WIWO Forward Plan 1999-2003

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1.1 WIWO: an initiative-supporting organisation of volunteers
1.2 Forward Plans 1989-1993 and 1994-1998
1.3 Forward Plan 1999-2003: no roads, only directions

2.1 Goals and working area of WIWO
2.2 African-Eurasian Migratory Waterbird Agreement
2.3 Organisation and working methods
2.4 Two lines of research: census work and migration ecology
2.5 Conservation and education

3.1 Introduction
3.2 Financial account
3.3 Evaluation of Forward Plan 1989-1993
3.4 Evaluation of Forward Plan 1994-1998
3.5 A cross-section of projects

4.1 General framework: the life-cycle approach
4.2 The annual life-cycle of migrant waders and waterfowl
4.3 Regional discussion of research priorities

ANNEX 1 References
ANNEX 2 List of WIWO-publications
ANNEX 3 Project participants



The Working Group on International Wader and Waterfowl Research (WIWO) was established in 1983, in order to create a well-defined and accessible intermediary between financing organisations and governmental bodies on the one hand, and volunteer ornithologists interested in studying waterbirds in countries with little own potential for such studies on the other. Starting with the first Netherlands Ornithological Mauritania Expedition (NOME 1980), a large number and diversity of projects have been carried out under WIWO's umbrella (figure 1.1, see chapter 3 for a review).
From the very start, the essence of WIWO has been its function as an intermediary: WIWO merely provides people willing to initiate waterbird studies abroad with assistance and guidance in logistic matters and with application for funds. The foundation's role as an initiative-supporting rather than an initiative-developing organisation bears upon the level of detail with which plans for the future can be specified. Which research issues are taken up and explored will primarily depend on the interest and availability of time and technical resources of the volunteers involved.


1.2 FORWARD PLANS 1989-1993 and 1994-1998

As the number and scope of WIWO's activities expanded in the 1980's, it became necessary to describe their scientific interrelations between and the motivation behind these activities, both as a means to provide a framework for guiding initiatives for future projects, and to provide financiers with information upon which a more stable form of funding could be based. This led to the Forward Plans 1989-1993 and 1994-1998 (WIWO 1989, 1994), in which such a framework was laid out. The Forward Plans identified the major gaps in our knowledge of the distribution, life cycle and population dynamics of migratory waterbirds, and translated these into a working scheme in which specific projects were listed that would especially merit carrying out. The previous Forward Plans served well in structuring initiatives and activities of the many people that together constitute the working force of WIWO (see review in chapter 3).



The current document reviews WIWO activities in the past period and sets the stage for the years to come. The second part of this document is a successor of the Forward Plan 1994-1998 in its intention to provide a guideline for future activities. Its primary aim is to provoke and lend some structure to initiatives by potential project organisers and participants. In short, the plan shows no specific roads, only general directions. Secondly, it may serve as a background document for funding organisations in evaluating the resulting project proposals.
In line with the initiative-supporting role of WIWO, it was felt that a detailed planning of future activities at the project level would be undesirable and unrealistic for an organisation relying so strongly on volunteers. Therefore, this document does not contain a timetable listing specific projects planned for the coming years, but rather tries to summarise the state of our current knowledge on the distribution and movements of migratory waterbirds and the resources and limitations involved. Based on this summary it then identifies topics that need to be addressed with priority in order to provide a level of understanding that is necessary for an effective management policy concerning these groups of birds.
The general lay out of the document is as follows. Chapter 2 restates the aims and describes the general organisation and work method of WIWO. Chapter 3 reviews activities carried out under its umbrella to date, and evaluates the extent to which the research issues identified in the Forward Plan 1989-93 has been addressed. In chapter 4, the current state of knowledge of waterbird distribution and biology is concisely reviewed and major gaps are identified. In this chapter, two complementary approaches are used. The first one takes the birds as the focal unit and evaluates our knowledge of the different phases in the annual life cycle (and important links between them). The second approach takes a geographical perspective, for a number of predefined regions reviewing to what extent we know where the most important wetlands are situated, which species use them in which stage of their annual life cycle, and what is the resource base on which the areas' importance rests.
The evaluation of previous WIWO-projects (see chapter 3) shows the following trends. (a) Long-term studies and repeated surveys at specific localities generate continuity in contacts with counterpart organisations. This leads to an increase in local support, interest and experience in waterbird and wetland studies ('expert capacity'). Ultimately, this may lead to more conservation awareness and action. (b) Life-cycle studies: following one species year-round identifies gaps in knowledge and gives insight in flyway connections, such as within the East Atlantic and Mediterranean Flyways (figure 1.2) and bottlenecks. (c) Distribution-oriented studies: identifying the international importance of wetlands with waterbird surveys contributes significantly to the identification of Important Bird Areas (e.g. Grimmett & Jones 1989).
In addition to the research priorities identified in chapter 4, the following suggestions could be made for future WIWO-studies. (1) To give more attention to priority species (conservation status 'Globally threatened' and 'Vulnerable' in Tucker & Heath 1996). (2) To include other wetland-dependent species, like hirundines and warblers. Often these studies can easily be combined with waterbird counts in the same area. Repeated surveys are needed due to the considerable variation in rainfall, water level and habitat availability within and among seasons and years, which is characteristic for many inland African wetlands. (3) To give more attention to intra-African migrants. Several species have migrations within Africa and are highly dependent on the wet and dry seasonality.




The goals of the foundation WIWO have been formulated as follows:

1. the stimulation, initiation and execution of scientific research into waders and other waterbirds and their haunts, for the benefit of nature conservation as well as the transfer of knowledge;

2. the transfer of knowledge of management of wetlands and wetland bird populations.

These goals allow a great variety of worldwide research activities on any species of waterbird. However, practical considerations have led WIWO to focus on migratory waterbirds in areas along the eastern coasts of the Atlantic Ocean. The focus on waders reflects the present personal interests of people most involved with WIWO activities, while the geographic focus arose from the connections between the areas studied and the Dutch Wadden Sea. More recently, activities have expanded both geographically (especially in the Mediterranean, Black Sea region and Middle East) and in the choice of subjects (e.g. breeding bird censuses of wetlands, non-passerines). With the focus on the research priorities identified in chapter 4, the range of WIWO's activities can be described as 'studies on waterbirds worldwide, with special attention to migratory species in wetlands in the Afro-Palaearctic migration system'.



The African-Eurasian Waterbird Agreement (AEWA) was formulated in 1995 and was signed by about 20 countries by May 1998. The agreement is based on the Bonn Convention, which acknowledges the importance of the conservation of migratory species. The signing parties of the Bonn Convention agreed to take action in this respect and in particular to endeavour the provision of immediate protection for migratory species that are endangered and to endeavour the conclusion of agreements for the conservation and management of species which have an unfavourable conservation status. The African-Eurasian Waterbird Agreement is such an agreement.

The African-Eurasian Waterbird Agreement aims to create a legal basis for the concerted conservation and management policy by the Range States (about 120) for migratory waterbird species. In total 170 species are covered. The agreement covers an area of about 60 million square kilometres consisting of whole Africa and Europe, North-eastern Canada, Greenland and Western Asia up to and excluding Pakistan, Afghanistan, China and the Lena basin of the Russian Federation.

The agreement has two legally binding parts consisting of the Agreement text describing the philosophy, legal framework and provisions and the Action Plan describing the conservation actions to be taken. The Action Plan is up to now limited to geese, swans, ducks, and spoonbills, ibises and storks. The implementation of the agreement is based on two fundamental principles consisting of to undertake co-ordinated measures to maintain migratory waterbird species in a favourable conservation status or to restore them to such a status and of taking into account the precautionary principle in implementing the measures.

The measures concern protection of endangered waterbird species, restriction of any use to a sustainable level, encouragement of protection of sites and habitats for those species, insurance or re-establishment of networks of suitable habitats, investigation of problems due to human activities, co-operation in emergency actions and developing appropriate emergency procedures, prohibition of introduction of non-native species, support and training concerning biology, ecology and monitoring, stimulation of awareness and understanding of conservation issues, exchange of information and results and co-operation in assisting each other to implement the agreement.

The Action Plan formulates three levels of protection:

- level A should provide strict protection for 32 species, partly only for regional populations;

- level B should provide such protection that a favourable conservation status is maintained or restored for 30 species, partly only for regional populations;

- level C concerns populations that do not need special attention but could benefit of the international co-operation.

Concerning research and monitoring the plan stipulates that parties should endeavour:

- the survey of poorly known areas which hold important concentrations of populations of the species listed in table 1 of the Action Plan; - the monitoring of those species; - the co-operation concerning those species to improve the measurement of population trends, the determination of the migration routes and season distribution, the research on ecology and population dynamics, the study of effects of wetland loss or degradation, and the study of the impact of hunting and trade; - the co-operation with relevant international organisations and to support research and monitoring projects.

It is evident that the geographical range of WIWO-activities and goals overlap considerably with the scope of the AEWA and WIWO-projects will continue to support it in the years to come.



In realising its scientific and conservation aims, WIWO follows a distinctly initiative-supporting or bottom-up approach. The basis of the organisation is formed by a large group of volunteers, consisting in part of scientifically trained biologists but also including many dedicated amateurs, initiating, organising, carrying out and reporting on particular research projects. The role of the foundation itself (i.e. the board) is limited to stimulating initiatives, providing guidance and practical assistance in the logistics of project organisation, maintaining quality standards, and the application for the necessary funds.

WIWO projects can have a wide range of forms, ranging from desk studies to large-scale field expeditions. The most common project form is that of field studies lasting between one and three months, carried out by volunteers. Such projects are adopted by WIWO after evaluation by its board of a written project proposal. In most cases, contacts between WIWO and project initiators already exist during the preparation of the proposal. For each project a WIWO board member is appointed as a mentor, maintaining the contacts between the board and the group carrying out the project.

When evaluating project proposals, criteria are used such as the scientific and/or conservation necessity of the work, its volunteer character, the feasibility of the project, and whether or not the subject is already being addressed by other organisations. Most projects therefore concern wetland areas of which little knowledge is available and which are situated in countries with few local ornithologists.

