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ARTICLE IN PRESS B I O L O G I C A L C O N S E RVAT I O N x x x ( 2 0 0 7 ) x x x –x x x available at www.sciencedirect.com journal homepage: www.elsevier.com/locate/biocon A quantitative method for evaluating the importance of marine areas for conservation of birds Henrik Skova,*, Jan Durinckb, Mardik F. Leopoldc, Mark L. Taskerd a DHI Water and Environment, Agern Alle 5, DK-2970 Hørsholm, Denmark Marine Observers, Svankjaervej 6, DK-7752 Snedsted, Denmark c Wageningen-IMARES, P.O. Box 167, Landsdiep 4, NRL-1797 SZ Den Hoorn (Texel), The Netherlands d Joint Nature Conservation Committee, Dunnet House, 7 Thistle Place, Aberdeen AB10 1UZ, United Kingdom b A R T I C L E I N F O A B S T R A C T Article history: Objective criteria are needed for ranking marine sites when examining candidate areas for Received 19 February 2006 protection measures. We suggest a Marine Classification Criterion (MCC) which allows the Received in revised form application of the widely used Ramsar 1% criterion for wetlands for seabirds with clustered 3 December 2006 distribution in offshore habitats. The maximum size of an area considered to be interna- Accepted 6 December 2006 tionally important has not been defined by the Ramsar Convention. Terrestrial and coastal sites generally have obvious hydrological or physical boundaries, whereas such boundaries are less obvious at sea. The smallest unit which would pass the demands set by the MCC is Keywords: 1% of the bio-geographic population of a particular species concentrated in an area (site) Marine classification criterion supporting a density exceeding a value equivalent of four times the average density of Seabirds the species in the investigated regional sea. The effect of choosing smaller or larger refer- Marine protected areas ence densities is tested. The results indicate that the chosen threshold density is a suitable Identification of concentrations requirement for the inclusion of the most important areas for seabird species with at least 25% of their bio-geographic population occurring in the studied regions of the North Sea and the Baltic Sea. The test cases indicate that provided the MCC is based on geo-statistical analyses of un-biased survey data the boundaries of areas holding large concentrations of seabirds can be estimated with confidence. The MCC could be used to identify concentrations of seabirds and other marine animals of conservation priority and to rank marine areas by their cumulative importance to different species. Ó 2007 Elsevier Ltd. All rights reserved. 1. Introduction The IVth World Congress on National Parks and Protected Areas held in Caracas, Venezuela, in 1992 (IUCN, 1993), highlighted the need for protected areas in marine as well as terrestrial environments in order to conserve biodiversity. In the following years, scientifically-based methods (e.g. gap and complementarity analysis) for prioritising conservation efforts have been developed and tested (Fjeldså and Rahbek, 1998; Williams, 1998). It has, however, been difficult to define important marine areas, and relatively few marine protected areas and integrated marine management zones exist today. The lack of obvious boundaries, of quantitative information and of jurisdiction has hampered the designation of areas for conservation in offshore habitats. This paper discusses the problems related to identifying and delineating areas for conservation for seabirds, and it proposes a procedure which ranks the conservation value of different marine areas for * Corresponding author: Tel.: +45 45169220; fax: +45 45169292. E-mail addresses: [email protected] (H. Skov), [email protected] (J. Durinck), [email protected] (M.F. Leopold), [email protected] (M.L. Tasker). 0006-3207/$ - see front matter Ó 2007 Elsevier Ltd. All rights reserved. doi:10.1016/j.biocon.2006.12.016 Please cite this article in press as: Skov, H. et al., A quantitative method for evaluating the importance ..., Biol. Conserv. (2007), doi:10.1016/j.biocon.2006.12.016 ARTICLE IN PRESS 2 B I O L O G I C A L C O N S E RVAT I O N seabirds based on existing quantitative criteria for selecting areas of international importance while taking the size of the areas into consideration. The method is applicable to all organisms for which reliable regional density and population estimates exist, but seabirds were chosen to illustrate the procedure, as these are highly visible organisms for which relatively much information is available. Over the last 15 years, scientifically-based methodologies