Herring and Sprat Abundance Indices Predict Chick Growth and Reproductive Performance of Common Terns Breeding in the Wadden Sea

Herring (Clupea harengus) and sprat (Sprattus sprattus) are the key prey resources of common terns (Sterna hirundo) breeding in the Wadden Sea. Breeding success of the terns has been below average since 2002, coinciding with exceptionally low herring recruitment and sprat abundance. Time series of herring and sprat abundance in the North Sea and in the Wadden Sea were analyzed to explain long-term breeding success and chick development at two common tern breeding colonies. North Sea herring recruitment and sprat abundance in the Wadden Sea explained the largest part of common tern breeding success, both as single variables and in a multiple regression approach. Breeding success showed stronger correlations with herring recruitment indices derived from the North Sea region compared to the Wadden Sea. Also, herring and sprat abundance data explained more variability in breeding success than of more directly responding measures such as growth rate and maximum weight of chicks. Despite spatial and temporal incoherences between fish surveys and the common tern breeding season, breeding success of common terns reflected the abundance of their key prey fish beyond their foraging range and breeding season. We argue that the ecological connectivity between large- and small-scale herring abundance and the responsiveness of common tern breeding success is strong enough to establish a fish–seabird indicator system to be potentially valuable in monitoring and conservation.     

Luxury in freshwater and stress at sea? The foraging of the Common Tern Sterna hirundo

We compared the foraging strategies of Common Terns Sterna hirundo in freshwater (Lake Jeziorsko, Brzeg, Poland) and marine environments (Minsener Oldeoog, German Wadden Sea). Body mass changes, nest relief and duration and number of feeding trips per day were studied by automatically weighing the adults, using electronic balances under the nests. At the freshwater site, adults were lighter both before and after feeding and gained less mass during a trip. in the Wadden Sea, single feeding trips lasted longer than at the freshwater site and the terns made fewer trips per day. To achieve the same mass gain per day as in birds in freshwater, trips at sea had to be longer and food intake per trip was higher. The daily duration of absence for feeding and the daily mass gain were about the same in both areas. The limnetic feeders finished foraging earlier in the evening than the terns foraging at sea. These differences are consistent with the hypothesis that limnetic prey availability was consistent, whereas the tides limited the availability of marine prey. In consequence, foraging over freshwater presents several advantages, such as higher colony attendance, better mate coordination and better parental care.     

Dark-bellied Brent Geese Branta bernicla bernicla, as recorded by satellite telemetry, do not minimize flight distance during spring migration

Nine Dark-bellied Brent Geese Branta bernicla bernicla were equipped with satellite transmitters during spring staging in the Dutch Wadden Sea in 1998 and 1999. The transmitters (in all cases less than 3% of body mass) were attached to the back by a flexible elastic harness. One juvenile female was tracked to the Yamal peninsula in 1998. Eight adult males were selected from a single catch of 75 to span the range of body mass observed on the date of capture (11 May 1999) and all but the lightest individual completed the first lap of the migratory flight to the White Sea, Russia, according to the time schedule normal for the species. Six birds were successfully tracked to Taymyr for a total distance averaging 5004 km (range 45775164) but judging from later movements none bred (although 1999 was a breeding year). Although the routes chosen during spring migration were closely similar, none of the tagged birds migrated together. On average the geese used 16 flights to reach their summer destinations on Taymyr. The longest uninterrupted flights during the first half of the journey (Wadden Sea to Kanin) covered 1056 km (mean of seven adult males, range 7681331), while the corresponding value for the second half of the migration (KaninTaymyr) was only 555 km (mean of six adult males). Only 7% of total time during spring migration was spent in active flight, as contrasted to c.  80% at long-term stopovers. Overall average travelling speed was 118 km/day (range 97148). Including fattening prior to departure the rate of travel falls to 62 km/day (range 4970), in keeping with theoretical predictions. Routes followed deviated from the great circle route, adding at least 700 km (16%) to the journey from Wadden Sea to Taymyr, and we conclude that the coastal route is chosen to facilitate feeding, drinking and resting en route instead of minimizing total flight distance.     

