DNA barcoding of some Pandeidae species (Cnidaria, Hydrozoa, Anthoathecata) Author Schuchert, Peter Muséum d'histoire naturelle, CP 6434, CH- 1211 Genève 6, Switzerland, E-mail: peter. schuchert @ ville-ge. ch text Revue suisse de Zoologie 2018 2018-03-31 125 1 101 127 journal article 3917 10.5281/zenodo.1196029 f34e9863-9f62-4f2a-8819-7e6e8fc09403 0035-418 1196029 Neoturris pileata ( Forsskål, 1775 ) Figs 1-7 Medusa pileata Forsskål, 1775: 110 . – Forsskål, 1776 : pl. 33, fig. D. Oceania Lesueurii Péron & Lesueur, 1810: 345 . – Goy, 1995: 244 , plate. Carybdea pisifera Oken, 1815: 125 . Oceania pileus de Blainville, 1830: 258 . Oceania ampullacea M. Sars, 1835: 22 , pl. 4 fig. 8. – Haeckel, 1879: 58 , synonym. Tiaria papalis Lesson, 1843: 287 . – Haeckel, 1879: 58 , synonym. Turris digitale Forbes, 1846: 286 . – Hartlaub, 1914: 324 , synonym. Turris digitalis . – Forbes, 1848: 21 , pl. 3 fig. 1. – Haeckel, 1879: 61 , pl. 4 figs 2-3. ‒ Kramp, 1955: 153 , revision of Haeckel’s material. Oceania episcopalis Forbes, 1848: 27 , pl. 2 fig. 1. – Haeckel, 1879: 58 , synonym. Oceania coccinea Leuckart, 1856: 20 , pl. 2 fig. 3.– Haeckel, 1879: 58 , synonym. Oceania constricta Patterson, 1859: 279 , figs. Tiara pileata . – Haeckel, 1879: 58 , pl. 3 figs 6-8. Turris coeca Hartlaub, 1892: 19 , fig. 1. – Hartlaub, 1914: 329 , synonym. in part Turris pileata . – Mayer, 1910: 123 , pl. 12 fig. 4, pl. 13 fig. 6. Tiara pileata . – Le Danois, 1914: 17 , fig. 4. Perigonimus abyssi G.O. Sars, 1874: 126 , pl. 5 figs 27-30. new synonym Neoturris pileata . – Hartlaub, 1914: 326 , figs 270, 273, 274- 281. – Kramp, 1926: 92 , fig. 37, pl. 2 figs 13-14, chart XVIII. – Russell, 1953: 203 , figs 104-106, pl. 12 fig. 1. – Edwards, 1965: 461 , figs 1-4, life cycle. ‒ Schuchert, 2007: 333 , figs 59-60, review. in part Leuckartiara brevicornis . – Hartlaub, 1914: 304 , figs 254-256. [incorrect subsequent spelling] in part Leuckartiara breviconis . – Kramp, 1926: 80 , pl. 2 fig. 8. ‒ Russell, 1953: 198 , pl. 12 fig. 2. – Kramp, 1959: 120 , fig. 121. [not Neoturris breviconis ( Murbach & Shaerer, 1902 ) ] Leuckartiara breviconis . ‒ Kramp & Damas, 1925: 280 [not Neoturris breviconis ( Murbach & Shaerer, 1902 ) ] Leuckartiara abyssi . – Rees, 1938: 19 , fig. 6a-d, part of life cycle. – Rees, 1956: 114 , re-examination of type material, lectotype designation. – Schuchert, 2007: 330 , fig. 57, redescription, status. Type locality: Mediterranean Material of N. abyssi : All specimens came from Bergen area in Norway . See also Table 1 for GenBank numbers. If no museum accession number is given, there is no material in a permanent collection. Hydroid stage : MHNG-INVE-54693; without gonophores on Nucula spec ; Herdlafjord, 60.503° 5.2152°, 375-440 m depth; collection date 20.04.2007 . ‒ MHNG-INVE-54695; without gonophores on Nucula spec. ; Hauglandsosen, 60.433°5.1167°, 180 m depth; collection date 15.08.2007 . ‒ Hydroid without gonophores on scaphopod of about 5 mm size; Raunefjord, Vatlestraumen, 60.33802° 5.18163°, 32-42 m depth; collection date 16.09.2008 ; DNA isolate 935. ‒ Hydroid without gonophores on Nucula spec. ; Raunefjord, Vatlestraumen, 60.338017° 5.181633°, 32-42 m depth, temperature °C; collection date 16.09.2008 ; DNA isolate 936. ‒ Hydroid without gonophores on sipuncule in Antalis entalis ; Raunefjord, Flesland, 60.30282° 5.2016°, 45-100 m depth; collection date 19.09.2008 ; DNA isolate 694. ‒ Hydroid without gonophores on Nucula