Spring arrival response to climate change in birds: a case study from eastern Europe

This paper analyses the dependence of the first spring arrival dates of short/medium- and long-distance migrant bird species on climate warming in eastern Europe. The timing of arrival of the selected species at the observation site correlates with the North Atlantic Oscillation (NAO) index, air temperature, atmospheric pressure, precipitation and wind characteristics. A positive correlation of fluctuations in winter and spring air temperatures with variations in the NAO index has been established in eastern Europe. Positive winter NAO index values are related to earlier spring arrival of birds in the eastern Baltic region and vice versa—arrival is late when the NAO index is negative. The impact of climate warming on the bird’s life cycle depends on local or regional climate characteristics. We tested the hypothesis that differences in climate indices between North Africa and Europe can influence the timing of spring arrival. Our results support the hypothesis that differences in first spring arrival dates between European populations occur after individuals cross the Sahara. We assume that the endogenous programme of migration control in short/medium-distance migrants synchronises with the changing environment on their wintering grounds and along their migration routes, whereas in long-distance migrants it is rather with environmental changes in the second part of their migratory route in Europe. Our results strongly indicate that the mechanism of dynamic balance in the interaction between the endogenous regulatory programme and environmental factors determines the pattern of spring arrival, as well as migration timing.     

Climate change, breeding date and nestling diet: how temperature differentially affects seasonal changes in pied flycatcher diet depending on habitat variation

1.?Climate warming has led to shifts in the seasonal timing of species. These shifts can differ across trophic levels, and as a result, predator phenology can get out of synchrony with prey phenology. This can have major consequences for predators such as population declines owing to low reproductive success. However, such trophic interactions are likely to differ between habitats, resulting in differential susceptibility of populations to increases in spring temperatures. A mismatch between breeding phenology and food abundance might be mitigated by dietary changes, but few studies have investigated this phenomenon. Here, we present data on nestling diets of nine different populations of pied flycatchers Ficedula hypoleuca, across their breeding range. This species has been shown to adjust its breeding phenology to local climate change, but sometimes insufficiently relative to the phenology of their presumed major prey: Lepidoptera larvae. In spring, such larvae have a pronounced peak in oak habitats, but to a much lesser extent in coniferous and other deciduous habitats. 2.?We found strong seasonal declines in the proportions of caterpillars in the diet only for oak habitats, and not for the other forest types. The seasonal decline in oak habitats was most strongly observed in warmer years, indicating that potential mismatches were stronger in warmer years. However, in coniferous and other habitats, no such effect of spring temperature was found. 3.?Chicks reached somewhat higher weights in broods provided with higher proportions of caterpillars, supporting the notion that caterpillars are an important food source and that the temporal match with the caterpillar peak may represent an important component of reproductive success. 4.?We suggest that pied flycatchers breeding in oak habitats have greater need to adjust timing of breeding to rising spring temperatures, because of the strong seasonality in their food. Such between-habitat differences can have important consequences for population dynamics and should be taken into account in studies on phenotypic plasticity and adaptation to climate change.     

Consequences of the settlement of migrant European Robins Erithacus rubecula in wintering habitats occupied by conspecific residents

In wintering areas where migrant birds meet sedentary conspecifics, early settlement of local residents in the best habitat patches might reduce the availability of suitable sites for arriving migrants. We studied how sympatric migratory and sedentary European Robins Erithacus rubecula occupy two wintering habitats of different quality (forests and shrublands) in southern Spain, and how such a distribution affects individuals of each population sector. In September, before migrants arrived, Robins were only found in forests, and they had already saturated these habitats, so that rather than increasing Robin abundance in these habitats, the arrival of migrants caused a massive occupation of the previously vacant shrublands. During the winter, we captured Robins and identified them as migrants or residents using a discriminant function based on morphological traits. Residents always predominated in forests, and migrants in shrublands, but through the winter around 35% of residents (mainly juveniles) moved to shrublands, having been replaced by some migrants in forests. Although food was more abundant in shrublands, Robins had better body condition in forests, suggesting that other factors determined habitat preferences (e.g. shelter availability or food diversity, which were higher in forests). In addition, we observed a greater variance in body mass relative to body size in forests, suggesting that energy management was less constrained in this habitat (for example owing to a lower exposure to predators or a higher food predictability). Our results suggest that sedentary Robins benefit from an early occupation of the best habitats in the wintering grounds, forcing migrants to colonize apparently less suitable sites. This would explain the persistence of these small southern populations despite the yearly flooding of the area by huge numbers of migrant conspecifics.     

