Paracaudina alta, Davey, Nicola & O’Loughlin, Mark, 2013

Davey, Nicola & O’Loughlin, Mark, 2013, The caudinid sea cucumbers of New Zealand (Echinodermata: Holothuroidea: Molpadida: Caudinidae), Zootaxa 3613 (4), pp. 357-368 : 361-366

Paracaudina alta sp. nov.

Figures 1 b, 3

Paracaudina species O’Loughlin et al. 2011: 63.

Material examined. Holotype. New Zealand: Kaikoura Basin: NIWA 68129, Stn TAN0705/277, 42.62 S, 175.92 E, 1202–1215 m, 26/04/2007. Paratypes. NIWA 43346 (7), station data as holotype.

Other material. NIWA registered specimens. Kaikoura Basin: 34288 (4), 34289 (9) Stn TAN0705/269, 42.59 S, 176.38 E, 1201–1203 m, 26/04/2007; 34291(1) Stn TAN0705/274, 42.62 S, 175.93 E, 1183–1203 m, 26/04/ 2007; 34294 (6), Stn TAN0705/279, 42.65 S, 175.5 E, 1180–1204 m, 27/04/2007; 34300 (2) Stn TAN0705/293, 42.54 S, 175.14 E, 1812–1813 m, 27/04/2007; 77383 (3) Stn G955, 42.68 S, 174.76 E, 1345 m, 31/08/2010. Hikurangi Margin: 63034 (1) Stn TAN1004/6, 41.7 S, 175.65 E, 1040–1080 m, 15/04/2010; 63195 (1) Stn TAN1004/21, 41.51 S, 175.72 E, 1076–1104 m, 17/04/2010; 63806 (5) Stn TAN1004/85, 41.95 S, 174.62 E, 964–1005 m, 23/04/2010; 77385 (3) Stn E438, 41.63 S, 174.88 E, 1064.0 m, 19/10/1965.

Description. Paracaudina species up to 98 mm long (NIWA 34289, preserved), main body 90 mm, tail 8 mm long, width up to 15 mm; cylindrical body with caudal taper and long distinct tail, tail up to full length of body but often broken off; body leathery, firm and parchment-like, prickly, smooth to extensively wrinkled, mainly transverse creasing, round to flattened body form (preservation), tentacles 15, equal with 2 tentacle digit lobes on each side.

Ossicles in mid-body wall and tail irregularly shaped closed knobbed cups with a central cross. The form is variable and can be twisted, round, triangular, elongate or octagonal, variable sizes up to 85 µ m long. Some closed cup variations have the chilensis -like central perforation with an elevated upper open bridged basal box with 4 supporting arms on one side, however these regular chilensis -like ossicles are rare; others variations are open and round in cross section. Surface of the closed cup ossicles variably knobbed, often 8 blunt projections, or smooth.

Colour (preserved). Off-white, cream, orange, often with anterior end more orange colour than rest of body.

Distribution. New Zealand: Hikurangi Margin, Kaikoura Basin, 964–1813 m.

Etymology. From the Latin alta (deep), referring to the deep water-only occurrence of this species.

Remarks. O’Loughlin et al. (2011) anticipated the possible occurrence of a new species of Paracaudina , east of Cook Strait (Hikurangi Margin) and noted that it was different from any known New Zealand species ( Paracaudina coriacea (Hutton, 1872) or Paracaudina chilensis (Müller, 1850)) . This was based on only two specimens they examined. Forty-two specimens have now been examined that show the characteristics described above and confirm O’Loughlin et al. ’s (2011) suspicions. Paracaudina alta sp. nov. is indistinguishable from other Paracaudina species at the macro level. However, P. alta sp. nov. is readily distinguished from the other New Zealand species of Paracaudina by the very irregular form, size and thickness of the mid-body and tail ossicles with only a rare presence of chilensis -like ossicle forms. The irregular form and often twisted shapes of the ossicles differ significantly from the regular symmetry of the P. c h i l e n s i s ossicles and the smooth, thick, round ossicles of the P. coriacea . P. a l t a sp. nov. only occurs in deep water east of Cook Strait, and is not sympatric with the other New Zealand Paracaudina species.

