marine research, resources and conservation in the azores

AQUATIC CONSERVATION: MARINE AND FRESHWATER ECOSYSTEMS, VOL. 5 , 31 1-354 (1995)

CASE STUDIES AND REVIEWS

Marine research, resources and conservation in the Azores

RICARDO SERRAO SANTOS Universidade dos &ores, Departamento de Oceanografia e Pescas, PT-9900 Horta (&ores), Portugal

STEPHEN HAWKINS University of Southampton. Centre for Environmental Sciences, Shackleton Building, Highfield, Southampton

SO17 IBJ, UK and

Universidade dos &ores, Departamento de Oceanografia e Pescas, PT-9900 Horta (AJores) Portugal LUIS ROCHA MONTEIRO, MARIO ALVES, EDUARDO JOSE ISIDRO

ABSTRACT

1. A history of marine research on the Azores is outlined. Until two decades ago most oceanic and littoral marine studies had been carried out by foreign scientists. Studies by Azorean scientists started to increase in the early 1980s when the University of the Azores was created.

2. Ocean circulation in this part of the Atlantic is described as a background for biogeography and diversity. The picture emerges that Azores is a 'meeting point' for shallow water marine fauna and flora of different origins.

3. The species composition of one of the best studied groups of organisms, the fish, is compared between locations in the northeastern Atlantic. The work on fish is also compared with other well studied groups (algae and hydroids) to highlight the interest of the Azores as a natural biogeographical experiment.

4. Studies of marine resources began less than two decades ago. The development of demersal fisheries is described focusing on the switch from small-scale artisanal fishing to more commercial fisheries.

5. Conservation of species and legislation in force for molluscs, crustaceans, fishes, marine turtles, seabirds and marine mammals are summarized.

6. Protected marine areas already designated are defined, as well as new areas recommended. 7. The paper concludes with a discussion of current threats and future management strategies.

INTRODUCTION

The Archipelago of the Azores consists of nine volcanic islands and several small islets, forming three groups along a tectonic zone running WNW-ESE between 37" and W N latitude, 25" and 32"W longitude, in the middle of the Atlantic (Figure 1). The oldest rocks on Santa Maria have been dated to the Miocene, c. 10 to 8 million years (Abdel-Monem et al., 1975; FCraud et al., 1980; see also Wilson, 1963; Nunn, 1994). Since then the islands have been in continuous formation as the result of the volcanic activity of the Mid- Atlantic Ridge. Thus each island is a mosaic of rocks of different ages. The youngest rocks are less than 40 years old (Capelinhos on Faial island) with most islands being between 3 and < 1 million years (Azevedo et al., 1991). There are also various seamounts including subsided islands (Ryall et al., 1983; Lambeck, 1984).

The Azores were uninhabited until colonized by the Portuguese in the 15th Century. On land many endemic species of plants suffered from the human influence-particularly agriculture and the introduction of exotic plants (Sjogren, 1973, 1991). The marine environment of the Azorean Archipelago and its

CCC 1052-7613/95/040311-44 01995 by John Wiley & Sons, Ltd

Received 21 September 1994 Accepted 22 September 1995

312 R. S . SANTOS ET AL.

e 2 3P,

0 @-- TWCQUA

- 0

Figure 1 . Archipelago of the Azores and surrounding area, with indication of the islands, main islets and seamounts. The lines represent the 200111 depth contour.

surrounding Economic Exclusion Zone (EEZ), of more than 1 million square kilometres, is of considerable conservation and marine biological interest-in large part because of its isolated position in the middle of the northeastern Atlantic and the recent age of the Archipelago.

Man has exploited littoral, nearshore and offshore living resources since the earliest colonization (Frutuoso, XV century--printed as Frutuoso, 1983); Ramos, 1869; Serpa, 1886; Sampaio, 1904). In recent years pressures on littoral and offshore resources have grown (see Martins et al., 1987; Santos et al., 1990; Menezes, 1991a; Isidro, H., 1988; Silva and Krug, 1992; Silva et al., 1994) with the switch from essentially subsistence or artisanal exploitation to more commercial operations (see below). Meanwhile the cessation of commercial whaling and greater environmental awareness, both at an international, national and regional level, have increased the pressure to protect marine wildlife.

In this review we first give a short account of the history of marine research in the Azores; this account also serves as an ‘entrke’ to the scattered literature on the Azores including an update of papers published since a recent comprehensive bibliography (Martins, 1990). The oceanographic conditions of the Azores are summarized to set the scene. The species composition of one of the best studied groups of organisms-the fish-are then compared between locations in the northeastern Atlantic. The work on fish is then compared with other well studied groups (algae and hydroids) to highlight the interest of the Azores as a natural biogeographical experiment. The development of demersal fisheries is outlined focusing on the switch from small-scale artisanal fishing to more commercial fisheries. Current legislation to protect marine wildlife, marine living resources and habitats of interest is then summarized. Existing and proposed marine protected areas are listed and briefly described. Finally, the few pollution problems are considered.

The review concludes with a discussion of future work, current threats and future management strategies. It is hoped that this review will attract attention to the Azores, emphasize their importance in the establishment of a network of protected sites in the North Atlantic and highlight the need for conserving

MARINE RESEARCH IN AZORES 313

selected species within the Azorean EEZ. Although this theme has also been briefly covered in recent years (Martins and Santos, 1991; Santos, 1992; Depledge et al., 1992) there has not been a comprehensive overview of the various aspects considered here. Throughout this article various reports and other items of ‘grey’ literature relevant to marine conservation and living resource management are cited. These are available through the library of the Departamento de Oceanografia e Pescas, Universidade dos Agores.

HISTORY OF RESEARCH ON THE MARINE ENVIRONMENT

A selected bibliography of the marine fauna and flora of the Azores has recently been published by Martins (1 990). An ornithological bibliography for the Azores, up to 1979, was presented by Le Grand (1 983). For the marine mammals, bibliographies are included in Galhardo (1990), Reiner (1990) and Reiner et al. (1993). Work on turtles has been summarized by Carr (1986), Eckert and Martins (1989), Bolten and Martins (1990) and Bolten et al. (1992, 1993). An outline of the major fisheries is given in Silva (1992) and Silva et al. (1994), with studies of selected major species by various authors (tuna: see Pereira, 1994 for a general review; squid: Martins, 1982; Porteiro, 1992; Porteiro and Martins, 1994; octopus: Gongalves, 1991 b; slipper lobster: Martins, 1985a; blackspot seabream: Krug, 1989, 1990; bluemouth rockfish: Isidro, E., 1987a,b; ocean blue horse mackerel: Isidro, H., 1990).

The list of Martins (1990) shows the evolution of littoral marine research in the Azores, particularly within the last 70 years. The end of the 19th and the beginning of the 20th century were particularly productive periods concerning marine research in the Azores, especially due to the cruises of Challenger and visits by the yachts of the Prince of Monaco, L’Hirondelle and Princesse Alice (see e.g. Monaco, 1905; Richard, 1910). An abbreviated account of the early work under the auspices of the House of Grimaldi is given by several authors (Bethoux, 1992; Carpine-Lancre, 1992; Fontaine, 1992; Klaveren, 1992; Saldanha, 1992). The results of the different expeditions were compiled in several articles which were published in various issues of the Bulletin du M w t e Octanographique du Monaco and in different volumes of the Rtsultats des Campagnes Scientifiques.

Also important were the oceanographic cruises in the eastern Atlantic by the French ships Travailleur and Talisman between 1880 and 1883, and the Norwegian steamer Michael Sars in 1910 (Koefoed, 1932). The results were compiled by Vaillant (1888) and by Murray and Hjort (1912). The importance of the deep- sea fishes from the region surrounding the Azores is also emphasized by Goode and Bean (1895) in their classical work on oceanic ichthyology. The Azores were also visited in 1903 by the German South-Polar Expedition. The fish species collected were listed by Lampe (1914). Important material was also collected in the region as part of the scientific cruises around the world of the Royal Danish research ship Dana, which are described in a large series of reports and monographs (for an introduction see Dana Report No. 1,1934).

Several Azoreans made original contributions to marine research in the Archipelago, and should be considered pioneers (see Saldanha, 1992): Arruda Furtado (Furtado, 1887), Afonso Chaves (Chaves, 1924, 1960), Ernest0 Ferreira (e.g. Ferreira, 1938, 1939, 1942) and Jos6 Agostinho (Agostinho, 1935, 1954). The Azores were also the subject of attention from other Portuguese from the mainland (Capello, 1871a,b; Guimariies, 1882, 1882; Nobre, 1924, 1930) and other workers (Drouet, 1858, 1861; Godman, 1866; Barrois, 1888; Hilgendorf, 1888; Dautzenberg, 1889; Collett, 1897, 1905; Piccone, 1889; Chavigny and Mayaud, 1932; Collins, 1954) during the 19th and early 20th centuries.

During the years of the Second World War very few papers were published (Martins, 1990). After the war there was an increase in activity due to some littoral expeditions organized by both the British, and also the Portuguese from the Faculty of Sciences (Lisbon).

The 1980s saw an upsurge in scientific research in the Azores. With the creation of the University of the Azores in 1976 scientists, based on the islands, initiated research on different aspects of the natural history of the Archipelago. The University also became a centre of attraction for scientific cooperation with other marine research institutions, both national and foreign.

314 R. S. SANTOS ET AL.

Since Martins' (1 990) comprehensive bibliography there has been a number of papers concerning the Azores. Many have resulted directly or indirectly from an Anglo-Portuguese Expedition in 1989 (Martins et al., 1992) jointly organized by the Department of Oceanography and Fisheries (University of the Azores) and the Port Erin Marine Laboratory (University of Liverpool) (Expediqiio Aqores '89, 1990). These include papers on fish (Patzner et al., 1992), sponges (Moss, 1992), decapods (Paula et al., 1992), hydroids (Cornelius, 1992a), seaweeds (Tittley and Neto, 1994) plus various papers on ecology and behaviour of nearshore fish (e.g. Nash ef al., 1991, 1994; Patzner and Santos, 1993; Nash and Santos, 1993; Santos et a[., 1994; Santos and Nash, 1995). Other work has stemmed from multidisciplinary expeditions to different islands organized by the Department of Biology of the University of the Azores (see the following reports: Graciosa '88, 1989, Flores '89, 1990, Santa Maria e Formigas '90, 1991 and Pic0 '91, 1992) and from two workshops organized in S2o Miguel by the Sociedade Afonso C h a w (see Martins, A. F. 1990, 1995). These include papers on zonation of the littoral (Hawkins et al., 1990a), various aspects of malacology (e.g. Gosliner, 1990; Houbrick, 1990; Morton, 1995; Backeljau et al., 1995; Bieler, 1995; Bullock, 1995; Jensen, 1995; Knudsen, 1995; Mikkelsen, 1995), crustacea (Moore, 1995; Moore and Weeks, 1995); ecology (Morton, 1990; Wells, 1995), and physiology and/or pollution (e.g. Moore et al., 1995; Vedel and Depledge, 1995; Weeks et al., 1995).

Interesting papers, including bathyscaphe studies in the Azores in the 1960s (Ptrks, 1992), are presented in a retrospective volume celebrating the centenary of the last oceanographic cruise by Prince Albert of Monaco (Saldanha et al., 1992). Other recent works on the systematics and ecology of the Azores include studies of intertidal algae (e.g. Neto, 1992, 1994) and algae collected during a recent expedition using the Sea Diver submersible (Reed, 1991; Fredericq et al., 1992), decapods (Fransen, 1991; Martins and Hargreaves, 1991), amphipods (Lopes et al., 1993), various little known invertebrates (Menezes, 1991b; Wirtz and Martins, 1993), octopods (Gonqalves, 1991a), inshore fish (e.g. Santos, 1995; Santos et al., 1995) and birds (Hamer et al., 1989; Monteiro et al., in press).