spec ; Hordaland , Hauglandosen, 60.435° 5.122°, 135-151 m depth; collection date 19.09.2008 ; DNA isolate 695. ‒ Hydroid without gonophores on Nucula spec ; Hordaland , Hauglandosen, 60.435° 5.122°, 135-151 m depth; collection date 19.09.2008 ; DNA isolate 704. Fig. 1. Neoturris abyssi , original illustration of Sars (1874) . Fig. 2. Neoturris abyssi (= Neoturris pileata ), preserved polyp specimens from Norway, Bergen area, Hauglandsosen. (A) MHNG-INVE-54695 on Nucula spec. (B) MHNG-INVE-62572, on a scaphopod, (DNA 695, see Table 1). Medusa stage : Raunefjord, 60.275° 5.200°, 10 m depth; collection date 22.05.2012 ; DNA isolate 953. ‒ MHNG-INVE-82129; Korsfjord, 60.20833° 5.20261°, 0-20 m depth; collection date 23.05.2012 ; DNA isolate 916. ‒ Korsfjord, 60.20833° 5.20261°, 0-20 m depth; collection date 23.05.2012 ; DNA isolate 917. ‒ Korsfjord, 60.20833° 5.20261°, 0-20 m depth; collection date 23.05.2012 ; DNA isolate 918. ‒ Korsfjord, 60.20833° 5.20261°, 0-20 m depth; collection date 23.05.2012 ; DNA isolate 919. ‒ Korsfjord, 60.20833° 5.20261°, 0-20 m depth; collection date 23.05.2012 ; DNA isolate 954. ‒ Fanafjord, 60.24079° 5.22941°, 0-20 m depth; collection date 24.04.2015 ; DNA isolate 1119. Material of N. pileata : MHNG-INVE-97957; France , Bay of Villefranche-sur-Mer, 43.685° 7.315667°, 0-30 m depth; collection date 11.04.2017 ; DNA isolate 1280. Additional examined material is given in Schuchert (2007) . Diagnosis: Neoturris medusa with bell that is usually higher than wide, height 2-4 cm , no exumbrellar nematocyst ridges, with or without apical projection, no apical canal, with up to 60-90 tentacles. Manubrium usually longer than half the subumbrella height, interradial gonad region large and without folds but with many gonadal pits (>20 per quadrant), eight adradial rows of horizontal gonads folds, folds appear directed towards interradii; no papillae on gonads, radial canals jagged, tentacle bases without abaxial spurs, no ocelli. Colours depending on age and environment, manubrium in younger ones yellow-orange, in fully grown medusae pink to ruby-red; tentacle-bases yellowish. Hydroids usually on scaphopods and Nucula shells, colonial, arising from creeping stolons; hydrocauli covered by perisarc, not branched, monosiphonic. Perisarc extends onto hydranth body as a more or less gelatinous pseudohydrotheca which does not envelop the tentacles. Hydranths with conical hypostome, one whorl of filiform tentacles. Gonophores develop on cauli or stolons, enclosed in thin perisarc membrane. Gonophores liberated as free medusae with four tentacles. Fig. 3. Neoturris abyssi (= Neoturris pileata ) medusae from Norway, photographs of living, relatively young stages. (A) Lateral view of medusa with bell size 7 mm (DNA 953, see Table 1). (B) Same as A, detail of radial canals and folds of stomach wall. (C) Same as A, oblique view from below. (D) Animal of bell size 9 mm, detail of tentacle bases, note the absence of ocelli. Description: See Schuchert (2007) . Remarks: As already suspected by Edwards (1965) , the 16S and COI sequence comparisons presented above are evidence that the hydroid Leuckartiara abyssi G.O. Sars, 1874 must belong to Neoturris pileata ( Forsskål, 1775 ) . The hydroid of L. abyssi from near the original collecting site of Sars belongs unambiguously to Neoturris medusae found at the same locality. These Neoturris medusae were smaller than those of adult Mediterranean ones (largest