Impact of different sheep grazing intensities on salt marsh vegetation in northern Germany

Abstract. Mainland salt marshes in Schleswig-Holstein (northern Germany) have been grazed intensively by sheep for several decades. In 1988 experimental sites were established in the lower and middle salt marsh of Sönke-Nissen-Koog and subjected to different grazing intensities. From 1989 to 1993 the impact of sheep grazing on the composition and structure of the salt marsh vegetation was studied through the yearly analysis of permanent plots, vegetation mapping and measurements of the vegetation height. The intensively grazed site (10 sheep/ha) is covered by a short monotonous Puccinellia maritima sward with Salicornia europaea and Suaeda maritima. Halimione portulacoides and Aster tripolium, especially flowering plants, are rare. On the sites with 1.5 and 3 sheep/ha Puccinellia maritima remained dominant. The population density of Salicornia europaea decreases after reduction of the grazing intensity, whereas Suaeda maritima finds optimal growing conditions. Stands of Halimione portulacoides and flowering Aster tripolium plants are rare near the sea dike but their cover and size increases further away from the dike. In the plot with 1.5 sheep/ha the height of the vegetation increases along the gradient from the dike towards the tidal flats, due to local differences in actual grazing intensity. When grazing is stopped, Puccinellia maritima is successively replaced by Festuca rubra, Halimione portulacoides and Aster tripolium. High variability of vegetation height indicates structural diversity. Patches of higher and lower vegetation correspond with the distribution pattern of different plant species. In terms of nature conservation cessation of grazing is recommended.     

Salt marshes along the coast of The Netherlands

The area of salt marshes does no longer increase. The recent erosion coincides with a rise in MHT-level in the last 25 years. Despite the decrease in area, sedimentation continues, especially in the lower salt marsh, which acts as a sink of nitrogen. Assimilation and mineralization of nitrogen are in balance in most plant communities along the gradient from lower to higher salt marshes. Mineralization of nitrogen increases towards the higher salt marsh, whereas the above-ground production and the mean nitrogen content of plants decrease. There is a positive correlation between quality of food plants in salt marshes and breeding success of Brent geese in the arctic tundra. Sedimentation on mainland salt marshes can compensate for the expected sea level rise. This is not the case for island salt marshes, if the relative sea level rise is more than 0.5–1.0 cm yr⁻¹. The natural succession on salt marshes results in an accumulation of organic material, which is related to the dominance of single plant species. It is not clear to which extent this process is enhanced by eutrophication from acid deposition and seawater. Human exploitation of unprotected salt marshes is old and heavy in the system of mound settlements. Reclamation rates by dikes in the last centuries were higher than the rate of area increase. Grazing by cattle as a management practice results in both a higher plant species-richness and community diversity than abandoning; hay-making is intermediate, but shows less structural diversity than grazing with low stocking density. The invertebrate fauna is favoured by a short period of abandoning, but eventually characteristic salt marsh invertebrates are replaced by inland species. Many bird species prefer grazed salt marshes. The final section gives some perspectives. Provided that no further embankments take place the optimal nature management option for plants and animals is a vegetation pattern, which includes areas with a low canopy (grazed) and areas with a tall canopy.     

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

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

Effects of stock grazing on the ground invertebrate fauna of woodland remnants

Habitat fragmentation can leave formerly widespread habitat types represented by only small habitat ‘islands’, and the conservation of these remnants is frequently compromised by ongoing disturbance. In northern Victoria, grazing of woodland remnants by sheep and cattle has profound effects on the vegetation structure of the woodland by removing understorey and ground vegetation. To investigate the effects of grazing pressure on remnant grey box Eucalyptus microcarpa woodland in northern Victoria, we surveyed the ground invertebrate fauna in ungrazed woodland remnants, grazed woodland remnants, and grazed pasture. The number of invertebrates caught increased from ungrazed woodland to grazed woodland to pasture, but this increase was due primarily to the most abundant orders (Hymenoptera, Coleoptera and Aranaea), and two abundant taxa characteristic of pasture (Orthoptera and Dermaptera). In contrast, most of the less abundant orders followed the opposite pattern, and were caught in higher numbers (and as a higher proportion of the total catch) in ungrazed woodland. Ungrazed woodland had a more diverse ground invertebrate fauna, most likely due to the greater diversity of food and habitat resources provided by the less disturbed vegetation. The differences in invertebrate communities corresponded to differences in vegetation and litter layers. The reduction in biodiversity of remnants due to grazing has implications for conservation management of remnant woodland in agricultural landscapes.     

