Hanleya nagelfar ( Lovén, 1846 ), Kaas and Van Belle, 1985

Sirenko, Boris, 2014, Composition of the genus Hanleya (Mollusca: Polyplacophora: Lepidopleurida), with the description of two new species, Journal of Natural History 48 (45 - 48), pp. 2913-2945 : 2931-2937

Hanleya nagelfar ( Lovén, 1846)

( Figures 9–19 )

Chiton nagelfar Lovén, 1846: 158 .

Hanleya nagelfar Kaas and Van Belle 1985: 196 , Figure 92, map 19 (synonymy); Waren and Klitgaard 1991: 51, Figures 1–6 ; Dell’ Angelo et al. 1998: 244, PL 1, Figure 10 .

Type material

Lectotype, NHRM, type collection 1329 (designated by Kaas and Van Belle (1985)).

Type locality

Finnmark, Norway.

Material examined

Barents and Norwegian Seas: R/V Andrey Pervozvaniy, 1899, stn 4, 70.0230° N, 31.49° E, 144–178 m, 2 spms; stn 38, 69.58° N, 32.40° E, 230 m, 1 spm; 1900, stn 102, 74.27° N, 22.04° E, 180–211 m, 1 spm ( Table 2 , No. 8); stn 462, 71.20° N, 33.30° E, 304– 280 m, 1 spm; stn 464, 71.21° N, 3159° E, 268 m, 1 spm. R/V Persey, 1924, stn 228, 70.3030° N, 31.13° E, 175 m, 1 spm ( Table 2 , No. 10); 1927, stn 677, 69.31° N, 34.39° E, 173 m, 1 spm; 1935, stn 3337, 71.35° N, 31.464° E, 313 m, 1 spm ( Table 2 , No. 11).

R/V SRT4225, cruise 1, 1955, stn 38/104, 61.308° N, 04.166° E, 300 m, 1 spm. R/ V Sevastopol. RT-97, cruise 8, 1958, Lofoten Ids., 68.08° N, 13.50° E, 1 spm ( Table 2 , No. 7); cruise 9, 1958, stn 1453, 62.002° N, 6.146° W, 112 m, 1 spm; stn 1488, 72.291° N, 23.072° E, 335 m, 2 spms. R/ V Maslov, cruise 5, 1969, stn 180/201, 70.40° N, 17.275° E, 510 m, 1 spm; stn 182/203, 71.30° N, 16.46° E, 300 m, 1 spm ( Table 2 , No. 13); stn 183/204, 72.02° N; 17.02° E, 360 m, 1 spm ( Table 2 No. 12); stn 188/209, 73.20° N, 15.25° E, 500 m, 1 spm. R/ V Tunets, cruise 105, 1978, stn 4, 70.06° N, 32.325° E, 230 m, 1 spm; stn 11, 70.110° N, 17.350° E, 225 m, 2 spms; stn 12, 70.330° N, 16.540° E, 1100 m, 5 spms. R/ V Dalnie Zelentsi, cruise, 16.1982, stn 64, 67.559° N, 10.080° E, 280 m, 3 spms. F.D. Behrens, 1948, Lofoten, 2 spm ( ZIM). F.D. Saturn, 1958, 64.44° N, 35.06° W, 500 m, 1 spm. ( ZIM) ; 1923, Folden, 50–100 m, 1 spm, ( TM 13350 ) ; Skattora, Tromso, 54 m, 2 spms, ( Table 2 , No. 5), ( TM 12923 ) ; 1951, Senja, Steinavar, 360–365 m, 1 spm, ( TM 13335 ) ; 1954, Morsdalsfjord, 50–110 m, 1 spm, ( TM 13196 ) ; 1954, Balsnes, 28 m, 1 spm, ( TM 13258 ) ; 1954, Brokskar, Rystr, 50–120 m, 1 spm ( Table 2 , No. 3), ( TM 12962 ) ; Beiarn, 40–90 m, 1 spm, ( TM 13237 ) ; 1955, Tjelsund, 60–110 m, 1 spm, ( TM 12955 ) ; 1955, Holandsfjord, 45–770 m, 1 spm, ( TM 13241 ) ; 1955, Eiet, 52–58 m, 6 spms, ( TM 13001 ) ; Masoy, 125– 90 m, 2 spms, ( TM 13299 ) ; 1955, Bjarandfjord, 76–92 m 1 spm, ( TM 12997 ) ; 1955, Rana, 35–42 m, 1 spm, ( TM 13304 ) .

