Phylogeography of the Black-tailed Godwit Limosa limosa: substructuring revealed by mtDNA control region sequences

Black-tailed (Limosa limosa) and Hudsonian Godwits (L. haemastica) are sometimes described as a superspecies. The Black-tailed Godwit is further split into three subspecies on the basis of morphological differences (L. l. limosa, L. l. islandica and L. l. melanuroides). We studied variation in partial mtDNA control region sequences among Black-tailed and Hudsonian Godwits which showed 5% divergence. Black-tailed and Hudsonian Godwits were thus clearly differentiated and the separate species status for the two taxa is validated. All three subspecies described for the Black-tailed Godwit had unique haplotypes but the genetic distances were small (0.3–0.6%). Despite small genetic differences we could not detect any substantial gene flow between any of the subspecies as haplotypes were private to each subspecies. Thus, genetic variation within Black-tailed Godwits showed a clear geographic structure. We found a high proportion of rare private haplotypes in three fringe populations of the nominate subspecies of the Black-tailed Godwit (L. l. limosa) where godwits breed in low numbers, but no genetic variation at all in a sample from the Netherlands where godwits are abundant. This suggests that Dutch Godwits may have been affected by a founder effect.     

Effects of green algal mats on bivalves in a New England mud flat

Concurrent with the spread of green algal mats on tidal flats, reports of macrofauna dieoffs under dense algal mats have increased in numbers. Bivalves seem to be particularly affected by persistent dense algal mats. Bivalve species with a long extendible siphon seem to be less affected underneath algal mats, but no distinction has been made in the past between species with short and those with long siphons,Mya arenaria andMacoma balthica, on an intertidal mudflat in New England. Abundances ofM. arenaria declined substantially during the study period when a thick green algal mat covered the mudflat for several months. Numbers of the small bivalveGemma gemma also decreased substantially, whereas abundances ofM. balthica showed minimal variation during the time of algal coverage. In algae removal/addition experiments numbers ofM. arenaria decreased, but effects were only significant in an algal addition to previously algal-free mudflat areas. Abundance ofM. balthica did not change significantly in the algal removal/additition experiments. Over the time period of the experiment (9 weeks),M. arenaria showed measurable size increase in uncovered mudflat areas, but not underneath algal mats. Similarly,M. balthica only increased in size in the uncovered mudflat area. From these results it is concluded thatM. balthica can survive time periods of dense algal coverage because it is able to penetrate through the algal mat with its long extendible siphon, and thus can reach well-oxygenated water layers above the mat.M. arenaria with its thick, less extendible, siphon cannot push through dense algal mats and therefore is more likely to die underneath persistent algal mats.     

Trends in breeding phenology across ten decades show varying adjustments to environmental changes in four wader species

Capsule: During 1928–2016, initiation of egg-laying advanced in two wader species, remained unchanged in one, and was delayed in one species. The changes across years and variation among species can be explained by climatic variables and differences in migratory strategies.

Aims: To document possible changes in initiation of egg-laying in common Danish wader species since the early part of the 20th century and seek possible correlations between egg-laying, timing of arrival and environmental factors.

Methods: Annual records of the first eggs and chicks found on the scientific reserve of Tipperne in western Denmark 1928–2016 were analysed using linear regression to determine patterns in timing of egg-laying, pre-breeding length and influence of climate factors.

Results: Two short/medium-distance migrant wader species, Northern Lapwing Vanellus vanellus and Common Redshank Tringa totanus advanced breeding initiation by about one week, with winter North Atlantic oscillation Index and spring temperature as important predictors. By contrast, two long-distance migrants, Black-tailed Godwit Limosa limosa and Ruff Calidris pugnax, did not advance egg-laying, and Ruff actually delaying it. As a result, the pre-laying period was significantly prolonged in both Black-tailed Godwit (21 days) and Ruff (52 days), while there was no significant change for Common Redshank.

Conclusion: Long-distance migrants are able to adjust spring arrival but unlike short/medium-distance migrants, do not necessarily adjust breeding initiation.     

