HELGOL,~NDER MEERESUNTERSUCHUNGEN Helgol~nder Meeresunters. 50, Suppl., 9-37 (1996) zgl.: SCHRIFTENREIHE FUR LANDSCHAFTSPFLEGE UND NATURSCHUTZ, H.

47 (1996)

II. Red List of Biotopes and Biotope C o m p l e x e s of the Wadden Sea Area*
CONTRIBUTORS: Germany: A. Ssymank N. Dankers

The Netherlands:

INTRODUCTION T h e R e d List of B i o t o p e s of t h e W a d d e n S e a A r e a " is b a s e d o n t h e e x i s t i n g lists for Germany and The Netherlands. T h e d e v e l o p m e n t of a r e d list for T h e N e t h e r l a n d s ( D a n k e r s & Wolff, 1994) w a s c o m m i s s i o n e d b y t h e N a t i o n a l R e f e r e n c e C e n t r e (IKC) of t h e D e p a r t m e n t for N a t u r e M a n a g e m e n t of t h e N e t h e r l a n d s M i n i s t r y of A g r i c u l t u r e , N a t u r e M a n a g e m e n t , a n d F i s h e r i e s . T h e list h a s b e e n p r e p a r e d b y N. D a n k e r s a n d W.J. Wolff of t h e I n s t i t u t e for F o r e s t r y a n d N a t u r e R e s e a r c h (IBN - DLO). T h e h a b i t a t m a p of D i j k e m a (1989) f o r m e d t h e s t a r t i n g p o i n t of t h e d e s c r i p t i o n of b i o t o p e s . L o c a l k n o w l e d g e a n d d i s c u s s i o n s w i t h s p e c i a l i s t s w e r e u s e d to c a t e g o r i z e t h e b i o t o p e s a n d t r a n s i t i o n z o n e s . T h e G e r m a n list of b i o t o p e s for t h e W a d d e n S e a a n d c o a s t a l r e g i o n s c o n s t i t u t e s a p a r t of a r e d list of all G e r m a n b i o t o p e s ( R i e c k e n et al., 1994). It h a s b e e n e l a b o r a t e d b y t h e F e d e r a l A g e n c y for N a t u r e C o n s e r v a t i o n ( B u n d e s a m t ffir N a t u r s c h u t z ) w i t h i n t h e D e p a r t m e n t for B i o t o p e P r o t e c t i o n a n d L a n d s c a p e Ecology. T h e r e s u l t s a r e b a s e d o n o w n e x p e r t i s e a n d a n e x t e n s i v e c o n s u l t a t i o n of s c i e n t i f i c e x p e r t s of t h e B u n d e s l ~ n d e r a n d of universities. B o t h n a t i o n a l lists h a v e b e e n b a s e d o n e x i s t i n g d a t a ; n o n e w s u p p l e m e n t a r y r e s e a r c h h a s b e e n c a r r i e d o u t in t h e field for t h i s p u r p o s e .

General

framework

of a R e d L i s t f o r B i o t o p e s

F o r a d y n a m i c s y s t e m like t h e W a d d e n Sea, it m a y b e v e r y u s e f u l to d e v e l o p a R e d List for b i o t o p e s . A g r e a t v a r i e t y of b i o t o p e t y p e s c a n b e r e c o g n i z e d , a n d l i m i t e d t a x o n o -

* This list forms part of the Report on the RED LISTS OF BIOTOPES, FLORA AND FAUNA OF THE TRILATERAL WADDEN SEA AREA. Although D e n m a r k could not contribute to this list due to lack of data on the situation of biotopes a n d biotope complexes, D e n m a r k at this stage agrees to the overall status of threat. 9 * While in the literature the terms habitat a n d biotope are often used as synonyms, the scientific term ,,habitat" m e a n s the place where a distinct species is living. ,,Biotope" (types) are ,, small" ecosystems with defined biotic and abiotic conditions. The scope of the trilateral W a d d e n Sea Conference was to elaborate a trilateral red list of biotopes. For definitions see chapter 'Spatial units'. 9 Biologische Anstalt Helgoland, H a m b u r g u n d Bundesamt ffir Naturschutz, Bonn

10

Biotopes and Biotope C o m p l e x e s of the Wadden Sea Area

mic k n o w l e d g e is required. Protection of biotopes will in most cases p r o v i d e the right conditions for d e p e n d e n t species to survive. Species protection alone is very difficult in m a n y cases b e c a u s e of high fluctuations in the populations from year to year or b e c a u s e of migrations. If it can be m a d e clear w h i c h biotope types are u n d e r pressure, t h e s e can be placed on a Red List Often it will also be clear w h i ch impacts are r e s p o n s i b l e for the deterioration of a Red List biotope, a n d m a n a g e m e n t can act accordingly. In this way, k n o w n but also u n k n o w n Red-List-species will be simultaneously protected. A Red List of biotopes serves m a n y purposes as for e x a m p l e listed in t h e g e n e r a l introduction, but focussing especially on: help for t h e i m p l e m e n t a t i o n of international conventions and Directives of the Europ e a n Union (for e x a m p l e Ramsar C o n v e n t io n . Habitats Directive 9 2 / 4 3 / E E C of the EUI as well as national laws that in c l u d e aspects of biotopes protection - introducing a more holistic l a n d s c a p e a p p r o a c h in nature conservation, w h i ch goes b e y o n d pure s p e c i e s - o r i e n t e d c o n s e r v a t i o n strategies. D e v e l o p m e n t of a Red List for biotopes confronts us with some p r o b l e m s w h i ch do not occur w h e n setting up a list for species. Without a clear definition for a biotope type it will prove difficult to m a k e a list of t h r e a t e n e d biotopes. W h e n o b s e r v e d in g r eat detail, each biotope is different and t h e r e f o r e u n i q u e. W h e n defined too b r o ad l y very few biotope types will fulfil the r e q u i r e m e n t s for inclusion in a Red List. A n o t h e r p r o b l e m is the fact that biotopes are in a d y n a m i c state. T h e y do not b e c o m e s u d d e n l y extinct like a species, but they usually c h a n g e their species composition and structural a p p e a r a n c e slowly w h e n the s u r r o u n d in g conditions c h a ng e. S o m e t i m e s they show a cyclic d e v e l o p ment. In a ' c o m p l e t e ' e c o s y s t e m all stages of the cycle should be p r e s e n t w h e n v i e w e d over a n u m b e r of years. The l e n g t h of this time span is d e p e n d e n t on the c y c l e - t i m e and the n u m b e r of successional stages. A d e c r e a s e of one particular stage does not h a v e to m e a n that the biotope is t h r e a t e n e d if earlier stages are allowed to d e v e l o p . This m e a n s that too strong 'protection' of one particular biotope type, b ei n g part of a succession series, m ay cause p r o b l e m s in the long run. The properties of threat (see criteria system), I Loss in area and, especially, II Loss in quality can only be quantified if r e f e r e n c e s are described. References can be d e s c r i b e d in three ways: 1. In a g e o g r a p h i c a l l y similar area, a c o m p a r a b l e system can be studied. 2. A historical situation can give an idea h o w a biotope can look like, t a k i n g into account d e v e l o p m e n t a n d natural fluctuations. For different variables, e x a m p l e s from different years m ay h a v e to be taken, or s o m e t i m e s m e a n s over a n u m b e r of y ear s are a better option. 3. On the basis of theoretical considerations, b a s e d on ecological k n o w l e d g e . In practice all th r e e m e t h o d s are u s e d together. From a n a t u r e m a n a g e m e n t point of v i e w the more important biotope t y p es in the W a d d e n Sea area are those w h i c h are f o r m e d an d m a i n t a i n e d by an i n t er p l ay of physical and biological processes. If only physical processes are important, the r e g e n e r a t i o n ability g e n e r a l l y will be good, unless the physical processes are p r e v e n t e d from acting. In the North Sea, the W a d d e n S e a a n d in the estuaries, tides an d w a v e s are responsible for the m i x i n g of the w a t e r column, the transport of s e d i m e n t a n d the transport of nutrients an d organisms. Light a n d t e m p e r a t u r e are responsible for the p r i m a r y production.
-

Biotopes and Biotope C o m p l e x e s of the W a d d e n Sea Area

11

In the es t u a r in e and coastal regions the tidal am p l i t u d e is responsible for the largescale m o r p h o l o g y of the coastal area. S a n d y coasts with a small tidal a m p l i t u d e d e v e l o p into a coastline with lagoons. Coasts with m e d i u m tidal amplitude d ev el o p into a system of barrier islands with intertidal flats b e t w e e n the islands and the mainland. A r e a s with a l arg e tidal a m p l i t u d e form an o p e n coast with intertidal sand banks. Th e tidal currents t o g e t h e r with w i n d and w a v e s are responsible for the m a i n t e n a n c e of gullies a n d tidal flats. Wind, and the availability of sand are the primary factors in d u n e an d island formation. In the tidal areas along the coast several characteristic structures h a v e d e v e l o p e d . A W a d d e n Sea system is characterized by c o m p l e t e gully systems. That means, a tidal inlet, ebb and flood systems and m a i n channels, w h i c h branch into small gullies and c r e e k s in sandy or silty areas or saltmarshes. Within a W a d d e n Sea system t h er e is a diversity of tidal flats with s e d i m e n t of different silt content and different e x p o s u r e times. In an interplay of physical and biological processes, saltmarshes and d u n e s are formed. Erosion a n d s e d i m e n t a t i o n processes cause small islands to m o v e in the direction of sand transport. S o m e typical structures especially in the marine e n v i r o n m e n t h a v e a biog e n i c origin such as oyster and mussel beds, reefs of tube building p o l y c h a e t e s or eelgrass fields. T h e area n e a r the tidal inlets is very dynamic. Sand banks in the tidal inlet m i g r a t e in a clockwise direction and cross over to the next island. The sand m o v es al o n g the island like a wave. T h e tip of the island shows cyclic g r o w t h and erosion, d e p e n d i n g on the availability of sand. On some high sand flats, v e g e t a t e d dunes m ay develop. Occasionally t h es e islands m o v e in the direction of the sand transport, and m ay e v e n t u a l l y disappear. A l t h o u g h the system as a w h o l e will contain the major e l e m e n t s of m a r i n e or estuarine systems, any specific part of the system m a y not always contain all of these elements. It is essential to d e t e r m i n e w h i c h e l e m e n t s of the eco sy st em must be r e g a r d e d as 'critical capital'. Critical capital are e l e m e n t s w h i ch are not r e p l a c e d w h e n destroyed. T h e s e can be processes, g e o m o r p h o l o g i c a l or b i o g e n i c structures, species, or e l e m e n t s like scenic beauty. In the following section a n u m b e r of biotopes will be d escr i b ed briefly. T h e s e biotopes h a v e b e e n s e l e c t e d on the basis of the most i m p o r t an t physical processes w h i c h w e r e responsible for their formation. As i m p o r ta n t Red List-processes w e consider horizontal an d vertical tides, wind, waves, primary production, grazing, p r ed at i o n an d b r e a k d o w n of organic matter. This does not m e a n that all biotopes will occur if all of t h ese p r o cesses occur in a region. If available, s o m e aspects relating to quantity, quality and possibilities for r e g e n e r a tion will be described, as well as the significance for some species w h i ch are c o n s i d e r e d important. Often t h e s e p a r a m e t e r s can only be q u a n t i f i e d properly if r e f e r e n c e s are available. For s ev er a l typical W a d d e n Sea biotopes, r e f e r e n c e areas h a v e b e e n s u g g e s t e d in the IKC-report by v a n L e e r d a m et al. (1993). Further studies are n e e d e d to d e v e l o p methods to describe a n d quantify the intrinsic v a l u e s of biotopes, an d to i m p l e m e n t this m et h o d o l o g y in the s u g g e s t e d biotopes. A g e n e r a l o v e r v i e w of coastal biotopes is g i v e n in the habitat atlas of D i j k e m a (1989). Biotopes are also listed on the s u g g e s t e d G e r m a n Red List (Riecken et al., 1994). It is m a i n l y b a s e d on characteristics of the substrate. T h e a d v a n t a g e is that it is easy to map, but the relation with physical a n d biological p r o cesses is not always clear. It is t h er ef o r e