OCEAN CIRCULATION IN THE CENTRAL NORTHEASTERN ATLANTIC

The basic average ocean circulation pattern of the North Atlantic is an asymmetric, large scale gyre that flows to the north, on the western side, with an intense thin jet (the Gulf Stream) and to the south, on the centre/eastern side, with a multibranched current system. The Gulf Stream very efficiently transports warm water of equatorial and tropical origin into the colder northern waters. The current patterns result in the high salinity, high temperature and low nutrient regime which typifies the Azores (Figure 2: salinity, Figure 3: temperature and Figure 4: oxygen and nutrients). pH is between 8.1 and 8.2 and dissolved silica is around 10pg-atomsL-I. During winter a deep mixed layer is present around 150m and in summer a seasonal thermocline develops around 40 to 100 m.

The Gulf Stream is also the source for many instability processes, meanders and eddies. This picture becomes particularly complicated when this current leaves the North American coast, at about 40" to 45"N, towards the central zone of the North Atlantic where the Azores are located (Crease et al., 1985; Gould, 1985; Kleine and Siedler, 1989).

The following consideration is based on some of the general ocean circulation patterns that have been calculated recently (Alves, 1990; Juliano, 1994) for the Central/Eastern North Atlantic, using all the hydrological data (temperature, salinity versus depth) available from National Oceanographic Data Center. Figures 5A and 5B represent the oceanic circulation in the area between 20"N to SOON and 0"W to 50"W (Azores inside the circle) at about lOOm depth and for two different average year seasons (respectively summer and winter). The averages have been derived for the 42 years between 1947 and 1988. Computation of ocean currents was obtained by the Beta-Spiral method (Stommel and Schott, 1977).

Even when considering the mean for the summer months (July, August, September) over 42 years (Figure SA), the complexity of the current system that surrounds the Azores is remarkable. This complex picture


LONClTWE W4

Figure 2. Ocean salinity (USP-"/co) in the north-eastern Atlantic (2Oo-5O0N and Oo-60"W) at Sm, lOOm and lOOOm deep in the winter ( A X ) and during the summer (D-F). Average values for the last 42 years (based on National Oceanographic Data Center-

NODC-data bases).

helps to explain what is observed locally in the Azores, in the region delimited by the circle on the chart (Figure 5A). The Gulf Stream (GS) can easily be identified entering the domain through the western boundary at about 40"N and then splitting into two main branches, the North Atlantic Current (NAC) and the Azores Current (AC). Furthermore each of these also divides into a further two branches (NACl and NAC2 and ACl and AC2). Clearly, at this time of the year, the northern part of Azores is partially


Figure 3. Ocean temperature ("C) in the noth-eastern Atlantic (20"-50"N and 0"-60°W) at Sm, lOOm and l000m deep in the winter (A-C] and during the summer (D-F). Average values for the last 42 years (based on NODC data bases).

influenced by the NAC2 system while the southern one is influenced by the ACl. It is also clear that the Canaries current (CC) is a continuation of the Madeira current (MC) and both are fed mainly by the ACl.

The above multibranching system is even more complicated than it appears because it changes with the time of the year. During winter for the same region (Figure 5B) climatological average conditions are very


60' H

v

02 3 0

0 60' 30' 0

Figure 4. Mean dissolved oxygen and nutrients at the surface, in July-September, in the north-eastern Atlantic, during the months of July to September. (A) dissolved oxygen (O'mL L-I); (B) alkali-chloride coefficient (pg-atoms L-'); (C) phosphates (pg-atoms L-l);

(D) maximum dissolved phosphates at around 800 m (pg-atoms L-I). Based on Gorshlov (1985).

different. The first notable difference is the clear formation of the Southwest European Current (SWEC) that emerges from the NACZ-AC1 confluence zone present to the east of the Azores. Of note is the Madeira current-Canaries current (MC-CC) system that also originates in this area. The second major difference is the intensity of currents (size of the arrows), especially on the ACl-MC-CC system. This is apparent because during summer the maximum intensity level of ACl-MC-CC is deeper than loom, while during winter it shallows.

A similar analysis for the other seasons and even for each month average reveals that there are many changes during the average year. An analysis of temporal series obtained at a fixed point of the domain is illustrated by Figure 6A-C. The horizontal axis of these figures represents the time of the average year in Julian days and the vertical axis the north-south intensity of the current at a fixed point (north is positive and south is negative). Each straight line drawn represents the current direction (left is west and right is east) at that fixed point and at the time instant where its origin is placed, with a length proportional to their intensity.

Figure 6B corresponds to the mean year time series at the point P3 (see Figure 5B) at 36"N, 25" to the southwest of Canaries. It can easily be seen that at 100 m depth the general current pattern at this point is a


A

...... * f l f t 1 1 1 . . . ,--. * - - ..--.. e.c. 2 ....... - .......... ,'...... .... . ( . . . . ...... ,,. . . . - . . - - * .

GS

............. ,.. ........

B

GS

Figure 5 . Ocean currents on the area surrounding the Azores: (A) summer months. (B) winter months (see text for explanation). Data from NODC.

flow coming from the north to the south (the CC) with a mean intensity of approximately 5 cm SKI. There are, however, two mean-events (marked A and B) in which current is clearly reversed and is flowing to the northwest-that is, from Africa towards the Azores. Similarly, Figure 6A shows the mean year time series at the point 34"N, 20"W, northwest of Madeira (point P2 on Figure 7A). Here it can be seen that AC1 dominates the general flow pattern coming in general from the west to the east, except for the mean events A and B, where the current is mainly from east to west (from Madeira towards Azores).

Finally, Figure 6C represents the mean year time series in a point well inside the NACZACI confluence zone east of Azores at 38"N, 22"W (point P1 on Figure 5A). The general regime is from the west to the east


A P2

NORlHWEST PIADEIRA

A *----*

LATITUOE: 34 N LONGITUDE, 20 W

c,

25 50 75 100 I25 150 175 200 295 250 275 300 375 350 375 4 NUflBER OF CAYS AVERAGE YEAR1

w g 10 SOUTHWEST CANARIAS 3 u

-

LATITUDE: 26 N A LONGIlUOE. 25 W

e--, B - 20kl ;5 ;0 ;5 A 115 I& 4;s 2L 245 2c, 215 4, 3;5 3L 3j5

NUYBER OF DAYS t AVERAGE YEAR)

EASl AZORES

LATITUDE: 3 8 N LONGITUOE. 22 U

25 50 75 ie0 125 1% 175 206 225 250 275 386 325 350 375 400 NUMBER OF DAYS I AVERAGE YEAR)

Figure 6 . Orientation and strength of water currents in the region of the Azores (see text for explanations). Data from NODC.

but there is a clear seasonal and half seasonal oscillation of the mean direction, with periods where NAC2 dominates (current coming from northwest) and periods where ACl is present (current coming from southwest).

This complex current system is the major influence on the hydrography and climate of the Azores. In particular, the large scale oceanic circulation in the Azores is dominated by the Azores Current flowing to


Linkage distance P I

o z - in N u 3

South Africa

Gulf of Guinea

Cape Verde

Mauritania

Morocco

Mediterranean

Portugal

Gulf of Biscay

Canaries

Madeira

Azores

I

3 Figure 7. Dendogram of marine icthyogeographic affinities of Atlantic regions and the Mediterranean. The analysis was based on 20

families of littoral fishes.

the east. This means that the subtropical thermohaline front, a major source of meanders and eddies, stays very near the islands. It generates considerable mesoscale variablity with direct consequences for the whole regional ecosystem.

Despite the fact that dominant average ocean current circulation reaches the Azores from the west, marine littoral flora and fauna have affinities with the eastern coasts of the Atlantic. This is probably due to the great distance separating the Azores from American coasts, making colonization from that side particularly difficult, especially with regard to fauna. An advective current trip between the point where GS leaves the American coast and the Azores would take more than three months, at an average speed of 30cms-' and following the calculated trajectory (Figure 5) . Another reason is that the littoral hydrological conditions found on the North American coast are very different (the northern side of the Gulf Stream is mainly of Labrador/Sub-Arctic origin) from those found at the Azores (which is predominantly sub-tropical). Thus, even if colonizers did arrive, it would be very unlikely for them to survive. This point highlights the importance of the episodic anomalies (Figure 6A and B, periods A and B) that are found in the general pattern of water movements described above (Figure 5) . These anomalies certainly indicate that there were significant time periods between, 1947 and 1988 (mainly in winter and autumn) when colonization, from east to west by the current systems from Europe or North Africa to the Azores, was possible. Larvae or algal propagules could follow those routes, thus reaching and colonizing the Azores. We believe that similar current anomalies in the past could have been the routes for


‘colonizing’ species from those regions. Furthermore, colonization by many species probably took place under very different current patterns than today, particularly after the Pleistocene (see below). Subsequent rare events would be sufficient to maintain gene flow and prevent or reduce the likelihood of allopatric speciation events.

BIOGEOGRAPHY AND DIVERSITY

Marine littoral fish: a case study of colonization, diversity and evolution

The fish fauna provides a discrete and reasonably well studied group to allow comparisons of biogeographical affinities and hence speculation on colonization, diversity and evolution. This speculation can then be broadened by reference to other groups, particularly seaweeds (see Prud’homme van Reine, 1988; Fredericq et al., 1992; Neto, 1994) and hydroids (Rees and White, 1966; Cornelius, 1992a), that have received recent attention.

Old references to fishes from the Azores (Hilgendorf, 1888; Collett, 1897; Regan, 1903; Collins, 1954) have been supplemented by a few recent papers on littoral fishes observed and collected around the islands (e.g. Arruda et al., 1992; Patmer et al., 1992; Azevedo et al., 1995). A complete provisional list was prepared for the third volume of the Red Book of Portuguese Vertebrates (Livro Vermelho 111, 1993). This checklist is part of a compilation of the marine fishes of the Azores (Santos et al., in press).

The area of the Azores has also been included recently in the Fishes of the North-Atlantic and the Mediterranean (Whitehead et al., 1984-1986), but several, even very common species, are not referred to, while others that do not occur in the Azores are cited for this Archipelago. Unfortunately these data have been used in recent studies of Macaronesia ichthyogeography. For example Lloris et al. (1991) do not include in their database between 145 and 152 species that occur in the Azores (33% to 37%) and cite around 53 species as present that do not occur here.

A preliminary ichthyogeographic general characterization of the Azores is attempted in this paper. The database of this study includes only 20 families of littoral species.

Colonization of the islands by littoral organisms is particularly difficult due to their isolation and the lack of extensive margins of shallow water. The greater distance from continental coasts is certainly the main factor contributing to the low number of shore fishes in the Azores, compared with Madeira (Livro Vermelho 111, 1993) and the Canaries (Brito, 1991). Dispersion and survival are particularly difficult for species with benthic and short-living planktonic stages and weak powers of locomotion when adults.

Preliminary cluster analysis (Figure 7), using our database, shows that ichthyogeographic affinities of the Azores are with Madeira and Canaries, forming a primary cluster. This joins a second cluster formed by Morocco, Mauritania and the Gulf of Guinea. A third cluster is formed by Portugal and the Mediterranean and a fourth by the Gulf of Biscay and the British Isles.

Dissimilarities between Madeira and the Canaries and between the more northernly Bay of Biscay and the British Isles are stronger than between the Azors and those two regions. This emphasizes the temperate affinities of the Azores. In contrast the Canaries are much more like the tropical African coast (Gulf of Guinea, Mauritania) than the Azores and Madeira. There are great dissimilarities between the Azores and the Mediterranean and the Gulf of Guinea.