for identifying and prioritising conservation areas have been developed. These methods define target areas based on species diversity or rarity, such as analyses of hotspots (Myers, 1990; ICBP, 1992) or more elaborately in analyses of complementarity (Vane-Wright et al., 1991; Fjeldså and Rahbek, 1998; Williams, 1998). However, these criteria may not be useful measures of the potential of sea areas for the conservation of birds, as large numbers of birds can be concentrated in species-poor marine seascapes such as estuaries and upwelling areas (Summerhayes et al., 1974). Kelleher and Kenchington (1992) proposed a set of general, qualitative criteria to be used for the selection of marine protected areas, but paid no attention to the specific conservation requirements of seabirds. Full, effective protection of widely ranging animals like seabirds needs the co-operation between several countries. Site-based conservation of birds has a long history in terrestrial, limnic and coastal ecosystems, and has played an important role in the development of today’s networks of protected areas worldwide. The overarching goal of site-based conservation of birds has been to identify those sites that are exceptionally important for birds at some period during the year, – ones of significance both nationally and internationally, and to try to ensure the conservation of the features that make these areas important. Examples of site-based bird protection systems are the EU network of Special Protected Areas (EU Birds Directive, 1979) and the global series of Important Bird Areas proposed by BirdLife International (Heath et al., 2001). Quantification of the importance of coastal and inland wetland areas for birds has long been based on a widely agreed set of international criteria under the auspices of the Convention on Wetlands of International Importance especially as Waterbird Habitat (the Ramsar Convention). To be considered internationally important for waterbirds, a wetland site has to meet any one of two criteria (Ramsar Convention Bureau, 1988): 1. It regularly supports 20,000 waterbirds. 2. It regularly supports 1% of the individuals in a population of one species or subspecies of waterbird. The 1% criterion has been particularly useful because it is comparatively easy to count waterbirds, and thereby derive both site counts and global population estimates. There is no biological reason to use 1% of a population as the threshold level for establishing international importance of a site. However, this percentage has gained wide acceptance and we see no reason to challenge it. The size of an area considered to be internationally important is, however, not defined. A larger site is by definition likely to contain more birds than a smaller site centred around the same location. Terrestrial and coastal sites generally have obvious hydrological or physical boundaries, whereas such boundaries are less obvious for x x x ( 2 0 0 7 ) x x x –x x x marine sites. Another restriction for the application of the Ramsar Convention criteria to the open sea is that ‘wetlands’ are limited to areas that are no deeper than 6 m. Despite these limitations, selection of conservation areas for birds by the application of the Ramsar criteria has taken place in truly marine areas, e.g. EU Special Protection Areas and Ramsar sites in the Danish part of the Kattegat (Jensen, 1993). Other approaches for selection of priority areas for conservation of birds in marine waters include seaward extensions of nationally and internally important breeding colonies (e.g. United Kingdom; Johnston et al., 2002), habitat features related to national marine sanctuary systems (e.g. Stellwagen Bank, USA; http://sanctuaries.noaa.gov/) or analyses of tagging data (e.g. Birdlife International’s ‘‘Ocean Wanderers’’ project (BirdLife International, 2004). The use of the 1% criterion requires the availability of estimates of total population size as well as estimates of the Table 1 – Total estimates of selected seabird species wintering in the Baltic Sea and the North Sea (1987–1995) Species Red-/black-throated diver Gavia stellata/arctica Great Northern Diver Gavia immer Great Crested Grebe Podiceps cristatus Red-necked Grebe Podiceps grisegena Slavonian Grebe Podiceps auritus Cormorant Phalacrocorax carbo Shag Phalacrocorax aristotelis Fulmar Fulmarus glacialis Mute Swan Cygnus olor Mallard Anas platyrhynchos Pochard Aythya ferina Tufted Duck Aythya fuligula Scaup Aythya marila Eider Somateria mollissima Steller’s Eider Polysticta stelleri Long-tailed Duck