose that the White Rhino acts as an influential ecosystem engineer, creating and maintaining short grass swards, which alter habitat for other grazers and change the fire regime. These results indicate the existence of context-dependent facilitation between White Rhino and other grazers in mesic, but not in semi-arid, savannas. Such top down effects on the ecosystem may have been much more widespread before the extinction of large grazers in the Pleistocene.     

The ecological significance of rapid wound-induced changes in plants: insect grazing and plant competition

Summary This paper investigates the hypothesis that a rapidly induced phytochemical response to grazing damage, such as that seen in tomato, serves to deflect insect herbivores away from leaves soon after damaging them (the grazing dispersal hypothesis). As a result, grazing damage is more dispersed than it otherwise would be, and young leaves, which may be of particular importance to a plant in competition for light, are not damaged excessively. In the first experiment, artificial removal of c. 15% of leaf area led to a significant reduction in plant performance compared with undamaged controls, but only when the plants were grown together in competition for light. The second experiment demonstrated that the distribution of grazing damage within the plant was an important factor in the outcome of competition; in those plants in which grazing was applied to the lower leaves there was no effect of damage upon performance compared with undamaged controls, whereas grazing to the upper leaves significantly reduced plant performance. A third experiment provided some insight into how this interaction between damage and competition comes about. It was shown that damage to leaves led to a rapid drop in the rate of extension growth of the main shoot, especially when the upper leaves were damaged, and normal rates of growth were not resumed for at least 3 days. It is argued that in a rapidly growing canopy, such an effect may mean that a damaged plant loses its position in the height hierarchy. The final experiment showed that previous damage to plants can affect the distribution of subsequent grazing by larvae of Spodoptera littoralis, apparently through a wound-induced reduction in leaf palatability. Plants which had been artificially damaged 48 h previously were grazed significantly less than controls, and the avoidance effect was greatest in the young leaves. These results are consistent with the grazing dispersal hypothesis, and suggest that rapid wound-induced responses may be of greatest significance in species characteristic of fertile environments where competition for light is particularly intense.     

Foraging ecology of roan antelope: key resources during critical periods

The use of landscape zones and grass species by roan antelope, a species threatened with local extirpation within South Africa's Kruger National Park, were investigated. Plant‐based observations of grazing were made within a 300 ha enclosure in the roan range, where 40 roan antelope were confined at high density in the absence of other grazers. The study spanned the dry seasons of two years, one with average rainfall and one with low rainfall. We recorded changes in the extent of grazing of different grass species, height differences between grazed and ungrazed tillers and intensity of cropping per tuft. In the average year, the grazing pressure in the bottomland grassland was twice that in the upland savanna, with two tall grass species bearing the brunt of the grazing through the dry season. Two highly palatable upland grasses were also extensively grazed by the mid dry season. In the dry year, the extent of grazing in the upland exceeded that in the bottomland, and several upland grass species little used the previous year became heavily grazed. Roan antelope appeared to be separated ecologically from more common grazers by their selective use of tall grasses growing in the drainage line grassland during the critical dry season months. However, their grazing expansion into the upland savanna during the dry year potentially brought them into competition with these grazers. Nevertheless, their population performance did not suffer despite the high‐density conditions. Heightened predation pressure following an influx of these grazers, rather than resource limitation, appeared to be primarily responsible for the drastic decline of this species in the park.     