The results are, however, preliminary since the data set includes only 397 species and needs to be reconfirmed and corrected for some regions. The degree of confidence is high for the British Isles (Wheeler, 1992), the Azores (Livro Vermelho 111, 1993; Santos et al., in press), Madeira (Livro Vermelho 111, 1993), the Canaries (Brito, 1991), continental Europe (Whitehead et al., 1984-1 986) and the Mediterranean (Whitehead et af. , 1984-1986). Confidence decreases concerning the check-lists of fishes from Africa, the best known work being Qutro et af . (1990). The Azores show the poorest representation, with only 61 species listed, while the Mediterranean has the greater representation, with 176 records of the 397 species

322 R. S . SANTOS ET A L .

considered. The primary ichthyogeographic affinity of the Azores with the Mediterranean, that was emphasized by Briggs (1970, 1974), has not been confirmed.

Interestingly, endemic marine fishes are almost absent in the Azores (Scorpaena azorica Eschmeyer, in a notoriously difficult group for taxonomy, is the only species considered so until now-Eschmeyer (1 969), and only the holotype is known). Little evolutionary divergence seems to be a common characteristic of the shore fish fauna of the North Atlantic Islands, as shown by Briggs (1966, 1970). However, in the Canaries (=0.9%, five species: Brito, 1991) the percentage of endemic fishes is higher than in the Azores (z0.0025%, possibly one species: Santos et al., in press).

Other groups

Some of the trends of the biogeographic picture described above for the fishes, are also evident for algae, according to the studies on phytogeography of the Azores by Prud’homme van Reine (1988). There is a lower number of species than in other Macaronesian Archipelagoes, and there are less endemic species. Cluster analysis shows that Madeira, Canaries and Salvage Islands form a distinct cluster, and the Euro- African Coast and Western Mediterranean a second cluster (Prud’homme van Reine, 1988). The flora of the Azores seems to be related, but not closely, to these two large clusters. Prud’homme van Reine (1988), follows Feldmann (1946) in including the seaweeds of the Azores in the Lusitano-African region. Western Mediterranean affinities, although weaker, are also present. He also makes the assumption that the eastern Atlantic species, which occur in the Azores, must have been transported, either directly from the African coast and/or from the other Macaronesian islands. However, in contrast to the fishes, the algae are capable of surviving long-range dispersal (Hoek, 1987; Prud’homme van Reine, 1988) through rafting or by transport attached to floating objects, such as ships.

Rafting and fouling are also the main methods of dispersal that could explain the colonization of the Azores by hydroids (Rees and White, 1966; Cornelius, 1992b). The major affinities of the shallow water Azorean hydroid fauna seem to be with the Caribbean and northwestern Atlantic coasts. A Mediterranean component of the hydroid fauna can also be identified (Cornelius, 1992a). Mediterranean affinities may seem stronger in another faunal group, the demospongiae (Boury-Esnault and Lopez, 1985; Weerdt, 1989). Moss (1992) found that, with the exception of one species, Cynachyrella alloclada (Uliczka), the sponge fauna consists of northeastern Atlantic and Mediterranean species. In both, demosponges and hydroids, endemisms are lacking in the Azores. From the 256 species of algae known in the Arquipelago, only 6 are endemic (Neto, 1994).

Concerning the biogeography, Rissoidae and Anabathridae (Mollusca), share with the fish predominant European/North-African and Macaronesian affinities (Gofas, 1990). In contrast, among crustaceans, Mediterranean affinities have been suggested (Wirtz and Martins, 1993). Among the Gammaridea and Caprellidae the fauna of the Azores is almost equally distant from the British, north-western French, Portuguese and Mediterranean, with a high proportion of species (48 of the 122 species recorded) which are only known for the Azores (Lopes et al., 1993).

The picture that emerges is that the Azores is at a ‘cross-roads’ where shallow marine fauna and flora of different origins meet. Geographic links of the different groups are related with many untested factors associated with rafting and dispersal capacity, length of larval stages, ability to disperse as adults, environmental conditions in the region of origin and physiological lability. Some sponges and crustaceans seem to be able to disperse from the Mediterranean in deep eddies. But this mode seems unlikely for, at least, the littoral fishes, thus explaining the lower Mediterranean affinities among these vertebrates. The waters of the Mediterranean penetrate into the region of the Azores at depths below 800 to l000m (Figures 4 and 5). The eddies circulating from western Africa and Macaronesian Islands, on one side, and from the Atlantic coasts of Europe, on the other, must be the main source of transport of eggs and fish larvae, and even young and adult fish. Small islets and shallow seamounts, which are a common topographic feature of the ocean between the Azores and the African continent, could have served as ‘stepping-stones’ for the


dispersal of organisms. It would be highly interesting to evaluate the role of the chain of seamounts in regard to dispersal of benthic fishes. This effect was evaluated by Leal and Bouchet (1991) along a seamount chain of the southern Atlantic for prosobranch gastropods.

The most outstanding detail of the global picture of the marine fauna and flora is the low number of endemic species. Reasons for the few endemisms in the Azores were first hypothesized for animals by Briggs (1 966, 1970, 1974), and could also be applied to the seaweeds (Prud’homme van Reine, 1988). The drop of sea temperatures (Emiliani, 1958; Crowley, 1981) that occurred during the Pleistocene probably resulted in mass extinctions. Most of the organisms now present would have reached the Azores in the last 17000 years-a too short period of time for species differentiation. Hence, the Azores has fewer species due to late arrival and has less endemism.

A word of caution is required as many so-called endemic fish species elsewhere, have been identified by meristic characteristics. Their status and degree of separation from other populations need to be reviewed, using genetic methods including allozyme analysis and more modern nucleic sequencing.

AZOREAN FISHERIES

Exploited fish stocks

In the Azores, most of the fishing activity falls into three main types. One fishery is directed to young blue horse mackerel and chub mackerel Trachurus picturatus (Bowdich) and Scomber japonicus (Houttuyn), performed with boats less than 12.5m long, using seine nets, dipnets and liftnets (Fernandes, 1984; Isidro, H., 1988). The second is a seasonal pole-and-line tuna fishery which begins in March/April and lasts until September/October, and is undertaken by boats between 15 and 30m long. The third is a bottom longline and handline multispecific fishery performed by boats less than 22m long. All these fisheries are interrelated, not only because there is a considerable mobility of fishermen between them, but also because tuna, longline and handline fisheries use blue horse mackerel and chub mackerel as bait. The captures of these latter species which are used as tuna bait are not reported in landing statistics (Isidro, H, 1988). Juveniles (0 + year old) of blackspot seabream (Pagellus bogaraveo) are used extensively as live bait in the tuna fisheries. This juvenile exploitation is a serious problem as this bream is economically the most important in the demersal long-line fishery (Krug and Silva, 1990).

Small scale artisanal fishing also occurs in the Azores largely on a part-time basis or as a very seasonal activity. This is also the case of the exploitation of crustaceans and molluscs, particularly the winter squid fishery directed to Loligo forbesi Streenstrup (Martins, 1982; Martins and Porteiro, 1992; Porteiro and Martins, 1992, 1994).

Biological studies on growth and reproduction of most exploited fish species from the Azores have been carried out (Silva, 1986a,b, 1987b; Isidro, E., 1987a,b; Krug, 1989, 1990; Isidro, H., 1990; Krug and Silva, 1990). A database with information on fleet and landings since 1979 has been developed. In 1988 a programme of length-frequency data sampling, covering the main fishing ports of the Azores was initiated.

Demersal fishery development

Economically, the most important fishery in the Azores, besides tuna which supports the canning industry, is the bottom longline and handline multispecific fishery. This fishery is considered in detail as it has most impact on marine communities. Besides blackspot seabream (Pagellus bogaraveo), the most significant catches consists of the bluemouth rockfish Helicolenus dacrylopterus dacryloprerus (Delaroche), the forkbeard Phycis phycis (Linnaeus) and Phycis blennoides (Brunnich) and the conger eel Conger conger ([Artedi] Linnaeus) (Silva and Krug, 1992). Longlines are mostly used by open-deck boats with lengths over


9 m and larger cabinned boats. Smaller open-deck boats usually use handlines. The longlines used are slightly different from boat to boat and are often modified seasonally, in terms of numbers and sequence of sinkers and floats, in order to change the distance of the hooks from the sea bottom. These changes in tactics allow a selection of certain species from the demersal assemblage.

Annual landings in the Azores from 1980 to 1992, are shown in Figure 8. The increase in landings since 1983 (Figure 8A) is due to the progressive arrival of new, more efficient, covered deck boats (Figure 8A-C).

I

79 80 81 82 83 8485 8687 88 89 9091 92

Years

B

O J . 79 80 81 82 83 84 85 86 87 88 89 90 91 92

Years

798081 8283 8485 86 87 88 8990

Years

Unknown

Closed deck

r_7 Open deck ~ > 9 m

0 Open deck L<=9m

79 80 81 82 83 84 85 86 87 88 89 90

Years Figure 8. (A) Yearly landings of the four most exploited demersal fish species. Data from LOTACOR data bases. (B) Number of boats that have landed the four most important demersal fish species in the island of Faial, by year and boat type. Data from Departamento de Oceanografia e PescasiUniversidade dos AGores-DOP/UA-databases. (C) Average yield/landing in the island of Faial by year and boat type using the four most important demersal species. Data from DOP/UA data bases. (D) Average first sell price in the island of Faial for the four most important species by year. Data from DOP/UA and LOTACOR databases. (E) Relative landings in the island of Faial by year and boat type using the four most important demersal species. Data from DOP/UA data bases. x,blackspot seabream;

+ ,bluemouth rockfish; A, forkbeard; 0, conger; *, open deck (length < = 9 m); 0, open deck (length > 9 m); 0, closed deck.


The expansion of fishing effort has been encouraged by better preservation facilities and expanded export markets. Both old and new areas have been exploited. There has been a general tendency for the decrease of small open-deck boats and an increase of covered-deck boats. As an example of these changes in the last decade, data from the Island of Faial, which accounted as 16% of total Azorean landings in 1992 for the four major taxa listed above is presented in Figure 8. Data were obtained from LOTACOR and from the Departamento de Oceanografia e Pescas databases.

The average weight per landing for each boat gives a good picture of the expected difference between the yields of the different boat types (Figure 8E). These differences must be interpreted carefully. Each landing represents for cabinned boats two or three fishing days (i.e. two or three line sets) whilst for open-deck boats bigger than 9m, one or two fishing days (i.e. one or two sets) and for the smaller open- deck boats only one fishing day. The number of hooks per set is very variable for each boat type, but in general it increases with boat length. In 1979 about 70% of total landings in Faial were performed by open-deck boats; while in 1989 about 90% of the landings were performed by cabinned vessels (Figure 8C). This very marked difference observed in Faial fisheries illustrates well the great changes that have taken place.

Rough estimates of the average yield per hook show that for blackspot seabream these figures have decreased about 50% from 1983 to 1989. The seriousness of this drop is accentuated by recent studies using VPA models (Silva et ul., 1994) indicating a considerable biomass drop of the blackspot seabream stock, a crucial component of the longline fishery. Other species have not declined so drastically but are still decreasing.

Price per kilogram and the yields per hook in PTE for the longline fishery have increased from 1983 to 1992 (Figure 8D). This means that with the increase in the average number of hooks per set the financial income for the fishermen has risen encouraging an increase in fishing effort.