Eastern Greenland Sea: R/ V Atlantida, cruise 10, 1973, stn 18/1668, 74.50° N, 16.00° E, 360 m, 1 spm .

Southeastern Greenland: R/ V Sevastopol, RT-97, cruise 5, 1957, stn 1168, 66.162° N, 31.356° W, 345 m, 2 spms; stn 1609, 65.526° N, 29.586° W, 425 m, 1 spm .

Central and North Atlantic Ocean: R/ V Akademik Keldysh, cruise 4, 1982 stn 513, 34.261° N, 30.030° W, 690 m, 1 spm; stn 444, 58.208° N, 31.363° W, 1635– 1535 m, 1 spm; stn 364, Reykjanes Ridge, 1330 m, 1 spm; st. 385, Reykjanes Ridge, 1680– 1800 m, 1 intermediate valve; R/ V Ichtiandr cruise 8, 1982, stn 39, 09.038° N, 21.101° W, 610–700 m, 1 spm; stn 50, 33.141° N, 29.425° W, 700–740 m, 1 spm .

Newfoundland Bank: R/V Sevastopol, RT-97, cruise 16, 1960, stn 2805, 49.477° N, 50.185° W, 318–324 m, 1spm; R/V Nereida, 2009 , cruise 0509, stn 6, cod 5, 48.2915° N, 44.0609° W, 1348 m, 1 spm; stn 10, cod 54, 48.0005° N, 43.7607° W, 1554 m, 1 spm; stn 12, cod 16, 47.7772° N, 43.5748° W, 1462 m, 1 spm; stn 19, cod 21, 47.1643° N, 43.5338° W, 1358 m, 2 spms ( Table 2 , No. 4), stn 22, cod 12, 46.8400° N, 43.6404° W, 956 m, 1 spm ( Table 2 , No. 6); stn 23, cod 22, 46.7749° N, 43.8651° W, 1127 m, 3 spms; stn 24, cod 27, 46.6942° N, 43.9686° W, 1104 m, 1 spm; 2010, cruise 0610, stn 63, cod 8, 46.3697° N, 44.9944° W, 1406 m, 2 spms; stn 67, cod 7, 46.2677° N, 46.5429° W, 613 m, 1 spm; stn 69, cod 10a, 46.1543° N, 46.4503° W, 502 m, 1 spm; stn 69, cod 10c, 46.1543° N, 46.4503° W, 502 m, 2 spms; stn 71, cod 10, 46.0800° N, 46.3421° W, 888 m, 1 spm; cruise 0710, stn 75, cod. 24, 45.8692° N, 46.8210° W, 1227 m, 1 spm ( Table 2 , No. 1); stn 96, cod. 30, 43.9894° N, 48.9512° W, 863 m, 1 spm; cruise 0810, stn 99, cod 11, 43.6147° N, 49.0781° W, 548 m, 1 spm; stn 100, cod 12, 43.4844° N, 49.1855° W, 694 m, 2 spm; stn 103, cod 20, 429433° N, 49.5621° W, 1154 m, 1 spm. FN3L, 2006, stn 2, 47.9415° N, 47.3725° W, 411 m, 1 spm; FN3L, 2006, stn 9, 47.9370° N, 47.1883° W, 460 m, 1 spm; FN3L, 2008, stn 37, 47.6480° N, 47.1340° W, 503 m, 2 spms; FN3L, 2008, stn 84, 46.4488° N, 47.2603° W, 396 m, 1 spm; FN3L, 2009, stn 89, 461100° N, 47.3333° W, 1005 m, 2 spms; PLA, 2008, stn 98, 43.6298° N, 48.9907° W, 1436 m, 4 spms ( Table 2 , No. 9).

North America: Off Eastport, Maine (Jeffreys coll.), 1 spm, ( USNM, 177511); Off Martha’ s Vineyard, Massachussetts, 1 spm, ( USNM); Georges Bank, 270 m, 1 spm ( USNM); Nova Scotia, 72 m, 1 spm ( USNM).

Distribution

North and Central Atlantic Ocean , the Barents , Norwegian and Greenland seas, near southern Greenland, North America , Europe and northern Africa, on Mid-Atlantic Ridge from 74.27° N to 09.038° N at depths from 28 to 1680 m , mostly deeper than 200 m.