Experimental disturbance by walkers affects behaviour and territory density of nesting Black-tailed Godwit Limosa limosa

In Europe, the number of areas supporting breeding Black-tailed Godwit Limosa limosa has halved over the last 30 years. Although the decline has been primarily attributed to habitat deterioration, human disturbance has also been implicated. We undertook a controlled experimental study at the Danish Special Protection Area (SPA) reserve at Tipperne, comparing bird behaviour and breeding densities from two baseline years with those in 3 years with two experimental levels of disturbance. Black-tailed Godwits flushed and showed mobbing behaviour significantly more often when disturbed. The duration of simultaneous flights by breeding pairs was greater when disturbed, leaving nests susceptible to predation. Behavioural observations suggested birds were highly sensitive to human disturbance and unlikely to habituate. Disturbance levels of seven walkers/day affected territory densities up to 500 m from routes taken by walkers, causing effective habitat loss to breeding Black-tailed Godwits. The species' sensitivity to disturbance may help explain why it has disappeared from many areas. Effective conservation of important breeding areas and maintenance of high densities of Black-tailed Godwit and other meadow birds necessitates control of public disturbance to breeding areas.     

No evident spatial genetic structuring in the rapidly declining Black-tailed Godwit <Emphasis Type="Italic">Limosa limosa</Emphasis><Emphasis Type="Italic">limosa</Emphasis> in The Netherlands

With 40% of the European Black-tailed Godwit population breeding in The Netherlands, this country harbours internationally significant numbers of this species. However, ongoing agricultural intensification has resulted in the fragmentation of the population and drastic population declines since 1967. By establishing genetic diversity, genetic differentiation and gene flow on the basis of 12 microsatellites, we investigated whether the population genetic structure of the Dutch Black-tailed Godwit bears the marks of these changes. Genetic diversity appeared to be moderate, and Bayesian model-based analysis of individual genotypes revealed no clustering in the Dutch populations. This was supported by pairwise F_ST values and AMOVA, which indicated no differentiation among the nine breeding areas. Gene flow estimates were larger than “one migrant per generation” between sample locations, and no isolation by distance was demonstrated. Our results indicate the maintenance of moderate levels of genetic diversity and genetic connectivity between breeding sites throughout the Dutch Black-tailed Godwit breeding population. We suggest that the Dutch Black-tailed Godwit breeding areas should be managed as a single panmictic unit, much as it is presently done.     

Effects of small‐scale environmental variation on metaphyton condition and community composition

1. Metaphyton mats typically originate as benthic algal biofilms that experience higher solar radiation and temperatures, and reduced access to nutrients, once they reach the water surface, but the impacts of these physicochemical changes on metaphyton condition and community composition have received little attention. 2. Using microprobes positioned at 0, 2, 4 and 6 cm depth, we recorded small‐scale differences in water chemistry within metaphyton mats constrained in floating nets during an 8‐week period. Concurrent weekly samples of filamentous algae and their diatom epiphytes were collected from the top, middle and bottom of the mats and were analysed for changes in ash‐free dry mass (AFDM) and chlorophyll‐a, nutrient (N, P, C, Si) content and taxonomic composition. 3. Light intensity, temperature and dissolved oxygen declined both with increasing depth in the mat and over the study period. The autotrophic index (=AFDM/chlorophyll‐a) was greatest at the top of the mats and increased over time; samples also had higher C/P and C/N ratios than samples deeper within the mat. 4. Pithophora was consistently the dominant algal filament throughout the study (representing 85% of all filaments averaged over time and depth); epiphytic diatom cover on Pithophora (calculated as epiphyte area index) declined over time, particularly at the top of the mat. 5. Densities of the diatom epiphytes Gomphonema, Cocconeis and Fragilaria increased with increasing depth within the mat, whereas Cymbella/Encyonema was more common in surface samples. 6. Our results indicate that metaphyton mats are highly dynamic communities, spatially organised in part by small‐scale environmental variation and subject to changes in taxonomic composition following their arrival at the water surface.     

