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A new cold-water coral (CWC) province has been identified in the Mediterranean Sea in the Capo Spartivento canyon system offshore the southern coast of Sardinia. The ‘Sardinia cold-water coral province’ is characterized in the Nora canyon by a spectacular coral growth dominated by the branching scleractinian Madrepora oculata at a depth of 380–460 m. The general biohermal frame is strengthened by the common occurrence of the solitary scleractinian Desmophyllum dianthus and the occasional presence of Lophelia pertusa. As documented by Remotely Operated Vehicle survey, this area is a hotspot of megafaunal diversity hosting among other also live specimens of the deep oyster Neopycnodonte zibrowii. The new coral province is located between the central Mediterranean CWC provinces (Bari Canyon, Santa Maria di Leuca, South Malta) and the western and northern ones (Melilla, Catalan-Provencal-Ligurian canyons). As for all the best developed CWC situations in the present Mediterranean Sea, the new Sardinian province is clearly influenced by Levantine Intermediate Water which appears to be a main driver for CWC distribution and viability in this basin.

Temnothorax lagrecai (Baroni Urbani, 1964) Figs 16–33 Diagnostic character combination Antennal clubs concolorous yellowish, antennae 12-segmented in females and 13 in males, eyes normal, metanotal groove very weak or absent, worker propodeal spines long, male propodeal spines dentiform, petiole upper profile usually with a short horizontal component, subpetiolar process toothlike, sculpturing mostly areolate and weak. Etymology Baroni Urbani (1964) dedicated this species to the Italian entomologist Marcello La Greca (1914–2001). Material examined We investigated 56 colony samples from 23 localities, consisting in a total of 212 ☿☿, 8 ♀♀, 2 ♂♂ from our collections, one colony under rearing in our possession, holotype queen and paratype workers from the MSNV and NHMW collections, and additional non-type workers from the personal collections of David Misfud and Stephen Schembri and from the MSNM. A detailed list is provided in the Supp. file 1: Table S1. Redescription Worker (Figs 19–27) MEASUREMENTS AND INDICES (13 specimens, 6 localities, 6 colonies). CL: 547.15 ± 33.51 (488–592); CW: 447.54 ± 31.37 (401–495); CS: 497.35 ± 31.84 (444–543); PoOC: 231.69 ± 14.9 (212–257); SL: 387.77 ± 24.95 (352–435); ML: 603.15 ± 57.6 (527–709); MW: 295.92 ± 26.65 (260–347); EL: 116.85 ± 16.1 (100–153); EW: 87.92 ± 11.19 (78–112); EYE: 102.38 ± 13.28 (89–132); SPST: 147.6 ± 17.97 (120–175); CL/CW: 1.22 ± 0.03 (1.18–1.29); PoOC/CL: 0.42 ± 0.01 (0.40–0.45); SL/CL: 0.71 ± 0.01 (0.68–0.73); ML/CS: 1.21 ± 0.05 (1.15–1.33); MW/CS: 0.59 ± 0.02 (0.56–0.65); EL/CS: 0.23 ± 0.02 (0.20–0.29); EW/CS: 0.18 ± 0.02 (0.15–0.21); EYE/CS: 0.20 ± 0.01 (0.18–0.25); SPST/CS: 0.30 ± 0.02 (0.27–0.32). COLORATION. Entirely yellowish with the exception of a variably large and often not clearly demarcated black transverse band on the gaster. HEAD. Subrectangular with rounded margins near the four angles, clypeus and mandibles rounded. Antennae of 12 segments, antennal clubs of 3 segments, antennal scapes relatively short (SL/CL: 0.71 ± 0.01). Compound eyes relatively small, ovoidal (EYE/CS: 0.20 ± 0.01). MESOSOMA. May present a slight metanotal depression, its dorsal profile being from almost straight to definitely rounded in lateral view. Propodeal spines are relatively long (SPST/CS: 0.30 ± 0.02), usually proportionally thin in minor worker but sometimes considerably thick in large ones. METASOMA. The petiole in profile view usually presents a short but visible horizontal section.A subpetiolar process is normally visible, consisting of a small tooth angled down at up to 90°. Postpetiole ordinarily roundish in lateral profile, subrectangular in dorsal view. SURFACE SCULPTURING. Most of the body is characterized by a very fine areolate-rugose sculpture, the longitudinal component of which tends to be notably more marked on the frontal side of the head only in larger specimens. The gaster and appendages are smooth, as well as the clypeus and a variable area extending around the frons. A central longitudinal stria visible in the lower portion of the clypeus. Very sparse, occasionally suberect but usually erect setae all over the body; dense, fine, and mostly adpressed pilosity on all appendages, especially abundant on the antennal flagelli. Queen (Figs 28–30) MEASUREMENTS AND INDICES (2 specimens, 2 localities). CL: 649–627; CW: 531–548; CS: 579–598; PoOC: 239–256; SL: 457–469; ML: 932–989; MW: 585–585; EL: 190–195; EW: 157–161; EYE: 173–178; SPST: 216–237; CL/CW: 1.08–1.08; PoOC: 0.37–0.41; SL/CL: 0.72–0.73; ML/CS: 1.65– 1.61; MW/CS: 0.97–1.00; EL/CS: 0.32–0.33; EW/CS: 0.27–0.27; EYE/CS: 0.30–0.30; SPST/CS: 0.37–0.39. COLORATION. Entirely yellowish with the main exception of parts of the gaster: at least the first tergite characterized by a blackish transverse band. Moreover, the mesoscutellar disk is darkened caudally. HEAD. Subrectangular, with rounded margins near the four angles, clypeus and mandibles rounded. Antennae of 12 segments, antennal clubs of 3 segments, antennal scapes relatively short (SL/CL: 0.72– 0.73). Compound eyes large and ovoidal (EYE/CS: 0.30); ocelli circular. MESOSOMA. Propodeal spines relatively long (SPST/CS: 0.37–0.39). METASOMA. Dorsum of petiole in profile view without a horizontal section. Subpetiolar process usually visible, consisting of a small tooth. Postpetiole ordinarily roundish in profile, subrectangular in dorsal view. SURFACE SCULPTURING. Most of the body is characterized by a variably fine areolate-rugose sculpture, strong longitudinal rugae on the head. Clypeus. gaster and appendages smooth; a variable area extending around the frons, most of the pronotum, anepisterna, katepisterna, mesoscutellar disk and mesoscutum. A central longitudinal stria visible in the lower portion of the clypeus. Very sparse, occasionally suberect but usually erect setae all over the body; dense, fine, and mostly adpressed pilosity on all appendages, especially abundant on the antennal flagelli. Male (Figs 31–33) MEASUREMENTS AND INDICES (3 specimens, 2 localities). CL: 443 ± 13.11 (429–445); CW: 405.67 ± 3.51 (402–409); CS: 424.33 ± 7.84 (415–430); PoOC: 189.67 ± 2.51 (187–192); SL: 203 ± 4 (199–207); ML: 836.67 ± 5.13 (832–841); MW: 498.67 ± 3.78 (496–503); EL: 186.67 ± 6.03 (181–193); EW: 165.67 ± 3.78 (163–170); EYE: 176.16 ± 3.62 (172–178); SPST: 126.33 ± 4.51 (122–131); CL/CW: 1.04 ± 0.01 (1.03–0.06); PoOC/CS: 0.43 ± 0.01 (0.42–0.43); SL/CL: 0.45 ± 0.01 (0.45–0.46); ML/CS: 1.97 ± 0.04 (1.93–2.01); MW/CS: 1.17 ± 0.03 (1.15–1.21); EL/CS: 0.44 ± 0.01 (0.42–0.45); EW/CS: 0.39 ± 0.02 (0.38–0.41); EYE/CS: 0.41 ± 0.01 (0.40–.043); SPST/CS: 0.30 ± 0.02 (0.28–0.31). COLORATION. Whole body yellowish, head slightly darker, appendages whitish, gaster dark and mesoscutellar disk caudally darkened. HEAD. Subrectangular, rounded especially above the eyes; clypeus and mandibles rounded. Antennae of 13 segments, antennal clubs of 4 segments. Compound eyes large and ovoidal (EYE/CS: 0.41 ± 0.01); ocelli circular. MESOSOMA. Propodeum armed with very short and thick spines (SPST/CS: 0.30 ± 0.02). METASOMA. Postpetiole ordinarily roundish in profile, subrectangular in dorsal view. SURFACE SCULPTURING. Sculpture relatively strong on the head, with well-developed longitudinal striae on its frontal side, very light all over the propodeum, petiole, postpetiole and in some areas near the mesosoma lateral sutures, all the remaining parts smooth. A central longitudinal stria visible in the lower portion of the clypeus. Sparse, erect setae all over the body; fine and mostly adpressed pilosity over the appendages. Phylogeny (Fig. 79) Sister species of T. marae Alicata, Schifani & Prebus sp. nov., which together are closely related to T. flavicornis. Distribution and biogeography (Figs 80–81) Occurs almost all over Sicily and is also found in the Maltese Islands. In the past, Sicily and the Maltese Islands have been connected via an extensive land bridge covering a large area of currently shallow waters, and the last time such a connection existed was during the Last Glacial Maximum (Foglini et al. 2016). Ecology and conservation Relatively thermophilous, collected from 5 to 840 m a.s.l. (see Fig. 82). Temnothorax lagrecai inhabits Mediterranean shrublands and relatively open forests like native and artificial forests of Pinus halepensis Mill., sometimes found under trees of Quercus ilex L., but was also collected in artificial gardens in the leaf litter of trees of Citrus L. Due to the wide ecological niche and distribution it may face a relatively positive situation in conservationist terms, but further assessment would be interesting. Nesting Nests are found in the soil, probably opportunistically exploiting several kinds of microhabitats when available. On Monte Etna entire nests were found several times under moss. Biology Monogynous in all documented cases. Social parasites One colony was found to be hosting T. muellerianus (Finzi, 1922) in the R.N.O. Pino d’Aleppo (Vittoria). Phenology Flying queens and males (often attracted by artificial lights) and mating were observed in Mondello (Palermo) from the early days of July to late August at least. Notes Baroni Urbani (1964) assembled the type series of this species from an unspecified number of worker specimens and a single queen collected in Bosco di S. Pietro (Hyblaean Plateau, SE Sicily) by Giovanni Sichel. No additional information on this taxon was published after its description, and its name was mentioned only on very few occasions (Baroni Urbani 1971; Salata & Borowiec 2019). Baroni Urbani decided to define the only queen he had as the holotype, despite the specimen being damaged (missing both antennae) and with the taxonomy of Temnothorax being mostly built around workers. He based the worker caste description on a specimen he defined as the ‘ergatotype’, but this definition has no legal value anymore according to the ICZN code, therefore that specimen must be considered as a simple paratype. We examined the holotype along with several paratypes preserved at the Natural History Museums of Verona and Vienna, but our efforts to find the ergatotype failed. Over 20 years ago, it was briefly observed at the home of Bruno Poldi (1920–2002) by one of us (AA), but it appears to be absent from the MNHM where Poldi’s collection is kept, as well as from the MSNV, NHMB and NHMW where other material from Baroni Urbani and specimens of T. lagrecai are stored. According to its description, the ergatotype shows several characters deviating from all type and non-type workers of T. lagrecai that we have found; however, it is unclear whether this is due to the actual features of this lost specimen or an imprecise description. Temnothorax marae sp. nov. cooccurs with T. lagrecai in the Bosco di S. Pietro, yet it is not characterized by the sharp petiole and mesoepinotal furrow of the drawing of the ergatotype of T. lagrecai found in Baroni Urbani's original description. In any case, the holotype of T. lagrecai, although partly damaged, is fully coherent with the morphology of the queens collected alongside workers with the same characters of T. lagrecai that we present here, so that we consider the issue over the identity of T. lagrecai to be resolved. Maltese records under Leptothorax rabaudi Bondroit, 1918 by Schembri & Collingwood (1981) are partly based on misidentified T. lagrecai according to an investigation of their voucher specimens., Published as part of Schifani, Enrico, Prebus, Matthew M. & Alicata, Antonio, 2022, Integrating morphology with phylogenomics to describe four island endemic species of Temnothorax from Sicily and Malta (Hymenoptera, Formicidae), pp. 143-179 in European Journal of Taxonomy 833 (1) on pages 152-156, DOI: 10.5852/ejt.2022.833.1891, http://zenodo.org/record/6981130, {"references":["Baroni Urbani C. 1964. Studi sulla mirmecofauna d'Italia. 