Spatially heterogeneous pressure raises risk of catastrophic shifts

Ecosystems may exhibit catastrophic shifts, i.e. abrupt and irreversible responses of ecosystem functions and services to continuous changes in external conditions. The search for early warning signs of approaching shifts has so far mainly been conducted on theoretical models assuming spatially-homogeneous external pressures (e.g. climatic). Here, we investigate how a spatially explicit pressure may affect ecosystems’ risk of catastrophic shifts and the associated spatial early-warning signs. As a case study, we studied a dryland vegetation model assuming ‘associational resistance’, i.e. the mutual reduction of local grazing impact by neighboring plants sharing the investment in defensive traits. Consequently, grazing pressure depends on the local density of plants and is thus spatially-explicit. We focus on the distribution of vegetation patch sizes, which can be assessed using remote sensing and are candidate early warning signs of catastrophic shifts in drylands. We found that spatially explicit grazing affected both the resilience and the spatial patterns of the landscape. Grazing impact became self-enhancing in more fragmented landscapes, disrupted patch growth and put apparently ‘healthy’ drylands under high risks of catastrophic shifts. Our study highlights that a spatially explicit pressure may affect the nature of the spatial pattern observed and thereby change the interpretation of the early warning signs. This may generalize to other ecosystems exhibiting self-organized spatial patterns, where a spatially-explicit pressure may interfere with pattern formation.     

Predator behaviour and predation risk in the heterogeneous Arctic environment

1. Habitat heterogeneity and predator behaviour can strongly affect predator-prey interactions but these factors are rarely considered simultaneously, especially when systems encompass multiple predators and prey. 2. In the Arctic, greater snow geese Anser caerulescens atlanticus L. nest in two structurally different habitats: wetlands that form intricate networks of water channels, and mesic tundra where such obstacles are absent. In this heterogeneous environment, goose eggs are exposed to two types of predators: the arctic fox Vulpes lagopus L. and a diversity of avian predators. We hypothesized that, contrary to birds, the hunting ability of foxes would be impaired by the structurally complex wetland habitat, resulting in a lower predation risk for goose eggs. 3. In addition, lemmings, the main prey of foxes, show strong population cycles. We thus further examined how their fluctuations influenced the interaction between habitat heterogeneity and fox predation on goose eggs. 4. An experimental approach with artificial nests suggested that foxes were faster than avian predators to find unattended goose nests in mesic tundra whereas the reverse was true in wetlands. Foxes spent 3.5 times more time between consecutive attacks on real goose nests in wetlands than in mesic tundra. Their attacks on goose nests were also half as successful in wetlands than in mesic tundra whereas no difference was found for avian predators. 5. Nesting success in wetlands (65%) was higher than in mesic tundra (56%) but the difference between habitats increased during lemming crashes (15%) compared to other phases of the cycle (5%). Nests located at the edge of wetland patches were also less successful than central ones, suggesting a gradient in accessibility of goose nests in wetlands for foxes. 6. Our study shows that the structural complexity of wetlands decreases predation risk from foxes but not avian predators in arctic-nesting birds. Our results also demonstrate that cyclic lemming populations indirectly alter the spatial distribution of productive nests due to a complex interaction between habitat structure, prey-switching and foraging success of foxes.     

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.     

Dynamics of intertidal marshes near shallow estuaries in Louisiana

A three-year (1991–1993) field investigation was conducted to quantify the hydrodynamics of intertidal marshes adjacent to tidal channels and shallow bays within two Louisiana coastal regions: (1) the sediment-rich Atchafalaya Basin, and, (2) the sediment-poor Terrebonne Basin with relatively minor riverine inflow. The Terrebonne Basin marsh is regularly inundated and flooding is characterized by sporadic draining interspersed by prolonged flooding events. The maximum water depth on the marsh surface exceeds 50 cm, the flow velocity across marsh surface reaches 10 cm sec^–1, and the sediment deposition rate varies from 10 to 90 g m^–2 per tidal cycle. This rather high sediment deposition rate occurs during winter storms with strong southerly winds. In contrast, the marsh site within the sediment-rich Atchafalaya Basin is irregularly inundated and characterized by sporadic flooding interspersed by prolonged draining. There the marsh flooding depth rarely exceeds 25 cm, the over-marsh flow velocity barely reaches 2.5 cm sec^–1, and the sediment deposition rate ranges from 5 to 50 g m^–2 per tidal cycle. The surprisingly low rate of sediment deposition in a marsh within a sediment-rich region is largely due to the man-made canals that alter the hydrologic regime in the upper reaches of the tidal channel.     