Foraging activity of Rhinolophus hipposideros on the Island of Herrenchiemsee, Upper Bavaria

We studied the foraging behaviour of Rhinolophus hipposideros on the island “Herrenchiemsee” in Lake Chiemsee (Upper Bavaria) during summer 2001. The island offers extensively managed woodlands, highly structured open landscapes and a broad reed belt around the shore. On average the flight activity of the 6 radio tracked females outside the roost lasted 229 min per night. The home range size varied between 6.8 and 62.7 ha (mean 25.2 ha). The size of the activity centres varied between 2.8 and 8.2 ha (mean 5.3) and all except one were located almost exclusively in woodland. Within woodlands the bats did not select for specific spatial structures (different age classes of the stands or canopy densities). Only two bats regularly foraged in additional habitats outside woodlands. One of these bats used orchards and tree rows; the other foraged over artificial ponds and gardens adjoining to its woodland foraging area. We never found the bats foraging over the lake or the reed belt. Longer linear landscape elements as tree lines were used during commuting flights but there was no indication of a continuous foraging activity along these elements. Two females left the island to forage on the mainland in August after the fledging of juveniles. To reach the mainland shore, the bats had to fly at least 1.2 km across the lake.

Assuming that most foraging flights on the island occur in woodlands, a bat density in this habitat type of 0.7 bats/ha can be calculated.     

Intercolony variation in foraging flight characteristics of black‐headed gulls Chroicocephalus ridibundus during the incubation period

Using GPS loggers, we examined the influence of colony, sex, and bird identity on foraging flight characteristics of black‐headed gulls Chroicocephalus ridibundus during the incubation period. We studied tracks of 36 individuals breeding in one urban and two rural colonies in Poland. Birds from both rural colonies performed the furthest flights (mean max distance 8–12 km, up to 27 km) foraging mainly in agricultural areas. Gulls from the urban colony performed shorter flights (mean 5 km, up to 17 km) visiting mainly urbanized areas and water bodies. We found that females performed longer flights and their flight parameters were less repeatable compared to males. Males from both rural colonies visited water bodies more frequently than females. In all colonies, males (but not females) used habitats unproportionally to their availability in the vicinity. Relatively low interindividual and relatively high intraindividual overlap in home ranges indicated considerable foraging site fidelity. Individuals specialized in the use of a particular type of habitat performed shorter foraging flights compared to individuals using diverse habitats during their foraging flights. Our results indicate diverse foraging strategies of black‐headed gulls, including generalists that explore various habitats and specialists characterized by high foraging site and habitat fidelity.     

Sex-specific foraging behaviour in a seabird with reversed sexual dimorphism: the red-footed booby

Most hypotheses attempting to explain the evolution of reversed sexual dimorphism (RSD) assume that size-related differences in foraging ability are of prime importance, but the studies on sex-specific differences in foraging behaviour remain scarce. We compare the foraging behaviour of males and females in a seabird species with a RSD by using several miniaturised activity and telemetry loggers. In red-footed boobies males are 5% smaller and 15% lighter than females, but have a longer tail than females. Both sexes spend similar time on the nest while incubating or brooding. When foraging at sea, males and females spend similar time foraging in oceanic waters, forage in similar areas, spend similar proportion of their foraging trip in flight, and feed on similar prey—flying fishes and flying squids—of similar size. However, compared to males, females range farther during incubation (85 km vs. 50 km), and furthermore feed mostly at the extremity of their foraging trip, whereas males actively forage throughout the trip. Males are much more active than females, landing and diving more often. During the study period, males lost mass, whereas females showed no significant changes. These results indicate that males and females of the red-footed boobies differ in several aspects in their foraging behaviour. Although some differences found in the study may be the direct result of the larger size of females, that is, the slightly higher speeds and deeper depths attained by females, others indicate clearly different foraging strategies between the sexes. The smaller size and longer tail of males confer them a higher agility, and could allow them to occupy a foraging niche different from that of females. The higher foraging effort of males related to its different foraging strategy is probably at the origin of the rapid mass loss of males during the breeding period. These results suggest that foraging differences are probably the reason for the differential breeding investment observed in boobies, and are likely to be involved in the evolution and maintenance of RSD.     