Climate and the complexity of migratory phenology: sexes,migratory distance,and arrival distributions

The intra- and inter-season complexity of bird migration has received limited attention in climatic change research. Our phenological analysis of 22 species collected in Chicago, USA, (1979–2002) evaluates the relationship between multi-scalar climate variables and differences (1) in arrival timing between sexes, (2) in arrival distributions among species, and (3) between spring and fall migration. The early migratory period for earliest arriving species (i.e., short-distance migrants) and earliest arriving individuals of a species (i.e., males) most frequently correlate with climate variables. Compared to long-distance migrant species, four times as many short-distance migrants correlate with spring temperature, while 8 of 11 (73%) of long-distance migrant species’ arrival is correlated with the North Atlantic Oscillation (NAO). While migratory phenology has been correlated with NAO in Europe, we believe that this is the first documentation of a significant association in North America. Geographically proximate conditions apparently influence migratory timing for short-distance migrants while continental-scale climate (e.g., NAO) seemingly influences the phenology of Neotropical migrants. The preponderance of climate correlations is with the early migratory period, not the median of arrival, suggesting that early spring conditions constrain the onset or rate of migration for some species. The seasonal arrival distribution provides considerable information about migratory passage beyond what is apparent from statistical analyses of phenology. A relationship between climate and fall phenology is not detected at this location. Analysis of the within-season complexity of migration, including multiple metrics of arrival, is essential to detect species’ responses to changing climate as well as evaluate the underlying biological mechanisms.     

Rapid change in host use of the common cuckoo Cuculus canorus linked to climate change

Parasites require synchrony with their hosts so if host timing changes with climate change, some parasites may decline and eventually go extinct. Residents and short-distance migrant hosts of the brood parasitic common cuckoo, Cuculus canorus, have advanced their phenology in response to climate change more than long-distance migrants, including the cuckoo itself. Because different parts of Europe show different degrees of climate change, we predicted that use of residents or short-distance migrants as hosts should have declined in areas with greater increase in spring temperature. Comparing relative frequency of parasitism of the two host categories in 23 European countries before and after 1990, when spring temperatures in many areas had started to increase, we found that relative parasitism of residents and short-distance migrants decreased. This change in host use was positively related to increase in spring temperature, consistent with the prediction that relative change in phenology for different migrant classes drives host-use patterns. These findings are consistent with the hypothesis that climate change affects the relative abundance of different host races of the common cuckoo.     

Climate change and fitness components of a migratory bird breeding in the Mediterranean region

The increase in spring temperatures in temperate regions over the last two decades has led to an advancing spring phenology, and most resident birds have responded to it by advancing their onset of breeding. The pied flycatcher (Ficedula hypoleuca) is a long‐distance migrant bird with a relatively late onset of breeding with respect to both resident birds and spring phenology in Europe. In the present correlational study, we show that some fitness components of pied flycatchers are suffering from climate change in two of the southernmost European breeding populations. In both montane study areas, temperature during May increased between 1980 and 2000 and an advancement of oak leafing was detected by using the normalized difference vegetation index (NDVI) to assess tree phenology. This might result in an advancement of the peak in availability of caterpillars, the main prey during the nestling stage. Over the past 18 yr, the time of egg laying and clutch size of pied flycatchers were not affected by the increase in spring temperatures in these Mediterranean populations. However, this increase seems to have an adverse effect on the reproductive output of pied flycatchers over the same period. Our data suggest that the mismatch between the timing of peak food supply and nestling demand caused by recent climate change might result in a reduction of parental energy expenditure that is reflected in a reduction of nestling growth and survival of fledged young in our study populations. The data seem to indicate that the breeding season has not shifted and it is the environment that has shifted away from the timing of the pied flycatcher breeding season. Mediterranean pied flycatchers were not able to advance their onset of breeding, probably, because they are constrained by their late arrival date and their restricted high altitude breeding habitat selection near the southern border of their range.     