Ecology of grazing lawns in Africa

Grazing lawns are a distinct grassland community type, characterised by short‐stature and with their persistence and spread promoted by grazing. In Africa, they reveal a long co‐evolutionary history of grasses and large mammal grazers. The attractiveness to grazers of a low‐biomass sward lies in the relatively high quality of forage, largely due to the low proportion of stem material in the sward; this encourages repeat grazing that concomitantly suppresses tall‐grass growth forms that would otherwise outcompete lawn species for light. Regular grazing that prevents shading and maintains sward quality is thus the cornerstone of grazing lawn dynamics. The strong interplay between abiotic conditions and disturbance factors, which are central to grazing lawn existence, can also cause these systems to be highly dynamic. Here we identify differences in growth form among grazing lawn grass species, and assess how compositional differences among lawn types, as well as environmental variables, influence their maintenance requirements (i.e. grazing frequency) and vulnerability to degradation. We also make a clear distinction between the processes of lawn establishment and lawn maintenance. Rainfall, soil nutrient status, grazer community composition and fire regime have strong and interactive influences on both processes. However, factors that concentrate grazing pressure (e.g. nutrient hotspots and sodic sites) have more bearing on where lawns establish. Similarly, we discuss the relevance of enhanced rates of nitrogen cycling and of sodium levels to lawn maintenance. Grazer community composition and density has considerable significance to grazing lawn dynamics; not all grazers are adapted to foraging on short‐grass swards, and differences in body size and relative mouth dimensions determine which species are able to convert tall‐grass swards into grazing lawns under different conditions. Hence, we evaluate the roles of different grazers in lawn dynamics, as well as the benefits that grazer populations derive from having access to grazing lawns. The effects of grazing lawns can extend well beyond their borders, due to their influence on grazer densities, behaviour and movements as well as fire spread, intensity and frequency. Variation in the area and proportion of a landscape that is grazing lawn can thus have a profound impact on system dynamics. We provide a conceptual model that summarises grazing lawn dynamics, and identify a rainfall range where we predict grazing lawns to be most prevalent. We also examine the biodiversity associated with grazing lawn systems, and consider their functional contribution to the conservation of this biodiversity. Finally, we assess the utility of grazing lawns as a resource in a rangeland context.     

Palatability factors and nutritive values of the food of buffaloes (Syncerus caffer) in Uganda

The food preferences of tame buffaloes grazing different vegetation types were recorded. The effects of fire and rainfall were investigated. By initiating a new growth cycle, fire altered the food habits of large herbivores and also the distribution of small mammals. Seasonal variations in the effect of rainfall on plant growth explains why preferences did not change in proportion to precipitation. The more important grasses were collected and their parts analysed physically and chemically. Their stem to leaf ratio is considered to have influenced the species eaten by coarse grazers such as buffalo. A broad correlation was obtained between buffalo preferences and the percentage green leaf of Hyparrhenia filipendula and Themeda triandra. This correlation was improved when preferences were compared with the number of grams green leaf carbohydrate per 100 g dry grass. There was a negative response to the silica present in dead leaves, but stem fibre appeared to have little effect on preferences. A grazing animal experiences a combination of attractants and repellants, i.e. green leaf carbohydrate and protein less stem fibre and dead leaf silica. Other palatability factors were noted, in particular tannins and oils in grasses of the tribe Andropogoneae. These may act as repellants, possibly only when the grazing animal has alternative vegetation from which to choose. The mineral content of grasses appeared to be adequate and calcium and phosphorus were present in about the required proportions. The observations support the belief that selective grazing enables a herbivore to consume a diet of significantly higher nutritive value than that of the average sward. Browsing and new growth after fire help compensate for the low nutritive value of pasture during the dry season. The nutritive value of the diet is considered to be the proximal factor in the seasonal cycle of reproduction of the buffalo and probably the ultimate factor in the reproduction of certain other ungulates and rodents.     

Grass competition is more important than seed ingestion by livestock for Acacia recruitment in South Africa

Although grazing livestock may have direct negative effects on woody species through herbivory and trampling, (heavy-)grazing is often associated with woody plant encroachment. Two main mechanisms can explain the positive effects of grazers on woody cover: (1) Grazers reduce the interspecific competition with trees and may reduce fuel load, and (2) gut passage through livestock increases seedling establishment by scarification (increased germination) and fertilization by dung (higher survival and growth of seedlings). We tested the effects of fire, grass, cattle ingestion (transit) and dung on germination, seedling height and survival as well as on recruitment of Acacia sieberiana in a sub-humid grassland of South Africa. About 8,000 seeds were planted in the field in a randomized block design. The removal of grass by grazing and/or fire had the most important effect on Acacia recruitment in savanna. Our findings highlight the hierarchy of the main factors affecting Acacia recruitment, which provides new insights to the understanding of woody plant encroachment.     