2. Formiche di Sicilia. Atti della Accademia Gioenia di Scienze Naturali in Catania 16: 25 - 66.","Foglini F., Prampolini M., Micallef A., Angeletti L., Vandelli V., Deidun A., Soldati M. & Taviani M. 2016. Late Quaternary coastal landscape morphology and evolution of the Maltese Islands (Mediterranean Sea) reconstructed from high-resolution seafloor data. Geological Society, London, Special Publications 411: 77 - 95. https: // doi. org / 10.1144 / SP 411.12","Baroni Urbani C. 1971. Catalogo delle specie di Formicidae d'Italia (Studi sulla mirmecofauna d'Italia X). Memoria della Societa Entomologica Italiana 50: 5 - 287.","Salata S. & Borowiec L. 2019. Preliminary division of not socially parasitic Greek Temnothorax Mayr, 1861 (Hymenoptera, Formicidae) with a description of three new species. ZooKeys 877: 81 - 131. https: // doi. org / 10.3897 / zookeys. 877.36320","Schembri S. P. & Collingwood C. A. 1981. A revision of the myrmecofauna of the Maltese Islands (Hymenoptera, Formicidae). Annali del Museo Civico di Storia Naturale \" Giacomo Doria \" 83: 417 - 442."]}

16 pages, 5 figures, 4 tables, supplementary material https://www.frontiersin.org/articles/10.3389/fmars.2021.698890/full#supplementary-material.-- Data Availability Statement: The original contributions presented in the study are included in the article/Supplementary Material, further inquiries can be directed to the corresponding author, Benthic habitats of the deep Mediterranean Sea and the biodiversity they host are increasingly jeopardized by increasing human pressures, both direct and indirect, which encompass fisheries, chemical and acoustic pollution, littering, oil and gas exploration and production and marine infrastructures (i.e., cable and pipeline laying), and bioprospecting. To this, is added the pervasive and growing effects of human-induced perturbations of the climate system. International frameworks provide foundations for the protection of deep-sea ecosystems, but the lack of standardized criteria for the identification of areas deserving protection, insufficient legislative instruments and poor implementation hinder an efficient set up in practical terms. Here, we discuss the international legal frameworks and management measures in relation to the status of habitats and key species in the deep Mediterranean Basin. By comparing the results of a multi-criteria decision analysis (MCDA) and of expert evaluation (EE), we identify priority deep-sea areas for conservation and select five criteria for the designation of future protected areas in the deep Mediterranean Sea. Our results indicate that areas (1) with high ecological relevance (e.g., hosting endemic and locally endangered species and rare habitats),(2) ensuring shelf-slope connectivity (e.g., submarine canyons), and (3) subject to current and foreseeable intense anthropogenic impacts, should be prioritized for conservation. The results presented here provide an ecosystem-based conservation strategy for designating priority areas for protection in the deep Mediterranean Sea, This study was supported by the DG ENV project IDEM (Implementation of the MSFD to the Deep Mediterranean Sea; contract EU No. 11.0661/2017/750680/SUB/EN V.C2). MC and QG-B acknowledge support from Generalitat de Catalunya autonomous government through its funding scheme to excellence research groups (Grant 2017 SGR 315), With the institutional support of the ‘Severo Ochoa Centre of Excellence’ accreditation (CEX2019-000928-S)

Acanthopleura vaillantii de Rochebrune, 1882 (Fig. 17) Chiton (Tonicia) sueziensis, non Reeve; Issel 1869, p. 235 (pars). Acantopleura (sic) vaillantii de Rochebrune 1882, p. 192. Acanthopleura gemmata; Ferreira 1983, p. 278, fig. 30. Acanthopleura haddoni; Vine 1986, p. 126, unnumbered figs p. 125, 126. Acanthopleura vaillantii; Ferreira 1986, p. 231, fig. 17; Kaas & Van Belle 1988, p. 123; Strack 1993, p.: 17, pl. 4, figs 3���4; Bosch et al. 1995, p. 190, fig. sp. 885; Dekker & Orlin 2000, p. 7; Slieker 2000, p. 50, fig. 24; Brooker 2003, p. 44, 153, 214, 486, figs 5.8, 6.5; Abubakr 2004, p. 73; Anseeuw & Terryn 2004, p. 14, figs 46���47; Kaas et al. 2006, p. 268, fig. 109, map 45; Dekker & Gemert 2008, p. 124; Dinapoli & Janssen 2009, p. 8, pl. 4a���c; Sadeghi & Loghmani 2010, p. 1, figs 2���3; Angeletti et al. 2018, p. 374, fig. 6, Blatterer 2019, p. 52, Fig. 3 f-g, pl. 3, fig. 6 a-e. Type material. MNHN, lectotype and paralectotype designated by Ferreira (1986). Type locality. Suez, Egypt. Material examined. Saudi Arabia, Gulf of Aqaba (Ash Shaykh Humayd): St. 3: 11 valves (1 head, width 18.5 mm, and 10 intermediate, maximum width 29 mm), Figs 17 A���C (MZB 50554); St. 6: 1 intermediate valve, width 24.5 mm (MZB 60261); St. 9: 1 intermediate valve (figured in Angeletti et al. 2018). Egypt, Shaw��r��ţ island.: St. 19: 1 intermediate valve, width 27.5 mm (MZB 60274). Description. Head valve nearly semicircular. Intermediate valves broadly rectangular to widely V-shaped, side margins rounded, apex blunt, lateral areas little or not raised, hardly marked. Tegmental sculpture often indistinguishable on account of erosion and encrustation, consists of roundish tubercles irregularly arranged, ocelli very small, abundantly distributed on end valves and more than half lateral areas of intermediate valves. Articulamentum glossy, apophyses large, rounded, connected across the sinus by a short, concave jugal plate, slit formula 10/I/ 9���10, teeth short, deeply grooved on the dorsal side, pectinate. Remarks. This is the only species of Acanthopleura known to live in the northern Red Sea, and is also the largest Red Sea chiton (reaching �� 100 mm in length). The determination of Acanthopleura species is often difficult due to their high degree of intraspecific variability, similarity between congeneric species, and frequent erosion on the dorsal parts of plates. Ferreira (1983, 1986) considered all the Red Sea specimens to belong to A. gemmata (de Blainville, 1825), while Kaas & Van Belle (1988) and Strack (1993) regarded the Red Sea, Yemen, Oman and Arabian Gulf populations as distinct from A. gemmata, A. vaillantii de Rochebrune, 1882 being the oldest available name for these. The Red Sea specimens were also determined in early publications as Chiton spiniger Sowerby, 1840 (now considered a synonym of A. gemmata) and Acanthopleura haddoni Winckworth, 1927 (Brooker 2003). Acanthopleura vaillantii inhabits the intertidal zone and is therefore an excellent sea-level indicator in paleoecological studies (Angeletti et al. 2018). However, on-land occurrences of loose plates could be misleading for geological purposes due to collection and displacement by humans rather than representing genuine ecological settings (M. Taviani, pers. observ.). Distribution. Late Pleistocene: Saudi Arabia, Gulf of Aqaba (Ash Shaykh Humayd: this study); Egypt (Hurghada: Issel 1869; Shawarit isl.: this study). Present-day: Red Sea, Yemen, Socotra Island, Oman, Iran, and the entrance of the Arabian Gulf (Kaas et al. 2006; Dinapoli & Janssen 2009; Blatterer 2019)., Published as part of Dell'Angelo, Bruno, Landau, Bernard M., Sosso, Maurizio & Taviani, Marco, 2020, Late Pleistocene Red Sea Mollusca: 1. Polyplacophora, pp. 401-449 in Zootaxa 4772 (3) on pages 428-429, DOI: 10.11646/zootaxa.4772.3.1, http://zenodo.org/record/3819654, {"references":["Rochebrune, A. T. de (1882) Monographie des especes fossiles appartenant a la classe des Polyplaxiphores. Annales des Sciences Geologiques, 14, 1 - 74, pls. 1 - 3.","Issel, A. (1869) Malacologia del Mar Rosso. Ricerche zoologiche e paleontologiche. Parte terza. Catalogo delle conchiglie fossili raccolte sulle spiagge emerse del Mar Rosso. Biblioteca Malacologica, Pisa, 61 pp. [pp. 243 - 303]","Ferreira, A. J. (1983) Researches on the coast of Somalia. The chiton fauna (Mollusca Polyplacophora). Monitore Zoologico Italiano, 9, 249 - 297. https: // doi. org / 10.1080 / 00269786.1983.11758574","Vine, P. (1986) Red Sea Invertebrates. IMMEL Publishing, London, 224 pp.","Ferreira, A. J. (1986) A revision of the genus Acanthopleura Guilding, 1829 (Mollusca: Polyplacophora). The Veliger, 28, 221 - 279.","Kaas, P. & Van Belle, R. A. (1988) Chitons (Mollusca: Polyplacophora) from the coasts of Oman and the Arabian Gulf. American Malacological Bulletin, 6, 115 - 130.","Strack, H. L. (1993) The Polyplacophora of the Red Sea. Journal of the Malacological Society of Australia, 14, 1 - 40. https: // doi. org / 10.1080 / 00852988.1993.11923901","Bosch, D. T., Dance, S. P., Moolenbeek, R. G. & Oliver, P. G. (1995) Seashells of Eastern Arabia. Motivate Publishing, Dubai, 290 pp.","Dekker, H. & Orlin, Z. (2000) Check-list of Red Sea Mollusca. Spirula, 47 (Supplement), 1 - 46.","Slieker, F. J. A. (2000) Chitons of the world. An illustrated synopsis of recent Polyplacophora. L'Informatore Piceno Ed., Ancona, 154 pp.","Brooker, L. R. (2003) Revision of Acanthopleura Guilding, 1829 (Mollusca: Polyplacophora) based on light and electron microscopy. Thesis, Murdoch University, Perth, xiii + 528 pp.","Abubakr, M. M. (2004) The Republic of Yemen Marine Biotic Ecosystem (Resources-Habitats and Species). Ministry of Water and Environment, Environment Protection Authority, Aden, 128 pp.","Anseeuw, B. & Terryn, Y. (2004) Intertidal chitons (Mollusca: Polyplacophora) from the coast of Jordan, Red Sea, with the description of a new species of Parachiton Thiele, 1909. Bollettino Malacologico, Supplement 5, 1 - 24.","Kaas, P., Van Belle, R. A. & Strack, H. L. (2006) Monograph of Living Chitons (Mollusca: Polyplacophora). Volume 6. Suborder Ischnochitonina (concluded): Schizochitonidae; Chitonidae. Additions to Volumes 1 - 5. E. J. Brill, Leiden-Boston, 463 pp.","Dekker, H. & Gemert, L. J. van (2008) A new list with corrections to the shells pictured in \" Red Sea Shells \" (1984) by Doreen Sharabati. De Kreukel, 44, 123 - 136.","Dinapoli, A. & Janssen, R. (2009) Polyplacophora (Mollusca) of the Socotra Arcipelago - Systematics and Biogeography. Fauna of Arabia, 24, 1 - 46.","Sadeghi, P. & Loghmani, M. (2010) First record of Acanthopleura vaillantii (Mollusca: Polyplacophora) from Iran-Chabahar Bay in the Oman Sea. Biodiversity Records, 3, e 7. https: // doi. org / 10.1017 / S 1755267209990984","Angeletti, L., Rasul, N. M. A. & Taviani, M. (2018) Mollusc fauna associated with Late Pleistocene coral reef systems of the Saudi Arabian side of the Gulf of Aqaba. In: Rasul, N. M. A. & Stewart, C. F. (Eds.), Geological Setting, Palaeoenvironment and Archaeology of the Red Sea, Springer, Cham, pp. 367 - 387. https: // doi. org / 10.1007 / 978 - 3 - 319 - 99408 - 6 _ 17","Blatterer, H. (2019) Mollusca of the Dahab region. Oberosterreichisches Landesmuseum, Linz, Austria, 480 pp.","Blainville, H. M. D. de (1825) Oscabrelle. Oscabrion. In: F. Cuvier (Ed.), Dictionnaire des Sciences Naturelles. Vol. 36. F. G. Levrault, Paris & Le Normant, Strasbourg, pp. 519 - 556.","Winckworth, R. (1927) New species of chitons from Aden and South India. Proceedings of the Malacological Society of London, 17, 206 - 208, pls. 28 - 29."]}