12

Biotopes a n d Biotope Complexes of the W a d d e n Sea Area

not easy to e v a l u a t e the level of threat which would justify that the biotope type be on a Red List. The G e r m a n list therefore contains clear biotopes, whereas the n a t i o n a l list of The N e t h e r l a n d s often contains biotope-complexes. W h e n attempts are m a d e at mapping, it s e e m s wise to follow the G e r m a n approach. D e l i m i t a t i o n of t h e a r e a The area i n v e s t i g a t e d for the preparation of the Red List of Biotopes has b e e n trilaterally a g r e e d on at the 1st m e e t i n g of the Working Group in N o v e m b e r 1993 ~for details see chapter G e n e r a l Introduction). The 20 m isobath line seaward d e l i m i t a t i o n corresponds m a i n l y to a water depth where the photic conditions are still sufficient for benthic macrophytes u n d e r favorable water conditions without pollutants. At the s a m e time this zone constitutes a part of the coastal system with sand transport by currents a n d waves. Elaborating a trilateral approach The G e r m a n list is based on biotopes as they can be recognized in the field. The biotopes are therefore easy to map, but b e c a u s e of interrelations b e t w e e n biotopes, more complex a n d specific c o m b i n a t i o n s of the biotope types are n e e d e d in some cases as man a g e m e n t units. The G e r m a n Red Data book gives only a very rough a p p r o a c h to these biotope complexes. The Dutch list is b a s e d on natural processes which form and m a i n t a i n recognizable biotope types. These biotope types often form clear m a n a g e m e n t units, b u t according to the G e r m a n system they often should be considered as biotope complexes. Although some of these complexes m a y not contain t h r e a t e n e d biotopes or species, the complex as a whole m a y be very characteristic for the W a d d e n Sea, a n d t h r e a t e n e d as a whole. In part I of the trilateral list, biotope complexes are listed which m i g h t b e considered functional l a n d s c a p e units. Examples are migrating islands, sheltered bays, estuaries, complete gully systems a n d bordering tidal flats, u n p r o t e c t e d tips of islands, saltmarshes, transition zones etc. These l a n d s c a p e units may contain all or a large n u m b e r of the threat e n e d biotopes listed later, a n d should be m a n a g e d as a whole. The b i g g e s t threats to these complexes that have led to a loss of area are coastal protection, u r b a n i z a t i o n , dev e l o p m e n t a n d m a i n t e n a n c e of shipping routes. The quality of these units m a y be affected by d i s t u r b a n c e (tourism, military exercises, m a n a g e m e n t services). In part II of this list, recognizable biotope types have b e e n listed. The structure of the G e r m a n list has b e e n followed, a n d the c o r r e s p o n d i n g biotope types of the Dutch list are indicated. For D e n m a r k , at the m o m e n t , it was not possible to get sufficient data that could be i n c l u d e d in the trilateral approach for the red list of biotopes. Thus the trilateral overall a s s e s s m e n t of threat is p r e l i m i n a r y in some cases, as far as possible c h a n g e s m i g h t occur after i n v e s t i g a t i n g the situation in the W a d d e n Sea area of Denmark. Nevertheless, in some cases, especially for biotope types which are characterized by certain well defined v e g e t a t i o n types, the already existing trilateral red list on p l a n t c o m m u n i t i e s of Westhoff et al. (1993) gives a good indication of the overall status of threat for the whole W a d d e n Sea area.

Biotopes a n d Biotope Complexes of the W a d d e n Sea Area Criteria system

13

The a s s e s s m e n t of threat a n d the e v a l u a t i o n of the regeneration ability follows the G e r m a n proposal (Blab et al., 1995) that has b e e n worked out by the Federal A g e n c y for Nature Conservation (Bundesamt ffir Naturschutz). Detailed definitions of all criteria used can be found in this paper. The system is b a s e d on two properties of threat, supplem e n t e d by a n assessment of r e g e n e r a t i o n ability. Criteria like international i m p o r t a n c e or species d e p e n d e n t on the biotope type have not b e e n used. The properties are: | T h r e a t e n e d by destruction (Loss of area). II T h r e a t e n e d by qualitative c h a n g e s (Loss of quality). III Assessment of r e g e n e r a t i o n ability. For each property an attempt is m a d e to use similar categories for the criteria as in the approaches for the Red List for species but a d a p t e d to the n e e d s of assessment of biotopes. The following categories according to Blab et al. (1995; for full definitions see this paper) are used: I. Loss of area, t h r e a t e n e d by direct d_eestruction (DE): 0 Completely destroyed (extinct). 1 T h r e a t e n e d by complete destruction without proper protection or m a n a g e m e n t (critical). 2 Heavily e n d a n g e r e d , indicated by heavy decline of area in nearly the whole W a d d e n Sea region or extinction in several subregions. 3 E n d a n g e r e d , indicated by a g e n e r a l decline or locally extinct at n u m e r o u s sites. P Potentially e n d a n g e r e d : Typical for a biotope only occurring within a confined area which is not t h r e a t e n e d as long as n o t h i n g goes wrong. * Presumably not e n d a n g e r e d at present. II. Loss of quality, t h r e a t e n e d by ci~_alitative c h a n g e s (QU) This is b a s e d on g e n e r a l degradation, such as decrease of biodiversity, especially the loss of typical species, or the destructions of specific variants. The quality can relate to abiotic conditions, structure, typical set of species, interactions of species etc. 0 Completely destroyed. Typical or n a t u r a l variants have disappeared. 1 T h r e a t e n e d by complete destruction, i n d i c a t e d by the fact that the quality is negatively c h a n g e d in the whole r a n g e of distribution within the W a d d e n Sea so that typical variants are t h r e a t e n e d by complete destruction. 2 Heavily e n d a n g e r e d , indicated by a decline of typical variants, or the extinction of variants in subregions. 3 E n d a n g e r e d , w h e n a decline of typical variants over a broad range can be stated, or typical variants b e c a m e extinct at n u m e r o u s sites. * Not e n d a n g e r e d . III. R e g e n e r a t i o n ability (RE) In a d y n a m i c system like the W a d d e n Sea, it is often possible for a biotope type to regenerate. Therefore, it is important to include this property w h e n assessing w h e t h e r a biotope has to be i n c l u d e d in a Red List. The r e g e n e r a t i o n ability is a characteristic fea-

14

Biotopes a n d Biotope C o m p l e x e s of the Wa dden Sea A r e a

ture of a certain b i o t o p e type, but the r e g e n e r a t i o n process itself d e p e n d s v e r y m u c h on the local a n d r e g i o n a l situation, i n c l u d i n g the distances to cover by r e c o l o n i z i n g species etc. For p r o p e r t y III ( r e g e n e r a t i o n ability, RE) the following criteria h a v e b e e n used: N R e g e n e r a t i o n impossible. K R e g e n e r a t i o n h a r d l y possible (more than 150 years). S R e g e n e r a t i o n difficult (15-150 years). B R e g e n e r a t i o n conditionally possible (less than 15 y e a r s u n d e r f a v o r a b l e conditions; certain typical s p e c i e s might n e v e r t h e l e s s n e e d l o n g e r time). X Classification not m e a n i n g f u l ( s p r e a d i n g i n t r o d u c e d or highly a n t h r o p o g e m c types, succession or d y n a m i c stages of a v e r y short occurrence). T h e criteria I a n d II w e r e m e r g e d to one overall evaluation, which h a s at least the s a m e level of the h i g h e s t r a n k given.

Spatial units used The Red List of Biotopes a n d Biotope C o m p l e x e s of the Trilateral W a d d e n Sea A r e a is b a s e d on the following definitions of spatial units that are classified as: PART I: T h r e a t e n e d biotope c o m p l e x e s a n d c o m p l e x r e g i o n a l l a n d s c a p e units; PART II: T h r e a t e n e d biotope types. Types of biotope complexes: characteristic, constant mosaic or s p a t i a l distribution of certain b i o t o p e types. This includes zonation of b i o t o p e s along an e c o l o g i c a l g r a d i e n t (e.g. g r a d i e n t of humidity), a n d a n t h r o p o g e n o u s distribution types c a u s e d b y the historical a n d cultural d e v e l o p m e n t of a certain l a n d s c a p e . " (Blab et al., 1995). Type of b i o t o p e : i d e a l i z e d type, d e r i v e d from similar biotopes in the field, h a v i n g specific ecological, u n i q u e a n d m o r e o r less constant e n v i r o n m e n t a l conditions for animal a n d p l a n t life .... Biotopes constitute the spatial c o m p o u n d of an e c o s y s t e m . " (Blab et al., 1995). Biotope t y p e s h a v e a certain m i n i m u m size, which e n a b l e s m a p p i n g in the field. F o l l o w i n g a hierarchical concept, also biotope c o m p l e x e s are often g r o u p e d t o g e t h e r in a v e r y specific way. T h e s e l a r g e r r e g i o n a l l a n d s c a p e units (see p r o p o s a l s of the IUCNCESP W o r k i n g G r o u p on Red lists for l a n d s c a p e s ) h a v e b e e n p l a c e d at the b e g i n n i n g of the r e d list of biotope complexes, if t h e y are t h r e a t e n e d a n d if they p l a y a n i m p o r t a n t role u n d e r functional a s p e c t s a n d in the m a n a g e m e n t of w h o l e d y n a m i c l a n d s c a p e systems. Small m i c r o s y s t e m s or biotope e l e m e n t s with m o r e or less specific e n v i r o n m e n t a l conditions like e.g. a stone covered with a l g a e in a s a n d y benthic b i o t o p e of the W a d d e n Sea, or a d e c a y i n g plant, are not i n c l u d e d in the trilateral list. T h e y m a y occur in different b i o t o p e t y p e s a n d in very characteristic combinations, but as t h e y c a n n o t b e t r e a t e d as s e p a r a t e m a n a g e m e n t units t h e y are omitted. I m p o r t a n t characteristic b i o t o p e elem e n t s are m e n t i o n e d in the definitions of the n a t i o n a l contributions (Riecken et al., 1994; D a n k e r s & Wolff, unpubl.).

Biotopes a n d Biotope Complexes of the W a d d e n Sea Area SHORT DESCRIPTIONS

15

PART I: B i o t o p e c o m p l e x e s a n d l a r g e r f u n c t i o n a l l a n d s c a p e u n i t s
Landscape complexes:

Migrating islands: Most W a d d e n Sea islands used to migrate, either in the direction of the sand drift, or i n l a n d with the rising sea level. The i n h a b i t e d islands are n o w kept in place by coastal protection measures. Only the u n i n h a b i t e d tips of these islands still show the characteristics of migrating islands. In the Dutch W a d d e n Sea there are two examples of m i g r a t i n g islands left (Rottumerplaat, Rottumeroog). The quality of these two must be considered low b e c a u s e of conservative coastal a n d d u n e m a n a g e m e n t practices. Restoration is relatively easy, but is partly h a m p e r e d by the occurrence of large quantities of rubble, rock a n d concrete which were used in the past for coastal protection. In the G e r m a n W a d d e n Sea several examples of small islands occur that recently d e v e l o p e d a n d are still without any coastal protection or h u m a n settlement (for example: Trischen). They usually have a typical mosaic of y o u n g d y n a m i c biotope types, p r e s e n t i n g a nowadays very rare complex functional system with very high nature conservation value. Major biotope complexes: A3, C1, C3. Sheltered bays: Sheltered bays with their great variety of biotopes used to be an integral part of the W a d d e n Sea. Since m e d i e v a l times the surface area of the bays has decreased due to e m b a n k m e n t s . A distinction should be m a d e b e t w e e n bays with and without freshwater inflow. The Dollart a n d Lauwerszee are examples of bays with freshwater inflow, the G e r m a n J a d e b u s e n an e x a m p l e of a bay without freshwater inflow. In The N e t h e r l a n d s only the Dollart is left. The quality of the bay is still impaired by pollution a n d large-scale d r e d g i n g activities in the seaward part. Restoration is in principle possible, but will cause loss of invested capital. Three areas with relative shelter which might be called bays are the bights of Terschelling, Vlieland a n d the Balgzand. Major biotope complexes: A2, A3, C1. Estuaries: A n estuary is defined as a semi-enclosed part of coastal waters in which there is a r e c o g n i z a b l e tide with a distinct inflow of freshwater by a river, c a u s i n g a gradient from fresh to saline water. The b o u n d a r y of the estuary is d r a w n as far as the brackish i n f l u e n c e is recognizable, yet tidal freshwater biotopes m a y go m u c h further inland. In the Dutch W a d d e n Sea the Doltard, Lauwerszee a n d Zuiderzee used to be estuaries. At p r e s e n t only the Dollard still has estuarine characteristics. The salinity gradient is caused by the runoff of the G e r m a n river Eros a n d the discharge of the Dutch Westerwoldse Aa. The Ems shows a gradient up to freshwater, b u t the Westerwoldse Aa discharges t h r o u g h a sluice at low tide. It can therefore be c o n c l u d e d that estuaries are extinct in the Dutch W a d d e n Sea. The r e m a i n i n g estuarine characteristics of the Dollard have b e e n of low quality in the past b e c a u s e of severe organic pollution. The situation has improved recently. Restoration of the estuarine situation in the Dutch W a d d e n Sea will only be possible after large-scale e n g i n e e r i n g activities a n d is not c o n s i d e r e d realistic. In some areas it m a y be possible to restore some of the characteristic conditions a n d give room for the species that are d e p e n d e n t on these characteristics. In Germany, the estuaries of the river Elbe a n d Weser are the most important ones, still h a v i n g in some small parts more or less n a t u r a l biotope types. Nevertheless, most of the e m b a n k m e n t s

