Mesothuria gargantua Deichmann, 1930

Gebruk, Andrey V., Solis-Marin, Francisco A., Billett, David S. M., Rogacheva, Antonina V. & Tyler, Paul A., 2012, Review of the genus Zygothuria Perrier, 1898 and the Atlantic group of species of the genus Mesothuria Ludwig, 1894 (Synallactidae: Holothuroidea) with description of the new species Mesothuria milleri sp. nov., Journal of Natural History 46, pp. 265-348 : 289-300

Mesothuria gargantua Deichmann, 1930

( Figure 8 )

Mesothuria gargantua – Deichmann, 1930: 95–96, pl. 7, fig. 1; 1940: 191; 1954: 386.

Mesothuria (Allantis) gargantua – Heding, 1940: 119.

Holothuria verrilli – Théel, 1886b: 6 (partim).

Material examined

See Table 7.

Type material

Holotype, MCZ 449 .

Type locality

Caribbean Sea , off Barbados, 720 m.

Diagnosis

After Deichmann (1954): large robust form, with thick skin. Stout, cylindrical feet over entire surface, except anterior part of ventrum; dorsal feet slightly smaller.

Ossicles very large and robust quadriradiate (rarely pentaradiate) tables; disk irregular ( Figure 8A–C ), diameter up to 0.25 mm; spire with numerous robust spines ( Figure 8D,E ). Feet with vestige of end plate or none; tables almost of same size as in skin.

Remarks

Mesothuria gargantua is characterized by very large and robust quadriradiate (rarely pentaradiate) tables (disk diameter up to 0.25 mm). It was described carefully by Deichmann, but a single illustration of ossicle she provided was insufficient. More clear illustrations, based on the preparations from the type material ( MCZ, cat. no. 449) are presented here ( Figure 8 ). A photograph of M. gargantua in situ is given in Rice and Miller (1991).

See also remarks for Mesothuria verrilli .

Distribution

Off Lesser Antilles (Blake); north of Cuba ( Atlantis ) ( Figure 6 ). This species may possibly be common in deep water of coasts of Cuba and Yucatan ( Deichmann 1954).

Bathymetric range

Depth 720–1343 m.

Mesothuria intestinalis ( Ascanius, 1805)

( Figures 1 , 9C–D )

Holothuria intestinalis – Ascanius, 1805: 5, pl. 45; Marenzeller, 1893a: 15; 1895: 21; Ludwig, 1893a: 174; Théel, 1886a: 209; Bell, 1892: 48–49, pl. 5, fig. 3; Koehler, 1895: 482; 1896: 106–108; Hérouard, 1896: 163.

Mesothuria intestinalis (Ascanius) – Östergren, 1896: 347–351, pl. 18, figs. 1–26; 1902: 6–7; Perrier, 1902: 304–307, text figs. 1–2, pl. 16, figs. 19–21; Ludwig, 1901: 139; Théel, 1901: 1–38, pls. 1–2, figs. 1–19, 12 textfigs.; Hérouard, 1923: 10, pl. 5, fig. 5–6; Mortensen, 1927: 381, text fig. 225, 228: 3; Koehler, 1927: 240, pl. 15, fig. 3 (partim); Deichmann, 1930: 94–95, pl. 6, figs. 9–10; 1954: 385–386; Sibuet, 1974: 795; Harvey et al., 1988: 184; Madsen & Hansen, 1994: 76–79, figs. 46–47, map 29; Massin, 1996: 43.

Mesothuria (Allantis) intestinalis (Ascanius) – Heding, 1942a: 7, text fig. 6, figs. 1–7; Panning, 1952: 123–125, figs. 1–3.

Allantis intestinalis var. verrilli – Hérouard, 1902: 18–21, pl. 1, figs. 3–6 (partim).

Fistularia mollis – M. Sars, 1835: 40.

Thyonidium scabrum – M. Sars, 1868: 19–20.

