AÇOREANA, 2014, Suplemento 10: 109-116

NEW RECORDS OF MARINE WOOD BORERS (: TEREDINIDAE AND : LIMNORIIDAE) FROM SÃO MIGUEL, AZORES, WITH A DISCUSSION OF SOME ASPECTS OF THEIR BIOGEOGRAPHY

Luísa M.S. Borges & Filipe O. Costa

Centre of Molecular and Environmental Biology (CBMA), Department of Biology, University of Minho, Campus de Gualtar, 4710-057 Braga, Portugal e-mail: [email protected] Helmhol -Zentrum Geesthacht, Zentrum für Material- und Küstenforschung GmbH, Max-Planck-Straße 1, 21502 Geesthacht, Deutschland/Germany

ABSTRACT Specimens of the morpho- Lyrodus pedicellatus (Bivalvia: Teredinidae) and Limnoria tripunctata and L. quadripunctata (Isopoda: Limnoridae) were extracted from wood samples collected during July 2012. Limnoria tripunctata and L. quadripunctata coexisted in the same wood piece, which is rare for limnoriids. All three species were recorded for the fi rst time from São Miguel waters and L. quadripunctata was also recorded for the fi rst time from the Azores. The recent recorded presence of these species in the Azores suggests a recent range expansion from southern Europe.

RESUMO Espécimes da morfo-espécie Lyrodus pedicellatus (Bivalvia: Teredinidae) e espécimes das espécies Limnoria tripunctata e L. quadripunctata (Isopoda: Limnoriidae) foram extraídos de pedaços de madeira coletados durante o Workshop Sabrina, Mosteiros, São Miguel, Açores. L. tripunctata e L. quadripuncta foram encontradas a coexistir no mesmo pedaço de madeira, o que é uma observação rara em limnorideos. Este é o primeiro registo para estas três espécies em São Miguel e também o primeiro registo da espécie L. quadripunctata nos Açores. A ocorrência destas três espécies nos Açores, coincidente com o aumento da actividade destes organismos na última década na Europa, sugere uma recente expansão com origem no Sul da Europa.

INTRODUCTION synonymised by Turner (1966) with (Linnaeus, 1758), Nototeredo arine wood borers are an norvagica (Spengler, 1792) and Mheteregeneous group of xylotrophic megotara (Hanley, 1848) and an isopod (wood feeding) organisms, including species, which was interpreted to be bivalves (Teredinidae [Distel, 2003]) and (Rathke, 1799) (Clapp & Crustacea (Limnoriidae [Cragg, 2003]). Kenk, 1963). They are considered serious pests due Since these early studies, a large to the destruction they infl ict on man- number of teredinids have been reported made wooden structures in the marine upon from European waters from cold environment (Turner & Johnson, 1971). Norway (Santhakumaran & Sneli, 1978; Indeed, economic and safety issues Borges, 2007) to the warm Mediterranean were the main motivation for the fi rst Sea (Bobat, 1995; Borges, 2007; Sivrikaya scientifi c studies of the species in the 18th et al., 2009). The large number of century, when outbreaks of wood borers publications describing new teredinid in the wooden dykes of the Netherlands species in the 18th, 19th and early 20th caused catastrophic destruction. Among centuries created many synonyms. the numerous reports produced on This situation left the systematics and wood borers at the time (Clapp & Kenk, of the Teredinidae in a chaotic 1963), the most important was that of state. One of the reasons for this was Sellius (1733) wherein he recognised probably the fact that teredinids are a three teredinid species, which were particularly diffi cult group of bivalves