Long-term changes in intertidal flat macrozoobenthos as an indicator of stress by organic pollution

Organic waste-water from beetsugar factories is being discharged by pipeline into the eastern part of the Dutch Wadden Sea since 1969 mainly in September–December. The organic waste load was high in the first four years and diminished considerably from 1974 onwards.From 1968 macrobenthos has been monitored at five permanent stations. Two of these stations were situated relatively close to the outfall of the pipe-line.Data on numerical density of seven macrobenthic species were analysed for synchrony of yearly fluctuations in order to discriminate between pollution-induced changes in population densities and changes due to variation of natural environmental factors.A great deal of concordance in density fluctuations was found between the five stations. Comparison with data from the western part of the Dutch Wadden Sea (150 km apart) revealed a relative good concordance of the density fluctuations in eastern (only 3 stations) and western Wadden Sea. The two stations close to the outfall showed an aberrant pattern of density variations due to the waste discharge.     

Visual tracking from a rigid‐hulled inflatable boat to determine foraging movements of breeding terns

ABSTRACT Defining the at‐sea foraging movements of seabirds is fundamental to understanding their ecology and can also be important in assessing the potential impact of marine developments such as offshore wind farms (OWFs). Surveys of predefined areas using aerial or boat‐based transect surveys may not allow adequate assessment of the relative importance of different areas to birds. Individual‐based satellite or radio‐telemetry can be effective in identifying foraging ranges and preferred areas, but may not be suitable for some species. We developed a method to determine the foraging movements of breeding terns (Sterna spp.) by visually tracking individuals using a rigid‐hulled inflatable boat (RHIB). Sandwich Terns (S. sandvicensis), Common Terns (S. hirundo), and Arctic Terns (S. paradisaea) were tracked from colonies in Norfolk and Anglesey, United Kingdom, from 2006 to 2008. The proportion of complete (from and to colony) trips varied from 29–60% among species, years, and colonies. Individual Sandwich Terns were tracked for periods up to 126 min over distances up to 72 km and as far as 54 km from the breeding colony, further than Arctic (up to 57 km and 29 km from the colony) and Common (to 29 km and <9 km from the colony) terns. Mean values were much smaller. Multivariate modeling of Sandwich Tern foraging trips indicated that flight speeds >50 km/hr coupled with greater distances from shore (>25 km) significantly reduced the likelihood of tracking a bird for an entire foraging trip. Use of different boats that differ in speed and performance may alleviate such issues. Visual tracking allowed us to collect data on foraging behavior, flight height, and prey capture rates, and also permitted comparisons between species. Our results indicate that visual tracking may be an effective means of determining the foraging movements and at‐sea behavior of a variety of short‐ranging, day‐active seabirds.     

A polar system of intercontinental bird migration

Studies of bird migration in the Beringia region of Alaska and eastern Siberia are of special interest for revealing the importance of bird migration between Eurasia and North America, for evaluating orientation principles used by the birds at polar latitudes and for understanding the evolutionary implications of intercontinental migratory connectivity among birds as well as their parasites. We used tracking radar placed onboard the ice-breaker Oden to register bird migratory flights from 30 July to 19 August 2005 and we encountered extensive bird migration in the whole Beringia range from latitude 64 degrees N in Bering Strait up to latitude 75 degrees N far north of Wrangel Island, with eastward flights making up 79% of all track directions.The results from Beringia were used in combination with radar studies from the Arctic Ocean north of Siberia and in the Beaufort Sea to make a reconstruction of a major Siberian-American bird migration system in a wide Arctic sector between longitudes 110 degrees E and 130 degrees W, spanning one-third of the entire circumpolar circle. This system was estimated to involve more than 2 million birds, mainly shorebirds, terns and skuas, flying across the Arctic Ocean at mean altitudes exceeding 1 km (maximum altitudes 3-5 km). Great circle orientation provided a significantly better fit with observed flight directions at 20 different sites and areas than constant geographical compass orientation. The long flights over the sea spanned 40-80 degrees of longitude, corresponding to distances and durations of 1400-2600 km and 26-48 hours, respectively. The birds continued from this eastward migration system over the Arctic Ocean into several different flyway systems at the American continents and the Pacific Ocean. Minimization of distances between tundra breeding sectors and northerly stopover sites, in combination with the Beringia glacial refugium and colonization history, seemed to be important for the evolution of this major polar bird migration system.     

