Actinarctus doryphorus doryphorus Schulz, 1935

Kaczmarek, Łukasz, Bartels, Paul J., Roszkowska, Milena & Nelson, Diane R., 2015, The Zoogeography of Marine Tardigrada, Zootaxa 4037 (1), pp. 1-189 : 117-118

124. Actinarctus doryphorus doryphorus Schulz, 1935

Actinarctus doryphorus nov. gen. nov. spec. ( Schulz 1935)

Actinarctus doryphorus E. Schulz ( Grell 1937)

Actinarctus doryphorus ( Grell 1939)

Actinarctus doryphorus Schulz ( Swedmark 1956, Swedmark & Teissier 1967) Actinarctus doryphorus Schulz, 1935 (Eibye-Jacobsen 1996, Villora-Moreno & Grimaldi de Zio 1996, D’Addabbo Gallo et al. 1999a, b, Kristensen & Neuhaus 1999, Grimaldi de Zio 2003, D’Addabbo Gallo et al. 2007, Accogli et al. 2011)

Terra typica : North Sea ( Germany, Europe)

Adriatic Sea:

• 40 ° 27.24′N, 18°14.63′E [40°22′N, 18°20′E]; 10 m bsl: [FAO37] Italy, Lecce Province, Ceisne (Marne protected area), coarse sand. Accogli et al. (2011)

• 40 ° 27.17′N, 18°14.58′E [40°22′N, 18°20′E]; 10 m bsl: [FAO37] Italy, Lecce Province, Ceisne (Marne protected area), coarse sand. Accogli et al. (2011)

Alboran Sea:

• 35°11′N, 02°27′W; 25 m bsl: [FAO37] Morocco, Chafarinas Islands, Congreso Island [ Isla del Congres], very coarse sand and gravels with biodendritic sediments. Villora-Moreno & Grimaldi de Zio (1996)

• 35°11′N, 02°26′W; 0 m bsl: [FAO37] Morocco, Chafarinas Islands, Congreso Island [ Isla del Congres], sublittoral mixed environment, Amphioxus gravel. Villora-Moreno & Grimaldi de Zio (1996)

Atlantic Ocean:

• 4°20′S to 4°50′S, 36°00′W to 37°15′W [04°35′S, 36°38′W]; 20 m bsl: [FAO41] Brazil, continental shelf off Ceará State and Rio Grande Do Norte State, medium sand. da Rocha et al. (2013), FAO: 41

• 8°25′10′′S to 8°25′20′′S, 34°55′00′′W [08°25′S, 34°58′W]; 3 m bsl: [FAO41] Brazil, Pernambuco State, Cupe beach, biogenic gravel. da Rocha et al. (2013)

Balearic Sea:

• 40°20′N, 08°22′E; 20 m bsl: [FAO37] Italy, Sardinia, Sassari or Oristano Province, Gulf of Alghero, medium sand. D’Addabbo Gallo et al. (2007)

Celtic Sea:

• 48°46′N, 04°05′W; 0 m bsl: [FAO27] France, Brittany Region, Roscoff, Trezen ar Skoden shell dune, 2.5 miles [4 km] WNW of Ile de Batz, shelly sand with mollusc shells, plates of barnacles, skeletons of bryozoans, and Ophiuroideans, with no organic debris. Swedmark & Teissier (1967)

• 48°44′N, 03°59′W; 0 m bsl: [FAO27] France, Brittany Region, Bloscon, Amphioxus sand. Swedmark (1956)

Ionian Sea:

• 39 ° 55.55′N, 18°02.81′E; 20 m bsl: [FAO37] Italy, Lecce Province, Torre San Giovanni Mancaversa, fine sand. Accogli et al. (2011)

• 39°55′55′′N, 18°02′71′′E [39°54′N, 18°06′E]; 0 m bsl: [FAO37] Italy, Lecce Province, San Giovanni [Torre San Giovanni]. Grimaldi de Zio et al. (2003)