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to identify. Whereas in the majority of respectively. The taxonomic status of the bivalves morphological identifi cation la er was, however, considered uncertain is based on shell features, in the case of (Cookson, 1990) but the species has now the Teredinidae, the shells show high been re-described (Castelló, 2011). intraspecifi c variation and interspecifi c Three teredinid species have been similarity and thus identifi cation is almost reported from the Azores in the early entirely based on the morphology of the literature. Nototeredo norvagica, from São pallets (Turner, 1966). In the catalogue Miguel, Faial (Dau enberg, 1889) and of the Teredinidae by Turner (1966), the Pico Islands (Nobre, 1924) and both N. taxonomy of the family was reviewed norvagica and were and six species were identifi ed from the reported as occurring commonly in Atlantic and the Mediterranean coasts of driftwood in the vicinity of the Azores Europe. In the last decade, however, the (Morton et al., 1998). In the last decade, number of teredinid species in Europe has although N. norvagica is still recorded increased to nine, including two invasive from São Miguel (Martins et al., 2009), the species (Borges et al., submi ed) and two species occurring in the Azores seems to cryptic species in the pedicellatus-like have changed. Teredora malleolus (Turton, Lyrodus complex of species (Borges et al., 1822) was reported for the fi rst time from 2012). São Miguel (Martins et al., 2009) and The Limnoridae includes wood and Lyrodus pedicellatus (Quatrefages, 1849) algae boring species. Of all the limnoriid was reported also for the fi rst time from species reported from Europe, fi ve are Terceira waters (Borges, 2007). In the case wood borers. Limnoria lignorum was of limnoriids, Limnoria tripunctata is the the fi rst limnoriid species described by only species recorded from the Azores, at Rathke (1799) as Cymothoa lignorum. Later Terceira (Borges, 2007). on, it was recognised by White (1857) as The malacological fauna of the Azores being synonymous with Limnoria terebrans has been the subject of several studies Leach, but subsequent authors referred to (Gofas, 1990; Morton, 1995; Ávila et al. the species as Limnoria lignorum, as it is now 1998, 2000; Martins et al., 2009). Similarly, known. For many years, it was thought there are a number of studies on the to be the only wood boring limnoriid fauna of the Azores (Paula et species, which led to several erroneous al., 1992; Southward, 1998; Costa & Jones, identifi cations (Neves, 1962). One and 2000; Rubal & Larsen, 2012; Torres et al., a half centuries later, a second wood 2012). No specifi c studies have, however, boring limnoriid, Limnoria quadripunctata been undertaken either on teredinids or Holthuis, 1949, was described from the on limnoriids, probably because, in the Netherlands. Soon after, a third species, Azores, contrarily to what happens in Limnoria tripunctata Menzies, 1951, was northern European countries, marine described from Southern California but wood borers do not pose a threat to was later found in southern Europe, in maritime structures, such as harbours, countries such as Portugal (Franco, 1962; pontoons and je ies, which are usually Neves, 1962) up to the English Channel constructed of concrete. These organisms (Eltringham & Hockley, 1958). The last are inconspicuous, due to the specifi c two limnoriid species, Limnoria borealis substrata they inhabit and thus they Kussakin, 1963 and Limnoria carinata might easily be missed in general surveys, Menzies & Becker, 1957 were reported which is probably the reason for the from Europe but only in Icelandic paucity of information in the literature. waters and in the Mediterranean Sea, The aim of the present work was, thus,

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to investigate the diversity of teredinids of the pleotelson (tubercles), according and limnoriids occurring in São Miguel to the keys in Menzies (1957) and the waters, as a contribution to knowledge of descriptions in Holthuis (1949). Complete the local fauna. specimens were preserved individually in 96% ethanol for future molecular analysis. MATERIALS AND METHODS RESULTS Wood pieces were collected during a dive at a depth of 10 metres at Aquário Twenty teredinid and twenty-nine (station SAB2011.DI3), Mosteiros, São limnoriid specimens were extracted Miguel, Azores (37º53´N; 25º48´W), in from the wood. All complete teredinid July 2011, during the ‘Sabrina’ Workshop. specimens were identifi ed as belonging to Limnoriid specimens were collected the morpho-species Lyrodus pedicellatus. from the wood surface, by carefully Some specimens were lacking pallets and spli ing the tunnels with forceps were identifi ed only to family level, as under a stereo dissecting microscope. without pallets further identifi cation is After the limnoriids were collected, the not possible. Limnoriid specimens were wood pieces were split up to extract the identifi ed as Limnoria quadripunctata and teredinids. In the majority of cases whole Limnoria tripunctata. Both species were specimens were removed but, in others, found co-existing in the same piece of the soft tissues had ro en and only pallets wood. Many specimens were identifi ed and shells were found. Morphological only to level, Limnoria sp., due identifi cations of teredinids were based to damage to the pleotelson tubercles. on pallet characters according to the keys All three species are reported for the in Turner (1971) and the illustrations and fi rst time from São Miguel and Limnoria descriptions in Turner (1966). In the case of quadripunctata is also reported for the fi rst limnoriids, morphological identifi cations time from the Azores. were based mainly on the ornamentations

