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Rhabditophora, Proseriata) in the Mediterranean

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Rhabditophora, Proseriata) in the Mediterranean UNIVERSITY OF SASSARI DISSERTATION FOR THE DEGREE OF DOCTOR OF PHILOSOPHY IN ENVIROMENTAL BIOLOGY UNIVERSITY OF SASSARI, 2008 Contribution to the knowledge of the family Otoplanidae Hallez, 1892 (Rhabditophora, Proseriata) in the Mediterranean Candidate: Valentina Delogu Tutor: Prof. Marco Curini Galletti Supervisor : Prof. Marco Apollonio UNIVERSITÀ DEGLI STUDI DI SASSARI TESI DI DOTTORATO IN BIOLOGIA AMBIENTALE UNIVERSITÀ DEGLI STUDI DI SASSARI, 2008 Contributo alla conoscenza della famiglia Otoplanidae Hallez, 1892 (Rhabditophora, Proseriata) nel Mediterraneo Candidata: Valentina Delogu Tutor: Prof. Marco Curini Galletti Supervisore : Prof. Marco Apollonio TABLE OF CONTENTS Abstract page 4 Introduction page 5 Chapter I - New species of the genus Parotoplana Meixner, 1938 (Proseriata, Otoplanidae) from southern Apulia (Italy). page 23 Chapter II - The genera Parotoplana Meixner, 1938 and Parotoplanella Ax, 1956 (Platyhelminthes: Proseriata) in southern Spain. page 40 Chapter III - Otoplanidae (Platyhelminthes, Proseriata) from the northern Adriatic sea page 61 Chapter IV - A Mediterranean radiation of interstitial flatworms: the Parotoplana jondelii (Platyhelminthes: Proseriata) species group. page 76 Italian Summary page 96 Acknowledgments page 99 4 Abstract A growing awareness is rising that present perception of marine biodiversity is inadequate, and based upon unrepresentative data. The inadequacy of knowledge on taxonomy and distribution of marine organisms is particularly acute for interstitial meiofauna, especially for ‘soft bodied’ taxa, which necessitate of observations on both living and fixed specimens. Among these groups, information on Platyhelminthes, a species-rich, ubiquitous and diverse taxon, is at present far from adequate, and is entirely absent for entire biogeographical regions. In this scenario, my research aimed to gather data in the composition and distribution of a family of Platyhelminthes Proseriata, the Otoplanidae Hallez, 1892, in biogeographically significant sectors along the coasts of Mediterranean. The most striking results of a series of sampling campaigns is the great number of new Otoplanidae species found. In this work fifteen new species, belonging to the genus Parotoplana Meixner, 1938 are formally described: P. pythagorae Delogu & Curini-Galletti, 2007; P. spathifera Delogu & Curini-Galletti, 2007; P. terpsichore Delogu & Curini-Galletti, 2007; P. jondelii Delogu & Curini-Galletti, 2007; P. cucullata Delogu et al., 2008; P. fretigaditani Delogu et al., 2008; P. varispinosa Delogu et al., 2008; P. mastigophora Delogu et al., 2008; P. geminispina sp. n., P. crassispina sp. n., P. carthagoensis sp. n., P. ichnusae sp. n., P. pulchrispina sp. n., P. obtusispina sp. n. and P. axii sp. n. Valentina Delogu "Contribution to the knowledge of the family Otoplanidae Hallez, 1892 (Rhabditophora, Proseriata) in the Mediterranean" Tesi di Dottorato in Biologia Ambientale, Università degli Studi di Sassari 5 Introduction Following the United Nations Conference on Environment and Development (UNCED) held at Rio de Janeiro in 1992, the value of biological diversity has been recognised also by governments and numerous national and international plans were launched to investigate global biodiversity both in terrestrial and marine environment. Traditionally marine environment is considered a low specific diversity habitat but at present a growing awareness is rising that this perception of marine biodiversity is inadequate and based upon unrepresentative data (Bouchet et al. 2002, Sala 2002). The inadequacy of data on taxonomy and distribution of marine organisms is particularly acute for interstitial meiofauna, where information may be absent altogether for entire biogeographical regions (Kennedy and Jacoby 1999). The lack of data about interstitial meiofauna is particularly worrisome if we consider in terrestrial habitat, the highest values of species diversity falls in the size class of 1-2 mm (Siemann et al. 1996) which, in the sea, is mostly represented by meiofaunal taxa. Knowledge of taxa belonging to the so-called “soft bodied” meiofauna, which necessitate observations on both living and fixed specimens, is particularly limited. Among meiofaunal taxa Platyhelminthes are most important for their contribution both in number of species and individuals. Their abundance is generally related to the granulometry of the sediments; they are well represented in sandy sediments (7 - 25% of total meiofauna in density), where their diversity, as well as the biomass, may reach and even exceed that of Nematodes. Moreover in high-coarse grained sediments, Platyhelminthes may represent the 95% of total meiofauna (Remane 