The Zoogeography of Marine Tardigrada Author Kaczmarek, Łukasz Author Bartels, Paul J. Author Roszkowska, Milena Author Nelson, Diane R. text Zootaxa 2015 4037 1 1 189 journal article 10.11646/zootaxa.4037.1.1 0dd60c86-99b2-43f8-a06e-18d012818103 1175-5326 233519 670819B1-3840-4C0A-ABFF-4D5AE3A263C0 82. Halechiniscus remanei remanei Schulz, 1955 Halechiniscus remanei nov. spec. ( Schulz 1955 ) Halechiniscus remanei Schulz ( Renaud-Debyser 1959b , Zio de 1964, Swedmark & Teissier 1967 , Fleeger 1978 ) Halechiniscus remanei (Zio de 1965, Grimaldi de Zio et al. 1980a, Accogli et al. 2009 ) Halechiniscus remanei Schulz, 1955 ( Rao & Ganapati 1968 , McGinty 1969 , Renaud-Mornant 1970c , 1971 , Renaud-Mornant 1981d , Renaud-Mornant & Gourbault 1981 , 1984, Grimaldi de Zio et al. 1987 , Pollock 1989 , Villora-Moreno & Grimaldi de Zio 1996 , Grimaldi de Zio & D’Addabbo Gallo 2001 , Chang & Rho 2002 , Grimaldi de Zio et al. 2003 , D’Addabbo Gallo et al. 2007) Halechiniscus remanei Schulz 1955 (D’Addabbo Gallo et al. 1978 , Grimaldi de Zio et al. 1979, Grimaldi de Zio et al. 1980b) H. remanei, Schulz 1955 (Grimaldi de Zio et al. 1983b) Terra typica : Tyrrhenian Sea ( Italy , Europe) 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, intertidal, fine-medium sand, mainly calcareous. Grimaldi de Zio et al. (1983b) • 41°57′20′′N , 15°03′23′′E ; 0 m bsl: [FAO37] Italy , Campobasso Province, Campomarino. Grimaldi de Zio et al. (2003) • 41°55′30′′N , 15°89′38′′E [ 41°55′N , 15°10′E ]; 0 m bsl: [FAO37] Italy , Foggia Province, Marina di Chieuti, intertidal, medium sand. Grimaldi de Zio et al. (1983b) • 41°33′35′′N , 15°53′36′′E ; 0 m bsl: [FAO37] Italy , Foggia Province, Frattarolo, intertidal, medium-fine sand. Grimaldi de Zio et al. (1983b) • 41°05′N , 17°01′E ; 0-3 m bsl: [FAO37] Italy , Bari Province, Calo Colombo, coarse sand. Accogli et al. (2009) • 41°02′30′′N , 17°08′00′′E ; 0 m bsl: [FAO37] Italy , Bari Province, Cozze, intertidal, medium sand. Grimaldi de Zio et al. (1983b) • 40°58′45′′N , 17°15′47′′E ; 0 m bsl: [FAO37] Italy , Bari Province, Torre Incina, intertidal, on the shoreline, coarse sand ( 0.841 mm ). Grimaldi de Zio et al. (1980b, 1983b) • 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°54′25′′N , 17°20′40′′E ; 0 m bsl: [FAO37] Italy , Bari Province, Capitolo Sud, intertidal, medium coarse calcareous sand. Grimaldi de Zio et al. (1983b) • 40°53′15′′N , 17°23′30′′E ; 0 m bsl: [FAO37] Italy , Brindisi Province, Torre Egnazia, intertidal, medium calcareous sand. 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, max 6 m from the sea to the river, salinity (<30‰), sand. Zio de (1965) , Grimaldi de Zio et al. (1979) • 40°46′20′′N , 17°39′30′′E ; 0 m bsl: [FAO37] Italy , Brindisi Province, Torre Pozzelle, intertidal, medium sand. Grimaldi de Zio et al. (1983b) • 40°22′54′′N , 18°18′30′′E ; 0 m bsl: [FAO37] Italy , Lecce Province, San Cataldo, beach, intertidal, mediumfine homogeneous sand ( 0.149–0.595 mm ). Zio de (1964) , Grimaldi de Zio et al. (1983b) Alboran Sea: • 35°11′N , 02°26′W ; 0 m bsl: [FAO37] Morocco , Chafarinas Islands, Isable II Island [ Isla Isabel II], shallow sublittoral sediment between rock blocks. Villora-Moreno & Grimaldi de Zio (1996) Atlantic Ocean: • 33°20′N , 79°10′W ; [FAO31] USA , South Carolina State, Georgetown County, North Inlet, subtidal sand. Fleeger (1978) • 33°20′N , 79°10′W ; 0 m bsl: [FAO31] USA , South Carolina State, Georgetown