FIGURE 1. Specimen of the morpho-species Lyrodus pedicellatus extracted from wood collected in São Miguel, Azores (A); detail of the pallets (B). bl, blade of pallet; cl, calcareous lining; pl, pallets; pt, periostracum of pallet; sp, siphon;st, stalk of pallet. Scale bars: A= 2 mm; B= 0.5 mm.

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FIGURE 2. Specimens of Limnoria quadripunctata (A) and Limnoria tripunctata (B). ml, Microfoliculina limnoriae (epibiotic heterotrich cilliate); plt, pleotelson; tb,tubercle. Scale bars= 0.5 mm.

Bivalvia: Teredinidae located on the median line, followed by a pair of tubercles; behind each of the Lyrodus pedicellatus (Quatrefages, 1849) posterior tubercles is a longitudinal ridge Diagnosis: pallets non-segmented, (Fig. 2B). composed of a single piece; calcareous portion of blade conical distally; upper DISCUSSION portion of calcareous blade (distal half) enveloped by a periostracal cap, In the present study, none of the which varies from light to dark brown, teredinid species reported upon by almost black; periostracal cap more or Martins et al. (2009) and Nobre (1924) less straight sided, with a distal margin were found. It is not, however, possible U-shaped, extending as lateral horns to ascertain, either from the present (Figs. 1A, B). study or from previous ones, if any of the reported species were or are established Isopoda: Limnoriidae in local waters. Both Lyrodus pedicellatus and Limnoria tripunctata were considered Limnoria quadripunctata Holthuis, 1949 to be established species in Terceira as Diagnosis: fi fth somite of pleon dorsally they were collected from test panels with an x-shaped carina on median line; exposed for a year in local waters. We do pleotelson with four sharp tubercles not know, however, whether the pieces on the median line; behind each of the of wood (present study) had been part posterior tubercles is a longitudinal of fi xed maritime structures or if they ridge which is more evident in female were driftwood and in previous studies specimens (Fig. 2A) teredinid shells were identifi ed from dredged sediment (Nobre, 1924; Martins Limnoria tripunctata Menzies, 1951 et al., 2009). To fi nd defi nitive evidence Diagnosis: dorsal surface of fi fth somite of established populations, therefore, of pleon with two elevated nodes located would require collecting specimens either anteriorly and a single posterior elevated from fi xed structures or from test panels area; pleotelson with an anterior tubercle exposed over a period of time.