Holothuria verrilli Théel, 1886 – Marenzeller, 1893b: 7–9, pl. 1, fig. 2, pl. 2, fig. 2.

Mesothuria triradiata – Heding, 1942b: 217–218, textfig. 1, figs. 1–6.

Mesothuria verrilli (Théel, 1886) – Tortonese, 1952: 228; 1961, pl. 1, text fig. 1.

Material examined

See Table 8.

Type material

Does not exist.

Type locality

Coast of Norway ( Deichmann 1954).

Diagnosis

After Madsen and Hansen (1994): body nearly cylindrical, only slightly flattened ventrally and slightly attenuated towards both ends. Body wall thin, soft, often becoming strongly wrinkled in preserved specimens. Mouth subventral, anus terminal. Tentacles 20 in adult, fewer in younger specimens, with slightly incised disks, completely retractile with rim of bodywall closed over them. Tubefeet small, especially scarce along mid-ventral radius, which appears almost naked. Tubefeet most strongly developed in ventrolateral ambulacra (two to four bands), but no distinct longitudinal series of large tubefeet. On dorsal side tubefeet fewer and smaller, very sparse. All tubefeet, including smallest ones, usually have sucking disk. One or two Polian vesicles. Sequential hermaphrodite gonads consist of many long tubules, arranged loosely inside body cavity

Ossicles tables of quadriradiate type, with spire of four rods, ending in crown of several thorns ( Figure 9D ); disk with typically eight regularly shaped small holes surrounding one circular central hole (0.7– 0.14 mm in diameter) ( Figure 9C ), sometimes with smaller peripheral holes, but many variations occur. In small specimens spire may be composed of only three rods. Tables of tubefeet well developed, with spire as in tables of bodywall, but slightly smaller, and with terminal plate, 0.15–0.30 mm in diameter. Ventral tables sometimes slightly smaller and more robust than dorsal ones. Spire of table 0.06–0.10 mm in height, consisting of four (very rarely three or five) vertical pillars rising from highly elevated primary cross and united by crossbeams distally where spire ends in low, spinous crown. In tentacles ossicles rod-shaped.

Museum catalogue No. of TL / W (mm) Locality number specimens

Date Depth (m)

UNAM-ICML 2 330 / 60 – – 5.116.0

USNM 8574 1 142 / 27 Norwegian Sea, Norway, Moldefjorden. – ZMUC Eh 319-747 1 – Ingolf, St. 85, 63 ◦ 21 ′ N, 25 ◦ 21 ′ W – ZMUC Eh 319-748 3 – – – ZMUC Eh 319-749 1 – – – ZMUC Eh 319-750 1 – – – ZMUC Eh 319-751 2 – Norway – ZMUC Eh 319-749 1 – – – ZMUC Eh 319-753 1 – Minch Straight, Scotland, NE Atlantic – ZMUC Eh 319-754 1 – – 31-07-1911 ZMUC Eh 319-755 2 – Gullmar Fjord, Skagerrak 14-01-1910 ZMUC Eh 319-756 8 – – 02-1918 ZMUC Eh 319-757 4 – – 07-1896 ZMUC Eh 319-758 3 – – – ZMUC Eh 319-759 2 – Gullmar Fjord, Skagerrak 15-01-1910 ZMUC Eh 319-760 1 – Thor, St. 134 15-07-1910 ZMUC Eh 319-761 5 – Gullmar Fjord, Skagerrak 07-1931 ZMUC Eh 319-762 2 – Wahrberg Eliasson St. 33, Koster Fjord, Boheslen 04-08-1925 ZMUC Eh 319-763 12 – – 08-1923 NHM 92.2.11.31 / 35 6 – – 17-09-1891 NHM 98.5.3.342 1 – Norway – NHM 88.6.21.1 / 2 2 – Kilbrennan Sound, NE Atlantic 03-03-1888 NHM 98.5.3.342 1 Norway – NHM 97.4.4.13 1 Bay of Biscay –