Influence of the Thin Drift Algal Mats on the Distribution of Macrozoobenthos in Kõiguste Bay, NE Baltic Sea

Macrozoobenthic communities within and outside of the drift algal mats were compared in Kõiguste Bay, NE Baltic Sea. The patches of the drift algae were on average 0.5–1 km wide in diameter covering about 25% of the total bottom area of the bay. Thickness of the mat did not exceed 6 cm. The biomass of the mat varied between 35 and 1391 g dw m^?2. The drift algal mats had no clear negative effect on macrozoobenthos except for a few infaunal species. The drift algae favoured several detrivorous, herbivorous and carnivorous species. Among the studied variables, the thickness of algal mat and oxygen concentration at near-bottom layer explained the best the structure of macrozoobenthos. Total number of invertebrate species increased curvilinearly with the thickness of algal mat having the peak value at 3–5 cm thick algal mat. To conclude, moderate drift algal mats increased habitat complexity and, thus, the diversity of benthic faunal assemblages in otherwise poorly vegetated coastal areas.     

Sex differences in digestive traits in sexually size-dimorphic birds: Insights from an assimilation efficiency experiment on Black-tailed Godwit

Digestive assimilation efficiency is considered a trait with important implications for animal ecology. However, practically all studies have ignored the importance of sex differences in food assimilation efficiency (AE). Here, we investigated sex differences in dietary and physiological parameters in the Black-tailed Godwit Limosa limosa limosa feeding on rice seeds, a species with sexual dimorphism in body size and body mass. Gross daily food intake, gross energy intake, gross energy output and metabolizable energy intake did not vary significantly between sexes, but godwit females showed lower faeces energy density and higher AE than males. Mass-specific AE was similar in males and females, and the difference in AE could be attributed to the females' greater body mass. We suggest that a differential AE could play a role in explaining sex differences in habitat or micro-habitat selection during the non-breeding season in bird species with sexual dimorphism in size. Finally, we addressed the question about assimilation efficiency accuracy in models that estimate prey acquisitions by declining shorebirds as the Black-tailed Godwit.     

THE WINTER FEEDING ECOLOGY OF THE REDSHANK TRINGA TOTANUS

The aim of the work was to find out how the Redshank which over-winter on the Ythan estuary, Aberdeenshire, adapt their feeding activities to the short daylength in winter. To achieve this, the feeding behaviour and daily routine in winter were compared with those in autumn and spring. Corophium volutator was the main prey on the estuary during the day. However, the temperature of the mud greatly affected the diet in the two areas where the feeding behaviour was studied in detail. At temperatures above 6°C, most of the biomass ingested consisted of Corophium. However, in one study area in one winter, Macoma balthica was taken more frequently at lower than at high temperatures. In the other area, Nereis diversicolor was taken more frequently at low than at high temperatures. These changes in diet appeared related to changes in the behaviour of the prey affecting their availability and not due to the birds changing their preference at low temperatures. The ingestion rate was not affected by the change in diet in the first area but decreased at low temperatures in the second. The sizes of Macoma and Nereis taken by the birds did not vary seasonally. The sizes of Corophium taken decreased in winter as a result of a reduction in the size of those present in the substrate. It was concluded that there was no evidence of Redshank increasing their ingestion rate in winter to compensate for the short daylength. The proportion of the time spent feeding on the estuary in daylight was greater in winter than in spring. Redshank continued feeding at high water in the surrounding fields and on the estuary at night during the winter but not in either autumn or spring. During the winter they obtained less than 50% of their daily food requirements from the estuary in daylight, mainly because of the short daylength. Consequently, they had to collect the balance at night and at high water. Seasonal changes in the numbers of Redshank were recorded. After an autumn decrease in numbers, no decrease in numbers could be detected during the winter. These findings are discussed in relation to the possible difficulties for Redshank in collecting their daily food requirements during the winter. The impact of the birds on their main estuarine prey Corophium is also discussed.     