A RADAR STUDY OF TERN MOVEMENTS ALONG THE COAST OF GHANA

During spring and autumn 1970 and 1971 the movements of terns off the coast of Ghana near Accra were studied by radar. In autumn, dense movements were recorded within 22 km of the coast both day and night from July to early December. Fluctuations in the magnitude of the movements suggest that the terns were on passage through the Gulf of Guinea. Simultaneous to these movements were roosting flights into shore roosts and both movements appeared independent.
In contrast, no spring movements along the coastline were detected either by radar or visually from the land. Although small tern flocks probably moved westwards within 22 km of the coast, but outside their detectable radar range, the results suggest that the majority of terns return through the Gulf of Guinea by a different route from that taken in autumn.
The difference between spring and autumn movements correlates well with the seasonal variation of the terns' food sources. In autumn, upwellings result in an abundant fish supply (sardines) in the surface waters close to land, whereas in spring, fish is abundant only well off shore such as at fronts (convergences) and upwellings (divergences) both of which have been located as far as 600 km from the coast.     

General Patterns in Invasion Ecology Tested in the Dutch Wadden Sea: The Case of a Brackish-Marine Polychaetous Worm

The success of invasive aquatic species is determined by a variety of attributes such as wide environmental tolerance, high genetic variability, short generation time, early sexual maturity, high reproductive capacity, and a broad diet. Usually, introduced species, after some time lag since inoculation, show an exponential population increase and expansion. Maintenance of the immigrant species at a high population level will be dependent on interspecific competition with native species and availability of habitat and food. Eventually, the immigrant population may decline, for instance due to increased predation pressure, parasite infestation or loss of genetic vigour. These characteristic patterns in invasive species are reviewed for the case of the North American spionid polychaete Marenzelleria cf. wireni in the Dutch Wadden Sea. This species was first recorded in estuaries and coastal waters of the European continent in the Ems estuary (eastern Dutch Wadden Sea) in 1983. In the western part of the Dutch Wadden Sea the first specimens were found in 1989. The Ems estuary population showed the typical lag-phase, explosive increase, stabilisation, and eventual decline. In the western part of the Dutch Wadden Sea the latter two phases have not yet developed. The strong development and stabilisation of the population in the Ems estuary may have been caused by the availability of a yet not utilised food source. The species' final decline remains largely unexplained.     

Population trend over 100?years and conservation needs of breeding sandwich terns ( Sterna sandvicensis ) on the German North Sea coast

The breeding population of the sandwich tern (Sterna sandvicensis) on the German North Sea coast has undergone substantial fluctuations throughout the last 100 years. Numbers of breeding birds were fluctuating quite strongly in the first 30 years of the twentieth century. From 1930 to the mid-1950s, a relatively steady decrease occurred. A minimum was reached in 1965 with 2,243 pairs. Around 1970, numbers increased quite markedly up to the mid-1990s and reached a centurial maximum with 10,138 pairs in 1996. Most recently, the numbers have dropped to only 5,681 pairs in 2005, the lowest number over the last 30 years. Some colonies have existed over long periods, others only for short periods, with often substantial and sudden changes. The distribution at sea was studied by transect counts from ships. During the reproductive period, high totals were found between the mainland coast and the islands, up to 30 km from the outer coast/island line. The seaward extent of the sandwich tern distribution coincided quite well with the 20-m depth line. Maximum foraging ranges for single colonies were estimated to be ca. 45 km for Trischen, ca. 35 km for Norderoog and ca. 30 km for both Scharh?rn/Nigeh?rn and Juist. Overall flight ranges for all colonies were estimated at 33.8 km for 95% of the birds. Germany has a high international responsibility for the protection of this species. Only a few colonies exist every year, making this species very vulnerable to anthropogenic disturbance, pollution events and fishing activities.

Spring migration strategies of two populations of bar-tailed godwits,Limosa lapponica,in the Wadden Sea: time minimizers or energy minimizers?