Ascorhiza mawatarii d���Hondt, 1983 ( Figs 2 A���E; 3A) Ascorhiza mawatarii d���Hondt, 1983: 24, fig. 12; Harmelin & d���Hondt 1992b: 609. Material examined. MNCN 25.03 /4194: DA10, Gazul MV, 390���410 m depth, 26 June 2010, INDEMARES CHI- CA project, Instituto Espa��ol de Oceanografia (IEO) coll., on shells; MNCN 25.03 /4195: BT05, Chica MVD, 607���665 m depth, 19 February 2011, INDEMARES CHICA project, IEO coll., on shells. Description. Colony erect (22���42 mm long), delicate, formed by a dilated head (7���18 mm long x 3.5���6 mm wide) and a narrower peduncle (15���24 mm long x 1���1.5 mm wide) (Fig. 2A, B), fixed to the substrate directly by expansion of the peduncle base or by rhizoids (Fig. 2C). Zooids from the head disposed irregularly, hexagonal with circular aperture (Fig. 2D); zooids from de peduncle (myoecia) with distinct form: those near the top polygonal, alternately arranged (Fig 2E), those near the base somewhat rectangular, slightly longer than wide, arranged in rows (Fig. 3A). ......continued on the next page ......continued on the next page ......continued on the next page Remarks. There are two described species of Ascorhiza: A. mawatari and A. occidentalis Fewkes, 1889 (Bock 2019; accessed 02.12.2019). The latter species is characterized by a narrow stalk composed of a number of segments, externally indicated by ferrules of uniform size with well-marked indentations. Ascorhiza mawatari was described from the Northern Pacific Ocean (d���Hondt 1983) and recorded later by Harmelin & d���Hondt (1992b) in European waters, precisely in the GoC and in the Mediterranean Sea (Bouche des Bonifacio). Colony length in the original material is 20 mm (peduncle: 12 mm long by 1.8 mm wide; head: 4.5 mm wide), while the material described by Harmelin & d���Hondt (1992b) from the GoC is more similar to our specimens (up to 55 mm long). The present material differs from that described by previous authors in having a narrower peduncle (often 1 mm wide) and smaller zooids (both length and width): d���Hondt (1983) reported that myoecia were almost 1 mm long by 0.4���0.5 mm width. Although the GoC material was poorly preserved, as all the peduncles were collapsed, the general features were similar. Two attachment modes were observed associated to the different type of substrate. In colonies attached to shells, the base was continuous and expanded, while colonies attached to sand grains used several rizhoids (Fig. 2C). Apparently, this species varies in both the base of the peduncle and the shape of the peduncle zooids, and further studies are required to understand the reasons behind this variability, as the knowledge on this species is still scarce. Habitat and associated species. Ascorhiza mawatari was found associated with the bathyal cold-water coral framework (mainly Madrepora oculata Linnaeus), colonized by small gorgonians (Acanthogorgia and Bebryce). It was also found in sand and muddy sea bottoms with sea pens [Kophobelemnon stelliferum (M��ller) and Funiculina quadrangularis (Pallas)], solitary corals (e.g. Flabellum chunii Marenzeller) and sea urchins, mainly Cidaris cidaris (Linnaeus)., Published as part of Ramalho, La��s V., L��pez-F��, Carlos M., Mateo-Ram��rez, Angel & Rueda, Jos�� Luis, 2020, Bryozoa from deep-sea habitats of the northern Gulf of C��diz (Northeastern Atlantic), pp. 451-478 in Zootaxa 4768 (4) on pages 455-462, DOI: 10.11646/zootaxa.4768.4.1, http://zenodo.org/record/3785758, {"references":["d'Hondt, J. - L. (1983) Tabular keys for identification of the Recent Ctenostomatous Bryozoa. Memoires de l'Institut Oceanographique, 14, 1 - 134.","Harmelin, J. - G. & d'Hondt, J. - L. (1992 b) Bryozoaires des parages de Gibraltar (campagne oceanographique BALGIM, 1984). 2 - Ctenostomes et cyclostomes. Bulletin du Museum National d'histoire Naturelle, 14, 605 - 621.","Alvarez, J. A. (1992) Briozoos de la Campana Fauna I. Parte Primera: Ctenostomida y Cheilostomida Anascina. Cahiers de Biologie Marine, 33, 273 - 297.","Alvarez, J. A. (1994) Briozoos de la Campana Fauna I (Sur de la Peninsula Iberica). Parte II: Cheilostomida Ascophorina y Cyclostomida. Graellsia, 50, 129 - 145.","Berning, B., Tilbrook, K. J. & Rosso, A. (2008) Revision of the north-eastern Atlantic and Mediterranean species of the genera Herentia and Therenia (Bryozoa: Cheilostomata). Journal of Natural History, 42, 1509 - 1547. https: // doi. org / 10.1080 / 00222930802109140","Calvet, L. (1906) Note preliminaire sur les bryozoaires recueillis par les expeditions du \" Travailleur \" (1881 - 1882) et du \" Talisman \" (1883). Bulletin du Museum National d'histoire Naturelle, 12, 154 - 166. [http: // biodiversitylibrary. org / page / 5021333]","Harmelin, J. - G. & Aristegui, J. (1988) New Cribrilinidae (Bryozoa, Cheilostomata) from the upper bathyal of the Atlanto-Mediterranean region. Journal of Natural History, 22, 507 - 535. https: // doi. org / 10.1080 / 00222938800770351","Harmelin, J. - G. & d'Hondt, J. - L. (1992 a) Bryozoaires des parages de Gibraltar (campagne oceanographique BALGIM, 1984). 1 - Cheilostomes. Bulletin du Museum National d'histoire Naturelle, 14, 23 - 67.","Harmelin, J. - G. (1973) Les Bryozoaires des peuplements sciaphiles de Mediterranee: le genre Crassimarginatella Canu (Chilostomes Anasca). Cahiers de Biologie Marine, 14, 471 - 492.","d'Hondt, J. - L. (1974) Bryozoaires recoltes par la \" Thalassa \" dans le Golfe de Gascogne (Campagnes de 1968 a 1972). Cahiers de Biologie Marine, 15, 27 - 50.","Lopez de la Cuadra, C. M. & Garcia-Gomez, J. C. (1994) Zoogeographical study of the Cheilostomatida from the Straits of Gibraltar. In: Hayward, P. J., Ryland, J. S. & Taylor, P. D. (Eds.), Biology and Palaeobiology of Bryozoans. Olsen & Olsen, Fredensborg, pp. 107 - 112.","Lopez de la Cuadra, C. M. & Garcia-Gomez, J. C. (2001) New and little-known ascophoran bryozoans from the Western Mediterranean, collected by ' Fauna Iberica' expeditions. Journal of Natural History, 35, 1717 - 1732. https: // doi. org / 10.1080 / 002229301317092414","Madurell, T., Zabala, M., Dominguez-Carrio, C. & Gili, J. M. (2013) Bryozoan faunal composition and community structure from the continental shelf off Cap de Creus (Northwestern Mediterranean). Journal of Sea Research, 83, 123 - 136. https: // doi. org / 10.1016 / j. seares. 2013.04.013","Prenant, M. & Bobin, G. (1966) Bryozoaires. deuxieme Partie. Chilostomes Anasca. 647 pp.","Ramalho, L. V., Lopez-Fe, C. M. & Rueda, J. L. (2018 a) Three species of Reteporella (Bryozoa: Cheilostomata) in a diapiric and mud volcano field of the Gulf of Cadiz, with the description of Reteporella victori n. sp. Zootaxa, 4375 (1), 90 - 104. https: // doi. org / 10.11646 / zootaxa. 4375.1.4","Reverter-Gil, O. & Fernandez-Pulpeiro, E. (1999) Some records of Bryozoans from NW Spain. Cahiers de Biologie Marine, 40, 35 - 45.","Reverter-Gil, O. & Fernandez-Pulpeiro, E. (2001) Inventario y cartografia de los Briozoos marinos de Galicia (N. O. de Espana). Monografias de Nova Acta Cientiifica Compostelana, Serie Bioloxia, 1, 1 - 243.","Reverter-Gil, O., Souto, J. & Fernandez-Pulpeiro, E. (2011) Revision of the genus Crepis Jullien (Bryozoa: Cheilostomata) with description of a new genus and family and notes on Chlidoniidae. Zootaxa, 2993 (10), 1 - 22. https: // doi. org / 10.11646 / zootaxa. 2993.1.1","Reverter-Gil, O., Souto, J. & Fernandez-Pulpeiro, E. (2014) Annotated checklist of Recent marine Bryozoa from continental Portugal. Nova Acta Cientifica Compostelana, Serie Bioloxia, 21, 1 - 55.","Reverter-Gil, O., Berning, B. & Souto, J. (2015) Diversity and Systematics of Schizomavella Species (Bryozoa: Bitectiporidae) from the Bathyal NE Atlantic. PLoS ONE, 10, e 0139084. http: // doi. org / 10.1371 / journal. pone. 0139084","Reverter-Gil, O., Souto, J. & Trigo, J. E. (2019 a) New species and new records of bryozoans from Galicia (NW Spain). Journal of Natural History, 53 (3 - 4), 221 - 251. https: // doi. org / 10.1080 / 00222933.2019.1582815","Reverter-Gil, O., Souto, J. & Trigo, J. E. (2019 b) Novos datos de Briozoos en Galicia (NW Espana). New data on Galician Bryozoa (NW Spain). Nova Acta Cientifica Compostelana, Serie Bioloxia, 26, 1 - 36.","Rosso, A. & Di Martino, E. (2016) Bryozoan diversity in the Mediterranean Sea: an update. Mediterranean Marine Science, 17 (2), 567 - 607. https: // doi. org / 10.12681 / mms. 1706","Rosso, A. & Di Martino, E. & Gerovasileiou, V. (2020) Revision of the genus Setosella (Bryozoa: Cheilostomata) with description of new species from deep-waters and submarine caves of the Mediterranean Sea. Zootaxa, 4728 (4), 401 - 442. https: // doi. org / 10.11646 / zootaxa. 4728.4.1","Rueda, J. L., Urra, J., Aguilar, R., Angeletti, L., Bo, M., Garcia-Ruiz, C., Gonzalez-Duarte, M. M., Lopez, E., Madurell, T., Maldonado, M., Mateo-Ramirez, A., Megina, C., Moreira, R., Moya, F., Ramalho, L. V., Rosso, A., Sitja, C. & Taviani, M. (2019) Cold-Water Coral Associated Fauna in the Mediterranean Sea and Adjacent Areas. In: Orejas, C. & Jimenez, C. (Eds.), Mediterranean Cold-Water Corals: Past, Present and Future-Understanding the Deep-Sea Realms of Coral. Coral Reefs of the World, 9, pp. 295 - 334. https: // doi. org / 10.1007 / 978 - 3 - 319 - 91608 - 8 _ 29","Souto, J., Reverter-Gil, O., De Blauwe, H. & Fernandez-Pulpeiro, E. (2014) New records of bryozoans from Portugal. Cahiers de Biologie Marine, 55, 129 - 150.","Souto, J., Berning, B. & Ostrovsky, A. N. (2016) Systematics and diversity of deep-water Cheilostomata (Bryozoa) from Galicia Bank (NE Atlantic). Zootaxa, 4067 (4), 401 - 459. https: // doi. org / 10.11646 / zootaxa. 