Grazers and soil moisture determine the fate of added 15NH4 + in Yellowstone grasslands

The influence of ungulate grazers on nutrient cycling and ecosystem productivity in grasslands has been shown to differ with moisture, nutrient availability, and feedbacks between above- and belowground activities. We examined the movement of nitrogen (N), applied as (¹⁵NH₄)₂SO₄, through both dry and mesic sites in the northern range of Yellowstone National Park to test the hypothesis that plants were more able to acquire added N in grazed relative to ungrazed sites. Previous studies showed enhanced N mineralization in grazed areas, and detritus removal by grazers was predicted to enhance early-season plant growth. Thirteen months after tracer addition, there were no differences in plant ¹⁵N as a function of grazing, but historically ungrazed sites retained more ¹⁵N in accumulated litter than at grazed sites. This result demonstrated the importance of detritus in regulating redistribution of incoming N and the role of grazers in this process. Site moisture status influenced ¹⁵N recovery in all pools—soils, microbial biomass, and plants—and greater plant ¹⁵N acquisition occurred in roots at dry relative to mesic sites. Understanding how grazers influence nutrient cycling at the landscape scale requires further investigation of interactions among soil moisture, plant production, litter accumulation, grazing intensity, and belowground processes.     

Interactions between local climate and grazing determine the population dynamics of the small herb Viola biflora

Plants of low stature may benefit from the presence of large herbivores through removal of tall competitive neighbours and increased light availability. Accordingly, removal of grazers has been predicted to disfavour small species. In addition to this indirect beneficial effect, the population dynamics of plants is strongly influenced by variation in external conditions such as temperature and precipitation. However, few studies have examined the interaction between large herbivores and inter-annual variation in climate for the population dynamics of small plant species not preferred by herbivores. We studied three populations of the perennial herb Viola biflora exposed to different sheep densities (high, low and zero) for 6 years in a field experiment. Plants were also impacted by invertebrate and small vertebrate herbivores (rodents). Rates of growth were marginally higher at high sheep densities, and during warm summers both survival and growth were higher when sheep were present. Thus, while the height of tall herbs was positively related to July temperature, it was less so in the treatments with sheep, suggesting that sheep reduce the negative effects of interspecific competition for this small herb. Life table response experiment analyses revealed that the population growth rate (λ) was slightly lower in the absence of sheep, but between-year variation in λ was larger than variation among sheep density treatments. λ was negatively related to July temperature, with an additional negative effect of vertebrate grazing frequency (sheep or rodent grazing). The evidence from this 6-year study suggests that the population dynamics of Viola biflora is determined by a complex interplay between climate and grazing by both large and small herbivores.     

Diet Switching by Mammalian Herbivores in Response to Exotic Grass Invasion

Invasion by exotic grasses is a severe threat to the integrity of grassland ecosystems all over the world. Because grasslands are typically grazed by livestock and wildlife, the invasion is a community process modulated by herbivory. We hypothesized that the invasion of native South American grasslands by Eragrostis plana Nees, an exotic tussock-forming grass from Africa, could be deterred by grazing if grazers switched dietary preferences and included the invasive grass as a large proportion of their diets. Bos taurus (heifers) and Ovis aries (ewes) grazed plots with varying degrees of invasion by E. plana in a replicated manipulative experiment. Animal positions and species grazed were observed every minute in 45-min grazing session. Proportion of bites and steps in and out of E. plana tussocks were measured and used to calculate several indices of selectivity. Both heifers and ewes exhibited increasing probability of grazing E. plana as the proportion of area covered by tussocks increased, but they behaved differently. In agreement with expectations based on the allometry of dietary preferences and morphology, ewes consumed a low proportion of E. plana, except in areas that had more than 90% E. plana cover. Heifers consumed proportionally more E. plana than ewes. Contrary to our hypothesis, herbivores did not exhibit dietary switching towards the invasive grass. Moreover, they exhibited avoidance of the invasive grass and preference for short-statured native species, both of which should tend to enhance invasion. Unless invasive plants are highly palatable to livestock, the effect of grazing to deter the invasion is limited, due to the inherent avoidance of the invasive grass by the main grazers in the ecosystem, particularly sheep.     