Paracaudina chilensis (Müller, 1850) Figures 1 c, 4

Synonymy. See O’Loughlin et al. (2011)

Material examined. NHMUK registered specimens: NHMUK 1938.8.23.68 (1), Antarctic expedition, no further station details; NHMUK 1875.10.27.22 (1), Wellington, New Zealand. On a ship’s anchor cable. Lieu. Praed. Jar label states: W. Wykeham-Perry, no further station details; NHMUK 1886.10.2.39 (17), Stn167A, 41.07 S, 174.32 E, Queen Charlotte’s Sound, near Long Island, Challenger Expedition, 10 fms.

NIWA registered specimens: New Zealand oceanic: 77289 (1) Stn B323, 40.25 S, 173.63 E, 88 m, 27/10/1960; 77284 (1) Stn B471, 43.33 S, 169.91 E, 62 m, 03/06/1961; 77287 (1) Stn B495, 44.56 S, 167.79 E, 120 m, 09/06/ 1961; 77291 (1) Stn B496, 40.60 S, 173.55 E, 54 m, 11/06/1961; 77290 (1) Stn C184, 39.83 S, 173.5 E, 95 m, 06/ 09/1959; 77288 (1) from Stn C223; 77268 (1), Stn C259, 41.10 S, 174.27 E, 18 m, 20/10/1959; 77286 (1) Stn C411, 41.57 S, 174.57 E, 150 m, 04/05/1960; 77191 (1) Stn C745, 36.66 S, 174.22 E, 35 m, 16/02/1962; 77263 (1), 77269 (1) Stn C787, 36.66 S, 175.10 E, 43 m, 22/02/1962; 77252 (1) Stn C846, 41.64 S, 174.50 E, 126 m, 01/03/ 1962; 77212 (1) Stn C852, 40.68 S, 174.33 E, 132 m, 02/03/1962; 77382 (1) Stn D285, 41.30 S, 174.80 E, 17 m, 18/11/1964; 77418 (1) Stn D868, 43.90 S, 179.7 W, 420 m, 24/03/1969; 77411 (1) Stn E225, 66.51 S, 162.43 E, 209 m, 12/02/1965; 77373 (1) Stn E293, 34.29 S, 172.41 E, 205 m, 08/04/1965; 77409 (3) Stn E389, 34.02 S, 172.72 E, 155 m, 16/04/1965; 77374 (1) Stn E704, 40.03 S, 177.14 E, 278 m, 21/03/1967; 77258 (1) Stn E747, 40.72 S, 176.80 E, 554 m, 29/03/1967; 77370 (1) Stn E780, 43.39 S, 169.44 E, 252 m, 16/10/1967; 77275 (1) Stn G171, 43.58 S, 179.4 W, 417 m, 17/01/1968; 77407 (1) Stn G186, 44.06 S, 179.6 W, 402 m, 18/01/1968; 77283 (1) Stn G192, 43.84 S, 179.6 W, 364 m, 18/01/1968; 77402 (1) Stn G210, 43.5 S, 179.93 E, 424 m, 19/01/1968; 77384 (1) Stn G219, 43.61 S, 179.83 E, 402 m, 21/01/1968; 77266 (1) Stn G276A, 43.58 S, 179.25 E, 410 m, 24/01/1968; 77277 (1) Stn G291, 43.70 S, 179.01 E, 402 m, 25/01/1968; 77379 (1) Stn G31, 43.71 S, 176.00 E, 348 m, 23/02/ 1967; 77386 (1) Stn G33, 43.66 S, 177.00 E, 457 m, 23/02/1967; 77372 (1) Stn G343, 43.79 S, 178.8 W, 408 m, 02/ 02/1968; 73975 (1) Stn G379, 43.63 S, 177.9 W, 380 m, 06/02/1968; 77220 (1) Stn G682, 45.72 S, 170.72 E, 33 m, 20/01/1970; 77406 (1) Stn H930, 43.47 S, 179.51 E, 402 m, 14/08/1975; 77207 (1) Stn I44, 34.99 S, 174.02 E, 102 m, 08/05/1975; 77272 (1) Stn J829, 35.84 S, 174.46 E, 0 m, 24/07/1975; 77366 (1) Stn K89, 77347 (1) Stn K90, 77352 (1) Stn K93, 77361 (1) Stn K95, 77336 (1) Stn K98, 77355 (1) Stn K99, 77360 (1) Stn K102, 77318 (1) Stn K105, 77193 (1) Stn K126, 77233 (1) Stn K133, 77242 (1) Stn K136, 77219 (1) Stn K137, 77199 (1) Stn K138, 77217 (1) Stn K151, 77246 (1) Stn K153, 77226 (1) Stn K173, 77255 (1) Stn K177, 41.17 S, 173.16 E, 14 