County, North Inlet, intertidal mud between Spartina alterniflora plants. Fleeger (1978) • 44°40′N , 01°14′W ; 0 m bsl: [FAO27] France , Aquitaine Region, Arcachon Bay, American beach, 20–40 cm deep in sand, sand (diameter 0.25–0.30 mm ). Renaud-Debyser (1959b) • 44°40′N , 01°10′W ; 1.5 m bsl: [FAO27] France , Aquitaine Region, Arcachon Bay, d’Eyrac beach, sand. Renaud-Debyser (1959b) • 44°03′N , 01°20′W ; 0 m bsl: [FAO27] Undefined locality, France , Aquitaine Region, coast of Landes Department. Renaud-Debyser (1959b) Balearic Sea: • 41°14′15′′N , 08°59′42′′E [ 41°14′N , 09°10′E ]; 0 m bsl: [FAO37] Italy , Sardinia, Olbia-Tempio Province, Santa Reparata. Grimaldi de Zio et al. (2003) • 41°09′15′′N , 09°31′48′′E [ 41°09′N , 09°07′E ]; 0 m bsl: [FAO37] Italy , Sardinia, Olbia-Tempio Province, Monte Russu, intertidal, coarse sand. Grimaldi de Zio et al. (1983b) • 40°49′N , 08°30′E ; 0 m bsl: [FAO37] Italy , Sardinia, Sassari Province, Gulf of Asinara, medium and medium fine sand. D’Addabbo Gallo et al. (2007) • 40°20′N , 08°22′E ; 0 m bsl: [FAO37] Italy , Sardinia, Sassari or Oristano Province, Gulf of Alghero, medium sand. D’Addabbo Gallo et al. (2007) Caribbean Sea: • 16°12.5′N , 61°29.4′W / 16°13′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 (1981 , 1984) Celtic Sea: • 48°01′N , 05°44′W ; 130 m bsl: [FAO27] France , Brittany Region, continental shelf, near Brest, coarse sand and shelly detritus with limestone and finer siliceous sand. Renaud-Mornant (1970c , 1971 ) Coral Sea: • 19°2′S , 147°30′E [ 19°15′S , 146°49′E ]; 0 m bsl: [FAO71] Australia , Queensland State, Townsville, beach, intertidal coralline sand. Renaud-Mornant (1981d) East China Sea: • 33°30′N , 126°58′E ; 2–3 m bsl: [FAO61] South Korea , Jeju Province, Jejudo Island , Udo Islet, sublittoral, coralline sand. Chang & Rho (2002) Indian Ocean: • 17°43′N , 83°20′E ; 0 m bsl: [FAO57] India , Andhra Pradesh, Bay of Bengal, Waltair coast, beach, mid-water level, medium sand. Rao & Ganapati (1968) • 20°11′S , 57°16′E [ 20°11′S , 57°46′E ]; 0 m bsl: [FAO51] Republic of Mauritius , Flacq Region, Belle Mar Beach, intertidal, interstitial, fine well-sorted coralline sand with Foraminifera. Grimaldi de Zio et al. (1987) Ionian Sea: • 40°29′00′′N , 17°03′25′′E ; 15 m bsl: [FAO37] Italy , Tarento Province, Secca dell’Armeleia, subtidal, organogenic, Amphioxus debris. Grimaldi de Zio et al. (1983b) • 40°24′30′′N , 17°13′28′′E ; 0 m bsl: [FAO37] Italy , Tarento Province, Lido Bruno, intertidal, medium coarse sand with high calcareous percentage (97%). Grimaldi de Zio et al. (1983b) • 40°23′07′′N , 17°18′29′′E ; 0 m bsl: [FAO37] Italy , Tarento Province, Lido Gandoli, intertidal, heterogenous sand with high calcareous content. Grimaldi de Zio et al. (1983b) • 40°22′03′′N , 17°22′29′′E [ 40°21′N , 17°22′E ]; 0 m bsl: [FAO37] Italy , Tarento Province, Gabbiano, intertidal, heterogenous sand mainly calcareous (98%). Grimaldi de Zio et al. (1983b) • 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°21′N , 17°23′E ; 0 m bsl: [FAO37] Italy , Taranto Province, Marina di Pulsano, Lido conche, beach, 2.5 m of the coast line, 15–20 cm deep in sand, medium coarse sand (max 0.841 mm ). D’Addabbo Gallo et al. (1978), Grimaldi de Zio et al. (1980b) • 40°17′45′′N , 17°33′38′′E ; 0 m bsl: [FAO37] Italy , Tarento Province, Campo Marino. Grimaldi de Zio et al. (2003) • 40°09′N , 17°58′E ; 0-20 m bsl: [FAO37] Italy , Lecce