Pre-nesting feeding selectivity of Pink-footed Geese Anser brachyrhynchus in artificial grasslands

The pre-nesting feeding ecology of Pink-footed Geese Anser brachyrhynchus was studied in southern Iceland in April and May 1989–1992. Extensive surveys showed that prior to movement to their nesting areas in the interior, Pink-footed Geese fed mainly on intensively managed grasslands of the southern lowlands. Faecal analysis showed that geese feeding in hayfields foraged almost exclusively on the most commonly reseeded species, Timothy grass Phleum pratense , which made up the majority (41% by number of green shoots) of new growth in studied hayfields. Dropping counts, as an indicator of goose use, were highly significantly correlated in all years with density of growing Phleum shoots in individual field units. Geese showed a response in feeding patterns to changes in Phleum density as a result of reseeding patterns. Analysis of forage quality in 1991 showed that Phleum shoots had a higher protein content than all other grasses present in hayfields, with the exception of Deschampsia caespitosa. The latter species showed similar levels of protein, but its tussocky nature precluded effective grazing by the geese. Phleum was similar in fibre content to other common grasses. Based on removal of tagged plants, geese grazed predominantly on the youngest leaves of Phleum , which also had higher protein content and lower fibre than older leaves and attached dead leaves, which were rarely taken by geese. Geese grazed shorter length classes of youngest leaves, which showed higher protein content than longer, older terminal leaves. Recently created grassland habitat has increased the opportunities for female geese to supplement their reserves during the crucial prelude to clutch initiation by selecting the highest quality plant species and the most nutritious parts of that forage.     

Do submerged aquatic plants influence their periphyton to enhance the growth and reproduction of invertebrate mutualists?

It has been suggested that submerged aquatic plants can influence the nutritional quality of the periphyton which grows on their surfaces, making it more nutritious for grazing invertebrates, particularly snails. In return, these grazers might preferentially feed on the periphyton and clear the plants of a potential competitor, with the plants and grazers both gaining from this mutualistic relationship. A highly replicated experiment was conducted, in which the nature of the plant (isoetid and elodeid types compared with similar-shaped inert substrata), the nutrient loading, and the influence of periphyton grazers (the bladder snail, Physa fontinalis) of similar size and history were controlled. Plant growth and survival significantly increased in the presence of the periphyton grazer. Whilst the presence of the grazers had the largest influence on periphyton abundance, nutrient availability and plant type also had effects. Plant type had little influence on the nutritional quality of the periphyton measured as carbohydrate, protein and C:N. Effects of treatment on snail growth, and the timing and extent of snail reproduction disappeared when they were compared with the quantity of periphyton available. There was no evidence of enhanced grazer success in the presence of the live plants compared with inert substrata. Although submerged plants affect the growth and reproduction of the grazers which feed on their surfaces, through differences in the amount of periphyton which grows there, we found no evidence that they manipulate the periphyton to encourage such grazers. Received: 1 September 1998 / Accepted: 12 May 1999     

Effect of defoliation intensity on aboveground and belowground relative growth rates

According to a simple growth model, grazed and ungrazed plants may have equal absolute growth rates provided that the relative growth rate (RGR) of grazed plants increases exponentially with grazing intensity (proportion of biomass removed). This paper reports results from an experiment designed to determine whether plants of two grass species subjected to a gradient of defoliation intensities, from 0 to 100% aboveground biomass removal, showed such a response. The relationship between aboveground RGR and defoliation intensity was exponential and closely matched the theoretical relationship of equal absolute growth rate. Thus, plants showed the same aboveground growth regardless of defoliation intensity thanks to an exponential stimulation of RGR by defoliation. Belowground RGR was depressed by defoliation of more than 20% of the above-ground biomass. In spite of the drastic modification imposed by the treatments on the relative proportions of different plant parts, after a 42-day recovery period basic allometric relationships, such as root:shoot and leafarea: weight ratios, were not affected by defoliation intensity. Exponential aboveground compensatory responses represent a key feedback process resulting in constant aboveground growth regardless of defoliation intensity and appear to be a simple consequence of strong commitments to certain allometric relationships.     