Interactions between macro-algal mats and invertebrates in the Ythan estuary, Aberdeenshire, Scotland

Blooms of opportunistic green macro-algae are a common feature of coastal areas and their effects on mudflat invertebrates can be dramatic. On the Ythan estuary, Aberdeenshire, Scotland, we have carried out a number of manipulative field experiments designed to evaluate the effects on invertebrates of different species of macro-algae with contrasting ecologies, and the effects of invertebrates on the development of the blooms. Macro-algal mats were found to have dramatic nega- tive effects on the density of the amphipod Corophium volutator, with higher algal biomasses having greater impact. The mechanism for this interaction seems to be interference by the algal filaments with the feeding behaviour of the amphipod. In contrast, the polychaete Capitella spp. increases in abundance under macro-algal mats due to enrichment of the sediment with organic material. These two interactions are seen at all scales, in areas of less than 1 m² to the scale of the entire estuary, irrespective of the species composition of the macro- algal mats. Bioturbation by Corophium and grazing by the snail Hydrobia ulvae had little effect on macro-algal biomass, but there were less algae when the polychaete Nereis diversicolor was present. The most significant interaction in this system is the pronounced negative impact of algal mats on the abundance of Corophium, probably the most important invertebrate species in the diets of the estuary’s shorebirds, fish and epibenthic crustaceans. Received: 14 December 1998 / Received in revised form: 26 May 1999 / Accepted: 28 May 1999     

REMOVAL OF ORGANIC AND INORGANIC MATTER FROM ANTARCTIC LAKES BY AERIAL ESCAPE OF BLUEGREEN ALGAL MATS1

Field studies on three perennially ice-covered Antarctic oasis lakes with little or no outflow disclosed a unique biological phenomenon. Benthic algal mats dominated by the prokaryotic Phormidium frigidum Fritsch and several pennate diatoms growing in shallower, more brightly illuminated areas beneath 4 to 5.5 m of ice accumulate and entrap bubbles of photosynthetically produced oxygen. Clumps of this gas-filled mat tear loose from the gravelly substrate, lift off and float to the bottom of the lake ice. Some of these floating mat pieces become frozen into newly forming ice with the onset of winter. Through the combination of ablation of ice from the upper surface and the formation of new ice from below, algal mat pieces reach the upper lake surface in 5–10 years. Here, they are lyophilized by polar winds and dispersed in at least a partially viable state. The process of mat lift-off and escape is important in removing nutrients and salts from these lakes and helps to perpetuate their oligotrophic state. Neutron activation and X-ray dispersive analyses of elements in the algal mats along with other analyses, field observations, and calculations suggest that significant quantities of organic matter, select minerals and salts are lost from the lakes annually through this process whose magnitude has not been recognized previously.     

BLUE-GREEN ALGAL MATS IN A SMALL STREAM1

Bedrock erosional features in a small stream (Little Schultz Creek, Bibb County, Alabama) created a variety of habitats for epilithic growth. One suck habitat was illustrated by the occurrence of small falls (<0.3 m) in the main channel of the stream and blue-green algal mats associated with them. The cohesive, laminar algal mats were found at 15 such sites along a 250-m reach of the stream. The primary mat matrix consisted of the blue-green alga Oscillatoria submembranacea Ardissone and Strafforella. The uppermost portion of each mat consisted of a thin (<1 mm thick) green layer of biologically active filaments. The lower layers were thicker (up to 2 cm thick) and consisted of brown laminae of Oscillatoria filaments, and associated sediments. In addition, numerous diatoms mere associated with the mat surface. Some were loosely attached (e.g. Achnanthes); others (Cymbella tumida (Bréb.) V. H.) were stalked. These mats were present throughout the year and showed a bimodal annual distribution with maxima hi February and July. In February, total mat coverage was higher than in July. This winter maximum may have been related to a mode of growth dependent upon sedimentation from storm events and subsequent upward growth of the alga. Mat primary productivity on an areal basis (432 mg C · m^?2· d^?1 in March and 907 mg C · M^?2· d^?1 in April) was 2–12 times the maxima measured on epizoic and cobbles surfaces and other bedrock surfaces in the same stream. The limited areal coverage of the mats, when compared to other surfaces available for algal colonization, made them less important than other epilithic and epizoic surfaces in terms of total primary production in this stream reach. However, we propose that the combination of their unique structure and high primary productivity may make these algal mats sites of high algal and bacterial metabolic activity, which may include anaerobic processes in midchannel, where such activity would not be expected to occur.     