Climate change effects on migration phenology may mismatch brood parasitic cuckoos and their hosts

Phenological responses to climate change vary among taxa and across trophic levels. This can lead to a mismatch between the life cycles of ecologically interrelated populations (e.g. predators and prey), with negative consequences for population dynamics of some of the interacting species. Here we provide, to our knowledge, the first evidence that climate change might disrupt the association between the life cycles of the common cuckoo (Cuculus canorus), a migratory brood parasitic bird, and its hosts. We investigated changes in timing of spring arrival of the cuckoo and its hosts throughout Europe over six decades, and found that short-distance, but not long-distance, migratory hosts have advanced their arrival more than the cuckoo. Hence, cuckoos may keep track of phenological changes of long-distance, but not short-distance migrant hosts, with potential consequences for breeding of both cuckoo and hosts. The mismatch to some of the important hosts may contribute to the decline of cuckoo populations and explain some of the observed local changes in parasitism rates of migratory hosts.     

Perspectives on Palaearctic and Nearctic bird migration; comparisons and overview of life-history and ecology of migrant passerines

In this paper we compare ecological attributes of tropical migrant passerines from the Nearctic and western Palaearctic, focusing particularly on habitat association patterns during both breeding and wintering seasons. Three regions were compared: Europe, western and eastern North America. Breeding bird census data from 32 studies (each including at least four stages of forest succession) were used to assess the association patterns of breeding habitats among tropical migrants. For each species we calculated an index of habitat diversity and habitat preference.
Tropical migrants preferred earlier successional stages than other birds in Europe. The opposite was true in eastern North America. In eastern North America, tropical migrants tended to be associated with a smaller range of serai stages than other passerine species. In their winter quarters, Palaearctic migrants live primarily in open habitats, such as savannas, whereas eastern Nearctic migrants make more frequent use of evergreen forests. Migrants from western North America show the greatest match between breeding and wintering habitats.
We relate the results to the taxonomy and probable history of contemporary avifaunas and vegetation formations of the Old and New World. Taxonomically, tropical migrants from different parts of the Holarctic are less closely related to each other than residents and short-distance migrants. Tropical and temperate avifaunas are more closely related to each other in the New World than in the Old World. Conservation implications of the between-continent differences are briefly discussed.     

Do Birds Select Habitat or Food Resources? Nearctic-Neotropic Migrants in Northeastern Costa Rica

Nearctic-neotropic migrant birds need to replenish energy reserves during stopover periods to successfully complete their semiannual movements. In this study we used linear models to examine the habitat use of 11 migrant species in northeastern Costa Rica to better understand the influence of food and structural resources on the presence of birds during stopover periods. Our models indicated that frugivorous migrants primarily used food abundance, while insectivorous migrants chiefly used vegetation structure as cues for habitat use during stopover. In addition to habitat use models, we documented fruiting plant phenology and found a general relationship between migrant arrival and the timing of ripe fruit availability. Our results suggest that insectivorous migrants probably rely on structural features when using habitat because it may be inherently difficult to assess cryptic-arthropod availability during a short period of time in a novel habitat, such as stopover periods.     

Climatic effects on timing of spring migration and breeding in a long-distance migrant, the pied flycatcher Ficedula hypoleuca

Climate change has advanced the breeding dates of many bird species, but for few species we know whether this advancement is sufficient to track the advancement of the underlying levels of the food chain. For the long-distance migratory pied flycatcher Ficedula hypoleuca the advancement in breeding time has been insufficient to maintain the synchrony with their main food sources. The timing of arrival in the breeding areas from their African wintering grounds is likely to constrain the advancement of breeding date. We hypothesise that this is because in Africa they cannot predict the advancement of spring in their breeding habitat. However, long-distance migrants may advance their arrival time by migrating faster when circumstances en route are favourable. In this study we show that both arrival and breeding date depend on temperatures at their main North African staging grounds, as well as on temperature at the breeding grounds. Male arrival and average laying date were not correlated, but the positive effect of temperature in North Africa on breeding dates suggests that breeding date is indeed constrained by arrival of females. Long-distance migrants thus are able to adjust arrival and hence breeding by faster spring migration, but the degree of adjustment is probably limited as timing schedules in spring are tight. Furthermore, as climate change is affecting temperatures differently along the migratory flyway and the breeding areas, it is unlikely that arrival dates are advanced at the same rate as the timing of breeding should advance, given the advancement of the underlying levels of the food chain.     