1933, Radziejewska & Stankowska-Radziun 1979, Martens & Schockaert 1986). Information on Platyhelminthes is at present far from adequate and it is considered one of the taxa (as well as Polychaeta, Olygochaeta, and Nematoda), that especially needs new identification guides and revised taxonomic lists (Costello et al. 2006) . In fact, only few areas are relatively well known, especially northern Europe and the Galápagos Islands (Ax & Schmidt 1973, Armonies & Reise 2000), with little and taxonomically limited research elsewhere. Recent evident suggest that marine-interstitial flatworms have limited geographical ranges (Casu & Curini-Galletti 2006) presumably linked to their lack of dispersal stage. For this reason these are the groups where modification in ranges and species composition, caused by phenomena of global change, could be more easily detectable. However, the absence of basic knowledge of most taxa for entire biogeographical regions has always prevented them utilisation for environmental monitoring. Furthermore the poor state of taxonomic knowledge may lead species to undetected cases of “silent extinction”. 1 Valentina Delogu "Contribution to the knowledge of the family Otoplanidae Hallez, 1892 (Rhabditophora, Proseriata) in the Mediterranean" Tesi di Dottorato in Biologia Ambientale, Università degli Studi di Sassari 6 Among Platyhelminthes, Proseriata (Neoophora), represent a taxon of high interest. Proseriata is a diverse and species-rich taxon of small (1-10 mm long), mostly free-living Platyhelminthes. They are particularly common in coastal habitats, from shallow water to the upper intertidal (Sopott 1972, 1973; Reise 1984, 1988). In some cases, they may be among the dominating organisms of interstitial assemblages, e.g., the Otoplanidae in the high-energy intertidal zone (Remane 1933). Many proseriates occupy narrow ecological niches. As a consequence, they show low α-diversity, i.e., in any given locality, each sediment type usually harbours a limited number of species. However larger scale data of proseriates suggest that γ- diversity of the group may be particularly high. In fact, different marine areas harbour quite distinct proseriate faunas. In Europe, North Atlantic has only the 7.5% of species in common with the Mediterranean (Curini-Galletti 2001), on the other hand among prosobranch gastropods, a group renowned as good biogeographical descriptors (Briggs 1974), shared species constitute about 18% of the mediterranean species (Poppe & Goto 1991). This implies that proseriates may experience physical and climate barriers to distribution to a finer degree than most macrofaunal taxa, resulting in “small-scale” biogeography, and ample opportunities for allopatric speciation. In this scenario, my research aimed to gather data in the composition and distribution of a family of Proseriata, the Otoplanidae Hallez, 1892, in the Mediterranean. The family Otoplanidae is the dominant taxon in the surf zone of the high energy beaches (the “Otoplana-zone” of Remane, 1933) (Reise 1988; Curini-Galletti 2001). It is known from marine environments of the North Atlantic (and adjacent seas: North Sea and Baltic Sea)(Ax 1956, 1959; Ax & Sopott-Ehlers 1987; Karling 1973; Giard 1904; Graff 1913; Luther 1960; Martens & Schockaert 1981; Meixner 1938; Sopott 1972; Sopott-Ehlers 1976, 1985; Sopott-Ehlers & Ehlers 1980), the Pacific Ocean (Karling 1964; Ax & Ax 1967; Tajika 1938a-b-c, 1984), the Atlantic coast of South America (Marcus 1949, 1950, 1952), the Galápagos Islands (Ax & Ax 1974), and the eastern Australia (Miller & Faubel 2003). Only five species has been reported from the freshwater or brackish-water environments (Riemann 1965; Gieysztor 1938; Ax 1951; Ax & Armonies 1990; Noreña et al., 2005). In Mediterranean only few areas, the Gulf of Lion, the Gulf of Naples and the Tuscany coast, has been studied ( Du Plessis, 1889; Ax 1956; Ax, Weidemann & Ehlers 1978; Lanfranchi 1969, 1978). Recently only two new species (Otoplana oxyspina Lanfranchi & Melai 2007; Parotoplana rosignana Lanfranchi & Melai 2008) have been reported for Mediterranean (Lanfranchi & Melai 2007, 2008), excluding my contributions. 2 Valentina Delogu "Contribution to the knowledge of the family Otoplanidae Hallez, 1892 (Rhabditophora, Proseriata) in the Mediterranean" Tesi di Dottorato in Biologia Ambientale, Università degli Studi di Sassari 7 During the three years of the Doctorate, a series of sampling campaigns has been undertaken, aimed to gather data on species composition and distribution of Proseriata, in biogeographically significant sectors along the coasts of Mediterranean and, particularly, I have dealt with Otoplanidae species. The sampling areas were been: 1 Porto Cesareo – Italy, Ionian sea; 2 Capo Caccia-I.la Piana – Italy, Tyrrhenian sea; 3 Tuscany coasts – Italy, Tyrrhenian sea; 4 Miramare-Gulf
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