4067.4.1","Zabala, M. & Maluquer, P. (1988) Illustrated keys for the classification of Mediterranean Bryozoa. Treballs del Museu de Zoologia de Barcelona, 4, 1 - 294.","Barroso, M. G. (1920) Notas sobres Briozoos espanoles. 8. Boletin de la Real Sociedad espanola de Historia Natural, 20, 353 - 362.","Hincks, T. (1880 a) Contributions towards a general history of the marine Polyzoa. Part I. Madeiran Polyzoa. Annals and Magazine of Natural History, Series 5, 6 (31), 69 - 80. https: // doi. org / 10.1080 / 00222938009458895","Hincks, T. (1862) A catalogue of the Zoophytes of South Devon and South Cornwall. The Annals and Magazine of Natural History; Zoology, Botany, and Geology, Series 3, 9 (49), 22 - 30. https: // doi. org / 10.1080 / 00222936208681181","Jullien, J. (1882) Dragages du \" Travailleur \", Bryozoaires. Especes draguees dans l'Ocean Atlantique en 1881. Especes nouvelles ou incompletement decrites. Extrait du Bulletin de la Societe zoologique de France, 7, 497 - 529. https: // doi. org / 10.5962 / bhl. part. 27178","Lagaaij, R. (1952) The Pliocene Bryozoa of the Low Countries and their bearing on the marine stratigraphy of the North Sea region. Mededelingen van de Geologische Stichting, 5, 6 - 233.","Busk, G. (1860) Zoophytology. Shetland Polyzoa. Collected by Mr. Barlee (continued and concluded). Quarterly Journal of Microscopical Science, 8, 213 - 214.","Lamouroux, J. V. F. (1816) Histoire des polypiers Coralligenes Flexibles, vulgairement nommes Zoophytes. F. Poisson, Caen, 559 pp. https: // doi. org / 10.5962 / bhl. title. 11172","Reuss, A. E. (1848) Die fossilen Polyparien des Wiener Tertiarbeckens. Haidingers Naturwissenschaftlichen Abhandlungen, 2, 1 - 109.","Bishop, J. D. D. & Househam, B. C. (1987) Puellina (Bryozoa; Cheilostomatida; Cribrilinidae) from British and adjacent waters. Bulletin of the British Museum (Natural History), Zoology, 53, 1 - 63.","Busk, G. (1858) Zoophytology. On some Madeiran Polyzoa. Quarterly Journal of Microscopical Science, 6, 124 - 130. https: // doi. org / 10.1111 / j. 1365 - 2818.1858. tb 04550. x","Busk, G. (1861) Description of new Polyzoa collected by J. Y. Johnston Esq., at Madeira. Quarterly Journal of Microscopical Science, New Series, 1, 77 - 80.","Norman, A. (1867) Report of the committee appointed for the purpose of exploring the coasts of the Hebrides by means of the dredge. Part 2. On the Crustacea, Echinodermata, Polyzoa, Actinozoa and Hydrozoa. Report of the British Association for the Advancement of Science London, 1866, 193 - 206.","Audouin, J. V. (1826) Explication sommaire des planches de polypes de l'Egypte et de la Syrie, publiees par Jules-Cesar Savigny. In: Audouin, J. V. (Ed.), Description de l'Egypte, ou recueil des observations et des recherches qui ont ete faites en Egypte pendant l'expedition de l'armee francaise ... Histoire naturelle, 1 (4), pp. 1 - 339.","David, L. & Pouyet, S. (1978) Le genre Herentia Gray, 1848 (Bryozoa, Cheilostomata). Systematique et phylogenese, biostratigraphie et biogeographie. Documents des Laboratoires de Geologie de la Faculte des Sciences de Lyon HS 4, Livre Jubilaire Jacques Flandrin, 1978, 167 - 193.","Linnaeus, C. (1767) Systemae naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differetiis, synonymis, locis Regnum Animale. Laurentii Salvii, Holmiae, 532 pp. https: // doi. org / 10.5962 / bhl. title. 158187","Hincks, T. (1860) Descriptions of new Polyzoa from Ireland. Quarterly Journal of Microscopical Science, 8, 275 - 280.","Smitt, F. A. (1867) Kritisk forteckning ofver Skandinaviens Hafs-Bryozoer. II. Ofversigt af Kongliga Vetenskaps-Akademiens Forhandlingar, 23, 395 - 534.","Meneghini, G. (1844) Polipidella famiglia dei Tubuliporiani finora osservati nell'Adriatico. Nuovi Annali delle Scienze Naturali, 3, 115 - 133.","Fewkes, J. W. (1889) A preliminary notice of a stalked bryozoan Ascorhiza occidentalis. Annals and Magazine of Natural History, Series 6, 3 (13), 1 - 6. https: // doi. org / 10.1080 / 00222938909460287","Bock, P. (2019) Recent and Fossil Bryozoa. Available from: http: // bryozoa. net / index. html (accessed 2 Dec. 2019)"]}

Biodiversity of coral forests and occurrence of Derelict Fishing Gears (DFGs) have been assessed in canyon systems of the western Ligurian Sea (Dramont, Monaco, Bordighera, Arma di Taggia and Bergeggi) exposed to different anthropic pressures. Arborescent cnidarians were elected as representative species due to their role as structuring organisms and their vulnerability to DFGs damage; hence, their occurrence, density and distribution were correlated to the presence of DFGs. The canyon systems were mapped using a Multibeam Echo Sounder and visually surveyed by means of a Remotely Operated Vehicle between 20 and 445 m depth. With the exception of the Bergeggi canyon system, all sites host rich assemblages of structuring anthozoans, accounting for more than 2000 colonies belonging to 11 species, predominantly Eunicella cavolinii, Paramuricea clavata, Corallium rubrum, Dendrophyllia cornigera, and E. verrucosa. We noticed that the large structuring gorgonians were affected most, with a high number of entangled colonies. DFGs represent 85% of the marine litter found, and are the most serious threat to resident sessile communities, most noticeably in the eastern canyons. Ports, size and fishing effort of local fleets, and socio-economical differences in the fishing activities, strongly influence the fishing footprints. Our study further confirms the role of submarine canyons as site of high coral biodiversity and vulnerability to the mechanical damages by fishing-related littering, calling for adequate management measures to reduce fishery pressure and concomitant DFGs discharge.

International audience; The Gulf of Naples is an example of the most beautiful and biodiverse marine regions of the Mediterranean Sea and of the most impacted areas in terms of industrial activities, large contaminated areas, resource exploitation, infrastructures at sea and maritime transportation. We conducted Remotely Operated Vehicle surveys in the Dohrn Canyon in the Tyrrhenian Sea at approximately 12 NM off Naples metropolitan area, and revealed a hotspot of deep-sea benthic biodiversity of sessile fauna at ca. 400 m depth. The hard bottoms are characterized by a high abundance of charismatic species, such as the habitat forming cold-water corals (CWC) Madrepora oculata, Lophelia pertusa, Desmophyllum dianthus in association with the large size bivalves Acesta excavata and Neopycnodonte zibrowii. This CWC-bivalve co-occurrence represents a novel biotope for the Mediterranean Sea, which coexists with the evidence of severe anthropogenic threats, such as illegal dumping and fishery malpractices that were visually documented during the survey. We recommend the adoption of specific protection measures to preserve these unique deep-sea assemblages showing the uncommon coexistence of such a number of deep-sea species in a single habitat. In the 19 th century, the beauty and biological richness of the Gulf of Naples (GoN) was such to convince Anton Dohrn to found there in 1872 the first marine station in the world to demonstrate the validity of the Darwin theories on evolution using marine organisms. The following 150 years have witnessed the unbridled growth of scientific studies in the gulf, along with the explosion of the industrial and urban development of the city of Naples and adjacent coast, especially after World War II 1 , which determined a progressive increase of the anthropogenic pressures. Today, the Naples metropolitan area accounts for ca. 4.5 million inhabitants and the entire population of the gulf is distributed along a mere 12 km coastal stretch. The presence of illegal dumping and untreated sewage , along with the presence of wide contaminated areas, intense maritime transportation, infrastructure at sea, other direct and indirect anthropogenic stressors 2-13 have progressively threatened the coastal ecosystems of this gulf, determining also a progressive loss of marine habitats. Despite a long tradition of biological studies that renders the GoN one of the most intensively investigated marine areas of the world, little is known about the ecology and environmental status of the deep-sea habitats of the gulf, whose hard bottom has been so far investigated only in terms of geological setting. A prominent submarine feature of the gulf is the Dohrn Canyon, a bifurcate structure (Fig. 1) that indents perpendicular to the coastline the continental shelf 12 NM from the Naples megalopolis, beginning at ca. −250 m and sharply declining down to ca 1300 m in the Tyrrhenian plain 14,15. This inactive canyon, part of the Magnaghi-Dohrn canyon system 16 and consisting of a western and an eastern branch, is thought to have been formed in response to relative

Solemyidae gen. indet. (Fig. 3 B) Material. Three specimens from Moonlight North (Y16/f0694), two medium sized, L4647, L4648, one larger, L4649 (Fig. 3 B), UOA. Description. Shell small (H up to ca. 12 mm, L up to ca. 21 mm), inequilateral, elongate oval; faint radial striae in anteroventral area, absent in other areas; posterior margin narrowly rounded; anterior margin more broadly rounded; dorsal and ventral margins gently convex; umbones prominent, angulated, situated ca. 20% along shell length from anterior margin; anterior portion of pallial line subparallel to ventral margin, adjoining anterior adductor scar directly medioventrally; anterior adductor scar rather large, situated with centre just in dorsal half of shell close to anterior margin. Remarks. New Zealand Miocene seep solemyids are known from only three poorly preserved internal moulds. A tenuous pallial line and anterior adductor scar were observed faintly in one of the three specimens. Critically, the posterodorsal region is never well enough preserved to show the presence or absence of an external ligament. Therefore, from the appearance of the shells and quality of preservation it is not possible to categorically place the specimens within any particular solemyid genus. Not only does the preservation of the New Zealand seep specimens hinder identification, but solemyid taxonomy is complex (e.g. Taylor et al. 2008; Kamenev 2009; Oliver et al. 2011) and external shell morphology in this group is conservative (e.g. Taviani et al. 2011; Hryniewicz et al. 2014). The Moonlight North locality is inferred as a bathyal deposit (Saether et al. 2010), and of those solemyid genera found in the New Zealand fossil record and at modern New Zealand seeps, Acharax is a deep-water genus. Solemya is found predominantly in shallower waters of the continental shelf and upper slope (Neulinger et al. 2006; Oliver et al. 2011), although Solemya has been recorded from bathyal depths (Kamenev 2009; Rodrigues et al. 2011; Sato et al. 2013)., Published as part of Saether, Kristian P., Jingeng, Sha, Little, Crispin T. S. & Campbell, Kathleen A., 2016, New records and a new species of bivalve (Mollusca: Bivalvia) from Miocene hydrocarbon seep deposits, North Island, New Zealand, pp. 1-26 in Zootaxa 4154 (1) on page 5, DOI: 10.11646/zootaxa.4154.1.1, http://zenodo.org/record/272151, {"references":["Taylor, J. D., Glover, E. A. & Williams, S. T. (2008) Ancient chemosynthetic bivalves: systematics of Solemyidae from eastern and southern Australia (Mollusca: Bivalvia). In: Davie, P. J. F. & Phillips, J. A. (Eds.), Proceedings of the Thirteenth International Marine Biological Workshop, The Marine Fauna and Flora of Moreton Bay, Queensland. Memoirs of the Queensland Museum. Nature, 54, 75 - 104.","Kamenev, G. M. (2009) North Pacific species of the genus Solemya Lamarck, 1818 (Bivalvia: Solemyidae), with notes on Acharax johnsoni (Dall, 1891). Malacologia, 51, 233 - 261. http: // dx. doi. org / 10.4002 / 040.051.0202","Oliver, P. G., Rodriguez, C. F. & Cunha, M. R. (2011) Chemosymbiotic bivalves from the mud volcanoes of the Gulf of Cadiz, NE Atlantic, with descriptions of new species of Solemyidae, Lucinidae and Vesicomyidae. ZooKeys, 113, 1 - 38. http: // dx. doi. org / 10.3897 / zookeys. 