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.     

Directional and non‐directional vegetation changes in a temperate salt marsh in relation to biotic and abiotic factors

Abstract. The effects of reduction and cessation of sheep grazing on salt‐marsh vegetation were studied on a formerly intensively grazed salt marsh in northern Germany. Plant species cover was recorded in 45 permanent plots from 1992 to 2000. In 1995, physical and chemical soil parameters were analysed. Results of Redundancy Analysis (RDA) indicated that salinity and the depth of anoxic conditions below the surface were the most important soil factors related to the spatial vegetation pattern. Furthermore, plant species distribution was influenced by present and past grazing intensity, by soil grain size and nitrogen content. Vegetation changes over 9 yr were analysed by non‐linear regression. The cover of Aster tripolium, Atriplex portulacoides, and, to a lesser extent, Artemisia maritima and Elymus athericus increased due to reduced grazing pressure, whereas the cover of Salicornia europaea decreased. After a strong increase in the first years Aster decreased 2 to 6 yr after abandonment. In the mid salt‐marsh zone Puccinellia maritima was replaced by Festuca rubra. The cover of Puccinellia, Festuca, Suaeda maritima, Glaux maritima and Salicornia fluctuated strongly, probably due to differences in weather conditions and inundation frequency. Species richness per 4 m² generally increased while vegetation evenness decreased during the study period. Only in the high salt marsh abandoned for 9 yr did the number of species decrease slightly. Thus far, cessation of grazing did not lead to large‐scale dominance of single plant species.     

Aggregative responses of brent geese on salt marsh and their impact on plant community dynamics

The aggregative responses and habitat preferences of a generalist herbivore, the dark-bellied brent goose Branta bernicla bernicla, feeding on salt marshes are examined in relation to vegetation community characteristics and the abundances of individual plant species. In the autumn, feeding was strongly concentrated on the low marsh, which had the highest biomass of the preferred food plant, Salicornia europaea. There was a strong aggregative response of the geese to the abundance of S. europaea. A decline in the availability of S. europaea led to an increase in the pattern of aggregation in relation to the two other major food plants on the low marsh, Aster tripolium and Puccinellia maritima. The availability of these food plants, however, reached critically low levels in mid-winter and the geese abandoned the low marsh for the high marsh. Within the high marsh, the plant communities selected tended to be dominated by the inedible species Limonium vulgare. The food plants selected were P. maritima in the winter and P. maritima and Triglochinmaritimum in the spring. On the high marsh, aggregative responses were shown to both P. maritima and T. maritimum, but in both cases, aggregation increased up to a critical level of biomass, and then declined. The prevention of grazing with exclosures for 3 years led to an increase in the abundance of P. maritima on both high and low marshes. This change was slight on high marsh but pronounced on low marsh, where S. europaea showed a decrease in abundance in the exclosures over this time. The implications of the aggregative responses for the population dynamics of P. maritima and S. europaea are discussed. Received: 11 September 1997 / Accepted: 12 December 1997     

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.     

Sulphide tolerance in coastal halophytes

The effect of sulphide on the growth of several species of salt-marsh plants was investigated. Relative growth rates were significantly reduced in two upper-marsh species, Festuca rubra and Atriplex patula, and in the lower-marsh species Puccinellia maritima. However the growth of Salicornia europaea, a species frequently associated with sulphide-containing sediments, was unaffected. In a separate experiment the wide ranging halophyte Aster tripolium, also appeared to be tolerant of sulphide at a concentration frequently encountered in salt marshes. Sulphide pretreatment inhibited the activity of two metallo-enzymes, polyphenol oxidase and external phosphatase, in plants from the upper marsh, but had no effect on enzymes from P. maritima or S. europaea. The rate of respiration by root tissue was significantly reduced in all of the species investigated but whereas the uptake of ⁸⁶rubidium was markedly inhibited in the other three species, uptake by S. europaea showed a significant stimulation. Similarly, whereas sulphide-grown plants of F. rubra, A. patula and P. maritima had a considerably reduced tissue iron content, the total iron concentration in S. europaea tissues was comparable to that of the controls. When the sulphide-tolerant species A. tripolium was grown in sulphide-containing media there was no significant effect on the tissue concentration of any of the elements investigated. These results are discussed in relation to possible mechanisms of sulphide toxicity and resistance.     

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.     