The increase in effort and fishing activity further offshore have put considerable pressure on the demersal fish assemblage (Silva, 1992). A variety of species (at least 50) are caught including some less favoured, low-priced (e.g. oilfish, Ruvettus pretiosus Cocco, bull’s eye, Epigonus telescopus (Risso), sailfin dory Zenopsis conchifer (Lowe) and John dory, Zeus fuber Linnaeus, bird fish, Polymixiu nobilis Lowe) and ‘trash’ species (e.g. rabbit fish, Promethichthys prometheus (Cuvier) and moon fish, Lumpris guttutus (Briinnich); the latter probably because it is highly occasional). This fishing activity will probably restructure the benthic food web. Officially, fishing is confined to traditional lining with the exception of large mesh nets used for the deepwater shark fishery (Dulutius lichu) (Silva, 1987a). Some illegal netting is now occurring on the offshore banks. This may have severe consequences on fish stocks particularly since gear is lost and continues to fish. Despite this being a recent problem, started by fishing boats from mainland, we believe that this will become a serious problem for living marine resources conservation in the Azores, which local authorities have to consider in the near future.

Nearshore netting of questionable legality was recently observed off Faial for tope with small hammerhead sharks also being caught. Rays are also caught in the mixed demersal line fishery. All elasmobranchs are low fecundity species, very susceptible to overexploitation (e.g. the virtually extinct common skate in the Irish Sea: Brander, 1981).

Further research in biology, ecology and fish stock assessment for the multi-species Azorean fisheries is required. This research must take into account the peculiarities of the Azorean Archipelago, namely its geographic isolation, the lack of a continental platform and the existence of particularly important fishing areas-the seamounts. At present little is known about the general physical and chemical oceanography and ecology in the Azores and of these, very important features, in particular. A programme was started in 1994 to study the physical and chemical oceanography, primary productivity, trophic relationships and dispersal of eggs and larvae around the seamounts. This programme should advance rapidly now that a research vessel is permanently available in the Azores.

Table 1. Azorean littoral marine fauna with special regulations. ~. ~~

Abundance and Further management Marine fauna Legislation Regulation accessibility Current status decisions

Moltuscs Ruditapes DLR 15/88/A decussatus 7 April

Port. 63/89 29 August

Patella spp. Port. in 1985/86/87/88

DLR 23/89/A 26 July

DLR 23/90/A renovated in 1991 and 1992

DLR 14/93/A 31 July

Catch and consumption Occurs in a small lagoon In 1988 the population prohibited (1985-1988) of SBo Jorge Island. was estimated on 1.5 and habitat

Only habitat in Azores million individuals, and a biomass of 46 000 kg. Limited local exploitation is permitted

Need to conserve lagoon

Regulates catches and commercial sales. Limited licensed collection P Catches not permitted in Formerly abundant from The stocks of limpets Need of more effective v the central group of intertidal to 6 m depth have recovered slowly control and fiscalization. g Islands on rocky shores between 1984 and 1992 Reinforcement of the role 5

8 of the protected areas.

sales prohibited in all after heavy exploitation. established regulations. Educational 2 actions r- Islands

Catches and commercial sales not permitted. Catches allowed only in Flores and Corvo. Regulates the catches of limpets in all islands. The catches are limited to licensed people. Minimum sizes for each species in the catches are defined. Creates protected areas in all Islands and defines a protected season

Catches and commercial They almost disappeared They seem now well Implementation of h

First in the central group, then in the eastern group

Crustaceans Scyllarides Port. 19/83 5 May l a m

Palinurus Por. 19/83 5 May elephas

Megabalanus Port. 81/84 (J.O.) tintinnabulum

Epinephelus DLR 15/83/A 11 March marginatus

Live bait

Fish

Others Port. 355/89 18 May

Spear gun fishing

DLR 5/85/A 8 May

Minimum length 17 cm. Access by skin and scuba The populations seem Total protection from diving on rocky shores, well established May 1 to August 31. and baited cages Fishing by baited cages is the allowed method Minimum length 17 cm. Total protection from October 1 to March 3 1. Fishing by baited cages is the method allowed Catches closed during the year of 1986

Catches by speargun fishing not allowed

Regulates the size of the nets to catch small pelagics to be used as live bait Maximum quantity allowed: 5 specimens/ person/day

Access by skin and scuba Healthy populations diving, and baited cages apparently. Needs

further studies

Occurs in very limited habitats further studies

Vulnerable. Needs

Common Healthy populations

Urgent implementation of more effective control of commercial sales

Urgent implementaiton of more effective control of commercial catches

Urgent control of commercial catches

Need further knowledge of fisheries biology

Two main species are the The demersal fisheries of Need further study of targets: Trachurus P. bogaraveo is probably these fisheries picturatus and Pagellus approaching the CPUE bogaraveo

Need of regulation of minimum sizes of some species

w c:

328 R. S. SANTOS ET A L .

SPECIES CONSERVATION AND LEGISLATION

Azoreans have always heavily exploited their littoral and nearshore waters, posing considerable problems for both resource management and nature conservation in recent years. In this section of the review we include only species that are caught very near to the shore using artisanal methods, with or without boats. Most of these species are specially regulated to manage human exploitation (see Table 1). Their importance for the local economy is, or has been, significant in some cases. In general, exploitation pressure increased with the ready availability of cheap snorkelling gear in the 1970s. Collection using SCUBA (Self Contained Underwater Breathing Apparatus) gear is banned, except for scientific purposes by authorized personnel. However, abuse is common.

Molluscs and Crustacea

Limpets (lapas) Two species of limpets occur in the Azores: ‘lapa brava’ the ‘wild limpet’ (Patella aspera Roding also

called P. ulyssiponensis Gmelin by Christiaens, 1973) and ‘lapa mama’ the ‘tame limpet’ (Patella candei D’Orbigny, see Christiaens, 1973). P. candei inhabits the intertidal, occasionally being found subtidally; P. aspera occurs low on the shore and in the upper sub-littoral to 6 m depth, very exceptionally reaching 15 m (see Hawkins et al., 1990a,b; Menezes, 1991a). Both species occur together low on the shore. P. candei is mainly exploited on a subsistence or recreational basis, whilst P . aspera has been exploited mainly by both licensed and unlicensed skin-divers depending on the island concerned. A full time professional fishery was present in Sgo Miguel until 1988.

The decline and recovery of the stocks have been followed in different islands since 1984 (Martins et al., 1987; Santos et al., 1990). The priority goals of research have been comparative assessment of the abundance of stocks, recruitment, change in catches and market, and conservation regulations. Their general biology, ecology and systematics have also been studied (e.g. Hawkins et al., 1990a,b; Corte-Real et al., 1992).

Most attention has been given to P. aspera which, in the mid 198Os, was the fifth or sixth most important commercially exploited species in the Azores. The commercial fishery in Slo Miguel crashed in 1988. Very low population levels were reached in the Central group between 1984 and 1987. In 1989 a ban on all collection of limpets was established for the Central and Eastern group. Currently, P . aspera populations are abundant in the western group where they have never been subjected to heavy exploitation. They are recovering in the Central group after various conservation measures. There is still a very critical situation in S. Miguel and to a slightly lesser extent in Sta. Maria. There were dense populations in Formigas in 1990, but recently they have been under illegal exploitation in what is meant to be a reserve. At present harvesting is strictly prohibited in selected areas in the islands. A closed season is also established (Table 1). Illegal harvesting is, however, frequent. Importations from Portugal and Spain during the last few years have made regulation still more difficult to distinguish illegally sold limpets from those which it is claimed have come from the continent.

P. aspera may be particularly vulnerable to over-exploitation leading to recruitment failure. These are protandrous hermaphrodites (Thompson, 1979). Removal of larger limpets can be at such a rate that few large females remain in the population and smaller limpets are removed before they have a chance to change sex. The proposed minimum size of 50mm would allow at least some females to reproduce in the population.

P. candei was never badly affected by exploitation. It was primarily the target of subsistence and recreational collection on the shore. This species becomes sexually mature at a much smaller size and there is little evidence for marked protandry. It grows rapidly and can occur in a variety of habitats, doing particularly well on large boulders. Although its abundance and average size have probably decreased in


recent years it is unlikely to suffer from recruitment overfishing leading to population decreases as found in P. aspera. In contrast, P . candei could be exploited on all islands providing a minimum size is enforced. It would also be prudent to have a closed season during which all limpet collecting is banned to allow reproduction to occur. This is not essential in the case of P . candei, but enforcement of regulations concerning P. aspera would be made easier, since when limpets are being illegally gathered the excuse of mistaken identity would be removed.

Clams Tapes decussatus (Linnaeus) is the only clam commercially exploited in the Azores. It occurs only in a

small lagoon in S. Jorge Island, which is a protected area. Its commercial importance was local until it gained a gastronomic status that also reached the other islands. Its exploitation increased dramatically in 1985/86. This, together with an obstruction of the channel connecting the lagoon with the sea, endangered this clam population. Studies were carried out (Santos and Martins, 1987; Santos et af . , 1989; Morton and Cunha, 1993) to evaluate water quality, sediment characteristics, distribution and clam biomass, population structure and growth. Exploitation is now strictly regulated by the number of licenses, quantity and minimum size (Table 1). The possibility of human introduction cannot be ruled out (Morton, 1967).

Crustacea The slipper lobster, Scyflarides latus (Latreille), locally called cavaco, is a highly esteemed delicacy in the

Azores. The study of its reproductive and growth biology and exploitation was initiated in 1982 (Martins, 1985a). After these studies it was possible to establish rules concerning its exploitation and conservation (Table 1). The reproductive biology and fisheries of the other lobster present, the spiny lobster, Palinurus efephus (Fabricius) (Table 1) requires study; there is a close season and size limit. The status of the stocks of this species is not clear, since the most of the catches, as with S . latus, do not pass through the fish auction.

The giant barnacle, Megabafanus tintinabulum (Linnaeus), is a species highly appreciated locally as seafood (Table 1). The populations of this species around the islands have not been studied. Data on fisheries effort and capture do not exist as the exploitation is very artisanal and it is sold directly from the harvesters.

Similar regulations as for spiny lobster apply to the spiny spider crab, Maja squinado (Herbst) (Table 1). Minimum size in fisheries is 20 mm more than allowed in Portugal. No studies have been done. Two deepwater crabs, ‘supateiru’, Cancer belliunus Johnson, and Chaceon affinb (Milne-Edwards and Bouvier), are locally believed to be abundant. The biology and potential for exploitation of the second one is now being studied (Gonqalves and Pinho 1994; Gonqalves and Santos, 1994). Both species have potential for fisheries and commercial interest (Zaferman and Sennikov, 1991).

The study of exploited crustacean (lobsters, crabs, shrimps, barnacles) is problematic as only a small proportion of the catches passes through the official fish auctions.

Common octopus (Octopus vulgaris Lamarck) Fourteen species of Octopoda occur in the Azores (Gonqalves, 1993). Only one (0. vulgaris) is

exploited and deserves attention in terms of conservation. The study of its biology and stock assessment is underway. As the catches traditionally escape control by the authorities, very little is known about the actual level of exploitation. However, there are reasons to believe that the stocks are diminishing in the main islands (Gonqalves, 1993). It is also likely that there has been a switch from catching limpets to catching octopus because of the ban introduced on collecting limpets in the central and eastern islands in 1989.


Fishes

Coastal fisheries There is considerable recreational and semi-subsistence fishing using rod and line or handlines from the

coast and close inshore from small boats, as well as considerable spearfishing activity. The dusky perch, Epinephelus marginatus, is the only fish under special regulation in the Azores (Table 1).

Catching this grouper by speargun is banned, although traditional fishing using hooks is permitted. This species is still quite abundant but the fishery needs monitoring.

Several other species of labrids, like Coris julis (Linnaeus), moray eels, e.g. Muraena helena Linnaeus, M . augusti (Kaup), Gymnothorax unicolor (Delaroche), sparids, Boops boops (Linnaeus), Pagellus acarne, Diplodus sargus cadenati de la Paz, Bauchot and Daget, Sarpa saZpa (Linnaeus), the red mullet, Mullus surmuletus Linnaeus, the parrot fish, Sparisoma (Euscarus) cretense (Linnaeus), are exploited. However, the most important commercial coastal fishery in the Azores, is for juvenile blue horse mackerel (Trachurus picturatus). Despite growing exploitation stocks were recently still assessed as good (Isidro, H., 1990).