Clangula hyemalis Common Scoter Melanitta nigra Velvet Scoter Melanitta fusca Goldeneye Bucephala clangula Smew Mergus albellus Red-breasted Merganser Mergus serrator Goosander Mergus merganser Coot Fulica atra Great Skua Catharacta skua Little Gull Larus minutus Common Gull Larus canus Herring Gull Larus argentatus Great Black-backed Gull Larus marinus Kittiwake Rissa tridactyla Guillemot Uria aalge Guillemot Uria aalge (Baltic form) Razorbill Alca torda Black Guillemot Cepphus grylle Black Guillemot Cepphus grylle (Baltic form) Little Auk Alle alle Puffin Fratercula arctica Baltic Sea North Sea 56 700 49 000 <50 11 300 5500 1800 19 400 <2500 <150 000 108 000 227 000 31 200 319 000 145 700 1 048 000 6850 4 272 000 783 000 933 000 123 000 17 000 44 000 73 000 212 000 <280 2250 72 000 310 000 21 000 76 000 66 000 20 000 156 000 1250 27 000 900 14 000 2000 <50 14 000 29 000 1872 000 <1800 <50 000 <3500 <7500 14 000 463 000 <150 <47 000 570 000 121 000 16 000 <250 9850 3200 <15 000 1000 5400 176 000 918 000 300 000 1 034 000 1 562 000 <500 324 000 6600 <500 <3000 <14 000 853 000 75 000 Estimates are only mentioned for species occurring in the relevant region in numbers exceeding 1% of the reference biogeographic population. Please cite this article in press as: Skov, H. et al., A quantitative method for evaluating the importance ..., Biol. Conserv. (2007), doi:10.1016/j.biocon.2006.12.016 ARTICLE IN PRESS B I O L O G I C A L C O N S E RVAT I O N number of birds of the particular species in the studied area. Population estimates for waterbirds inhabiting coastal environments have recently been reviewed (Wetlands International, 2002), while those for pelagic species of seabirds vary from estimates with confidence limits to educated guesses. Estimates of numbers of birds offshore and their distribution are now becoming available through large-scale surveys using line transect techniques (Tasker et al., 1984; Webb and Durinck, 1992). Due to the wide range of marine habitats occupied by birds it is frequently necessary to merge data from multiple sources (e.g. counts from land, ships, aeroplanes) in order to obtain a complete description of birds using an area based on unbiased population estimates (e.g. Skov et al., 1995). 2. Materials and methods 2.1. Study regions and survey data This paper is based on European work, but the concept should be applicable to all offshore marine areas including oceanic and shelf seas, coastal zones and archipelagos. The methodology has gradually been developed and tested during a large number of projects from feasibility studies in relation to marine conservation areas within the European Union (Durinck et al., 1994), to analyses of marine important bird areas (Skov et al., 1995, 2000) to designation of potential EU Special Protection Areas (SPAs, Johnston et al., 2002; ICES, 2006). In this paper, the selection procedures used in Skov et al. (1995, 2000) have been refined and updated in order to strengthen their general application. The revised selection procedures have been tested on an international seabird database (ESAS, European Seabirds at Sea database) containing winter distributions of waterbirds for the Baltic Sea and year-round seabird distributions for the North Sea at large (Stone et al., x x x ( 2 0 0 7 ) x x x –x x x 3 1995). Although the surveys contained in the ESAS database were not specifically designed for this analysis, the use of it made it possible to test the methods on a large, basin-wide database that includes seabird densities in virtually all types of habitats, including all coastal environments. The ESAS data used here were collected by many observers between 1987 and 1995 (1993 in the Baltic Sea) using standard methods (Tasker et al., 1984). Although more recent data have been collected in the two regions, comprehensive coverage of both regions was only achieved during the described period. All data collected under relatively calm conditions (Beaufort sea state 6 4) were processed to describe seabird distributions in the Baltic Sea during winter and in the North Sea during the whole year. Approximately 150,000 km of line transect observations is represented in the data used for analysis. In addition to offshore line transect data, information was obtained from national databases on birds in near-shore waters during the same periods. These data mainly cover observations made from aerial or land-based surveys in connection with the internationally co-ordinated Wetlands International Midwinter Census of waterbirds. Information on numbers of birds presumed