The role of spatio‐temporal heterogeneity in the establishment and maintenance of Suaeda maritima in salt marshes

Abstract. The effects of disturbance and microtopography on the organization and dynamics of plant communities were studied in a European salt marsh located in the Bay of Mont St. Michel, France. The existence of seed trapping mechanisms was also tested. The study took place in the lower and middle marsh plant communities dominated by the perennials Puccinellia maritima and Halimione portulacoides, respectively and associated with the annual Suaeda maritima. Three treatments were used in series of plots placed in each community: (1) vegetation removal and root destruction to a depth of 10 cm and refilling, (2) non‐remnant herbicide treatment without vegetation removal and (3) creation of depressions (20 cm deep). These treatments were compared with adjacent control plots. The first year of the experiment showed that the perennials facilitated the establishment of Suaeda by trapping its seeds. Estimation of cover, density and biomass over 5 yr following the disturbances showed that in the first 2 yr Suaeda dominated the disturbed plots. Thereafter Suaeda was gradually eliminated by competitive exclusion after ca. 3 yr in the zone originally dominated by Puccinellia maritima and after 4 yr in the zone occupied by Halimione portulacoides. Depressions constituted refuge habitats for Suaeda by limiting competition with the perennials but also led to a high risk of mortality with temporal fluctuations in density. Despite a period of investigation limited to 5 yr, our study demonstrated that natural disturbances of various types occurred and influenced the dynamics of Suaeda, Halimione and Puccinellia. We deduced that natural disturbances and microtopography are responsible for the maintenance of the habitat in a state of non‐equilibrium by favouring the establishment of both spatial and temporal environmental heterogeneity. These conditions appear to be particularly favourable for the maintenance of annual species such as Suaeda maritima.     

Factors affecting the population dynamics of Suaeda maritima at initial stages of development

This study describes life history and population dynamics of the annual halophyte Suaeda maritima, at initial stages of seedling development, in salt-marshes of the Mont-Saint-Michel Bay. Effects of salinity on seed germination and seedling development were studied both in the laboratory and in the field. Salinity was measured as sediment electrical conductivity. Experiments in the laboratory showed that Suaeda maritima was relatively tolerant to salinity during the germination stage. In the field, germination occurred when soil salinity was minimal. Almost all seeds germinated in the field; this may explain the absence of a persistent seed bank. Salinity did not seem to affect the growth of seedlings either. Mortality in the field was attributed to physical factors such as anoxia during temporary immersion, burial by sediment and tidal uprooting. The intensity of these effects varied according to the location of Suaeda maritima seedlings: in hollows, along the edge of tidal creeks, in gaps among dense vegetation cover, or in pattern with Puccinellia maritima. Perennial vegetation did not restrict Suaeda maritima germination. In contrast, perennials such as Puccinellia maritima and Halimione portulacoides limited the development of Suaeda maritima seedlings since only a small number of this annual emerged beneath their canopies. The fate of seedlings depended to a great extent on the permeability of the perennial canopy to light. Thus, disturbance resulting in bare patches inside the perennial vegetation seemed essential for the development of this annual species. Suaeda maritima can grow in the presence of Puccinellia maritima so long as the latter present as an open matrix. Suaeda maritima may also benefit from protection against desiccation and tidal action where this occurs.     

Germination and early establishment of lower salt-marsh species in grazed and mown salt marsh

Abstract. Lower salt-marsh species occur more in the grazed than in the mown sites of a salt marsh in Schiermonnikoog in the Netherlands. This was demonstrated by a sowing experiment which determined which characteristic of the stand structure, height of the canopy or percentage of bare soil, is responsible for this difference. The total number of seedlings which emerged was negatively related to the height of the canopy and positively to the percentage of bare soil. The survival of seedlings of Suaeda maritima and Plantago maritima could be explained by the height of the canopy and for the latter species also by the percentage of bare soil. The survival of Aster tripolium and Plantago maritima was higher in grazed than in mown sites. Since the amount of bare soil was higher than seemed necessary for germination and survival, it was concluded that the taller canopy was responsible for the absence of lower salt-marsh species in the mown sites.     

The effect of sodium chloride on enzyme activities from four halophyte species of chenopodiaceae

The in vitro effect of sodium chloride on the enzyme activity of four halophytes, Beta vulgaris ssp. maritima (L.) Thell., Halimione portulacoides (L.) Aell., Salicornia ramosissima Woods and Suaeda maritima (L.) Dum. was investigated. The activity was, in general, affected by sodium chloride in a similar manner to that reported for salt sensitive species. The most notable exceptions were the sodium chloride stimulated ATPases of Beta and Salicornia.     

Food preferences of Shorelarks Eremophila alpestris,Snow Buntings Plectrophenax nivalis and Twites Carduelis flavirostris wintering in the Wadden Sea

Capsule Seeds of plants from lower salt marsh communities are preferred, with insects less important.

Methods Droppings of Shorelarks Eremophila alpestris, Snow Buntings Plectrophenax nivalis and Twites Carduelis flavirostris sampled in the German Wadden Sea were analysed and compared with food abundance to assess preferences.