Alteration of salt marsh plant community composition by grazing snow geese

A notable omission from wetlands ecology has been the study of the influence of herbivores on vegetation. Reported here are the effects of grazing by snow geese Anser caerulescens altantica on the vegetation of salt marshes along the mid-Atlantic coast of the United States, Exclosures were used to compare total and species percent cover between grazed and ungrazed areas in three marshes (Salt Flats, South Pond, and Bodie Island) with differing vegetation communities from 1978 to 1980.
Spartina alterniflora was reduced by ⅔ in grazed versus ungrazed areas of Salt Flats, S. S. patens was reduced by ½ in grazed portions of South Pond but recovered when grazing ceased. In grazed portions of Bodie Island total plant cover was reduced by 16%. Scirpus robustus and S. patens reacted in opposite ways to grazing pressure with S. robustus increasing and S. patens decreasing, Elocharis was found only in grazed areas of Bodie Island. Echinochloa crusgali appeared in grazed portions of this marsh in 1978 but decreased in abundance during subsequent years. Scirpus americanus was unaffected by grazing, maintaining a nearly constant percent cover in grazed and ungrazed areas at Bodie Island.
Differences in responses to grazing are discussed in terms of each species' growth and reproductive strategies as tempered by the physical and biological environment within each marsh.     

Response of galling invertebrates on Salix lanata to reindeer herbivory

Browsing and defoliation often increase the densities of insect herbivores on woody plants. Densities of herbivorous invertebrates were estimated in a long-term grazing manipulation experiment. More then 30-yr-old fences allow us to compare densities of invertebrate herbivores on Salix lanata in areas heavily grazed and areas lightly grazed by reindeer. The number of gall-forming insects ( Pontania glabrifons) and gall-forming mites were higher on the heavily grazed shrubs than on lightly grazed shrubs. In contrast to most short-term studies, the heavily grazed S. lanata had shorter current annual shoots. No difference in leaf size, leaf nitrogen content, or C:N ratio between grazing intensities were detected. However, the enhanced natural δ15N value indicates that heavily grazed shrubs get a higher proportion of their N directly from reindeer faeces. Leaf weight per unit area and relative fluctuating asymmetry of leaf shape increased in heavily grazed S. lanata . Enhanced relative fluctuating asymmetry might indicate higher susceptibility to herbivores. Long-term grazing seems to increase the density of invertebrate herbivory in the same way as short-term grazing, even if the plant responses differ substantially.     

Long-term changes of salt marsh communities by cattle grazing

Over a period of 9 years a grazing experiment was carried out in the mainland salt marsh of the Leybucht (Niedersachsen) with three stocking rates, namely, 0.5 ha^-1, 1 ha^-1, and 2 cattle ha^-1. These were also compared with an abandoned area. The results are based on sampling of the invertebrates in 1980, 1981, 1982, and 1988, and of the vegetation in 1980 and 1988. The rate of sedimentation is highest in the Puccinellia maritima-zone and decreases with the increase of stocking rates. The Elymus pycnanthus vegetation type becomes dominant in the higher salt marsh in the abandoned site. The canopy height decreases with increasing stocking rate, whereas a gradient in the structure of the vegetation develops with the lowest stocking rate. The population densities, the species-richness and the community diversity of invertebrates increases after the cessation of grazing. The high rate of sedimentation in the abandoned site promotes the immigration of species from higher salt marsh levels and adjacent grasslands, and eventually halotopophilous species and communities may disappear. On the other hand grazing reduces numerous species living both in or on upper parts of the vegetation or being sensitive to trampling by cattle. The community structure shows that the salt marsh ecosystem changed from a food web dominated by plant feeding animals to a food web dominated by animals foraging on detritus. The salt marsh management has to be differentiated into both ungrazed and lightly grazed areas (each 50%) or an overall grazing in large areas with less than 0.5 cattle ha^-1.     