High gene flow in epiphytic ferns despite habitat loss and fragmentation

Tropical montane forests suffer from increasing fragmentation and replacement by other types of land-use such as coffee plantations. These processes are known to affect gene flow and genetic structure of plant populations. Epiphytes are particularly vulnerable because they depend on their supporting trees for their entire life-cycle. We compared population genetic structure and genetic diversity derived from AFLP markers of two epiphytic fern species differing in their ability to colonize secondary habitats. One species, Pleopeltis crassinervata, is a successful colonizer of shade trees and isolated trees whereas the other species, Polypodium rhodopleuron, is restricted to forests with anthropogenic separation leading to significant isolation between populations. By far most genetic variation was distributed within rather than among populations in both species, and a genetic admixture analysis did not reveal any clustering. Gene flow exceeded by far the benchmark of one migrant per generation to prevent genetic divergence between populations in both species. Though populations are threatened by habitat loss, long-distance dispersal is likely to support gene flow even between distant populations, which efficiently delays genetic isolation. Consequently, populations may rather be threatened by ecological consequences of habitat loss and fragmentation.     

Habitat use,abundance, and persistence of Neotropical migrant birds in a habitat matrix in northeast Belize

ABSTRACT To ensure adequate protection of nonbreeding habitats used by Neotropical migratory landbirds, we must first address questions about habitat use and quality. On the Yucatan peninsula, migrants use many habitats, several of which remain unstudied, and methodological differences preclude interhabitat comparisons based on studies to date. We used distance sampling along line transects in six habitats in northeast Belize to examine use of previously unstudied habitats (e.g., salt marsh) by Neotropical migrants and to permit comparison across habitats. We calculated unadjusted and adjusted (for detectability) density estimates for individual migrant species and for all species combined to generate hypotheses about habitat quality based on the assumption that density and quality are positively correlated. Adjusted density estimates for all migrants were highest in black mangrove habitat (1799 ± 110 ind/km²), intermediate in three forest types and milpa (range 598–802 ind/km²), and lowest in salt marsh (207 ± 32.3 ind/km²). By combining density estimates with habitat availability in our study region, we estimated that evergreen forest and black mangrove supported 70% and 9% of the region's migrant population, respectively. At the species level, five of the 10 most common species had habitat preferences (>50% detections in one habitat). Given the diversity of habitat preferences among species and apparent seasonal movements, our results indicate that Neotropical migrants in northeast Belize are dependent on a matrix of interconnected habitats.     

Long-term population declines in Danish trans-Saharan migrant birds

Capsule Long-distance migrant birds show less favourable trends than sedentary/short-distance species.

Aims To use breeding bird surveys to contrast population trends amongst common species according to their migration pattern.

Methods Changes in abundance of 62 Danish breeding sedentary, short-distance (Europe/North Africa) or long-distance (trans-Saharan) migrants were described by fitting log linear regression models to point-count census data gathered during 1976–2005.

Results Trans-Saharan migrants declined by 1.3% per annum during this period, while short-distance migrants and sedentary species increased by 1.4% and 1.0% per annum, respectively. There were no significant decadal declines amongst species using different summer breeding habitats, except for wetlands, and there was no consistent variation in trends associated with wintering regions or habitats or diet.

Conclusions More information is urgently needed on diet, feeding ecology, habitat requirements, winter distribution and intra-African movements of the commoner European summer visitors to identify causes of the declines and highlight when in the annual cycle detrimental effects occur. Studies linking individuals on their breeding, staging and wintering grounds are especially needed. Danish trends resemble those from elsewhere in Europe, confirming that restoration to favourable conservation status requires inter-continental action to meet European and global targets to reduce or halt biodiversity loss.     

Broad-scale patterns in geographical and habitat distribution of migrant and resident passerines in Britain and Ireland

We analysed patterns of breeding habitat use in migrant and resident passerines on a broad geographical scale in Britain. Migrants were less widely distributed than residents and used fewer habitats. Although resident genera that were more widespread used more nesting habitats, this did not hold for migrant genera though it did at the species level. Although migrancy status in Europe has been associated with the use of 'open' structural habitat types (e.g. scrubland), evidence from Britain was contradictory. Within lowland woodlands the ratio of migrants to residents and the density of migrants was highest in habitats with large areas of young growth. The highest migrant ratios of all, however, were recorded in mature upland woods, with little or no undergrowth. These upland woods carried the lowest densities of residents. Regression models accounting for variation in the migrant ratios and densities in woodland habitats varied according to the geographical scale on which they were constructed. The extent of young growth was a strong predictor of migrant ratios and densities in south-east England but not for woods in the north and west. Resident densities in northern and western woods were inversely related to altitude.     