The volunteers generally invest in the organisation and execution of projects large amounts of time and, in many cases, also money. Although most projects are initiated from the ranks of WIWO, in cases they may be instigated by, or executed in co-operation with other organisations such as Wetlands International (the former International Waterfowl and Wetlands Research Bureau, IWRB), BirdLife International (the former International Council for Bird Preservation), or organisations in the countries involved.

WIWO has a small financial income from report sales, these hardly covering the organisation costs, and has always been completely dependent on external financing for the realisation of projects. The voluntary character of WIWO's activities should not be viewed as a source of cheap labour for projects that could reasonably involve the payment of salary.

In the organisation of field projects, a formal procedure is followed to obtain permission for research from the authorities. Counterparts from institutions and universities in the countries visited are invited to participate in the projects and local Embassies and the Dutch Ministry of Foreign Affairs is informed of our presence and activities.



In broad terms, two basic lines can be identified in the studies conducted by WIWO. The first line is census work and population identification, the second is research into migration strategies and feeding ecology of waterbirds in relation to the resources available in wetlands.

2.4.1 Census work

Census work basically consists of counting waterbirds in wetland areas. For population identification, ringing of birds is in many cases an important source of information, either by producing recoveries linking breeding and wintering areas or different stopover sites used en route, or by producing biometric data which can sometimes be used to identify birds from different breeding populations (e.g., Wymenga et al. 1990).

This type of work is important in two ways. Firstly, it is the primary tool to identify which wetlands is of the highest importance for (certain populations of) waterbirds, and therefore need protection. Secondly, counts in as many areas as possible are necessary to obtain estimations of the total size of waterbird populations. Estimates of population size are important since they identify species or populations that are vulnerable because of a limited population, or decreasing in number, and because they form the basis of the so-called '1%-threshold' that are used to evaluate the importance of areas, e.g. for inclusion in international conservation agreements such as the Ramsar Convention or for population conservation and management under the future African/Eurasian Waterbird Agreement and the Bonn Convention. Thus, the relevance of census work for conservation and management is large and direct. In addition, knowledge of numbers and distribution of waterbirds forms the basis of all further work on their biology.

2.4.2 Migration ecology

Research into migration strategies and feeding ecology, although less directly, ultimately is as important to conservation as census work. For migratory birds, critical conditions for population survival may occur at any time during their annual cycle, and anywhere in their migratory routes. Conditions in one area or stage of the annual cycle may profoundly influence what happens elsewhere. For a comprehensive conservation policy, therefore, knowledge is required of the factors limiting a bird's performance during all stages of the annual cycle.

The subjects of this type of studies as conducted during WIWO projects can be tentatively classified into closely related topics:

- describing strategies used during the migratory journey. The number of steps in which the distance between wintering and breeding sites is covered is the central issue and determines the number of necessary stopover sites, fuel loads and the speed with which they must be stored;

- quantifying the resource base on which the success of these strategies depends. Feeding conditions are obviously a very important aspect in this respect, but e.g., wind patterns facilitating migratory flights can also be regarded as a resource;

- identifying physiological constraints, which determine whether or not the resource base can be fully exploited by the birds.

In comparison to census work, this second type of studies generally involves a greater input of technical equipment, research experience, time, and in many cases also laboratory facilities for processing and analysing materials collected during the field work. Often therefore, this type of work is carried out in co-operation with professional biologists and by using facilities of universities or scientific institutions. It is important for WIWO to maintain close contacts with these institutions in order to secure that studies into migration strategies and feeding ecology will be part of future field activities.



WIWO aims to collect scientific information. The practical and political conservation and management actions to be undertaken on the information collected or interpreted by WIWO are considered to be better dealt with by local or international conservation bodies and to governments. However, through the research activities, much relevant information on the importance, threats and conservation needs of wetland areas become available. In the past, this information has led to practical conservation measures in several cases.

One of the ways in which WIWO tries to communicate information critical for conservation is through contacts and co-operation with authorities, universities and private organisations in the countries visited, and the invitation of local counterparts to participate in the projects. No special training is normally provided for counterparts, but by participating in all research activities, they become familiar with research techniques and problems, as well as the areas studied and the threats they face.

It is hoped that the participation of counterparts will promote field studies and conservation awareness in the countries visited in the period after the expeditions have left. However, WIWO lacks the resources and the manpower to maintain long-term support for local activities after each project and has not been set up for that reason. In practice, the follow-up depends largely on the personal interest and involvement of project participants. Notwithstanding this fragile basis, WIWO activities, especially when repeated for several years in the same country as in the case of Turkey, have in several cases produced lasting local research and conservation spin-offs.




During the past 18 years in total 63 WIWO - and co-operative expeditions were carried out (figure 1.1, table 1). These were the projects for which funds were obtained. Besides these, two desk-studies and some contract work was carried out and several reports were published reporting counting or other results of wetland studies during 'vacations' or other types of free time. All sites visited during the WIWO expeditions, co-operation projects or during 'vacation' studies are indicated in figure 1.1. The distribution pattern of the projects indicates the flyways and regions where most work was performed. Concentration of activities was initially along the East Atlantic flyway (figure 1.2) in particular coastal West Africa, but later spread out to the Mediterranean, Black Sea and Middel East and recently to the breeding grounds in the Arctic. The projects included a wide range of different research subjects ranging from base-line studies (e.g. counting) to studies of feeding ecology in relation to migration behaviour. At the end of this chapter seven typical examples of WIWO-studies are highlighted to give an impression of the scientific 'yield' of 18 years of research. Lists of publications (c.140) and reports (63) are given in Annex 1 and 2, respectively. Lists of captured birds (c. 50,000) are given in Annex 3-5.



Generally, funding is only requested for part of the project costs, i.e. the logistics of the fieldwork and the printing of a short and a final report. Part (or sometimes all) of the subsistence costs are covered by the participants. The preliminary activities prior to the fieldwork and the output part, the time consuming analysis of data and the preparation of papers and the report, are never parts of the project budget for which funds are requested.

Table 2 and figure 2 show the development of funds received between 1980 and 1998. Included are a list of the countries where projects were carried out, and the distribution of the yearly costs over the four main regions of activity (West Africa, Mediterranean, Black Sea, and Arctic). Table 2 also includes a list of all participating funding agencies over the years. In this brief evaluation of the years 1980-1998 the following conclusions can be drawn:

* The supportive financial commitment from the Ministry of Agriculture, Nature Management, and Fisheries, International Affairs, proved to be of great help to support, continue and expand WIWO activities.

* The spectrum of funding organisations is still expanding. This is believed to reflect the growing appreciation of the work done. Hence, the procedures to apply for funding shortened with some organisations. However, they remain laborious with others.

* Despite the increased number of funding organisations, the amount of funding acquired averaged around DFL 150,000 per year in the period 1980-1993 to DFL 100,000 in the period 1994-1998.

* Projects in Africa are complicated, relatively expensive and need years of preparation. Despite these obstacles, the interest to organise projects fortunately remains, as the key information to many problems is situated in this continent.

* The interest of organisations other than the Ministry of Agriculture, Nature Management and Fisheries in funding studies in the Arctic has so far been limited.



In Forward Plan 1989-1993 many research priorities for the different regions were identified. In this period a lot of these aims were fulfilled, others remain to be tackled. The following section briefly reviews the identified priorities and indicates the progress.

* Completing the series of counts of waders wintering along the shores of the Gulf of Guinea, W-Africa. Complete surveys were made of the coastal wetlands in Guinea-Conakry (1988, 1990) and Gabon (1992). During the project in Guinea-Bissau in 1992/93, a repeat count was made by Danish ornithologists of the Archipelago dos Bijagos.

* Repeat count of wintering waterbirds on the Banc d'Arguin, Mauritania. This challenge has not yet been taken up.

* Surveys of wintering waders in inland wetlands in western Africa. Little progress was made in this field. Counting dispersed waterbirds in areas of low surveyability is laborious and time-consuming (and therefore expensive), while there is still a lack of a good methodology. Some work on this topic was done during a WIWO project in Cameroon (1993). Perennou (1991) reviewed existing data on waterbird distribution in coastal and inland tropical Africa.

* Surveys of inland wetlands in east Africa and coastal wetlands along the Red Sea. Because of the political and logistic difficulties in many countries in this region, organising expeditions is still very difficult. Recently, local ornithologists have started regular counts of wetlands in Kenya.

* Migration strategies of waders in West Africa. Autumn migration and moult of waders on the Banc d'Arguin and preparation for spring migration from Guinea-Bissau were studied during trapping studies in 1988 and 1993.

* Migration patterns, migration strategies, and resources for migration in the eastern Mediterranean / Black Sea region. A large proportion of WIWO activities during the past five years has focused on spring migration in the eastern Mediterranean region. Projects were carried out in Tunisia (1990), Greece (1990), Turkey (1989, 1990), Egypt (1990), Israel (1989), and the northern coasts of the Black Sea and Azov Sea in the Ukraine (1990, 1992, and 1993). The projects in 1990 were part of the Eastern Mediterranean Wader Project 1990, in which also a network of local observers in a number of Mediterranean countries were involved. Our knowledge of distribution, phenology, and body mass development as well as turnover of waders during spring migration has considerably increased.

Relatively little work has as yet been done on the food resources for staging waders, with the exception of the 1992-1993 projects in the Sea of Azov.

* Breeding distribution, densities, breeding success and population characterisation of arctic waders. Somewhat contrary to expectation at the time of writing of the Forward Plan 1989-93, distinct progress has been made in this field. As a result of 'perestroika', WIWO was able to participate in international expeditions to the Taimyr Peninsula in Siberia in 1991, 1992, and 1993. In two different areas breeding waders were censused, birds were ringed, and nesting success was measured in relation to the abundance of lemmings and predators (see 3.4.7).

* Identifying arctic staging areas of migrating waterbirds. No particular project was organised to address this topic. Some observations of staging waders were made during the Taimyr expeditions, but these revealed no marked concentrations of birds in specific areas. During the past years, other groups have quantified numbers of waders stopping over in Iceland during spring migration, and made a first reconnaissance trip to the White Sea. In addition to the projects envisaged in the Forward Plan 1989-93 and reviewed above, a number of other initiatives were developed in the past five years. These concern projects in Turkey, Botswana, Libya, Albania, and Iceland (see table 1 for details).