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Biotopes a n d Biotope C o m p l e x e s of the W a d d e n Sea A r e a

a r e heavily d e g r a d e d or d e s t r o y e d by industry a n d housing estates a n d both w a t e r pollution a n d transport cause serious threats. M a j o r biotope complexes: A2, A3. C1, C3.
Biotope complexes
A2 Natural coastal sea complexes : This biotope complex includes both the benthic and pelagic ecosystems of the shallow water zone of the North Sea. The limit being a water depth where sand can still be transported by waves and currents and where the photic conditions are still sufficient for benthic macrophytes. This is to some extent dependent on water conditions and pollutants and can vary from about 12 m to a maximum of 20 m water depth. ,,Natural" means relatively low disturbance, for example no modification of the substrate by dredging or extraction of material, and relatively low water pollution. Types of biotopes present'. 03.01, 03.02 all subtypes. A3 Natural complexes of tidal flats: The diversity of tidal flats, vegetated or without vegetation, including different subtrate-types, as far as the water dynamics is natural and not reduced or heavily modified by coastal defence or shipping routes. It includes highly dynamic patterns of zonation and complete gully systems, as well as biogenic structures like blue mussel beds or congregations of tube-building polychaetes etc. The quality of the different flats has decreased greatly, mainly because of the impact of mussel and cockle fishery which has an impact on the biological quality, but also on the sediment dynamics and sediment composition. Locally, disturbance may lower the quality for foraging birds or seals. With proper policies the quality of most biota will be restored quickly, with the exemption of sea grass beds and stable mussel beds which may take decades to redevelop. There are three subtypes that deserve special attention, as they are functional subunits to be taken into account for management plans and conservation: A3.01 Tidal divide: Between two tidal inlets in general a clear tidal divide develops. These regions are characterized by tidal flats, small gullies and creeks, low current speeds, small median grain size and extensive mussel beds. The majority of these characters are dependent on physical processes and are therefore not threatened, as long as no dams to islands are constructed. Fisheries, and local disturbance, have decreased the quality of these areas, both biologically as well as from a sediment point of view. Restoration is simple by proper policies. Stable mussel beds and seagrass beds will take a long time to develop. Unstable, temporary mussel beds will probably restore the typical sediment characteristics of these areas.

M a j o r t y p e s of b i o t o p e s present: 05.01.01-.03, 05.01.07, 09.01. A3.02 E b b delta: T h e e b b deltas in the s e a w a r d part of the tidal inlets a r e f o r m e d by p h y s i c a l forces, a n d are therefore not t h r e a t e n e d from a quantitative p o i n t of view. The s a n d b a n k s b e t w e e n the gullies m i g r a t e in a clockwise direction a n d e v e n t u a l l y join the next island. Some s a n d b a n k s m a y b e so h i g h that t h e y are not c o v e r e d at h i g h tide. T h e s e b a n k s serve as roosting p l a c e s for birds a n d seals. Occasionally v e g e t a t i o n or small dunes m a y develop, but only t e m p o r a r i l y as w i n t e r storms will flood t h e s e b a n k s . O n l y dist u r b a n c e has d e c r e a s e d the quality of this biotope. Restoration follows a l m o s t i m m e d i a tely after the i m p l e m e n t a t i o n of regulations. M a j o r t y p e s of b i o t o p e s present: 05.01.01-.03, 05.03.

Biotopes a n d Biotope Complexes of the W a d d e n Sea Area

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A3.03 C o m p l e t e gully systems: Complete gully systems stretching from the e b b - a n d flow gullies in the tidal inlet to secondary, tertiary a n d higher-level creeks in s a n d a n d m u d fiats or saltmarshes, a n d occurring in a natural state. Only locally are they i m p a c t e d by physical processes. The quality of the biota in these systems is i n f l u e n c e d by fisheries in the whole area. The impact is expected to increase with the o n g o i n g increase in the power of the fishing vessels. Because of the dynamic character of the biotope a n d its biota, restoration will be quick. Major types of biotopes present: 03, 05 especially 05.03.01, 05.03.02. C1 Saltmarsh complexes (natural and seminatural): Along coastlines w h e r e sheltered conditions prevail, a typical zonation of saltmarsh complexes develop. In the v e g e t a tion belt, s e d i m e n t a t i o n occurs resulting in a sediment composition with a particle size finer than sand, often i n c l u d i n g relatively high amounts of organic matter a n d distinct layers of shell fragments. From the sea to the land the vegetation belts usually i n c l u d e different pioneer types of saltwort stands {Salicomia europaea agg.), Puccinellia-domin a t e d grassland (Puccinellion), species-rich saltmarshes of Asteretea tripolii, for e x a m p l e J u n c e t u m gerardii, up to more or less brackish marsh vegetation. Many other biotope types like gully systems, brackish reed v e g e t a t i o n etc. are intermingled. Major types of biotopes present: 07, 05, 09, 10.01-10.03. C1.01 Island saltmarsh complex: Island saltmarshes have almost all d e v e l o p e d within sheltered regions; in the N e t h e r l a n d s b e h i n d artificial sand dikes. The s e d i m e n t contains a larger fraction of sand than the m a i n l a n d marshes. Island saltmarsh complexes are often found in close contact with transition zones to d u n e systems or sand fiats covered with Salicornia dobchstachya ssp. (decumbens) with strong airborne sand drift. The protection levels are such that the quality is in most cases g u a r a n t e e d . In some of the island marshes, a too high grazing pressure has lowered the quality. Also the use of four-wheel driven vehicles for m a i n t e n a n c e has locally influenced the quality of the marsh. W h e n island marshes are lost, restoration will often be difficult in a period with sea level rise a n d lack of s u s p e n d e d sediment. Major types of biotopes present: 07.01.01, 07.02.01, 05.01.01, 05.02, 05.03. C1.02 Mainland saltmarsh complex, natural: M a i n l a n d saltmarshes usually contain a larger fraction of silty a n d m u d d y material than the island saltmarsh complex. N a t u r a l zonation complexes from p i o n e e r glasswort vegetation up to the transition zones towards freshwater marshes are very rare nowadays. In the Netherlands, especially those transition zones up to low-lying freshwater marshes have completely disappeared. Here, there may be some possibilities for restoration w h e n n e w saltmarshes are d e v e l o p e d in the s u m m e r polders. Because the l a n d b e h i n d the sea dike lies lower than the saltmarsh, no natural flow of freshwater to the marsh will occur. Water will have to be p u m p e d to the restored area. A completely n a t u r a l d e v e l o p m e n t will therefore not be possible. Only with the help of large-scale projects in areas w h e r e the m a i n l a n d is higher t h a n the high tide level (southeast G r o n i n g e n a n d Drente) are n a t u r a l d e v e l o p m e n t s possible. Major types of biotopes present: 07, 05, 09, 10.01, especially 07.01.01, 07.02.02. C1.03 Sluiter: A slufter is a d u n e valley in more or less o p e n c o n n e c t i o n to the North Sea. They typically develop w h e n the sea breaks t h r o u g h a d u n e ridge, or w h e n a n estuary along a s a n d y coast gradually fills up. It is not certain if slufters are n a t u r a l l y occurring long lasting biotopes. Most of the time, the elevation of the d u n e valley will be such that the tidal volume is not sufficient to keep a tidal inlet in existence. If the d u n e

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Biotopes a n d Biotope Complexes of the W a d d e n Sea Area

ridge closes again, the slufter will evolve into a primary d u n e valley. Two e x a m p l e s of slufters in The Netherlands are the ,,Zwin" on the Belgian/Dutch border a n d the ,0Slufter" on Texel. The Zwin has formed in the previous estuary of Brugge, a n d the Slufter dev e l o p e d after a b r e a k in the s a n d dike b e t w e e n the islands Texel a n d Eyerland. The flooded area consisted of previous tidal flats. Within slufters a great variety of biotopes will occur, r a n g i n g from intertidal fiats a n d saltmarsh vegetation to areas covered with s w a m p - or f e n - v e g e t a t i o n that varies with the salinity a n d the a m o u n t of lime in the soil. As slufters a p p e a r to be a stage in a succession, they are severely t h r e a t e n e d if n e w slufters do not develop. Major types of biotopes present: 10.01-10.03, 07.05. C3 S a n d c o a s t c o m p l e x e s (natural): Along coastlines with s e d i m e n t a t i o n of sands in sufficiently large quantities, a zonation complex with s a n d beaches, drift lines, embryonic d u n e s a n d the whole system of d e v e l o p i n g white, grey a n d brown d u n e s arises. D u n e formation a n d succession series m a i n l y d e p e n d on s e d i m e n t a t i o n rates of sand, average w i n d conditions a n d from the C a - c o n t e n t of the deposited material. Natural u n d i s t u r b e d complexes are rare, as in most cases either coastal defence limits the d y n a m i c s of sand transportation, or part of the biotope types have already b e e n destroyed b y b u i l d i n g activities, tourism etc. Major types of biotopes present: 05, 09, 10, 11.03. Three Subtypes, forming functional units, are of special importance: C3.01 D u n e - t i d a l flats t r a n s i t i o n z o n e : M a n y island transitions b e t w e e n d u n e s a n d tidal flats can be found, but both quantitatively a n d qualitatively they c a n n o t be compared with the n a t u r a l situation. Especially the transition from old d u n e s to tidal flat has become rare. The biggest threat is coastal defence. In i n h a b i t e d areas restoration will be difficult, but with proper m a n a g e m e n t prospects for restoration are promising. Major types of biotopes present: 05, 09, 10. C3.02 P l e i s t o c e n e t i d a l f l a t s / s a l t m a r s h t r a n s i t i o n z o n e : Only in a few places in The N e t h e r l a n d s a n d in G e r m a n y does a n a t u r a l transition b e t w e e n the pleistocene (geest) a n d tidal flats or saltmarshes still exist (NL: Texel, Wieringen, Zuiderzee islands a n d coast, a n d Bergen op Zoom; D: Nordfriesische Inseln, esp. Sylt, Amrum). In NL these places are now separated from sea by polders, stretches of freshwater or dikes. The only place where restoration is feasible is along parts of the n o r t h e r n shore of Wieringen. Major types of biotopes present: 09, 05, 07. C3.03 ,,Washover": Washovers occur w h e n the sea breaks through a single d u n e ridge which separates the North Sea beach from the beach plain along the W a d d e n Sea. Perman e n t gullies are not formed, but eolian erosion will create a diversity in biotopes. Sand will become available for the d e v e l o p m e n t of n e w biotopes. Conservative coastal defence a n d d u n e m a n a g e m e n t are the biggest threat for washovers from a quantitative point of view. Major types of biotopes present: 09, 10.
P A R T II: B i o t o p e t y p e s

The definitions of biotope types are m a i n l y b a s e d on the Red Data book of Biotopes for G e r m a n y (Riecken et al., 1994) a n d have b e e n adapted, i n c l u d i n g the concepts of The Netherlands. The list is a r r a n g e d in groups of biotope types in a hierarchical way.