Birds can optimize their migration either by minimizing time of transport, energy expenditure, or predation risk during migration. For each of these optimization criteria different fattening and stopover strategies are predicted. The first two of these optimization criteria are examined here for the bar-tailed godwit ( Limosa lapponica ). In the European Wadden Sea two populations of bar-tailed godwits stop over during spring migration between their wintering and breeding areas. The European population winters mainly in Great Britain and the western part of the Wadden Sea and breeds in Fennoscandia. The Afro-Siberian population winters in West Africa and breeds in Siberia. The European wintering population migrates to the eastern parts of Wadden Sea in March where it stays until early May. During this time birds gain 1.9 g d⁻¹ in body mass for a 1500–2000-km non-stop flight to the breeding areas. Afro-Siberian birds stay only for one month in May in the Wadden Sea where they gain on average 9.4 g d⁻¹ in mass for a 4000-km non-stop flight. Intake rates in April/May did not differ between the two populations (1.5 kJ min⁻¹ and 1.8 kJ min⁻¹ for Siberian and European migrants, respectively) but total energy intake was higher for the Siberian migrants, since they spend 50% of the day foraging vs 30% in the European birds. In contrast to European migrants, Afro-Siberian birds start to moult into breeding plumage already in their winter quarters. During their stopover in the Wadden Sea thermostatic costs are lower than at times when European birds are present. Thus, the higher energy demands of the Afro-Siberian birds seem to be fulfilled by a combination of physiological adaptations and a high working level. European birds seem to adopt an energy-minimized migration strategy whereas Afro-Siberian birds appear to follow a time-minimized migration.     

On a wing and a prayer: the foraging ecology of breeding Cape cormorants

The Cape cormorant Phalacrocorax capensis is unusual among cormorants in using aerial searching to locate patchily distributed pelagic schooling fish. It feeds up to 80 km offshore, often roosts at sea during the day and retains more air in its plumage and is more buoyant than most other cormorants. Despite these adaptations to its pelagic lifestyle, little is known of its foraging ecology. We measured the activity budget and diving ecology of breeding Cape cormorants. All foraging took place during the day, with 3.6 ± 1.3 foraging trips per day, each lasting 85 ± 60 min and comprising 61 ± 53 dives. Dives lasted 21.2 ± 13.9 s (maximum 70 s), attaining an average depth of 10.2 ± 6.7 m (maximum 34 m), but variability in dive depth both within and between foraging trips was considerable. The within-bout variation in dive depth was greater when making shallow dives, suggesting that pelagic prey were targeted mainly when diving to <10 m. Diving ecology and total foraging time were similar to other cormorants, but the time spent flying (122 ± 51 min day⁻¹, 14% of daylight) was greater and more variable than other species. Searching flights lasted up to 1 h, and birds made numerous short flights during foraging bouts, presumably following fast-moving schools of pelagic prey. Compared with the other main seabird predators of pelagic fish in the Benguela region, Cape gannets Morus capensis and African penguins Spheniscus demersus , Cape cormorants made shorter, more frequent foraging trips. Their foraging range while feeding small chicks was 7 ± 6 km (maximum 40 km), similar to penguins (10–20 km), but less than gannets (50–200 km). Successful breeding by large colonies depends on the reliable occurrence of pelagic fish schools within this foraging range.     

Impact of wind turbines on birds in Zeebrugge (Belgium)