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The occurrence of L. pedicellatus in now, the most common limnoriid species São Miguel seems to confi rm the trend (Borges, 2007). It was also reported for that this is the most common teredinid the fi rst time along the Atlantic coast occurring in southern Europe, occurring of France (Le Roux, 2009), in the Tagus as far north as the English Channel Estuary (Borges et al., 2010) and in Viana (Lebour, 1946; Morton, 1971). Indeed, do Castelo in 2009, northern Portugal this was the dominant species in test (Borges, unpublished data). panels exposed in Terceira and Algarve Of the two limnoriid species found (Borges, 2007) and in wooden piles in São Miguel, the most abundant was retrieved from the Tagus Estuary (Borges Limnoria tripunctata. This was expected, et al., 2010). Lyrodus pedicellatus is also as L. tripunctata is the most common the dominant teredinid species in the limnoriid species in southern Europe. Mediterranean, south of Turkey (Borges, Indeed, it is the most destructive wood 2007; Sivrikaya et al., 2009). A recent study boring species in southern Turkey, in the by Borges et al. (2012), using molecular south of Portugal and in Terceira (Borges, markers in addition to morphology, has 2007). In Portugal, this species is found as shown, however, that the Atlantic and far north as Aveiro (Borges, unpublished Mediterranean forms of L. pedicellatus data). The recent observed proliferation are in fact cryptic species. Hence, in of limnoriid activity in southern Europe, order to determine the probable origin which rivals that of teredinids, might of L. pedicellatus occurring in the Azores, be related to an increase in salinity we plan to obtain DNA sequences to (Einsenreich, 2005) as limnoriids appear compare with homologous sequences of to be restricted to waters with salinities L. pedicellatus from the Atlantic and the close to that of full seawater (Cragg et al., Mediterranean (Borges et al., 2012) and 1999). with GenBank sequences of specimens obtained from Florida (Distel et al., 2011). Biogeographic considerations on the Finding Limnoria quadripunctata and occurrence of wood borers in the Azores Limnoria tripunctata coexisting in the Although prevailing currents in the same piece of wood was unexpected. As Azores are from the Western Atlantic, far as we are aware, the coexistence of several studies have shown that limnoriids in the same piece of wood has molluscan species in the Islands have only been reported upon by Eltringham greater affi nity with European than & Hockley (1958) in Southampton, with Western Atlantic species (Wir , southern England. In the Tagus Estuary, 1998; Ávila, 2000; Sá-Pinto et al., 2008; L. quadripunctata and L. tripunctata co- Malaquias et al. 2009). It is probable occur in the same area, Porto Brandão, that Azorean specimens might be the but not in the same piece of wood eastern Atlantic form of L. pedicellatus and (Borges et al., 2010). Since its description have their origin in mainland Portugal. by Holthuis (1949), L. quadripunctata Indeed, a phylogeographic study on was known to occur only in the coastal species of Patella (: Gastropoda) waters of the Netherlands and England in Macaronesia and mainland Portugal (Eltringham & Hockley, 1958). In the last indicates that species of this genus, such decade, however, the distribution of L. as Patella aspera, in the Azores seem to quadriunctata seems to have expanded. have originated from mainland Portugal The species was reported for the fi rst populations (Sá-Pinto et al., 2008). time in Portsmouth, Bournemouth and Lyrodus pedicellatus is a long-term Swanage, southern England, where it is, brooder, thus dispersal of this species