Major consequences of minor damage: impacts of small grazers on fast-growing kelps

Damage by small herbivores can have disproportionately large effects on the fitness of individual plants if damage is concentrated on valuable tissues or on select individuals within a population. In marine systems, the impact of tissue loss on the growth rates of habitat-forming algae is poorly understood. We quantified the grazing damage by an isopod Amphoroidea typa on two species of large kelps, Lessonia spicata and Macrocystis pyrifera, in temperate Chile to test whether non-lethal grazing damage could reduce kelp growth rates and photosynthetic efficiency. For L. spicata, grazing damage was widespread in the field, unevenly distributed on several spatial scales (among individuals and among tissue types) and negatively correlated with blade growth rates. In field experiments, feeding by A. typa reduced the concentration of photosynthetic pigments and led to large reductions (~80 %) in blade growth rates despite limited loss of kelp biomass (0.5 % per day). For M. pyrifera, rates of damage in the field were lower and high densities of grazers were unable to reduce growth rates in field experiments. These results demonstrate that even low per capita grazing rates can result in large reductions in the growth of a kelp, due the spatial clustering of herbivores in the field and the selective removal of photosynthetically active tissues. The impacts of small herbivores on plant performance are thus not easily predicted from consumption rates or abundance in the field, and vary with plant species due to variation in their ability to compensate for damage.     

Tree size and herbivory determine below-canopy grass quality and species composition in savannahs

Large single-standing trees are rapidly declining in savannahs, ecosystems supporting a high diversity of large herbivorous mammals. Savannah trees are important as they support both a unique flora and fauna. The herbaceous layer in particular responds to the structural and functional properties of a tree. As shrubland expands stem thickening occurs and large trees are replaced by smaller trees. Here we examine whether small trees are as effective in providing advantages for grasses growing beneath their crowns as large trees are. The role of herbivory in this positive tree-grass interaction is also investigated. We assessed soil and grass nutrient content, structural properties, and herbaceous species composition beneath trees of three size classes and under two grazing regimes in a South African savannah. We found that grass leaf content (N and P) beneath the crowns of particularly large (ca. 3.5 m) and very large trees (ca. 9 m) was as much as 40% greater than the same grass species not growing under a tree canopy, whereas nutrient contents of grasses did not differ beneath small trees (<2.3 m). Moderate herbivory enhanced these effects slightly. Grass species composition differed beneath and beyond the tree canopy but not between tree size classes. As large trees significantly improve the grass nutrient quality for grazers in contrast to smaller trees, the decline of the former should be halted. The presence of trees further increases grass species diversity and patchiness by favouring shade-tolerant species. Both grazing wildlife and livestock will benefit from the presence of large trees because of their structural and functional importance for savannahs.     

Defoliation of a grass is mediated by the positive effect of dung deposition,moss removal and enhanced soil nutrient contents: results from a reindeer grazing simulation experiment

