Orzeliscus belopus subsp. sensu Kaczmarek, Bartels, Roszkowska & Nelson, 2015, sensu

85. Orzeliscus belopus Du Bois-Reymond Marcus, 1952 sensu lato

Orzeliscus belopus, n. sp. (Du Bois-Reymond Marcus 1952)

Orzeliscus belopus (Du Bois-Reymond-Marcus, 1952) (Morgan & Lampard 1986a, Renaud-Mornant & Jouin 1965)

Orzeliscus belopus du Bois-Reymond Marcus (Coull 1970)

Orzeliscus belopus Bois-Reymond M., 1952 (Renaud-Mornant 1970b)

Orzeliscus belopus Bois-Reymond Marcus, 1952 (Pollock 1971, Renaud-Mornant & Gourbault 1984) Orzeliscus belopus Bois-Reymond Marcus (McIntyre & Murison 1973)

Orzeliscus cf. belopus du Bois-Reymond Marcus, 1952 (McKirdy et al. 1976, Bertolani 1987, Suzuki & Kristensen 2014)

Orzeliscus belopus du Bois-Reymond Marcus 1952 (Grimaldi de Zio et al. 1979) Orzeliscus belopus, Du Bois Reymond Marcus 1952 (Grimaldi de Zio et al. 1980b, Grimaldi de Zio et al. 1983b)

Orzeliscus belopus (Grimaldi de Zio et al. 1980a, Kristensen & Higgins 1989, Accogli et al. 2009) Terra typica: Atlantic Ocean (Brazil, South America)

Adriatic Sea:

• 42°07′18′′N, 15°29′49′′E; 0 m bsl: [FAO37] Italy, Foggia Province, Tremiti Islands, San Domino Island, Cala delle Arene. Grimaldi de Zio et al. (2003)

• 40°57′30′′N, 17°17′30′′E [40°58′N, 17°16E]; 0 m bsl: [FAO37] Italy, Bari Province, Cala Corvina [Cala Corvino], intertidal, medium sand, mainly calcareous. Grimaldi de Zio et al. (1983b)

• 40°51′20′′N, 17°25′12′′E [40°50′N, 17°28′E]; 0 m bsl: [FAO37] Italy, Brindisi Province, Torre Canne, intertidal, medium-fine sand. Grimaldi de Zio et al. (1983b)

• 40°50′N, 17°28′E; 0 m bsl: [FAO37] Italy, Brindisi Province, Torre Canne, Masseria Fiume Morello, beach, tidal zone to minimum level of low tide, sand. Grimaldi de Zio et al. (1979)

• 40 ° 27.17′N, 18°14.58′E [40°22′N, 18°20′E]; 20 m bsl: [FAO37] Italy, Lecce Province, Ceisne (Marne protected area), fine sand. Accogli et al. (2011)

• 40°15′12′′N, 18°26′30′′E; 0 m bsl: [FAO37] Italy, Lecce Province, Torre S. Andrea [Sant’andrea], intertidal, medium-fine sand, mainly siliceous (60%). Grimaldi de Zio et al. (1983b)

• 40°13′42′′N, 18°27′40′′E; 0 m bsl: [FAO37] Italy, Lecce Province, Frassanito, intertidal, medium-fine sand, mainly siliceous (61%). Grimaldi de Zio et al. (1983b)

Atlantic Ocean:

• 57°50′N, 05°41′W; 0 m bsl: [FAO27] UK, Scotland, Highland County, Loch Ewe, Firemore Bay, sand. Pollock (1971)

• 57°50′N, 05°41′W; mean high water neaps to mean low water neaps: [FAO27] UK, Scotland, Highland County, Loch Ewe, Firemore Bay, flatfish nursery ground, small pocked beach, beach or subtidal zone, 4–16 cm deep in sand, sand. McIntyre & Murison (1973)

• 55°52′N, 04°53′W; 0 m bsl: [FAO27] UK, Scotland, North Ayrshire County, Clyde Sea, Meigle Bay, Cunninghame, sand from low and mid-tide level. Morgan & Lampard (1986a)

• 54°48′N, 04°57′W; 0 m bsl: [FAO27] UK, Scotland, Dumfries and Galloway County, Clyde Sea, Sandhead, sand from low and mid-tide level. Morgan & Lampard (1986a)