to be associated with the breeding colonies of seabirds in the region were derived from the United Kingdom Seabird Colony Register (Lloyd et al., 1991), Grimmett and Jones (1989) and Hälterlein and Steinhardt (1993). No extrapolation from these numbers into waters surrounding the breeding colonies took place. The distributions of the 30 most common species have been analysed. Some species of seabirds are best surveyed by one type of platform (e.g. aeroplane) while the distribution of others may best be described using a combination of methods from several platforms. Thus, in order to achieve reliable figures for the entire seabird fauna all data were combined using a Geographical Information System (GIS) to produce Fig. 1 – Study regions marked by solid lines; I: the North Sea and II: the Baltic Sea. Please cite this article in press as: Skov, H. et al., A quantitative method for evaluating the importance ..., Biol. Conserv. (2007), doi:10.1016/j.biocon.2006.12.016 ARTICLE IN PRESS 4 B I O L O G I C A L C O N S E RVAT I O N Fig. 2 – Example variogram (spherical) used for description of spatial variance for black guillemot (Cepphus grylle) in the Baltic Sea. x x x ( 2 0 0 7 ) x x x –x x x integrated maps of bird numbers per unit area. The procedure implied integration of densities obtained from aerial and ship-based line transects with densities converted from total counts (land-based and aerial total counts). Densities were converted from total counts in near-shore areas by assuming a detection range of 1 km during these counts. Densities obtained during aerial and different types of ship-based line transects were standardised using correction factors for birds missed in the searched area determined from estimated detection probabilities in the software Distance (http:// www.ruwpa.st-and.ac.uk/distance/). Comparison of the total numbers of all species of seabirds occurring in the whole Baltic Sea and the North Sea was made for the winter season (December–February). During winter the total number of seabirds in the two regions is of the same order of magnitude (106), but the communities are clearly different (Table 1). The Baltic avifauna is dominated by benthic-feeding species (diving ducks), while in the North Sea rather more pelagic-feeding species are dominant (fulmar, gannet, gulls and auks). Hence, the two study areas provided a wide range of different habitats to test the criteria on. Fig. 3 – The effect of choosing different degrees of concentration reflected by density levels on the classification of areas of importance to seabirds in the North Sea (a and b) and the Baltic Sea (c and d). The four graphs show the proportion of the total estimated number of birds at sea covered by the classified areas in the two regions as determined by increasing minimum densities from 2 to 7 times the average regional density of the species in question. The species selected are species for which at least 25% of the total biogeographic population are estimated to occur in the North Sea and the Baltic Sea. In the North Sea, the season has been chosen in which the largest number of birds in the region was found. Please cite this article in press as: Skov, H. et al., A quantitative method for evaluating the importance ..., Biol. Conserv. (2007), doi:10.1016/j.biocon.2006.12.016 ARTICLE IN PRESS B I O L O G I C A L C O N S E RVAT I O N 2.2. Identification of concentrations Different species of seabirds show different degrees of clustering within their preferred habitats depending on the scale analyzed. Aggregating seabirds typically occur in few large patches or more commonly in multiple smaller patches. In this study the spatial variation of each species was analysed by applying b-spline approximation techniques (North Sea, see Skov et al., 1995 for details) and interpolations (Baltic Sea, see Skov et al., 2000 for details) using kriging to the database of integrated densities from all survey platforms, see (Fig. 1). In the case of multiple patches, our study shows that gradients are found in the density between strata marking the background density of birds and strata in which patches of birds occur frequently at short relative intervals (extended aggregations). The preferred scientific method for stratifying areas of different densities of birds is fine-scale geo-statistical analyses (kriging) and interpolation of sampled densities by application of experimental variograms (Cressie, 1991). The use of variograms allows for a model-based determination of the scale of spatial autocorrelation (variogram range), which