Herbivory is one of the key drivers shaping plant community dynamics. Herbivores can strongly influence plant productivity directly through defoliation and the return of nutrients in the form of dung and urine, but also indirectly by reducing the abundance of neighbouring plants and inducing changes in soil processes. However, the relative importance of these processes is poorly understood. We, therefore, established a common garden experiment to study plant responses to defoliation, dung addition, moss cover, and the soil legacy of reindeer grazing. We used an arctic tundra grazed by reindeer as our study system, and Festuca ovina, a common grazing‐tolerant grass species as the model species. The soil legacy of reindeer grazing had the strongest effect on plants, and resulted in higher growth in soils originating from previously heavily‐grazed sites. Defoliation also had a strong effect and reduced shoot and root growth and nutrient uptake. Plants did not fully compensate for the tissue lost due to defoliation, even when nutrient availability was high. In contrast, defoliation enhanced plant nitrogen concentrations. Dung addition increased plant production, nitrogen concentrations and nutrient uptake, although the effect was fairly small. Mosses also had a positive effect on aboveground plant production as long as the plants were not defoliated. The presence of a thick moss layer reduced plant growth following defoliation. This study demonstrates that grasses, even though they suffer from defoliation, can tolerate high densities of herbivores when all aspects of herbivores on ecosystems are taken into account. Our results further show that the positive effect of herbivores on plant growth via changes in soil properties is essential for plants to cope with a high grazing pressure. The strong effect of the soil legacy of reindeer grazing reveals that herbivores can have long‐lasting effects on plant productivity and ecosystem functioning after grazing has ceased.     

Interactive effect of flooding and grazing on the growth of Serengeti grasses

Summary Grazing and flooding may potentially interact in particular habitats of many grassland regions around the world. We tested the hypothesis that grazing and flooding induce different and largely opposed allocation responses in individual plants. As a result, their combined effect on plant growth would be negative. We studied the response of three grass species from the Serengeti ecosystem (Tanzania) to the effects of flooding and clipping. Plants under the combined effect of flooding and clipping had lower growth rates than plants growing under the effect of either of the two factors individually. Plants under flooding grew taller and allocated more resources to stem growth than controls; for two of the three species, flooded plants also generated a new root system above soil level. All these morphological and physiological responses conflict with the ability of a plant to respond to defoliation with minimum reduction in growth rates. The three species showed a response to flooding reflecting their distribution ranges in the field: the species from the most flood-prone habitat showed a positive effect of flooding on growth, whereas the species from dry uplands showed a strong negative effect of flooding. Flood-tolerant species were taller and less tolerant of clipping than flooding sensitive species. Our results suggest that, in ecological time, individuals subjected to both flooding and grazing have their growth reduced to a greater extent than by either of the two factors acting individually, whereas in evolutionary time, species adapted to flooding are poor grazing tolerators and species adapted to grazing are poor flooding tolerators.     

Herbivore-induced expansion of <Emphasis Type="Italic">Helianthemum nummularium</Emphasis> in grassland–scrub mosaic vegetation: circumstantial evidence for zoochory and indirect grazing impact

Extensive grazing often has a strong influence on the structure and composition of herbaceous plant communities with increasing population sizes for some species and decreasing presence in others. Herbivores affect plant communities directly by selective grazing of plant species, and indirectly by either epizoochory or endozoochory. Helianthemum nummularium is considered an increasing species because its distribution increased after the introduction of large, free-ranging grazers in at least two coastal dune grassland areas in Belgium. However, its seeds lack any obvious adaptations for epizoochory, and direct observations of plant/seed consumption are scarce. Through field and lab experiments, we assessed the dispersal ability of H. nummularium via endozoochory and epizoochory. In a differentiated grazer exclusion experiment, evidence was found that plants are grazed by large domestic ungulates and small wild herbivores although these incidences were rare. Direct endozoochory evidence remained scarce. No seeds were found germinating in field-collected dung, and only few seedlings emerged following a seed feeding experiment. However, once deposited, we found higher growth rates when seeds were mixed with dung and decreased establishment success when seeds were sown in combination with competitively superior species. Epizoochory was plausible because both fur and hooves of cattle and horses were potentially capable of contributing to the transport of H. nummularium seeds. We conclude that herbivores play a role in seed dispersal, while their selective grazing behaviour most probably creates an appropriate environment for Helianthemum establishment and maintenance.     