The Impact of Sheep Grazing on Net Nitrogen Mineralization Rate in Two Temperate Salt Marshes

Abstract: Nitrogen mineralization rate was studied in grazing trials with three different stocking rates (0, 3, 10 sheep ha^-1) in two man-made salt marshes, viz. a Puccinellia maritima -dominated low salt marsh and a high salt marsh dominated by Festuca rubra. Mineralization rates were derived from the amounts of mineral N which accumulated in situ during six-week incubation periods in tubes containing undisturbed soil cores from the upper 10 cm soil layer. The annual rates of net N mineralization were significantly higher in the better drained, high salt marsh (71 - 81 kg ha^-1 yr^-1) than in the low salt marsh (39 - 49 kg ha^-1 yr^-1). High amounts of belowground litter accumulated in the low salt marsh due to frequent water logging. Both N mineralization and nitrification rate were negatively correlated with soil water content. In the Puccinellia maritima salt marsh, grazing had neither an effect on N mineralization rates during any of the incubation periods nor on annual mineralization rates. In the Festuca rubra salt marsh, N mineralization rates increased earlier during spring at the intensively grazed site than at the moderately grazed and the ungrazed site. N mineralization and nitrification rates were significantly higher at the ungrazed site than at the intensively grazed site during the period of peak net N mineralization from the end of April until mid-June. Although sheep grazing affected the seasonal pattern of N mineralization in the high marsh, grazing did not affect the annual rate of net N mineralization.     

Field studies on the variability of populations of Aster tripolium L. in relation to saltmarsh zonation

1. Aster tripolium is a very variable species of which a number of types have been described both on a morphological and ecological level.
2. In permanent plots along the height gradient in the salt marsh it appeared that differences in the A. tripolium subpopulations occur.
3. In the lowest zone of the marsh, dominated by Spartina anglica (Spartinion), the mean life expectancy of individual adult plants is relatively high but it is relatively low for seedlings.
4. In the higher zones of the marsh the mean life expectancy is relatively lower, but for the seedlings it is higher (Puccinellion maritimae).
5. In the Spartina-zone individual A. tripolium plants have about twice the number of shoots as the plants from the other zones.
6. The plants from the Spartina zone produce per individual more generative shoots (absolute) but these have less capitula than elsewhere in the marsh. The number of ripe seeds per head is almost constant everywhere in the marsh.

     

Interactions between hare and brent goose in a salt marsh system; evidence for food competition?

In this study we accumulate evidence that brown hare competes with brent goose for food resources in a temperate salt marsh. We show that both species overlap in habitat use and share food plants. The two herbivores mainly used the common habitat at different times of the day, with hares active in the dark and geese during the daylight. During the morning and evening, however, the habitat was exploited simultaneously. Food availability was manipulated by excluding brent geese on both small-scale (30 m²) and large-scale (0.96 ha) plots, while hares had free access everywhere. Exclusion of brent geese enhanced the level of utilisation by hares in both Festuca and Puccinellia dominated marshes, which are among the most intensively grazed parts of the salt marsh. The increase in hare grazing pressure following goose exclusion was stronger, when the adjacent control plots had attracted more goose visitation. When geese were excluded, the decrease in Festuca consumption by geese was completely matched by increased hare grazing, while for Puccinellia only part of the `surplus' was harvested. Enhanced levels of hare utilisation were not due to geese interfering directly with hare, nor due to hares avoiding goose droppings. Considering the interaction from the other perspective, hares were observed to disturb geese effectively in every spring. This might have reduced exploitation by geese of the shared resources. On the basis of our experimental results, we conclude that in this salt- marsh system competition for food with brent geese plays a role in the habitat use of hares, and that hares can reduce goose exploitation of shared habitats. Received: 30 March 1998 / Accepted: 6 July 1998     

Production estimates of the benthic invertebrate community in a small Danish stream

Quantitative samples were used to investigate density, biomass and annual production of the benthic invertebrate fauna in a small Danish stream. Forty-eight taxa were found and the total invertebrate densities varied from 3 810 m^?2 in July to 20 040 m^?2 in December. The total mean annual biomass of the invertebrate fauna was 6.1 g ash-free dry wt m^?2. The annual production of the invertebrates was estimated from their mean annual biomass and their annual P/B ratio. Production of the primary consumers (herbivores and detritivores) was 21.4 g ash-free dry wt m^?2 y^?1 and of secondary consumers (carnivores) 1.1 g m^?2 y^?1. The amount of invertebrate production available to the trout population and the importance of the species as food for trout are discussed.     

