Biogeographical Homogeneity in the Eastern Mediterranean Sea. II
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Vol. 19: 75–84, 2013 AQUATIC BIOLOGY Published online September 4 doi: 10.3354/ab00521 Aquat Biol Biogeographical homogeneity in the eastern Mediterranean Sea. II. Temporal variation in Lebanese bivalve biota Fabio Crocetta1,*, Ghazi Bitar2, Helmut Zibrowius3, Marco Oliverio4 1Stazione Zoologica Anton Dohrn, Villa Comunale, 80121, Napoli, Italy 2Department of Natural Sciences, Faculty of Sciences, Lebanese University, Hadath, Lebanon 3Le Corbusier 644, 280 Boulevard Michelet, 13008 Marseille, France 4Dipartimento di Biologia e Biotecnologie ‘Charles Darwin’, University of Rome ‘La Sapienza’, Viale dell’Università 32, 00185 Roma, Italy ABSTRACT: Lebanon (eastern Mediterranean Sea) is an area of particular biogeographic signifi- cance for studying the structure of eastern Mediterranean marine biodiversity and its recent changes. Based on literature records and original samples, we review here the knowledge of the Lebanese marine bivalve biota, tracing its changes during the last 170 yr. The updated checklist of bivalves of Lebanon yielded a total of 114 species (96 native and 18 alien taxa), accounting for ca. 26.5% of the known Mediterranean Bivalvia and thus representing a particularly poor fauna. Analysis of the 21 taxa historically described on Lebanese material only yielded 2 available names. Records of 24 species are new for the Lebanese fauna, and Lioberus ligneus is also a new record for the Mediterranean Sea. Comparisons between molluscan records by past (before 1950) and modern (after 1950) authors revealed temporal variations and qualitative modifications of the Lebanese bivalve fauna, mostly affected by the introduction of Erythraean species. The rate of recording of new alien species (evaluated in decades) revealed later first local arrivals (after 1900) than those observed for other eastern Mediterranean shores, while the peak in records in conjunc- tion with our samplings (1991 to 2010) emphasizes the need for increased field work to monitor their arrival and establishment. Finally, the scarce presence (or total absence) in the most recent samples of some once common habitat-forming species, as well as of some other native taxa, con- firmed their recent rarefaction (or local extinction), possibly related to their replacement by the aliens Brachidontes pharaonis, Spondylus spinosus and Chama pacifica. KEY WORDS: Mediterranean Sea · Lebanon · Mollusca · Bivalvia · Alien species · Faunal changes Resale or republication not permitted without written consent of the publisher INTRODUCTION the basin gave rise to the peculiar and variegate Medi - terranean assemblage, with the co-occurrence of tem- The present-day autochthonous Mediterranean perate and subtropical organisms and a main transi- marine fauna is mostly of Atlantic origin, having tional zone between the 2 seas (the Alboran Sea), originated with the re-establishment of the Atlanto- hosting a mix of Mediterranean and Atlantic species Mediterranean connection (5.33 million years ago), (Bianchi & Morri 2000, Oliverio 2003, Coll et al. 2010). after the Messinian Salinity Crisis (from 5.971 to 5.33 The opening of the Suez Canal in the south-eastern million years ago; Manzi et al. 2013) had probably corner of the Mediterranean in 1869 contributed nearly exterminated the stenoecious marine biota (Sa - to changes in the local biodiversity and provided belli & Taviani in press). The subsequent 5 million years an additional human-induced transitional zone. The of evolution within the framework of the complexity of spreading of alien species (used here as defined by *Email: [email protected] © Inter-Research 2013 · www.int-res.com 76 Aquat Biol 19: 75–84, 2013 the International Union for Conservation of Nature, Shiber & Shatila 1978, Shiber 1980, Zibrowius & Bitar see Crocetta 2012) has become one of the main local 1981, 2003, Bogi & Khairallah 1987, Bitar 1996, 2010, phenomena affecting species distribution and com - 2013, Bitar & Kouli-Bitar 1995a,b, 1998, 2001, Nakhlé position within assemblages (Por 1978, Galil 2009). et al. 2006, Bitar et al. 2007, Bariche 2012, Crocetta Mainly confined for over a century along the Levant & Russo 2013). However, wide-ranging overviews Sea, alien species of thermophilic origin are cur- of the temporal trends and the long-term faunal rently spreading further to the western and northern changes occurring in the area are still lacking. Mediterranean, presumably favoured by the general Using material obtained on field trips within the warming of the area (Occhipinti-Ambrogi 2007, Rait- Programme CEDRE (French-Lebanese cooperation) sos et al. 2010). Present-day local biodiversity, which from 1999 to 2002, enriched by some additions from is currently threatened by habitat loss and degrada- earlier and later years, this study is the most com- tion, fishing impacts, pollution, climate change and plete overview of the marine Bivalvia of Lebanon. It eu trophi cation, is also impacted by alien introduction aims at: (1) providing an updated