Biotopes and Biotope Complexes of the Wadden Sea Area

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03. Shallow coastal water zone: The shallow coastal water zone includes the sublittoral, i.e. the sea bottom from the MLT-lineto the deepest distribution of benthic macrophytes (ca 15-20m below MLT, for practical convenience the 20 m depth line can be taken) and the water body on top. This complex is roughly divided into the pelagic zone and the benthic zone. 03.01 Pelagic zone of the shallow coastal waters: Water body of the shallow coastal water zone. The coastal sea is considered to be the region between the islands and ca the 20 m depth line. There are several types of impact. The majority of impacts does not seriously threaten the quality as a whole. Fisheries and pollution have an influence on the area as a whole. Part of the direct fishery impact can be stopped by closing areas. Changes in fish stocks or species composition are to a large extent caused by fishery activities in the rest of the North Sea. Improving the quality in this aspect may be possible by closing large areas of the North Sea for fisheries, includingthe adjoining parts of the nurseries inside the Wadden Sea. Factors of threat: eutrophication and pollution (waste-water inlets, heavy-metal input, oil pollution, dumping) intensive fishery, shipping. 03.02 Benthic zone of the shallow coastal waters of the North Sea: Sublittoral sea bottom of the shallow coastal water zone of the North Sea between MLT and ca 15m depth with benthic plant and animal communities. In this area macrophytes and macroalgae can occur due to favourable or sufficient light conditions; further division according to substrate types and vegetation cover. Vegetation communities: Ruppion maritimae Br.-Bl. 31 em Den Hart. et Seg. 64, Zosteretum marinae Harms. 36. Factors of threat: eutrophication and pollution (waste-water inlets, heavy-metal input, oil pollution, dumping), groundnet fishery, oil exploitation. 03.02.01 Benthic zone of the shallow coastal waters with muddy bottom and few macrophytes: Sea bottom with poor vegetation of the shallow coastal waters of the North Sea with a substrate of clay and mud (prevailing particle size < 0.06mm). Factors of threat: eutrophication and pollution (waste-water inlets, heavy-metal input, oil pollution, dumping), groundnet fishery, dyking, oil exploitation. 03.02.02 Benthic zone of the shallow coastal waters with fine to medium sand bottom, few macrophytes: Sea bottom, poor in vegetation, of the shallow coastal waters of the North Sea with a substrate of fine to medium sand (prevailing particle size between 0.06 and 0.6ram). Factors of threat: eutrophication and pollution (waste-water inlets, heavy-metal input, oil pollution, dumping, mud accumulation), groundnetfishery, dyking, extraction of sand, oil exploitation. 03.02.02.a Sand bank (permanently submerged) 03.02.03 Benthic zone of the shallow coastal waters, coarse sand, gravel or shells bottoms, few macrophytes: Sea bottom of the shallow coastal waters, without vegetation or with few macrophytes only, with a substrate of coarse sand - gravel or mussel shells (prevailing particle size between 0.6 - > 2mm). Factors of threat., eutrophication and pollution (waste-water inlets, heavy-metal input, oil pollution, dumping, mud accumulation), groundnet fishery, dyking, extraction of sand and gravel, oil exploitation.

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Biotopes a n d Biotope Complexes of the W a d d e n Sea Area

03.02.04 Benthic z o n e of the s h a l l o w coastal waters w i t h stony and h a r d bottom, f e w macrophytes: Sea bottom, with hard substrates, of the shallow coastal w a t e r zone without v e g e t a t i o n or with few macrophytes only, mostly with rock, p e b b l e or coarse stone. Hard substrates can be considered rare in the Dutch W a d d e n Sea. N a t u r a l hard substrates are fossil shell banks, clay a n d peat b a n k s or gravel a n d boulders w h i c h come to the surface if gullies cut through old sediments. These biotopes often c o n t a i n o p e n spaces which are u s e d as shelter by a variety of organisms. They also act as a substrate for sed e n t a r y organisms, a n d some fish species require hard substrate for d e p o s i t i n g a n d attaching their eggs. Several organisms which drill holes in peat, clay or w o o d are fully dep e n d e n t on this biotope type. Factors of threat: eutrophication a n d pollution (waste-water inlets, h e a v y - m e t a l input, oil pollution, d u m p i n g , m u d accumulation), g r o u n d n e t fishery, d y k i n g , oil exploitation. 03.02.05 Benthic z o n e of the s h a l l o w coastal waters w i t h m u d d y and s a n d y bottom, rich in macrophytes: Upper parts of the sublittoral of the North Sea with m u d d y or s a n d y bottom, covered with Zostera m a r i n a or Ruppia maritima agg. - vegetation. The m a i n vegetation consists of subtidal seagrass beds (dense stands of Zostera marina). The subtidal seagrass beds in the Dutch W a d d e n Sea area have b e e n extinct since the early 1930 s b e c a u s e of a disease. In the Dutch W a d d e n Sea the area might have b e e n up to 5000 ha, a n d the only other big stand in the W a d d e n Sea occurred in K 6 n i g s h a f e n in Sylt. In G e r m a n y subtidal seagrass beds still exist and have partly recovered from their decline. The seagrass stands form a biotope for a n u m b e r of rare species especially for b r e e d i n g of fishes. Reduced currents, m u d d y sediments, a n d cover favours egg l a y i n g a n d larval development. Vegetation communities: Zosteretum m a r i n a e Harms. 36, Ruppion m a r i t i m a e Br.-BI. 31 em. Den Hart. et Seg. 64. Factors of threat: eutrophication a n d pollution (waste-water inlets, h e a v y - m e t a l input, oil pollution, d u m p i n g , m u d accumulation), g r o u n d n e t fishery, d y k i n g , oil exploitation. 03.02.06 Benthic z o n e of the s h a l l o w coastal waters w i t h hard b o t t o m and rich in macrophytes: Upper parts of the sublittoral of the North Sea with hard bottom, mostly covered with large b r o w n algae (Laminaria saccharina, L. digitata, L. hyperborea, partly Fucus serratus). In d e e p e r areas with i n c r e a s i n g a m o u n t s of small red algae; in G e r m a n y naturally only outside the W a d d e n Sea region on the rock-base of H e l g o l a n d (blue mussel beds covered with algae s. 03.02.07). Hard substrates can be c o n s i d e r e d rare in the Dutch W a d d e n Sea as well. Natural hard substrates are fossil shell banks, clay a n d peat b a n k s or gravel a n d boulders which come to the surface if gullies cut t h r o u g h old sediments. These biotopes often c o n t a i n o p e n spaces which are used as shelter by a variety of organisms. They also act as a substrate for s e d e n t a r y organisms, a n d some fish species require hard substrate for d e p o s i t i n g a n d attaching their eggs. Several o r g a n i s m s which drill holes in peat, clay or wood are fully d e p e n d e n t on this biotope type. Factors of threat: eutrophication a n d pollution (waste-water inlets, h e a v y - m e t a l input, oil pollution, d u m p i n g , m u d accumulation), g r o u n d n e t fishery, extraction of sand, oil exploitation. 03.02,07 B l u e - m u s s e l b e d of the sublittoral: Colony of blue mussels (Mytilus edulis) in the subtidal region. Blue mussel beds also occur in the intertidal region (see 05.01.07).

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Mytilus edulis is a species that doesn't n e e d to live directly on the sea floor b u t builds thick, multi-layered mussel beds with its byssus. The mussel bed creates a hard substrate in the sandy or m u d d y surroundings, partly with a more or less d e n s e cover of b r o w n algae (mostly Fucus vesiculosus a n d F. serratus). Stocks are partly initiated by the mussel culture; t h r e a t e n e d mussel beds are several years old a n d naturally grown. T h e r e are indications that the beds in the subtidal zone are not very stable a n d do not form old structures. Factors of threat: pollution (heavy-metal input, oil pollution, dumping, m u d a c c u m u l a tion), particularly old, n a t u r a l stocks are t h r e a t e n e d by intensive fishery. 03.02.08 Sublittoral oyster reef: Oyster reefs have b e e n extinct since last century. Like mussel beds, oyster reefs are the basis for complex biological communities. A study on an oyster reef in the W a d d e n Sea resulted in the introduction of the term 'biocenosis' in ecology, well over a h u n d r e d years ago. The flat oyster Ostrea edulis has d i s a p p e a r e d from the W a d d e n Sea. Attempts at reintroduction have not b e e n successful. In some places the introduced J a p a n e s e oyster Crasostrea virga is surviving a n d reproducing. Reefbuilding has b e e n observed in the Eastern Scheldt, but not in the W a d d e n Sea. It is not clear whether the p r e s e n t conditions in the W a d d e n Sea are suitable for the d e v e l o p m e n t of reefs of the flat oyster. 03.02.09 Sublittoral Sabellaria reef: The reefbuilding polychaete Sabellaria sp. is still occurring in the W a d d e n Sea. Extensive reefs can be considered extinct for the Dutch W a d d e n Sea, a l t h o u g h it is not clear w h e t h e r they ever occurred in the Dutch part of the W a d d e n Sea. In G e r m a n y the reefs have largely d i s a p p e a r e d b e c a u s e of fishing activities with s e d i m e n t - d i s t u r b i n g trawls a n d dredges. R e g e n e r a t i o n may be possible in areas which are closed for fishery with bottom gear. It is not k n o w n w h e t h e r these reefs will redevelop if fishery is stopped. Factors of threat: pollution (waste-water inlet, h e a v y - m e t a l input, oil pollution, dumping), intensive fishery, g r o u n d n e t fishery. 03.02.10 ,,Seamoss" stand: Colony of diverse species of hydropolyps (i.e. Sertularia spp., Hydralmannia spp.) on hard bottom, also on mussel beds or shell substrates. Factors of threat: pollution (waste-water inlet, h e a v y - m e t a l input, oil pollution, dumping), intensive fishery, g r o u n d n e t fishery. 03.09 Artificial benthic hard substrates: A special type of biotope are artificial rocky shores, (dikes a n d breakwaters) a n d wrecks of w o o d e n or metal vessels often with very rich growth of macrophytes. They m a y be of great value as they provide secondary biotopes substituting rare primary ones a n d they increase the diversity. T h e y m a y have a dense cover of macrophytes. 05 Tidal flats (eulittoral): Flats of the North Sea, periodically falling dry at low tide, also occurring in estuaries, situated b e t w e e n MLT a n d MHT. Typical are great amplitudes of temperature, light a n d salinity, high eutrophication a n d i n p u t of organic s u b s t a n c e s from rivers a n d the open sea. Multiple forms of organisms, from microscopic diatomes over mussels, worms to g r e e n algae a n d seagrass, settle in this area within or on the substrate. They play a n essential role in the trophic p y r a m i d a n d support a high variety of birds and fish. This type is s u b d i v i d e d into substrate types, v e g e t a t i o n cover a n d salinity of the water. 05.01 Tidal flats, free of vegetation: Tidal flats of the North Sea without macrophytes

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Biotopes a n d Biotope C o m p l e x e s of the W a d d e n Sea A r e a