Province, Nard, Corvine cave, coarse sand. Accogli et al. (2009) • 40°08′38′′N , 17°59′00′′E ; 0 m bsl: [FAO37] Italy , Lecce Province, Santa Caterina, intertidal, medium sand, mainly calcareous. Grimaldi de Zio et al. (1983b) • 40°08′N , 17°59′E ; 0 m bsl: [FAO37] Italy , Lecce Province, Santa Caterina, beach, 1 m from the shoreline, 35 cm deep in sand, medium sand (max 0.354 mm ). D’Addabbo Gallo et al. (1978), Grimaldi de Zio et al. (1980b) • 40°03′00′′N , 18°00′00′′E ; 0 m bsl: [FAO37] Italy , Lecce Province, near Gallipoli, Lido Conchiglie, intertidal, medium sand, mainly calcareous (97%). Grimaldi de Zio et al. (1983b) • 40°02′50′′N , 17°59′30′′E ; 0 m bsl: [FAO37] Italy , Lecce Province, near Gallipoli, Baia Verde, intertidal, medium sand, mainly calcareous. Grimaldi de Zio et al. (1983b) • 40°02′N , 18°01′E ; 0 m bsl: [FAO37] Italy , Lecce Province, near Gallipoli, Baia Verde, beach, 1 m of the coast line, 50 deep cm deep in sand, sand ( 0.354 mm ). D’Addabbo Gallo et al. (1978) • 39°51′05′′N , 18°12′20′′E [ 39°50′N , 18°12′E ]; 0 m bsl: [FAO37] Italy , Lecce Province, Torre Pali, intertidal, medium-fine sand, mainly calcareous. 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) • 39°50′40′′N , 18°10′15′′E ; 50 m bsl: [FAO37] Italy , Lecce Province, Lido Marini, subtidal. Grimaldi de Zio et al. (1983b) Mediterranean Sea: • 36°45′00′′N , 02°53′00′′E [ 36°46′N , 02°53′E ]; 0 m bsl: [FAO37] Algeria , Tipaza Province, near Staouéli. Grimaldi de Zio et al. (1980a) • 36°38′00′′N , 02°40′00′′E ; 0 m bsl: [FAO37] Algeria , Tipaza Province, near Khemisti. Grimaldi de Zio et al. (1980a) • 35°54′N , 14°32′E ; 41 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, coarse organogenic detritus. Grimaldi de Zio & D’Addabbo Gallo (2001) North Sea: • 59°17′N , 11°21′E ; 0 m bsl: [FAO27] Sweden , Västra Götalands Iän Region, Kattegat, Gunnarskär, shelly sand with shells, fragments of sea urchin tests and pieces of bryozoans. Swedmark & Teissier (1967) • 58°35′N , 11°04′E ; 0 m bsl: [FAO27] Sweden , Västra Götalands Iän Region, Kattegat, Väderöarna, shelly sand with shells, fragments of sea urchin tests and pieces of bryozoans. Swedmark & Teissier (1967) • 58°17′N , 11°30′E : [FAO27] Undefined locality, Sweden , Västra Götaland County, Bohuslän Province, Skagerrak, Gullmarsfjord. Renaud-Mornant (1971) • 58°06′N , 11°20′E ; 0 m bsl: [FAO27] Sweden , Västra Götalands Iän Region, Kattegat, Bonden Island near Käringön, shelly sand with shells, fragments of sea urchin tests and pieces of bryozoans. Swedmark & Teissier (1967) Pacific Ocean: • 38°17.0′N , 123°2.0′W [ 38°18′N , 123°03′W ]; 0 m bsl: [FAO77] USA , California State, Sonoma County, N of the parking area at Bodega Head, 0–60 cm deep in sand, well-sorted coarse or fine sand (mean phi grains = 2.545). Pollock (1989) • 36°38′N , 121°56′W ; 0–20 cm / 0–0.2 m bsl: [FAO77] USA , California State, Monterey County, Pacific Grove, SW of Hopkins Marine Station, sandy beach, mid- to high tide, medium grain sand. McGinty (1969) • 35°59.0′N , 121°30.0′W / 35°59′N , 121°30′W ; 0 m bsl: [FAO77] USA , California State, Monterey County, Los Padres National Forest, Mill Creek Picnic Area, Big Sur coastline, 0–60 cm deep in sand, sand. Pollock (1989) • 33°35.0′N , 117°53.0′W / 33°35′N , 117°53′W ; 0 m bsl: [FAO77] USA , California State, Orange County, Corona del Mar, Corona del Mar State Park, Little Corona Beach, 0–60 cm deep in sand, sand with shell