The effect of cattle and sheep grazing on salt-marsh vegetation at Skallingen,Denmark

The aggregated effect of cattle and sheep grazing on Puccinellion maritimae and other salt-marsh vegetation has been studied together with changes in species composition, the percentage cover of each species, total cover and the percentage of bare ground, six years after grazing had been prevented by construction of experimental exclosures. The results of these experiments are discussed in relation to the natural development of the vegetation that occurred in a permanent plot in the ungrazed part of the salt marsh. Six years without grazing caused a marked increase in total cover and a dramatic reduction in the amount of bare ground. The species composition of the Puccinellia maritima community did not change during these six years, but all species increased in cover. Whereas Artemisia maritima migrated into the Festuca rubra community and only two species Festuca and Halimione portulacoides, increased in cover, all other species showed reduced cover and Salicornia europaea disappeared from the plot after six years without grazing. During the same period of time, three species, Salicornia europaea, Suaeda maritima, and Glaux maritima, disappeared from the plot in the ungrazed marsh as a result of natural development. During thirty-five years the vegetation originally dominated by Puccinellia maritima and Salicornia europaea has changed into a community dominated by Halimione portulacoides, whereas the grazed salt marsh is still dominated by Puccinellia maritima and Salicornia europaea. These results are discussed in relation to the literature on the effect of cattle and sheep grazing on salt-marsh vegetation.     

Is livestock grazing compatible with amphibian diversity in the High Mountains of Córdoba, Argentina?

The High Mountains of Córdoba, Argentina have a long evolutionary history of grazing by large herbivores. However, about 400?years ago, European livestock were introduced and gradually replaced native herbivores. Since the 1920s, domestic herbivores have been the only large herbivores present in the area, causing severe soil erosion and a threat to the system diversity. The endemic fauna of the region includes four amphibian species. We evaluated the effect of livestock rearing on amphibian diversity and water bodies in woodlands and grasslands of the High Mountains of Córdoba. The work was conducted on stream stretches and ponds in two contrasting grazing situations: an area with livestock presence and another area where livestock was excluded 14?years ago. The following variables were recorded at each sampling site: amphibian richness and abundance, percentage of emergent, submerged and peripheral vegetation in areas surrounding the water bodies, water pH, and water dissolved O₂. No significant differences were detected in amphibian diversity between streams of both grazing situations, whereas a greater diversity (p?<?0.01) was observed in ponds in grazed grasslands. Our results suggest that livestock rearing, qualitatively measured as grazing and 14?years of livestock exclusion, in the study area would not have negative effects on amphibian diversity. This finding might be due to the long evolutionary grazing history of the area, large-scale livestock exclusion exhibiting a novel scenario. The ongoing reintroduction of native grazers may provide the benefits of grazing without the consequent soil erosion and habitat degradation associated with domestic livestock.     

Impact of grazing management on hibernating caterpillars of the butterfly Melitaea cinxia in calcareous grasslands