can be used to inform about the chord length of extended aggregations. In the analysis of the Baltic Sea the range was also used to set the maximum distance of extrapolation away x x x ( 2 0 0 7 ) x x x –x x x 5 from sample points. In this way the obvious danger of using arbitrary borders, e.g. by expanding an area until numbers within it reach 1%, was avoided. Further, the use of variograms as a basis for kriging made it possible to determine nugget effects and the orientation of anisotropies in the data. For most of the species of conservation concern (i.e. divers, grebes, seaducks and auks) radial and spherical variograms fitted best to the sampled data, and strong anisotropies were apparent. A sample variogram is depicted in Fig. 2. 2.3. Identification of important areas per species – the Marine Classification Criterion (MCC) Geo-statistical analyses of the distribution of a seabird species can be coupled with two further parameters to determine the importance of an area holding an aggregation of that species: the percentage of the bio-geographic population of the bird living within the area and the degree of concentration displayed by the aggregation. Due to characteristic pattern of dispersal of many of the key marine bird species in the two sea regions (species with more than 25% of the bio-geographic population occurring in either region), only a small number of areas can be considered as of high importance to seabirds and thus the target of conservation efforts. In the Fig. 4 – Sketch of the GIS-procedure for merging boundaries of selected species areas into priority multi-species polygons. The example is from Pomeranian Bay showing nine species areas meeting the MCC being merged into the multi-national priority area for conservation marked with a blue line. EEZ boundaries for Germany, Poland and Denmark are indicated. Please cite this article in press as: Skov, H. et al., A quantitative method for evaluating the importance ..., Biol. Conserv. (2007), doi:10.1016/j.biocon.2006.12.016 ARTICLE IN PRESS 6 B I O L O G I C A L C O N S E RVAT I O N North Sea, the six most important areas together account for 80% of the estimated total importance of areas to seabirds (Skov et al., 2000), and in the Baltic Sea the ten most important areas, covering less than 5% of the Baltic Sea area, host about 90% of the total estimated number of wintering seabirds in the region (Durinck et al., 1994). Accordingly, the choice of the critical size of the reference basin-wide density level (4*D, Fig. 3) was determined both by the requirement to secure the inclusion of the globally important aggregation areas and by the need to exclude peripheral areas characterised by moderate densities dispersed over wide areas. The two parameters; percentage of the total bio-geographic population and the degree of concentration were combined into a Marine Classification Criterion (MCC) for selecting the most valuable sites. The percentage of the total bio-geographic population living in the area was calculated from figures provided by Rose and Scott (1994), Rose and Scott (1997). In order to be considered ‘important’, this proportion needed to be equal to or exceed one percent. The degree of concentration displayed by the aggregation should exceed four times the average basin-wide (North Sea/ Baltic Sea) density. Accordingly, the MCC can be written as MCC : ðn=NÞ P 0:01; and d > ð4 DÞ where N is the total biogeographic population, n is the estimated number of birds within the aggregation, D is the regio- x x x ( 2 0 0 7 ) x x x –x x x nal average density and d is the local density of birds within the aggregation. 2.4. Selection of priority areas across species The boundaries of the combined areas were computed by a GIS procedure which (a) merged the borders of all overlapping specific areas of international importance meeting the MCC and (b) stratified merged areas by their total value calculated as the sum of proportions of the total populations of the species meeting the MCC. The procedure for merging borders of selected species areas according to the MCC into multi-species priority areas is outlined in Fig. 4. 