Flower power: tree flowering phenology as a settlement cue for migrating birds

1. Neotropical migrant birds show a clear preference for stopover habitats with ample food supplies; yet, the proximate cues underlying these decisions remain unclear. 2. For insectivorous migrants, cues associated with vegetative phenology (e.g. flowering, leaf flush, and leaf loss) may reliably predict the availability of herbivorous arthropods. Here we examined whether migrants use the phenology of five tree species to choose stopover locations, and whether phenology accurately predicts food availability. 3. Using a combination of experimental and observational evidence, we show migrant populations closely track tree phenology, particularly the flowering phenology of honey mesquite (Prosopis glandulosa), and preferentially forage in trees with more flowers. Furthermore, the flowering phenology of honey mesquite reliably predicts overall arthropod abundance as well as the arthropods preferred by migrants for food. 4. Together, these results suggest that honey mesquite flowering phenology is an important cue used by migrants to assess food availability quickly and reliably, while in transit during spring migration.     

Waterfowl of a subtropical African floodplain

Many water bodies in arid southern Africa are seasonal and waterfowl habitats become restricted during the dry winters. Nevertheless, some tropical species, such as the white-faced duck, are expanding their ranges into more temperate regions where summer habitats are available, thus increasing pressure on overwintering habitats. The subtropical Pongolo River floodplain is an important overwintering area for a diverse water-fowl community that includes 13 of the 16 species that breed in South Africa and one of the few regular migrants. Historical evidence suggests that numbers of white-faced duck, which represent over 70% of the floodplain community, have increased 10-fold over the last two decades. An abundant, nutritious food (Potamogeton crispus turions) attracts waterfowl to the Pongolo floodplain during winter and an increase in bird fat reserves suggests that they leave for summer breeding grounds in good condition. Unseasonal flood releases from an upstream reservoir may cause low food supplies in late spring before the onset of the summer rainy season when breeding habitats become available. This study emphasizes the need for a better understanding of the role of winter habitats in maintaining waterfowl populations.     

Population trends of breeding birds in British woodlands over a 32-year period: relationships with food, habitat use and migratory behaviour

This paper outlines population trends (with confidence intervals) for 49 species in woodland habitats in Britain as monitored by the British Trust for Ornithology's (BTO) Common Bird Census (CBC) between 1967 and 1999. Additionally, the possible causes of these population trends are investigated by relating the ecological characteristics of species to the degree of population change they have undergone over different time periods. Over the whole period, 17 species showed significant decreases in abundance and 12 species showed significant increases. Whilst population trajectories were diverse, long-distance migrants showed more negative trends than other species and the timing of the changes in their populations was related to their wintering latitude, suggesting that these species may be suffering from environmental changes in the non-breeding season. There was also support for habitat specializations being related to population changes, with species classified as scrub and understorey specialists declining on average, but this was only evident across the entire study period. Additionally, species eating seeds in summer declined and those eating vegetation and making use of the agricultural landscape matrix increased. Therefore wide-scale factors such as landscape-scale processes or processes operating outside of Britain appear to be important in addition to local habitat change, especially for long-distance migrants.     

Disturbance regime and limits on benefits of refuge use for fishes in a fluctuating hydroscape

Refuge habitats increase survival rate and recovery time of populations experiencing environmental disturbance, but limits on the ability of refuges to buffer communities are poorly understood. We hypothesized that importance of refuges in preventing population declines and alteration in community structure has a non‐linear relationship with severity of disturbance. In the Florida Everglades, alligator ponds are used as refuge habitat by fishes during seasonal drying of marsh habitats. Using an 11‐year record of hydrological conditions and fish abundance in 10 marshes and 34 alligator ponds from two regions of the Everglades, we sought to characterize patterns of refuge use and temporal dynamics of fish abundance and community structure across changing intensity, duration, and frequency of drought disturbance. Abundance in alligator ponds was positively related to refuge size, distance from alternative refugia (e.g. canals), and abundance in surrounding marsh prior to hydrologic disturbance. Variables negatively related to abundance in alligator ponds included water level in surrounding marsh and abundance of disturbance‐tolerant species. Refuge community structure did not differ between regions because the same subset of species in both regions used alligator ponds during droughts. When time between disturbances was short, fish abundance declined in marshes, and in the region with the most spatially extensive pattern of disturbance, community structure was altered in both marshes and alligator ponds because of an increased proportion of species more resistant to disturbance. These changes in community structure were associated with increases in both duration and frequency of hydrologic disturbance. Use of refuge habitat had a modal relationship with severity of disturbance regime. Spatial patterns of response suggest that decline in refuge use was because of decreased effectiveness of refuge habitat in reducing mortality and providing sufficient time for recovery for fish communities experiencing reduced time between disturbance events.     