Foraging patterns of voles at heterogeneous avian and uniform mustelid predation risk

Temporal variation of antipredatory behavior and a uniform distribution of predation risk over refuges and foraging sites may create foraging patterns different from those anticipated from risk in heterogenous habitats. We studied the temporal variation in foraging behavior of voles exposed to uniform mustelid predation risk and heterogeneous avian predation risk of different levels induced by vegetation types in eight outdoor enclosures (0.25 ha). We manipulated mustelid predation risk with weasel presence or absence and avian predation risk by reducing or providing local cover at experimental food patches. Foraging at food patches was monitored by collecting giving-up densities at artificial food patches, overall activity was automatically monitored, and mortality of voles was monitored by live-trapping and radiotracking. Voles depleted the food to lower levels in the sheltered patches than in the exposed ones. In enclosures with higher avian predation risk caused by lower vegetation height, trays were depleted to lower levels. Unexpectedly, voles foraged in more trays and depleted trays to lower levels in the presence of weasels than in the absence. Weasels match their prey's body size and locomotive abilities and therefore increase predation risk uniformly over both foraging sites and refuge sites that can both be entered by the predator. This reduces the costs of missing opportunities other than foraging. Voles changed their foraging strategy accordingly by specializing on the experimental food patches with predictable returns and probably reduced their foraging in the matrix of natural food source with unpredictable returns and high risk to encounter the weasel. Moreover, after 1 day of weasel presence, voles shifted their main foraging activities to avoid the diurnal weasel. This behavior facilitated bird predation, probably by nocturnal owls, and more voles were killed by birds than by weasels. Food patch use of voles in weasel enclosures increased with time. Voles had to balance the previously missed feeding opportunities by progressively concentrating on artificial food patches.     

Below-ground biomass in healthy and impaired salt marshes

Twelve salt marshes in south Louisiana (USA) were classified as either impaired or healthy before a summer sample collection of above- and below-ground biomass and determination of sediment accretion rates. The above-ground biomass of plant tissues was the same at both impaired and healthy salt marshes and was not a good predictor of marsh health. However, below-ground root biomass in the upper 30cm was much lower in the impaired marshes compared to the healthy marshes. Compromises to root production apparently occur before there is an obvious consequence to the above-ground biomass, which may quickly collapse before remedial action can be taken. The subsequent change in vertical position of the marsh surface may be equivalent to many years of accretion, and be irreversible within decades without considerable effort. These results are consistent with the hypothesis that it is the plants below-ground accumulation of organic matter, not inorganic matter that governs the maintenance of salt marsh ecosystem in the vertical plane. Reversing the precursor conditions leading to marsh stress before the collapse of the above-ground biomass occurs is therefore a prudent management objective and could be easier than restoration.     

Cellulose-decomposing fungi of salt marshes in Egypt

Seventy-five species and three varieties which belong to thirty-four genera were identified from 74 soil samples collected from salt marshes in Egypt. The most frequent fungi wereAspergillus fumigatus, Aspergillus niger, Cladosporium herbarum andAlternatia alternata, followed byAspergillus terreus,Curvularia spicifera andPenicillium notatum. Six genera were of moderate occurrence:Penicillium, Futarium, Curvularia, Rhizopus, Stachybotrys, andChaetomium. Five genera were of low occurrence:Paecilomyces, Oephalosporium, Epicoccum, Mucor andMyrothecium.     