Large specimens of Hanleya nagelfar are easily distinguished from H. hanleyi (see Kaas and Van Belle, 1985), but small chitons (<10 mm) are difficult to identify.

In order to study age variability in H. nagelfar , seventeen specimens of different sizes (BL = 5.0–73.0 mm), from different regions of the Atlantic Ocean were measured ( Table 2 , Figures 8–18 ). These measurements revealed that important changes occur in the shape of the valves, the thickness of the girdle, and the shape, size and ribbing of the dorsal spicules of the girdle.

With increasing body length, the ratio of head valve width/tail valve width decreased from 1.27 (BL = 5.0 mm) to 0.84 (BL = 70.0 mm); the ratio of valve V width/valve V length decreased from 2.56 (BL = 5.0 mm) to 1.36 (BL = 70.0 mm); and the ratio of valve V width/width of adjacent girdle also decreased from 5.9 (BL = 5.0 mm) to 1.36 (BL = 70.0 mm) ( Table 2 ).

Small and medium size specimens (BL = 5.0–29.0 mm) have flattened, short and pointed spicules with 6–7 ribs arranged around the spicule ( Figures 10A, C , 12A, B , 14A, C, D, E , 16A–C , 17D , 18A, C ). With increasing age (specimen size) the spicules become longer and the ribs become less prominent or obsolete ( Figures 18B, D , 19E ).

Figures

Figure 1. Distribution of the genus Hanleya. Black circle: specimens studied by author; empty circle: literature data; triangle: fossil specimens.

Figure 2. Hanleya harasewychi sp. nov., holotype (A–E, G, H) and paratype, USNM 454684, Florida off Foway Light, BL 3 mm (F). (A) valve I, dorsal view; (B) valve V, dorsal view; (C) valve VIII, dorsal view; (D) valve VIII, lateral view; (E) valve V, front view; (F, H) whole specimen, dorsal view; (G) valve V, dorsal view. Scale bars: 1mm.

Figure 3. Hanleya harasewychi sp. nov., holotype. (A) valve I, front view; (B) valve V, jugal area; (C) valve VIII, ventral view; (D) valve VIII, pleural are; (E) valve V, ventral view; (F) dorsal, marginal and ventral spicules and needles. Scale bars: A, B, D, F: 100 μm; C, E: 1 mm.

Figure 4. Hanleya harasewychi sp. nov., holotype (A–H), paratype, USNM 323067, Florida, off Key West, BL 5.0 mm (I, J). (A) dorsal and marginal needles; (B) dorsal spicules; (C) ventral spicules near margin; (D) ventral spicules; (E) aesthete group; (F) central, first lateral and major lateral teeth of radula; (G, H) head of major lateral teeth of radula; (I) dorsal spicules; (J) dorsal and marginal needles. Scale bars: A–D, F–J: 100 μm.

Figure 5. Hanleya harasewychi sp. nov., paratype, USNM 323067, Florida, off Key West, BL 5 mm. (A) valve I, dorsal view; (B) valve III, dorsal view; (C) valve III, ventral view; (D) valve I, front view; (E) valve III, jugal and pleural areas; (F) valve III, pleural area. Scale bars: A–D: 1 mm, E, F: 100 μm.

Figure 6. Hanleya mediterranea sp. nov., holotype, ZISP 2201, Mediterranean Sea, Spain, Gerona, Bagur, depth 200–300 m, BL 5.4 mm. (A) valve I, dorsal view; (B) valve V, dorsal view; (C) valve VIII, dorsal view; (D) valve V, dorsal view; (E) valve V, central and lateral areas; (F) valve VIII, lateral view; (G) valve IV, ventral view.

Figure 8. Hanleya mediterranea sp.nov., paratype, ZISP 2202, Prinkipo, Mediterranean Sea, depth 50 m, BL 4.7 mm (A–J). Hanleya tropicalis (Dall, 1881), paratype, USNM 95172, Florida, Sands Key Pourtales, depth 230 m, BL 10 mm (K). (A) valve V, frontal view; (B) tail valve, lateral view; (C) dorsal and marginal needles; (D) dorsal scales; (E) ventral scales; (F) central and first lateral tooth of radula; (G, H) head of major lateral tooth of radula; (I, J) aesthete group. Scale bars: A, B: 1 mm, C, H, K: 100 μm.