Semi-natural grasslands are increasingly grazed by large herbivores for nature conservation purposes. For many insects such grazing is essential for the conservation of their habitat, but at the same time, populations decrease at high grazing intensity. We hypothesised that grazing management may cause increased butterfly mortality, especially for life-stages with low mobility, such as hibernating caterpillars. To test this, we measured the effect of sheep grazing on overwinter larval survival. We used the Glanville fritillary (Melitaea cinxia), which has gregarious caterpillars hibernating in silk nests, as a model species. Caterpillar nests were monitored throughout the hibernating period in calcareous grassland reserves with low and high intensity sheep grazing and in an ungrazed control treatment. After grazing, 64?% of the nests at the high intensity grazing treatment were damaged or missing, compared to 8 and 12?% at the ungrazed and low intensity grazing treatment, respectively. Nest volume and caterpillar survival were 50?% lower at the high intensity grazing treatment compared to both ungrazed and low intensity grazing treatments. Nest damage and increased mortality were mainly caused by incidental ingestion of the caterpillars by the sheep. It is likely that grazing similarly affects other invertebrates, depending on their location within the vegetation and their ability to actively avoid herbivores. This implies that the impact of grazing strongly depends on the timing of this management in relation to the phenology of the species. A greater focus on immature and inactive life-stages in conservation policy in general and particularly in action plans for endangered species is required to effectively preserve invertebrate diversity.     

Temporal and spatial variation of arbuscular mycorrhizas in salt marsh plants of the Tagus estuary (Portugal)

The factors which may influence temporal and spatial variation in plant arbuscular mycorrhizal (AM) colonization and propagule occurrence were evaluated in a Portuguese salt marsh poor in plant diversity. Two distinct sites were studied: a more-flooded (low marsh) and a less-flooded zone (high marsh). AM root colonization, AM fungal spore number and inoculum potential, soil edaphic parameters and tidal flooding time periods were analysed. Levels of AM colonization were considerable in Aster tripolium and Inula crithmoides but very low in Puccinellia maritima and non-existent in Spartina maritima, Halimione portulacoides, Arthrocnemum fruticosum and Arthrocnemum perenne. Fungal diversity was very low, with Glomus geosporum dominant at both marsh zones. Colonization showed no spatial variation within marsh zones but temporal variation was observed in the high marsh, dependent on plant phenological phases. In the low marsh, no significantly seasonal variation was observed. Apparently, plant phenological events were diluted by stressful conditions (e.g. flooding, salinity). Spore density was significantly different between marsh zones and showed temporal variation in both zones. This study showed that distribution of mycorrhizas in salt marsh is more dependent on host plant species than on environmental stresses.     

Plant—herbivore interactions in streams near Mount St Helens

1. In four separate field experiments near Mount St Helens (Washington, U.S.A.) during 1986, the grazing effects of two large benthic herbivores, tadpoles of the tailed frog Ascaphus truei and larvae of the caddisfly Dicosmoecus gilvipes, were investigated using streamside channels and in-stream manipulations. In the experimental channels, abundances of periphyton and small benthic invertebrates declined significantly with increasing density of these larger herbivores. 2. In eleven small, high-gradient streams affected to varying degrees by the May 1980 eruption, in-stream platforms were used to reduce grazing by A, truei tadpoles on tile substrates. Single platforms erected in each tributary and compared to grazed controls revealed only minor grazing effects, and no significant differences among streams varying in disturbance intensity (and, consequently, tadpole density). However, results probably were confounded by high variability among streams in factors other than tadpole abundance. 3. Grazing effects were further examined in two unshaded streams with different tadpole densities, using five platforms per stream. In the stream with five tadpoles m^?2, grazing reduced periphyton biomass by 98% and chlorophyll a by 82%. In the stream lacking tadpoles, no significant grazing effects were revealed. Low algal abundance on both platforms and controls, and high invertebrate density in that stream (c. 30000m^?2) suggests that grazing by small, vagile invertebrates was approximately equivalent to that of tadpoles. 4. The influence of large benthic herbivores on algal and invertebrate communities in streams of Mount St Helens can be important, but reponses vary spatially in relation to stream disturbance history, local environmental factors, and herbivore distributional patterns and abundance.     

Does collembolan grazing influence nitrogen fixation by cyanobacteria in the high Arctic?