Demersal fisheries discussed above constitute one of the most important commercial activities in the Azores. There are no special regulations, such as minimum, sizes or close seasons, for demersal exploited species in the Azores (in mainland Portugal, 25 demersal fishes have special regulations). Officially fishing is confined to traditional lining, except in the case of the juveniles of the species (Pagellus spp. and T . picturatus), used as live bait, and the deep-water shark Dalatius licha. It is considered that regulations need to be introduced on a species by species basis.

Threatened marine fishes The third volume of the Red Book of Portuguese Vertebrates (Livro Vermelho 111, 1993) determines the

conservation status of 15 Azorean fish species: four are commercially threatened (CT), four are rare (R), two of indeterminate status (I) and five are insufficiently known (K) (Table 2). The pipefishes and seahorses (Syngnathidae), which were known to be abundant some years ago, are now extremely rare, despite being unexploited. Finally three Serranidae were included: one may be affected by speargun fishing (Mycteroperca fusca), the second (Epinephelus marginatus) by both speargunning and commercial fishing and, the third (Polyprion americanus) by commercial fishing using illegal netting.

Marine turtles

Five species of turtles have been recorded in the Azores: Caretta caretta (Linnaeus) (loggerhead), Chelonia mydas (Linnaeus) (green turtle), Eretmochelys imbricatu (Linnaeus) (hawksbill), Lepidochelys kempii (Garman) (Kemp’s ridley) and Dermochelys coriacea (Linnaeus) (leatherback), of which only C . caretta is encountered frequently. The green turtle and the leatherback are reported every year while the other two have been recorded only once or twice. None of these species breeds in the Azores. Only one of the species, L. kempi, was considered threatened in the Red Book of Portuguese Vertebrates (Livro Vermelho I, 1990). The status of ‘vulnerable’ was given, not because of threats in the area but regarding threats at nesting beaches distant from the Azores.

Conservation of marine turtles in the Azores is ruled by international conventions (e.g. Bern Convention) which establish total protection of the species. Some local exploitation for food and curios was common until a few years ago. This has stopped as public opinion has changed.

Research is carried out in the Azores on loggerheads, the most common species, which make regular transatlantic migrations. Individuals reaching the Azores come from the Western Atlantic, travelling along the Gulf Current, according to tagging studies (e.g. Eckert and Martins, 1989). Nesting beaches also exist in


Table 2. Azorean marine fishes with threatened status.

Family Species IUCN status Azorean status*a

Rhincodontidae Lamnidae Cethorhinidae Squalidae Syngnathidae

Phycidae Serranidae

Acropoma tidae Spar i d a e

Blenniidae

Gobiesocidae

Rhinchodon typus Smith Charcharodon carcharias (Linnaeus) Cethorhinus maximus (Gunnerus) Dalatias licha (Bonnaterre) En telurus aequoraeus (Linnaeus) Hippocampus hippocampus (Linnaeus) Hippocampus ramulosus Leach Phycis phycis (Linnaeus) Epinephelus margina tus (Lowe) Mycteroperca fusca (Lowe) Polyprion americanus (Schneider) Pagellus bogaraveo (Briinnich) Pagrus pagrus (Linnaeus) Blennius ocellaris Linnaeus Lipophrys pholis (Linnaeus) (giant race) Lipophrys trigloides (Valenciennes) Parablennius ruber (Valenciennes) Diplecogaster bimaculata pectoralis Briggs

1 K K R

CT K R R

CT V I I

CT CT K R K K K

R, = rare: taxa with small world populations that are not at present ‘endangered’ or ‘vulnerable’, but are at risk. These taxa are usually localized within restricted geographical areas or habitats or are thinly scattered over a more extensive range. I, indeterminate: taxa known to be ‘endangered’, ‘vulnerable’, ‘rare’ or ‘commercially threatened’ but where there is not enough information to say which of three categories is appropriate. CT, commercially threatened: taxa not currently threatened with extinction, but most of all of whose populations are threatened as a sustainable commercial resource, or will become so, unless their exploitation is regulated. K, insufficiently known: taxa that are suspected but not definitively known to belong to any of the above categories, because of lack of information. ‘Livro Vermelho dos Vertebradm de Portugul 1993 (Vol. 111: Peixes Marinhos). ICN (Lisboa).

the Mediterranean and Cape Verde Islands, Kemp’s ridley, green turtles and hawksbill, which also make long migrations along the ocean currents, are only accidental visitors.

A joint programme of research on pelagic turtles (mainly loggerheads), by the University of Florida (Dept of Zoology) in cooperation with the University of the Azores is underway (Bolten et a/., 1992, 1993). From 1984 to 1994 over 1500 turtles were tagged. They were captured at sea, by tuna boats, tagged at the University and then released again in the sea. Approximately E3 ($5) for each turtle is paid to the fisherman. In 1990 a more ambitious programme was implemented. With the help of the tuna fleet, turtles are measured, tagged at sea and immediately released. Payment is made on the basis of the number of individuals that were tagged and measured (Bolten et al., 1993). In 1994 a programme of satellite tracking of turtles marked and released in the Azores was initiated.

The main threat for sea turtles at sea are oceanic pollution (e.g. oil and tar) and floating rubbish (e.g. plastics, polyethylene, Styrofoam, etc.) which they eat (Nierop and Hartog, 1984; Carr, 1987). Unfortunately such marine debris is common in the waters surrounding the Azores.

Seabirds

Species The breeding seabirds assemblage of the Azores comprises five species of Procellariiformes, four

Charadriiformes and one Pelecaniform. The regular breeders are: Bulwer’s petrel Bulweria bulwerii, Cory’s shearwater Calonectris diomedea borealis, probably Manx shearwater Puffinus puffinus, little shearwater


Puffinus assimilis baroli, Madeiran storm petrel Oceanodroma castro, yellow-legged gull Larus cachinnans atlantis (Dwight), common tern Sterna hirundo and roseate tern Stern dougallii (Bannerman and Bannerman, 1966). These are two occasional breeders, red-billed tropicbird Phaethon aethereus Linnaeus (Furness and Monteiro, 1995) and sooty tern Sternafuscata Linnaeus (Monteiro et al., in press), a possible breeder, Fea’s petrel Pterodroma feae (Salvadori) (Monteiro and Furness, 1995) and a possible former breeder, white-faced storm petrel Pelagodroma marina (Latham) (Monteiro et al., in press).

Status and populations The distribution and total populations of all of these seabirds are poorly known, except for the roseate

tern (c. 1000 pairs, mainly on Flores, Graciosa and Santa Maria) and the common tern (c. 4000 pairs, which breed on all islands) (del Nevo et al., 1993). The Azores hold over 60% of the total European population of the roseate tern, being the most important breeding site in Europe for that declining species.

The most abundant seabird is the Cory’s shearwater. It breeds on all islands with an estimated population of 30 000-70 000 pairs, possibly representing the majority of the subspecies borealis (Monteiro et al., in press).

The little shearwater, Bulwer’s petrel and Madeiran storm petrel have tiny populations on a few islets that do not have resident populations of rats. The extremely rare gadfly petrel may represent a remnant population reduced almost to extinction, probably due to predation by rats. As this is the northerly edge of the range for these species of predominantly tropical distribution, this may in part explain the small populations. However, judging from the historical records, intense exploitation by man, habitat damage and predation by introduced mammals seem to have been the main threats (Monteiro et al., in press).

The Madeiran storm petrel has two temporally segregated populations breeding on the islets of Graciosa, thought to represent sibling forms (Monteiro and Furness unpublished).

The present status of the Manx shearwater in the Azores is doubtful. The species has bred in the western group Gust two records; Flores, 1865 and Corvo, 1929; Bannerman and Bannerman, 1966), where it possibly still breeds (Grimmet and Jones, 1989).

Research and conservation Recent inventories established the importance and priority for conservation of the Azores seabird fauna

in Europe. All the regular breders, except the yellow-legged gull and the common tern, are ‘Species of European Conservation Concern’ with an Unfavourable Conservation Status (categories 1 to 3; Tucker and Heath, 1994); six species are classified as threatened in the Red Book of Portuguese Vertebrates (Table 3). Six sites were selected as ‘Important Bird Areas’ because of their seabird populations (Grimmet and Jones, 1989) and fulfil the criteria for designation as ‘Seabird Sanctuaries’ (Duffy, 1994).

Table 3. Azorean sea birds with threatened status.

Family Species Bird’s directive Azorean statusa

Procellariidae

Hydrobatidae Laridae

Bulweria bulwerii (Jardine and Selby) included K

Puffinus puffinus (Briinnich) included K Puffinus assirnilis baroli Bonaparte included K Oceanodroma castro (Hartcourt) included K Sterna dougaflii Montagu included v Sterna hirundo Linnaeus included K

Calonectris diomedea (Cory) included NT

K, insufficiently known: taxa that are suspected but not definitively known to belong to any of the above categories, because of lack of information. NT, not threatened. V, vulnerable: taxa believed likely to move into the ‘endangered’ category in the near future if the causal factors continue operating. aLiwos Vermelho dos Vertebrados de Portugal 1990. (Vol. I:. Mamiferos, Aves, Repteis e Anfibios). SNPRCN (Lisboa).


This has some parallel in the legal status of both the species and breeding sites. All the main breeders except the yellow-legged gull are included in Annex 1 of the EU Wildbirds Directive 79/409 and Annex 2 of the Bern Convention, both transcripted to the national law. Virtually all the major known seabird sites in the Azores have been listed, as ‘Special Protection Areas’ under the EU Wildbirds Directive (Anon., 1991) but national regulations are still lacking. The most important seabird sites are not classified locally as Nature Reserves, although some relevant seabird colonies lie in the following Reserves: Vulcio dos Capelinhos, Monte da Guia, Ilheu do Topo, Lagoa do Fogo, IlhCu de Vila Franca. Overall, protective measures for species and sites are still poorly enforced.

Human exploitation (for oil, food and feathers), mammal predation and habitat modification were major threats in historical times (Monteiro et al., in press). Current threats are diverse and may affect differentially the various species. These range from predation by gulls and introduced mammals, human disturbance and exploitation, habitat loss due to invasive alien plants and overgrazing by introduced herbivores, and potentially competition with fisheries (Le Grand et al., 1984; Monteiro et al., in press).

Current seabird research involves status and distribution, breeding biology, feeding ecology and pollutant levels (Martin, 1986; Bibby and del Nevo, 1991; Klomp and Furness, 1992; del Nevo et al., 1993; Moore, 1994; Hamer et al., 1994; Furness and Monteiro, 1995; Monteiro and Furness, 1995; Monteiro et al., 1995; Ramos and del Nevo, in press; Monteiro et al., in press) and has been conducted by the University of the Azores in collaboration with the University of Glasgow (Applied Ornithology Unit), the Royal Society for the Protection of Birds and the Instituto de Conservagiio da Natureza.

Marine mammals

Occurrences Between 20-23 species of cetaceans are known from the Azorean waters belonging to five families (Table

4). Also two species of pinnipides have been recorded-Phoca vitulina Linnaeus: Reiner, 1990 and P. hispida (Schreber): Le Grand, 1981, but these were very exceptional. Of these sea mammals, 10 are included in the Red Book of Portuguese Vertebrates (Livro Vermelho I, 1990).

Exploitation and conservation The 18th century whaling around the Azores was practised by British ships based in New England. In the

middle of last century whaling based in the Azores was initiated as commercial activity. Up to 200 small boats were hunting the sperm whale (Physeter macrocephalus) at one time.