113.1402","Taviani, M., Angeletti, L. & Ceregato, A. (2011) Chemosynthetic bivalves of the family Solemyidae (Bivalvia, Protobranchia) in the Neogene of the Mediterranean Basin. Journal of Paleontology, 85, 1067 - 1076. http: // dx. doi. org / 10.1666 / 10 - 119.1","Hryniewicz, K., Little, C. T. S. & Nakrem, H. A. (2014) Bivalves from the latest Jurassic - earliest Cretaceous hydrocarbon seep carbonates from central Spitsbergen, Svalbard. Zootaxa, 3859 (1), 1 - 66. http: // dx. doi. org / 10.11646 / zootaxa. 3859.1.1","Saether, K. P., Little, C. T. S., Campbell, K. A., Marshall, B. A., Collins, M. & Alfaro, A. C. (2010) New fossil mussels (Bivalvia: Mytilidae) from Miocene hydrocarbon seep deposits, North Island, New Zealand, with general remarks on vent and seep mussels. Zootaxa, 2577, 1 - 45.","Neulinger, S. C., Sahling, H., Suling, J. & Imhoff, J. F. (2006) Presence of two phylogenetically distinct groups in the deep-sea mussel Acharax (Mollusca: Bivalvia: Solemyidae). Marine Ecology Progress Series, 312, 161 - 168. http: // dx. doi. org / 10.3354 / meps 312161","Rodrigues, C. F., Duperron, S. & Gaudron, S. M. (2011) First documented record of a living solemyid bivalve in a pockmark of the Nile Deep-sea Fan (eastern Mediterranean Sea). Marine Biodiversity Records, 4, e 10. http: // dx. doi. org / 10.1017 / S 175526721100008 X","Sato, K., Watanabe, H. & Sasaki, T. (2013) A new species of Solemya (Bivalvia: Protobranchia: Solemyidae) from a hydrothermal vent in the Iheya Ridge in the Mid-Okinawa Trough, Japan. The Nautilus, 127, 93 - 100."]}

Leiopathes annosa sp. nov. (Fig. 1 a���e, 2 a���d, 3 a���h, 4 a���o, 5 a, 6 a���d) not Leiopathes glaberrima, (Esper 1972) (and later authors; see Opresko and Baron-Szabo, 2001 for synonymy) Leiopathes glaberrima, Grigg and Opresko, 1977: 242 ���261, fig. 7. Leiopathes glaberrima, Grigg 1988: 142. Leiopathes glaberrima, Chave and Malahoff, 1998: 40, fig. 93. Leiopathes glaberrima, Roark et al., 2006 *: 1���14. Leiopathes glaberrima, Parrish and Baco, 2007 *: 186. Leiopathes sp. Roark et al. 2009 *: 5204���5208. Leiopathes sp. Wagner et al., 2011: 211���225, fig. 1 k, 2 d. Leiopathes sp. Wagner et al., 2012: 76, 80, 84, 88, 90. Leiopathes sp. P 4-227 - 2 Brugler et al., 2013: 325, 327, 335, fig. 2���3. Leiopathes USNM 1070976 Brugler et al., 2013: 335, fig. 2���3. * Re-identification based on in situ photographs in HURL archives only. For all others, both specimens and in situ photographs were examined. Type locality. South of Ni���ihau Island, Main Hawaiian Islands, 21 �� 49 ��� 21 ���N 160 �� 6 ��� 37 ���W, 382 m. TABLE 1. Leiopathes specimens examined as part of this study (* = type material; NR = not reported; 1 molecular sequence data for specimen BPBM D 1871 (as P 4-227 - 2) was reported in Brugler et al. (2013) and is available from GenBank under accession numbers KF054599, KF054665 and KF054466; 2 molecular sequence data for specimen USNM 1070976 was reported in Brugler et al. (2013) and is available from GenBank under accession numbers KF054593, KF054663 and KF054467). Diagnosis. Colonies are up to 1 m in height or more, with most branches located in an approximately single plane like a fan (Fig. 1 a���c, 3 a���h). In some colonies, branches overlap considerably producing a rather thick corallum that is still planar, whereas in others all the branches are in a single plane (Fig. 1 a���c, 3 a���h). Most of the smaller branches are directed distally relative to the branches from which they arise, as well as upwards towards the top of the corallum. Branching is mostly irregularly bilateral, but uniserial in some places, with the branches tending to be directed vertically. The highest order branches are typically 1.5���6 cm in length, 1.4 mm in diameter at their midpoint with tissue (range = 1.0��� 1.8 mm), and on average spaced 0.8 cm apart (range = 0.4���1.7 cm). Polyps are variable in size (0.88���3.35 mm), arranged irregularly on all sides of the corallum on thicker branches and uniserially on terminal branches with adjacent polyps spaced 1.2���3.2 mm apart, resulting in 5���6 polyps/cm (Fig. 1). Spines are typically 75 ��m tall (range = 30���225 ��m) and 140 ��m wide at the base (range = 35���315 ��m). On the terminal branchlets (~ 0.07 mm in diameter excluding the spines), the spines can be hemispherical, balloon-shaped, irregularly knobby or with one or more round or misshapen lobes (Fig. 2 d, 4 a���o). On such terminal branchlets, 3���5 spine rows are visible in lateral view (Fig 2 d, 4 a���o). On thicker branches (0.15 mm in diameter excluding spines), the spines are more hemispherical to mound-like with fewer, but larger round lobes, and the spine arrangement in rows becomes obscured (Fig. 2 b���c). On branches thicker than 0.27 mm in diameter excluding the spines, the spines become smaller and more knob-like or somewhat conical to deltoid in shape, with little remaining evidence of lobes (Fig 2 a). The living tissues are colored bright orange (Fig. 1 a, 3 a���h). Material examined. Holotype (BPBM D 1871), south of Ni��ihau Island, Main Hawaiian Islands, 21 �� 49 ��� 21 ���N 160 �� 6 ��� 37 ���W, 382 m, HURL submersible Pisces IV, dive P 4-227, specimen 2, 2 December 2009, ethanol. Paratype (USNM 77480), bank north of Necker Island, Northwestern Hawaiian Islands, no coordinates recorded, 457 m, R/ V Townsend Cromwell, cruise 76 -06- 73, fish trawl, 15 October 1976, dry. For all other non-type material examined as part of this study see Table 1. Description of holotype. The holotype (BPBM D 1871) consists of two separate ethanol-preserved fragments collected from the distal part of a colony (Fig. 1 a���b). The colony from which the holotype fragments were collected, branches sparsely towards the center of the corallum, and becomes more dense and irregular distally, with multiple orders of branching (Fig. 1 a���b, 3 a). The first holotype fragment has a basal branch diameter of 5 mm and is approximately 31 cm long and 16 cm wide, whereas the second fragment has a basal branch diameter of 6 mm, measures 33 cm in length and 18 cm in width. On both fragments polyps are arranged irregularly on all sides of the axis on the main branch and larger branches, and uniserially on the terminal branches. Polyps are 0.88���3.35 mm in transverse diameter, and spaced 1.2���3.2 mm apart on the terminal branches, resulting in 5���6 polyps/cm. Terminal branches are typically 3���4 cm in length (range = 1.5���6 cm), 1.4 mm in width at their midpoint with tissue (range = 1.0��� 1.8 mm) and 280 ��m in width without tissue (range = 165���400 ��m). On the terminal branches, 3���4 spine rows are visible in lateral view; however, on thicker branches the spine arrangement in rows becomes obscured (Fig. 2 a���b). Spines are smooth and typically hemispherical, but spines with multiple lobes occur often (Fig. 2 c���d). On terminal branches spines are generally 70 ��m tall (range = 30���100 ��m) and 145 ��m wide (range= 70���310 ��m). On thicker branches (width = 400���810 ��m without tissue), spines range between 30���110 ��m in height and 130���315 ��m in width (Fig. 2 a). Description of paratype. The paratype (USNM 77480) consists of a dried colony that is 63 cm tall, 17 cm wide, 8 mm in basal diameter and was broken off above the base (Fig. 1 c, e). The branching is sparse towards the center of the corallum, and becomes more dense and irregular distally, with nine orders of branching (Fig. 1 c). Most of the smaller branches are directed distally relative to the branches from which they arise, as well as upwards towards the top of the corallum (Fig. 1 c, e). Branching is mostly irregularly bilateral, but uniserial in some places, with the branches tending to be directed vertically (Fig. 1 c, e). Branches coming off the main axis range between 3���5 mm in diameter. The terminal branches are 1.5���6 cm long, tend to be curved distally, and measure 400���650 ��m at their midpoint with remaining tissue. On portions without remaining tissue, terminal branches measure 280���440 ��m at their midpoint. Despite being preserved as a dry specimen, polyp remnants remain on a few terminal branchlets of the paratype (Fig. 1 e) and measure 2���2.5 mm in diameter. On the terminal branches, 4���5 spine rows are visible in lateral view; however, on thicker branches the spine arrangement in rows becomes obscured. Spines on terminal branches are smooth and range between hemispherical, balloon-shaped and irregularly knobby. On terminal branches (250���400 ��m at their midpoint without spines), spines are 55���140 ��m tall and 75���240 ��m wide at their base. On thicker branches (width = 450���675 ��m without tissue), spines range between 50���120 ��m in height, 120���260 ��m in width and tend to be mound-like, conical or deltoid in shape. Intraspecific variability and a unique morphotype. The holotype and paratype of Leiopathes annosa sp. nov. both have skeletal spines that are smooth, hemispherical, often times multi-lobed, 30���225 ��m in height and 35���310 ��m in width. In the examination of the 14 additional specimens assigned to this species (see Table 1), skeletal spine morphology ranged widely even within the same specimen, and varied from hemispherical, balloonshaped, irregularly knobby, multi-lobed, mound-like, conical and