A field experiment was performed to test whether cyanobacterial nitrogen fixation rates increase or decrease when exposed to low and moderate microarthropod grazing intensities. The densities of naturally occurring Collembola, i.e. Hypogastrura viatica, were manipulated on a salt marsh covered with cyanobacteria in the high Arctic, Svalbard. Nitrogen-fixation rates in grazed cyanobacterial crust were measured and used as an indirect measurement of cyanobacterial biomass on three dates during one summer. After 30 days, a second order polynomial regression gave a good fit to the data, indicating an increase in nitrogen-fixation rates at low/intermediate grazing pressures and a decrease at high grazing pressures. Thus, grazing collembolans may influence the nitrogen-fixation rates in an arctic salt-marsh community. Although based on a small set of data, the study indicates a compensatory fixation at low grazing pressures followed by a reduction at high grazing pressures. Accepted: 21 March 2000     

Herbivory and ramet performance in the clonal herb Trientalis europaea L.

1 The intensity of leaf damage caused by invertebrate herbivores and grazing by vertebrates and their effects on clonal growth, survival and reproduction were examined in a population of the forest herb Trientalis europaea during 4 years.
2 Levels of herbivory were low and varied between years in the studied population. Larger ramets were exposed to a greater risk of both grazing and leaf defoliation. Ramets that suffered leaf damage in 1 year experienced an increased probability of leaf damage in succeeding years. The probability of suffering herbivore attack was independent among the ramets belonging to the same clonal fragment, suggesting that clonal propagation might operate as a risk-spreading strategy in this species.
3 Leaf damage did not affect any measure of plant performance, probably due to the low amounts of leaf area removed by invertebrate herbivores. In contrast, vertebrate grazing affected all phases of the pseudo-annual life cycle of T. europaea . Grazing prevented flowering and fruiting, increased ramet mortality during summer and decreased tuber production. Furthermore, grazed ramets produced shorter stolons and smaller tubers, which in turn had a lower winter survival and produced smaller ramets in the following growing season. The large impact of grazing was due to the consumption of the whole of the single shoot of ramets of T. europaea . Although regrowth was possible, secondary shoots were significantly smaller and assimilation was delayed.
4 Tubers originating from grazed ramets were placed shallower than tubers from ungrazed ramets. This could be due to an alteration in the growth pattern of stolons of the grazed ramets.     

Mobile dunes and eroding salt marshes

The paper deals with general outlines of salt marsh and dune vegetation in the Ellenbogen and Listland area on Sylt (Schleswig-Holstein, FRG). The composition of current salt marsh vegetation is considered to be mainly the result of a long-lasting process of tidal inundation, grazing, and a permanent influence of groundwater seepage from the surrounding dunes. The lower salt marsh communities have shown constancy for 67 years, due to the effect of heavy grazing. The mid-upper salt marsh communities demonstrated a succession from aPuccinellia maritima-dominated community of the lower marsh to aJuncus gerardii-dominated community of the mid-upper salt marsh, which may be due to the transport of sand — over a short time — on the surface of the marsh. The area covered by plant communities of annuals below Mean High Water (MHW) seemed to diminish. Salt marsh soils, especially of the mid-upper marsh, indicate sandy layers resulting from sand drift of the dunes. Dry and wet successional series of the dunes in the Listland/Ellenbogen area both show grassy stages shifting to dwarf shrubs as final stages. White primary dunes can only be found on the accreting shoreline of the Ellenbogen, which is also grazed by sheep; vegetation cover therefore remains dominated by grasses, mosses and lichens. Three mobile dunes (as the most prominent features of this landscape) have been left unaffected by seeding and planting by local authorities. Grazing is considered to be an inadequate tool in nature conservation as long as natural processes are to prevail in the landscape as major determinants.     

Grazing by Talorchestia longicornis on an algal mat in a new England salt marsh

Grazing experiments using ¹⁴C and an analysis of fecal pellets and gut contents established that the gammaridean amphipod, Talorchestia longicornis Say, ingests blue-green algae on algal mats in a Massachusetts salt marsh. This grazing had a measurable effect on the lower algal mat, where the density of T. longicornis was high. Exclusion of amphipods resulted in increases in chlorophyll a content, carbon incorporation, and nitrogen fixation. This effect was not seen on the upper mat where T. longicornis was less abundant. The assimilation efficiency of T. longicornis feeding on a diet consisting mainly of blue-green algae was surprisingly high (67 %) considering that blue-green algae are usually considered as a poor quality food for herbivores. The population of T. longicornis seems to be annual, with growth of the overwintered juveniles in spring and early summer.