We studied the impact of a wind farm (line of 25 small to medium sized turbines) on birds at the eastern port breakwater in Zeebrugge, Belgium, with special attention to the nearby breeding colony of Common Tern Sterna hirundo, Sandwich Tern Sterna sandvicensis and Little Tern Sterna albifrons. With the data of found collision fatalities under the wind turbines, and the correction factors for available search area, search efficiency and scavenging, we calculated that during the breeding seasons in 2004 and 2005, about 168 resp. 161 terns collided with the wind turbines located on the eastern port breakwater close to the breeding colony, mainly Common Terns and Sandwich Terns. The mean number of terns killed in 2004 and 2005 was 6.7 per turbine per year for the whole wind farm, and 11.2 resp. 10.8 per turbine per year for the line of 14 turbines on the sea-directed breakwater close to the breeding colony. The mean number of collision fatalities when including other species (mainly gulls) in 2004 and 2005 was 20.9 resp. 19.1 per turbine per year for the whole wind farm and 34.3 resp. 27.6 per turbine per year for 14 turbines on the sea-directed breakwater. The collision probability for Common Terns crossing the line of wind turbines amounted 0.110–0.118% for flights at rotor height and 0.007–0.030% for all flights. For Sandwich Tern this probability was 0.046–0.088% for flights at rotor height and 0.005–0.006% for all flights. The breeding terns were almost not disturbed by the wind turbines, but the relative large number of tern fatalities was determined as a significant negative impact on the breeding colony at the eastern port breakwater (additional mortality of 3.0–4.4% for Common Tern, 1.8–6.7% for Little Tern and 0.6–0.7% for Sandwich Tern). We recommend that there should be precautionary avoidance of constructing wind turbines close to any important breeding colony of terns or gulls, nor should artificial breeding sites be constructed near wind turbines, especially not within the frequent foraging flight paths.     

Long-term changes in Wadden Sea nutrient cycles: importance of organic matter import from the North Sea

The Wadden Sea is a shallow tidal area along the North Sea coast of The Netherlands, Germany and Denmark. The area is strongly influenced by rivers, the most important of which are the rivers Rhine, Meuse and Elbe. Due to the increased nutrient load into the coastal zone the primary production in the Wadden Sea almost tripled during the past few decades. A conceptual model is presented that links nitrogen input (mainly nitrate) via Rhine and Meuse with the annual nitrogen cycle within the Wadden Sea. Three essential steps in the model are: (1) nitrogen limits the primary production in the coastal zone, (2) a proportional part of the primary produced organic matter is transported into the Wadden Sea and (3) the imported organic matter is remineralized within the Wadden Sea and supports the local productivity by nitrogen turn-over. The conceptual model predicts that during years with a high nutrient load more organic matter is produced in the coastal zone and more organic matter is transported into and remineralized within the Wadden Sea than during years with low nutrient loads. As a proxy for the remineralisation intensity ammonium plus nitrite concentrations in autumn were used. Based on monitoring data from the Dutch Wadden Sea (1977–1997) the above mentioned model was statistically tested. In autumn, however, a significant correlation was found between autumn values of ammonium and nitrite and river input of nitrogen during the previous winter, spring and summer. The analysis supports that in years with a high riverine nitrogen load more organic matter is remineralized within the Wadden Sea than in years with a low nitrogen load. A comparison with older data from 1960 to 1961 suggests that the remineralisation intensity in the Wadden Sea has increased by a factor of two to three. This is not reflected by a two to three-fold increase in riverine nitrogen load from 1960 to present. It is suggested that the increased remineralisation rates in the Dutch Wadden Sea between the 1960s and the 1980s/1990s are largely caused by an increased nitrogen flux through the Channel and the Strait of Dover and by an increased atmospheric nitrogen input.     