Figure 9. Hanleya nagelfar (Lovén 1846), Newfoundland Bank, R/V NEREIDA, cruise 0710, stn. 75, 24, depth 1227 m, BL 5.0 mm. (A) head valve, dorsal view; (B) valve V, dorsal view; (C) valve VIII, dorsal view; (D) valve V, front view; (E) pleural and lateral areas of valve V; (F) valve IV, ventral view; (G) tail valve, lateral view.

Figure 10. Hanleya nagelfar (Lovén, 1846), Newfoundland Bank, R/V NEREIDA, cruise 0710, stn. 75, 24, depth 1227 m, BL 5.0 mm. (A) dorsal and ventral spicules and needles; (B) mucro of tail valve; (C) dorsal spicules; (D) radula.

Figure 11. Hanleya nagelfar (Lovén, 1846), Newfoundland Bank, R/V NEREIDA, cruise 0610, stn. 69, 10a, depth 502–504 m, BL 7.0 mm. (A) valve I, dorsal view; (B) valve V, dorsal view; (C) valve VIII, dorsal view; (D) valve I, front view; (E) valve IV, ventral view; (F) valve V, central and lateral areas; (G) valve VIII, lateral view; (H) radula.

Figure 12. Hanleya nagelfar (Lovén, 1846), Newfoundland Bank, R/V NEREIDA, cruise 0610, stn. 69, 10a, depth 502–504 m, BL 7.0 mm. (A, B, D) dorsal specules and needles; (C) ventral scales.

Figure 13. Hanleya nagelfar (Lovén, 1846), Northern Norway, depth 52–58 m, BL 7.0 mm. (A) valve I, dorsal view; (B) valve V, dorsal view; (C) valve VIII, dorsal view; (D) valve V, front view; (E) valve V, central area; (F) valve VIII, lateral valve; (G) valve IV, ventral valve.

Figure 14. Hanleya nagelfar (Lovén, 1846), Northern Norway, depth 52–58 m, BL 7.0 mm. (A) dorsal, marginal and ventral spicules and needles; (B) radula; (C) dorsal spicules and needles.

Figure 15. Hanleya nagelfar (Lovén, 1846), Reykjanes sea mounts, R/V Akademik Keldish, stn. 364, depth 1330 m, BL 14.0 mm. (A) valve I, dorsal view; (B) valve V, dorsal view; (C) valve VIII, dorsal view; (D) valve IV, ventral view; (E) valve V, central and lateral areas; (F) valve VIII, lateral view; (G) valve I, dorsal view; (H) valve V, front view.

Figure 16. Hanleya nagelfar (Lovén, 1846), Reykjanes sea mounts, R/V Akademik Keldish, stn. 364, depth 1330 m, BL 14.0 mm. (A) dorsal, marginal and ventral, spicules, needles and scales; (B, C) dorsal spicules and needles; (D) radula.

Figure 17. Hanleya nagelfar (Lovén, 1846), Eastport, Maine, USNM 177511, BL 15 mm (A–H); (A) valve V, frontal view; (B) valve VIII, lateral view; (C) dorsal and marginal spicule; (D) dorsal spicule; (E) ventral spicule; (F, G) head of major lateral tooth of radula; (H) aesthete group. Scale bars: A, B: 1 mm, C–G: 100 μm.

Figure 18. Hanleya nagelfar (Lovén, 1846), Northern Norway near Tromsø, depth 54 m, BL 28.0 mm (A); Norway, depth 300 m, BL 70 mm (B); Newfoundland Bank R/V NEREIDA, cruise PLA08, stn. 98, depth 1436 m, BL 29.0 mm (C); Central Atlantic, R/V Ikhtiandr, st. 50 depth 700–740 m, BL 73 mm (D). (A,D) dorsal spicules and needles.

Figure 19. Hanleya nagelfar (Lovén, 1846), Southeastern Barents Sea, depth 300 m, BL 36 mm. (A) valve V, frontal view; (B) valve VIII, lateral view; (C) aesthete group; (D) dorsal and marginal needles; (E) dorsal spicules; (F) ventral spicule; (G) central and first lateral tooth of radula; (H) head of major lateral teeth of radula. Scale bars: A, B: 1 mm, D–H: 100 μm.