–

– 320 – – – – – – 200 100 400 – – – – – 600

200–300

72–109

72–109 91 – –

NHM 66.11.23.2 1 Minch Straight, off Hebrides, NE Atlantic –

NHM 88.4.15.27-30 5 – – 17-03-1888

NHM 98.5.3.306-8 21 – Norman Expedition St. 30, Norway 1879

NHM 2 – Norway –

NHM 95.3.3.303-5 5 – – –

NHM 4 – – –

98.5.3.309-311

NHM 2 – – –

1900.4.1.21 / 22

ZMUC 1 130 / 29 – –

ZMUC 1 85 / 20 – –

ZMUC HOL 221 ∗ 1 – Valdivia St. 33, NE Atlantic, off Morocco 25-08-1898 24 ◦ 35 ′ N, 17 ◦ 04 ′ W

ZMUC 1 – Atlantide, St. 120, 02 ◦ 09 ′ N, 09 ◦ 27 ′ E, off Africa 1946

MNHN 1103 1 – – 1963

MNHN 1260 5 – – 1962

MNHN 1261 1 – – 1963

MNHN 1262 1 – – 1951

MNHN 1263 2 – – 1963

MNHN 1349 2 – – 14-08-1962

MNHN 1350 2 – – –

MNHN 1723 6 – – 24-08-1962

MNHN 1755 1 – Talisman, St. 48, off Canary Islands, 29 ◦ 1 ′ N, 26-06-1883 14 ◦ 51 ′ W

MNHN 1756 2 – Travailleur St. 38, 34 ◦ 13 ′ 30 ′′ N, 10 ◦ 3 ′ W 30-07-1882

MNHN 1757 2 – Talisman, St. 49, coast of Morocco, 28 ◦ 37 ′ N, 27-06-1883 15 ◦ 22 ′ W

MNHN 1811 1 – Princesse de Monaco St. 1275 1910

MNHN 39 1 – Prince Napoleon 1856, Bergen Museum –

MNHN 40 1 – Bergen Museum, id.?Loven 1852 –

MNHN 1776 1 – Talisman St. 17, off Morocco, 33 ◦ 33 ′ N, 11 ◦ 19 ′ W 14-06-1883

– 54–69

–

–

–

–

–

– 2480

650–260 350

–

– 280–400 –

450

150

–

1180

636 865

200–300 – – 550

(Continued)

Museum catalogue No. of TL / W (mm) Locality number specimens

Date Depth (m)

MNHN 2544 1 – Thalassa St. U 863, 47 ◦ 11.4 ′ N, 5 ◦ 39 ′ W – MNHN 3482 1 – Bauguls 23-08-1968 MNHN 7401 3 – Europe, Depro 96, St. 3, Golf du Lion, 48 ◦ 43.5 ′ N, 19-04-1996 4 ◦ 29.4 ′ E

MNHN 8058 1 – Europe, Depro 96, St. 4, 42 ◦ 34.6 ′ N, 4 ◦ 07.4 ′ E 19-04-1996 MNHN 8059 5 – Europe, Depro 96, St. 6, 42 ◦ 20.7 ′ N, 3 ◦ 58.9 ′ E 20-04-1996 MNHN 8060 1 – Europe, Depro 96, St. 8, 42 ◦ 23.5 ′ N, 3 ◦ 43.5 ′ E 21-04-1996 MNHN 8061 2 – Europe, Depro 96, St. 14, 42 ◦ 15.6 ′ N, 3 ◦ 42.9 ′ E 21-04-1996 MNHN 1758 4 – Off Baleares Islands, Mediterranean – MNHN 1759 1 – Talisman, St. 12, coast of Morocco, 35 ◦ 11 ′ N, 02-06-1883 9 ◦ 31 ′ W

