18 Breeding Birds
18 Breeding Birds
18 Breeding Birds
Breeding Birds
Kees Koffijberg
Lieuwe Dijksen
Bernd Hlterlein
Karsten Laursen
Petra Potel
Stefan Schrader
2009
Common Wadden Sea Secretariat
Trilateral Monitoring and Assessment Group
Colophon
Publisher
Common Wadden Sea Secretariat (CWSS), Wilhelmshaven, Germany;
Trilateral Monitoring and Assessment Group (TMAG).
Editors
Harald Marencic, Common Wadden Sea Secretariat (CWSS)
Virchowstr. 1, D - 26382 Wilhelmshaven, Germany
Language support
Seabury Salmon
Graphic support
Gerold Leren
Published
2009
ISSN 0946896X
1. Introduction
Little Tern
(Photo: Jan van de Kam)
In addition to 10-12 million migratory birds, complete counts across the entire Wadden Sea.
the Wadden Sea also supports large numbers In the past, a total count of all species, including
of breeding birds. For several breeding bird spe- common species, has been carried out every five
cies like Eurasian spoonbill Platalea leucorodia, years (1991, 1996, 2001, 2006). However, the
oystercatcher Haematopus ostralegus, avocet interval has now been changed to six years, so
Recurvirostra avosetta, Kentish plover Charadrius the next survey is due in 2012). The monitoring
alexandrinus, common redshank Tringa totanus, scheme aims to assess and detect population
lesser black-backed gull Larus fuscus, gull-billed size, distribution and population trends in Wad-
tern Gelochelidon nilotica and sandwich tern den Sea breeding birds. Fieldwork is standardized
Sterna sandvicensis, the salt marshes, dunes and and carried out according to trilaterally harmo-
outer sands of the Wadden Sea are among the nized methods (Hlterlein et al., 1995) by nearly
most important breeding sites in Northwest-Eu- 500 ornithologists, mainly consisting of staff of
rope. Several species are included in Annex I of the NGOs, governmental bodies, site managers and
EU-Bird Directive or listed as Species of European volunteers. A so-called Quality Assurance Meet-
Concern (SPEC). At national level, many Wadden ing (QAM) is organized regularly to provide a
Sea breeding birds represent an important share of platform for exchange of field experience among
national breeding bird populations and are listed participants and for discussion of specific count-
as Red List species. ing pitfalls (e.g. Blew, 2003).
Monitoring of breeding birds in the Wadden Sea This chapter presents an update of the QSR
has been carried out by the Joint Monitoring Group 2004 (Koffijberg et al., 2005). It mainly focuses on
for Breeding Birds (JMBB) in the framework of the trends in numbers of breeding birds 1991-2006
Trilateral Monitoring and Assessment Program and highlights some recent developments and
(TMAP) since 1991 (Fleet et al., 1994; Melter et related management issues. Finally an evaluation
al., 1997; Rasmussen et al., 2000; Koffijberg et al., of the targets of the Wadden Sea Plan (1997) is
2006). The monitoring scheme currently focuses made and recommendations are given regarding
on 35 species that are considered characteristic monitoring, research and assessment of manage-
of the Wadden Sea ecosystem. Common breeding ment issues.
birds (eight species) are counted annually in 103
representative census areas evenly distributed
over all regions and habitats. Colonial and rare
breeding birds (27 species) are difficult to survey
with census areas, so they are counted by annual
The results presented here were derived from the carried out in several parts of the Wadden Sea,
data that currently are being processed for the but failed to halt further declines. Kentish plover
next trilateral breeding bird report on the total is still thriving only in Denmark, and hopefully
count of 2006 (Koffijberg et al., 2009). All trend further investigations will provide some clues
analyses have been carried out with the commonly as to what measures could improve the situa-
used package TRIM from Statistics Netherlands tion for both plovers. Furthermore, management
(Pannekoek and van Strien 1999). Significance was measures in the Het Tij Geleerd program in The
tested for p <0.05 by use of a Wald test. Netherlands might be able to restore some of