deltoid (Fig. 4 a���o). However, spines with multiple lobes were present on all specimens assigned to L. annosa sp. nov. (see Table 1). Similar to skeletal spine morphology, the dimensions of skeletal spines varied between the 16 specimens assigned to L. annosa sp. nov. (Table 1), and ranged from an average height of 55 ��m and average basal width of 125 ��m for specimen USNM 1071417, to an average height of 95 ��m and average basal width of 190 ��m for specimen USNM 99500. However, the skeletal spines of all specimens assigned to L. annosa sp. nov., ranged between 30���225 ��m in height and 35��� 310 ��m in width at the base. In contrast, one specimen from Hawaiian waters (Leiopathes sp.; USNM 52278) was unique in that its skeletal spines were (1) much smaller (average = 40 ��m; range = 20���65 ��m), (2) narrower (average = 90 ��m; range = 30���160 ��m), and (3) had almost no multi-lobed spines (Fig. 4 p). Additionally, while the polyp spacing of specimen USNM 52278 (1.2���3.05 mm) is similar to those of the specimens assigned to L. annosa sp. nov (1.2���3.2 mm), its polyps are smaller (0.70���2.10 mm vs. 0.88���3.35 mm). Furthermore, the 16 Hawaiian specimens assigned to L. annosa sp. nov. were all collected at depths ranging between 295���536 m, whereas specimen USNM 52278 was collected in much deeper waters (966 m; Table 1). Based on these substantial differences, specimen USNM 52278 represents a distinct species that is morphologically more similar to L. montana and L. glaberrima. However, a more detailed study, ideally using both morphological and molecular markers, will be needed to determine the taxonomic identity and affinities of specimen USNM 52278. Comparisons. In Hawaiian waters, Leiopathes annosa sp. nov. has previously been referred to as Leiopathes sp. (Roark et al. 2009; Wagner et al. 2011; Wagner et al. 2012; Brugler et al. 2013) and L. glaberrima (Grigg and Opresko 1977; Grigg 1988; Chave and Malahoff 1988; Roark et al. 2006; Parrish and Baco 2007). As part of this study, we reexamined in situ photographs and specimens referenced in those previous studies, and as a result reassigned those records to L. annosa sp. nov. The previous misidentification of the Hawaiian specimens as L. glaberrima was due to: (1) Hawaiian specimens were not compared to specimens from other geographic locations, and (2) diagnostic characters of L. glaberrima were largely unknown because Esper's (1792) original species description was quite brief and type material remained without further reexamination until recently (Opresko & Baron-Szabo 2001). The recent redescription of L. glaberrima has established the diagnostic characters of this species which include (1) triangular, simple and smooth spines which are 20���60 ��m tall, (2) polyps that are up to 1 mm in diameter, (3) terminal branches with a length of 1���2 cm, and (4) an irregularly branched corallum (Opresko & Baron-Szabo 2001). Leiopathes annosa sp. nov. differs from L. glaberrima in that its spines are hemispherical and frequently multi-lobed, and substantially larger (30���225 ��m) (Fig. 5 a. e). Additionally, polyps of L. annosa sp. nov. reach a larger maximum size (3.35 mm) than those in L. glaberrima (1 mm), and its terminal branches are longer (L. annosa sp. nov. colonies are fan-like and bright orange in situ (Fig. 1 a, 3 a���h), whereas living colonies of L. glaberrima are grayish-white, white, pale orange or bright orange, and branched irregularly (Bo 2008; Vertino et al. 2010; Mastrototaro et al. 2010; Bo et al. 2015). Leiopathes glaberrima has been reported to occur at depths as shallow as 37 m in the Gulf of Mexico (Opresko 2009) and as deep as 1532 m off Morocco (Grasshoff 1989), whereas L. annosa sp. nov. has a limited bathymetric range (295���536 m). Leiopathes annosa sp. nov. can be easily differentiated from all but one of the remaining seven described species of Leiopathes on the basis of the morphology of its skeletal spines (Fig. 5 a���e). In L. expansa, L. grimaldi, L. valdiviae, L. secunda, L. acanthophora and L. montana, the spines are predominantly conical, subdeltoid or relatively tall (height greater than width at the base) with a rounded apex. In none of these seven species are the spines hemispherical or multi-lobed (see summary of species characteristics as given on Table 1 in Molodtsova 2011). Hemispherical spines are, however, found in L. bullosa (Fig. 5 b, 6 f���h). The major morphological difference between L. annosa sp. nov. and L. bullosa is the fact that in the former the spines, especially those on the terminal branches, can be multi-lobed (Fig. 4 a���o, 6 c���d), whereas in L. bullosa multi-lobed spines have not been reported (Fig. 6 f���h). Other differences between the two species exist in the maximum thickness of the terminal branches (0.2 mm in L. bullosa and 0.4 mm in L. annosa sp. nov.), maximum length of the terminal branches (2 cm in L. bullosa and 6 cm in L. annosa sp. nov.), maximum size of the spines (140 ��m in L. bullosa and 225 ��m in L. annosa sp. nov.), arrangement of the spines (clearly in rows in L. bullosa, and not in discernable rows in L. annosa sp. nov.), maximum size of the polyps (2 mm in L. bullosa and 3.35 mm in L. annosa sp. nov.), and in the density of the polyps (3���5 /cm in L. bullosa and 5���6 /cm in L. annosa). L. annosa sp. nov. forms colonies that are large, flabellate, with relatively thick branches (Fig. 1 a, 3 a���h). In comparison, the branches of L. bullosa are much thinner, and even though individual clusters of branches of the holotype were described as being somewhat planar, the holotype consists of small fragments, and therefore the size, shape, and branching pattern of intact colonies of L. bullosa are unknown. Previous studies in Hawai��i. Several studies have reported on different ecological aspects of L. annosa sp. nov. in Hawai���i (identified at the time as L. glaberrima or Leiopathes sp.), including on its: (1) growth rate and longevity (Roark et al. 2006; Roark et al. 2009), (2) population density (Grigg 1988), (3) associated fish assemblages (Parrish 2006), and (4) sexual reproduction (Wagner et al. 2011). Roark et al. (2006, 2009) used highresolution skeletal radiocarbon measurements and estimated average radial growth rates of 5���13 ��m/year for this species, corresponding to longevities ranging between 350���4,265 years, thus making this the longest living marine organism studied to date. Furthermore, Roark et al. (2009) concluded that growth rates are not uniform throughout the lifespan of this species, with faster initial growth rates that gradually decrease with age. Grigg (1988) examined the population densities of various dominant deep-water corals (365���400 m) off O��ahu Island, and noted average densities of 0.002���0.003 colonies/m 2 for this species. Parrish (2006) studied fish assemblages within deep-water corals (300���500 m) across various Hawaiian Islands, and noted that various fish occasionally use taller colonies of L. annosa sp. nov. as shelter. Wagner et al. (2011) used histological techniques to study the reproductive anatomy of eight Hawaiian antipatharian species, including a sample of the L. annosa sp. nov. holotype, and noted that it had both previtellogenic and vitellogenic oocytes within its primary mesenteries. Finally, as part of a study on the evolutionary history of the order Antipatharia, Brugler et al. (2013) provided mitochondrial and nuclear DNA sequence data for two specimens (USNM 1070976 and BPBM D 1871 as P 4-227 - 2) that are assigned to L. annosa sp. nov. here (see Table 1 for GenBank accession numbers). Molecular sequences of these two specimens had identical sequences in the intergenic region between nad 5 and nad 1, and between cox 3 -cox 1, but did differ in a single position in the intergenic region between trnW and nad 2 (Brugler et al. 2013). Future studies will need to include a molecular component in order to determine whether the morphological analysis presented in this study holds up, as well as to elucidate the taxonomic relationships among the various species within the genus Leiopathes. Etymology. From the latin ��� annosa ��� (long-lived) in reference to the extreme longevity estimates reported for this species (350���4,265 years; Roark et al. 2006; Roark et al. 2009). Distribution. Under the name L. glaberrima or Leiopathes sp., L. annosa sp. nov., has previously been reported throughout the Hawaiian Islands (Grigg & Opresko 1977; Chave & Malahoff 1998; Parrish and Baco 2007; Roark et al. 2006; Roark et al. 2009; Wagner et al. 2011; Wagner et al. 2012; Brugler et al. 2013), as well as from Johnston Atoll (Chave & Malahoff 1998) at depths between 300��� 500 m. Specimens examined as part of this study were collected throughout the Hawaiian Archipelago from Hawai���i Island (19 �� 30 '00''N 155 ��00'00''W) to Salmon Bank (27 ��01' 10 ''N 176 �� 31 ' 33 ''W) at depths between 295���536 m (Table 1). Leiopathidae biogeography. Within the monogeneric family Leiopathidae, seven species have been recorded from a limited geographic region, including (1) L. expansa from Madeira (Johnson 1899), (2) L. montana from the Great Meteor Seamount (Molodtsova, 2011), (3) L. grimaldii from two locations in the northeastern Atlantic (Roule 1905), (4) L. valdiviae from the Nicobar Islands (Pax 1915; Pax 1922), (5) L. secunda from Tasmania and New Zealand (Opresko 1998; Cairns et al. 2009), (6) L. acanthophora from South Australia and New Zealand (Opresko 1998; Cairns et al. 2009), and (7) L. bullosa from South Australia and New Zealand (Opresko 1998; Cairns et al. 2009), (Fig. 7). In contrast, L. glaberrima has been recorded from widespread geographic locations, including from throughout the Mediterranean (Gray 1857; Opresko & Baron-Szabo 2001; Bo 2008; Carlier et al. 2009; Sinninger & Pawlowski 2009; Deidun et al. 2010; Vertino et al. 2010; Mastrototaro et al. 2010; D���Onghia et al. 2012; Brugler et al. 2013; Angeletti et al. 2014; Bo et al. 2014 a, 2014 b, 2015; Fabri et al. 2014; Ingrassia et al. in press), the northeastern Atlantic (Johnson 1899; Grasshoff 1985, 1989; Brito & Oca��a 2004), the northwestern Atlantic and Caribbean (Brook 1889; Cairns et al. 1993; Williams et al. 2006, 2007; Opresko 2009) (Fig. 7). To, Published as part of Wagner, Daniel & Opresko, Dennis M., 2015, Description of a new species of Leiopathes (Antipatharia: Leiopathidae) from the Hawaiian Islands, pp. 277-289 in Zootaxa 3974 (2) on pages 278-287, DOI: 10.11646/zootaxa.3974.2.11, http://zenodo.org/record/244528, {"references":["Opresko, D. M. & Baron-Szabo, R. C. 