MNHN 1760 1 – Travailleur, St. 27, 38 ◦ 0 ′ N, 11 ◦ 34 ′ W 26-07-1882 MNHN 1761 1 – Travailleur, St. 32, 36 ◦ 36 ′ N, 9 ◦ 46 ′ W 25-07-1882 MNHN 1762 fragments – Talisman, St. 78, off Sahara, 23 ◦ 58 ′ – 23 ◦ 55 ′ N, 11-07-1883 19 ◦ 32 ′ – 19 ◦ 35 ′ W

MNHN 2432 1 – Thalassa, St. W 393, 44 ◦ 06.9 ′ N, 04 ◦ 04.7 ′ W 08-10-1970 MNHN 2436 1 – Thalassa, St. W 415, 43 ◦ 55.1 ′ – 43 ◦ 52.5 ′ N, 11-10-1970 06 ◦ 11.3 ′ – 6 ◦ 10.9 ′ W

MNHN 2439 2 – Thalassa, St. W 444, 44 ◦ 10.5 ′ – 44 ◦ 10.1 ′ N, 13-10-1970 08 ◦ 38.4 ′ – 8 ◦ 38.5 ′ W

MNHN 2441 1 – Thalassa, St. W 413, 43 ◦ 50 ′ N, 06 ◦ 08.9 ′ W 11-10-1970 MNHN 2472 1 – Thalassa, St. W 440, 44 ◦ 11.1 ′ N, 08 ◦ 38.8 ′ W 13-10-1970 MNHN 2477 1 – Thalassa, St. W 452, 44 ◦ 11.1 ′ N, 08 ◦ 36 ′ W 14-10-1970 MNHN 2484 1 – Thalassa, St. W 434, 44 ◦ 11.8 ′ N, 08 ◦ 40.4 ′ W 13-10-1970 MNHN 2485 1 – Thalassa, St. W 445, 44 ◦ 11.7 ′ N, 08 ◦ 39.9 ′ W 13-10-1970

450–455 – 1129–1293

1179 1600 1321 1153–1275 – 958

470 440

1400–1435

970–590 1150–860

580–452

500–540

– 500–495 500–540 500–600

MNHN 2486 10 – Thalassa, St. W 394, 44 ◦ 04.6 ′ N, 04 ◦ 51.5 ′ W 08-10-1970

MNHN 2512 3 – Thalassa, St. U 863, 47 ◦ 11.4 ′ N, 05 ◦ 39.8 ′ W 1968

MNHN 2531 2 – Thalassa, St. U 862, 47 ◦ 10.3 ′ N, 05 ◦ 41.4 ′ W 1968

MNHN 2536 3 – Thalassa, St. U 836, 43 ◦ 29.5 ′ N, 09 ◦ 18 ′ W 1968

USNM E2582 1 – Albatross, St. 2754, north Pacific, Caribbean Sea, 05-12-1887 north-east of Trinidad and Tobago, 11 ◦ 40 ′ N,

58 ◦ 33 ′ W

USNM E2587 1 – Fish Hawk, St. 7514, north Atlantic, Cape 25-03-1903 Florida, Off Fowey Rocks light

USNM E2588 1 – Fish Hawk, St. 7285, north Atlantic, Gulf of 19-02-1902 México, Florida, Florida Keys, west of key

west, 24 ◦ 15 ′ N, 81 ◦ 47 ′ 30 ′′ W

USNM E8574 1 – Artic Ocean, Norwegian Sea, Norway, – Moldefjorden

USNM E10591 1 – Coast of Norway –

USNM E17020 4 – Norway, Hardangerfjorden –

USNM E18292 1 – Pola, St. 229, Mediterranean Sea , Aegean Sea , – Greece, Sporades, Samos Island, 37 ◦ 54 ′ N ,

26 ◦ 43 ′ E

USNM E19576 8 – North Atlantic , United States, off Georgia, 30-08-1979 30 ◦ 59 ′ N, 80 ◦ 8 ′ W

USNM E34354 1 – Icita , St. 13 / 8, north Atlantic , Gulf of Guinea, 02-05-1964 Off of Liberia, 6 ◦ 32 ′ 30 ′′ N, 11 ◦ 29 ′ 30 ′′ W

