Angursa bicuspis n. g., n. sp. (Pollock 1979)
Angursa bicuspis abyssalis n. ssp. (Renaud-Mornant 1981 b)
Angursa bicuspis Pollock, 1979 (D’Addabbo Gallo et al. 1987, Grimaldi de Zio & D’Addabbo Gallo 1987) Angursa bicuspis abyssalis (Sandulli et al. 1999, Romano III et al. 2011)
Angursa bicuspis abyssalis Renaud-Mornant 1981 (D’Addabbo Gallo et al. 2001, Grimaldi de Zio & D’Addabbo Gallo 2001, Grimaldi de Zio et al. 2003)
Angursa bicuspis Renaud-Mornant, 1981 (Accogli et al. 2011)
Terra typica: Atlantic Ocean (USA, North America)
Adriatic Sea:
• 42 °06′N, 15 ° 29 ′E; 10 m bsl: [FAO 37] Italy, Foggia Province, Tremiti Islands, southern area of San Domino Island, in the proximity of Punta Zio Cesare, submarine cave Grotta delle Viole, external cave’s entrance, coarse organogenic detritus. D’Addabbo Gallo et al. (2001), Grimaldi de Zio & D’Addabbo Gallo (2001)
• 42 °06 ′ N, 15 ° 29 ′E; 7 m bsl: [FAO 37] Italy, Foggia Province, Tremiti Islands, southern area of San Domino Island, in the proximity of Punta Zio Cesare, submarine cave Grotta delle Viole, ca. 20 m from the cave’s entrance, coarse organogenic detritus. D’Addabbo Gallo et al. (2001), Grimaldi de Zio & D’Addabbo Gallo (2001)
• 42 °06′N, 15 ° 29 ′E; 5–10 m bsl: [FAO 37] Italy, Foggia Province, Tremiti Islands, San Domino Island, Punta Zio Cesare, cave Grotta delle Viole, 0–10 cm depth, 80 % coarse organogenic sand, 3 % silt and clay, (mean diameter = 0.125 mm). Sandulli et al. (1999), Grimaldi de Zio & D’Addabbo Gallo (2001)
• 40 ° 31.59 ′N, 18 °05. 51 ′E; 20 m bsl: [FAO 37] Italy, Brindisi Province, Lendinuso, coarse sand in Posidonia meadows. Accogli et al. (2011)
• 40 ° 19.36 ′N, 18 ° 23.16 ′E ; 50 m bsl: [FAO 37] Italy, Lecce Province, Torre Specchia, mud. Accogli et al. (2011)
Atlantic Ocean:
• 47 ° 33.8 ′N, 08° 38.6 ′W; 2205 m bsl: [FAO 27] W Atlantic, ca. 300 km from the coast of France, abyssal. Renaud-Mornant (1981 b)
• 41 ° 31 ′ 41 ′′N, 70 ° 40 ′ 41 ′′W; 0 m bsl: [FAO 21] Type Locality: USA, Massachusetts State, Barnstable County, Woods Hole, Buzzard’s Bay side of Penzance Point, Crane’s Beach, 38 cm above mean low water level, 27 cm deep in sand, medium coarse sand. Pollock (1979)
• 34 ° 07.3′N, 75 ° 57.7 ′W / 34 °07′N, 75 ° 58 ′W; 400 m bsl: [FAO 31] USA, North Carolina State, ca. 100 km off the coast of North Carolina, fine sand with silt-clay. Pollock (1979)
• 27 °07′N, 92 ° 20 ′W; 1401 and 2020 m bsl: [FAO 31] Undefined locality, USA, Gulf of Mexico. Romano III et al. (2011)
• 12 °03′ 3 S, 12 ° 20 ′ 5 E [12 °03′03′′S, 12 ° 20 ′05′′E]; 2063 m bsl: [FAO 47] SE Atlantic, ca. 150 km from the coast of Angola, abyssal. Renaud-Mornant (1981 b)
Ionian Sea:
• 40 ° 0 3.92 ′N, 17 ° 59.64 ′E ; 50 m bsl: [FAO 37] Italy, Lecce Province, Gallipoli, mud. Accogli et al. (2011)
• 39 ° 59.38 ′N, 18 ° 25.01 ′E ; 50 m bsl: [FAO 37] Italy, Lecce Province, Castro Marina, Mussel Farm, mud. Accogli et al. (2011)
• 39 ° 52 ′00′′N, 16 ° 42 ′ 30 ′′E; 20–200 m bsl: [FAO 37] Italy, Cosenza Province, High Ionian Sea offshore, Calabrian Coast, Amendolara Shoal, coralligenous sand, coralligenous sand mixed with debris, coarse medium or fine coralligenous sand (few samples). D’Addabbo Gallo et al. (1987), Grimaldi de Zio & D’Addabbo Gallo (1987), Grimaldi de Zio et al. (2003)