Whaling in the Azores evolved as a nearshore activity. Methods were always traditional using small open boats (Clarke, R., 1954, 1981; Esteves, 1984). The contribution of the Azorean whaling in the world context, solely concerning sperm whales, was always very low (Galhardo, 1990). In the 1960s whaling began to decrease (Martin and Melo, 1983) and commercial whaling ceased completely in 1984 (Galhardo, 1990). However, two individuals were caught in 1987. At first the declining importance of whaling was mainly related to the growth of other fishing activities, like tuna fishing and demersal fishing (Clarke, R., 1981). In the last years there were also international pressures against whaling products, and a growing difficulty in finding markets for the oil, for which artifical substitutes had existed for many years. Only teeth (‘scrimshaw’) or whale bone carving was really profitable latterly.

Dolphins were also hunted in the Azores, both for human consumption and fishing bait. In 1983 a law was approved (DLR#2/83/A) by the Regional Parliament protecting four species of

cetaceans: Delphinus delphis, Stenella coeruleoalba, Tursiops truncatus and Grampus griseus. In continental Portugal and Madeira all sea mammal species were already protected. Total protection of sea mammals in the Azores was only defined in 1989, by a Central Government law (DL#316/89, 22 September), which adopted and extended the Bern Convention to continental Portugal, the Azores and Madeira.

Common dolphins (e.g. D . delphis and T . truncatus) are highly abundant in the area (Clarke, 1981).

334 R. S. SANTOS ET A L .

Table 4. Cetaceans which occur in the Azores (Reiner et al., 1993; Steiner, 1995) and their IUCN and Azorean status.

Family Species IUCN status Azorean statusa

Balaenidae ? Eubalaena glacialis (Miiller) Balaenopteridae Megaptera novaenglia (Borowski)

? Balaenoptera musculus (Linnaeus) Balaenoptera physalus (Linnaeus) Balaenoptera borealis Lesson Balaenoptera acutorostrata LacCpede

Phoceinidae ? Phocoena phocoena (Linnaeus) Delphinidae Delphinus deiphis Linnaeus

Tursiops truncatus (Montagu) Stenella coeruleoalba (Meyen) Stenella jirontalis (Cuvier) Pseudorca crassidens (Owen) Orcinus orca (Linnaeus) Grampus griseus (Cuvier) ? Globicephala melaena (Traill) Globicephala macrorhynchus (Gray)

Mesoplodon europeus Gervais Mesoplodon bidens (Sowerby) Ziphius cavisrostris Cuvier

Kogiidae Kogia breciceps (de Blainville) Stenidae Steno brenadensis Ph yseteridae Physeter macrocephalus Linnaeus

Ziphiidae Hyperoodon ampullatus (Forster)

E I V I E V V

I R

K I

R

V

K

K

~~ ~~ ~~~~~

E, endangered: taxa in danger of extinction and whose survival is unlikely if the causal factors continue operating. V, vulnerable: taxa believed likely to move into the ‘endangered’ category in the near future if the causal factors continue operating. R, rare: taxa with small world populations that are not at present ‘endangered’ or ‘vulnerable’, but are at risk. These taxa are usually localized within restricted geographical areas or habitats or are thinly scattered over a more extensive range. I, indeterminate: taxa known to be ‘endangered‘, ‘vulnerable’ or ‘rare’ but where there is not enough information to say which of three categories is appropriate. K, insufficiently known: taxa that are suspected but not definitively known to belong to any of the above categories, because of lack of information. ‘Livro Vermelho dos Vertebrados de Portugal 1990 (Vol. 1: Mamiferos, Aves, Rkpteis e Anfibios). SNPRCN (Lisboa). ? species records considered as dubious.

Research To our knowledge, the first studies of sea mammals around the Azores were made by the research team

of Prince Albert I of Monaco, and were compiled by Richard (1936); but reference must also be made to Girard (1892) and, in the first half of this century, to Chaves (1924), Ferreira (1935) and more recently Clarke, R. (1954, 1956, 1981).

Current research involves feeding habits, physiology and social behaviour (Martin, 1986; Martins et a f . , 1985; Arnbom e t af., 1988; Steiner and Gordon, 1990; Clarke, M. R. e t a f . , 1993). Since 1987 Jonathan Gordon and his team have become regular visitors to the Azores with the yacht ‘Song of the Whale’. Their research has greatly increased the knowledge of sperm whales’ social behaviour; communication, abundance and distribution of several species of cetaceans; acoustics applied to cetacean detection; calculations of group size and individual sizes, and photo identification (Gordon et a f . , 1987, 1988, 1989). Another aim of the programme was to study the interest and biological feasibility of whale watching in the Azores (Henson et al., 1989). Enterprises are now operating, mostly, around Pic0 Island. There is an urgent need to produce adequate regulation of whale watching around the Azores, since the activity is growing fast and has outstanding potential (WDCS, 1995).


HABITAT CONSERVATION

Measures protecting individual species in the Azores are normally related to habitat management-with the exception of turtles, cetaceans and birds. The geomorphology of the Azores makes the coastal zone rather narrow as a consequence of the high gradient of the slope (Martins, J. A., 1986). This restricts the extent of littoral habitats. Habitat conservation by creation of Marine Protected Areas is an important condition for the conservation of marine fauna and flora, and of whole ecosystems. In an archipelago with scattered islands, the creation of a network of protected areas must comprise representative littoral areas in each of them. They are particularly important for various reasons.

Several littoral species are heavily exploited e.g. limpets: Patella aspera and P. candei; barnacles: Megabalanus tintinnabulum; lobsters; Palinurus elephas and Scyllarides latus; octopus: Octopus vulgaris; several fish species and some algae, particularly Pterocladia capiffacea (Gmelin). Reproductive output for some of these species is little known and some of them are protrandrous (e.g. Patella aspera) or protogynous (e.g. Epinephelus marginatus) hermaphrodites. Females need to be protected.

Scope for colonization from elsewhere is probably low due to limited dispersal for some species (e.g. Patella species) and strongly dependent on weather and sea conditions. Recruitment probably varies from year to year in some species. Hence, recovery from disruption of habitats and species populations, mainly due to human predation can only occur slowly by the reproductive output from other islands under less intense exploitation.

In addition to their habitat preservation roles, marine reserves located in the different islands will have a major resource conservation function: to serve as reproductive reserves as a source of recruitment for exploited species on their own and adjacent islands.

Designated marine protected areas in the Azores

For several years the University of the Azores has been lobbying for the implementation of a plan for conservation of marine areas in the Azores (Martins and Santos, 1991; Santos, 1992; see also Saldanha, 199 1) which should essentially follow criteria of selection based on: (i) physical criteria; (ii) ecological criteria; (iii) cultural and educational criteria (cultural value, scientific value); (iv) pragmatic criteria (value for research or monitoring, degree of threat or fragility, feasibility, redundancy, regional, national or international value, educational, recreational and economic value) and to the peculiarities of the Archipelago. This section briefly summarizes various areas currently enjoying protection or areas that are proposed as Marine Protected Areas (MPA).

At present there are nine MPA, distributed on four islands and one isolated group of islets. Seven are designated as Marine Reserves (MR); one is a Protected Landscape; and another is a Special Ecological Area. Six are located in the Eastern group and three in the Central group. There are currently no protected areas in the Western group. Apart from these, there are fifteen Special Protection Areas for the Birds of the Azores (Anon., 1991). Most of them, excepting two, have special interest for the marine birds but they only include terrestrial areas. They were created to conform to the EEC directive 79/409. Descriptions of the protected areas follow.

Eastern group

Sunfa Maria (Figure 9) Four MRs (Nature Reserves of Baia da Maia-between 36" 57.1'N, 25" 0l.l'W and 36" 55.7'N, 25" 09W, Baia de S5o Lourenqo-between 36" 59.9'N, 25" 03.1'W and 36" 58.8", 25" 02.4'W, Baia dos Anjos- between 37" 02'N, 25" 13'W and 37" 08'N, 25" 08.6W and Baia da Praia-between 37" 56.8'N, 25" 07.3'W and 36" 55.7", 25" 09W). These were designated (DLR 7/87/A of May 29) mainly on recreational criteria, but are also biologically and ecologically very important for this most southern island in the Azores, and

336 R. S . SANTOS ET AL. L S.MIGUEL

10 km L

S. M A R I A

W

FORMIGAS

Figure 9. Existing (dotted pattern) and proposed (lined pattern) marine protected areas for the eastern group of islands.

representative of the littoral habitats. Each of the MPA extends 1.5 km from coast line. The protective legislation is not particularly clearly defined and not yet enforced.

Formigas (Figure 9) The isolated Formigas Islets (37", 19", 24" 34'W) and Dolabarat Bank (37" 13.1'N, 24" 48'W), around 50 miles away from Siio Miguel and 20 miles from Santa Maria form another important MR (created by DLR 11/8/A of April 14). They constitute representative shallow to medium depth rocky (maximum depth around 400 m) habitats in the North Atlantic. Their isolation provides some protection but even recently their populations of limpets and barnacles have been threatened by illegal exploitation. Artisanal fishing with boats of less than 14m is allowed. Regulations are not yet published.

Scio Miguel (Figure 9) A small islet, Ilhtu de Vila Franca, close to the coastline has been established as a MPA (DLR 3/83/A of March 3). It is an extinct volcanic crater, with ecological and recreational importance. Hamer et al. (1 989) noted that the Vila Franca Islet 'is exceptional in the density' of nests of Cory's Shearwater (Calonectris diomedea borealis) and 'in the range of habitats used as nest sites'. According to Hamer et al. (1989) there is a need for implementation of existing special regulations concerning the protection of the nesting colony of Cory's shearwater. This area has considerable recreational use and regulations need to be implemented, coupled with public information, to encourage observance of voluntary codes of practice to minimize litter and disturbance ot wildlife, particularly birds.


Central group

Srio Jorge (Figure 10) One N R (Reserva Natural do Zlhe'u do Topo, created by DLR 13/84A of February 20) is located on the eastern extremity of the Island. It has great ornithological and botanical interest. Unfortunately, until very recently, cows were allowed on the islet affecting the colonies of marine birds. This was solved when the regional administration bought the islet. The N R should be slightly enlarged to include the coast of the island at 38" 33.2'N, 27" 45.5'W and 38" 32.6", 27" 45.5'W. Despite its great biological interest, regulations are not yet published.

One Special Ecological Area (Area Ecoldgica Especial da Lagoa do Santo Cristo created and regulated by DLR 6/89/A of July, 18 and Port. 63/89 of August 28). It constitutes a singular and unique habitat in the Azores where the clam Ruditapes decussatus occurs. Its exploitation is strictly regulated. The SEA comprises a coastal lagoon, with soft bottom, encircled by a ring of boulders on the sea side. Area protected: 18ha. and the adjacent sea coast to the 30m bathymetric.

Faial (Figure 10) One Protected Landscape, Monte da Guia, includes a small marine reserve, confined to the crater of an old volcano opened to the sea (created and regulated by DLR 1/80/A of January 31 and DLR 13/87/A of March 3 1). The volcano is formed by two 'caldeiras' breached seaward. The inside depths vary from 6 m to

39

N

G R A C I O S A c

FAIAL a S JORGE \ TERCEIRA a

Figure 10. Existing (dotted pattern) and proposed (lined pattern) marine protected areas for the central group of islands.


40 m, with rocky and gravelly bottoms. Only the inside of the volcano is a marine reserve. It is comprised of bedrock (soft tuff) giving way to boulders, submarine caves and eventually, sediments. This area is very close to the main town of Faial. It should be extended to include Porto Pim, a shallow sandy bay, and ought to include buffer zones. The terrestrial part is particularly interesting for its endemic flora. It is close to the laboratory of the Department of Oceanography and Fisheries and therefore ideal for study. This area also includes an important terrestrial reserve.

