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Patterns of Diversity of the North-Eastern Atlantic Blenniid Fish

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Patterns of Diversity of the North-Eastern Atlantic Blenniid Fish GEB244.fm Page 411 Tuesday, June 5, 2001 3:59 PM Global Ecology & Biogeography (2001) 10, 411–422 RESEARCH LETTER PatternsBlackwell Science, Ltd of diversity of the north-eastern Atlantic blenniid fish fauna (Pisces: Blenniidae) VÍTOR C. ALMADA, RUI F. OLIVEIRA, EMANUEL J. GONÇALVES, ARMANDO J. ALMEIDA†, RICARDO S. SANTOS‡ and PETER WIRTZ‡ Unidade de Investigação em Eco-Etologia, Instituto Superior de Psicologia Aplicada, Rua Jardim do Tabaco 34, 1149–041 Lisboa, Portugal, E-mail: [email protected] †Laboratório Marítimo da Guia — IMAR, Universidade de Lisboa, Estrada do Guincho, 2750 Cascais, Portugal; and ‡Departamento de Oceanografia e Pescas — IMAR, Universidade dos Açores, 9900 Horta, Portugal ABSTRACT formed by the islands of Ascension and St Helena. Within the north temperate group, the sub- This paper presents an analysis of the distribu- groups with higher similarities were: Azores and tional patterns of blenniids (Pisces: Blenniidae) Madeira, Canary Islands and Morocco, and the in the north-eastern Atlantic. Two peaks of Mediterranean and the Atlantic coast of the species diversity were found, both in terms of Iberian Peninsula. Based on affinity indices, the number of species and number of endemics: one probable directions of faunal flows were inferred. in the tropical African coast and another in the The tropical coast of Africa and the Mediter- Mediterranean Sea. A cluster analysis of similar- ranean emerged from this analysis as probable ity values (Jaccard coefficient) among the eastern speciation centres of the north-eastern Atlantic Atlantic zoogeographical areas, revealed the blenniid fauna. The Mediterranean may have following groups: a north temperate group, a also acted as a refuge during glacial periods. tropical group formed by the tropical African coast and Mauritania, another group formed by Key words Blenniid fishes, distributional patterns, the islands of Cape Verde, a south temperate group eastern Atlantic, Macaronesia, Mediterranean, (South Africa), and a southern Atlantic group speciation centres, species diversity, zoogeography. invasions, leading to a higher level of diversity in INTRODUCTION the Mediterranean area (Wirtz, 1978). The blennioids have served as models for differ- Zander (1980) emphasized the possible role of ent scenarios that deal with the evolution and the Macaronesian islands (i.e. Azores, Madeira and zoogeography of the ichthyofauna of the north- Canary Islands), especially the Azores, as sites eastern Atlantic and the Mediterranean (Zander, of speciation of eastern Atlantic blennioids, from 1973, 1980; Wirtz, 1978; Santos et al., 1995). which the new species would have subsequently In a study on the behaviour of Mediterranean migrated to the west European shores. tripterygiid fishes, Wirtz (1978) proposed that the Santos et al. (1995) emphasized the strong three species that currently occur in the Mediter- connections between the fish faunas of the Azores, ranean may be derived from a common Atlantic Madeira and Canary Islands. They discussed ancestor. According to this model, several inva- the possible oceanographic patterns of circulation sions of the Mediterranean by an Atlantic stock, that may explain the affinities between the faunas interspersed with periods of relative isolation, of the Macaronesian islands and those of the would have promoted speciation and ecological Mediterranean and the eastern Atlantic shores, segregation among descendants from previous despite the prevalence, at least in the Azores, of © 2001 Blackwell Science Ltd. http://www.blackwell-science.com/geb 411 GEB244.fm Page 412 Tuesday, June 5, 2001 3:59 PM 412 V. C. Almada et al. eastward-flowing currents. Similar considerations because it was felt that, due to its transitional concerning the marine fauna of the Azores have character, this division would provide a better been presented for other animal groups (e.g. gobiid picture of the relationships between the Mediter- fishes: Miller, 1984; Rissoidae and Anabathridae ranean and the north-eastern Atlantic faunas. molluscs: Gofas, 1990). Other areas, namely the north-west coast of In recent years, knowledge of the blenniid Africa, should also be further subdivided but the faunas of Macaronesia and the West African lack of adequate information made this unfeasible. coast has substantially progressed (Wirtz, 1980, The similarity among different areas was 1994; Dooley et al., 1985; Bath, 1990; Brito, 1991; assessed using a cluster analysis procedure. The Santos et al., 1997). Similar improvements have Jaccard coefficient of similarity was used and the occurred in relation to the shores of the Iberian resulting similarity matrix was analysed through Peninsula (Nieto & Alberto, 1990, 1993–94; Oliveira a sequential, agglomerative, hierarchical and et al., 1992; Almada et al., 1993; Gonçalves et al., nested clustering algorithm described by Sneath 1993). & Sokal (1973). The option used was In this paper, the distributional relationships (unweighted pair-group method, arithmetic aver- of the blenniids from the eastern Atlantic and the ages). This option uses a linkage criterion inter- Mediterranean are analysed, and the different mediate between the single and the complete hypotheses on the origin of the blenniid ichthy- ones, in which the decision to include a new ofauna of the north-eastern Atlantic are re-evaluated. element in a cluster is taken with reference to the arithmetic average of the similarity values of the elements already in the cluster. The PC METHODS program used was version 1.60 (©Applied The division of the main zoogeographic areas Biostatistics, Inc.). of the eastern Atlantic and the Mediterranean To further the analysis of the relationships shores considered in this study is summarized in between areas the following indices (XA and XB) Table 1. The database used in the present work were computed for each pair of areas (A and B): (Table 2) was compiled based on this division. The information on the fauna of the Black Sea XA = A∩B/A; XB = A∩B/B is also provided for completeness, although it was not subsequently analysed since it is an impover- where: A∩B is the number of species present ished subset of the Mediterranean fauna. in both areas, A is the total number of species Nomenclatural authorities for most species present in area A, and B is the total number of present in Table 2 can be found in Zander (1986) species present in area B. If, for a given pair of and Bath (1990). The species Parablennius sangui- areas, there was a marked flux of fauna in a given nolentus include populations ascribed to another direction, we would expect that the target area species (P. parvicornis) by some authors (Zander, should present a subset of the species present 1979; Bath, 1996). Since the populations of the in the source area rather than the reverse. Thus, Atlantic islands, namely those of the Azores, are an asymmetry between the two indices would be intermediate between those of the Mediterranean predicted with the smaller value occurring in the and the west coast of Africa (Santos, 1992), it is source area. not yet clear to us if the two species are valid, It is important to stress that we are dealing and the distributional limits of the two morphs with historical hypotheses that are not necessar- are also not yet clearly defined. A species that is ily in contradiction with other approaches to the referred to in the literature (Lipophrys sabry: Bath, problem of diversity, namely those that relate 1982) was not included since it has been shown area and diversity, or energy base and species to be a synonym of L. trigloides (Bath, 1996). richness. We are aware that our divisions do not Finally, the species Petroscirtes ancylodon was have the same area but, for this type of fish, area not included since it is a recent immigrant from is probably not a very relevant issue, since they the Red Sea (Goren & Galil, 1989). live in an almost linear band, the shoreline, a few The Atlantic shore of the Iberian Peninsula hundred metres wide at most. Our comparisons was subdivided in more detail than other areas refer to points located in different geographical © 2001 Blackwell Science Ltd, Global Ecology & Biogeography, 10, 411–422 GEB244.fm Page 413 Tuesday, June 5, 2001 3:59 PM Blenniid zoogeography in the Atlantic 413 Table 1 Definition of the zoogeographic areas considered in the present study and references used for the description of their blenniid fauna Area Zoogeographic Area borders References region (Briggs, 1974) Boreal Eastern Atlantic From the North Sea to the Wheeler, 1969, 1992 region western entrance of the English Channel Biscay Gulf Mediterranean- From the western entrance Regueiro et al., 1979; Atlantic region of the English Channel to Cape Ibañez & Motos, 1977; Finisterra (north-west Spain) Motos & Ibañez, 1979; Almeida & Ibañez, 1981; Ibañez et al., 1989 North Iberian Mediterranean- From Cape Finisterra to Cape Regueiro et al., 1979; Oliveira Peninsula Atlantic region Carvoeiro (west Portugal) et al., 1992; ICN, 1993 West Iberian Mediterranean- From Cape Carvoeiro to Cape Oliveira et al., 1992; ICN, 1993 Peninsula Atlantic region S. Vicente (south-west Nieto & Alberto, 1990, Portugal) 1993–94 South Iberian Mediterranean- From Cape S. Vicente to the Oliveira et al., 1992; Peninsula Atlantic region Strait of Gibraltar ICN, 1993 Western Mediterranean- From the Strait of Gibraltar to Patzner, 1984 Mediterranean Atlantic region the Strait of Messina (Sicily) Central and Mediterranean- From the Strait of Messina to Steinitz, 1950; Ben-Tuvia, Eastern Atlantic
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