the coastal dynamics and provide new breeding
2.1 Overall trends opportunities.
Reliable trend estimates are available for 29 spe- Among the other wader species, black-tailed
cies for a period of 16 years (other species were godwit Limosa limosa and Northern lapwing
too rare to allow proper trend estimates). Between Vanellus vanellus have declined as well, demon-
1991 and 2006 nearly half of the monitored spe- strating that the Wadden Sea does not always
cies (13) have been subject to declines (Table 1). function as a last refuge as has been suggested
Furthermore, 8 species have increased whilst 7 earlier (Rasmussen et al., 2000). Even if breeding
species have remained stable. Gull-billed tern, performance in the Wadden Sea is enhanced by
for which the Wadden Sea is the only known less intensive farming (as shown e.g. on the island
breeding site in Northwest-Europe, is the sole of Mand in Denmark, which supports the best
species for which no significant trend could be breeding site in Denmark), population trends in
detected. Today, this species is only breeding in both species are highly dependent on the situation
the Schleswig-Holstein part of the Elbe Estuary, in agriculturally-managed marshland areas behind
where recorded numbers fluctuate, although not the seawall (Thorup and Laursen, 2008). Here,
significantly. habitat changes due to intensification of farming
The rate of population change differs between and increased predation rates (see below) have had
species (Figure 1). With 11 out of 13 declining a serious impact on breeding birds. In Denmark,
species, the strongest declines have been observed where the coastal marshes are part of the trilateral
in waders. Dunlin Calidris alpina schinzii, ruff cooperation area, recent analyses show a decline
Philomachus pugnax and common snipe Gallinago of 60% in all breeding bird species, including six
gallinago still teeter on the verge of extinction Annex 1 species that have disappeared recently
and might well disappear before the next total (Laursen and Thorup, in press.).
count in 2012. All three species have also suf- Largest increases have been observed in
fered major losses in other parts of their breeding a number of colonial breeding birds, notably
ranges (Zckler, 2002; Thorup, 2006), and their Mediterranean gull Larus melanocephalus, great
Wadden Sea breeding sites (mainly in Denmark) cormorant Phalacrocorax carbo sinensis, Eurasian
are only maintained by taking specific conserva- spoonbill, lesser black-backed gull, great black-
tion measures (Thorup, 2003). Declines in dunlin backed gull Larus marinus and common gull
are especially worrying as the small population in Larus canus. In particular, Mediterranean gull and
the Wadden Sea is part of the vulnerable Baltic Eurasian spoonbill have expanded their breeding
population of the subspecies schinzii. range from the southwest to the northeast in
Of the regularly breeding wader species, Kent- the past decade and have colonized most parts
ish plover and great ringed plover Charadrius of the Wadden Sea. In coming years, these spe-
hiaticula show the highest rate of decline and cies will probably extend their breeding range
have abandoned many breeding sites in the past further north. Great cormorant has shown signs
decade. Both species are known to suffer from of stabilization in most inland colonies in Europe
disturbance by beach recreation (Schulz, 1998; (Bregnballe et al., 2003) but it still explores new
Tulp, 1998). In addition many temporarily used breeding and feeding sites in the Wadden Sea.
breeding sites (embankments, industrial areas) Only settlement in Denmark has been halted so
have been deserted as vegetation succession made far, as nests and eggs are controlled annually to
habitat unsuitable to breed (Hlterlein, 1998). prevent successful breeding and expansion of
Besides, natural coastal dynamics a prerequisite the population (Bregnballe and Eskildsen, 2009).
for creating suitable habitat for these species Elsewhere, a lack of safe breeding sites (mainly
is lacking at most sites due to coastal defense islands) might become a limiting factor, resulting
policies. Conservation measures (i.e. prevention of in a saturation of the current population level.