In the Forward Plan 1994-1998 about 30 research priorities were listed. In this period 24 field projects were carried out which addressed to more than one-third (12) of the listed priorities. Only one project did not address one of the priorities (591). This implies that the Forward Plan 1994-1998 performed very well to provide 'directions' rather than 'roads' for new initiatives. In the period 1994 -1998 21 WIWO-reports (45-67) were published, in addition to papers in scientific and popular journals, and reviews of WIWO-reports in ornithological journals (Appendix 2).

The results of many WIWO-projects (1980-1998) have contributed significantly to the identification of Important Bird Areas (IBA) in Europe (Grimmett & Jones 1989), Middle East (Evans 1994) and Africa (BirdLife International, in preparation). Waterbird counts have also contributed to the African and Asian Waterbird Census (e.g. Dodman et al. 1997) and updated population estimates of waterbirds (e.g. Rose & Scott 1997). Several projects collected data on Globally Threatened Species as a major aim: Slender-billed Curlew (29, 54, Boere & Yurlov 1998), Pygmy Cormorant (60), or indirectly: Sociable Lapwing (Boere & Yurlov 1998), Audouin's Gull (63, 47); Lesser White-fronted Goose in Turkey (mainly negative counts, 18, 21, 31, 37); White-headed Duck, Marbled Teal, Ferruginous Duck: most Mediterranean projects.

Two WIWO-initiatives were awarded with ICBP/BirdLife International Expedition Competition Award: a breeding bird survey (including Dalmatian Pelican) in the Kizilirmak Delta, Turkey, in 1991 (45) and the 'rediscovery' of the Lesser Crested Tern colony in Libya (46).

3.4.1 The arctic

The following WIWO-projects were carried out in the Arctic and are listed with year, locality, authors and number of WIWO-report: 1991 Northeast Taimyr, Prochishcheva Lake, Schekkerman & van Roomen (55); 1992 Northeast Taimyr, Prochishcheva Lake, Spiekman & Groen (33); 1993 Taimyr, Medusa Bay van Dijk & Venema (in prep.); 1993 Iceland, Gerritsen & Groen (51); 1994 Taimyr, Piassina Delta, Vonk & Duiven (in prep.); 1994 Northern Taimyr, Cape Sterlegova, Tulp et al. 1998 (61); 1995 No project in the Arctic; 1996 West Taimyr, Medusa Bay, Tulp et al. 1997 (57); 1996 Northern Norway, Finnmark, van der Have et al. (in prep.); 1997 West Taimyr, Medusa Bay, Komenko et al. (66, in prep.); 1998 West Taimyr, Medusa Bay, van Turnhout et al. (in prep).

* Censuses of densities and habitat choice of breeding birds in a (large) number of sites throughout the arctic. The geographical spread in the arctic has increased when studies were started in Northern Norway in co-operation with NINA-NIKU, Tromsø, Norway (33, 51, 55, 57, 61, 66, van Dijk & Venema (in prep), van der Have et al. (in prep), van Turnhout et al. (in prep).

* Describing food resources of arctic breeding birds in relation to energy requirements and distribution. The relationship between temperature, invertebrate availability and chick growth rate was studied in 1991 (Taimyr, Prochishcheva Lake) by Schekkerman & van Roomen (55).

* Quantification of factors influencing breeding success, especially food availability and weather. Two studies in particular adressed this topic in N. and W. Taimyr (57, 61). Species with suitable densities were Little Stint and Curlew Sandpiper.

* Studies on mating systems, parental care, site tenacity and philopatry, in different parts of a species' breeding range. Working in the same area for two or more years is crucial to part of these questions. The studies carried out at Medusa Bay 1994-1998 are particular suitable to address this topic, especially after the Willem Barentz Station was build in 1995.

3.4.2 Black Sea and Sea of Azov

* Waterbird surveys of coastal and inland freshwater marshes along the Sea of Azov. A pilot study was carried out in August 1996 in the Sivash, exploring the numbers and distribution of marsh terns. A complete waterbird count of the Sivash was carried out in August 1998, and habitat availability, food resources, biometrics and moult of terns and waders was studied.

3.4.3 Mediterranean

* Monitoring of waterbird numbers, habitat availability and food resources at a selected number of sites: in February 1994 Kneiss, Gulf of Gabès, Tunisia was surveyed to compare with the count of February 1984 (54).

* Methodological studies of monitoring wintering waders and waterfowl; in February-March 31 wetlands were surveyed in search for Slender-billed Curlew; a monitoring method was proposed (at least two days per wetland, including checking evening roosts, 54).

* More studies on the migration ecology of waterbirds, especially of vulnerable species using localised and/or unpredictable food resources. Foraging ecology of Broad-billed Sandpipers was studied in spring 1993 in Sivash, Ukraine (Verkuil et al. 1993).

* More surveys of inland wetlands (e.g. Algeria, Morocco, Tunisia, Turkey). 31 inland and coastal wetlands were surveyed in Tunisia in February-March 1994 (54).

* Increasing the geographical scope of basic waterbird surveys in all seasons for infrequently-visited countries (e.g. Albania, Libya); A wetland survey of Albania was carried out in 1996.

* Desk studies to analyse and combine the results of the various spring projects. Several papers have summarised the results of the Mediterranean projects in spring (van der Have & van den Berk 1996, van der Have, in prep, de Goeij et al. 1994).

* Feeding studies of fish-eating birds (grebes, cormorants). Although not in this region, an ecological study of Pygmy Cormorants was carried out in Lake Prespa, Greece, in 1996 (60).

3.4.4 Caspian and Arabian

* Feeding ecology studies of wintering waders, especially Broad-billed Sandpipers in UAE and Oman. A waterbird survey of the UAE (in co-operation with NARC) was carried out in autumn 1994 and spring 1995. The feeding ecology of Broad-billed Sandpipers was studied in Khor Dubai in spring 1995 (62).

3.4.5 Inland wetlands in the Sahel

* Ground surveys in the Senegal Basin, Inner Niger Delta, and Lake Chad Basin (Chad, Mali, Niger, and Senegal). A survey of the Waza-Logone floodplains and the Lake Chad Basin in Chad was planned for 1997, but was postponed to 1998.

* Desk study on relation between rainfall patterns and shifts in distribution of waterbirds. Although not in this region, three January surveys (1993-1995) of the Boteti River, Botswana, were very illustrative as the Boteti River progressively dried out during these years with dramatic effects on waterbirds and European Swallows (56).

3.4.6 Coastal West Africa

* Regular counts of core sites, in particular during winter. The long awaited, complete winter count of Banc d'Arguin, Mauritania, was carried out in January-February 1997 (Zwarts et al. 1998). The Sine-Saloum Delta in Senegal was counted in January 1997 (63).

3.4.7 East Africa

* Censuses of wintering waders along the Red Sea and Indian Ocean coasts. A winter count of Pemba and Zanzibar Island, Tanzania, was carried out in February 1998 (Geene et al. in prep.).

3.4.8 Southern Africa

* A waterbird census of the Okavango Delta. A (very) limited survey of the Okavango Delta, Botswana, was carried out in January 1995 (56).

* Inventory of wintering waders and terns on coast between Gabon and South Africa. Although not a WIWO-project, but to be published as WIWO-report, was a survey of the West Coast of South Africa (Underhill et al. in prep.).



3.5.1. Between Taimyr and Table Mountain: cross-continental links by weather and lemmings

Seven WIWO-expeditions studied breeding waders in Taimyr since 1991 (see 33, 55, 57, 61). All these studies have gathered information on wader breeding success. The large variation in breeding productivity of arctic geese and waders has since long caught the attention of ornithologists along the East-Atlantic Flyway, and was a major incentive for taking up studies in the Arctic. By measuring hatching success of wader clutches in a number of years, the expeditions have contributed direct field data relating to the hypothetical relationship between cyclic variations in the abundance of lemmings, and shifts in the numbers and prey choice of predators (mainly Arctic Foxes Alopex lagopus and skuas Stercorarius spp.) that affect the survival of birds' clutches and chicks. In general, results have supported the hypothesis (e.g. Underhill et al. 1993), although there seems to be significant local variation in the dynamics of lemming populations, and hence predation pressures. Although hatching success is one important aspect of reproduction, the survival of chicks may be as important. Two expeditions have addressed the chick-rearing period. Wader chicks forage for themselves and are not fed by the parents. In the cold arctic climate, this exposed and active lifestyle leads to very high-energy expenditure, as was shown by measurements on Knot chicks. During frequent spells of cold weather, finding the necessary amount of food becomes difficult due to the facts that small chicks cannot maintain their body temperature and need more brooding, and that the availability of surface-active insects decreases under cold conditions.

Effects of weather conditions on the growth rate of chicks were demonstrated in Curlew Sandpipers (55) and Knots (61). That these effects may translate into chick survival and breeding productivity was shown in Curlew Sandpipers. The proportion of juveniles in the population wintering in South Africa, measured over 18 years, was found to be correlated (after accounting for lemming abundance effects) with weather conditions on Taimyr in the second decade of July, the period in which most small chicks are present in the tundra (Schekkerman et al. in press; figure 3.2).

3.5.2. The Broad-billed Sandpiper: world's smallest Nereis-specialist?

The work on this intriguing species started in 1990 during simultaneous surveys in the Mediterranean. Detailed studies of the stopover ecology were carried out in spring 1992 and 1993 (Van der Winden et al. 1993, Verkuil et al. 1993). Attention shifted closer to the wintering grounds in autumn 1994 and spring 1995 in the United Arab Emirates (Keijl & Ruiters 1998). A small wintering population was discovered in Tunisia in February 1994 (van der Have et al. 1997 [54]). A recapture of a female, ringed in May 1993 in the Sivash, on the nest in Northern Norway in 1995, motivated breeding biology studies in 1996 and 1997, started earlier by others in Northern Norway (Rae et al. 1998).