(normally c o v e r e d b y thin layers of blue a l g a e a n d diatoms). This t y p e c a n b e s u b d i v i d e d a c c o r d i n g to s u b s t r a t e types a n d the salinity of the water. Factors of threat: eutrophication a n d pollution ( w a s t e - w a t e r inlets, h e a v y - m e t a l input, oil pollution), g r o u n d n e t fishery, dyking, poldering, oil/gas exploitation, military target areas, shipping, intensive r e c r e a t i o n a l use. 05.01.01 M u d flats, free of vegetation: Mud flats free of vegetation, of h i g h e r plants or m a c r o a l g a e , covers of m i c r o a l g a e are frequent: with s e d i m e n t s of finest material, at least half of the s e d i m e n t consisting of silt and clay (grain size < 0.06mm). Factors of threat: eutrophication a n d pollution ( w a s t e - w a t e r inlets, h e a v y - m e t a l input, oil pollution), g r o u n d n e t fishery, dyking, poldering, oil/gas exploitation, military target areas, shipping, intensive recreational use. 05.01.02 M i x e d flats, free of vegetation: M i x e d m u d a n d s a n d flats with g r a i n fractions of s a n d b e t w e e n 50 - 85% fine sand; mostly free of m a c r o p h y t i c v e g e t a t i o n ; covers of m i c r o a l g a e m a y be present. Factors of threat: e u t r o p h i c a t i o n a n d pollution ( w a s t e - w a t e r inlets, h e a v y - m e t a l input, oil pollution), g r o u n d n e t fishery, dyking, poldering, oil/gas exploitation, military target areas, shipping, intensive recreational use. 05.01.03 S a n d flat, tree of vegetation: Flats with a s e d i m e n t consisting of m o r e than 85% of fine s a n d (0.06 - 0.2mm), mostly free of m a c r o p h y t i c vegetation. E s p e c i a l l y in sand flats, t u b e - b u i l d i n g p o l y c h a e t e s like Lanice conchilega m a y d e v e l o p d e n s e colonies, giving rise to specific structures (biotope-elements) with a quick s e d i m e n t a t i o n in the order of m o r e than 0.5 m p e r year, c o m p a r e d to s u r r o u n d i n g parts of the flat. Factors of threat: e u t r o p h i c a t i o n a n d pollution ( w a s t e - w a t e r inlets, h e a v y - m e t a l input, oil pollution), g r o u n d n e t fishery, dyking, poldering, oil/gas exploitation, military target areas, shipping, intensive r e c r e a t i o n a l use, extraction of sand. 05.01.03.01 S a n d flat, free of vegetation: Pure tidal s a n d flats without s p e c i a l i z e d algal communities. 05.01.03.02 ,,Farbstreiien" sand flat (,,colour-striped sand flat"): S p e c i a l s u b t y p e of tidal s a n d flats, typically with coloured stripes of p h o t o s y n t h e t i c a l l y active layers of m i c r o b e s with a thickness of 2 - 20mm; consisting of a surface s a n d l a y e r (by a c c u m u l a t e d sand) that g l i m m e r s b l u e g r e e n d u e to c y a n o b a c t e r i a , p u r p l e to p i n k d u e to sulfurp u r p l e b a c t e r i a a n d b l a c k c a u s e d by sulfur- a n d i r o n - r e d u c i n g bacteria. Factors of threat: e u t r o p h i c a t i o n a n d pollution ( w a s t e - w a t e r inlets, h e a v y - m e t a l input, oil pollution), g r o u n d n e t fishery, d y k i n g , poldering, oil/gas exploitation, military target areas, shipping, intensive r e c r e a t i o n a l use. 05.01.03.03 S a n d b a n k (eulittoral zone): S a n d b a n k s isolated from t h e coast, b e n e a t h the MHT, falling dry at low tide. Due to their state of relatively low d i s t u r b a n c e they are of i m p o r t a n c e as resting p l a c e for seals. Factors of threat: e u t r o p h i c a t i o n a n d pollution ( w a s t e - w a t e r inlets, h e a v y - m e t a l input, oil pollution), military t a r g e t areas, shipping, intensive r e c r e a t i o n a l use b y boats. 05.01.05 Brackish tidal flats of estuaries: Tidal flats of estuaries p e r i o d i c a l l y falling dry at low tide, with typical l o w e r e d a n d v a r y i n g salinity d u e both to i n f l u e n c e of raina n d river-freshwater. With the loss of the estuaries, also the floodplain a l o n g the e s t u a r y a n d the river h a v e b e c o m e extinct in the Dutch W a d d e n Sea. Restoration m a y b e possible a l o n g the W e s t e r w o l d s e Aa, but only after l a r g e - s c a l e e n g i n e e r i n g a n d m a j o r d e struction. In G e r m a n y there are g o o d possibilities a l o n g the Weser, Elbe a n d Eider.

Biotopes a n d Biotope C o m p l e x e s of the W a d d e n Sea A r e a

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Vegetation communities: T h e r o - S a l i c o r n i e t e a Pign. 53 em. Tx. 55, Ruppion m a r i t i m a e Br.-B1.31 era. D e n Hart. et Seg. 64 u.a. Factors of threat: D y k i n g of river marshes, coastal protection, constructions of e m b a n k ments, eutrophication a n d pollution (fertilizer- and pesticide input by intensive agriculture, w a s t e - w a t e r inlets, h e a v y - m e t a l input, oil pollution), shipping. 05.01.06 Freshwater tidal flats of estuaries: Flats of estuaries, co v er ed with shallow f r e s h w a t e r at high tide a n d periodically falling dry at low tide; no influence of brackish or saltwater e x c e p t during e x t r e m e storms. T h e f r e s h w a t e r tidal regions are extinct in the Dutch part of the W a d d e n Sea. In G e r m a n y possibilities for restoration exist al o n g the rivers Elbe, Weser, H u n t e a n d Eider. Factors of threat: D y k i n g of river marshes, coastal protection, constructions of e m b a n k ments, eutrophication a n d pollution (fertilizer- and pesticide input by intensive agriculture, w a s t e - w a t e r inlets, h e a v y - m e t a l input, oil pollution), shipping. 05.01.07 Eulittoral (old) blue m u s s e l beds: Colony of blue mussels (A4ytilus edulis) in the intertidal region. Blue mussel beds also occur in the subtidal region (see 030207). Mytilus edulis is a species that does not n e e d to live directly on the sea floor but builds thick, multi-layered mussel beds with its byssus. The mussel b ed creates a hard substrate in the sandy or m u d d y surroundings, partly with a more or less d e n s e cover of b r o w n al g ae (mostly Fucus vesiculosus and K serratus). Mussel beds, several years of ag e an d naturally grown, are t h r e a t e n e d . Intertidal beds h a v e b e e n k n o w n to survive for decades. An intertidal bed contains mussels of different a g e classes. Species richness and diversity are positively correlated with a g e and structural complexity of the bed. In a m a t u r e b e d in the W a d d e n Sea 96 associated species h a v e b e e n recorded. In the Dutch W a d d e n S ea it was s h o wn that 25 % of the birds occurred on the m u s s e l beds while these c o v e r e d only 5 % of the area. Intertidal mussel beds h a v e b e e n u n d e r s e v e r e stress in the W a d d e n Sea. In the 1950s f i s h e r m e n tried to destroy them, b e c a u s e the beds w e r e t h o u g h t to be a vector in the spread of a parasite w h i c h was c o n s i d e r e d to be a threat for the mussel culture. Mature beds can n o w be c o n s i d e r e d almost extinct. It has b e e n shown that formation of n e w beds is a rare event, b e c a u s e s e t t l e m e n t in g e n e r a l only takes place on existing beds or remains of old beds. Restoration of intertidal m a t u r e mussel beds may be possible if a complete b an is imposed on destruction of intertidal beds. T h e majority of the d e v e l o p i n g beds will not be stable, but a small part of t h e m m a y d e v e l o p into m a t u r e beds. With the p r esen t fishery regulations and m a n a g e m e n t plans, the m u s s e l b ed should be considered an e n d a n g e r e d biotope. Factors of threat: pollution ( h e a v y - m e t a l input, oil pollution); particularly old, natural stocks are t h r e a t e n e d by intensive fishery. 05.02 Tidal flats with vegetation: Usually m u d flats, with a more or less d e n s e cover of macrophytes. V e g e t a t i o n communities: T h e r o - S a l i c o r n i e t e a Pign. 53 em. Tx. 55, S p a r t i n e t e a Tx. 61, z. T. Zosteretea m a r i n a e Pign. 53. Factors of threat: eutrophication a n d pollution ( w a s t e - w a t e r inlets, h e a v y - m e t a l input, oil pollution), dyking, poldering. 05.02.01 Intertidal Seagrass beds: S l e n d e r sea grass m e a d o w s (Zostera noltii an d Z. marina ssp. angustifolia) in the lowest tidal zone close to MLT or in shallow d ep r essi o n s of the m e d i u m or low tidal flat zones; m o r e or less periodically falling dry for a short time;

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Biotopes a n d Biotope Complexes of the W a d d e n Sea Area

Intertidal seagrass beds have decreased in a b u n d a n c e over the last decades. In a few parts of the W a d d e n Sea the decrease seems to have halted during the last ten years. The beds are very susceptible for s e d i m e n t - d i s t u r b i n g activities. C o m p a r e d with the G e r m a n W a d d e n Sea. only a very small p e r c e n t a g e of the Dutch part is covered b y seagrass. The intertidaI seagrass used to be a n i m p o r t a n t food source for Brent geese. As it is not clear what caused the decline of the intertidal seagrass, it is difficult to predict w h e t h e r the fields will regenerate. Vegetation communities: e.g. Zosteretum noltii Harms. 36, Z o s t e r e t u m m a r m a e partly. Factors of threat: pollution Iwaste-water inlets, h e a v y - m e t a l input, oil pollution), dyking, intensive recreational use, bottom d i s t u r b i n g fishery Icockle fishery). 05.02.02 Tidal flats with saltwort stands (Salicornia): Situated a b o v e the seagrass zone, often monospecific, by root competition very g a p p y stands of different species of Salicornia (S. europaea agg.), g r o w i n g either upright or prostrate. Vegetation communities: Thero-Salicornietea Pign. 53 era. Tx. 55: z.B. S a l i c o m i e t u m dolichostachyae (strictae) K n a u e r 52, Salicornietum ramosissimae Christians. 55 era. Tx. 74. Factors of threat: pollution (waste-water inlets, h e a v y - m e t a l input, oil pollution), dyking. 05.02.03 Tidal flats with Spartina anfflica stands: Thickets of Spartina anfflica in m u d d y flats usually b e t w e e n -40cm M H T a n d +20cm MHT, in protected locations, e.g. at sea e m b a n k m e n t s . Partly located in the higher Salicornia zone and in the Puccinelia zone a n d displacing Salicornia a n d Puccinellia. Often, but in most cases w i t h o u t success, planted for land reclamation purposes. Vegetation communities: Spartinetea Tx. 61: S p a r t i n e t u m anglicae Corill. 53. 05.02.04 Bogs in tidal e r o s i o n (e.g. Sehestedter Moor): The only e x a m p l e on the Germ a n North Sea coast is the ,, Sehestedter Moor", the rest of a S p h a g n u m bog, located in the , , J a d e b u s e n " (Niedersachsen) s e a w a r d of the dike. Its step by step erosion is b a s e d on the d y n a m i c s of tidal currents. Factor of threat: n a t u r a l erosion a n d possibly eutrophication. 05.03 Tidal g u l l i e s and channels: C h a n n e l s a n d gullies formed by tidal currents with steep slopes a n d u n d e r c u t b a n k s located in tidal flats of the North Sea a n d e x t e n d i n g into the shallow water zone; differentiated by width a n d length; periodically or always flooded a n d u n d e r constant changes. Vegetation communities: at c h a n n e l slopes: H a l i m i o n e t u m portulacoides Kuhnh.-Lord. 27, S u a e d a macrocarpa C o m m u n i t i e s e.g. (Asteretea tripolii Westh. et Beeft. in Westh. et al. 62, Thero-Salicornietea Pign. 53 em Tx. 55). Factors of threat: pollution (waste-water inlets, h e a v y - m e t a l input, oilpollution), g r o u n d net fishery, eutrophication, dyking. 05.03.01 Tidal g u l l i e s and c h a n n e l s , p e r m a n e n t l y s u b m e r g e d 05.03.02 C h a n n e l s and gullies, dry at l o w tide 07 Saltmarshes and - m e a d o w s : I n c l u d e s n a t u r a l a n d a n t h r o p o g e n i c meadows, pastures a n d marsh areas that are no l o n g e r grazed or m o w e d on episodically flooded m u d d y a n d s a n d y bottoms on the North Sea coast. Located above MHT, different zones d e p e n d on f r e q u e n c y a n d lasting of flooding a n d on the salinity of the soil water. Mostly composed of p e r e n n i a l grass species a n d h e r b s in compact stands.