3. Results and discussion Extended aggregations in the North Sea and the Baltic Sea typically measured more than 500 km2 (Table 2). The total number of birds within each stratum was estimated. The amount of aggregation displayed by species of seabirds wintering in the Baltic Sea and the North Sea is indicated in Table 3. Contrary to common belief, benthivorous species like seaducks were generally not found to be more concentrated than offshore, pelagic species. Species showing a high degree of clustering were found among a wide range of groups from divers to auks and seaducks. Table 2 – The proportion (%) of selected species of wintering seabirds in the Baltic and North Seas estimated to occur within areas of different categories of size <500 km2 500–1000 km2 1000–3000 km2 3000–10 000 km2 >10 000 km2 Baltic Sea Red-/black-throated diver Great Crested Grebe Red-necked Grebe Slavonian Grebe Eider Long-tailed Duck Common Scoter Velvet Scoter Common Gull Herring Gull Great Black-backed Gull Razorbill Guillemot Black Guillemot, Baltic 42.97 86.21 13.80 7.10 1.76 1.15 20.91 47.09 1.04 1.33 1.06 21.52 0.00 44.78 11.46 0.00 31.65 55.46 0.64 31.71 6.54 2.86 10.88 0.78 1.47 0.67 34.31 22.57 42.32 13.79 17.20 0.00 8.81 7.95 5.25 47.43 7.57 4.70 12.93 46.99 0.00 5.76 3.25 0.00 37.35 37.43 80.03 45.01 67.30 2.61 40.71 19.24 46.25 23.52 30.93 17.42 0.00 0.00 0.00 0.00 8.76 4.19 0.00 0.00 39.80 73.96 38.29 7.30 34.76 9.47 North Sea Red-/black-throated diver Great Northern Diver Great Crested Grebe Shag Fulmar Gannet Morus bassanus Eider Common Scoter Common Gull Herring Gull Great Black-backed Gull Kittiwake Razorbill Guillemot Little Auk 8.59 85.23 30.17 68.99 0.50 0.00 19.11 10.16 0.11 0.00 0.00 0.21 0.00 0.26 0.03 22.31 14.77 0.00 1.13 0.00 2.09 2.60 6.18 12.61 0.00 0.00 0.86 0.00 0.56 0.04 22.89 0.00 0.00 20.09 0.13 33.90 1.09 31.23 20.55 15.57 12.99 8.15 72.23 5.32 0.09 21.31 0.00 69.83 9.79 0.67 17.54 57.31 51.99 38.29 18.25 9.08 9.95 21.50 13.87 1.52 24.90 0.00 0.00 0.00 98.70 46.47 19.89 0.43 28.44 66.18 77.94 80.83 6.27 79.99 98.32 Species Please cite this article in press as: Skov, H. et al., A quantitative method for evaluating the importance ..., Biol. Conserv. (2007), doi:10.1016/j.biocon.2006.12.016 ARTICLE IN PRESS B I O L O G I C A L C O N S E RVAT I O N Table 3 – The table shows the intensity of aggregation among species of seabirds distributed in the Baltic Sea and the North Sea during winter Species Intensity of aggregation Baltic Sea Red-/black-throated diver Razorbill Cormorant Common Scoter Long-tailed Duck Scaup Smew Steller’s Eider Guillemot Velvet Scoter Tufted Duck Mute Swan Pochard Coot Black Guillemot Eider Mallard Great Crested Grebe Red-breasted Merganser Goosander Herring Gull Goldeneye Common Gull Red-necked Grebe Little Gull Great Black-backed Gull Slavonian Grebe 8.15 12.14 17.29 18.12 18.35 21.97 24.38 25.00 25.06 28.30 29.89 30.49 31.42 32.84 35.90 37.32 47.59 51.54 54.62 56.19 56.27 62.71 73.44 83.81 84.43 84.79 92.83 North Sea Razorbill Little Auk Velvet Scoter Common Scoter Fulmar Goldeneye Cormorant Kittiwake Red-breasted Merganser Herring Gull Scaup Shag Guillemot Great Crested Grebe Gannet Eider Great Black-backed Gull Common Gull Red-/black-throated diver 9.5 12.7 19.4 23.7 30.5 31.4 51.2 52.0 58.8 59.5 59.5 63.0 66.6 71.8 72.8 74.2 76.5 76.5 83.7 The intensity of aggregation is calculated as the proportion (%) of the area of the sea surface inhabited by the 75& of the total number of the species compared to the area of the sea surface which embraces the total range of the species in the two regions. In other words the measure of the intensity of aggregation increases as the part of the range occupied by 75% of the population increases. The species are ranked by (descending) degree of concentration. In Fig. 3 it can be seen that the selection of key areas to many of the globally important species with their main occurrence in offshore waters (red-/black-throated diver, long- x x x ( 2 0 0 7 ) x x x –x x x 7 tailed duck in the Baltic Sea; great skua and common guillemot in the North Sea) are sensitive towards the application of higher reference density levels than four times the regional density. During the selection process it was further evident that the 20,000 birds criterion (Ramsar Convention criterion 3a) is generally inapplicable in marine habitats, as any larger unit of area within our study regions proved to meet the criterion at any one season! Our study in the North Sea and the Baltic Sea showed that most areas are of value to more than one species. The ranking of selected marine areas in the North Sea by total conservation value is shown in Fig. 5a and for the Baltic in Fig. 5b. The classified marine areas in the Baltic Sea and North Sea cover large proportions (30% and 34% respectively, Fig. 5a and b of the studied regions. Yet, of the 9.19 million seabirds found wintering in the Baltic Sea the selected areas cover 8.40 million birds, – equivalent to 91.4%! In the Baltic Sea, the