Recommended marine protected areas

The following areas have not been designated as yet, but they are considered potentially suitable for designation, in whole or part, as marine protected areas. The areas suggested below for designation should extend to the depth of 30m.

Eastern group

Santa Maria (Figure 9) Lagoinhas This site (37" 01.3'N, 25" 05.5'W), which holds a regular and important colony of roseate terns Sterna dougallii with up to c. 80 pairs, is included in the inventory of Grimmet and Jones (1989). This justifies the urgent classification of the islets as a protected area.

Ilheu da Vila This islet is an important bird area holding internationally significant seabird populations (seven species occur in this islet, among them over two hundred breeding pairs of roseate terns, see also Grimmet and Jones, 1989). It should be urgently classified as a nature reserve and access should be restricted.

S6o Miguel (Figure 9) Caloura There should be implementation of a Protected Landscape in the region of Caloura on the south coast of S. Miguel-from Ponta da Agua de Pau (37" 42.8'N, 25" 31.8'W) through Caloura, Ponta da Galera until Vila Franca (37" 42.6", 25" 26.3'W), including the Ilheu da Vila and the Baixa das Cracas. This area is characterized by sea beds of great geological beauty, formed by lava arcs. The coast is very indented and sinuous forming small protected bays with considerable habitat variation. Zonation patterns in the area have been described by Hawkins et al. (1990a). Other faunistic and floristic work in the area include descriptions of mollusc communities (see Martins, 1990) and birds (Hamer et al., 1989). At present the owner of the hotel at the shore front operates it as a voluntary marine reserve; spearfishing by guests being banned.

Mosteiros Mosteiros is the most unusual fishing port in the Azores, since the fishing harvest is mostly of inshore resources not systematically sold in other fish auctions. Octopus, limpets, lobsters, wrasses, giant barnacles and whelks are heavily exploited in the surrounding littoral that for long stretches does not go deeper than 20m. The flat shallow submarine basaltic platforms are not common in the Azores, where much of the coast tends to fall steeply to great depths.

This locality should be urgently classified as a Special Ecological Area, and properly regulated, in part to ensure the future of the local fishing industry. The area to be considered is from Ponta da Ferraria (37" 51.5'N, 25" 51.2'W) to Ponta da Bretanha (37" 54.5'N, 25" 47.1'W), including, for total protection, the Ilheus dos Mosteiros.


Porto Formoso From Porto Formoso (37" 49.4", 25" 25.7'W) to Porto da Maia (37" 50'N, 25" 23.1'W) on the north coast of SHo Miguel are a variety of sandy and rocky habitats. These have considerable ecological interest as well as potential for tourism.

Nordeste From Ponta do Arne1 (37" 49.4", 25" 08.3'W) to the Ponta da Madrugada (37" 47", 25" 08.7'W) is a rocky exposed coast on the northeast side of the island. This is a site with great potential as a reserve for populations of important species of invertebrates and fish inhabiting the rocky coasts.

Ribeira Quente Sandy areas with some boulders close to the village of Ribeira Quente (37" 44", 25", 18'W): from Ponta do Garajau to Ponta da Albufeira. A large area occurs with gaso-hydrothermal activity (from the intertidal sandy beach down to 15 m and deeper). Removal of sand for industrial purposes should be prohibited.

Central group

Terceira (Figure 10) No Marine Reserves presently exist on this Island. We propose the implementation of at least three of the following areas.

Ilheus das Cabras (and Ilheus dos Fradinhos) This is a group of small islets and rocks from 1 to 5 km away from the southern coastline. They possess a good variety of marine fauna, from molluscs to crustaceans and fish. The Cabras islets (38" 38'N, 27" 02.8'W) give shelter to an important breeding colony of common terns and to a few pairs of the vulnerable roseate tern (del Nevo et al., 1990).

Monte Brad (and surroundings) The proposal is to extend seawards the terrestrial reserve already implemented, in order to protect the shore fauna and flora and create a protected feeding area and buffer zone for the nesting colony of common terns. All the coastal area between 38" 39.2'N, 27" 14.8'W and 38" 34", 27" 12.8'W should be delimited.

Serreta The area from Baia Grande (38" 45.5'N, 27" 22.4'W, Ponta da Serreta to Ponta do Cavalo (38" 48'N, 27", 18.2'W) is particularly interesting for littoral fish and crustaceans.

Other Special measures should be taken to protect colonies of breeding terns, particularly the roseate tern located in IlhCu de Mbs (southwest coast).

Graciosa (Figure 10) Ilheu da Praia This small islet (approximately 700 x 300 m) is on the west coast and in front of Baia da Praia at a distance from coast line of 250 m. It has a diverse fauna and flora, paricularly fish. The area to be designated should also include the coast from Baixa do Redondo (39" 04", 27" 58.8'W) to Ponta dos Fenais (39" 02.2'N, 27" 57.2'W). This is an important area for internationally important seabird populations (Grimmet and Jones, 1989). Rabbits and lizards should be eradicated, access restricted and management of vegetation initiated to counter severe problems of erosion.

Ponta do Carapacho and Ilheu de Baixo From Baia do Carapacho (39" 06", 27" 58.2'W) to Ponta do Feliciano (39" 01.2'N, 27" 57'W), on the southwest coast, includes several islets. It has important submarine habitats. Additionally there are two


important colonies of breeding roseate terns at Ilheu de Baixo and Gaivota. Common terns also breed there. These two islets are important bird areas (Grimmet and Jones, 1989). The Ilheu Gaivota holds a regular and important colony of roseate terns, with up to c. 50 breeding pairs. This justifies a maximum level of protection and prohibition of access.

Ponta Branca and Ilheu This stretch on the southeast coast-Baixa Afonso Correia to Ponta Branca (39" 01.9", 28" 03.8'W; 39" 01.4", 28" 02.4W) is of great relevance for the colonies of common and roseate terns.

Baia da Vit6ria to the Baia das Diagaves (Ponta da Barca, IlhCus de Barro Velho) This is a very interesting and irregular rocky coastline exposed to the north between 39" 05.4'N, 28" 03.2'W and 39" 05.7'N, 28" 01.4'W. A variety of habitats support a diverse marine flora and fauna.

Scio Jorge (Figure 10) Fajii dos Cuberes to Fajii do Santo Cristo Fajii do Santo Cristo possesses an already classified area that should be integrated in a Nature Reserve including the neighbouring Fajii dos Cuberes. Both contain lagoons unique in the Azores. The surrounding littoral is particularly rich in fish, molluscs, crustacea and many other invertebrates. The lagoon of Santo Cristo is rich in clams and serves as a nursery for some fish species, such as the grouper Epinephelus marginatus (GonGalves et al., 1993). The biological diversity of this area is allied to the beauty of the landscape, to cultural attributes and tourism potential. The protected coast should include the area between 38" 38.6'N, 27" 58.2'W and 38" 37.1'N, 27" 55.2'W. Fajii dos Cuberes holds a small, but regular, mixed colony of roseate and common terns.

Morros das Velas From Morro de Lemos at 38" 40.6", 28" 12.5'W to Morro Grande at 38" 40.6", 38" 12.5'W are a pair of coastal hills surrounding a bay open to the sea at the south coast of the island. These mounts hold regular and important colonies of common terns.

Ponta dos Rosais The area surrounding the Ponta dos Rosais on the north coast at 38" 45.3'N, 28" 18.5'W and on the south coast at 38" 44.9", 28" 18.6'W on the far west part of the island has rocks emerging steeply from deep water as well as shallow reefs. It is an area very rich in fish.

Pic0 (Figure 10) Despite its size (the highest island and the second largest of the Archipelago) and marine biological diversity and richness, Pic0 does not possess any protected marine area. At least two should be designated.

Ilhtus da Madalena and Baixa do Canal The first are two beautiful islets, part of an old volcano, located on the west coast. Surrounding depths do not exceed 20m. The second is a shallow bank between Pic0 and Faial. Both are highly rich in resident fish, e.g. Epinephelus marginatus, Mycteroperca fusca, Sparisoma (Euscarus) cretense, Muraena spp., etc.-and also large itinerant fish--e.g. Sphyraena viridensis Cuvier, Pseudocaranx dentex (Block and Schneider), Seriola dumerili (Risso), S. rivolina Cuvier, Mobula mobular (Bonnaterre). The islets give shelter to breeding pairs of both species of terns present in the Azores and to other seabirds. Due to the easy access, both from Faial and Pico, they have potential for underwater tourism, and hence some regulation or codes of practice are needed. On Pico's coast the delimited area should include from Pi. do Monte (38" 30.1'N, 28" 32.6'W) to Ponta do Cachorro (38" 33.4", 28" 26.9'W). The shallow bank is located at 38" 30.07'N, 28" 35.02'W).

MARINE RESEARCH IN AZORES 34 1

Lajes do Pic0 From the far west of Baia das Lajes do Pic0 (38" 24.7'N, 28" 16.7'W) to Ponta da Queimada (38" 23.2'N, 28" 13.4'W) are flat littoral platforms, with an extended intertidal area. These are not very common in the Azores. One is located close to the small whaling village of Lajes do Pic0 (south coast), and requires urgent designation as a Nature Reserve to preserve the habitats present which are being degradate due to human pressure. The bay at these location constitutes an interesting nursery for Pagrus pagrus and Epinephelus marginatus (Patzner, 1990; Azevedo, 1992).

Baia das Canas The coast from Cedros Ponta dos Misttrios (36" 29.9'N, 28" 15.1'W) to Farol da Prainha (38 " 28.3'N, 28" 11.9'W) on the north coast of Pic0 has mixed habitats of sand and rock. This area is rich in fish and invertebrates. Baia das Canas holds an important colony of Cory's shearwater.

Fuiul (Figure 10) Capelinhos This area was formed between 1957 and 1959, when a series of volcanic eruptions occurred in the far northwest of the island of Faial. The terrestrial area has already been designated as a Nature Reserve. This protection should be extended to the submarine surroundings of the volcano, not only for the conservation of the unusual habitats but also as a feeding and buffer area to benefit one of the most important colonies of common terns of Faial. It should be delimited by the coordinates of 38" 36.3'N, 28" 49.1'W and 38" 35.4", 28" 49.5'W. The area also holds an important colony of Cory's shearwater.

Morro de Castelo Branco The terrestrial nature reserve should also be extended to the surrounding sea in an area between 38" 31.9'N, 28" 45.1'W and 38" 31.2", 28" 44.2'W. There is an unusual cliff located at the south coast of Faial. It is an interesting site for breeding of seabirds. The shallow sea that surrounds the cliff is rich in fish. Dolphins come frequently to the Baia do Varadouro, on the west side of the cliff. In front of Morro de Castelo Branco is located a shallow bank with, apparently, an important population of a grouper, Mycteroperca fusca. An important colony of Cory's shearwater is also present.

Cedros/SalHo From Ponta dos Cedros (38" 38.6'N, 28" 43'W) to Ponta do SalLo (38" 37.6'N, 28" 39.6W) there is an exposed rocky coast on the north coast of the island. It is very rich in fish, crustacean and molluscs.

Feteira to Horta Harbour An area beginning at Feteira (38" 31.3'N, 28" 39.9'W) and finishing at the wall of Horta harbour (38" 31.4", 28" 37.4'W) and including the Protected Landscape of Monte da Guia and the Baia do Porto Pim, with its shallow sandy beach which is an important fish nursery. The Caldeiras do Monte da Guia are already classified as reserves, but the exterior cliffs of this old volcano also contains important and unique habitats, such as caves. At Feteira there is also an interesting intertidal platform with many rock-pools. The coast in between is highly irregular with many furnas (cells in the rock made by volcanic activity).

Western group These islands are the farthest west and north of the Azores on the western side of the Mid-Atlantic Ridge.

