Shallow-water and not deep-sea as most plausible origin for cave-dwelling Paramisophria species (Copepoda: Calanoida: Arietellidae), with description of three new species from Mediterranean bathyal hyperbenthos and littoral caves

Japan (Tanaka, 1967; Ohtsuka, 1985; Ohtsuka & Mitsuzumi, 1990; Ohtsuka et al., 1991), Australia (McKinnon & Kimmerer, 1985), and New Zealand (Othman & Greenwood, 1992). Records of species far from their ordinary ranges, such as those of Paramisophria cluthae T. Scott, 1897 from the Mediterranean (Sars, 1924-1925) and the western Pacific (Tanaka, 1967), or that of P. ammophila Fosshagen, 1968 from the Mediterranean (Carola et al., 1995), should be considered with caution since they are based on females only, and the major diagnostic characters at the species level rely on male morphology. In this paper, we describe three new species of Paramisophria from the Spanish zone of the Mediterranean, where they apparently follow an allopatric depth zonation pattern extending from the upper littoral to depths in excess of 1200 m. Additionally, the discovery of deep-water species has provided new evidence relevant to the deep-sea / shallow-water controversy on the origin of anchialine cave faunas (see Stock, 1986) from our analysis of the phylogenetic relationships between Paramisophria species and of the type of habitat exploited by each.