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07.01 Lower saltmarshes: Lower zone of the saltmarsh, b e g i n n i n g at the zone of glassworts (Salicornia), ca - 20 cm to + 25 cm MHT, mainly with Puccinella - grassland; + p e r m a n e n t l y u n d e r saltwater conditions, h e l p i n g land reclamation carrying more or less d e n s e vegetation, in most cases d o m i n a t e d by Puccinella maritima. 07.01.01 Lower saltmarshes, natural or seminatural: Lower saltmarsh of the North Sea, rich in structure, not or only very extensively used. Vegetation communities: Puccinellion maritimae Christians. 27 em. Tx. 37, and: SaginoC o c h l e a r i e t u m d a n i c a e Tx. et Gilln. 57. Factors of threat: eutrophication a n d pollution (waste-water inlets, h e a v y - m e t a l input, oilpollution), dyking, poldering, recreational use, sheep grazing. 07.01.02 Lower saltmarshes, a n t h r o p o z o o g e n i c : Lower saltmarsh of the North Sea, due to g r a z i n g or m o w i n g poorer in species a n d structure than natural marshes; mostly very low growth with d o m i n a t i n g Puccinellia maritima. In the whole W a d d e n Sea m a i n land marshes were increased in area by stimulating sedimentation. These m a r s h e s served a function in coastal protection a n d as grazing area for livestock. The m a n - m a d e marshes c o n t a i n the same v e g e t a t i o n types as the natural marshes. Vegetation communities: Puccinellion maritimae Christians. 27 em. Tx. 37, and: SaginoC o c h l e a r i e t u m d a n i c a e Tx. et Gilln. 57. Factors of threat: pollution (waste-water inlets, h e a v y - m e t a l input, oilpollution), dyking, poldering, recreational use, intensification of agriculture. 07.02 Upper saltmarshes oi the North Sea: Following the P u c c i n e l l a - m a r s h l a n d further inland, a b o u t 20-30 cm above MHT, a m a r s h l a n d with less flooded g r a s s l a n d vegetation exists; better air conditions in the soil a n d initial freshwater influence in the grou n d water table; naturally very rich in both structures a n d species. 07.02.01 Upper saltmarshes, natural or seminaturah Upper saltmarsh of the North Sea (e.g. in the ,,Halligen", G e r m a n part of the W a d d e n Sea), rich in structure, no or only very extensive use. Vegetation communities: Armerion maritimae Br.-B1. et de L. 36, v.a. J u n c e t u m gerardii (Warming 06) Du Rietz 23, and: S a g i n o - C o c h l e a r i e t u m d a n i c a e Tx. et Gilln. 57. Factors of threat: eutrophication a n d pollution (waste-water inlets, h e a v y - m e t a l input, oilpollution), dyking, poldering, intensive recreational use, intensification of agriculture. 07.02.02 Upper saltmarshes, a n t h r o p o z o o g e n i c : Upper saltmarsh of the North Sea, due to grazing or m o w i n g poorer in species a n d structure t h a n n a t u r a l marshes. In the whole W a d d e n Sea m a i n l a n d marshes were increased in area by stimulating s e d i m e n t a tion. These m a r s h e s served a function in coastal protection a n d as grazing area for livestock. W h e n the marsh surface had g a i n e d sufficient height, the area was e m b a n k e d by dikes. In the past centuries, e m b a n k m e n t s surpassed n e w accretion so m u c h that the saltm a r s h area today is only a small part of that expected in a reference situation. The m a n m a d e marshes contain the same v e g e t a t i o n types as the n a t u r a l marshes. The m a i n difference is the u n n a t u r a l structure of the d r a i n a g e c h a n n e l s a n d the high density of these channels. The quality of the marsh d e p e n d s greatly on the type of active m a n a g e m e n t . Studies are u n d e r way to develop m a n a g e m e n t t e c h n i q u e s which m a k e a n optimal use of the n a t u ral processes.

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Vegetation communities: A r m e r i o n m a r i t i m a e Br.-B1. et de L. 36, v.a. J u n c e t u m g e r a r d i i (Warming 06) Du Rietz 23. Factors of threat: e u t r o p h i c a t i o n a n d pollution ( w a s t e - w a t e r inlets, h e a v y - m e t a l input, oilpollution), dyking, poldering, r e c r e a t i o n a l use, intensification of agriculture, s h e e p - g r a zing. 07.03 G r e e n b e a c h (Strandwiese): Beach. richly structured by g u l l y - s y s t e m s a n d embryonic d u n e systems, which c a n n o t be a s s i g n e d to the lower or u p p e r s a l t m a r s h b e c a u s e of its small s t r u c t u r e d biotope mosaic. T h e s e biotopes develop b e h i n d d u n e r i d g e s a n d b e a c h barrier systems, but are r e g u l a r l y flooded by seawater. The v e g e t a t i o n is similar to the saltmarshes, but also j u v e n i l e s t a g e s of d u n e s occur. Relics of these d y n a m i c b i o t o p e s occur in slufters a n d on the b e a c h plains. The b i g g e s t threat is coastal p r o t e c t i o n a n d d u n e m a n a g e m e n t both for the a r e a a n d for the quality of the biotope. A n a d d i t i o n a l threat for the quality is disturbance. As is the case with most d y n a m i c b i o t o p e s , restoration is relatively easy. Vegetation communities: e.g. Puccinellion m a r i t i m a e Christinas. 27 em. Tx. 37, A r m e r i o n m a r i t i m a e Br.-B1. et Del. 36, C a k i l e t e a m a r i t i m a e Tx. et Prsg. 50. Factors of threat: e u t r o p h i c a t i o n a n d pollution ( w a s t e - w a t e r inlets, h e a v y - m e t a l input, oilpollution), dyking, poldering, i n t e n s i v e recreational use, intensification of agriculture. 07.04 Brackish and saltwater r e e d marsh: Reeds u n d e r salt- or b r a c k i s h - w a t e r influence on the shores of the North Sea islands, on the mainland, along coastal lakes, in estuaries, on slopes of gullies or in w e t d u n e valleys. The most c o m m o n p l a n t s are Phragmites australis, Bolboschoenus m a r i t i m u s ssp. compactus, S c h o e n o p l e c t u s t a b e r n a e m o n tani .

Vegetation communities: Scirpion maritimi Dahl et Had. 41: S c i r p e t u m m a r i t i m a e Tx. 37, partly P h r a g m i t i o n communis W. Koch 26: P h r a g m i t e t u m vulgaris S o g u l l i e s F a c t o r s of threat: e u t r o p h i c a t i o n a n d pollution ( w a s t e - w a t e r inlets, h e a v y - m e t a l input, oilpollution), dyking, intensive r e c r e a t i o n a l use, agriculture. 07.05 Brackish tall h e r b stands: Tall h e r b stands at coastal l a k e s w h e n s e d i m e n t a t i o n fills t h e m up, in b r a c k i s h fallows, in w e t d u n e valleys or on shores of estuaries. V e g e t a t i o n communities: partly S e n e c i o n fluviatilis Tx. 50, e.g. C o n v o l v u l o - A n g e l i c e t u m a r c h a n g e l i c a e litoralis Pass. 59. Factors of threat: e u t r o p h i c a t i o n a n d pollution ( w a s t e - w a t e r inlets, h e a v y - m e t a l input, oilpollution), dyking, intensive r e c r e a t i o n a l use, agriculture. 07.06 Brackish saltmarsh of estuaries: C o m m o n l y grazed, b r a c k i s h (poldered) m e a dows in tidal zones of estuaries or in n e w polders that are still i n f l u e n c e d b y salt water. The latter u n d e r l i e a n o n g o i n g d e c r e a s e in saiinity a n d evolve in most c a s e s to intensiv e l y u s e d grassland, or b e c o m e a r a b l e land. V e g e t a t i o n communities: A r m e r i o n m a r i t i m a e Br.-BI. et De L. 36, Potentillion a n s e r i n a e Tx. 47 (Agropyro-Rumicion). Factors of threat: e u t r o p h i c a t i o n a n d pollution ( w a s t e - w a t e r inlets, h e a v y - m e t a l input, oilpollution), dyking, intensification of agriculture. 09 Sands and beaches: S a n d y a n d g r a v e l b e a c h e s in the coastal zones of North S e a islands a n d the m a i n l a n d ; l o c a t e d a b o v e the p e r i o d i c a l l y flooded zone. C o m m o n l y free of vegetation, often with drift lines or a c c u m u l a t i o n s of d e a d a l g a e a l o n g the shore, p a r t l y c o v e r e d with vegetation.

Biotopes and Biotope Complexes of the W a d d e n Sea Area

27

09.01 S a n d b a n k s and sands of the backshore: Sands above MHT that are only episodically flooded a n d isolated from the coast; d u n e evolution possible w h e n s a n d supply is high. Factors of threat: intensive recreational use, shipping, pollution (e.g. oil, other harmful organic matter, garbage, h e a v y - m e t a l input). 09.02 Sandy beach: Sand flats at the coast, i n f l u e n c e d by salt a n d airborne s a n d drift. S a n d y b e a c h e s are in most cases free of vegetation, but often with drift lines of organic matter (e.g. algae a n d sea grass) along the shore, the u p p e r ones are often covered with characteristic v e g e t a t i o n (Atriplicon littoralis); i n c l u d e d are also sand flats covered with Salicornia dofichstachya ssp. (decumbens) with strong airborne sand drift; on n a t u r a l beaches d u n e s occur landward. The quality of the r e m a i n i n g n a t u r a l a n d u n d i s t u r b e d beaches has shown a considerable deterioration, m a i n l y because of disturbance. W h e l p i n g areas for seals, a n d b r e e d i n g areas for spezialized birds have b e c o m e rare or extinct. With proper policies a n d m a n a gement, restoration will be simple a n d promising. Migrating sandbars as a very d y n a m i c biotope can cover part of the sand beach. Few organisms will use these as a p e r m a n e n t biotope. Only locally will m a n a g e m e n t activities (dredging, coastal defence etc) influence the r e l e v a n t physical processes. The biotope is considered qualitatively a n d q u a n titatively healthy a n d will restore quickly if affected. Vegetation communities: Cakiletea maritimae Tx. et Prsg. 50, H o n k e n y o - E l y m i o n Tx. 66 em. G~hu et Tx. in G 6 h u 75, Salicornietum x d e c u m b e n t i s Schwabe et Tx. 77. Factors of threat: intensive recreational use, pollution (e.g. oil, other harmful organic matter, garbage, h e a v y - m e t a l input), r e m o v i n g of drift line material (cleaning of the beach). 09.05 Beach ridge: Ridges in different mixtures of sand, pebbles, shingles, algae a n d water plants, s e d i m e n t e d on the b e a c h by d y n a m i c processes, such as currents, b r e a k e r s a n d ice pressing. D e p e n d i n g on its degree of succession, free of vegetation or covered with drift line communities. At erosive coasts w h e n fossil peat layers are eroded, b e a c h ridges of peat or clay may occur. Factors of threat: intensive recreational use, coastal defence measures. 09.06 Coastal lake: Lakes with brackish- or saltwater, separated from the sea at low water levels by b e a c h ridges or large spits, in most cases poor in vegetation. Location a n d habitus can often be c h a n g e d by flooding. With growing beach ridges a n d spits, such lakes can become totally separated from seawater inflow and will develop into brackisha n d later into freshwater lakes of the hinterland. Brackish i n l a n d waters are in most cases not n a t u r a l biotopes in the W a d d e n Sea region, but they can be important from a n a t u r e conservation point of view. In G e r m a n y some large saline or brackish still waters occur. In The N e t h e r l a n d s it would be possible to develop the L a u w e r s m e e r into such an area. Before the dikes were of the s t a n d a r d as we k n o w them today, s e e p a g e of saline water caused small brackish pools or terrestrial biotopes to develop b e h i n d the dikes. These biotopes are rare. They m a y contain the last r e m n a n t s of a n earlier a b u n d a n t brackish-water fauna. Restoration is relatively easy by letting in salt water, but this m a y interfere with agricultural practice. Vegetation communities: e.g. Ruppion maritimae Br.-B1. em D e n Hart. et Seg. 64. Factors of threat: intensive recreational use, pollution (e.g. waste-water inlet, oil, heavymetal input), d u m p i n g of e.g. garbage.