debris (mean phi grains = 0.655). Pollock (1989) Tyrrhenian Sea: • 40°50′N , 14°16′E ; 0 m bsl: [FAO37] Type Locality: Italy , Province of Naples, Naples, Gulf of Naples, coarse sand. Schulz (1955) • 40°36′N , 14°52′E ; 0 m bsl: [FAO37] Italy , Salerno Province, Battipaglia. Grimaldi de Zio et al. (2003) • 40°13′N , 09°38′E ; 0–7 m bsl: [FAO37] Italy , Sardinia, Nuoro Province, Gulf of Orosei, coarse and medium fine 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°27′N , 14°55′E [ 38°27′N , 14°56′E ]; 1.5–5 m bsl: [FAO37] Italy , Aeolian Islands, Lipari Island , Valle Muria, subtidal zone, siliceous sand. Grimaldi de Zio & D’Addabbo Gallo (2001) , Grimaldi de Zio et al. (2003) • 38°26′N , 14°55′E [ 38°26′N , 14°57′E ]; 1.5–5 m bsl: [FAO37] Italy , Aeolian Islands, Lipari Island , Pietra Lunga South, subtidal zone, siliceous sand. Grimaldi de Zio & D’Addabbo Gallo (2001) , Grimaldi de Zio et al. (2003) • 38°01′40′′N , 13°56′38′′E [ 38°02′N , 14°01′E ]; 0 m bsl: [FAO37] Italy , Sicily, Palermo Province, Cafal (La Plaia), intertidal, coarse sand, mainly siliceous (93%). Grimaldi de Zio et al. (1983b) • 37°34′50′′N , 12°50′38′′E [ 37°35′N , 12°50′E ]; 0 m bsl: [FAO37] Italy , Sicily, Trapani Province, Selinunte, intertidal, medium sand, mainly calcareous. Grimaldi de Zio et al. (1983b) Record numbers (Sea/Ocean classification): Adriatic Sea: 14, Alboran Sea: 1, Atlantic Ocean: 5, Balearic Sea: 4, Caribbean Sea: 1, Celtic Sea: 1, Coral Sea: 1, East China Sea: 1, Indian Ocean: 2, Ionian Sea: 16, Mediterranean Sea: 3, North Sea: 4, Pacific Ocean: 4, Tyrrhenian Sea: 9; total: 66. Record numbers (FAO classification): FAO27: 8, FAO31: 3, FAO37: 47, FAO51: 1, FAO57: 1, FAO61: 1, FAO71: 1, FAO77: 4; total: 66. Remarks: This is the most widely distributed member of the genus Halechiniscus , but known almost exclusively from the northern hemisphere. Halechiniscus r. remanei was reported from warm waters of temperate and tropical zones (northern parts of tropical zone and southern parts of temperate zone). However, it was also reported from cold waters of the North Sea on the coast of Sweden . Halechiniscus r. remanei is known mainly from intertidal and shallow subtidal zones, although it was also found deeper (to 130 m bsl.). It has been reported from 0–40 cm deep in different types of sand. Although it is a rather common species, it has never been found in marine caves. Based on Renaud-Mornant (1984b) , it is cosmopolitan but in the process of speciation (the changes are mainly visible in modifications of the caudal appendages). Speciation is less visible in the Australian region and more advanced in the Caribbean region where a new subspecies H. r. antillensis was described (see remarks for Halechiniscus r. antillensis above). Also specimens from California clearly differ from typical forms ( McGinty 1969 , Renaud-Mornant 1984b ) and should probably be considered a new subspecies or even a new species. In the light of these statements and new findings of cryptic species in marine tardigrades (see e.g. Faurby et al. 2011 ), we question the cosmopolitan character of this species and consider it as a group of species or subspecies. Thus, the geographical range of H. r. remanei and its ecological requirements need thorough revision. It also should be noted that Chang & Rho (2002) in their relatively recent treatment of Halechiniscus do not even mention the two subspecies of H. remanei .