m, March-April 1971; 77310 (1) Stn K199, 41.16 S, 173.16 E, 18 m, 01/07/1971; 77196 (1) Stn K289, 77209 (1) Stn K290, 77230 (1) Stn K293, 77198 (1) Stn K296, 77213 (1) Stn K298, 77194 (1) Stn K299, 77256 (1) Stn K300, 77216 (1) Stn K304, 77237 (1) Stn K309, 77229 (1) Stn K310, 77236 (1) Stn K313, 77214 (1) Stn K314, 77245 (1) Stn K319, 77249 (1) Stn K316, 77234 (1) Stn K317, 77244 (1) Stn K323, 41.17 S, 173.16 E, 15 m, 14/12/1971; 77294 (1) Stn K326, 77297 (1) Stn K327, 77330 (1) Stn K329, 77304 (1) Stn K331, 77327 (1) Stn K333, 70956 (2) Stn K334, 70956 (1) Stn K334, 77324 (1) Stn K335, 77312 (1) Stn K336, 77313 (1) Stn K338, 77302 (1) Stn K339, 77323 (1) Stn K340, 77309 (1) Stn K341, 77311 (1) Stn K346, 77274 (1) Stn K347, 77296 (1) Stn K354, 77306 (1) Stn K355, 77301 (1) Stn K357, 77325 (1) Stn K359, 77308 (1) Stn K362, 77201 (1) Stn K387, 77210 (1) Stn K395, 77215 (1) Stn K400, 77251 (1) Stn K401, 77240 (1) Stn K412, 77205 (1) Stn K413, 77235 (1) Stn K419, 77247 (1) Stn K420, 77254 (1) Stn K427, 77206 (1) Stn K428, 77208 (1) Stn K502, 77222 (1) Stn K507, 77243 (1) Stn K508, 77221 (1) Stn K510, 77232 (1) Stn K512, 77225 (1) Stn K520, 77211 (1) Stn K526, 77298 (1) Stn K544, 77316 (1) Stn K545, 77320 (1) Stn K546, 77328 (1) Stn K550, 77307 (1) Stn K552, 77299 (1) Stn K555, 77300 (1) Stn K558, 77317 (1) Stn K559, 77436 (1) Stn K590, 77315 (1) Stn K608, 77332 (1) Stn K610, 77305 (1) Stn K613, 77314 (1) Stn K618, 77321 (1) Stn K620, 77253 (1) Stn K667, 77227 (1) Stn K669, 77248 (1) Stn K670, 77241 (1) Stn K672, 77231 (1) Stn K676, 77250 (1) Stn K677, 77228 (1) Stn K678, 77204 (1) Stn K679, 77195 (1) Stn K681, 41.17 S, 173.16 E, 0 m, January-June 1972; 77343 (1) Stn K57, 40.66 S, 172.86 E, 31 m, 26/02/1971; 77358 (1) Stn K63, 40.58 S, 173.08 E, 55 m, 27/02/1971; 77353 (1) Stn K67, 77303 (1) Stn K70, 77337 (1) Stn K73, 77363 (1) Stn K80, 77362 (1) Stn K82, 77350 (1) Stn K86, 40.75 S, 173.33 E, 47 m, 28/02/1971; 77413 (2) Stn KAH9401/21, 36.82 S, 176.30 E, 502–510 m, 09/01/1994; 77189 (1) Stn N924, 41.46 S, 175.04 E, 65 m, 28/08/1977; 77417 (1) Stn N926, 41.54 S, 175.02 E, 100 m, 28/08/1977; 77341 (1) Stn P6153, 34.86 S, 173.21 E, 18 m, 27/08/1978; 77349 (1) Stn P6274, 34.80 S, 173.19 E, 22 m, 29/08/1978; 77357 (1) Stn P6441, 34.98 S, 173.46 E, 12 m, 30/08/ 1978; 77218 (1) Stn Q12, 43.85 S, 179.8 W, 410 m, 15/03/1978; 77200 (1) Stn Q40, 44.49 S, 176.5 W, 345 m, 24/ 03/1978; 76069 (1) Stn Q691, 42.96 S, 170.20 E, 109 m, 18/02/1982; 77261 (1) Stn Q692, 42.89 S, 170.30 E, 245 m, 19/02/1982; 77381 (7) Stn S121, 43.50 S, 175.96 E, 335 m, 20/10/1979; 76070 (1) Stn S122, 43.59 S, 175.95 E, 322 m, 20/10/1979; 77190 (1) Stn S124, 43.51 S, 175.99 E, 363 m, 20/10/1979; 77389 (1) Stn S125, 43.53 S, 175.97 E, 365 m, 20/10/1979; 77197 (1) Stn S127, 43.59 S, 175.95 E, 322 m, 20/10/1979; 77223 (1) Stn S128, 43.51 S, 175.98 E, 365 m, 21/10/1979; 77257 (1) Stn S130, 