disturbance at potential breeding sites) have been
Table 1:
Summary of trends of breeding birds in the Wadden Sea in 1991-2006. Trend classification of significant trends is ex-
pressed as moderate increase (+), strong increase (++), stable (=), moderate decline (-) or strong decline (--). Non-signifi-
cant trends are indicated by F (fluctuating). For 2001-2006 trends have been classified as increasing (INC) or declining
(DEC= when difference in indices between 2001 and 2006 were >10%; changes of <10% have been classified as stable
(STA). This classification merely gives an indication on recent developments. Not included are little egret, barnacle goose,
Eurasian wigeon and pintail, i.e. species that have been added to the list of counted species recently.
Mediterranean gull
Figure 1:
Trends in breeding birds Great cormorant
1991-2006, expressed as Red-breasted merganser
the rate of annual popula-
tion change (in %). Non- Eurasian spoonbill
significant changes are Lesser black-backed gull
marked light-blue. Popula- Great black-backed gull
tion changes in common
snipe, ruff and dunlin are Common gull
estimated from the data Shelduck
of the total counts 1991,
Sandw ich tern
1996, 2001 and 2006.
Little tern
Hen harrier
Short-eared owl
Arctic tern
Black-headed gull
Oystercatcher
Common eider
Avocet
Common redshank
Common tern
Eurasian curlew
Gull-billed tern
Herring gull
Northern lapw ing
Black-tailed godw it
Great ringed plover
Kentish plover
Common snipe
Ruff
Dunlin
2.2 Recent developments are rather small and it is not clear if the observed
For many species, the trends in 1991-2006 are trends are part of natural fluctuations or indicate
consistent throughout the whole 16-year period. a structural decline. On the other hand, rates of
However, changes in indices between 2001 and decline in oystercatcher and avocet have been
2006 suggest that some formerly thriving spe- lower from 2001 onwards, indicating that the
cies or species with a long-term stable trend are long-term negative trend in these species seems
tending to decline recently. This especially applies to have leveled off. Increases in common gull and
to common eider, arctic tern Sterna paradisaea lesser black-backed gull have leveled off as well,
and little tern Sterna albifrons. All three species suggesting that existing colonies are at saturation
have shown nearly annual declines since 2001, level and that perhaps density-dependent factors
and their populations have suffered losses of are starting to operate.
20-40% in the last five years. In common eider, 2.3 Regional differences in
downward trends have been reported already for
a longer term in the important breeding areas in
trends
The Netherlands, due to limited mussel stocks Given the large geographical range and differences
that were depleted by fisheries (see below). The in management, it is not surprising that many
small populations of red-breasted merganser, species do not show comparable trends within the
gull-billed tern and great black-backed gull have four sections of the Wadden Sea (Table 1, Figure
tended to go down as well recently, but numbers 2). Regional differences in trends are important
in several aspects. They might provide a first clue
Figure 2:
Niedersachsen Summary of regional
Denmark trends 1991-2006. Given
10
10 is the number of species
in each trend category for
8
8 the four respective Wadden
number of species
Sea countries. Fluctuating
number of species
6 6 represents non-significant
trends. Only species that
4 4 occur in relevant numbers
in all four countries have
2 2 been included (N = 17).
0 0
increase decrease stable fluctuating increase decrease stable fluctuating
Schleswig-Holstein
The Netherlands
10
10
8
8
number of species
number of species
6 6
4 4
2 2
0 0
increase decrease stable fluctuating increase decrease stable fluctuating
about the mechanisms that have caused bird or stable numbers elsewhere), great ringed plover
populations to decline, especially when also com- (stable in The Netherlands, declining elsewhere)
paring breeding success in the next years (after and Kentish plover (increase in Denmark, declining
implementation of this parameter in TMAP has elsewhere). All three species are declining on the
been completed). When analyzing the 17 species scale of the Wadden Sea. Furthermore, a south-
that breed all over the Wadden Sea, it is obvious north gradient appears in population trends of
that most populations breeding in the Danish shelduck (increasing to stable towards the north),
Wadden Sea are generally doing well. The large avocet (declining to increasing, apart from small
number of fluctuating trends in this section of the fluctuating population in Denmark) and herring
Wadden Sea is probably a result of the small size gull (declining to increasing). Further analysis
of the Danish part of the Wadden Sea (and thus is necessary to unravel the mechanisms causing
smaller breeding populations in many species). In these patterns.