The Broad-billed Sandpiper appears to be highly selective with characters unique among the smaller waders. During the non-breeding season Broad-billed Sandpipers specialise on large worms usually taken from soft mud. Strong head muscles and a broad bill are thought to be an adaptation to large prey items compared to its size (slightly larger than Little Stint). Marine wetlands are preferred in winter (e.g. Arabian Gulf) and brackish wetlands, such as the Sivash, Ukraine, are selected in spring.

The flyway is also rather unusual being almost northwest - southeast oriented. After short stopovers in the Baltic, the breeding starts late and almost exclusively in mountain bogs and aapa mires (peatlands with palsa's, i.e. peat hill with ice core). At hatching females desert the broods, the males leave perhaps well before fledging one or two weeks later. The moulting period is the least known part of the life cycle. A complete moult takes place probably exclusively in the Arabian Gulf. During stopovers of adult birds in July-August in the Sivash, Ukraine, no birds in primary moult have been observed. Birds finishing primary moult were caught in Khor Dubai, United Arab Emirates, in October 1986 (Uttley et al. 1988).

The Broad-billed Sandpiper is vulnerable due to its small population size (< 25,000 birds, Fennoscandian subspecies), highly selective habits and localised occurrence during its whole life cycle. To date, no population trends are available, but the increased cover of the wintering areas did not reveal numbers exceeding the breeding population estimate yet. Breeding habitat loss due to drainage of aapa mires and bogs in Northern Scandinavia in relation to reindeer husbandry is a matter of concern.

3.5.3 European Swallow in Botswana: survival of the fittest

Twice yearly European Swallows cross the Sahara to and from the non-breeding areas in trans-Saharan Africa. During the non-breeding season European Swallows are highly dependent on wetlands: they roost almost exclusively in reedbeds throughout Africa. They undergo a complete moult starting in September - October in West Africa and in November in Southern Africa.

The rate of moult of European Swallows in Botswana was studied during December - January of 1992/93, 1993/94 and 1994/95 to investigate the effects of variability in rainfall and roosting habitat availability (van den Brink et al. 1998 [56]). In January 1994, 2-3 million European Swallows were counted at a traditional roost along the Boteti River. The rate of moult was relatively slow; about one feather (primary, secondary or tail feather) was replaced every two weeks in both adults and juveniles. Moult started more than a month later than in Ghana, West Africa (van den Brink et al. 1998 [58])

The speed of moult in juveniles was generally lower than in adults, in particular of secondaries and tail feathers. Moulting rate of both primaries and tail feathers was lowest in 1994/95 during a period of drought and coincided with the almost complete destruction of roosting habitat. Body mass dropped dramatically in particular in juveniles of which many perished during the drought (Fig. 3.4). In 1992/93, moulting rate was highest when rainfall was moderate and roost habitat abundant. Moulting rate was intermediate in 1993/94 when rainfall was frequent but roosting habitat reduced because of the low water level in the Boteti River. Indirect evidence suggested that during these droughts large numbers of mainly juvenile swallows perished.

The combined effect of reduced food availability during droughts and higher densities and longer foraging flights when roosting habitat is scarce might explain the annual variation in moulting rate. From the second week of January onwards many adults started moulting the outermost tail feather before the penultimate feathers. This phenomenon could indicate the importance of long tail streamers in aerial manoeuvring when foraging during the return migration to the breeding grounds. Staying alive in Africa and preparing for the return migration to arrive in time clearly selects for swallows best adapted to these harsh and varying conditions, also known as 'survival of the fittest'.

3.5.4. Dunlins in Kneiss, Tunisia: moon cycle effects

Two previous WIWO-projects had identified the intertidal areas of Kneiss in the Gulf of Gabès, Tunisia, as a wetland of considerable international importance in winter (van Dijk et al. 1986 [11]) and spring (Spiekman et al. 1993 [38]). For example, it harbours the largest concentration of wintering Dunlins Calidris alpina in the Mediterranean. The last complete waterbird count of Kneiss in winter was in 1984 and an update was long overdue. Therefore, a project was initiated by the WIWO Foundation and organised in 1994 in close co-operation with Nicola Baccetti (INFS, Italy), the Tunisian Ministry of Agriculture and the Tunisian bird protection society 'Les Amis des Oiseaux'. This project was also motivated by the common interest of researchers studying the Mediterranean Flyway population of the Dunlin, which has stopover sites in autumn in Poland and Italy and in spring in Ukraine (Gromadzka 1989).

A discriminant analysis (with POSCON) of size variation of Dunlins (710 caught with mistnets) showed that 42% could be identified as the larger Siberian subspecies Calidris alpina centralis, and 58% as the nominate subspecies. 'Adult buff' wing coverts, characteristic for Dunlins of eastern origin, were found in 37 birds (5%). Two ring recoveries (one recaptured as breeding female in Sweden and one ringed as pullus in Finland) proved that birds from the Baltic schinzii population do winter in Kneiss. Although the proportion of Baltic schinzii in Kneiss may be too small (<5%) to identify with certainty with biometrical analysis, it may represent a significant proportion of the small Baltic population of this small-sized subspecies. The sex ratio, estimated with POSCON discriminant analysis (Engelmoer & Roselaar 1998), was significantly male-biased (60.4% males) which was similar in a sample of 24 dissected casualties (67% males). This may suggest that males do winter more southerly than females.

The Dunlins captured in Kneiss were generally low in weight and slightly leaner than expected from the relationship between mid-winter temperature and body mass found in Dunlins wintering in Britain. The amount of visible fat was also small. A principal component analysis showed that females (sexed with POSCON) were 1-2 grams heavier than males even after correction for differences in structural size. This may have been caused by a slightly higher non-fat body mass in females.

Body mass in males, but not in females, varied significantly during the study period and was lowest around new moon (Fig. 3.5A). This was confirmed in a multiple regression analysis including variation in structural size, time of catching relative to the time of high water and the number of days before or after new moon (Fig. 3.5B). Several not mutually exclusive explanations could account for this effect. The short-billed males might be more constrained by light conditions, if foraging depends more on visual clues, than the longer-billed females, which may forage more with tactile clues.

3.5.5. Trouble in paradise? Banc d'Arguin, Mauritania, revisited

The Banc d'Arguin is one of the most remote and unspoilt coastal wetlands in the world and seems to have remained unchanged over recent decades. Around two million waders, over a quarter of all waders found wintering along the entire European and West-African coast, are concentrated here. This natural wetland of shallow seas, low islands and vast tidal flats is host to many other waterbirds too. This impressive number amounts to twice the total number of all wintering coastal waders found in the Danish-German-Dutch Wadden Sea or in all combined British estuaries. Despite its huge importance, few attempts have been made to completely count the waders on the Banc d'Arguin.

The counts carried out during the winters of 1978-79 (Trotignon et al. 1980) and 1979-80 (Altenburg et al. 1982 [1], Engelmoer et al. 1984) were the first (nearly) complete ones. Apart from a count in the summer of 1988 (Van Dijk et al. 1990), it took more than ten years before a new winter count occurred (Gowthorpe et al. 1996). The two counts in 1979 and 1980 respectively arrived at about the same number of over two million waders and there was no reason to doubt that the wintering number would have changed since then. Instead of confirming the status quo, their results were alarming. They pointed to the fact that, during the intervening years, there must have been a significant decline in the number of wintering waders and other waterfowl, as well as in the breeding population of Long-tailed Cormorant and six different tern species.

They also suggested a possible common explanation for these negative trends: over-exploitation of fish causing a disturbance of the entire ecosystem. The Banc d'Arguin is the shallowest part of an extensive upwelling zone that appears along the Mauritanian coast (e.g. Wolff et al. 1993). This attracts, further offshore, an international fleet of many fishing boats, using huge fine-mesh nets. Local fishermen on the Banc d'Arguin who use small sailing boats and standing nets to catch fish, mainly mullet, complain that their catch has decreased dramatically over recent years. Indeed, it is conceivable that the decline in fish numbers on the Banc d'Arguin is a direct consequence of the increase in the industrial fisheries further offshore.

The new count of the waders and waterbirds on the Banc d'Arguin (Mauritania) in 1997 confirms the decline in numbers suggested by recent winter counts (Gowthorpe et al. 1996), although not on the same scale and not in all species (Zwarts et al. 1998). A total of two million waders was similar to the total numbers counted in 1979 and 1980. However, more Dunlins Calidris alpina and Curlew Sandpipers Calidris ferruginea were counted, and fewer Knot Calidris canutus and Bar-tailed Godwits Limosa lapponica than nearly two decades ago. The same trend was found in coastal waders in Guinea-Bissau, but Redshank Tringa totanus and Greenshank Tringa nebularia showed an increase on the Banc d'Arguin, possibly because there are more shrimp and young fish now due to the over-exploitation of fish further offshore, and a decrease in Bissau. In Oystercatchers the reverse was true: they decreased on the Banc d'Arguin, due to the disappearance of their main prey: the Bloody Giant Cockle, and increased in Bissau. The most notable declines were in Kentish Plover Charadrius alexandrinus, Turnstone Arenaria interpres, Little Stint Calidris minuta and Knot on the Banc d'Arguin as well in Bissau.

It is recommended (1) to monitor the wader numbers on the Banc d'Arguin on a more regular basis, (2) to subdivide the count into the sections as used in 1980 and 1997 and (3) to accept, at least for the moment, that the only way to monitor waders numbers on the Banc d'Arguin is by performing a complete count of the entire area.




Migrant waders and waterfowl are characterised by their itinerant life-cycle, which means that they are constantly on the move between breeding areas, stopover and moulting sites and wintering haunts figure 4.1). All factors which influence survival of waders and waterfowl throughout their annual life-cycle and their reproductive success during the breeding season in particular will eventually, across seasons, determine population size (e.g. van der Have 1991). Therefore, conservation and management can only be effective if this fundamental knowledge of population size and life cycles of waders and waterfowl is available. As population size and population stability usually determine conservation and management priorities, this implies that for every species the key sites have to be mapped and monitored, and the key factors in population regulation throughout the annual life cycle have to be determined. In this framework, all information from all seasons is relevant and their value can only be weighted afterwards. To date, this approach has pervaded virtually all WIWO-projects.