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Family Solemyidae Gray, 1840 Genus Solemya Lamarck, 1818 Type species. Solemya mediterranea (Lamarck, 1818) = Tellina togata Poli, 1795 (by subsequent designation, Children, 1823). Remarks. Our species is included in Solemya Lamarck, 1818, because of its internal ligament (Taylor et al. 2008; Kamenev 2009; Oliver et al. 2011). It is not a member of the genus Acharax Dall, 1908, because this taxon has an external ligament positioned above the thickened shell margin, developed probably as a support for the ligament nymphs (Amano & Ando 2011; Oliver et al. 2011; Taviani et al. 2011). Solemya has an internal ligament with a distinct resilium supported on a chondrophore. Fossil solemyids are morphologically very conservative, having a cylindrical shape with inequilaterally positioned beaks and an elongated shell anterior (e.g. Logan 1967; Duff 1978; Liljedahl 1984; Cope 1996; Kiel et al. 2008a; Bailey 2011). This conservatism (e.g. Taylor & Glover 2010) is maintained not only at the genus, but also at the species level, as testified by the presence of cryptic modern species of Acharax (Neulinger et al. 2006). Thus, the taxonomy of solemyids is difficult, and there are few characters useful for species discrimination. Subgenus Petrasma Dall, 1908 Type species. Solemya borealis Totten, 1834 Remarks. We have included our species in the subgenus Petrasma Dall, 1908, because of the presence of a chondrophore supporting an oblique buttress adjoining the anterior side of the posterior adductor muscle scar, and small ligament demipads elongated along the dorsal margin and developed in front of the chondrophore. A buttress in front of the posterior adductor muscle scar is also present in the subgenera Solemyarina Iredale, 1931, Zesolemya Iredale, 1939, and Austrosolemya Taylor, Glover & Williams 2008, all of which have large ligament demipads perpendicular to the dorsal margin in front of the chondrophore (e.g. Kamenev 2009, fig. 4–6). For more detailed discussion of subgeneric differences in the genus Solemya see Taylor et al. (2008), Kamenev (2009) and Oliver et al. (2011)., Published as part of Hryniewicz, Krzysztof, Little, Crispin T. S. & Nakrem, Hans Arne, 2014, Bivalves from the latest Jurassic-earliest Cretaceous hydrocarbon seep carbonates from central Spitsbergen, Svalbard, pp. 1-66 in Zootaxa 3859 (1) on pages 7-8, DOI: 10.11646/zootaxa.3859.1.1, http://zenodo.org/record/4930112, {"references":["Gray, J. E. (1840) Shells of molluscan animals. In: Synopsis of the contents of the British Museum, ed. 42. Woodfall & Son, London, pp. 105 - 152.","Lamarck, J. B. (1818) Histoire naturelle des animaux sans vertebres. Vol. 5, Deterville, Paris, 612 pp.","Poli, G. S. (1795) Testacea utriusque Siciliae eorumque historia et anatome tabulis aenis illustrata. Rusconi & Parma, Chiaje, Neapoli, i - lxxvi, 1 - 264, pls. 9 - 39.","Children, J. G. (1823) Lamarck's genera of shells. Quaterly Journal of Science, Literature & Arts, 14, 298 - 322.","Taylor, J. D., Glover, E. A. & Williams, S. T. (2008) Ancient chemosynthetic bivalves: systematics of Solemyidae from eastern and southern Austalia. Memoirs of Queensland Museum - Nature, 54, 75 - 104.","Kamenev, G. M. (2009) North Pacific species of the genus Solemya Lamarck, 1818 (Bivalvia: Solemyidae) with notes on Acharax johnsoni (Dall, 1891). Malacologia, 51, 233 - 261. http: // dx. doi. org / 10.4002 / 040.051.0202","Oliver, G. P., Rodrigues, C. F. & Cunha, M. R. (2011) Chemosymbiotic bivalves from the mud volcanoes of the Gulf of Cadiz, NE Atlantic, with descriptions of new species of Solemyidae, Lucinidae and Vesicomyidae. ZooKeys, 113, 1 - 38. http: // dx. doi. org / 10.3897 / zookeys. 113.1402","Dall, W. H. (1908) Reports on the dredging operations off the west coast of Central America to the Galapagos to the west coast of Mexico, and in the Gulf of California, in charge of Alexander Agassiz carried on by the U. S. Fish Commission Steamer \" Albatross \" during 1891, Lieut. - Commander Z. L. Tanner, U. S. N., commanding. XXXVII; Reports on the scientific results of the expedition to the Eastern tropical Pacific, in charge of Alexander Agassiz by the U. S. Fish Commission Steamer \" Albatross \", from October, 1904 to March 1905, Lieut. - Commander L. M. Garrett, commanding. XUV; Reports on the Mollusca and Brachiopoda. Bulletin of the Museum of Comparative Zoology, 43, 205 - 487, pls. 1 - 22.","Amano, K. & Ando, H. (2011) Giant fossil Acharax (Bivalvia: Solemyidae) from the Miocene of Japan. The Nautilus, 125, 207 - 212.","Taviani, M., Angeletti, L. & Ceregato, A. (2011) Chemosynthetic bivalves of the family Solemyidae (Bivalvia, Protobranchia) in the Neogene of the Mediterranean Basin. Journal of Palentology, 85, 1067 - 1076. http: // dx. doi. org / 10.1666 / 10 - 119.1","Logan, A. (1967) The Permian Bivalvia of Northern England. Palaeontographical Society Monographs, 121, 1 - 72, pls. 1 - 10.","Duff, K. L. (1978) Bivalvia from the English lower Oxford Clay (Middle Jurassic). Palaeontographical Society Monograph s, 132, 1 - 137, pls. 1 - 13.","Liljedahl, L. (1984) Janeia silurica, a link between nuculoids and solemyoids (Bivalvia). Palaeontology, 27, 693 - 698.","Cope, J. C. W. (1996) Early Ordovician (Arenig) bivalves from the Llangynog Inlier, South Wales. Palaeontology, 39, 979 - 1025.","Kiel, S., Amano, K. & Jenkins, R. G. (2008 a) Bivalves from Cretaceous cold seep deposits on Hokkaido, Japan. Acta Palaeontologica Polonica, 53, 525 - 537. http: // dx. doi. org / 10.4202 / app. 2008.0310","Bailey, J. B. (2011) Paleobiology, paleoecology, and systematics of Solemyidae (Mollusca: Bivalvia: Protobranchia) from the Mazon Creek lagerstatte, Pennsylvanian of Illinois. Bulletins of American Paleontology, 382, 1 - 72, pls. 1 - 5.","Taylor, J. D. & Glover, E. A. (2010) Chemosymbiotic Bivalves. In: Kiel, S. (Ed.), The vent and seep biota. Aspects from microbes to ecosystems. Topics in Geobiology, Springer, Dordrecht, Heidelberg, London, New York, pp. 107 - 133.","Neulinger, S. C., Sahling, H., Suling, J. & Imhoff, J. F. (2006) Presence of two phylogenetically distinct groups in the deep-sea mussel Acharax (Mollusca: Bivalvia: Solemyidae). Marine Ecology Progress Series, 312, 161 - 168. http: // dx. doi. org / 10.3354 / meps 312161","Totten, J. G. (1834) Description of some new shells, belonging to the coast of New England. American Journal of Science and Arts, 26, 366 - 369.","Iredale, T. (1931) Australian Molluscan notes. No. 1. Records of the Australian Museum, 18, 201 - 235, pls. 22 - 25. http: // dx. doi. org / 10.3853 / j. 0067 - 1975.18.1931.725","Iredale, T. (1939) Mollusca. Part I. Scientific Reports of the Great Barrier Reef Expedition 1928 - 1929, 5, 209 - 245, pls. 1 - 7."]}

Genus Calliax de Saint Laurent, 1973 Type species. — Callianassa (Callichirus) lobata de Gaillande & Lagardère, 1966. Extant species included. — Three species (including one referred species but not formally named): Calliax doerjesti Sakai, 1999 (Figs 3A–B); Calliax lobata (de Gaillande & Lagardère, 1966) (Figs 3C–D); Calliax sp. sensu Taviani et al. (2013). Fossil species included.– Calliax michelottii (A. Milne Edwards, 1860) comb. nov. More fossil occurrences in open nomenclature are recognized (see Table 1). Diagnosis. Carapace lacking dorsal oval; rostrum short, with blunt tip, rostral spine absent. Pleonal segment 2 longest, no lateral tufts of setae on segments 3–5. Telson slightly wider than long, lateral margin curved, posterior margin straight or slightly convex. Eyestalk about twice as long as wide, slightly flattened dorso-ventrally; cornea small, weakly pigmented. A1 peduncle shorter than that of A2. Mxp1 epipod tapering anteriorly. Mxp2 with small, leaf-like epipod. Mxp3 subpediform (sensu Ngoc-Ho 2003), propodus and dactylus rounded, exopod absent. P1 unequal, dissimilar. Major P1 propodus rectangular, usually longer than high, fixed finger shorter than manus, with a double ridge accompanied by a furrow extending onto manus and parallel to the lower margin of propodus. Fixed finger as long as dactylus in major P1, shorter than dactylus in minor P1, with wide proximal gap and large triangular proximal tooth on cutting edge. Major P1 carpus shorter than high, distinctly shorter than propodus. Major P1 merus longer than high, keeled, lower margin armed with small spines. P3 with small proximal heel on propodus, P5 subchelate. Paired arthrobranch on Mxp3 and P1–4. Male and female Plp1 uniramous male and female Plp2 biramous, all lacking appendix interna, male Plp2 with appendix masculina overreaching endopod. Plp3–5 biramous, foliaceous, appendix interna finger-like in both sexes. Uropodal endopod and exopod slightly longer than telson, with rounded posterior margin; exopod with dorsal plate terminating in short distal setal row [emended from Ngoc-Ho (2003: 489) with characters on major P1]. Remarks on the taxonomy. Calliax has a complex taxonomic history. The genus was erected by de Saint Laurent (1973) with Callianassa lobata de Gaillande & Lagardère, 1966, as the type species. Since then the concept of the genus has been changed several times (cf. Manning & Felder 1991; Sakai 1999, 2005, 2011; Ngoc- Ho 2003; Hyžný 2012; see also Dworschak 2007). Here the view of Ngoc-Ho (2003) and Sakai (2011) is adopted, and thus, only two formally described extant species are recognized. Discussion on distinguishing Calliax from related taxa based on soft-part morphology was provided by Ngoc-Ho (2003) and will not be repeated here. When dealing with chelipeds the two known extant species of Calliax can be characterized by unequal and dissimilar chelae, from which the minor one has „fixed finger shorter than and separated from the dactylus by a wide gap, bearing a large triangular proximal tooth (Ngoc-Ho 2003: 490)“. This morphology approaches a subchelate cheliped state. Comparison of the illustrated major P1 propodus of C. lobata (de Gaillande & Lagardère 1966: fig. 2a; de Saint Laurent & Božić 1976: fig. 23; Ngoc-Ho 2003: fig. 17D), C. doerjesti (Sakai 1999: figs. 28, 29b) and Calliax sp. (Taviani et al. 2013: fig. 8) clearly shows consistency in its general shape, i.e. propodus is rectangular and usually longer than high and on its lateral surface the fixed finger possesses two ridges accompanied by furrow (or furrows) parallel to the lower margin of propodus and extending onto manus. The ridges are visible especially when viewing under low angle light (Fig. 2C). There are several distinct setal pores (accompanied by tubercles) arranged obliquely across the lateral surface of propodus. In Calliax the major P1 carpus is always much shorter than manus, with rounded proximo-lower margin (Fig. 3). The merus is longer than high with a distinct meral keel and its lower margin is usually armed with small spines. The above mentioned combination of the characters of minor chela and major propodus, carpus and merus is unique for Calliax; thus, the genus can be identified on the basis of chelipeds alone. The number of spines on the