43.56 S, 175.96 E, 335 m, 21/10/1979; 77279 (1) Stn S130, 43.56 S, 175.96 E, 335 m, 21/10/1979; 77331 (1) Stn S371, 42.94 S, 170.02 E, 200 m, 28/01/1983; 77273 (1) Stn S893C, 42.51 S, 170.41 E, 491 m; 77264 (1) Stn T468, 41.04 S, 174.32 E, 37 m, 06/12/1983; 77276 (1) Stn T475, 41.04 S, 174.29 E, 33 m, 07/12/1983; 77364 (1) Stn T476, 41.03 S, 174.30 E, 81 m, 07/12/1983; 77344 (1) Stn T482, 41.11 S, 174.31 E, 28 m, 08/12/1983; 77280 (1) Stn T486, 41.09 S, 174.36 E, 21 m, 08/12/1983; 77278 (1) Stn T487, 41.11 S, 174.35 E, 21 m, 08/12/1983; 77369 (1) Stn T490, 41.15 S, 174.30 E, 45 m, 08/12/1983; 77348 (1) Stn T517, 40.98 S, 174.27 E, 53 m, 12/12/1983; 77342 (1) Stn T537, 40.91 S, 174.13 E, 33 m, 13/12/ 1983; 77354 (1) Stn T560, 41.04 S, 173.98 E, 23 m, 15/12/1983; 41504 (1) Stn TAN0705/163, 42.78 S, 176.7 W, 996–1009 m, 16/04/2007; 34265 (2) Stn TAN0705/169, 43.15 S, 178.3 W, 478–487 m, 17/04/2007; 34267 (2) Stn TAN0705/176, 43.52 S, 178.6 W, 425– 425 m, 17/04/2007; 34272 (1) Stn TAN0705/187, 42.96 S, 178.7 W, 527–528 m, 18/04/2007; 34281 (1) Stn TAN0705/241, 43.10 S, 177.7 W, 533–537 m, 23/04/2007; 34280 (1) Stn TAN0705/241, 43.10 S, 177.7 W, 533–537 m, 23/04/2007; 34282 (1) Stn TAN0705/244, 43.02 S, 179.37 E, 518–528 m, 24/04/2007; 34284 (1) Stn TAN0705/248, 42.99 S, 178.99 E, 530–538 m, 24/04/2007; 34283 (1) Stn TAN0705/248, 42.99 S, 178.99 E, 530–538 m, 24/04/2007, 34295 (1) Stn TAN0705/281, 43.03 S, 175.23 E, 309–310 m, 27/04/2007; 30594 (1) Stn TAN0705/29, 43.79 S, 175.31 E, 414–419 m, 04/04/2007; 30595 (1) Stn TAN0705/29, 43.79 S, 175.31 E, 414–419 m, 04/04/2007; 34869 (1) Stn TAN0705/4, 44.18 S, 175.35 E, 536–539 m, 02/04/2007; 30546 (1) Stn TAN0705/58, 43.80 S, 178.11 E, 497–502 m, 07/04/2007; 34248 (4) Stn TAN0705/ 90, 44.10 S, 178.5 W, 459–460 m, 10/04/2007; 30717 (1) Stn TAN0707/140, 39.64 S, 172.14 E, 266–267 m, 07/06/ 2007; 30698 (1) Stn TAN0707/26, 38.62 S, 168.94 E, 478– 478 m, 29/05/2007; 77829 (4) Stn TAN0906/134, 34.46 S, 173.21 E, 140–141 m, 13/07/2009; 54966 (1) Stn TAN0906/38, 35.21 S, 174.40 E, 99–105 m, 06/07/200; 54961 (1) Stn TAN0906/38, 35.21 S, 174.40 E, 99–105 m, 06/07/2009; 77322 (1) Stn U264, 42.58 S, 173.78 E, 940 m, 16/12/1982; 77345 (1) Stn V361, 43.50 S, 178.64 E, 340 m, 06/09/1989; 77356 (1) Stn V365, 43.74 S, 179.0 W, 397 m, 08/09/1989; 77346 (1) Stn V366, 43.49 S, 178.9 W, 499 m, 08/09/1989; 77340 (1) Stn V367, 43.24 S, 179.0 W, 467 m, 08/09/1989; 77359 (1) Stn V372, 43.33 S, 178.98 E, 418 m, 13/09/1989; 77335 (1) Stn V380, 43.58 S, 176.99 E, 334 m, 15/09/1989; 77338 (1) Stn V390, 43.08 S, 177.00 E, 330 m, 17/09/1989; 77326 (1) Stn V391, 42.83 S, 176.99 E, 476 m, 17/09/1989; 77292 (1) Stn V411A, 44.49 S, 171.23 E, 14 m, 19/05/1992; 77293 (1) Stn V412A, 44.50 S, 171.23 E, 14m, 19/05/1992; 77365 (1) Stn W251, 43.83 S, 178.99 E, 442–469 m, 15/09/1993; 77380 (26) Stn W258, 42.96 S, 178.99 E, 549–602 m, 19/09/1993.