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to the Azores was probably achieved by collecting the wood. We thank also Brian rafting adults in driftwood either carried Morton for the constructive comments by currents or in ballast water. Once the that benefi ted the manuscript. species is present in a certain area with suitable environmental conditions, which LITERATURE CITED is the case of the Azores (Borges et al., submi ed), the release of the larvae in ÁVILA, S.P., 2000. Shallow-water marine molluscs such a late stage of development certainly of the Azores: biogeographical relationships. maximises their chances of survival and Arquipélago. Life and Marine Sciences, Universidade dos Açores, Supplement 2 (Part se lement, when appropriate substrata A): 99-131. are available. It is, therefore, probable that ÁVILA, S.P., J.M.N. AZEVEDO, J.M. this species will become established in the GONÇALVES, J. FONTES & F. CARDIGOS, area. 1998. Checklist of the shallow-water marine The dispersal capability of limnoriids molluscs of the Azores: 1- Pico, Faial, Flores e is small compared to that of teredinids, Corvo. Açoreana, 8(4): 487-523. as they have direct development and ÁVILA, S.P., J.M.N. AZEVEDO, J.M. GONÇALVES, J. FONTES & F. CARDIGOS, reduced swimming capabilities (Cragg et 2000. Checklist of the shallow-water marine al., 1999). Mechanical transport by drift- molluscs of the Azores: 2- São Miguel Island. infested wood (Johnson, 1935) or chance Açoreana, 9(2): 139-173. introductions due to human activities BOBAT, A., 1995. Marine wood-borer test (Cragg et al., 1999) are, hence, still the with preservatives on the coast of Turkey. most probable mechanisms for spread of International Research Group on Wood limnoriid species to remote areas such Preservation, IRG/WP/95-10091. 7 pp. BORGES, L.M.S., 2007. Biogeography of wood boring as the Azores. Rafting, for instance, is organisms in European Coastal waters and new known to be of particular importance approaches to controlling borer a ack, xi+253 for many coastal organisms without pp. Unpublished PhD thesis, Portsmouth pelagic dispersal stages (Thiel & Gutow, University, Portsmouth, UK. 2005). In algae boring limnoriids, all life BORGES, L.M.S., A.A. VALENTE, P. PALMA & stages and both sexes have been found L. NUNES, 2010. Changes in the wood boring community in the Tagus Estuary: a case in fl oating holdfasts. This suggests that study. Marine Biodiversity Records, 3(e41): 1-7. limnoriids can reproduce successfully BORGES, L.M.S., H. SIVRIKAYA, A. LE ROUX, during rafting, facilitating long-distance J.R. SHIPWAY, S.M. CRAGG & F.O. COSTA, dispersal (Miranda & Thiel, 2008). 2012. Investigating the taxonomy and systematics of marine wood borers (Bivalvia: ACKNOWLEDGEMENTS Teredinidae) combining evidence from morphology, DNA barcodes and nuclear locus sequences. Invertebrate Systematics, 26: This work was supported by FEDER 572-582. through POFC-COMPETE and by national BORGES, L.M.S., L. MERCKELBACH, S.M. funds from “Fundação para a Ciência e a CRAGG & Í. SAMPAIO. Biogeography of Tecnologia (FCT)” in the scope of the grants bivalve wood borers in European coastal FCOMP010124FEDER-007381 and PEstC/ waters: using niche-based modelling to BIA/UI4050/2011. FOC benefi ed from predict the potential distribution of species. a Marie Curie European Reintegration Submi ed. CALLOWAY, C.B., & R.D. TURNER, 1983. Grant PERG02GA2007224890 provided Documentation and implications of rapid by the European Commission. We thank successive gametogenic cycles and broods the accommodation provided to LMSB in the Lyrodus fl oridanus (Bartsch) during the workshop by the organizing (Bivalvia, Teredinidae). Journal of Shellfi sh commi ee and Gonçalo Calado for Research, 3: 65-69.