the other three sections of the Wadden Sea dif-
ferent developments are more or less balanced, 2.4 Comparison with the QSR
although in Niedersachsen and The Netherlands 2004
declining species are clearly dominating. The last QSR 2004 (Essink et al., 2005) included a
In most species no specific pattern in regional trend assessment of Wadden Sea breeding birds
trends could be detected. However, remarkable for 1991-2001. In several species, recent develop-
differences in regional trends were found in oys- ments have caused trends to change when looking
tercatcher (increase in Niedersachsen, declining at the whole 1991-2006 period and decreases
have become more pronounced (Figure 3). This tion). Thus, several typical Wadden Sea breeding
is mainly due to negative trends in populations birds, including Annex 1 species (avocet, arctic
of oystercatcher and avocet. Even if the rate of tern, little tern) and/or species that breed in the
decline has tended to level off recently, popula- Wadden Sea in internationally important numbers
tions in both species have declined by 20-30% (oystercatcher, common redshank) are currently
since 1996. Common redshank has changed its showing negative trends. Of the 14 Annex 1 spe-
previous status as stable into a declining trend, cies breeding in the Wadden Sea, five species
mainly as a result of declines of up to 40-50% in show a long term decline, and an additional four
The Netherlands and Niedersachsen in some recent species show a recent tendency to decline. Hence,
years. Besides, former increases in arctic tern and the conservation status of many breeding birds
little tern have now turned into stable trends, with has become worse compared to QSR 2004.
a recent tendency to decline (see previous sec-
Figure 3: 15
Summary of trends in
the Wadden Sea based
QSR 2004
on assessment in 1991-
2001 (QSR 2004) and
QSR 2009
1991-2006 (QSR 2009). 12
Compared to the previous
number of species
0
increase decrease stable fluctuating
3. Target evaluation
The current Wadden Sea Plan (1997) puts forward of available food stocks in other species groups
two targets that are considered relevant for breed- is largely unknown and has not been investigated
ing birds: (1) favorable food availability and (2) in detail so far. In shellfish-eating birds, the long-
natural breeding success. Two other bird targets, term decline and low current population level
natural flight distances and sufficiently large demonstrate that their conservation status is not
and undisturbed moulting sites, are aspects that satisfactory at the moment.
are mainly dealt with in migratory and wintering Breeding success is another important target
birds (see Laursen et al., 2009). Being designed as in assessing the status of breeding birds. An im-
a monitoring scheme, the data from the breeding portant issue that has been pointed at in several
bird surveys provide excellent information on the recent assessments is the lack of breeding success
state of the Wadden Sea breeding birds, but they due to presumed high predation rates. Declines in
do not always provide information on the mecha- avocet at least for some sites in Niedersachsen
nisms that have caused bird numbers to change. and the eastern Wadden Sea in The Netherlands,
Hence, it is mainly specific research carried out in seem to be associated with low reproduction
the past decade, triggered by reports on declines, rates due to high predation risk, mainly of red fox
that has pointed at some processes going on in Vulpes vulpes (Willems et al., 2005, de Boer et al.,
the Wadden Sea. 2007; Melter and Vaas, 2008). Predation is also
Regarding the target favorable food avail- know to play a role in common redshank (Thyen
ability, especially shellfish-eating birds received et al., 2005), black-headed gull (Oltmanns, 2003;
much attention, particularly because of declines Koffijberg et al., 2006) and common tern Sterna
in the Dutch Wadden Sea in the 1990s. For hirundo (Dijksen and Koks 2003), and also has con-
oystercatcher and herring gull Larus argentatus, tributed to the declines in many farmland-breed-
downward trends in breeding birds also coincide ing waders (Teunissen et al., 2005; Langgemach
with negative trends observed in migratory birds and Bellebaum, 2005, Thorup and Laursen, 2008).