28

Biotopes a n d Biotope Complexes of the W a d d e n Sea Area

10 C o a s t a l dunes: S a n d y hills of different height, a c c u m u l a t e d by w i n d along sandy beaches. D e p e n d i n g on its degree of succession, poor in vegetation, c o v e r e d with fragm e n t a r y grass stands, h e a t h e r or shrubberies. 10.01 P r i m a r y dunes: Beginning of d u n e succession: e m b r y o m c or p r i m a r y dunes. These are low, often strongly w i n d s w e p t a n d saltwater-influenced sand a c c u m u l a t i o n s on the u p p e r beach; in most cases, only f r a g m e n t a r y cover of tufts of Agropyrum junceum is present. W h e n series of b e a c h d u n e s c o n n e c t with each other, d u n e ridges are formed. A y o u n g d u n e ridge is p r e s e n t for e x a m p l e at the Hors on Texel. but to a l a r g e extent it should be considered m a n - m a d e . D u n e ridges are not threatened, but due to their scale, r e g e n e r a t i o n will be difficult. Vegetation communities: Agropyrion lunceiformis G~hu 75. Factors of threat: trampling through intensive recreational use. 10.02 White dunes: Secondary dune: already higher d u n e with preliminary developm e n t of freshwater lentils, groundwater in most cases still rich in lime (shell fragments); undergoes p e r m a n e n t sand accumulation, mostly fragmentary cover with b e a c h grass. (Ammophila arenana, Ammocalamagrostis x baltica); The term secondary refers to the stage of d u n e development, while the d u n e formation itself may be primary or secondary due to erosion on coastlines with break-throughs in older d u n e ridges or due to h u m a n degradation. Vegetation communities: A m m o p h i l i o n a r e n a r i a e (Br.-BI. 33) Tx. 55, Honkenyo-Elymion Tx. 66 era. GShu et Tx. in G 4 h u 75. Factors of threat: protective m e a s u r e s against erosion, coastal defence m e a s u r e s , recreational use (camping, trampling). 10.03 G r e y d u n e g r a s s l a n d : Last stage of d u n e d e v e l o p m e n t before d u n e scrub invades with more or less d e n s e v e g e t a t i o n cover; in most cases higher d u n e , with a n already decalcified, acidic u p p e r soil layer with low moderate h u m u s a c c u m u l a t i o n ; only little sand supply, covered with n u t r i e n t poor grassland on sand; in some cases d e n s e moss or lichen carpets can develop. Vegetation communities: Koelerion a r e n a r i a e Tx. 37 corr. G u t e r m a n n et M u c i n a 93 (Koelerion albescentis), C o r y n e p h o r i o n canescentis Klika 31, Thero-Airion Tx. et Oberd. 57, u.a. Factors of threat: protective m e a s u r e s against erosion, coastal defence m e a s u r e s , recreational use (camping, trampling). 10.04.01 D u n e s a n d c o a s t a l h e a t h w i t h Empetrum nigrum: Primary, dwarf h e a t h e r on b r o w n d u n e s with cranberries (Empetrum nigrum) at locations that u n d e r g o only little a c c u m u l a t i o n of w i n d - b l o w n sand. The soil is acidic with distinct a c c u m u l a t i o n of humus, a b e g i n n i n g podsolization is visible. Vegetation communities: Empetrion nigri B6ch. 43. Factors of threat: protective m e a s u r e s a g a i n s t erosion, coastal defence m e a s u r e s , recreational use (camping, trampling), g a m e bite off. 10.04.02 D u n e h e a t h w i t h Calluna vulgaris: Brown d u n e , in most cases p r e s e r v e d by extensive g r a z i n g with d o m i n a t i n g c o m m o n h e a t h e r (Calluna vulgaris). T h e soil is acidic with distinct a c c u m u l a t i o n of humus, a b e g i n n i n g podsolization is visible. Vegetation communities: Genistion pilosae Duv. 42 em. Schub. 60: v.a. G e n i s t o anglicaeC a l l u n e t u m Schwick. 33 em. Tx. 75. Factors of threat: protective m e a s u r e s a g a i n s t erosion, coastal defence m e a s u r e s , recreational use (camping, trampling), g a m e bite off esp. by rabbits.

Biotopes a n d Biotope Complexes of the W a d d e n Sea Area

29

10.05 Wet dune valleys (incl. coastal fens): Valley within the d u n e belt of the coast, formed by deflation or coastal erosion. Due to g r o u n d w a t e r contact covered with swampor f e n v e g e t a t i o n that varies with the salinity a n d the a m o u n t of lime in the soil. Because precipitation is higher than evaporation, the v e g e t a t i o n loses the salt-tolerant taxa. Dune valleys occur on all islands, a n d in The N e t h e r l a n d s the majority is m a n a g e d as nature reserve. W h e n d u n e valleys are not completely separated from the sea or w h e n the sea breaks through the ridge, a slufter is formed. Within wet d u n e valleys d u n e lakes m a y occur: The first type of d u n e lake develops w h e n eolian erosion in d u n e valleys proceeds until the soil water level is reached. This type of d u n e lake develops on bare s a n d a n d is poor in nutrients. M a n y of these lakes are temporary, a n d contain water only in winter. The second type of d u n e lake develops w h e n a d u n e system enlarges a n d the g r o u n d w a t e r level rises. These d u n e lakes develop in v e g e t a t e d valleys a n d are relatively rich in nutrients. P e r m a n e n t d u n e lakes are rare a n d their biggest threat is exploitation of freshwater for i n h a b i t a n t s a n d tourists. Vegetation communities: Asteretea tripolii Westh. et Beetf. in Westh. et al. 62, Phragmitetalia Koch 26, I s o e t o - N a n o j u n c e t e a Br.-B1. et Tx. ex Westh. et al. 46, Ericetum tetralicis Jonas 32, Bolboschoenetalia maritimi H e j n ' y in Holub e t a l . 67. Factors of threat: trampling through recreational use, exploitation of freshwater or groundwater, eutrophication. (23.01) Dune brooks: Because the a m o u n t of rainfall exceeds the a m o u n t of evaporation, and the g r o u n d water level is higher t h a n the s u r r o u n d i n g area or sea, water seeps out of dunes. W h e n d u n e systems are of sufficient size, brooks or creeks m a y be formed. T h e s e can either flow towards land or towards the sea. If the latter is the case, a n extremely rare s u b t y p e develops, In its optimal form it r e s e m b l e s a small estuary c o n t a i n i n g the whole r a n g e from m u d flat, saltmarsh to freshwater marsh. The only relatively well preserved e x a m p l e in The Netherlands is the Moksloot on Texel, although saltwater intrusion is p r e v e n t e d there. Further restoration seems possible. 10.00 Dune scrub {natural): With o n g o i n g succession on d u n e s (with e.g. sand consolidation, e n r i c h m e n t of humus, decalcified soil), shrubs a n d thickets of Salix repens, Hippophae rhamnoides, or Rosa pimpinellifolia develop; Thickets of n o n a u t o c h t h o n o u s species, e.g. Rosa rugosa, are not included. Vegetation communities: e.g. Salicion a r e n a r i a e Tx. 52, z.T. P r u n o - R u b i o n fruticosi Tx. 52 corr. Doing 62. Factors of threat: p l a n t a t i o n of n o n a u t o c h t h o n o u s species, trampling t h r o u g h recreational use, game bite off. 10.07 Natural shifting dune: D u n e m o v i n g due to strong airborne sand drift in the m a i n w i n d direction; almost free of vegetation. This type of d u n e can be primary w h e n no v e g e t a t i o n has developed. It can also b e of a secondary nature after v e g e t a t i o n has disappeared. In G e r m a n y it usually develops after d e g r a d a t i o n of white-, grey- a n d b r o w n dunes. Because of intensive m a n a g e m e n t a n d stimulation of v e g e t a t i o n development, this type has become rare; restoration is relatively simple. Factors of threat: protective m e a s u r e s a g a i n s t erosion (conservative m a n a g e m e n t ) 11.03 P l e i s t o c e n e cliffs: Smaller pleistocene cliffs {Geest) along the coast of the North Frisian islands. Factors of threat: direct destruction by b u i l d i n g s a n d constructions.

30

Biotopes a n d Biotope Complexes of the W a d d e n Sea Area LEGEND TO THE TABLES

The t h r e e - p r o n g e d criteria system used in the red list of biotopes a n d biotope complexes comprises two criteria of threat a n d an assessment of the r e g e n e r a t i o n ability. The criteria of threat are g i v e n for each s u b r e g i o n separately, while the a s s e s s m e n t of the reg e n e r a t i o n ability, together with a n overall evaluation of threat, is given for the region of the W a d d e n Sea as a whole.The 'Red List of Biotopes a n d Biotope Complexes of the Wadd e n Sea Area' is an alphabetical list g r o u p e d by the overall status of biotope types a n d biotope complexes.The 'List of t h r e a t e n e d Biotopes a n d Biotope Complexes of the Wadd e n Sea Area' gives a complete hierarchical red list of the biotope complexes a n d biotopes of the W a d d e n Sea area with regional evaluations, m a i n threats a n d further information. DE: Threat by direct destruction (Loss of area) QU: Threat by gLU_alitative changes (loss of quality, i.e. adverse c h a n g e s of abiotic conditions, of the typical structural a p p e a r a n c e , a n d of the typical set of p l a n t a n d a n i m a l species present). 0 completely destroyed 1 t h r e a t e n e d by complete destruction 2 heavily e n d a n g e r e d 3 e n d a n g e r e d (vulnerable) P potentially e n d a n g e r e d (susceptible); this category is only used for criterion DE (threat by direct destruction) * p r e s u m a b l y not e n d a n g e r e d at p r e s e n t not p r e s e n t OE: O v e r a l l e v a l u a t i o n of threat: Both criteria ldirect destruction a n d qualitative change) are c o m b i n e d to give a n overall classification of threat. This has at least the same level of the highest r a n k given in one of the criteria I a n d II

RE: Assessment of regeneration ability

N K S B X r e g e n e r a t i o n impossible r e g e n e r a t i o n hardly possible ~> 150 years necessary) r e g e n e r a t i o n difficult (15-150 years) r e g e n e r a t i o n conditionally possible E less t h e n 15 years) classification not m e a n i n g f u l

Further information G e r m a n codes a n d assessments of threat a n d r e g e n e r a t i o n ability are according to the National Red Data Book of Biotopes (Riecken et al., 1994). The trilateral approach, d e a l i n g with one specific n a t u r a l region, m a d e it possible a n d necessary to i n c l u d e some specific t h r e a t e n e d subtypes of biotopes. T h e y h a v e b e e n listed in addition to the lists of the National G e r m a n Red Book.

The EEC Habitats Directive Code The habitats directive (Council Directive 92/43/EEC of 21st May 1992 on the conservation of n a t u r a l habitats a n d of wild f a u n a a n d flora) aims at a E u r o p e a n n e t w o r k of pro-

Biotopes and Biotope Complexes of the Wadden Sea Area

31

tected sites of Community importance called NATURA 2000. In Annex I natural habitat types are listed ,,whose conservation requires the designation of special areas of conservation". To facilitate the implementation of this Directive and to give transparency for political decisions and in m a n a g e m e n t plans of the Wadden Sea Area, the officially used codes of the Standard Data Entry Form (Doc. Habitats 94/4, Commission of the EEC) are given in this column. *: An asterisk refers to ,,priority" habitats with ,,particular responsibility for their conservation" according to Annex I of the Habitats Directive. Main Threat Factors Under this heading a rough indication of the main negative impacts on biotope complexes and biotopes is given, using a system worked out by Ole Norden, Ministry of Environment, C o p e n h a g e n (Denmark). A different system from that devised for the red lists of species was necessary to meet the special needs involved in an assessment of the biotopes. Biotope loss (permanent~ irreversible: HL) HLB Construction, dredging, dumping HLM Mineral extraction (prospecting, mining, dredging) HLW Water regulation (land reclamation, drainage, coastal defence, dyking) Biotope change (semi-permanent, reversible: HC) HCA Agriculture (intensive, changing, land reclamation, stop of traditional farming) HCB Construction, dredging, dumping HCC Climate change (largely irreversible to man) HCE Eutrophication (fertilization, sewage, combustion) HCF Fishing HCG Wear (traffic, tourism) HCM Mineral extraction HCP Pollution (non-eutrophication) of air, earth and water (pesticides, waste disposal, sewage, combustion) HCT Forestry HCW Water regulation (drainage, rerouting, extraction Biotope disturbance (temporary: HD) HDA Agriculture, forestry HDB Construction, dredging, dumping, mineral extraction HDF Fishing Specimen loss (SL) SLC Displacement, competition, disease, parasites SLE Exploitation SLP Pest control SLT Traffic Specimen disturbance (SD) SDR Recreation (hunting, tourism, sports) SDT Traffic

32

B i o t o p e s a n d B i o t o p e C o m p l e x e s of t h e W a d d e n S e a A r e a

Acknowledgements: Gratitude should be expressed to D. Boedeker, BfN INA Insel Vilm for a preliminary translation of the German biotope definitions, H. v. Nordheim (BfN, INA Insel Vilm), U. Riecken (BfN, Bonn), B. Reineking (CWSS, Wilheimshaven), Dr. H. Farke {Wadden Sea National Park of Lower Saxony), D. Fleet (Wadden Sea National Park of Schleswig-Holstein), J. Thissen (National Reference Centre for Nature Management, Wageningen) for critical discussion of the manuscript. O. Norden Andersen, Copenhagen, was so kind as to provide a suitable classification of the main threat factors.