ten most important areas covered only 5% of the region. For the species with clustered distribution of high regional importance found wintering in the Baltic Sea more than 70% of the estimated total number of individuals occur within the proposed areas, and in the North Sea more than 40–80% of key species occur within the proposed areas. Another striking general feature about the selected areas is that for the majority of species, more than half of the cumulated sum of proportions is held within the top four (Baltic Sea) or five (North Sea) areas. In addition, more than 75% of the cumulative sum of proportions is held within the top ten areas in both regions. Thus, the case studies indicate that the areas selected on the basis of the Marine Classification Criterion possess large conservation potentials. This is corroborated by Hägerhäll and Skov (1998) who showed that a satisfactory level of protection of many seabird species in the Baltic Sea may be accomplished by conserving a smaller part of the classified areas which overlaps coherent concentrations of other predators or other biota. It should be noted however, that our analyses refer to complete basins and do not take national jurisdictions into account. Obviously, within an international sea, important seabird areas may not be distributed equally over the respective waters of the different countries around that sea. Although from a biological perspective, selecting the most important areas within a whole basis, or even within larger, coherent biogeographical regions is clearly defendable, this approach is currently not followed by the EU Bird Directive, which only considers national jurisdictions. Our analysis did not include assessments of the fine-scale temporal variability of selected areas or the distributions of individual species. Although not a requirement for selection of potential areas for conservation of seabirds, analysis of the variability of the number of birds related to the selected areas between seasons and years would form an important prerequisite for drafting detailed management plans for these areas. In spite of the fact that the suggested Marine Classification Criterion (MCC) is a modified version of the MCC criteria implemented by Skov et al. (1995, 2000) the differences in the resulting selection of areas are minor. Most areas selected using the early version of the MCC were retained during this analysis, and the boundaries were only modified slightly as an effect of adding or deleting the boundary of a few concentra- Please cite this article in press as: Skov, H. et al., A quantitative method for evaluating the importance ..., Biol. Conserv. (2007), doi:10.1016/j.biocon.2006.12.016 ARTICLE IN PRESS 8 B I O L O G I C A L C O N S E RVAT I O N x x x ( 2 0 0 7 ) x x x –x x x Fig. 5 – The location and extent of areas of international importance for seabirds selected on the basis of the MCC in (a) the North Sea and (b) the Baltic Sea. The numbers refer to the priority ranking of the areas based on the sum of proportions of total populations of seabirds supported. Please cite this article in press as: Skov, H. et al., A quantitative method for evaluating the importance ..., Biol. Conserv. (2007), doi:10.1016/j.biocon.2006.12.016 ARTICLE IN PRESS B I O L O G I C A L C O N S E RVAT I O N tions of moderate importance. As can be seen in Fig. 3, adjustments of the acceptance criteria for defining a concentration of seabirds (2–7*D) have only a limited impact on the selection of the top ten areas to seabirds in both study regions. This reflects the fact that the gradients in area importance and thus in seabird densities play a significant role in shaping the boundaries of the selected areas. It should be underlined, however, that picking a value of four times the average density has been based on practical site selection processes rather than scientific approaches. Hence, this element of the MCC follows the history of selection of wetlands of international importance, in which an arbitrary 1% criterion has proven a practical tool in wetland management. The MCC was developed in view of the protected areas systems and seabird databases available or potentially available for shallow shelf seas, like the North Sea and the Baltic Sea. Thus, although the MCC is applicable for similar environments provided seabird line transect data are available, for most marine areas, and certainly the deeper parts of the ocean outside the shelf seas the MCC needs to be supplemented by techniques like habitat modeling based on tracking data. With the recent