Palatability of macrophytes to the invasive freshwater snail Pomacea canaliculata: differential effects of multiple plant traits

1. By selective grazing, invasive grazers can alter macrophyte‐herbivore relationships in shallow freshwater bodies. Evaluating the palatability of macrophytes and understanding the determinants of plant palatability can help predict grazing impact. In no‐choice feeding assays, we tested the palatability of 21 species of freshwater macrophytes to the invasive freshwater apple snail Pomacea canaliculata. 2. Daily feeding rate varied greatly with plant species, ranging from 1.1 to 22% of snail body mass. We assessed six plant properties and examined their correlation with feeding rate. Total nitrogen content was positively related, and C:N ratio and dry matter content (DMC) negatively related, to snail feeding rate. There was no significant correlation between snail feeding rate and plant phenolic content, but the feeding rate on Myriophyllum aquaticum (the plant with the highest phenolic content) was very low. 3. We repeated the feeding assays for 15 species that were not palatable as fresh leaves with reconstituted plant tissues formed by mixing ground up dried leaves with agar. The feeding rate still differed greatly among macrophyte species. Phragmites australis and Vallisneria natans (two species with the highest DMC) were eaten much more as reconstituted plant than as fresh leaves, indicating that structure (i.e. DMC) may be important in their defence against snail herbivory. For two plants (M. aquaticum and Alternanthera philoxeroides) that had moderate amounts of nitrogen/phosphorus but were consumed very little as fresh and reconstituted tissues, we incorporated their extracts into a palatable agar‐based food. The extracts from both species greatly reduced snail feeding rate, indicating the presence of chemical defences in these two species. 4. These results indicated that feeding was affected by several plant traits. The snail favoured plants with a high nitrogen content and avoided plants with a high DMC. Only a few plants possessed chemical feeding deterrents that were effective against this snail. Given the invasive spread of P. canaliculata in Asia, ecologists and managers should consider plant palatability when selecting plants for use in wetland restoration and when predicting the impact of further invasion by this species.     

The effects of grazing by gastropods and physical factors on the upper limits of distribution of intertidal macroalgae

Summary The cover of foliose algae is sparse to non-existent above a low-level algal zone on many shores in N.S.W., except in rock-pools. Above this algal zone, encrusting algae, mostly Hildenbrandia prototypus, occupy most of the primary substratum on sheltered shores. Experimental manipulations at midtidal levels were used to test hypotheses about the effects of grazing by molluses and of physical factors during low tide on this pattern of algal community structure.Fences and cages were used to exclude grazers: molluscs grazed under roofs and in open areas. Cages and roofs provided shade, and decreased the harshness of the environment during low tide: fences and open areas had the normal environmental regime.In the absence of grazers, rapid colonization of Ulva and slower colonization by other foliose algae occurred in all experimental areas. The rate of colonization by Ulva sporelings was initially retarded on existing encrusting algae, but after a few months, cover of Ulva equalled that on cleared rock.Most species of algae only grew to maturity inside cages, and remained as a turf of sporelings inside fences. No foliose algae grew to a visible size in open, grazed areas. Grazing thus prevents the establishment of foliose algae above their normal upper limit on the shore, but the effects of physical factors during low tide prevent the growth of algae which become established when grazers are removed. Physical factors thus limit the abundance of foliose algae at mid-tidal levels.The recolonization of cleared areas by Hildenbrandia was not affected by the presence of a turf of sporelings, nor by the shade cast by roofs, but was retarded in cages where mature algae formed a canopy. Even under such a canopy, Hildenbrandia eventually covered as much primary substratum as in open, grazed areas. This encrusting alga is able to escape from the effects of grazing by having a tough thallus, and by its vegetative growth which allows individual plants to cover a lot of substratum, and by the tendency for new individuals to start growing from small cracks and pits in the rock, which are apparently inaccessible to the grazers.Mature foliose algae are removed from the substratum by waves, and many individual plants died during periods of hot weather. Sporelings in a turf were eliminated, after experimental fences were removed, by the combined effects of macroalgal grazers, which invaded the areas, and microalgal grarers which ate the turt from the edges inwards.The results obtained here are discussed with respect to other studies on limits to distribution of intertidal macroalgae, and the role of grazing in the diversity and structure of intertidal algal communities. Some problems of these experimental treatments are also discussed.