REFERENCES Blab, J., Riecken, U. & A. Ssymank, 1995. Proposal on a criteria system for a National Red Data Book of Biotopes. - Landscape Ecol. 10, 43-50. Dijkema, K. S. & W. J. Wolff (Eds), 1983. Flora and vegetation of the Wadden Sea islands and coastal areas. In: Ecology of the Wadden Sea. Ed. by W. J. Wolff. Balkema, Rotterdam, 3, 9/1-9/413. Dijkema, K. S., 1989. Habitats of the Netherlands, German and Danish Wadden Sea. Research Institute for Nature Management, TexeL 24 maps, 30 pp. Leerdam, A. van, Wassen, M. J. & N. Dankers, 1993. Onderzoek nagenoeg-natuurlijke referentieecosystemen. Rijksuniversiteit, Utrecht, 84 pp. Riecken, U., Ries, U. & A. Ssymank, 1994. Rote Liste der Biotoptypen der Bundesrepublik Deutschland. SchrReihe Landschaftspfl. Natursch. 4 I, 184 pp. Westhoff, V., Hobom, C. & J. H. J. Schaminee, 1993. Rote Liste der Pflanzengesellschaften des Naturraums Wattenmeer unter Berticksichtigung der ungef~ihrdeten Vegetationseinheiten. - Tuexenia 13, 109-140.

RED LIST O F B I O T O P E S A N D B I O T O P E C O M P L E X E S O F T H E W A D D E N S E A AREA" Code

0 - Completely destroyed:

Type B

sublittoral oyster reefs

1 - Threatened by complete destruction: estuaries (natural & seminatural) migrating islands s h e l t e r e d b a y s ( r e d list c a t e g o r y 1-2) d u n e - t i d a l fiats t r a n s i t i o n z o n e mainland saltmarsh complex, natural p l e i s t o c e n e - t i d a l flats t r a n s i t i o n z o n e B e n t h i c z o n e , m u d d y a n d s a n d y b o t t o m s , r i c h in m a c r o p h y t e s b o g s in t i d a l e r o s i o n (e.g. S e h e s t e d t e r M o o r ) b r a c k i s h s a l t m a r s h of e s t u a r i e s b r a c k i s h t i d a l fiats of e s t u a r i e s coastal lake f r e s h w a t e r t i d a l flat of e s t u a r i e s

03.02.08

C3.01 C1.02 C3.02 03.02.05 05.02.04 07.06 05.01.05 09.06 05.01.06

L L L C C C B B B B B B

*Note that types that are threatened in only one subregion are not included b e c a u s e according to the a g r e e m e n t (see 'Introduction') the overall evaluation was only given if at least two regional evaluations were present. Types: L: Landscape complex; C: Biotope complex; B: Biotope.

Biotopes a n d Biotope Complexes of the W a d d e n Sea Area lower saltmarshes, natural or s e m i n a t u r a l sublittoral sabellaria reefs subtidal seagrass beds u p p e r saltmarshes, n a t u r a l or s e m i n a t u r a l 07.01.01 03.02.09 03.02.05.01 07.02.01
B B B B

33

2 - Heavily endangered:

n a t u r a l complexes of tidal fiats slufter (dune valleys in open c o n n e c t i o n to the sea) washover benthic zone, m u d d y and s a n d y bottoms with macrophytes other than seagrass benthic zone, stony a n d hard bottoms, few macrophytes benthic zone, stony a n d hard bottoms, rich in macrophytes brackish a n d salt water reed m a r s h d u n e brooks eulittoral (old) blue mussel beds g r e e n b e a c h (Strandwiese) grey d u n e grassland intertidal seagrass beds m u d flats, free of vegetation n a t u r a l shifting d u n e pleistocene cliffs sublittoral (otd) blue mussel beds tidal flats with saltwort stands (Salicornia) wet d u n e valleys (incl. coastal fens)
3 - Endangered:

A3 C1.03 C3.03 03.02.05.02 03.02.04 03.02.06 07.04

C C C B
B B B B B B B B B B B B B B

(23.01)
05.01.07 07.03 10.03 05.02.01 05.01.01 10.07
11.03

03.02.07 05.02.02 10.05

complete gully system ebb delta island saltmarsh complex tidal divide b e n t h i c zone of the coastal w a t e r flat s a n d y areas, free of v e g e t a t i o n mixed flats, free of vegetation pelagic zone of the coastal w a t e r 'priels' a n d gullies, dry at low tide primary d u n e s sand b a n k a n d sands of the backshore s a n d flats, free of vegetation sandy b e a c h tidal gullies, p e r m a n e n t l y s u b m e r g e d u p p e r saltmarshes, anthropozoogenic white d u n e s

A3.03 A3.02 C1.01 A3.01 03.02 05.01.03.01 05.01.02 03.0I 05.03.02 10.01 09.01 05.01.03 09.02 05.03.01 07.02.02 10.02

C C C C
B B B B B B B B B B B B

34

Biotopes and Biotope Complexes of the Wadden Sea Area LIST OF THREATENED BIOTOPES AND BIOTOPE COMPLEXES OF THE WADDEN SEA AREA

Status o(,.;eat in the

subregions of the I J

Biotope type

Main Threats

Complex functional landscape units:

Mrnns gil d iag ta s

Sheltered bays Estuaries (natural and seminatural)

A3, C1, C3 ] HLW (coastal .,T'-"'.////~-., A2, A3, C1 I . . H L W . . I 1160

Biotope complexes: 03.01, 03.02 all subtypes HCB, HCE, HCF, HCP, HLM HCP, SLE, HCF, HDF :]5.01.01-.03, 05.01.07, 09.01 :)5.01.01-.03, 05.03 03, 05, esp. 05.03 07, 05, 09. 10.01-10.03 07.01.01, 07.02.01, 05.01.01, 05.02, 05.03 HDA, HLW HCA SDR, SLE, HCF, HDF HLB HLB, HCF, HDF

07, 05, 09, 10.01. HDA, HCA .=sp. 07.01.01, HCC, SDR 07.02.01 10.01-10.03. 07, 05 05, 09, 10, 11.03 05, 09, 10 HLW (coast= protection) HLW

Biotopes and Biotope Complexes of the Wadden Sea Area

35

Code:

Status of threat in the subregions of the

]----7 I I Main biotope types

Biotope type

Main
Threats

Pleistocene- tidal flats transition zone "Washover"

09, 05 09, 10

HLW (coastal protection) HLW (coastal protection)

Part I1: Biotope types

Biotope type

[----r

Main Threats

Habitat codDieleCotifEEC

Coastal Water Pelagic zone of the coastal Benthic zone of the coastal Benthic zone, muddy bottorr few macrophytes Benthic zone, sandy bottom (fine-middle sand), few macrophytes Benthic zone, coarse sand, gravel and shell bottoms, few macrophytes Benthic zone, stony and bar bottoms, few macrophytes Benthic zone, muddy and s~ bottoms, rich in macrophyte~ Subtidal seagrass beds Benthic zone, muddy and s~ bottoms with other macroph' Benthic zone, stony and har bottoms, rich in macrophyte~ incl. artificial substrates Sublittoral (old) blue mussel beds Sublittoral oyster reefs Sublittoral sabellada reefs 1170 1170 1170 ~artly incl. (1110)

36

Biotopes and Biot0pe Complexesof the Wadden Sea Area

Part Ih B i o t o p e t y p e s Status of threat in the subregions of the Wadden Sea Area NL D DK

Code:

Biotope type
(NL) "Seamoss"-stands (Sertularia, Hydralmannia) Artificial benthic hard substrates

OE

RE

Main Threats

Habitat Directive Codes of EEC

(D) 1

DE IQU DE QU DE QU

03.02.10
03.09 Tidal fiats

2
* 3 *

2
*

B X

HCF, HCP

idal fiats free of vegetation Mud fiats, free of vegetation Mixed fiats, free of vegetation Sand fiats, free of vegetation Flat sandy areas, free of vegetation "Farbstreifen"-sand fiat Sandbank (eulittoral zone) Brackish tidal flats of estuaries 1 0

l I
2 3 3 3

* 2 2 2 2

2 3 3 3 3 1 2 3 3 3 2 2-3 2

S S S

HCF, SDR
HCF, SDR

1140 1140 1140

HCF, SDR ,I B B HCE, HCA, HLB, HLW, HCP HCE, HCA, HLB, HLW. HCP HCF, HCP

1140 1140 1140 1130

/
L

* 2

Freshwater tidal flat of estuaries Eulittoral (old) blue mussel beds Tidal fiats with vegetation Intertidal seagress beds Tidal fiats with saltwort stands (Salicornia) ]idal fiats with Spartina anglica stands Bogs in tidal erosion (e.g. Sehestedter Moor) Tidal gullies, pdels

1 2

0 1 1

2 3

2 2-3

S S

1130 1170

2 2

2 3 *

K B X

HCF, HCE

partly incl. 1310 1320

'
1 1

"
1

I
N HL (natural) 7110 u.a.

,,, ,,
HCF, HLB
!

Tidal gullies, permanently submerged 'Pdels' and gu{lies, dry at low tide ~adows 07.0~.01 07o201 10.1 10.1 Lower saltmarshes, natural or seminetural Upper saltmarshes, natural or seminatural

3 3

" *

3 3

* 3

2 3

B B

HCF, HLB HCF, HLB

1 1

1 1

1 1

1-2 1-2

2 2

HLW HLW

1330 1330

Biotopes and Biotope Complexes of the Wadden Sea Area

IPart Ih Biotope types
Status of threat in the subregions of the Wadden Sea Area L DEIQU Upper saltmarshes, anthropozoogenic Green beach (Strandwiese) Brackish and salt water reed marsh Brackish tall herb stands 10.1 Brackish saltmarsh of estuaries 0 3 2 1 3 2 2 ]E QU D[ !-3 1 2 2 2 3 2 2 2 2 < QU X S HCA, HLW SDR, HLW HDA, HCA, HLW SDR, HLW SDR, HLW

37

Code:

Biotope type
(NL) 10.3 10.4

RE

Main Threats

Habitat Directive Codes of EEC

(D) 1 o7.02.02 07.03 07.04 07.05 07.06

1330 1330, 2110

B B B

1130

~ands and beaches

09.01 09.02 09.05 09.06 18 3 4 Sand bank and sands ofthe backshore Sandy beach Beach ridge (Strandwall) Coastallake 2 3 3 3 2-3 2 2 HLW SDR, HLW SDR, HLW partly incl. partly incl. partly incl. "1150

Coastal dunes
10.01 Prima~ dunes 3 3 J 3 2 HCG, SDR HLW (coastal protection), HCG HLW (coastal protection), HCG HLW, HCG HLW, HCG HLW, HCG HLW, HCG 2 SLC, HCG HLW (coastal protection), HCG HLB, HLW, HCG 1230 2190 2190 2160, 2170 2110

10.02

21

White dunes

2120

10.03

(24)

Grey dune grassland

2!3

"2130, 2320

1004C

Dunes and coastal heath with

EmpetnJm nigrum
Dune heath with Calluna vulgaris 22, 2 26 Wet dune valleys (incl. coastal fens) Dune brooks Dune scrub (natural)

2 ~-3

!-3 2 I-2

*2140

1004 C

10.05 (23.0" 10.06

*
2

3
2

2
? 3

10.07

28

Natural shifting dune

11.03

~ 12:1

Pleistocene cliffs

1 C o d e s a c c o r d i n g to R i e c k e n et al., 1994.