development of spatial prediction modeling methods and the increasing electronic access to vast sets of oceanographic and occurrence records from satellite tracking and other seabird telemetry studies, as illustrated by the Birdlife International’s ‘‘Ocean Wanderers’’ project (BirdLife International, 2004), the potential for achieving reliable estimates of gradients in some seabird distributions for vast expanses of ocean is rapidly increasing. Novel habitat modeling methods, including both data-driven and machine-learning methods, have the capacity to use presence-only records as tracking data and clearly outperform more established statistical methods for eco-geographical predictions (Elith et al., 2006). Thus, an application of the MCC in ocean areas might be developed for tracking data using statistics on time budgets rather than bird densities. In 1990, BirdLife International launched the Dispersed Species Project with the aim of developing wide-scale habitat conservation measures for European bird species that are in need of conservation action. Seabird species of conservation concern were identified by Tucker and Heath (1994) and strategies for the integration of habitat conservation and management have been drawn up for these species (Tasker and Canova, 1997). Ours and other studies show that although seabirds have large or very large ranges of occurrence at sea, only a minority of species show truly dispersed distributions throughout their range, at least in shelf seas. The size of aggregations and the difficulty of enforcement in offshore waters, however entail that rather than establishing networks of strict reserves, a realistic goal would be to develop networks of integrated management zones. Acknowledgements A significant part of the field work on which the initial MCC tests were based was carried out within the frames of the EPAS (Establishment of a Protected Areas network at Sea) project, funded through the European Union Directorate General XI, ACE Contract No. 2242/90-09-011994. Additional funding came from Netherlands Institute for Sea Research, the Nordic x x x ( 2 0 0 7 ) x x x –x x x 9 Council of Ministers and the National Environmental Research Institute, Denmark. The study on the ESAS database was funded by BirdLife International. The EPAS surveys were undertaken by Ornis Consult A/S, Netherlands Institute for Sea Research and the National Environmental Research Institute, Denmark, in collaboration with the following coordinators of seabird surveys in the Baltic and North Seas: Leif Nilsson and Per Andell, University of Lund, Sweden, Martti Hario, Finnish Game and Fisheries Research Institute, Finland, Andres Kuresoo, Institute of Zoology and Botany, Estonia, Patrick Meire, Belgium Institute of Nature Conservation, Antra Stipniece, Institute of Biology, Latvia, Saulius Svazas, Institute of Ecology, Lithuania, Wlodzimiers Meissner, University of Gdansk, Poland, Hans Wolfgang Nehls, Rostock Zoological Garden, Germany and Jan Meissner, University of Kiel, Germany. We greatly acknowledge their help and support during the whole project. The surveys would not have been possible without the support from hundreds of observers participating in the International Waterfowl Census around the Baltic Sea. Furthermore, a number of ornithologists from most Baltic and southern North Sea countries took part in the air and ship surveys. We thank them all for their great effort during the long and often cold surveys. We also acknowledge the assistance and support from Wetlands International. The North Sea data were kindly supplied by the European Seabirds at Sea database (ESAS) Danish National Environmental Research Institute, The Dutch National Institute for Coastal and Marine Management, Norwegian Institute for Nature Research, University of Lund, Sweden, Institut für Vogelforschung, Germany, Free University Brussels, Belgium, Institute for Nature Conservation, Belgium, Netherlands Institute for Sea Research, Joint Nature Conservation Committee, Central Institute for Waterfowl Research and Wetland Protection, Germany, Ligue pour la Protection des Oiseaux, France, Office National de la Chasse, France, The Wildfowl & Wetlands Trust, United Kingdom, Samenwerkende Organisaties Vogelonderzoek Nederland (SOVON), The Netherlands and Institut für Meereskunde an der Universität Kiel, Germany. An early version of the manuscript was improved by Professor Jon Fjeldså, Zoological Museum, Copenhagen and Ole Norden Andersen, Danish Forest and Nature Agency. 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