Throughout their annual life-cycle, migrant waterbirds have to make choices such as whether to choose one mate or another, whether to breed early or late, whether to moult or to suspend moult, whether to forage or to look for predators, whether to leave or to stay and so on. The study of these choices and their relation to survival and reproduction, whether carried out on the wet Taimyr tundra or in a shrinking lake in Africa, forms the heart of the annual life cycle approach.

4.2.1 Waterbirds

Table 3 summarises very roughly the extent of our knowledge concerning some basic constituents of the annual itinerary of different waterbird groups in the Palaearctic-African migration system. Flamingos, Cormorants, Pelicans and Geese are relatively well-known groups, in contrast to Divers, Grebes and inland-wintering waders (see also table 4). It is clear that further studies on these groups are needed and initiatives are welcomed.

4.2.2 Waders

Waders have attracted special attention from the very first WIWO-project and this focus has remained to date. As they are relatively easy to study, have relatively specialised feeding habits, usually frequent remote areas and display an intriguing variety of migration strategies it is not difficult to see why the fascination for this waterbird group has remained throughout the years. In fact, wader enthusiasm proved to be contagious and inspired new generations of 'wader-workers'.

In table 4 the current state of knowledge with respect to various aspects of the annual cycle is summarised. The 40 wader species of the East Atlantic and Mediterranean Flyway are sorted according to their non-breeding habitat and ranked according to the amount of knowledge available. The topics covered in the table columns are derived from figure 4.1. For five phases in the annual cycle, a number of important aspects are identified and evaluated. These refer to the geographical (where are the main sites, what migration routes are taken), phenological (what is the timing of different activities within the annual cycle) aspects of each phase, and to the resources upon which the birds subsist (food stocks), as well as the constraints involved (climate, risk of starvation, predation). Four basic phases are distinguished (breeding, autumn migration, wintering and spring migration, plus the annual moult of flight feathers which is fitted into different phases in different species. A column summarising our knowledge of the total population size of each species precedes these phases.


It is clear from the table that available information is heavily biased in favour of the coastal waders migrating along the East Atlantic Flyway. The majority of the waders occurring inland and dispersing over many smaller wetlands in Africa are still relatively unknown. The breeding biology of part of these species is however relatively well known, as they tend to breed at lower latitudes than the - often arctic-breeding - coastal species, and are thus more easily accessible during this phase (Piersma 1986, Thompson & Thompson 1991).

The lack of knowledge of the non-breeding season in these species is of course explainable by the difficulties of studying them in small, structurally complex areas where they occur in small numbers only. It remains to be seen if their scattered distribution is a result of their adaptability in habitat choice and feeding ecology, before it can be taken as an indication that these birds are perhaps less sensitive to local habitat losses than relatives which occur more concentrated.

During the breeding phase, the least-known aspect is that of resources and constraints. Food choice and food availability have been studied in relatively few species during this time of the year (e.g. Holmes 1966), as are the constraints that the arctic climate may impose on the birds' reproductive effort (Piersma & Morrison in press).


With the exception of inland waders, the wintering phase is perhaps the best-known phase of the annual life cycle of many waders. Distribution and numbers are relatively well known, while studies on food availability and feeding ecology have been carried out in a variety of areas ranging from Northwestern Europe to South Africa (Kaletja 1992). A gap still exists in the most tropical parts of coastal Africa, although it is now partly being filled by studies in Guinea-Bissau, Guinée Conakry and Gabon (15,23,25,26,27,35,41). A lot of work has also been done on the implications of climatic differences between wintering areas at different latitudes (Piersma, Drent & Wiersma 1992). Relatively little is known yet of predation pressure in different wintering areas, though this might be a factor of importance in shaping migration strategies.


The spring migration from wintering grounds to breeding grounds has been studied in comparatively great detail. For several species of coastal waders, e.g. Knot, Bar-tailed Godwit and Whimbrel, it is now well known what are the main staging areas and which distances are covered in single flights (25, Piersma & Davidson, 1992), while considerable progress has been made in describing the feeding conditions and constraints allowing the birds to make these flights. Even for these species a final uncertainty remains in that high-arctic breeding birds may use one extra stopover between the Waddensea and the breeding grounds. This area - quite possibly the eastern White Sea - needs to be identified and the feeding conditions there need to be studied in order to obtain a complete picture of the spring migration. This is made extra important by the possibility that conditions in this last stopover site influence breeding success through the size of energy stores upon arrival on the breeding grounds. Just how important this possible mechanism is for waders still needs to be established.


Compared to spring migration, the phase of autumn migration is a poorly explored part of the annual life cycle. Although the general timing and routes are relatively clear, only in few species do we know which staging areas are used successively and what distances are covered by single flights. Although it is possible that autumn migration is not as critical in the annual cycle as spring migration because the pressure to reach the breeding grounds in time is absent, this might not be true since the resources for autumn migrants generally decrease with time, and because autumn migration is often closely linked with moult. Furthermore, birds may have to compete for wintering resources as well It is certainly untrue for juveniles which lack experience, and which have to perform their first autumn migration while the location of suitable stopover sites is probably unknown on detailed scale, and in the meanwhile have to compete for resources with more experienced older birds (Van der Have et al. 1981).


There are several indications that the flight feather moult is a sensitive phase in the annual cycle of waterbirds. It interferes with migration directly because flight is impaired. The fact that moult rarely overlaps more than partly in time with pre-migratory fattening, suggests that it is also a costly process in terms of energy expenditure. Thirdly, many waders moult in large concentrations in very specific areas before spreading out to winter in a larger number of sites. Many waterfowl species even undertake long 'moult migrations', the direction of which may be quite different from that of the movement between breeding and wintering areas. This indicates that moult poses special demands on the areas where the birds stay, either with respect to food or safety and disturbance (Smit & Piersma 1989). Identification of the most important moulting sites and the resources that make them suitable therefore deserves attention.



4.3.1. The Arctic

A review of WIWO-activities WIWO has been active in arctic regions since 1991, when a first team of two persons joined the arctic expeditions to the Taimyr Peninsula, organised by the (then) Institute of Evolutionary Morphology and Animal Ecology of the Russian Academy of Sciences. Since then, six WIWO-expeditions have worked at four different sites on Taimyr in five years (see 33, 55, 57, 61). In addition, teams have studied the breeding biology of Broad-billed Sandpipers Limicola falcinellus in northern Norway in the summers of 1996, 1997 and 1998, in co-operation with Norwegian ornithologists.

The two basic types of research can both be recognised in the activities carried out in these expeditions. Besides area-directed census work, most expeditions have addressed aspects of the breeding ecology of arctic waders. In general, the ecological studies aim to identify critical periods in the breeding cycle and the factors that are most important in influencing their outcome, based on the idea that knowledge of the critical factors in a species' population dynamics is important for understanding its conservation needs. The account given below summarises the work that has been done since 1991 and highlights research opportunities that have appeared from this work. At the same time, issues mentioned in the previous forward plan (e.g. densities and habitat choice, mating systems, early stages of autumn migration) remain relevant.

In the first years, organising expeditions to the Russian Arctic was relatively simple due to the logistic network provided by the IEMAE. Later on, it has become more difficult, as this network has fallen away and transportation infrastructure has decreased in extent and quality while prices have increased enormously. The costs of expeditions to the Russian Arctic now approach those involved in getting to Greenland or northern Canada. On the other hand, the development of a few permanent research stations such as the Willem Barentz Station on Taimyr, has provided useful facilities for future work, especially for studies spanning several years.

Census work All past expeditions have censused breeding waders and other arctic birds in fixed study plots. Besides information on geographical distribution of species, which is still quite valuable in the vast Eurasian tundra, the fact that teams have visited the same areas in several years has given some insight into annual variations in their breeding bird fauna. These variations appear to be quite substantial, especially due to the fact that some of the most abundant species (e.g. Little Stint Calidris minuta and Curlew Sandpiper C. ferruginea) lack breeding site fidelity and react to snow conditions upon arrival in the tundra. Other species (e.g. Grey Plover Pluvialis squatarola and Knot C. canutus) are more site-faithful and numbers seem more stable from year to year. Although the lack of site fidelity devaluates long-term census programs of breeding numbers as a tool for monitoring population size, the question what factors affect the distribution of breeding birds on a more local scale is interesting in itself. Such longer-term censuses at single sites have been rarely conducted in the tundra's of Russia.

At the Willem Barentz Station on Taimyr, activities are currently undertaken to start a long-term monitoring program for breeding waders. One of these is the development of a suitable census method; in view of the substantial between-year variation in local distribution of birds, quick mapping censuses or transect counts over a wide area surrounding the station may yield more useful information than a complete census based on nest searches in a single (and inevitably small) area. It is intended to repeat censuses at Medusa Bay on a regular, if possible annual, basis.

Research priorities:

* Developing a suitable technique for long-term monitoring of breeding waders in the Eurasian tundras.

* Regular (annual) census projects to implement this technique at a single site.

Breeding ecology Given the extreme climate of the polar region (Meltofte 1985), it seems surprising that so many waders migrate to arctic latitudes to reproduce. In fact, chicks of arctic-breeding wader species show higher growth rates than more temperate species. Some work was done in Knots to explore what makes this fast growth possible. Rather than an increased food supply, arctic Knot chicks seem to profit from the combination of continuous daylight and an increased cold-hardiness in comparison with temperate-breeding waders (61). A scarcity of parasites in the high Arctic may be a further contributing factor (Piersma 1997).

In the same study (61), indications were found that species may differ in their chicks' ability to grow under cold conditions, species with a more northerly distribution being more tolerant of cold than less high-arctic ones even within the same area. This topic would be a very interesting one to explore further, as it could provide insights in the factors that delineate a species' breeding range (Castro et al. 1992, Gratto & Cooke 1987), and is thus highly important for predicting potential effects of climate change on arctic birds. It would involve measuring effects of weather and insect availability on growth rate in different species and different latitudes, including the relation between insect abundance and climate.