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CALLOWAY, C.B., & R.D. TURNER, 21553, European Commission- Joint Research 1987. Species pairs in the Teredinidae. Centre, Ispra, Italy, 253 pp. International Research Group on Wood ELTRINGHAM, S.K., & A.R HOCKLEY, 1958. Preservation, IRG/WP/4142 . 2 pp. Coexistence of three species of the wood- CASTELLÓ, J., 2011. The genus Limnoria boring isopod Limnoria in Southampton (Limnoriidae, Isopoda, Crustacea) in Europe, water. Nature, 4624: 1659-1660. including a key to species. Zootaxa, 2968: 1-25. FRANCO S.E., 1962. Os ataques de xilófagos CLAPP, W.F., & R. KENK, 1963. Marine borers marinhos em estruturas de madeira an annotated bibliography, 1136 pp. Offi ce of submersas. Ensaios de durabilidade natural Naval Research Department of the Navy, de ensaios de madeira—II. Separata da Revista Washington D.C. Agronómica XLV—Tomo III. COOKSON, L.J., 1990. Annotated check-list of the GOFAS, S. 1990. The li oral Rissoidae and Limnoriidae. International Research Group on Anabathridae of São Miguel, Azores. In: MARTINS, A.M.F. (ed.), The marine fauna Wood Preservation, IRG/WP/4160. 11 pp. and fl ora of the Azores. Proceedings of the First COSTA, A.C., & M.B. JONES, 2000. The genus Internaltional Workshop of Malacology São Tesseropora (Cirripedia: Tetraclitidae) from Miguel, Azores. Açoreana, supplement [2]: 97- São Miguel, Azores. Arquipélago. Life and 134. Marine Sciences, Supplement 2 (Part A): 71-78. HOLTHUIS, L.B., 1949. The Isopoda and CRAGG, S.M., 2003. Marine wood boring Tanaidacea of the Netherlands, including : ecology, functional anatomy the description of a new species of Limnoria. and control measures. In: GOODELL, T.P., Zoologische Mededelingen, 30: 163-190. B. NICHOLAS & D.D. SCHULTZ (eds.), JOHNSON, M.W., 1935. Seasonal migrations of Wood deterioration and preservation: advances the wood-borer Limnoria lignorum (Rathke) in our changing world, pp 272-288. American at Friday Harbor, Washington. Biological Chemical Society, Washington D.C Bulletin, 69: 427-438. CRAGG, S.M., A.J. PITMAN & S.M. LE ROUX, A., 2009. Aperçu de la faune des HENDERSON, 1999. Developments in the xylophages marins du golfe du Morbihan. understanding of the biology of the marine Bulletin Société de Sciences Naturelle du Ouest de wood boring and in methods of la France, 31(2): 75-83. controlling them. International Biodeterioration LEBOUR, M.V., 1946. The species of Teredo and Biodegradation, 43: 197-205. from Plymouth waters. Journal of the Marine DAUTZENBERG, P., 1889. Contribution à la Biological Association of the United Kingdom, 26: faune malacologique des Îles Açores. Révision 381-389. des mollusques marins des Açores. Résultats MALAQUIAS, M.A.E., G.P. CALADO, V. des campagnes scientifi ques Albert 1er de PADULA, G. VILLANI & J.L CERVERA, Monaco, 112 pp, 4pls. 2009. Molluscan diversity in the North DISTEL, D.L., 2003. The biology of marine Atlantic Ocean: new records of opisthobranch gastropods from the Archipelago of the wood boring bivalves and their bacteria Azores. Marine Biodiversity Records, 2(e38): endosymbionts. In: GOODEL, T.P., B. 1-9. NICHOLAS & D.D. SCHULTZ (eds.), Wood MARTINS, A.M.F., J.P. BORGES, S.P. ÁVILA, deterioration and preservation: advances in A.C. COSTA, P. MADEIRA & B. MORTON, our changing world, pp 253-271. American 2009. Illustrated checklist of the infrali oral Chemical Society, Washington D.C. molluscs off Vila Franca do Campo. In: DISTEL, D.L., M. AMIN, A. BURGOYNE, E. MARTINS, A.M.F. (ed.), The marine fauna LINTON, G. MAMANGKEY, W. MORRILL, and fl ora of the Azores. Proceedings of the Third J. NOVE, N. WOOD & J. YANG, 2011. International Workshop of Malacology São Molecular phylogeny of Pholadoidea Miguel, Azores. Açoreana, supplement [6]: 15- Lamarck, 1809 supports a single origin for 103. xylotrophy (wood feeding) and xylotrophic MENZIES, R.J., 1957. The marine borer family bacterial endosymbiosis in Bivalvia. Molecular Limnoriidae (Crustacea, Isopoda). Part I: phylogenetics and evolution, 61: 245-54. northern and central America: systematics, EISENREICH S.J., 2005. Climate change and the distribution, and ecology. Bulletin of Marine European water dimension. EU report no Science of the Gulf and Caribbean, 7: 100-200.