Ecological studies of Chloroflexis,a gliding photosynthetic bacterium

Summary Chloroflexis, a gliding, filamentous, photosynthetic bacterium, is present in the stratified algal-bacterial mats which occur in the 50°–70°C temperature range of alkaline hot spring effluents. The organism is in association with the alga in the upper, algal layer, and also forms thick, orange mats beneath the algal layer. Natural populations of Chloroflexis from these mats demonstrated light-stimulated uptake of some ¹⁴C-labelled organic compounds. Photosynthetic ¹⁴CO₂ fixation by natural samples of Chloroflexis was investigated with respect to temperature, light intensity and mat depth. Bacterial photosynthesis was determined in samples in which algae were present by use of the inhibitor 3-(3,4-dichlorophenyl)-1,1-dimethylurea (DCMU). Bacterial photosynthesis was maximal at depths down to about 3 mm and then decreased rapidly to very low levels at greater depths. The greatest amounts of bacteriochlorophyll pigments were also concentrated in the top 3–4 mm of the mat. The optimum light intensity for bacterial photosynthesis (about 400 ft-c) was considerably lower than the normal summer light intensity at the surface of the mat (5000-8000 ft-c).The temperature optima for photosynthesis by the bacterial component of natural mat samples from several sites of different temperatures in a hot spring thermal gradient were determined. Temperature optima approximated the environmental temperatures, indicative of the occurrence of strains of Chloroflexis adapted to different temperatures. Although bacterial standing crop was greatest in the temperature range 50°–55°C, maximum photosynthetic efficiency was observed at about 45°C. Sulfide was stimulatory to photosynthetic ¹⁴CO₂ fixation by naturally occurring populations of Chloroflexis under field conditions. These data are consistent with the hypothesis that Chloroflexis may utilize sulfide as an electron donor for photosynthetic CO₂ reduction. However, it is also likely that Chloroflexis grows photoheterotrophically in these mats, obtaining organic compounds from algal excretory products.     

洱海漂浮草垫的物种组成及分布

研究通过调查洱海2017年夏季和冬季漂浮草垫的物种组成及分布模式, 探究洱海在水位上升过程中, 哪些挺水植物更易形成漂浮草垫而存活下来。夏季共调查到漂浮草垫18个, 植物分属15科, 18属, 24种, 冬季共调查到漂浮草垫22个, 植物分属13科, 17属, 22种, 形成洱海漂浮草垫的植物共15科, 19属, 26个物种。草垫下水体中总氮(TN)、总磷(TP)、溶解性总磷(TDP)都明显高于开阔水域的(P<0.05), 而开阔水域的溶氧(DO)明显高于草垫下水体的(P<0.05)。结果表明在夏季和冬季漂浮草垫面积大小与其组成草垫的物种数和漂浮草垫生物量都呈显著正相关(P<0.01), 而在夏季和冬季漂浮草垫面积大小与漂浮草垫单位面积平均生物量相关性都不显著(P>0.05), 漂浮草垫面积大小与漂浮草垫下最长根在夏季相关性显著(P<0.01), 而在冬季相关性不显著(P>0.05)。漂浮草垫主要分布在离岸距离60 m以内以及水深不超过2 m的水域, 漂浮草垫面积大部分都小于600 m2(夏季占87%, 冬季占95%), 每个漂浮草垫的物种数都小于10个, 漂浮草垫下最长根主要分布在40—120 cm。组成漂浮草垫的物种中菰(Zizania latifolia)的频度和相对生物量在夏季和冬季(频度: 夏季-73.33%, 冬季-66.67%; 生物量: 夏季-43.38%, 冬季-41.91%)都是最高的, 说明菰相比其他挺水植物更容易形成漂浮草垫从而避免洱海水位上升对其生长和繁殖的影响, 这可能是菰成为洱海单优挺水植物的主要原因。而菰比其他挺水植物更容易形成漂浮草垫的机制则需要更进一步的研究。     