USNM E51998 1 – Ancon, St. C 19, north Atlantic , Caribbean Sea, 11-1998 Colombia, Guajira, Cabo de la Vela ,

12 ◦ 19 ′ 5 ′′ N, 72 ◦ 42 ′ 52 ′′ W

NOCS 1 – RRS Challenger, St. 50503 #1, 01-06-1979 51 ◦ 37.1 ′ – 51 ◦ 35.0 ′ N, 13 ◦ 14.6 ′ – 13 ◦ 15.34 ′ W

500 450 630 510–488 1609

365

559

–

–

– 580

46

400

462

104–992

(Continued)

Museum catalogue No. of TL / W (mm) Locality number specimens

Date Depth (m)

NOCS 1 – RRS Challenger, St. 50505 #1, 01-06-1979 51 ◦ 44.1 ′ – 51 ◦ 43.2 ′ N, 12 ◦ 46.3 ′ – 12 ◦ 50.6 ′ W

NOCS 1 – RRS Challenger, St. 50519 #1, 08-06-1979 49 ◦ 29.5 ′ – 49 ◦ 29.4 ′ N, 12 ◦ 48.9 ′ – 12 ◦ 43.8 ′ W

NOCS 1 – RRS Challenger, St. 50704 #1, 49 ◦ 40 ′ N, 12 ◦ 07 ′ W 13-10-1979

NOCS 3 – RRS Challenger, St. 50709 #1, 49 ◦ 23.5 ′ N, 16-10-1979 12 ◦ 21.5 ′ W

NOCS 2 – RRS Challenger, St. 50802 #1, 49 ◦ 39.5 ′ N, 30-07-1980 12 ◦ 36.9 ′ W

NOCS 2 – RRS Challenger, St. 50809 #1, 49 ◦ 31.9 ′ N, 01-08-1980 12 ◦ 09.7 ′ W

NOCS 2 – RRS Challenger, St. 50815 #1, 51 ◦ 36.1 ′ N, 05-08-1980 13 ◦ 04.2 ′ W

NOCS 1 – RRS Challenger, St. 50819 #1, 52 ◦ 04.5 ′ N, 06-08-1980 13 ◦ 29.2 ′ W

NOCS 1 – RRS Challenger, St. 51025 #1, 49 ◦ 26.5 ′ N, 10-05-1981 11 ◦ 25.1 ′ W

NOCS 1 – RRS Challenger, St. 51313 #1, 49 ◦ 32.1 ′ N, 12-02-1982 12 ◦ 11.7 ′ W

NOCS 1 – RRS Challenger, St. 51403 #1, 25-03-1982 51 ◦ 37.7 ′ – 51 ◦ 36.6 ′ N, 12 ◦ 59.8 ′ – 13 ◦ 00.0 ′ W

NOCS 1 – RRS Challenger, St. 51403 #5, 26-03-1982 51 ◦ 37.8 ′ – 51 ◦ 37.3 ′ N, 12 ◦ 58.9 ′ – 12 ◦ 59.0 ′ W

1250

1465–1431

1260–1265 1260

1857–1910

1250–1260

1280–1344

512–500

460–480

1250

1292–1314

1289–1297

NOCS

NOCS

NOCS

NOCS

7 – RRS Challenger, St. 51403 #7, 26-03-1982 51 ◦ 36.4 ′ – 51 ◦ 39.2 ′ N, 12 ◦ 59.6 ′ – 12 ◦ 58.8 ′ W

1 – RRS Challenger, St. 51420 #1, 02-04-1982 51 ◦ 37.3 ′ – 51 ◦ 36.9 ′ N, 12 ◦ 58.6 ′ – 12 ◦ 58.6 ′ W

1 – RRS Challenger, St. 51420 #4, 02-04-1982 51 ◦ 37.9 ′ – 51 ◦ 37.5 ′ N, 12 ◦ 59.5 ′ – 12 ◦ 59.6 ′ W

1 – RRS Challenger, St. 51707 #1, 12-05-1983 51 ◦ 39.9 ′ – 51 ◦ 39.2 ′ N, 13 ◦ 00.0 ′ – 13 ◦ 3.4 ′ W

1320–1247

1326–1328

1279–1287

1230–1205

Note: ∗ Labelled as “ Holotype of M. triradiata ”.