Restoration of salt marshes in the Netherlands

The conquest of land from the sea has been a long tradition in the Netherlands. When salt marshes were high enough, they were embanked when it was economically feasible, and transformed into intensively exploited agricultural land. This resulted in the transformation of halophytic communities to glycophytic communities. Often as an alternative, a low levee, a summerdike was built, which greatly reduced the flooding frequency of the landward summerpolder, hence creating a sedimentation deficit therein. Such summerpolders now cover 1200 ha in the Netherlands, 2100 ha in NW-Germany and small areas in England. Due to continuous embankments, the present salt-marsh area is relatively small with respect to the tidal basins. Discussions have been started how to increase the salt-marsh area. Two options will be discussed, firstly de-embankment of summerpolders and maintenance of the protective seawall, secondly increase of the effects of saline seepage behind the seawall by top soil removal. Both options include the restoration of salt-marsh communities (target communities) in intensively agriculturally exploited sites that have been salt marshes before. From the few examples abroad and experiments it is discussed (1) to which extent the sedimentation deficit in summerpolders could be compensated for, (2) if the soil seed bank is likely to contribute to re-establishment of salt-marsh communities, (3) if the dispersal of propagules of halophytic plants will be possible by hydrochory when the summerdike is breached, (4) to what extent is dispersal by endozoochory through waterfowl important in case re-establishment in a saline seepage area behind the seawall without open connection to the sea is envisaged. Two case studies of de-embanked summerpolders in the Netherlands revealed that the sedimentation deficit can be counteracted by rapid sedimentation, provided enough transport is possible from the foreshore. Dispersal by incoming tidal water from the nearby salt-marsh source area into the target area is possible for many salt-marsh plant species. The rate of success seems to depend on the relative position of source area and target area. A case study in a saline seepage area after top soil removal in the Netherlands, showed that the number of viable seeds dispersed by droppings from waterfowl is limited. Hence the possibilities for restoration of inland halophytic plant communities seem much lower than after de-embankment of summerpolders.     

Zonation of shrubs in western Atlantic salt marshes

We explored the generality of the processes mediating shrub zonation in western Atlantic salt marshes by comparing the results of our experiments in Georgia, USA with previous studies from Rhode Island, USA. The shrub Borrichia frutescens dominates the terrestrial border of many Georgia salt marshes. Within the shrub zone, physical stress increased at lower elevations, shrubs at lower elevations were stunted, and experimentally reducing physical stress reduced shrub stunting. Below the shrub zone, physical stress increased further, and the grass Spartina alterniflora dominated. Transplant and neighbor-removal experiments indicated that the lower border of the shrub zone was set more by physical stress than by competition, but that the upper border of the grass zone was set primarily by competition with shrubs. Laboratory experiments indicated that S. alterniflora seedlings survived best and shrub seedlings worst in the flooded, salty treatment that mimicked low-marsh conditions. These processes are similar to those maintaining zonation patterns between the shrub Iva frutescens and the rush Juncus gerardi in Rhode Island salt marshes. However, markedly different processes appear to occur further to the north, where woody shrubs are absent from coastal marshes, and further to the south, where woody plants (mangroves) dominate coastal wetlands.     

The coastal salt marshes of California and Baja California

A comparative study of the North American salt marshes between Magdalena Plain (Baja California, 24° N)_and the north of San Francisco (California 38° N) is made in order to establish relationships between floristic and zonal gradients. The sigmatistic-phytosociological method along with Average Linkage Clustering are used to describe the zonation of these salt marshes and to define the optimum zone of several halophilous plants and communities.Abbreviations (As) Arthrocnemum subterminalis - (Ds) Distichlis spicata - (Fg) Frankenia salina - (Jc) Jaumea carnosa - (MI) Monanthochloe littoralis - (S1) Sarcocornia pacifica, prostrate biotype - (S2) Sarcocornia pacifica, erect biotype - (Sp) Spartina foliosa     

Environmental gradients and herbivore feeding preferences in coastal salt marshes

Current theories of plant-herbivore interactions suggest that plants may differ in palatability to herbivores as a function of abiotic stress; however, studies of these theories have produced mixed results. We compared the palatability of eight common salt marsh plants that occur across elevational and salinity stress gradients to six common leaf-chewing herbivores to determine patterns of plant palatability. The palatability of every plant species varied across gradients of abiotic stress in at least one comparison, and over half of the comparisons indicated significant differences in palatability. The direction of the preferences, however, was dependent on the plant and herbivore species studied, suggesting that different types of stress affect plants in different ways, that different plant species respond differently to stress, and that different herbivore species measure plant quality in different ways. Overall, 51% of the variation in the strength of the feeding preferences could be explained by a knowledge of the strength of the stress gradient and the type of gradient, plant and herbivore studied. This suggests that the prospects are good for a more complex, conditional theory of plant stress and herbivore feeding preferences that is based on a mechanistic understanding of plant physiology and the factors underlying herbivore feeding preferences.     