(that are part of the same population dealt with But, its impact at Wadden Sea scale is not clear
in the breeding birds scheme). Migratory and win- as breeding success has not been monitored yet
tering common eider Somateria mollissima have and specific studies to quantify predation rates
been subject to declines as well, although only in the Wadden Sea are scant. Birds breeding on
breeding bird numbers in the Netherlands (i.e. the the mainland coast are especially susceptible
core breeding area within the Wadden Sea) have to predation, as usually important mammalian
gone down significantly so far. All three species predators like red fox or mustelids are absent on
have suffered from the limited stocks of food. This most islands. This probably also explains that spe-
especially applies to blue mussels Mytilus edulis in cies like avocet and black-headed gull are doing
the Dutch Wadden Sea, that were depleted mainly better on island-breeding sites (trend 1991-2006:
by shellfish fisheries at the beginning of the 1990s stable) than along the mainland coast (trend:
(Desholm et al., 2002; Rappoldt et al., 2003; Ens 1991-2006 decline). However, in common red-
et al., 2004; Leopold et al. 2004; Verhulst et al., shank only, trends in Niedersachsen differ from the
2004; Kats 2007). In the eastern part of the Dutch expected pattern (stable on islands and declining
Wadden Sea some recovery of mussel beds has mainland), but at the scale of the entire Wadden
been observed recently, but overall stocks remain Sea, populations both on islands and along the
much lower than before 1990 and a population mainland coast are going down. Trends in com-
recovery in shellfish eating (breeding) birds has mon tern are opposite the picture that would be
not been observed so far. For oystercatcher, Van de expected from predation risk: populations on the
Pol et al. (2007) have also shown that the chance mainland coast are performing better than those
of flooding (due to more stormy weather) in the on islands. In this species, food availability might
breeding season has increased and has had a nega- be a better explanation for the observed trends
tive impact on breeding performance. This clearly (Brenninkmeijer et al. 1997; Stienen et al., 2009;
demonstrates the possible impact of changing cf. recent declines in arctic tern and little tern),
weather patterns as part of global climate change but again, this aspect has not been investigated
(also affecting other breeding birds breeding on in detail. Breeding on the fringe of land and water,
the fringe of water and land). In herring gull, the terns are also susceptible to flooding during the
general abandonment of open rubbish dumps will breeding season, as has been demonstrated fre-
have contributed to the negative trend as well quently for breeding sites like the island of Griend
(Koffijberg et al., 2006). Apart from the rather in the Dutch Wadden Sea (Stienen et al., 2009; cf.
well-investigated shellfish-eating birds, the role oystercatcher mentioned previously).
5. References
trend information on the TMAP website. Jong, F. de, Bakker, J.F., van Berkel, C. J. M., Dankers, N. M. J.
A, Dahl, K., Gtje, C., Marencic, H. and Potel, P., 1999. Wad-
Becker, P. and Muoz-Cifuentes, A., 2004. Contaminants in den Sea Quality Status Report. Wadden Sea Ecosystem No. 9,
bird eggs: recent spatial and temporal trends 1991-2000. Common Wadden Sea Secretariat, Trilateral Monitoring and
Wadden Sea Ecosystem No. 18: 5-25. Common Wadden Sea Assessment Group, Quality Status Report Group. Wilhelm-
Secretariat, Trilateral Monitoring and Assessment Group, shaven, Germany.
Wilhelmshaven.