The arctic climate affects not only chicks, but adult birds too (Norton 1973, Ashkenazie & Safriel 1979). Measurements on Knots (61) and Little Stints (57) have shown that incubating waders incur high energetic costs, close to hypothetical energetic limits. In combination with the time demands posed by incubating the clutch, this may lead to problems with energy balance during cold periods, especially in the smaller species. This was studied in the Little Stint, in which the problem is aggravated by the multi-clutch breeding system which leads to parents having to incubate clutches alone. Using small field-proof temperature loggers placed in the nests, It was found that Little Stints spend more time foraging, instead of incubating, during cold weather, but still need to draw on extensive energy stores built up during more favourable periods (57). These stores are at least as large as found in birds wintering on the seaboards of Europe. In larger wader species, incubation stores are smaller, which makes it interesting to perform comparative time allocation studies in such species.

In addition to factors operating in the Arctic itself (Holmes 1966, 1970), breeding performance of arctic birds may be influenced by factors operating elsewhere during the annual cycle, in wintering (departure) areas or along the migration routes (Prop et al. 1984). Quantity and quality of spring staging areas is an example of such a potential factor, and may be affected by human activities such as habitat destruction, pollution and disturbance or man-induced climate change. In recent years, theoretical models have been developed which try to predict the impact of changes in staging conditions on the fitness of migratory birds (Weber et al. 1994, in press). These models depend on the assumption that there is a fitness cost of arriving on the breeding grounds late (relative to other birds or to the peak in food supply) or lean (in poor body condition). This assumption has not yet been tested on arctic birds other than geese, and this is another issue that could be addressed by future workers. Such studies could benefit greatly from the existence of permanent field stations in the Arctic, as they should span several years to make use of natural variation in arrival conditions. They should preferably make use of an individually marked population, and a species with strong breeding site fidelity.

In all three aspects of breeding ecology mentioned above, (variation in) the availability of food plays a major role. Some work has been done on the surface activity of arthropods in most of the past WIWO-expeditions, but this topic could be much further developed. Surface-active spiders and insects certainly are a major food source for many arctic-breeding waders, but little is known yet about the additional importance of soil fauna, or about interspecific differences in prey choice, or about spatial variation of arthropod abundance within the tundra. Such data may help explain differences between species in habitat preference within the tundra. Also, while all studies have found a strong effect of weather on surface activity of arthropods, there are no direct observations relating this to the intake rate of feeding waders, either adults or chicks. These are obviously not the easiest observations to make, but they are needed for a further understanding of arctic breeding patterns. The tameness of many arctic birds should be of some help in this respect.

The work that was done on breeding of Broad-billed Sandpipers in northern Norway provides a somewhat special case. This species has a restricted breeding area centred on northern Scandinavia. Population size (small), habitat requirements and breeding biology are all poorly known. The two summer projects, and other work by Norwegian counterparts of NINA-NIKU have provided information on several of these aspects. The species seems to require a specific type of wet bogs for breeding. These bogs may be recognisable from satellite images which could then be used to identify potential breeding sites and, in combination with ground work, predict population size. Bogs with Broad-billed Sandpipers also held a high diversity of other waders and the species could thus provide a useful handle for conserving important wader breeding areas in northern Scandinavia. Several of these wet bogs were under threat of drainage (deliberately for creating reindeer pasture or accidentally by driving with off-road vehicles).

Research priorities:

* Describing breeding season diets of arctic-breeding waders and the relation between foraging success (food availability) and weather conditions.

* Further work on the relationship between weather, arthropod availability and growth rate of arctic wader chicks of different species and under varying conditions.

* Further comparative studies of the energetic limitations of incubation and the role of energy stores among wader species.

* Studies of the effects of arrival date, arrival condition and prelaying weather conditions and food availability on breeding success.

* Studies of mating systems, breeding site fidelity and philopatry in arctic, migratory birds.

* Describing the early stages of autumn migration of waders from the tundra.

* Further work on the distribution and habitat requirements of Broad-billed Sandpipers, and threats to their breeding habitat.

4.3.2. Northwest Europe

Various long-term monitoring projects on the number and distribution of wader and waterfowl are institutionalised (e.g. several BTO projects in the British Isles, IWRB mid-winter counts, OAG-Münster) and will be further developed (e.g. Waddensea). Ecological research considering the entire flyway, or on human disturbance, damage to agricultural crop etc. is focused on several target species. Potential research capacity is relatively large when compared to other regions. Therefore, WIWO will focus its attention primarily on other regions.

4.3.3. Central and Eastern Europe

The monitoring of wader and waterfowl (mainly goose and duck species) is relatively well organised and established in Northwest Europe. In eastern Europe, several interesting wader species (e.g. Great Snipe Gallinago media, Terek Sandpiper Tringa terek, Marsh Sandpiper Tringa stagnatilis) breed in bogs and riparian wetlands in Russia (west of Ural), Belorussia and Ukraine riparian wetlands in particular. The distribution and biology of these species is not well known.

* Breeding wader surveys of riparian wetlands in Belorussia, Ukraine and Russia (west of Ural).

4.3.4. Black Sea and Sea of Azov

Wetland types Several large deltas occur in the northern part of the region (e.g. Danube, Dnestr, Dnepr and Kuban) and one large coastal lagoon (Sivash Saliv). The north coast of the Black Sea and Azov Sea is, apart from the deltas, dominated by an array of small, so called limans, which are comparable to small estuaries. Above this, extensive shallows along land spits or peninsulas (Biruji Peninsula, Arabatskaja Strelka) are typical for the northern shores of Black and Azov Sea west of the Crimea. The lagoons, shallows and limans are highly productive (extensive eelgrass beds, rich in macrozoobenthos) and support large numbers of waterbirds during most of the year.

Extensive steppe areas with extensive cattle grazing are present in the northern part of the area. Small parts of natural, prime steppe vegetation still remain in several nature reserves. Along the southern coastline of the Black Sea, the coastal wetlands are mainly deltas (e.g. Kizilirmak and Yesilirmak). Inland wetlands like riparian river ecosystems, freshwater or brackish (playa) lakes or inland marshes occur on both sides of the area.

Current knowledge Wilson & Moser (1994) provide a good overview of current knowledge and conservation status of all major wetlands in the region. The area is important for huge numbers of waterbirds. Not less than eight globally-threatened bird species occur in the Black Sea region. The coastal and marine wetlands along the northern Black Sea coast and Sea of Azov are relatively well studied and protected (winter counts of Wetlands International, IBA programme of the Ukrainian bird protection society UTOP). Numbers of migrating waders and waterbirds as well as numbers of breeding and wintering waterbirds are known for parts of the region (e.g. Brehme et al. 1992 [43], Chernichko et al. 1991, Van der Have et al. 1993 [36], Korzyukov et al. 1991, Lysenko 1991, and Siokhin et al. 1988).

Wetlands along the north-eastern coast of the Sea of Azov and Black Sea (e.g. Kuban Delta remain relatively unknown. Wetlands along the Turkish coast are relatively well studied although information on some areas is scarce. To date, WIWO contributed in the knowledge of birds of the Sivash (spring counts and migration ecology of waders) and Kizilirmak Delta (breeding bird census). In the period 1994-1998 a complete waterbird count was organised in the Sivash, and the feeding ecology, distribution, biometrics and moult of terns and waders was studied.

Future research For almost all wetlands in the area bird counts are missing for at least some periods of the year. This includes breeding bird counts as well as counts outside the breeding season. Breeding bird counts are especially needed for wetlands in the eastern part of the Sea of Azov and Black Sea (e.g. Kuban Delta). Counts outside the breeding season are needed for almost all wetlands. This means that almost all wetlands need at least in one part of the season additional information on bird numbers. For the entire region there is a general lack of knowledge of the function of wetlands for birds during their annual life cycles. The importance as a moulting area for ducks (e.g. feeding on eelgrass), gulls and terns is largely unknown.

Very little is known about the impact of fisheries on birds feeding on fish in lagoon systems or at sea. Moreover the ecological background of wetlands in this continental climate is relatively unknown. The saline and hypersaline lagoons are usually dry in summer and autumn, which means that some species likely have to adapt their migration strategy. They may choose different migration routes compared with spring (e.g. Western Europe, with stopovers in the Wadden Sea) or skip the area during migration to moult more southerly (e.g. Broad-billed Sandpiper).

In addition, very little is known about the importance for birds of the steppe along the north coast of the Black Sea. The impact of agriculture and cattle grazing on steppe habitats is relatively unknown. Some rare bird species likely are influenced by deterioration of steppe vegetation (e.g. Great Bustard Otis tarda, Black-winged Pratincole Glareola nordmanni).

Research priorities

* Breeding bird surveys in wetlands along the eastern part of the Northern Black Sea.

* Waterbird surveys in almost all wetlands with emphasis on spring and autumn.

* Identification of most important food types for the majority of waterbirds or guilds of waterbirds;

* Moult studies of waterbirds (e.g. terns) and foraging during this vulnerable period.

* The importance of eelgrass for waterbirds in coastal lagoons and shallows.

* The impact of the dry summer periods on wetlands and migration strategies of waterbirds.

* The importance of steppe habitats for wetland birds.

* Breeding bird surveys of riparian freshwater areas.

4.3.5. Mediterranean

Wetlands types In the Mediterranean coastal as well as inland areas occur. Coastal areas are formed by large or small river deltas (e.g. Nile or Göksu) or estuaries (e.g. Gulf of Gabès). Especially in Turkey large inland wetlands occur (e.g. Tuz Gölü, Magnin & Yarar 1992).

Current knowledge Because knowledge of wetlands in the northwestern part of the Mediterranean (Spain, France, and Italy) is good and potential research capacity is large, WIWO has focused its attention primarily to other regions in the Mediterranean. Since the last Forward Plan the research capacity of other countries increased (e.g. Turkey). Winter surveys (Wetlands International) are executed in most countries and information about the winter period is therefore rather good. Spring surveys were the main WIWO projects in the Mediterranean, while autumn projects are still needed at most places. In the period 1994-1998 WIWO projects have been executed in Albania (breeding bird survey and study of Pygmy Cormorant in 1996) and Tunisia (early spring migration of waders 1994).