Soil stabilization by a prokaryotic desert crust: Implications for Precambrian land biota

A cyanophyte dominated mat, desert crust, forms the ground cover in areas measuring hundreds of square meters in Utah and smaller patches in Colorado. The algal mat shows stromatolitic features such as sediment trapping and accretion, a convoluted surface, and polygonal cracking. Sand and clay particles are immobilized by a dense network of filaments of the two dominating cyanophyte species,Microcoleus vaginatus andM. chthonoplastes, which secrete sheaths to which particles adhere. These microorganisms can tolerate long periods of desiccation and are capable of instant reactivation and migration following wetting. Migration occurs in two events: 1. immediately following wetting of dry mat, trichomes are mechanically expelled from the sheath as it swells during rehydration, and 2. subsequently, trichomes begin a self-propelled gliding motility which is accompanied by further production of sheath. The maximum distance traveled on solid agar by trichomes ofMicrocoleus vaginatus during a 12 hour period of light was 4.8 cm. This corresponds to approximately 500 times the length of the fastest trichome, and provides a measure of the potential for spreading of the mat in nature via the motility of the trichomes.Dehydration resistence of the sheath modifies the extracellular environment of the trichomes and enables their transition to dormancy. Following prolonged wetting and evaporative drying of the mat in the laboratory, a smooth wafer-like crust is formed by the sheaths ofMicrocleus trichomes that have migrated to the surface. Calcium carbonate precipitates among the algal filaments under experimental conditions, indicating a potential for mat lithification and fossilization in the form of a caliche crust. It is suggested that limestones containing tubular microfossils may, in part, be of such an origin.The formation of mature Precambrian soils may be attributable to soil accretion, stabilization, and biogenic modification by blue-green algal land mats similar to desert crust.     

Dynamics of benthic algal vegetation and environment in Dutch estuarine salt marshes studied by means of permanent quadrats

Summary Knowledge of the spatial structure of benthic algal vegetations in salt marshes, consisting of about 100 species of blue-green, green, brown and red algae, formed the basis of a detailed study into the dynamics of the algal mat.The temporal changes in algal vegetations and concomitant processes in their environment were studied in 27 permanent quadrats (PQs), plotted in tidal salt marshes and along brackish inland waters in the SW Netherlands. Once a month vegetation and environment of the PQs were examined over the period March 1968-February 1971. From the Spearman rank-correlation between vegetational and environmental parameters it appeared that in semiterrestrial and terrestrial environments the average number of algal species in a relevé and the average similarity between the species composition of successive relevés obtained from one PQ have a significant positive correlation with the stability of the substrate, and hardly any correlation with fluetuations in soil-moisture content and salinity of the soil moisture and with the maximum cover percentage of the phanerogams.Stable substrates tend to bear stable algal mats, as appeared from quantitative data. Small and moderate fluctuations in soil-moisture content and salinity of the soil moisture have no limiting influence on the algal cover degree. During periods of extremely high salinity and concomitant extremely low soil-moisture contents that coincide with evaporation surpluses, the green algal mat bleaches and decreases in cover in semi-terrestrial and terrestrial environments. Below 20% the soil-moisture content is limiting for the expansion of a mat of green algae (Rhizoclonium riparium, Percursaria percursa, Enteromorpha torta, E. prolifera) and Vaucheria species. The mat desiceates and bleaches but recovers after increase of the soil-moisture content above 20% within one month. Two weeks of severe frost (temperature down to – 10°C) do not influence the cover degree of the algal mat in aquatic branckish PQs, but the cover degree in semi-terrestrial PQs decreases. Short lasting soil-moisture salinities of 130–190 Cl^– have no limiting effect on the expansion of the algal mat.Nomenclature authorities are given in Table 3.Contribution to the Symposium of the Working Group for Succession Research on Permanent Plots, held at Yerseke, The Netherlands, October 1975.Communication No. 163 of the Delta Institute for Hydrobiological Research, Yerseke.     