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MENZIES, R.J., & G. BECKER, 1957. Holzzer- International Research Group on Wood störende Limnoria-Arten (Crustacea, Isopoda) Preservation, IRG/WP/435. 20 pp. aus dem Mi lmeer mit Neubeschreibung SELLIUS, G., 1733. Historia naturalis teredinis seu von L. carinata. Zeitschfi rt für Angewandte xylophagi marini tubulo-conchoidis speciatim Zoologie, 44: 85-92. Belgici, xxxiii+366 pp., 2 pls. Trajecti ad MIRANDA, L., & M. THIEL, 2008. Active and Rhenum. passive migration in boring isopods Limnoria SİVRIKAYA, H., S.M. CRAGG & L.M.S. BORGES, spp. (Crustacea, ) from kelp 2009. Variation in resistance to marine holdfasts. Journal of Sea Research, 60 (3): 176- borers in commercial timbers from Turkey, 183. as assessed by marine trial and laboratory. MORTON, B., 1971. The functional anatomy of Turkish Journal of Agriculture and Forestery 33: the organs of feeding and digestion of Teredo 569-576. navalis Linnaeus and Lyrodus pedicellatus SOUTHWARD, A.J., 1998. New observations (Quatrefages). Proceedings of the Malacological on barnacles (Crustacea: Cirripedia) of the Society of London, 39: 151-167. Azores region. Arquipélago. Life and Marine MORTON, B., 1995. The biology and functional Sciences, 16A: 11-27. morphology of Trichomusculus semigranatus SVAVARSSON, J., 1982. Limnoria borealis (Isopoda, (Bivalvia: Mytiloidea) from the Azores. In: Flabellifera) and its commensal, Caecijaera MARTINS, A.M.F. (ed.), The marine fauna borealis (Isopoda, Aselota), found in Icelandic and fl ora of the Azores. Proceedings of the waters. Sarsia, 67: 223-226. Second International Workshop of Malacology THIEL, M., & L. GUTOW, 2005. The ecology of and Marine Biology, Vila Franca do Campo, São rafting in the marine environment. II. The Miguel, Azores. Açoreana, Supplement [4]: 279- rafting organisms. Oceanography and Marine 295. Biology: an Annual Review, 43(1): 279-418. MORTON, B., J.C. BRITTON & A.M.F. MARTINS, TORRES, P., A.C. COSTA & M.A. DIONÍSIO, 1998. Ecologia costeira dos Açores, x+249 pp. 2012. New alien barnacle species in the Sociedade Afonso de Chaves, Ponta Delgada. Azores and some remarks on the invasive NEVES, C.M.B., 1962. Sobre a não existência potential of Balanidae. Helgoland Marine de Limnoria lignorum Rathke em Portugal. Research, 66: 513-522. Separata da Revista Agronómica, XLV (II). TURNER, R.D., 1966. A survey and illustrated NOBRE, A., 1924. Contribuições para a fauna catalogue of the Teredinidae, 256 pp. The dos Açores. Anais do Instituto de Zoologia da Museum of comparative Zoology, Harvard Universidade do Porto, 1: 41-90. University, Cambridge, M.A. PAULA, J., A. CARTAXANA & H. QUEIROGA, TURNER, R.D., 1971. Methods of identifi cation 1992. Decapod crustaceans collected by the of marine borers and fungi. In JONES, S.K., “Expedition Azores 1989”. Arquipélago. Life & G.E.B. ELTRINGHAM (eds.), Marine borers, and Marine Sciences, 10: 67-74. fungi and fouling organisms of wood, pp 18-63. RUBAL, M., & K. LARSEN, 2012. A new species Organisation for Economic Co-operation and of Ingolfi ellidae (Peracarida, Amphipoda, Development, Paris. Crustacea) from the Azores, Portugal. TURNER, R.D., & A.C. JOHNSON, 1971. Biology Helgoland Marine Research, doi:10.1007/ of marine wood boring molluscs. In: JONES, s10152-012-0311-6. S.K., & G.E.B., ELTRINGHAM (eds.), Marine SÁ-PINTO, A., M. BRANCO, D. SAYANDA borers, fungi and fouling organisms of wood, & P. ALEXANDRINO, 2008. Pa erns of pp. 259-296. Organisation for Economic Co- colonization, evolution and gene fl ow operation and Development, Paris. in species of the genus Patella in the TURTON, W., 1822. Conchylia inusularum: the shells Macaronesian Islands. Molecular ecology 17: of the British Islands, systematically arranged, 519-32. xvii+279+1 pp. London. SANTHAKUMARAN, L.N., & J.-A. SNELI, WHITE A., 1857. A popular history of British 1978. Natural resistance of diff erent species Crustacea, 358 pp. L. Reeve (ed.), London of timber to marine borer a ack in the WIRTZ, P., 1998. Opisthobranch molluscs from Trondheims ord (Western Norway). the Azores. Vita Marina, 45(1-2): 1-6.

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