Osmophilous fungi in the salt marshes of Kuwait.

During an investigation of the mycoflora inhabiting saline soils in Kuwait, special attention was focused on osmophilous fungi. A total of 101 species belonging to 46 genera were encountered from 40 soil samples collected from salt marshes using Czapek's agar supplemented with 40% sucrose. Soil samples were collected from different habitats at different distances from the water edge. Soils near to the water edge were poor in their fungal content, while those taken from areas covered by Juncus arabicus contained highest fungal populations. The recorded genera were classified as follows: 7 were of high frequency of occurrence, 8 moderate, 17 low, and 13 were rare. The order of dominance was Aspergillus, Alternaria, Penicillium, Cephalosporium, Fusarium, Stachybotrys, and Drechslera. Comparison between our results and those in other studies showed that there is no fungal flora characteristic of saline soils.     

The importance of predation,substrate and spatial refugia in determining lotic insect distributions

An experiment was conducted to evaluate the interaction between predation, substrate, and spatial refugia in the organization of a stream insect community (Reeds Creek, Pendleton Co., West Virginia). Patterns of insect colonization were compared between fish exclusion cages and open controls that allowed access to vertebrate predators. Each cage contained 4 different substrates that varied in the relative amount of spatial refugia. Fish had little influence on the diversity or abundance of any insect taxa, even when spatial refugia were limited. The only significant effect due to predation, was an increased diversity of large (>8 mm) invertebrates in the absence of predators. However, because these taxa were relatively rare, the overall role of fish predation on insect community structure was minimal.In contrast, substrate had a marked effect on insect colonization. Insects were always more abundant (number/basket) on loose substrates containing large numbers of interstitial spaces, compared to cement-embedded substrates with few refuges available. In addition, invertebrates were more abundant on loose gravel compared to loose cobbles. Howver, when substrate preferencesrd were examined according to insect density (number/m²), loose cobbles were generally the preferred substrate. The present experiment rejects the hypothesis that patterns of substrate colonization can be explained as differential insect mortality by predators, due to varying amounts of refugia. Alternative mechanisms such as differing amounts of trapped detritus and substrate surface area may account for substrate preference.     

Spatially heterogeneous stochasticity and the adaptive diversification of dormancy

Diversified bet‐hedging, a strategy that leads several individuals with the same genotype to express distinct phenotypes in a given generation, is now well established as a common evolutionary response to environmental stochasticity. Life‐history traits defined as diversified bet‐hedging (e.g. germination or diapause strategies) display marked differences between populations in spatial proximity. In order to find out whether such differences can be explained by local adaptations to spatially heterogeneous environmental stochasticity, we explored the evolution of bet‐hedging dormancy strategies in a metapopulation using a two‐patch model with patch differences in stochastic juvenile survival. We found that spatial differences in the level of environmental stochasticity, restricted dispersal, increased fragmentation and intermediate survival during dormancy all favour the adaptive diversification of bet‐hedging dormancy strategies. Density dependency also plays a major role in the diversification of dormancy strategies because: (i) it may interact locally with environmental stochasticity and amplify its effects; however, (ii) it can also generate chaotic population dynamics that may impede diversification. Our work proposes new hypotheses to explain the spatial patterns of bet‐hedging strategies that we hope will encourage new empirical studies of this topic.     

Sounds recorded in salt marshes and attributed to the European eel

The sounds were recorded in coastal ponds off the French Atlantic coast. They are characterized by continuous series of regularly spaced sounds, each sound being composed of one to four pulses of 7.8-ms duration. The main frequencies of these sounds are under 350 Hz. Due to the special characteristics of these sounds, their temporal patterning, their coastal pond provenance and their close similarity to sounds made by the American eel (Anguilla rostrata), we attribute them to the European eel (Anguilla anguilla).     