• 54°44′N, 04°55′W; 0 m bsl: [FAO27] UK, Scotland, Dumfries and Galloway County, Clyde Sea, New England Bay, sand from low and mid-tide level. Morgan & Lampard (1986a)

• 44°41′N, 01°03′W; 0 m bsl: [FAO27] France, Aquitaine Region, NE Arcachon Bay, Graveyron channel, Amphioxus sand. Renaud-Mornant & Jouin (1965)

• 44°35′N, 01°14′W; 0 m bsl: [FAO27] France, Aquitaine Region, entrance to Arcachon Bay, lagoon of Banc d'Arguin, Amphioxus sand. Renaud-Mornant & Jouin (1965)

• 27°30′N, 80°20.0′W / 27°30′N, 80°20′W; 5 m bsl: [FAO31] USA, Florida State, St. Lucie County, the channel of the Intracoastal Waterway in the Indian River near marker 175 about 1.6 km N of Fort Pierce, medium to coarse sand mixed with shell hash. Kristensen & Higgins (1989)

• 27°30.0′N, 80°17.0′W / 27°30′N, 80°17′W; 15 m bsl: [FAO31] USA, Florida State, St. Lucie County, 8.4 km E of North Hutchinson Island, near Fort Pierce, coarse quartz sand mixed with shell hash (Amphioxus sand). Kristensen & Higgins (1989)

• 4°20′S to 4°50′S, 36°00′W to 37°15′W [4°35′S, 36°38′W]; 20–100 m bsl: [FAO41] Brazil, continental shelf off Ceará State and Rio Grande Do Norte State, fine to coarse quartz sand with detritus, and terrigenous mud. da Rocha et al. (2013)

• 8°25′10′′S to 8°25′20′′S, 34°55′00′′W [08°25′S, 34°58′W]; 0 m bsl: [FAO41] Brazil, Pernambuco State, Cupe beach, biogenic gravel or sandy-muddy sediment. da Rocha et al. (2013)

• 23°47′S, 45°22′W; 3–5 m bsl; [FAO41] Type Locality: Brazil, São Paulo State, coast of Island of São Sebastião, near Ilhabela, medium fine detritus-rich sand. Du Bois-Reymond Marcus (1952)

Balearic Sea:

• 40°49′N, 08°30′E; 5 and 7 m bsl: [FAO37] Italy, Sardinia, Sassari Province, Gulf of Asinara, medium and medium fine sand. D’Addabbo Gallo et al. (2007)

Caribbean Sea:

• 17°47′30′′N, 64°37′W; 3–4 m bsl: [FAO31] U.S. Virgin Islands, N of the eastern end St. Croix Island, between patch reefs on the NE side of Buck Island, coral sand. Pollock (1982)

• 16°12′N, 61°29′W; 0 m bsl: [FAO31] Guadeloupe Islands, Grande-Terre Island, Le Gosier, beach, intertidal, fine sand (mean diameter 0.225 mm). Renaud-Mornant & Gourbault (1984)

Celtic Sea:

• 48°44′N, 03°59′W; 0 m bsl: [FAO27] France, Brittany Region, Roscoff, in front of the Biological Station, upper tidal zone, area of minimum oil pollution, 6–10 cm deep in sand, beach sand (diameter 0.24–0.26 mm). Renaud-Mornant & Gourbault (1980)

Coral Sea:

• 22°00′30′′S, 165°56′48′′E; 4 m bsl: [FAO71] New Caledonia, Saint Vincent Bay, Grand Ténia Island, coarse sand, gray sand or sediment in seagrass. Renaud-Mornant (1967b)

• 22°02′40′′S, 166°01′00′′E; 8 m bsl: [FAO71] New Caledonia, Saint Vincent Bay, “Coupe V 13 B de la radiale, coarse sand, gray sand or sediment in seagrass. Renaud-Mornant (1967b)

Indian Ocean:

• 03°57′N, 73°25′E; 9 m bsl: [FAO51] Republic of the Maldives, Male Sud Atoll, coarse sand. Gallo et al. (2007)

Ionian Sea:

• 40°22′25′′N, 17°18′18′′E [40°21′N, 17°22′E]; 0 m bsl: [FAO37] Italy, Tarento Province, Marina di Pulsano. Grimaldi de Zio et al. (2003)