Identifying new buffer areas for conserving waterbirds in the Mediterranean basin: the importance of the rice fields in Extremadura,Spain

The impact of wetland loss on migratory waterbirds can be mitigated by the presence of anthropogenic habitats such as rice fields. In the Mediterranean basin, wetlands have been drained and altered to such a degree that their very existence is threatened. It is, therefore, essential to identify key buffer areas in the basin to develop conservation strategies for migratory waterbirds. In Extremadura, continental SW Spain, 30,000 ha of new rice fields have created since the 1960s. The present study describes for the first time the waterbird community associated with these inland rice fields, and addresses the question of the potential value of this area in buffering the loss of natural wetlands for populations of waterbirds using the East Atlantic Flyway (EAF). The waterbird community studied consisted of 45 species, with population levels reaching values of international importance (>1% of the biogeographical population using EAF) for the Common Crane Grus grus (19%), Black-tailed Godwit Limosa limosa (1.2%), Black-winged Stilt Himantopus himantopus (1.2%), and Cattle Egret Bubulcus ibis (1%). The presence of these rice fields may be modifying the wintering and/or feeding sites of some waterbird species such as the Common Crane and Black-tailed Godwit. Unfortunately, there is no legal protection for this area. We propose the designation of these rice fields as a Special Protection Area for birds.     

The significance of spatial and temporal patterns of algal mat deposition in structuring salt marsh vegetation

Question: Are there hot spots of algal mat deposition in space and time at the marsh scale and, if so, how does this affect the coexistence of a dominant (Spartina anglica) and gap dependent (Salicornia europaea) species? Location: The Rattekaai salt marsh in the Scheldt estuary in the southwestern Netherlands (NW Europe). Methods: Mat cover and the abundance of the gap dependent species Salicornia europaea were monitored at the scale of a marsh. The effects of mat cover on the vegetation structure were studied by applying three mat removal treatments over three growing seasons. Results: The low marsh border was found to be a hot spot of algal mat deposition during the growing season, which had a correlated spatial pattern between two successive years at a 20 m X 20 m scale. The combination of duration, timing and repetition of mat cover determined growth inhibition of the competitive dominant Spartina anglica, and thereby the abundance of subordinates such as Salicornia europaea. Mat cover reduced the storage of carbon reserves in Spartina and our results imply that repetition of non‐lethal mat cover can lead to ‘gap creation’. Gaps gave only temporary habitat to less dominant species since Spartina quickly re‐invaded them. The gap dependent annual Salicornia was most abundant at intermediate levels of disturbance measured as a function of both space and time. Conclusions In addition to disturbance level, the spatial and temporal distribution of disturbance are important in creating and maintaining habitat for gap dependent species. Relatively small disturbances will have a large effect on diversity if the spatial and temporal distribution of the disturbances leads to ‘disturbance hot spots’.     

Effects of grass field margin management on food availability for Black-tailed Godwit chicks

Over the last six decades, populations of wader species like the Black-tailed Godwit (Limosa limosa) have sharply decreased in the Netherlands. Agricultural intensification has led to reduced habitat quality for meadow birds. As a consequence, reproductive success has declined. One of the main drivers of this decline in reproductive success is reduced food availability for meadow bird chicks. Agri-environment schemes (AES), designed to halt this decline, have so far been insufficient. Most of these AES focus on entire fields, but recent research suggests that differences in suitability exist within fields. Grass field margins may be more suitable for meadow bird chicks than the center of intensively managed grass lands. To improve existing meadow bird AES it could be beneficial to implement additional management in field margins of intensively managed grass fields. An already existing type of field margin AES with additional management is the botanical field margin. Here, we evaluate four different types of field margin management, including botanical field margins, focusing on aerial insects (an important part of the diet of Black-tailed Godwit chicks and Redshank chicks) in field centers and margins. Grass field margins contained more large aerial insects than field centers and, more importantly, additional management of the grass field margin increased the number of aerial insects in the margin. We conclude that combining meadow bird AES and botanical field margin management may enhance meadow bird food availability and improve the efficacy of AES.