The nature of migration in the red admiral butterfly Vanessa atalanta: evidence from the population ecology in its southern range

1. The migrant Vanessa atalanta (L.) occurs throughout Europe and North Africa. In autumn, populations emigrate from northern and central Europe to the Mediterranean region to overwinter. In the spring, the northern range is recolonised by migrants from the south. The dynamics of the species in the winter range is poorly known. 2. From 1994 to 1999, adults and immatures of V. atalanta were monitored all year round in Mediterranean habitats in north‐east Spain. 3. Data showed that the Catalonia lowlands is an area to which V. atalanta migrates to breed during the winter. Migrants arrive in October and early November and initiate a period of intensive breeding. Larval development occurs throughout the winter until a first annual generation of adults appears in early spring. 4. Most of the butterflies emerging in the spring emigrate and leave the area without breeding. The data suggest strongly that recolonisation of the northern range is by these butterflies not by wintering adults. Altitudinal migration also seems to be a common phenomenon, allowing a further summer generation of adults to occur at high elevations within the Mediterranean region. 5. The complex phenology of V. atalanta in its southern range has evolved as a strategy to track larval resources through space and time. Autumn migration coincides with the greatest availability of the main food plant, Urtica dioica L. Late spring migration occurs by the time food quality is decreasing.     

Habitat selection of wintering passerines in salt marshes of the German Wadden Sea

The salt marshes of the Wadden Sea are important wintering areas for some species of granivorous passerines, which have declined considerably since the 1960s. We investigated the habitat choice of all wintering passerines in eight study areas in German salt marshes with special consideration of human impact on these habitats. Granivorous species that almost exclusively winter in salt marshes, Shorelark (Eremophila alpestris), Snow Bunting (Plectrophenax nivalis) and Twite (Carduelis flavirostris) were concentrated in the lower salt marsh vegetation and in the driftlines, while all other species preferred the high upper salt marsh communities, although Rock Pipits (Anthus petrosus littoralis) fed in muddy areas along ditches. Shorelarks switched habitat in conditions where seeds were scarce to feed instead on arthropods in upper salt marshes. Intensively sheep-grazed upper salt marshes resemble lower salt marshes in their vegetation and were therefore mainly visited by Shorelarks, Snow Buntings and Twites. In winter, the driftline is preferred by the two former species, while in autumn and spring more birds foraged in the salt marshes. Twites prefer to feed mainly on seeds of Salicornia. Areas with S. europaea are visited mainly in late autumn and early winter, while areas with S. stricta are used throughout the winter because of a steady supply of seeds. Several years after embankment, polders are hardly used any more by the lower salt marsh species as the habitat changes into freshwater marshes. Large embankment projects since the early 1960s have included salt marshes and intertidal flats, and the resultant loss of habitat is responsible for the decline of lower salt marsh species. For other passerine species the effects of reclamation are unknown. The effects of intensified grazing on the wintering populations of Shorelark, Snow Bunting and Twite are still unresolved. Although grazing supports lower salt marsh vegetation, the seed production per plant is much lower there and some important seed producers hardly occur. Since grazing was reduced and embankment projects have been stopped, the salt marsh areas (especially lower salt marshes) have increased and so have the wintering populations of Shorelark, Snow Bunting and Twite. For the other species, the consequences of habitat changes are unknown, although it is suggested that reduced grazing will support them. Reducing the human impact on salt marshes will, in the long run, probably lead to a natural salt marsh with much variety in elevation and in its corresponding vegetation and bird communities. Meanwhile, management by grazing might be required in parts of the salt marshes.