• 39 ° 58.71′N, 18°18.28′E [39°49′N, 18°18′E]; 10 m bsl: [FAO37] Italy, Lecce Province, San Gregorio, coarse sand. Accogli et al. (2011)

• 39°48′40′′N, 18°40′38′′E [39°49′N, 18°19′E]; 0 m bsl: [FAO37] Italy, Lecce Province, San Gregorio. Grimaldi de Zio et al. (2003)

Ligurian Sea:

• 43°32 ′ N, 10°12′E; 7 m bsl: [FAO37] Italy, Livorno Province, 300 m NW of Meloria Shoals Lighthouse, pit bottom, salinity 38 PSU, coarse coralligenous detritus (granulometric max. = 1.4 mm). D’Addabbo Gallo et al. (1999a, b)

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°54′51.78′′N, 10°48′55.38′′E – 58°54′59.04′′N, 10°48′51.90′′E; 0 m bsl: [FAO27] Sweden, Västra Götaland County, Tjärnö area, Grisbådarna, sublittoral samples from dredge, carbonate sand. Willems et al. (2009)

• 58°52′34.02′′N, 11°06′40.56′′E – 58°52′33.96′′N, 11°06′45.06′′E; 0 m bsl: [FAO27] Sweden, Västra Götaland County, Tjärnö area, Inre Vattenholmen, sublittoral samples from dredge, carbonate sand. Willems et al. (2009)

• 58°49′02.16′′N, 11°04′24.24′′E – 58°49′25.62′′N, 11°04′28.62′′E; 0 m bsl: [FAO27] Sweden, Västra Götaland County, Tjärnö area, Ullvillarna, sublittoral samples from dredge, shell gravel or carbonate sand ( Amphioxus sand). Willems et al. (2009)

• 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°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)

• 57°30.2′N, 10°39.4′W / 57°30′N, 10°39′E; 25 m bsl: [FAO27] Denmark, Nordjylland Region, Kattegat of Fredrikshavn, NE of Hirsholmene, Amphioxus sand (sand with shelly gravel). Jørgensen et al. (1999)

• 57°28′N, 10°42′E [57°28′N, 10°42′E]; 25 m bsl: [FAO27] Denmark, Nordjylland Region, Kattegat, Fredrikshavn, NE of Hirsholmene, Amphioxus sand. Kristensen & Neuhaus (1999)

• 57°26′N, 10°32′E; 0 m bsl: [FAO27] Denmark, Nordjylland Region, Fredrikshavn, Kattegat, Amphioxus sand. Eibye-Jacobsen (1996)

• 54°11′N, 07°53′E; 0 m bsl: [FAO27] Type Locality: Germany, Schleswig-Holstein State, Helgoland Island, in Amphioxus sand on echinoderm ( Echinocyamus pusillus ) and in “Schell [shell]. Schulz (1935)

• 54°11′N, 07°53′E; 0 m bsl: [FAO27] Germany, Schleswig-Holstein State, Helgoland Island, Skitt-Gatts, dune, coarse biogenic detritus ( Polygordius Bruchschill ). Grell (1937, 1939)

Record numbers (Sea/Ocean classification): Adriatic Sea: 2, Alboran Sea: 2, Atlantic Ocean: 2, Balearic Sea: 1, Celtic Sea: 2, Ionian Sea: 4, Ligurian Sea: 1, North Sea: 11; total: 25.

Record numbers (FAO classification): FAO27: 13, FAO37: 10, FAO41: 2; total: 25.

Remarks: This species is reported mainly from the temperate zone of the northern hemisphere; however, recently it has also been found in the tropical region in the southern hemisphere, but these records probably need confirmation. It is mainly an intertidal species, but it has also been found in the shallow subtidal zone often in Amphioxus sand. D’Addabbo Gallo et al. (1999b) redescribed this species in detail based on a large population (adults and larvae) from Livorno Province. These authors showed a large variability of some morphological features of A. d. doryphorus and suggested that the subspecies A. d. ocellatus should be considered as nonvalid. However, they did not state this definitively and we feel that this case needs further research and a reexamination of specimens from both type localities.