Results Shorelarks prefer seeds of Salicornia sp., Suaeda maritima, Atriplex sp., Halimione portulacoides and unidentified small grass seeds. Insects are eaten mainly in periods of seed shortage, but are consumed in smaller amounts during the winter. The food composition of Snow Buntings is very similar, but additionally Triglochin maritimum is commonly eaten. Twites are specialized on seeds of Salicornia sp. and Suaeda maritima and rarely ingest other seeds and insects. All seeds consumed were of plants from lower salt marsh communities. Many halophyte seeds, and especially those that birds feed on, are rich in energy. Large seeds and those which need a long handling time are avoided.

Conclusion Changes in the lower salt marshes of the Wadden Sea by embankments and intensified grazing might have been responsible for the rapid population decline from the 1960s to 1980s.     

Growth strategies of four salt marsh plants on Mankyung River estuary in Korea

The emergence of seedlings, the length of roots and shoots, and the biomass of four dominant plant species and shore height were measured to investigate the growth strategy of these plants on the salt marsh of Mankyung River estuary. Four salt marsh plants showed a distinctive zonation, for example, Suaeda japonica was predominantly spread around the low salt marsh, Atriplex gmelini and Aster tripolium were in the middle, and S. asparagoides was in the upper part of the marsh. In terms of emergence of seedlings, S. japonica appeared first followed by A. gmelini, S. asparagoides, and A. tripolium. The growth strategies of halophytes were as follows: S. japonica germinated earlier than the other halophytes so that its root grew rapidly and extensively at the beginning of growth. This species adopted a continuous germination strategy, allowing growth whenever favorable conditions were provided. A. gmelini germinated later than S. japonica, as a quasi-simultaneous germination type, it showed the highest germination rate within the shortest time. Aster tripolium germinated later than any other halophyte. Since this species exhibited characteristics between the continuous germination type and the quasi-simultaneous germination type, it did not show a very high germination rate. Instead, it showed continuous germination and consistent growth of both above-ground and underground parts. Suaeda asparagoides showed an especially high emergence rate at the beginning of its growth. However, the high density retarded its growth until the middle stage. Its roots extended longer than the other halophytes, allowing it to grow well in the dry conditions of the upper marsh.     

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.     

Aspects of population dynamics in Halimione portulacoides communities

Summary Studies on sample plots inHalimione portulacoides communities show that environmental disturbances, either natural or induced by man, start a sequence of partly overlapping density maxima inSuaeda maritima, Aster tripolium andPuccinellia maritima successively, before the originalHalimione community totally recovers. When succession time before recovering is long enough, there are tendencies in redundancy of this sequence stressing the unilinear character of the succession. Minor environmental impacts induce a longer time-lag period of theSuaeda density maximum, suggesting threshold values of these impacts for the species to maintain minimal population densities or to become locally extinct. This sequence of interim species starting after an environmental disturbance, suggests also a gradient character in various biological attributes, for instance in life-time, propagation, nutrient and genetic plasticity strategies. The mechanism described can therefore be interpreted as a complex of mostly well-adapted and well-integrated inherent species strategies capable of absorbing environmental shocks. It is suggested that in the salt-marsh ccosystem the pattern of spatial variation in densities and that of temporal variation in fluctuations of the three species populations under natural conditions reflect corresponding patterns of environmental disturbances in the vegetation taking into account a timelag associated with the magnitude of the impact concerned.Contribution to the Symposium on Plant species and plant communities held at Nijmegen, 11–12 November 1976, on the occasion of the 60th birthday of Professor Victor Westhoff.Nomenclature follows Heukels-van Ooststroom. Flora van Nederland, 18e druk, 1975. Wolters-Noodhoff, Groningen.The authors are greatly indebted to Dr K.F. Vaas (Yerseke) for reviewing the English text.Communication Nr. 160.     

Vegetation change in a man‐made salt marsh affected by a reduction in both grazing and drainage