The Dutch Wadden Sea: a changed ecosystem

Since 1600 the surface area of the Dutch Wadden Sea decreased considerably by successive reclamations of saltmarshes. In 1932 the Zuiderzee (3200 km²) was closed off from the Wadden Sea causing in the remaining part an increase in tidal range and current velocities. In 1969 the Lauwerszee (91 km²) was closed off and turned into a freshwater lake as well. Man's use of the Wadden Sea changed simultaneously. Dredging in harbours and shipping routes, and extraction of sand and shells became common practice. Extraction of sand increased manifold between 1960 and 1985. These activities did contribute to the turbidity of the Wadden Sea water. Discharge of nitrogen and phosphorus compounds into the western Wadden Sea increased also manifold since 1950, causing an increase in phytoplankton production, duration of phytoplankton blooms, and intertidal macrozoobenthic biomass. Loads of metals and organochlorine contaminants entering the Wadden Sea were hard to estimate. Fisheries changed drastically since the 1930's. Fishing for ‘Zuiderzee’ herring came to an end shortly after closing off the Zuiderzee. The anchovy fishery ceased in the 1960’, that for flounder in 1983. Undersized brown shrimps were fished until 1971. Selective shrimptrawls and sorting devices with flushing seawater were introduced to reduce mortality among young flatfish and shrimps. Oysters became extinct in the 1960's due to over-exploitation of the natural beds. Production of mussels increased more than ten times between 1950 and 1961 due to ‘culturing’, catches of cockles increased slowly between 1955 and 1984. Whelks were fished until 1970. The most important changes in the biotic system of the Wadden Sea, increased production of microalgae and intertidal macrozoobenthos, can be attributed to increased nutrient loads. Eutrophication provided ample food supply for mussels which are harvested mainly by man and eider duck, and may have caused also increased growth in juvenile plaice. Increased turbidity may have impaired life conditions for adult dab, and have presented also recovery of sublittoral eelgrass beds after their disappearance in the 1930's due to the ‘wasting disease’. Increased turbidity in the Wadden Sea is probably caused by closing off the Zuiderzee (1932), a significant increase of dredge spoil disposal near Hoek van Holland between 1970 and 1983, and a more than 10-fold increase of mussel culturing in the Wadden Sea since 1950. Stocks of several bird species breeding in the Wadden Sea area suffered great losses in the early 1960's due to a pesticide accident. Most of the breeding populations have recovered. PCB's caused a reproduction failure among harbour seals in the 1970's. Since 1980 official Dutch policy aims at multiple use of the Wadden Sea, with emphasis on protection and restoration of the natural environment. The 3rd Water Management Plan (1989) aims at a development of the Wadden Sea ecosystem towards the situation of ca. 1930, i.e. without undoing present sea dikes and reclaimed polders. Management of the Dutch Wadden Sea will therefore have to focus mainly on reduction of eutrophication, pollution and turbidity. Some management options are discussed.     

Effect of brood size and hatching sequence on prefledging mortality of Sandwich terns: why lay two eggs?

The mortality of Sandwich tern Sterna sandvicensis chicks held in enclosures was studied in colonies on Griend, in the Dutch Wadden Sea, from 1992 to 1999, and on Hirsholm, in the Danish Kattegat, in 1997. Survival of chicks until fledging was 73% for chicks hatching from first-laid eggs or single-egg clutches and 59–64% for partially hatched two-egg clutches, whereas 6% of second hatchlings survived until fledging. Less than 2% of all two-chick broods actually fledged two chicks. Because 18% of the two-egg clutches only hatched one egg, 7% of fledglings of two-egg clutches originated from a second-laid egg. In nests where both eggs hatched, the number of chicks was usually reduced soon after hatching. Within five days of hatching more than 50% of the second hatchlings died of starvation or were preyed upon. It seems that overproduction commonly occurs in Sandwich terns and that investment in a surplus egg mainly serves as an insurance mechanism. On Griend and Hirsholm, chick productivity of two-egg clutches was somewhat higher than for one-egg clutches. Undernourishment was an important cause of death, either directly by starvation or by selective predation of chicks in poor condition. This, in combination with earlier, studies suggests that Sandwich tern parents on Griend are exposed to severe food stress.     

Optimal trajectories for the short-distance foraging flights of swans

Optimal flight theory relates body measurements (wing span, body cross-section, body mass) and aerodynamic variables (air density, drag, profile and induced power ratios) to the most energy-efficient velocity for long distance migration. For short-range (2-10 km) foraging flights the theory is expanded to include non-negligible costs for take-off and energy savings/losses for climbing to altitude (drag decreases with air density and therefore with altitude). The theory predicts clear differences between Tundra and Trumpeter swans. Generally speaking, for flights between 2 and 10 km Trumpeter swans can be expected to fly approximately 5-10 m lower in altitude and 1-2 ms(-1)more slowly than Tundra swans. Moreover, the total energy required for these foraging flights is approximately 150% larger for a Trumpeter than a Tundra swan (80 vs. 120 kJ of direct mechanical energy for a 5 km flight), suggesting that Trumpeter swans may be less inclined to take-off than Tundra swans. These factors indicate that even Trumpeters native to the area (as opposed to recently translocated) would be more vulnerable to hunting than native Tundra swans. The expanded theory is compared to observations made in Utah's Bear River Migratory Bird Refuge.