In general, levels of human predation on littoral species are much lower than in central and eastern groups.

Flora (Figure 11) Ponta Delgada to Santa Cruz This stretch of the northeast coast encompasses Ponta Ruiva (39" 29.8'N, 31" 09.7'W). Jlhtus de Alvara, IlhCu Garajau, Ilhtu Garajau, IlhCu Alagoa, Baixa do Moinho, Baixa Vermelha to Santa Cruz at 39"

R. S . SANTOS ET AL. 342

N

3930

CORVO D 5 km -

\ FLORES /

31' W

Figure 1 1 . Proposed marine protected areas for the western group of islands.

27.4", 31" 07.1'W. This, nearly 7 km, coastline consists of small islets and sinuous rocky coasts with a very diverse marine fauna. Being relatively close to the village of Santa Cruz, this area could attract underwater tourism.

This coastline with cliffs, numerous islets and stacks, gives shelter to the largest concentration of roseate terns in the eastern Atlantic, with up to 350 breeding pairs, and it holds more than 300 breeding pairs of the common tern (del Nevo et al., 1990). The tern colonies are located in small islets with easy access. Serious episodes of disturbance with complete desertion of the colonies have occurred recently. The area requires urgent designation. Recreational activities must be strictly controlled and wardening implemented.

Ponta dos Bredos to Ponta Lop0 Vaz Around lOkm of coastline on the southwest coast, from Ponta dos Bredos (39" 25.6", 31" 15.6'W) to Ponta Lop0 Vaz (39" 22.3'N, 31" 12'W) could be designated for the same reasons as previously. Being geographically opposed they would complement each other in cases of asymmetry of recruitment, due to climatic and oceanographic conditions, in sessile or semi-sessile organisms like Megabalunus tintinnabulum and Patella spp.. This coastline gives shelter to an important concentration of roseate terns, with up to 150 pairs (del Nevo et al., 1990).

Ilhtu da Gadelha and adjacent coast An interesting area for molluscs, crustaceans, fishes and seabirds occurs from Baixa Rosa (39" 29.1'N, 31" 15.3'W) to Ponta Delgada (39" 31.1" to 31 " 12'W), including the Ilhtu da Gadelha and other islets.


Coruo (Figure 11) The small island of Corvo with its low population of around 450 people presents an opportunity for the most radical approach to conservation: all the island should be designated as a protected area. Within the reserve, areas should be zoned for integral protection. The main function would be the protection of populations of the exploited species of molluscs and crustaceans. The following three represent important habitats of local populations of rocky invertebrates and fishes. Many cliffs of the coastline are considered important bird areas (Grimmet and Jones, 1989).

PBo de Aquar, Porto da Areia, Ponta Negra Delimited by the coordinates 39" 40.5'N, 31" 07'W and 39 40.1'N, 31" 06.6'W.

Ponta do Marco Delimited by the coordinates 39" 43'N, 31" 07'W and 39" 43.5'N, 31" 06.2'W.

Pedra do Atlas, Canto da Carneira Delimited by the coordinates 39" 43.2'N, 31" 05.6'W and 39" 42.5'N, 31" 04.8'W.

MARINE POLLUTION

Until now pollution has not represented a great threat to the marine environment in the Azores. The only oil spill known in the Azores region occurred in 1970, although recent spills in Madeira and Shetland emphasize the vulnerability of islands. According to the Times Atlas of the Oceans, the Azores are situated in one of the safest (in terms of oil and other sources of marine pollution) parts of the Atlantic ocean. Organotin pollution due to leachates from visiting yacht occurs in the harbour of Faial but is extremely localized (Spence et al., 1990). Some effects of the pollution on whelks (Thais haemostoma Linnaeus) occur but are restricted to the harbour itself. Similar effects may occur in the other major harbours on Terceira and SBo Miguel.

Locally derived pollution has not had much impact due mainly to the relatively little industrial development. However, during the last few years, sewage pollution has been increasing through discharges to sea. This problem mainly affects areas close to the main towns (Lobo, 1987). It is likely to be a greater public health problem than a major ecological impact-the collection of shellfish on the shore, very close to sewage pipes discharging to the mid-intertidal, has been observed at Vila Franca de Campo in S5o Miguel and, no doubt, may occur elsewhere.

Despite little industrial development, it is possible to detect some localized impact. Since most of the factories are located on the coast, effluents from these are discharged indirectly into the sea. One of the recent cases involved the tuna fish factory on Faial island, situated close to an important bay and to a Protected Landscape. Other effluents having localized impacts are wastes from agar-agar processing, on Terceira island.

Despite the remote geographical location of the Azores, levels of heavy metals and other chemicals within the marine environment of the Archipelago do not seem to differ significantly from levels observed in other areas of the North Atlantic. This is true for levels of heavy metals in seawater (Gardner, 1975; Mart and Niirnberg, 1986; Helmers et al., 1990), levels of mercury in large migratory fish (Monteiro and Lopes, 1990), levels of organochlorines and mercury in resident fish (Magalhiies and Barros, 1987; Monteiro et al., 1991), and levels of mercury in cephalopods (Monteiro et al., 1992) and seabirds (Monteiro et al., 1995).

A recent study (Monteiro et al., 1992) has shown, surprisingly, mercury levels five times higher in octopus from coastal locations under urban influence, such as harbours, compared with levels in remote sites. This is perhaps due to sewage disposal and sloppy disposal of used batteries, lamps and antifouling paints. However, the absolute values were considered low.


The Azores offer an unique opportunity for the measurement of the historical variations in heavy metal contamination within the North Atlantic marine environment, using seabirds as bioindicators. This study is currently underway and will add to the assessment of patterns of atmospheric deposition of metals in the northern hemisphere.

Depledge et al. (1992) have emphasized the importance of the Azores as a site for comparison with levels of contamination on the coasts of Europe. A limited suite of indicator species are available-particularly as mussels (Mytilus) are absent from the Azores. Candidate bioindicator species include Puchygrapsus spp., Octopus vulgaris, Chthamalus stellatus (Poli) and Patella aspera, all of which have mainland conspecifics. Enteropmorpha spp. and Fucus spiralis Linnaeus are two types of algae which may also be useful (see also Moore et al., 1995, Vedel and Depledge, 1995).

DISCUSSION

Further work

Much basic marine ecological, systematical and biogeographic research are required to place the Azores into its proper context in the northeast Atlantic. Some groups have been well studied (e.g. algae, fish, amphipods, decapods, tunicates) and biogeographical conclusions can be made; in other groups there are huge information gaps. Comprehensive catalogues of the flora and fauna would be immensely useful for future studies on the conservation of biodiversity in this area of the northeast Atlantic. There is also a need for further basic survey work to describe major habitats and communities (biotopes) throughout the Archipelago. This work needs to be extended beyond the limits of diving, using remote vehicles and submarines. Of particular interest are the various seamounts in the area. Some interesting communities surrounding volcanic springs and vents were recently discovered at around 1100 to 1700 m deep for the Lucky Strike (Langmuir et al., 1992, 1993; Gonqalves, 1994) and at 860-970m for the Menez Gwen (Desbruykres et af., 1994; GonGalves, 1994). The Azores also offer the best potential for studying both shallow and deep vents in the Mid-Atlantic Ridge. Shallow water hydrothermal vents are known on the Jog0 de Castro seamount (our observations) and at SZo Miguel (our observations).

The littoral zone is under extremely heavy pressure from subsistence, semi-commercial and recreational collectors, and also rod and line fishermen. The offshore banks and seamounts are now being increasingly exploited by line fishermen which will inevitably affect top predators. Snagged or lost gear and anchoring may also have impacts on benthic communities. Various regulations have been enacted to prevent the loss of some littoral shellfish in the Azores. This may have happened already in the Canaries with one type of limpet (Patella cundei candei) whose specific status is uncertain (see Christiaens, 1973). Certainly P . ferruginea is facing extinction in the Mediterranean. Given the cultural importance attached to ‘lapas’ in the Azores similar losses must be prevented by enforced regulations and littoral reserves. The same considerations apply to various species of lobsters and the large barnacles ( M . tintinnabulum). Biochemical genetic studies of limpets suggest that the Azorean population of P. candei can be considered sufficiently genetically different to be elevated to a specific status (Corte-Real e t al., 1992). Although clearly conspecific, P. aspera shows considerable genetic differentiation between the Azores, Madeira, the Canaries and mainland Europe. If these and similar species were acutely overexploited, then the chances of recolonization would be limited. The exception is the lobster, Scyllarides latus which has a teleplanic larva (Martins, 1985a,b). It could also be argued that there is a case for designating at least one or two seamounts as reserves. In effect the Formigas, which barely jut above the surface, only partially fulfil this need. Other suitable seamounts need to be described and proposed as potential submarine reserves. Although, given their commercial importance, it will be difficult to persuade the fishing community of the necessity. However, this measure could contribute to the conservation of exploited demersal species, and be of primary interest for the future of the demersal fisheries in the Azorean EEZ.


Another aspect deserving close attention is the potential impact of climate change. It is now well established that global climate is changing and that these changes will have impacts on the marine fauna and flora, both at global and regional scales (Peters and Lovejoy, 1992). It is probable that in the northeast Atlantic there will be an increase in the sea temperature (Southward et al., 1995). An increase of the temperature of coastal waters, even if only by a few degrees, will lead to the loss of many cold temperate forms which reach the southern limit of their distribution here. If in continental seas these losses may be followed by a concomitant increase of warm temperate and sub-tropical species, in the case of the most isolated Atlantic islands of the Azores, it is unlikely that losses of ‘northern forms’ will be followed by the concomitant substitution (or increase) of ‘southern forms’. Most adult coastal fishes have a limited capacity to cross the great extensions of deep Ocean that separates the islands from the nearest coastal zones. Recolonization would depend on larva dispersion. Given the distances involved, and the differences of larval biology of the coastal and benthic forms, it could be expected that the loss of Northern forms, would reduce the biodiversity in these islands. This would only be mitigated by colonization by species with great capacity for dispersal. There is a need for detailed investigation, both on historical data bases and on comparative and empirical research, to characterize the composition and variations of both coastal and pelagic assemblages. Monitoring programs should be established.

There is a profusion of legislation for conservation in the Azores. However, it needs to be implemented and enforced. It also needs to be respected. Therefore the most important way of promoting marine conservation in the Azores is by education-both formally in schools and informally for adults and visitors. Steps have been taken to increase awareness, such as a TV series produced about a joint Anglo-Portuguese Expedition in 1989 (Zlhas Vivus), exhibitions and lectures. Interest in promoting eco-tourism is growing. Whale-watching and a museum of whaling attract visitors in the former whaling town of Lajes do Pico. Realization is growing that marine life needs to be seen in the wild-as well as on display, and in restaurants. The Department of Environment and Tourism and the Department of Fisheries of the Regional Government of the Azores must continue to protect wildlife and provide its appreciation by a wider audience, both locally and through tourism.

ACKNOWLEDGEMENTS

This paper has benefited from the collaboration of Paul Cornelius (Natural History Museum, London), H. R. Martins, H. A. Isidro, J. M. Gonqalves, M. Juliano, J. A. Martins, J. G. Pereira, F. M. Porteiro, H. Rodrigues (Departamento de Oceanografia e Pescas, Universidade dos Aqores) and L. Gallagher (Gallagher Marine) and JosC Constincia (Museu Carlos Machado, Ponta Delgada) to whom we are deeply grateful. Manuela Juliano provided figures 3 and 4. Helen Rost Martins is specially acknowledged for critical revision of the manuscript. This paper has resulted indirectly from Expedition Aqores ’89. The authors also acknowledge the support by SRAP/DRP and SRTA/DRA (R. A. dos Aqores), JNICT/Cihcia (Lisboa) and The British Council (Lisboa).

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