Arbuscular mycorrhizal colonization of halophytes in Central European salt marshes

 Halophytes from both coastal and inland Central European salt marshes were examined for colonization by arbuscular mycorrhizal (AM) fungi. Plants from different families were strongly colonized but the degree of colonization varied with the individual plant and apparently during the vegetation period, too. Members of the typical non-mycorrhizal families like Armeria maritima of the Plumbaginaceae and Salicornia europaea of the Chenopodiaceae were found to be colonized, particularly in the drier salt marshes. High numbers of Glomus spores were found in the saline soils, especially those of the inland locations examined. Approximately 80% of these spores were from Glomus geosporum as shown by a typical restriction fragment length polymorphism (RFLP) pattern of the amplified internal transcribed spacer regions. The present study demonstrates that RFLP analysis is useful when screening habitats for the occurrence of mycorrhizal fungi which can be identified only with difficulty by morphological criteria. Accepted: 25 August 2000     

Trematodes associated with mangrove habitat in Puerto Rican salt marshes

Batillaria minima is a common snail in the coastal estuaries of Puerto Rico. This snail is host to a variety of trematodes, the most common being Cercaria caribbea XXXI, a microphallid species that uses crabs as second intermediate hosts. The prevalence of infection was higher (7.1%) near mangroves than on mudflats away from mangroves (1.4%). Similarly, there was a significant positive association between the proportion of a site covered with mangroves and the prevalence of the microphallid. The association between mangroves and higher trematode prevalence is most likely because birds use mangroves as perch sites and this results in local transmission to snails.     

Ecophysiological adaptations of coastal halophytes from foredunes and salt marshes

Ecophysiological strategies of coastal halophytes from foredunes and salt marshes are discussed. A comparison is made of the factors that limit growth in salt marshes and sand dunes. In salt marshes, zonation and succession are primarily governed by variation in soil salinity, which strongly depends on inundation with seawater. Results are described of experiments which aim at separating salinity and inundation effects on growth, osmotic and mineral relations in a comparison of salt-marsh halophytes. The growth response of plants cannot simply be correlated (and causally explained) with the concentration of Na, Cl, and K in the tissues. Also, the compatible osmotic solutes proline and methylated quaternary ammonium compounds may accumulate both in species with a positive response to increased salinity and in species with a growth reduction under seawater inundation. More likely inadequate adaptation of the plants water potential with these components is partly the cause of retarded growth. Disfunctioning of the plant in this respect may be at three levels: (a) total water potential of the plant, (b) (loss) of turgor pressure potential; (c) regulation at the cellular level. The ecological importance of some factors in seawater other than sodium chloride is considered. In coastal sand dunes airborne rather than soil salinity limits plant growth, together with the effects of abrasion, sand accretion, drought and the poor nutrient status of the dune sand. Adaptations of sand-dune species to these factors may consist of: large seeds with storage tissue germinating in the dark and seedling growth enough to emerge through the accreted sand. Aerial parts must be resistant to mechanical damage (high wind speed and abrasion), possibly by a sclerophyllous and tough structure. Efficient nutrient uptake, translocation and retranslocation seem to help survive sand-dune species in a nutrient-poor rooting medium.     

Latitudinal variation in plant–herbivore interactions in European salt marshes

Ecological interactions often vary geographically. Work in salt marshes on the Atlantic Coast of the United States has documented community-wide latitudinal gradients in plant palatability and plant traits that may be driven in part by greater herbivore pressure at low latitudes. To determine if similar patterns exist elsewhere, we studied six taxa of saltmarsh plants ( Atriplex , Juncus , Limonium , Salicornia , Spartina and Suaeda ) at European sites at high (Germany and the Netherlands) and low (Portugal and Spain) latitudes. We conducted feeding assays using both native and non-native consumers, and documented patterns of herbivore damage in the field. As in the United States, high-latitude plants tended to be more palatable than low-latitude plants when offered to consumers in paired feeding assays in the laboratory, although assays with grasshopper consumers were less consistent than those with crab consumers, and plants in the field at low-latitude sites tended to experience greater levels of herbivore pressure than plants at high-latitude sites. Similarly, high-latitude leaf litter was more palatable than litter from low-latitude plants when offered to consumers in paired feeding assays in the laboratory. Latitudinal gradients in plant palatability and herbivore pressure may be a general phenomenon, and may contribute to latitudinal gradients in decomposition processes.