lower margin of P1 merus may vary between respective members of Calliax and may help in distinguishing taxa at the species level, although possible variation has not been studied in detail yet. Regarding the number of meral spines, there are discrepancies in the literature. Sakai (1999: 114) in the description of C. doerjesti mentioned that the lower margin of the merus was “armed with three interspaced denticles”. One of the figures (Sakai 1999: fig. 29b) indeed shows three small spines, however, in the other one (Sakai 1999: fig. 28) depicting the same specimen (holotype) the merus is armed with seven spines (Fig. 3a). Ngoc-Ho (2003: fig. 17D; note that the published figure depicts the right major chela, whereas the caption refers to it as the left one) figured the holotype (male) of C. lobata with seven spines on the merus and de Saint Laurent & Božić (1976: fig. 23a) figured a female specimen of C. lobata also with seven spines. Calliax cf. C. lobata, examined and figured herein (Figs 2, 3E–F), possesses only four blunt spines, presumably mirroring its small size (Fig. 4). Remarks on the fossil record. Articulated chelipeds are relatively sparse in the fossil state and often only isolated propodi are at hand. In this respect, the major P1 propodus of Calliax is distinct enough to be differentiated from all other ghost shrimp genera. It must be stressed, however, that the more cheliped elements are found, the more secure assignment at the genus level can be provided. The best preserved and most numerous remains of Calliax in the fossil record belong to species originally described as Callianassa michelottii (Figs 5–10). It is discussed in detail below. Feldmann et al. (2005) reported several isolated cheliped elements from the Miocene of the Navidad Formation of Chile as Callianassoidea sp. 1. Although the authors stated that it does not resemble any callianassoid genus (Feldmann et al. 2005: 431), the figured material (Feldmann et al. 2005: fig. 2A) exhibits striking similarities with Calliax as discussed herein. Interestingly, Callianassa szobensis Müller, 1984, which is herein considered a junior subjective synonym of C. michelottii (see below) and hence a member of Calliax, is mentioned by Feldmann et al. (2005) as similar to their Callianassoidea sp. 1. The same locality also yielded another specimen which has been identified as Callichirus sp. The figured propodus (Feldmann et al. 2005: fig. 2A) shows a relatively short manus; however, the fixed finger with ridges and a furrow indicates its affinities to Callianassoidea sp. 1 (Feldmann et al. 2005: fig. 2D). Both specimens are treated here as Calliax sp. 1. Feldmann et al. (2011) reported fragmented material from the Miocene of Tierra del Fuego (Argentina) as “Cheliped Form B” of indeterminate callianassoid. As already noted by Hyžný & Hudáčková (2012: 13), the minor chela exhibits remarkable similarities to Calliax, (Feldmann et al. 2011: fig. 5E) showing a fixed finger shorter than the dactylus and separated from it by a wide gap with a proximal tooth (cf. Ngoc-Ho 2003: 490). The major P1 propodus (Feldmann et al. 2011: fig. 5B), however, does not possess the ridges on the fixed finger. It is fairly likely that the two specimens do not belong to the same taxon, as they were not found associated with each other. For the purposes of this contribution only the minor chela is referred to here as Calliax sp. 2. Charbonnier et al. (2013) reported a single near-complete propodus from the Paleocene of Pakistan identified as a minor chela of Calliax. Indeed, the specimen shows all features typical for minor chelae of the genus (Charbonnier et al. 2013: fig. 2) as discussed above. This occurrence is considered the oldest confirmed fossil record of the genus, treated here as Calliax sp. 3. Callianassa whiteavesi Woodward, 1896 from the Campanian of Canada (Woodward 1896; Feldmann & McPherson 1980; Schweitzer et al. 2003) was assigned to Calliax by Schweitzer et al. (2003). The material is rich and sufficiently preserved to reconstruct both chelipeds (Feldmann & McPherson 1980). The species differs markedly from any Calliax species. It does not possess the typically shaped minor cheliped as discussed above, nor has it parallel ridges on the base of the fixed finger. Moreover, some specimens exhibit a rather deep dactylus, a character not observed in Calliax. As a result, the species is excluded from Calliax herein. Until the type material is restudied we suggest to keep the species under Callianassa sensu lato. Swen et al. (2001) reported a single fragmentary right propodus from the Maastrichtian of the Netherlands as “ Calliax ? sp.”. The material is too fragmentary for resolving its generic status. The oblique development of the ridge at the base of fixed finger (Swen et al. 2001: fig. 5.3), however, points to closer affinities to Eucalliax or Calliaxina rather than Calliax. Van Bakel et al. (2006) listed in a table of Cenozoic decapods from Belgium the presence of Calliax in the Miocene strata. The material was recently described as a new member of the family Axiidae (Fraaije et al. 2011). Occurrence and distribution. Paleocene–Holocene. Two formally described extant species are known from West Atlantic (Florida) and Mediterranean (Sakai 2011). Based on the reports discussed above (Feldmann et al. 2005, 2011), the geographical distribution of the genus was much wider during the Miocene than today, and the genus was apparently also present in the East Pacific (see below). All occurrences are reviewed in Table 1., Published as part of Hyžný, Matúš & Gašparič, Rok, 2014, Ghost shrimp Calliax de Saint Laurent, 1973 (Decapoda: Axiidea: Callianassidae) in the fossil record: systematics, palaeoecology and palaeobiogeography, pp. 37-57 in Zootaxa 3821 (1) on pages 42-45, DOI: 10.11646/zootaxa.3821.1.3, http://zenodo.org/record/4919918, {"references":["Saint Laurent, M. de (1973) Sur la systematique et la phylogenie des Thalassinidea: definition des familles des Callianassidae et des Upogebiidae et diagnose de cinq genres nouveaux (Crustacea Decapoda). Comptes Rendus Hebdomadaires de Seances de l'Academie des Sciences, Serie D, 277, 513 - 516.","de Gaillande, D. & Lagardere, J. - P. (1966) Description de Callianassa (Callichirus) lobata nov. sp. (Crustacea Decapoda Callianassidae). Recueil des Travaux de la Station Marine d'Endoume, 40, 259 - 265.","Sakai, K. (1999) Synopsis of the family Callianassidae, with keys to subfamilies, genera and species, and the description of new taxa (Crustacea: Decapoda: Thalassinidea). Zoologische Verhandelingen, Leiden, 326, 1 - 152.","Taviani, M., Angeletti, L., Ceregato, A., Foglini, F., Froglia, C. & Trincardi, F. (2013) The Gela Basin pockmark field in the strait of Sicily (Mediterranean Sea): chemosymbiotic faunal and carbonate signatures of postglacial to modern cold seepage. Biogeosciences, 10, 4653 - 4671. http: // dx. doi. org / 10.5194 / bg- 10 - 4653 - 2013","Milne Edwards, A. (1860) Histoire des Crustaces podophthalmaires fossiles et Monographie des Decapodes Macroures de la famille des Thalassinens. Annales des Sciences Naturelles, 4 e Serie 14, 129 - 357.","Ngoc-Ho, N. (2003) European and Mediterranean Thalassinidea (Crustacea, Decapoda). Zoosystema 25, 439 - 555.","Manning, R. B. & Felder, D. L. (1991) Revision of the American Callianassidae (Crustacea: Decapoda: Thalassinidea). Proceedings of the Biological Society of Washington, 104, 764 - 792.","Sakai, K. (2005) Callianassoidea of the world (Decapoda: Thalassinidea). Crustaceana Monographs, 4, 1 - 285.","Sakai, K. (2011) Axioidea of the World and a Reconsideration of the Callianassoidea (Decapoda, Thalassinidea, Callianassida). Crustaceana Monographs, 13, 1 - 520. http: // dx. doi. org / 10.1163 / 9789047424185","Dworschak, P. C. (2007) Book review. Sakai, K. 2005. Callianassoidea of the world (Decapoda: Thalassinidea). Crustaceana Monographs 4. Journal of Crustacean Biology, 27 (1), 158 - 169.","Saint Laurent, M. de & Bozic, B. (1976) Diagnoses et tableau de determination des Callianasses de l'Atlantique nord oriental et de Mediterranee (Crustacea, Decapoda, Callianassidae). Thalassia Jugoslavica, 8, 15 - 40.","Feldmann, R. M., Schweitzer, C. E. & Encinas, A. (2005) New decapods from the Navidad Formation (Miocene) of Chile. Journal of Crustacean Biology, 25, 427 - 449. http: // dx. doi. org / 10.1651 / c- 2547","Muller, P. (1984) Decapod Crustacea of the Badenian. Geologica Hungarica, Series Palaeontologica, 42, 3 - 317.","Feldmann, R. M., Schweitzer, C. E., Casadio, S. & Griffin, M. (2011) New Miocene Decapoda (Thalassinidea; Brachyura) from Tierra del Fuego, Argentina: Paleobiogeographic implications. Annals of Carnegie Museum, 79, 91 - 123. http: // dx. doi. org / 10.2992 / 007.079.0202","Hyzny, M. & Hudackova, N. (2012) Redescription of two ghost shrimps (Decapoda: Axiidea: Callianassidae) from the Middle Miocene of the Central Paratethys: systematics, intraspecific variation, and in situ preservation. Zootaxa, 3210, 1 - 25.","Charbonnier, S., Garassino, A., Pasini, G., Metais, G., Merle, D., Bartolini, A., Brohi, I. A., Solangi, S. H., Lashari, R. A., Welcomme, J. - L. & Marivaux, L. (2013) Early Paleogene decapod crustaceans ftom the Sulaiman and Kirthar Ranges, Pakistan. Annales de Paleontologie, 99, 101 - 117. http: // dx. doi. org / 10.1016 / j. annpal. 2012.12.003","Woodward, H. (1896) On some podophthalmatous Crustacea from the Cretaceous Formations of Vancouver and Queen Charlotte islands. Quarterly Journal of the Geological Society of London, 52, 221 - 228. http: // dx. doi. org / 10.1144 / gsl. jgs. 1896.052.01 - 04.11","Feldmann, R. M. & McPherson, C. B. (1980) Fossil decapod crustaceans of Canada. Geological Survey of Canada, Paper 79 - 16, 1 - 20.","Schweitzer, C. E., Feldmann, R. M., Fam, J. M., Hessin, W. A., Hetrick, S. W., Nyborg, T. G. & Ross, R. L. M. (2003) Cretaceous and Eocene decapod crustaceans from southern Vancouver Island, British Columbia, Canada. NRC Research Press, Ottawa, Ontario, 66 pp.","Swen, K., Fraaije, R. H. B. & van der Zwaan, G. J. (2001) Polymorphy and extinction of the Late Cretaceous burrowing shrimp Protocallianassa faujasi and first record of the genera Corallianassa and Calliax (Crustacea, Decapoda, Thalassinoidea) from the Cretaceous. Contributions to Zoology, 70, 85 - 98.","Van Bakel, B. W. M., Fraaije, R. H. B. & Jagt, J. W. M. (2006) Synopsis of Ceonozoic decapods crustaceans from Belgium. Revista Mexicana de Ciencias Geologicas, 23, 370 - 374.","Fraaije, R. H. B., van Bakel, B. W. M., Jagt, J. W. M. & Mollen, F. H. (2011) A new axiid (Crustacea: Decapoda: Axiidea) from the Neogene of Belgium and the Netherlands. Neues Jahrbuch fur Geologie und Palaontologie, Abhandlungen, 260 (2), 157 - 163. http: // dx. doi. org / 10.1127 / 0077 - 7749 / 2011 / 0167"]}