Mediterranean Sea:
• 43 °03.30′N, 05° 15.10 ′E / 43 °03′N, 05° 15 ′E; 460 m bsl: [FAO 37] France, Provence-Alpes-Côte d'Azur Region, Cassis Canyon, bathyal. Renaud-Mornant (1981 b)
Tyrrhenian Sea:
• 40 ° 20 ′ 30 ′′N, 09° 40 ′ 15 ′′E; 41–60 m bsl: [FAO 37] Italy, Sardinia Island, Nuoro Province, Orosei Gulf, sand in Posidonia seagrass meadow and coralligenous detritus. Grimaldi de Zio & D’Addabbo Gallo (2001), Grimaldi de Zio et al. (2003)
• 38 ° 36 ′N, 14 ° 55 ′E; 40 m bsl: [FAO 37] Italy, Aeolian Islands, Secca del Capo, coralligenous bank, subtidal zone, coarse coralligenous detritus. Grimaldi de Zio & D’Addabbo Gallo (2001)
• 38 ° 24 ′N, 14 ° 59 ′E; 1.5–15 m bsl: [FAO 37] Italy, Aeolian Islands, Vulcano Island, Punta Lucia, subtidal zone, siliceous sand. Grimaldi de Zio & D’Addabbo Gallo (2001), Grimaldi de Zio et al. (2003)
Record numbers (Sea/Ocean classification): Adriatic Sea: 5, Atlantic Ocean: 5, Ionian Sea: 13, Mediterranean Sea: 1, Tyrrhenian Sea: 3; total: 17.
Record numbers (FAO classification): FAO 21: 1, FAO 27: 1, FAO 31: 2, FAO 37: 12, FAO 47: 1; total: 17.
Remarks: Pollock (1979) described this taxon (which is also the type species for the genus Angursa) based on two specimens, one from the intertidal zone and one from the deep sea. He described some differences between these specimens, but he decided they were not sufficient to assign these specimens to two different taxa (some differences were very small and some structures were not visible in the deep sea specimen probably due to preservation). Pollock (1979) pointed out that such a wide range of depths for tardigrade species is unusual, as is the difference in substrates where the two specimens were found. Based on modern taxonomy we should probably consider specimens described by Pollock (1979) as belonging to two different taxa, but reexamination of the type material is necessary to confirm this statement. Type specimens are available at the Smithsonian Institute’s National Museum of Natural History. Renaud-Mornant (1981 b) described the subspecies Angursa bicuspis abyssalis based on some minor differences, however Villora-Moreno (1998) stated that this description was a redescription of Pollock’s species. He also stated that Pollock’s description was not sufficient to separate nominal species from other Angursa species, but the new characters described by Renaud-Mornant (1981 b) for A. bicuspis were very valuable and adequate for such separation (Villora-Moreno 1998). We think the situation is still unclear because this species has a very peculiar distribution from the intertidal zone to the abyss (2200 m bsl). Temporarily, we decided to accept Villora-Moreno’s (1998) point of view but we think that the issue needs more detailed study. This species has also been found in submarine caves. It has been reported from different substrates mainly in southern parts of the northern hemisphere, however it is also known from one locality in the southern hemisphere.