Description. See O’Loughlin et al. (2011).

Distribution. New Zealand: Chatham Rise, Challenger Plateau 0–1009 m depth (this study). Antarctica: Balleny Islands (1 record, NIWA 77411).

Remarks. An extensive diagnosis and images of this species have been recently presented by O’Loughlin et al. (2011). Prior to this study all Paracaudina specimens in the NIWA collections were identified as Paracaudina chilensis (Müller, 1850) . Further to the observations of O’Loughlin et al. 2011 where Paracaudina coriacea (Hutton, 1872) was raised out of synonymy with P. chilensis and the possibility of new species noted, we sampled all material. Much of this material remained as P. chilensis however P. coriacea and our two new species described here were recognised in this collection.

Both P. chilensis and P. coriacea are sympatric in New Zealand waters. Externally these species are not able to be distinguished; however the ossicle types are distinctive with P. c h i l e n s i s having regular shaped, octagonal closed knobbed cups with a regular central cross, predominantly with marginal bluntly pointed projections or knobs, and open spaces within the ossicles. However, P. coriacea has thick closed cups with either bulbously knobbed projections or smooth and completely lacking marginal knobs, and reduced or no spaces within ossicles.

Paracaudina coriacea (Hutton, 1872) Figures 1 d, e, 5, 6

Synonymy. See O’Loughlin et al. (2011)

Material examined. Lectotype (established here). NHMUK:1898.10.8.1 off New Brighton, S. Island, New Zealand (no further station data). Paralectotypes (2; established here). NHMUK:1898.10.8.2a, 1898.10.8.2b (known station data as lectotype).