Kats, R., 2007. Common Eiders Somateria mollissima in the
Blew, J. 2003. New ways to improve and test methods Quality Netherlands: The rise and fall of breeding and wintering
Assurance activities in breeding bird monitoring. Wadden Sea populations in relation to the stock of shellfish. Dissertation,
Newsletter 2003-2, 18-23. Rijksuniversiteit Groningen, Groningen.
De Boer P., Oosterbeek, K.H., Koffijberg, K., Ens, B.J., Smit, C.J. Koffijberg, K., Dijksen, L., Hlterlein, B., Laursen, K., Potel, P. and
and De Jong M.L., 2007. Broedsucces van kustbroedvogels in Sdbeck, P., 2005. Breeding birds. In: Essink, K., Dettmann, C.,
de Waddenzee in 2006. Wageningen, Alterra, Alterra-rapport Farke, H., Laursen, K., Leren, G., Marencic, H. and Wiersinga,
1745. W. (eds.). Wadden Sea Quality Status Report 2004. Wadden
Bregnballe, T. and Eskildsen, J., 2009. Forvaltende indgreb i Sea Ecosystem No. 19. Common Wadden Sea Secretariat,
danske skarvkolonier i Danmark 1994-2008. Omfang og ef- Trilateral Monitoring and Assessment Group, Wilhelmshaven,
fekter af oliering af g, bortskrmning og beskydning. Dan- Germany.
marks Miljundersgelser, Aarhus Universitet. Arbejdsrapport
fra DMU nr. 249. Koffijberg, K., Dijksen, L., Hlterlein, B., Laursen, K., Potel, P. and
Sdbeck, P., 2006. Breeding birds in the Wadden Sea in 2001.
Bregnballe, T., Engstrm, H., Knief, W., van Eerden, M.R., van Results from the total survey in 2001 and trends in numbers
Rijn, S., Kiekbusch, J.J. and Eskildsen, J., 2003. Development of between 1991-2001. Wadden Sea Ecosystem No 22. Common
the breeding population of Great Cormorants Phalacrocorax Wadden Sea Secretariat, Trilateral Monitoring and Assessment
carbo sinensis in The Netherlands, Germany, Denmark and Group, Joint Monitoring Group of Breeding Birds in the Wadden
Sweden during the 1990s. Die Vogelwelt 124 supplement, Sea, Wilhelmshaven, Germany.
15-26.
Koffijberg K., Dijksen, L., Hlterlein, B., Laursen, K., Potel, P. and
Brenninkmeijer, A., Klaassen, M. and Stienen, E.W.M., 1997. Schrader, S., 2009. Breeding birds in the Wadden Sea in 2006.
Sandwich Terns Sterna sandvicensis feeding on shell fractions. Results of the total survey in 2006 and trends in numbers
Ibis 139, 397-400. between 1991 and 2006. Wadden Sea Ecosystem. Common
Desholm, M., Christensen, T.K., Scheiffarth, G., Hario, M., Wadden Sea Secretariat, Trilateral Monitoring and Assessment
Andersson, ., Ens, B.J., Camphuysen, C.J., Nilsson, L., Waltho, Group, Wilhelmshaven, Germany. In prep.
C. M., Lorentsen S.-H., Kuresoo, A., Kats, R. K. H., Fleet, D.M. Langgemach, T. and Bellebaum, J., 2005. Prdation und Schutz
and Fox, A.D. 2002. Status of the Baltic/Wadden Sea popula- bodenbrtender Vogelarten in Deutschland. Die Vogelwelt
tion of the Common Eider Somateria m. mollissima. Wildfowl 126, 259-298.
53, 167-203.
Laursen, K., Blew, J., Ens, B., Eskildsen, K., Gnther, K., Hlterlein,
Dijksen, L. and Koks, B. 2003. Broedvogelmonitoring in het B., Koffijberg, K., Potel, P. and van Roomen, M., 2009. Migratory
Nederlandse Waddengebied in 2002. Sovon-monitoringrapport birds. QSR 2009 Thematic Report No. 18.