Future research The most important gaps in knowledge identified in the last WIWO Forward Plan still remain. In winter the irregular coverage of the Eastern Mediterranean and the Maghreb precludes accurate population estimates and trends, whereas waterfowl are better covered than waders are. Recently the importance of inland wetlands in the Maghreb increased with the discovery of large numbers of Marbled Teal. Counts of these wetlands, especially simultaneously in different countries are urgently needed. In spring wetlands studied in 1990 should be studied again. The impact of rapid habitat changes in the Mediterranean wetlands, on birds can be identified in this way.

Moreover studies in autumn and late summer are needed in almost all wetlands. Especially important stopover sites like the Gulf of Gabès, the Çukurova deltas, the Nile Delta and some smaller wetlands in Turkey and Greece need attention. As for wetlands in the Black Sea region the autumn situation is completely different from the spring situation in terms of water availability. The demands of water by agriculture can be competitive with the need for water for nature. All future activities should fit in programs of bird research in the Mediterranean.

Research priorities

* Breeding bird surveys in wetlands in the Maghreb countries.

* Monitoring waterbird numbers, habitat availability and food resources at a selected number of sites: Gulf of Gabès, Lake Kelbia (Tunisia), Nile Delta (Egypt), Çukurova deltas (Turkey).

* Repeated counts of major Mediterranean wetlands in spring (monitoring concept).

* The importance of inland wetlands in the Maghreb (e.g. dam lakes and oasis) for Marbled Teal, in particular in winter.

* Autumn counts of waterbirds in Gulf of Gabès (Tunisia), Nile Delta (Egypt), Çukurova deltas Turkey) and other wetlands.

* Moult studies of waders and terns (e.g. autumn Gulf of Gabes).

* The impact of water level variability on waterbirds: comparisons of dry periods (summer and autumn) with wet periods (spring).

* Identification of foraging guilds of waterbirds.

* General studies of fish-eating birds in coastal lagoons and intertidal wetlands.

* Surveys of colour-marked waterbirds and checking harbours and fishermen for ring recoveries.

4.3.6. Caspian and Arabian

Marine wetlands without tidal range, deltas and brackish lagoons occur along the Caspian Sea (Northern Iran, Azerbeijan, Volga Delta, Russia) including some wetlands with considerable international importance (Evans 1994). The Arabian coastline of the Gulf is fringed with extensive intertidal areas, which are very shallow and sandy, numerous small islands and locally mangroves. The tidal range is relatively small and often overshadowed by wind force. Due to the sheer size, combined with high temperature, high humidity and almost continuous, unhindered sunshine, there have been relatively few surveys of these wetlands (Aspinall 1995). Along the Iranian coast several estuaries occur with increasing coverage of mangroves to the east. There are hardly any quantitative data on migrating and wintering waders in these Iranian wetlands.

Gulf A number of studies have been carried out in Bahrain, United Arab Emirates (UAE) and Oman (Uttley et al. 1988, Zwarts et al. 1991, Scott & Rose 1989, Keijl & Ruiters 1998) and a long-term programme has been started in Saudi Arabia (Symens pers. comm.). Especially the Gulf War triggered much interest in this region and revealed the numerical importance of this region for wintering waders (Zwarts et al. 1991, Evans & Keijl 1993a,b), although predominantly for the East African Flyway. However, the European populations of Broad-billed Sandpiper and Red-necked Phalarope migrate almost exclusively through this region and probably moult and winter here (Broad-billed Sandpiper). No key moulting sites for the Fennoscandian population of Broad-billed Sandpiper are yet known.

Red Sea Nothing is known of the Red Sea area, although extensive mudflats and mangroves are present in the Southern part (e.g. Yemen). The political situation makes work in this region difficult in some countries.

Inland wetlands Lake Uromiyeh is one of the most important inland wetlands in Iran (Evans 1994) and among the largest saline lakes in the world. It is shallow and fringed with extensive reedbeds near river mouths, saline mudflats and tens of small islands. A large Flamingo colony is present on small islands and the coastline of the lake is probably an important stopover site in particular for waders foraging on brine shrimps (e.g. Red-necked Phalarope and Calidris-species, Verkuil et al. 1993).

Research priorities

* Waterbird census of Lake Uromiyeh, northwest Iran, in spring and autumn.

* Waterbird census of the Gulf estuaries Hilleh and Monde River deltas in autumn to check for moulting Broad-billed Sandpipers.

* Waterbird census of coastal wetlands in winter to check for Slender-billed Curlew.

* Feeding ecology studies of wintering waders, especially Broad-billed Sandpipers in UAE and Oman.

* Studies of waterbird migration ecology in wetlands along northern coast of Caspian Sea.

* Feeding studies of fish-eating birds (grebes, cormorants).

4.3.7. Central Asia

Several globally threatened or vulnerable waterbirds breed in wetlands in Central Asia, such as Pygmy Cormorant Phalocrocorax pygmeus, Dalmatian Pelican Pelecanus crispus, Marbled Duck Marmaronetta angustirostris, Corn Crake Crex crex, Slender-billed Curlew Numenius tenuirostris, Asiatic Dowitcher Limnodromus semipalmatus, Sociable Plover Vanellus gregarius. Wetland types range from permanent bogs and peatlands in the north, forest steppe at intermediate latitudes and ephemeral saline lakes with salt marshes and temporary wet grasslands near rivers. Riparian wetlands occur mainly in the north. Stopover sites for waders are probably widespread and few sites with concentrations are known.

Research priorities

* Breeding bird surveys of forest steppe and wet grasslands in steppe areas, with focus on Slender-billed Curlew and other globally-threatened species.

* Migration ecology studies of waterbirds at stopover sites.

4.3.8. Inland wetlands in the Sahel

Census work The inland wetlands of the Sahelian Region (i.e. savannah and steppe habitats of West and East Africa), and the East African Nile Basin and Rift Valley are the main wintering grounds for a number of Palaearctic waterbird species. The main wader species are Little Stint, Ruff, Black-tailed Godwit, Little Ringed Plover, Common Ringed Plover, Black-winged Stilt, Collared Pratincole, Common Snipe and most Tringa species.

For waterfowl, the region is particularly important for Garganey, Pintail and Shoveler (Perennou 1991, 1992). Their number in this region is estimated to be about 4 million (Monval & Pirot 1989). Core areas where mass aggregations of wintering waders and waterfowl have been found are the Senegal Basin, Inner Niger Delta, Lake Chad Basin and Nile Basin.

Although considerable progress has been made in counting wintering waterfowl by means of aerial surveys at the Sahelian wetlands in West Africa, these counts have been irregular and not well synchronised, whereas coverage of counts between years was such that no accurate calculations of trends or population estimates are possible yet (Rüger et al. 1986). No such counts have been conducted for the Nile Basin and Rift Valley (Perennou 1991, 1992, Rose & Pirot 1990).

Few numerical data are available on wader numbers inland. The method of aerial surveys in counting ducks is not applicable for counting waders. Waders are small, rarely aggregate as ducks do and occur dispersed over large areas. The efficiency of ground surveys in terms of effort and results have proved to be poor. Logistical problems are enormous. Erratic flooding and unpredictable variability of circumstances adds further difficulty. The estimates of wintering waders are therefore to be considered very unreliable.

Apart from the known extensive key sites in the Sahelian and East Africa, comparatively small sized temporary floodplains can host important numbers of waders and waterfowl (e.g. van Wetten & Spierenburg 1998 [67]). During a ground survey in five southern districts of Niger, January 1992 and January 1993, twenty such temporary wetlands were visited and produced wintering numbers of 18,000 in 1992 and 42,000 in 1993 (Mullié et al. 1993).

Rainfall variability The condition, availability and distribution of the inland wetlands throughout the Sahel, Sudan and Ethiopia are dependent on rainfall patterns. Distribution of wintering waders and waterfowl can vary greatly within the wintering season and between years depending on the condition and amount of available habitats. These species seem to be able to search and find favourable circumstances in this vast area. Circumstantial evidence suggests strong movements between the major wintering resorts. Winter movements might include this entire west to east belt of the African continent. This mechanism might be the practical explanation for loop-migration as has been documented for several species from this region, i.e. species having their main breeding distribution in Central Russia and Siberia, moving to North-western Europe in autumn from which they enter Tropical Africa. In search of suitable wintering conditions they might be forced eastward in the Sahelian from where they return to their breeding grounds.

The last two decades have shown severe droughts in the West African Sahel and in Sudan and Ethiopia. Rainfall data showed a clear decline in mean annual rainfall values, a clear shift in the onset and ending of the rains during the drought years and the rains becoming less frequent. As a result, the isohyets in Niger for example, have shifted 100-150 km southwards (Sivakumar 1992). Rainfall figures improved during the past two years. Winter conditions and the occurrences of flood plains are part of a fragile system affecting the entire savannah and steppe zone of this continent. Hardly anything is known on the strategy, capacity, and constraints of species to cope with these circumstances, whereas these unpredictable winter conditions might well be a major bottleneck in the life-cycle of wintering species involved (e.g. van der Have 1991, Peach et al. 1992).

Desertification At least 20% of the African continent consists of deserts and another 60% are seriously affected by erosion. The steady population growth is causing increased land pressure. The custom of fallow land in the traditional farming system is disappearing. As a result, the recent increased rainfall added serious water erosion to the long list of environmental problems in the region. Over the past ten years, overgrazing increased with 20%. Vast irrigation schemes are launched and include the manipulation of all river systems.

These developments will no doubt have an accumulated impact on the amount and distribution of wetlands and on the wintering conditions for waders and waterfowl of the region in particular. Hence, there is a need to get a better notion on the functioning of this fragile system for wintering waders and waterfowl.

Research priorities

Given the rapid environmental changes in this region, and the need for increased agricultural development affecting the water management of the entire region, basic information on waterbirds is urgently required. Hence, methods to tackle and overcome the immense organisational, logistic and topographical obstacles are to be developed before research can be conducted. For the moment support for local initiatives in this field is needed.

* Ground surveys of waterbirds in the Senegal Basin, Inner Niger Delta, and Lake Chad Basin (Chad, Mali, Niger, and Senegal).

* Desk study on relation between rainfall patterns and shifts in distribution of waterbirds.

