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Mediterranean Bioconstructions Along the Italian Coast        83


              2.2.8 Serpulid Reefs, Including Biostalactites
              Serpulidae are a large family of polychaetes that produce a calcareous tube in
              which they live (Bianchi, 1981). Tubes typically encrust any hard substrate
              and may be considered “secondary framework builders”, filling crevices and
              cementing rubble in coralligenous reefs and other marine bioconstructions
              (Bianchi et al., 1995). The scissiparous species Salmacina dysteri (Huxley,
              1855) may form large “pseudocolonies”, which are nevertheless very fragile
              and do not exhibit any significant constructional capacity (Bianchi and
              Morri, 2000). In contrast, some gregarious species can grow erect attached
              to each other, thus acting as “primary frame builders” and forming huge
              aggregates (Aliani et al., 1996). The individual tubes are small (rarely longer
              than 10cm or wider than 1cm), but their aggregations can form reef-like
              structures more than 1m thick that may cover tens of square metres
              (Bianchi et al., 1995). Serpulid reefs are well known in the geological past
              (Fagerstrom, 1987), and mass occurrences of recent serpulid species have
              been reported from many sites (Ten Hove, 1979). In Italy, major serpulid
              reefs, formed by Ficopomatus enigmaticus (Fauvel, 1923) or Hydroides dianthus
              (Verrill, 1873), have been described from brackish lagoons (Bianchi and
              Morri, 1996, 2001), but these will not be further discussed here.
                 In fully marine waters, serpulids may develop conspicuous assemblages
              in submarine caves, even encrusting preexisting speleothemes, as first
              described for the Mediterranean by Ten Hove and Van Den Hurk (1993)
              and in Italy by Bianchi and Morri (1994) and Antonioli et al. (2001).
              Particular serpulid structures in submarine caves are the so called bio-
              stalactites. They develop in completely dark parts based on single (e.g. Cape
              of Otranto) or few (e.g. Plemmirio Islands) (Fig. 7C) serpulid species,
              whose aggregations become substrate for smaller invertebrates and bacteria
              (Belmonte et al., 2009; Guido et al., 2012). Biostalactites grow far from the
              entrance on rocky walls where generally Serpulidae dominate (Bussotti
              et al., 2006) and where other benthic species are less competitive. Bio-
              stalactites can protrude a few cm from the basal substrate (Plemmirio) but
              also up to 2m (Cape of Otranto, Fig. 7A) and they can be also oblique or
              horizontal. Freshwater inflows (even intermittent) seem to favour their
              formation. The engineering species always have been referred to the genus
              Protula, but recent genetic analyses of the Otranto species showed the
              existence of a still unnamed species. The growth of Otranto biostalactites
              started about 6000 years ago, corresponding with the sea level during its
              postglacial rise (Belmonte et al., 2009, 2016). The outer surfaces of the
              biostalactites are encrusted by living and dead fauna of several invertebrate
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