• 40°17′45′′N, 17°33′38′′E; 0 m bsl: [FAO37] Italy, Tarento Province, Campo Marino. Grimaldi de Zio et al. (2003)

• 40°16′10′′N, 17°53′08′′E; 0 m bsl: [FAO37] Italy, Lecce Province, Porto Cesareo, intertidal, on the shoreline, medium-fine sand (max 0.354 mm). Grimaldi de Zio et al. (1980b, 1983b)

• 40°11′N, 17°48′E; 0-20 m bsl: [FAO37] Italy, Bari Province, Calo Colombo, coarse sand or fine sand mixed with mud. Accogli et al. (2009)

• 39 ° 59.38′N, 18°25.01′E ; 20 m bsl: [FAO37] Italy, Lecce Province, Castro Marina, Mussel Farm, coarse sand. Accogli et al. (2011)

• 39°59′34′′N, 18°25′01′′E; 0 m bsl: [FAO37] Italy, Lecce Province, Castro Marina. Grimaldi de Zio et al. (2003)

• 39°50′40′′N, 18°10′15′′E; 50 m bsl: [FAO37] Italy, Lecce Province, Lido Marini, subtidal. Grimaldi de Zio et al. (1983b)

• 39°50′40′′N, 18°10′25′′E [39°51′N, 18°11′E]; 0 m bsl: [FAO37] Italy, Lecce Province, Lido Marini, intertidal, medium coarse sand. Grimaldi de Zio et al. (1983b)

Mediterranean Sea:

• 39°06′58′′N, 08°15′36′′E; 0 m bsl: [FAO37] Italy, Carbonia-Iglesias Province, San Pietro Island, La Caletta, intertidal, medium sand. Grimaldi de Zio et al. (1983b)

• 37°10′00′′N, 10°11′30′′E; 0 m bsl: [FAO37] Tunisia, Bizerte Governorate, Ghar El Melh Lake, near Ghar El Melh. Grimaldi de Zio et al. (1980a)

• 36°47′30′′N, 03°18′00′′E; 0 m bsl: [FAO37] Algeria, Algiers Province, near Aïn Taya. Grimaldi de Zio et al. (1980a)

• 36°43′00′′N, 02°51′00′′E [36°43′N, 02°50′E]; 0 m bsl: [FAO37] Algeria, Tipaza Province, near Zéralda. Grimaldi de Zio et al. (1980a)

• 35°54′N, 14°32′E; 35 m bsl: [FAO37] Republic of Malta, SE coast of Malta, Sicily Channel, 1.75 km SE of the entrance of Grand Harbour and 1.4 km NW of the coastal settlement Xghajra, fine sand. Grimaldi de Zio & D’Addabbo Gallo (2001)

Pacific Ocean:

• 00°33′S, 90°31′W; 0 m bsl: [FAO87] Galapagos Islands, N of Santa Cruz Island, Playa Borreo [Borrero], damp sand. McKirdy et al. (1976)

• 00°54′S, 89°37′W; 0 m bsl: [FAO87] Galapagos Islands, San Cristóbal Island, Wreck Bay. McKirdy et al. (1976)

Red Sea:

• 27°16′N, 33°49′E; 1.5–3 m bsl: [FAO51] Egypt, Red Sea Governorate, El Hurghada, 4 km S of the Marine Biological Station, clean, coarse coralline sand. Bertolani (1987), Suzuki & Kristensen (2014)

Sargasso Sea:

• 32°22′N, 64°40′W; 4.5 m bsl: [FAO31] Bermuda, Saint David’s Island, very coarse gravel (5–6 mm). Renaud-Mornant (1970b)

• 32°22′N, 64°39′W; 8 m bsl: [FAO31] Bermuda, Saint David’s Island, Saint David’s Head, Castle Harbour, fine coralline sand. Suzuki & Kristensen (2014)

• 32°21′N, 64°40′W; 8 m bsl: [FAO31] Bermuda, Castle Island, Castle Harbour, fine coralline sand. Kristensen & Neuhaus (1999)

• 32°20′N, 64°40′W; 0 m bsl: [FAO31] Bermuda, Castle Island, Castle Harbour, shallow intertidal zone, sediment sample. Coull (1970)