* Field and desk studies of the relation between rainfall variation, habitat changes and abundance and condition of birds depending on wetlands during the non-breeding season (waterbirds and passerines).

4.3.9. Coastal West Africa

Census work With regard to winter and spring, baseline information on distribution, species and numbers involved in the region are relatively well known. However, some important staging areas are still gaps in our knowledge. For these areas, the first midwinter counts still have to be carried out to judge their importance for staging and/or migrating Palaearctic waterbirds as well as Afrotropical species. Surveys in these areas are welcomed.

On the other hand, regular census activities are lacking in many key site areas. Although the African Waterfowl Census (AfWC), co-ordinated by Wetlands International is growing rapidly in terms of participating countries and areas covered (Dodman et al., 1997), it is still difficult to perform regular counts by local birdwatchers and professional ornithologists. Especially for key areas, such as the Banc d'Arguin (Mauritania), the Archipelago dos Bijagos (Guinea Bissau) and other deltas and estuaries of the larger West African rivers, it is of great importance to carry out complete censuses on a regular basis (monitoring). These counts must be set up in close co-operation with local organisations in order to build a network and capacity of local birdwatchers.

This is very well possible within the framework of the Wetlands International-EAME Project 'Building the Capacity in West Africa of a regional network for wetland and waterbird management (1997/98-2001)'. This project comprises a mixture of institutional support, capacity building and the development of lasting programmes of research, especially monitoring, in environmental, ecological and socio-economic fields. Activities performed by WIWO are possible and can be financed under this programme.

To address and work out the collaboration between WIWO and Wetlands International concerning projects in West Africa, both organisations have signed a 'Terms of Reference' (1998). In 1998, projects have been carried out already successfully (Senegal, Mauritania). During the African Waterfowl Census Meeting in 1996 (Djoudj, Senegal), where WIWO presented its work and aims, it has become clear that there is still a great need for assistance in bird census and training of local people (Dodman, 1997).

Research priorities

The research priorities for 1999-2003 are a continuation of earlier activities as well as some possible new activities:

Waterbird censuses

* Regular waterbird counts in areas, which are extremely important for Palaearctic waterbirds, during northern winter (e.g. Banc d'Arguin, Archipelago dos Bijagos, and other coastal wetlands).

* Census of possible key areas, that have not previously censused. Examples are Casamance Delta (Senegal), Gambian Delta and others.

* Census of Afrotropical waterbirds and colonial breeding bird surveys in West-African wetlands.

* Surveys of endangered and vulnerable Palaearctic and/or Afrotropical waterbird species.

* Studies of the importance of coastal wetlands for summering waterbirds, especially for waders with respect to the high variability in breeding success of arctic species.

Ringing projects

* Ringing of Palaearctic waterbirds and marshland birds on a regular basis, in order to set up bird ringing centres and/or constant effort sites in close co-operation with local organisations;

* Once-only ringing activities of waterbirds in combination with waterbird censuses (especially waders and terns).

Feeding ecology

* Feeding ecology studies in relation to food availability in various wetlands and/or for certain species, for wintering and (colonial) breeding bird species.

Training and transfer of knowledge

* To assist in building capacity of local bird watchers in order to maintain field work activities by training in bird counts;

* To transfer knowledge on organising, collecting, processing and reporting field data.

4.3.10. East Africa

Since the previous reviews in the Forward Plan 1989-1993 and 1994-1998, very little progress has been made in this region of which knowledge was already quite limited. The most notable exception has been an inventory of wintering waders along the coast of Tanzania by a Danish team (Bregneballe et al. 1990). This study was very worthwhile both in establishing wintering numbers in a previously little-studied area, and in developing a census technique which can be applied much more widely in the region, where coral flats bordered by mangroves on the landward side form the major coast type from Somalia down to Mozambique. This work has been continued on a larger scale in the Tanzanian Waterbird Census in 1995 (Baker 1997). As part of this wintering waterbirds on the islands of Zanzibar and Pemba were counted in January 1998 (Geene et al. in prep.)

Counts in major wintering and staging sites of waterfowl are available only for some countries in the region, notably Kenya and Tanzania (e.g. Schekkerman & van Wetten 1987). The prospects for filling the gaps in the near future are not good, considering the political situation in the majority of countries in the region, which is at least as unstable as it was five years ago.

Exceptions are Ethiopia and Madagascar, where some work might be possible in the near future. Two Ethiopian endemics (apparently not threatened) occurring almost exclusively in small, highland wetlands are little studied: and Blue-winged Goose Cyanochen cyanopterus and Spot-breasted Plover Vanellus melanocephalus. Seven endemic, and globally-threatened waterbirds occur in Madagascar: Alaotra Grebe Tachybaptus rufolavatus, Madagascar Grebe T. pelzelnii, Madagascar Heron Ardea humbloti, Madagascar Teal Anas bernieri, Madagascar Pochard Aythya innotata (probably extinct), Madagascar Pratincole Glareola ocularis, Black-banded Sandplover Charadrius thoracicus.

Research priorities

When political complications are left out of consideration, the following research priorities can be listed:

* Censuses of wintering waders along the Red Sea and Indian Ocean coasts of Sudan, Eritrea, Djibouti, Somalia, Kenya, and Mozambique.

* Censuses of wintering waterbirds in inland Sudan and Ethiopia.

* Studies of endemic waterbirds of Ethiopia and Madagascar.

* Invertebrate food resources of mangrove-coral flats.

4.3.11. Southern Africa

The coastal wetlands of Namibia (Walvisbaai) and the Republic of South Africa (e.g. Langebaan Lagoon) are relatively well studied and hold important numbers of predominantly high arctic waders. It is increasingly recognised also as an important wintering area for Palaearctic terns, in particular Black Tern, Common Tern and Sandwich Tern. The inland wetlands of Zimbabwe and Botswana, however, are relatively unknown. The huge Makgadikgadi Saltpans in central Botswana, for example, can hold Palaearctic waders in years with abundant rainfall, as well as huge numbers of flamingo's, but little information is available on its numerical importance.

Other huge but permanent wetlands in this region, like the Okavango delta and the floodplains of the Chobe-Lyanti river in system in northern Botswana and the Zambezi floodplains in Zimbabwe probably hold relatively small numbers of Palaearctic waders like Ruff, Wood Sandpiper and Little Stint. They can be potentially important for relatively unknown species like Black-winged Pratincole and Great Snipe (see also table 4). These areas, however, are probably very important for intra-African migrants and African waterbirds, but are relatively understudied despite their international reputation (e.g. Okavango delta). Good logistics and a relatively stable economy in parts of this region will greatly facilitate future projects.

Research priorities

* An integrated waterbirds census of the Okavango Delta combining aerial surveys, with ground transects by boat and information from satellite maps in co-operation with local organisations.

* Inventory of waders and terns staying the non-breeding season along the coast between Gabon and Namibia.

* Studies of the geographical origin and feeding ecology of Afro-Palaearctic terns staying the non-breeding season in Namibian coastal wetlands.



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Zwarts, L. & T. Piersma. 1990. How important is the Banc d'Arguin, Mauritania, as a temporary staging area for waders om spring? Ardea 78: 113-122.

Zwarts, L. & A.-M. Blomert. 1990. Selectivity of Whimbrel feeding on fiddler crabs explained by component specific digestibilities. Ardea 78: 193-208.

Zwarts, L., A-M. Blomert, B.J. Ens, R. Hupkes & T.M. van Spanje. 1990. Why do waders reach high feeding densities on the intertidal flats of the Banc d'Arguin, Mauritania? Ardea 78: 39-52.

Zwarts, L. & S. Dirksen. 1990. Digestive bottleneck limits the increase in food intake of Whimbrels preparing their departure from the Banc d'Arguin, Mauritania, in spring. Ardea 78: 257-278.

Zwarts. L. 1988. Numbers and distribution of coastal waders in Guinea-Bissau. Ardea 78, (1/2), 42-55.

Zwarts, L. J. van der Kamp, O. Overdijk, T. van Spanje, R. Veldkamp, R. West & M. Wright. 1998. Wader count of the Banc d'Arguin, Mauritania, in January/February 1997. Wader Study Group Bull. 86: 53-69.

Zwarts, L. J. van der Kamp, O. Overdijk, T. van Spanje, R. Veldkamp, R. West & M. Wright. 1998. Wader count of de Baie d'Arguin, Mauritania, in February 1997. Wader Study Group Bull. 86: 70-73.



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ANNEX 3. Projects with reference number

(see table 1):

1. Mauritania 1980
2. Morocco 1981
3. Tunisia 1984
4. Mauritania 1985
5. Mauritania 1986
6. Guinea-Bissau 1987
7. Greece 1987
8. Turkey 1987
9. Mauritania 1988
10. Morocco 1988
11. Guinea Conakry 1988
12. Turkey-2 1988
13. Turkey-3 1988
14. Turkey-1 1988
15. Israel 1989
16. Turkey-2 1989
17. Turkey-1 1989
18. Guinea-Conakry 1990
19. Greece 1990
20. Tunisia 1990
21. Turkey 1990
22. Egypt 1990
23. Ukraine 1990
24. Morocco 1990
25. Russia 1991
26. Gabon 1992
27. Guinea-Bissau 1992
28. Ukraine 1992
29. Russia 1992
30. Cameroon 1993
31. Botswana 1993
32. Libya 1993
33. Albania 1993
34. Ukraine 1993
35. Iceland 1993
36. Russia 1993
37. Turkey-2 1992
38. Turkey-1 1990
39. Turkey-1 1992
40. Botswana 1994
41. Tunisia 1994
42. Russia-Sterl. 1994
43. Russia-Piass. 1994
44. UAE 1994
45. Botswana 1995
46. UAE 1995
47. Senegal 1996
48. Greece 1996
49. Albania 1996
50. Norway 1996
51. Russia-Taimyr 1996
52. Russia-Kargopol 1996
53. Senegal 1997
54. Mauritania 1997
55. Ghana-1 1997
56. Russia-C.Siberia 1997
57. Ghana-2 1997
58. Mauritania 1998
59. Tanzania 1998
60. Russia-Moscow 1998
61. Senegal 1998
62. Russia-Taimyr 1998
63. Ukraine 1998