NIWA registered specimens: 70954 (1) Stn N529, 40.58 S, 175.08 E, 61 m, 26/05/1975; 70955 (1) Stn T566, 41.12 S, 173.86 E, 18 m, 15/12/1983; 77188 (1) Stn C846, 41.64 S, 174.50 E, 126 m, 01/03/1962; 77192 (1) Stn G30, 43.76 S, 175.41 E, 366 m, 23/02/1967; 77202 (1) Stn M13, 41.1 S, 174.88 E, 2 m, 16/01/1975; 77259 (1) Stn B37, 39.42 S, 176.93 E, 17 m, 02/09/1957; 77262 (1) Stn G381, 43.53 S, 177.9 W, 377 m, 06/02/1968; 77271 (1) Stn C209, 41.16 S, 174.74 E, 40 m, 12/09/1959; 77285 (1) Stn C412, 41.59 S, 174.57 E, 225 m, 04/05/1960; 77334 (1) Stn K66, 40.58 S, 173.08 E, 55 m, 27/02/1971; 77367 (1) Stn T488, 41.13 S, 174.29 E, 37 m, 08/12/1983; 77368 (1) Stn F189, 40.5 S, 173.49 E, 73 m, 21/03/1965; 77375 (2) Stn C945, 41.24 S, 174.05 E, 41 m, 13/02/ 1963; 77376 (1) Stn C814, 37.66 S, 178.94 E, 157–209 m, 25/02/1962; 77387 (1) Stn E55, 43.16 S, 173.35 E, 70 m, 06/03/1964; 77388 (1) Stn C803, 37.66 S, 177.39 E, 133 m, 24/02/1962; 77390 (1) Stn C925, 41.00 S, 173.91 E, 39 m, 10/02/1963; 77403 (1) Stn G193, 43.78 S, 179.5 W, 366 m, 18/01/1968; 77405 (1) Stn B629, 42.66 S, 170.98 E, 20–21 m, 20/10/1962; 77408 (1) Stn B624, 43.97 S, 168.63 E, 13 m, 19/10/1962; 77415 (2) Stn B525, 42.29 S, 173.91 E, 77 m, 13/02/1962; 77417 (1) Stn N926, 41.54 S, 175.02 E, 100 m, 28/08/1977; 68130 (1) Stn S121, 43.50 S, 175.96 E, 335 m, 20/10/1979.

Description. See O’Loughlin et al. (2011)

Distribution. New Zealand: Chatham Rise, Kermadec Ridge 2–377 m depth (this study)

Remarks. Further to the observations of O’Loughlin et al. (2011) we confirm that Paracaudina coriacea (Hutton, 1872) is common on the New Zealand coast and has been previously identified as Paracaudina chilensis (Müller, 1850) . Extensive tissue sampling of the mid-body and tail ossicles of all specimens at NIWA has produced a clear distinction between the two species.

O’Loughlin et al. (2011) discusses the type status of Paracaudina coriacea and the fact that no type material had been located. Subsequent to that paper, three specimens labelled Caudina coriacea (Hutton) and ‘syntype’ from New Brighton, New Zealand were located in the British Natural History Museum (NHMUK) and borrowed for this work. We confirm these specimens as P. coriacea and establish here the three specimens in the New Brighton lot lectotype and paralectotypes and illustrate ossicles from the lectotype.

Under light microscopy the lectotype fits clearly into the type description with the distinct thick, smooth midbody octagonal closed cup ossicles with little space within the ossicle. However, the SEM ossicle photos indicate some similarities between Paracaudina chilensis and Parcaudina coriacea not clear in the light microscopy. The type ossicles are still thicker and smoother than the P. c h i l e n s i s angular open spaced ossicles, but blunt surface and marginal projections (not as obvious in light microscopy) have become visible. This type material clearly justifies the historic debate around the recognition of two species. We judge that the SEM ossicle figures for P. coriacea and P. chilensis in O’Loughlin et al. (2011) and this paper validate the distinction between the two species.

O’Loughlin et al. (2011) noted that Heding (1933) illustrated ossicles from a specimen from New Brighton as P. c o r i a c e a. While Heding does not refer to the material as type, his ossicle illustrations concur with our observations so we believe they are the same specimens.

Théel (1886) discussed two specimens of Molpadia coriacea (Hutton, 1872) in the Stockholm Museum and noted that he considered them close to or a synonym of Paracaudina ransonnetti Marenzeller, 1881 . The issue of the status of P. ransonnetti being a junior synonym of Paracaudina chilensis has been extensively discussed in O’Loughlin et al. (2011) and we judge that the specimens in Stockholm must be P. c h il en s is.

Figures

FIGURE 1. Photos of preserved specimens of species in the family Caudinidae Heding, 1931: a, Hedingia glabra (Théel, 1886) (NIWA 64078); b, Paracaudina alta sp. nov. (NIWA 68129); c, Paracaudina chilensis (Müller, 1850) (NIWA 77411); d, Paracaudina coriacea (Hutton, 1872) (NIWA 70954); e, Paracaudina coriacea (Hutton, 1872) lectotype (NHMUK 98.10.8.1.2); f, Paracaudina reductia sp. nov. (NIWA 70957).