Tyrrhenian Sea:

• 40°20′30′′N, 09°40′15′′E; 0–60 and> 80 m bsl: [FAO37] Italy, Sardinia Island, Nuoro Province, Orosei Gulf, pure sand, coastal organogenic debris, coralligenous debris mud or Posidonia seagrass meadows. D’Addabbo Gallo et al. (1989), Grimaldi de Zio & D’Addabbo Gallo (2001)

• 40°13′32′′N, 09°37′46′′E; 0–7 m bsl: [FAO37] Italy, Sardinia, Nuoro Province, Orosei Gulf, Cala Luna, coarse and medium coarse sand. D’Addabbo Gallo et al. (2005)

• 40°13′N, 09°38′E; 5 and 7 m bsl: [FAO37] Italy, Sardinia, Gulf of Orosei, coarse and medium sand. D’Addabbo Gallo et al. (2007)

• 38°44′N, 16°06′E [38°43′N, 16°06′E]; 0 m bsl: [FAO37] Italy, Vibo Valentia Province, Vibo Valentia, Trainiti beach. Grimaldi de Zio et al. (2003)

• 38°37′N, 15°03′E [38°38′N, 15°04′E]; 1–4 m bsl: [FAO37] Italy, Aeolian Islands, Panarea Island, Baia del Porto, subtidal zone, siliceous sand. Grimaldi de Zio & D’Addabbo Gallo (2001), Grimaldi de Zio et al. (2003)

• 38°38′N, 15°04′E [38°38′N, 15°05′E]; 1–4 m bsl: [FAO37] Italy, Aeolian Islands, Panarea Island, Porto, subtidal zone, siliceous sand. Grimaldi de Zio & D’Addabbo Gallo (2001), Grimaldi de Zio et al. (2003)

• 38°29′N, 14°57′E [38°29′N, 14°58′E]; 1.5–5 m bsl: [FAO37] Italy, Aeolian Islands, Lipari Island, Canneto, subtidal zone, siliceous sand. Grimaldi de Zio & D’Addabbo Gallo (2001), Grimaldi de Zio et al. (2003)

• 38°25′N, 14°56′E [38°25′N, 14°57′E]; 1.5–15 m bsl: [FAO37] Italy, Aeolian Islands, Vulcano Island, West Porto, subtidal zone, siliceous sand. Grimaldi de Zio & D’Addabbo Gallo (2001), Grimaldi de Zio et al.

(2003)

Record numbers (Sea/Ocean classification): Adriatic Sea: 7, Atlantic Ocean: 12, Balearic Sea: 1, Caribbean Sea: 2, Celtic Sea: 1, Coral Sea: 2, Indian Ocean: 1, Ionian Sea: 8, Mediterranean Sea: 5, Pacific Ocean: 2, Red Sea: 1, Sargasso Sea: 4, Tyrrhenian Sea: 8; total: 54.

Record numbers (FAO classification): FAO27: 8, FAO31: 8, FAO37: 29, FAO41: 3, FAO51: 2, FAO71: 2, FAO87: 2; total: 54.

Remarks: This is a widely distributed species in the northern hemisphere with isolated reports in the southern hemisphere. It has been reported from warm tropical waters to cold waters of the temperate zone on the Atlantic coast of Scotland, but it has never been reported from marine caves. It is an intertidal or subtidal species reported from different types of sand and sediments. In our opinion the status of this taxon is similar to H. r. remanei (see remarks for H. r. antillensis and H. r. remanei) and a major revision is needed. For example, McKirdy et al. (1976) reported it as Orzeliscus cf. belopus from the Pacific Ocean on the coast of the Galapagos Islands, and they suggested that the description of this taxon is insufficient and some characters may have been missed by previous authors. The same authors also found other differences between typical specimens and specimens from the Galapagos, but they stated that they were not sufficient to describe a new taxon. Bertolani (1987) and recently Suzuki & Kristensen (2014) reported this species as Orzeliscus cf. belopus (but did not explain why), which suggests that their specimens do not agree fully with the original description or that the original description is not sufficient to do a complete identification. In light of these statements and new findings of cryptic species in marine tardigrades (see e.g. Faurby et al. 2011), we should probably question the present geographic distribution of this species and consider it as a group of species or subspecies.