2003/03. SOVON, Beek-Ubbergen.
Laursen, K. and Thorup, O., 2008. Breeding birds in the Danish
Ens, B.J., Smaal, A.C. and de Vlas, J., 2004. The effects of Wadden Sea Region 1983-2006, assessment of SPAs. Dansk
shellfish fishery on the ecosystems of the Dutch Wadden Sea Ornitologisk Forenings Tidskrift, in press.
and Oosterschelde. Final report on the second phase of the
scientific evaluation of the Dutch shellfish fishery policy (EVA- Leopold, M.F., Smit, C.J., Goedhart, P.W., van Roomen, M., van
II). Alterra-rapport 1011, RIVO-rapport C056/04, RIKZ-rapport Winden, E. and van Turnhout, C., 2004. Langjarige trends in
2004/03. Alterra, Wageningen. aantallen wadvogels in relatie tot de kokkelvisserij en het
gevoerde beleid in deze. EVAII deelrapport C2. Alterra, Wage-
Essink, K., Dettmann, C., Farke, H., Laursen, K., Lerssen, G., ningen, Alterra rapport 1-139.
Marencic, H. and Wiersinga, W. (Eds.), 2005. Wadden Sea
Quality Status Report 2004. Wadden Sea Ecosystem No. 19. Melter J. and Vaas, A., 2008. Untersuchungen zum Schlupf- und
Trilateral Monitoring and Assessment Group, Common Wadden Bruterfolg des Sbelschnblers Recurvirostra avosetta in der
Sea Secretariat, Wilhelmshaven, Germany. Leybucht. Vogelkdl. Ber. Niedersachs. 40, 305-319.
Exo, K-M., Becker, P.H., Hlterlein, B., Htker, H., Scheufler, Melter, J., Sdbeck, P., Fleet, D.M., Rasmussen, L-M. and Vogel,
H., Stiefel, A., Stock, M., Sdbeck, P. and Thorup, O., 1996. R.L., 1997. Changes in breeding bird numbers in census areas in
Bruterfolgsmonitoring bei Kstenvgeln. Die Vogelwelt 117, the Wadden Sea 1990 until 1994. Wadden Sea Ecosystem No.
287-293. 4. Common Wadden Sea Secretariat, Wilhelmshaven.
Fleet, D.M., Frikke, J., Sdbeck, P. and Vogel, R.L., 1994. Breeding Oltmanns, B., 2003. Von der Hellerweide zur Salzwiese:
birds in the Wadden Sea 1991. Wadden Sea Ecosystem No. 1. Vernderungen der Brutvogelgemeinschaft in der Leybucht
Common Wadden Sea Secretariat, Wilhelmshaven. durch die Nutzungsaufgabe. Vogelkdl. Ber. Niedersachs. 35,
157-166.
Hlterlein, B., 1998. Brutvogelbestnde im Schleswig-Holstei-
nischen Wattenmeer. UBA-Texte 76-97. Umweltbundesamt, Pannekoek J. and van Strien, A.J., 1999. Missen is gissen. Ont-
Berlin. brekende tellingen in vogelmeetnetten. Limosa 72, 49-54.
Hlterlein, B., Fleet, D.M., Henneberg, H.R., Mennebck, T., Pol, M. van de, Pen, I., Heg, D. and Weissing, F. J., 2007. Variation
Rasmussen, L-M., Sdbeck, P., Thorup, O. and Vogel, R.L., 1995. in habitat choice and delayed reproduction: adaptive queuing
Anleitung zur Brutbestandserfassungen von Kstenvgeln im strategies or individual quality differences? American Natural-
Wattenmeerbereich. Wadden Sea Ecosystem No. 3. Common ist 170, 530-541.
Wadden Sea Secretariat, Wilhelmshaven. Rappoldt, C., Ens, B.J., Dijkman, E. and Bult, T., 2003. Voedsel-
reservering voor Scholeksters in de Nederlandse Waddenzee.