checklist of marine through human activities (e.g. aquaculture, shipping bivalve species recorded in Lebanon; (2) investigat- and leisure boating). Prevention and/or control of ing historical taxon names based on Lebanese type alien species diffusion have therefore become a major material and checking their availability as a contri- challenge for conservationists (Galil 2009, Zenetos et bution to Mediterranean bivalve taxonomy; (3) focus- al. 2012), and the cumulative impact of these biotic ing on alien bivalves in Lebanon, providing a check- and abiotic changes is resulting in the disruption of list of published and unpublished records, with date the original bio geographic pattern within the basin, of the first record, most plausible vector(s) of intro- with several authors drawing attention to the pro- duction and the establishment status; and (4) tracing gressive homo genization of Mediterranean marine temporal variations and qualitative modifications of biota (Bianchi 2007, Lejeusne et al. 2010, Philippart the bivalve fauna composition in the framework of et al. 2011). current biological pressures. The Mediterranean molluscan fauna is considered as the best known in the world, and numerous de- tailed taxonomic inventories now exist, most of which MATERIALS AND METHODS are specific to sub-regions or countries (Oliverio 2003, Coll et al. 2010). Mollusca, as a ‘popular’ and thus fre- Study area quently investigated group, comprise the highest number of known introduced species in the Mediter- Lebanon is an area of particular significance for the ranean. There are now an estimated ~200 introduced study of marine biodiversity and its recent changes. species, more than half of which are with established Lying at the north-eastern tip of the Mediterranean populations, indicating a high rate of introduction and Sea, ~450 km north of the Suez Canal, it is located accounting for ~10% of the Mediter ran ean Mollusca along the natural pathways of Indo-Pacific taxa spread- (slightly more than 2000 species) (Coll et al. 2010, ing from the Red Sea via the prevailing Mediterranean Zenetos et al. 2012). Despite continuous efforts to pro- currents (Bergamasco & Malanotte-Rizzoli 2010). Al- vide updated data sets for the entire basin, our knowl- though the main coastal features and the respective edge of the Levantine area still remains considerably communities are similar to those observed along all the poor when compared with the central and western Mediterranean shores (Bitar et al. 2007, Bitar 2010), basin. This is probably due to undersampling due to several major engineering species are lacking, such the absence of recent faunal projects. as the seagrass Posidonia oceanica (Linnaeus) Delile, Our knowledge of Lebanese marine fauna has a 1813 and the sea fans Paramuricea clavata (Risso, 1826) similar history to the rest of the Levant basin. Early pi- and Eunicella spp. (Harmelin et al. 2009). oneering papers provided accurate data on shelled molluscs from 1856 to 1938, while the following decades were rather unfertile until 1971. From then Bibliographic data up until 2013, less than 30 papers, notes, abstracts and non-peer-reviewed articles have provided additional An extensive literature survey was conducted using scattered information on Lebanese marine bivalve the same methods as Crocetta et al. (2013). Indexed molluscs (Puton 1856, Brusina 1879, Pallary 1911, papers were searched, but an attempt to cover the 1912a, 1919, 1933, 1938, Gruvel & Moazzo 1929, Gru- grey literature as much as pos sible (i.e. non-peer- vel 1931, Moazzo 1931, Spada 1971, Fadlallah 1975, reviewed and/or non-indexed papers) was also per- Crocetta et al.: Marine bivalves in Lebanon 77 formed. Most of the historical journals are not in- Table 1. Localities where bivalve species were found, with dexed, and many malacological records are still being co ordinates and depth ranges (see also Fig. 1). Details of bio - topes from which the collections were made are reported in published in non-indexed journals, thus allowing only the Supplement under each species manual searching. In addition, the availa bility of all historical taxon names based on Lebanese type mate- No. Site Latitude Longitude Depth rial was checked. (°N) (°E) (m) 1 Ramkine Island 34° 29’ 47’’ 35° 45’ 38’’ 0−15 Sampling 2 Tripoli 34° 27’ 28’’ 35° 49’ 34’’ 0−5 3 Anfeh 34° 21’ 43’’ 35° 43’ 36’’ 10−24 In order to evaluate the current molluscan fauna, 4 Chekka 34° 19’ 12’’ 35° 43’ 14’’ 3−6 the taxonomic composition was the main target of in- 5 El Heri 34° 18’ 37’’ 35° 41’ 51’’ 1−5 terest. Several sampling localities, covering more or 6 Ras El Chakaa 34° 18’ 47’’ 35° 40’ 59’’ 0−20 less all of the Lebanese shores, were investigated by 2 7 Chak El Hatab 34° 17’ 36’’ 35° 40’ 17’’ 9−14 of the authors (G. Bitar and H. Zibrowius) between 8 Selaata 34° 17’ 03’’ 35° 39’ 31’’ 0−35 1999 and 2002 within the framework of the French- 9 Batrou 34° 15’ 13’’ 35° 39’ 19’’ 2−9 Lebanese Programme CEDRE (see Zibrowius & Bitar 10 Kfar Abida 34° 14’ 02’’ 35° 39’ 15’’ 1−12 11 El Barbara 34° 11’ 32’’ 35° 37’ 19’’ 26−28 2003, Morri et al.