Abstract. In order to restore natural salt marsh in a 460‐ha nature reserve established in man‐made salt marsh in the Dollard estuary, The Netherlands, the artificial drainage system was neglected and cattle grazing reduced. Vegetation changes were traced through two vegetation surveys and monitoring of permanent plots over 15 yr after the management had been changed. Exclosure experiments were started to distinguish grazing effects from effects of increased soil waterlogging caused by the neglect of the drainage system. Both vegetation surveys and permanent plots demonstrated a dichotomy in vegetation succession. The incidence of secondary pioneer vegetation dominated by Salicornia spp. and Suaeda maritima increased from 0 to 20%, whereas the late‐successional (Phragmites australis) vegetation from 10 to 15%. Grazing intensity decreased towards the sea. The grazed area contracted landward, which allowed vegetation dominated by tall species to increase seaward. Grazing and increased waterlogging interacted in several ways. The impact of trampling increased, and in the intensively grazed parts soil salinity increased. This can probably be explained by low vegetation cover in spring. Framework Ordination, an indirect‐gradient‐analysis technique, was used to infer the importance of environmental factors in influencing changes in species composition. Many changes were positively or negatively correlated with soil aeration and soil salinity, whereas elevation was of minor importance. Grazing accounted for only a few changes in species frequency. Changes in permanent plots were greater during the first than during the second half of the study period. In exclosures that were installed halfway through the study period, there was a relatively rapid recovery of previously dominant species that had decreased during the first half of the study period. Species richness per unit area in the reserve increased. At the seaward side of the marsh, the altered management allowed succession to proceed leading to establishment of stands of Phragmites australis, whereas on the landward side, the combination of moderate grazing with neglect of the drainage system appeared an effective measure in maintaining habitats for a wider range of halophytic species.     

High occurrence of rare inland halophytes on post‐mining sites in western Ukraine

Rare inland halophytic vegetation including the associations Salicornietum prostratae, Spergulario marginatae‐Suaedetum prostratae, Puccinellietum limosae, com. Puccinellietum limosae – a variant of Aster tripolium subsp. pannonicus and Astero pannonici‐Bolboschoenetum compacti have been found on 3 localities in the Drohobych region of western Ukraine. These are isolated (‘island’) localities, almost unknown until today. Halophytes grew there on salted sites in the vicinity of the old salt industry. Species of halophytes previously not known from this region were discovered: Aster tripolium subsp. pannonicus, Bolboschoenus maritimus, Gypsophila paniculata, Limonium gmelinii, Lotus tenuis, Puccinellia distans, Sonchus palustris, Suaeda prostrata, Trifolium fragiferum var. bonannii, Triglochin maritima, Typha laxmannii and Zannichellia palustris subsp. pedicellata. The occurence of these plants strictly depends on the presence of salt in the soil and they are endangered by shrinkage of salted sites following the termination of the salt industry in Drohobych region.     

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.     

Biomass and aboveground production of four angiosperms in Cantabrian (N. Spain) salt marshes

New data of aboveground biomass and production of four angiosperms over a 12 month period for the Cantabrian Sea salt marshes (Bay of Biscay, N. Spain) are presented. Based on harvest methods, maximum aboveground total biomass values for Spartina maritima (Curtis) Fernald, Spartina alterniflora Loisel, Salicornia ramosissima J. Woods and Halimione portulacoides (L.) Aellen were 628, 1109, 480 and 1267 gm^-2, respectively. We conclude that although a slight latitudinal gradient in biomass is revealed in the data compiled with reference to some of the species studied, more work is neccesary in order to assess the potential productivity of these ecosystems on the coasts of Europe and/or to make comparisons with salt marshes of the American coasts. Annual net aerial primary production estimates using Smalley's method were: 296, 1160, 486 and 952 gm^-2yr^-1, for Spartina maritima, Spartina alterniflora, Salicornia ramosissima and Halimione portulacoides, respectively. These results together with turnover rate estimates point to the lack of vigour of the native S. maritima, while the exotic S. alterniflora, which seems to be spreading along the Cantabrian estuaries, behaves like a veritable pionner throughout the low marshes in this region.     

Utilisation of Wadden Sea salt marshes by geese in relation to livestock grazing

To arctic breeding geese, the salt marshes of the International Wadden Sea are important spring staging areas. Many of these marshes have always been grazed with livestock (mainly cattle and sheep). To evaluate the influence of livestock grazing on composition and structure of salt-marsh communities and its consequences for habitat use by geese, a total of 17 pairs of grazed and ungrazed marshes were visited both in April and May 1999, and the accumulated grazing pressure by geese was estimated using dropping counts. Observed grazing pressure was related to management status and to relevant vegetation parameters.The intensity of livestock grazing influences the vegetation on the marsh. Salt marshes that are not grazed by livestock are characterised by stands with a taller canopy, a lower cover of grasses preferred by geese, and a higher cover of plants that are not preferred.Overall goose-dropping densities are significantly lower in ungrazed marshes compared to marshes grazed by livestock. Some ungrazed marshes had comparatively high goose grazing pressure, and these were all natural marshes on a sandy soil, or artificial mainland marshes with a recent history of intensive livestock grazing. Goose grazing is associated with a short canopy. The plant communities with short canopy, dominated by Agrostis stolonifera, Festuca rubra and Puccinellia maritima, together account for 85% of all goose droppings in our data.The sites that were not visited by geese differed very little from those that were visited, in the parameters we measured. This might indicate that there was no shortage of available habitat for spring staging geese in the Wadden Sea, in the study period.