ones seen about 15 mm high), but the morphology of the manubrium with its numerous interradial pits and the adradial folds ( Fig. 4B ) comes close to the ones in more southern waters (comp. Figs 6-7 ). The colour of the manubrium was, however, never as red as found in medusae south of Norway to the Mediterranean. A Neoturris medusa from Sweden ( Fig. 5 ) had a much darker manubrium, despite being not much larger than the Norwegian ones. The yellowish Neoturris medusae occur regularly in the Bergen region (see also Hosia & Båmstedt, 2007 ; as N. pileata ) and must also have been seen by Kramp & Damas (1925) who attributed them to N. breviconis . The sequence comparison made here ( Figs 8-9 ), however, show that this cannot be the case as N. breviconis is well separated from the N. abyssi + N. pileata clade. Fig. 4. Neoturris abyssi (= Neoturris pileata ) medusae from Norway, photographs of the most advanced stages found. (A) Lateral view of manubrium of a medusa with bell size 9 mm (DNA 918, see Table 1). No gametes could be seen when examining the gonad fold under a compound microscope. (B) Manubrium of a medusa with bell size 12 mm (DNA 916, see Table 1); note the increased number of gonadal folds and pits. Small oocytes were present in the gonads folds. Except for the colour this animal closely resemble Mediterranean specimens (Figs 6-7). (C) Medusa of about 10 mm height (DNA 919, see Table 1) cut open and spread to visualise anatomical details (inner side of stomach facing observer). Fig. 6. Neoturris pileata , preserved specimen (MHNG-INVE-35522) from the Mediterranean, collected before 1895 and identified by C. Hartlaub. Note that the bell shape is not elongated as often seen in other illustrations (e.g. Fig. 7), but nevertheless lies within the range of variation for Mediterranean specimens. Moreover, the bell is somewhat flattened in this preserved sample. Fig. 5. Neoturris abyssi (= Neoturris pileata ), living medusa from the Swedish coast, photo taken by Fredrik Pleijel and reproduced with the permission of the author. The manubrium is contracted, feigning a horizontal gonadal fold on the manubrium resembling the permanent one seen in some Leuckartiara species. This photo is copyright protected and it must not be reproduced without the consent of the author. The hydroid of N. pileata without medusa buds is not readily distinguishable from Leuckartiara octona , the only other pandeid known from the region ( Hosia & Båmstedt, 2007 ). The only character to reliably distinguish the two is found in the newly released medusae, which have four tentacles instead of the two tentacles present in L. octona . A less reliable character is the absence of branching of the stems, which in fully grown colonies of L. octona are quite regularly branched once, but not so in N. abyssi . The Norwegian hydroids here assigned to L. abyssi lacked medusa buds, but were nevertheless assigned to L. abyssi because they came from close to the type locality, they grew on the typical substrate, the pedicels were never branched, and they occurred in relatively deep waters. Their sequences separated them immediately from L. octona medusae collected at the same locality ( Figs 8-9 ). An infertile pandeid hydroid on a Nucula shell collected in 5-50 m depths along the Swedish coast (DNA 1055, Table 1 ) was initially also identified as L. abyssi , but the DNA data clearly identified it as L. octona and it was reclassified accordingly.