Remarks

Mesothuria intestinalis is a well-defined species, although in the old publications it was often confused with M. verrilli and possibly with M. gargantua . The type specimen does not exist. The species was described in detail by Östergren (1896) and Heding (1942a). The age variation of ossicles is similar to M. verrilli , i.e. ossicles in young specimens are larger than in older specimens. It is a large form, growing up to 300 mm in length. Mesothuria intestinalis can be easily separated from the other Mesothuria species by looking at the gonads: the sequential hermaphrodite gonads of M. intestinalis consist of many long tubules, arranged loosely inside the body cavity.

Rathke has often been cited as a co-author of Holothuria intestinalis , but he merely edited the volume after the death of Ascanius in 1803. Koehler (1896) regarded Mesothuria verrilli (Théel, 1886) from the West Indies as a synonym of M. intestinalis , however, Östergren (1896) found that they were distinct species. Mesothuria verrilli differs from M. intestinalis in (1) having more slender and very crowded tubefeet, (2) the spire of the tables ending in four simple prolongations, and (3) reduced tables in the tubefeet. Madsen and Hansen (1994) pointed out that the tables of M. intestinalis resemble those of the dendrochirotid Ekmania barthi , which are rather heavier, shorter and thicker than those of M. intestinalis .

Hérouard (1923) reported that some specimens found in the Mediterranean Sea by the Hirondelle II had 22 tentacles. The two extra tentacles were smaller than the rest and were placed ventrally.

The holotype of M. triradiata at ZMUC has been examined by F. Solis-Marin and found to correspond with M. intestinalis .

Distribution

This species is distributed all over the North Atlantic, including West Indian seas and the Mediterranean ( Figure 3 ) (see also Madsen and Hansen 1994). Records in the Mediterranean include Zavodnik and Simunovic (1985), Tortonese (1952, 1961).

Bathymetric range

This is believed to be 20–2480 m. Shallower records tend to occur in more northerly waters: the shallowest records come only from cold waters off Norway. In the Porcupine Seabight the species is found between 460 and 1465 m depth ( Billett 1988). The deepest record (2480 m) corresponds to type locality of M. triradiata off Morocco ( Heding 1942b). Other reports of M. intestinalis occurring deeper than 2000 m ( Tortonese 1949) appear to be the result of confusion with M. milleri sp. nov. (in the north-east Atlantic) and M. verrilli (in the western Atlantic).

Biological information

The species is hermaphroditic ( Théel 1886a, 1901; Östergren 1896; Hérouard 1923). Eggs large and yolky (> 0.6 mm in diameter); the development, therefore, probably direct and lecithotrophic, without a pelagic larval stage. Lives on a variety of bottoms, from sand with pebbles to soft clay and usually covers itself with shell fragments and other bottom material. Théel (1886a) was the first to suggest continuous reproduction in this species. It is rather difficult to preserve, as it usually eviscerates during capture.

Figures

Figure 1. External morphology of Mesothuria intestinalis (from Ascanius, 1805).

Figure 3. Distribution of Mesothuria verrilli, Mesothuria milleri sp. nov. and Mesothuria intestinalis.

Figure 6. Distribution of a group of Mesothuria and Zygothuria species.

Figure 8. Mesothuria gargantua Deichmann, 1930. Ossicles from type specimen (MCZ cat. Nr 449).

Figure 9. Ossicles of Mesothuria maroccana Perrier, 1902 (A,B) and Mesothuria intestinalis (Ascanius, 1805) (C,D). (A,B) Specimen from St. 50519 #1; (C,D) specimen from St. 51403 #7.