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Dendronotus zakuro sp. nov. (Figures 1, 3, 5B) http://zoobank.org/ 3B54EB14-A55E-42C5-B962-82D6A40B9BF5 Type material. Holotype, ZMMU Op-700, 30 mm long live, 15 mm preserved, dissected, Russia, Kamchatka, Starichkov Island, 17.07.2015, depth— 12 m, on stones, collector N.P. Sanamyan. Two paratypes, KSNHM-OP0485, Japan, Hokkaido, Usujiri, 10 and 12 mm long (preserved), 13.03.2016, depth 10–20 m, stones, collector Sho Kashio One paratype, ZMMU Op-664, 25 mm long live, Japan, Honshu, Echizen Coast, Fukui Prefecture, 19.04.2018, depth 7.1 m, collector Chihiro Dairi. Type locality. Russia (Kamchatka) and Japan (Hokkaido; Honshu: Sea of Japan). Etymology. From zakuro (ẈŽo, HAE×ďAE), meaning in Japanese “pomegranate”, after a striking red to red-brownish colouration of the new species. Diagnosis. Body elongate, 6–7 pairs dorsolateral appendages, colour bright reddish to reddish-brownish with opaque white spots, central tooth completely smooth (in adults), lacking denticles and furrows, vas deferens moderate in length, penis massive, long, twisted. Description. Body elongate, up to 30 mm in length (Fig. 3 A–E), 5–7 branched appendages of oral veil, ca. 5 appendages of rhinophoral stalks, 10–11 rhinophoral lamellae, branched rhinophoral lateral papilla present, 6–7 pairs dorsolateral appendages, 25–30 lip papillae. Dorsolateral appendages with elongate primary stalk, moderately relatively highly branched secondary branches, and attenuated tertiary branches (Fig. 3 A–E). Reproductive and anal openings placed laterally on right side. General colour bright red to reddish-brownish with thin white broken lines between dorsolateral appendages, also scattered opaque white dots and speckles on dorsal and lateral sides, and on various appendages (Fig. 3 A, D, E). The jaws are ovoid with strong dorsal processes, denticles present (Fig. 3 F–G). Masticatory processes bear up to 120 denticles. Radula formula is 36 × 3–11.1.11–3 (holotype), 34 × 3–12.1.12–3 (paratype, 12 mm). Central tooth completely devoid of denticles and furrows in most parts of the radula (except for the anterior-most juvenile radula in some specimens) only faint traces of the furrows on the surface appear on some teeth in the holotype (Fig. 3H, J); the studied paratype possesses almost completely smooth central teeth throughout the whole radula. Lateral teeth are long, distinctly curved, bearing up to 7 long denticles (Fig. 3I, K). Reproductive system triaulic (Fig. 5B), ampulla thin, twice folded (Fig. 5B, am), prostate consisting of ca. 25 alveolar glands (Fig. 5B, pr), vas deferens relatively long (Fig. 5B, vd) expanding to voluminous penial sheath (Fig. 5B, psh), penis long and twisted (Fig. 5B, p), vagina long and straight (Fig. 5B, vg), bursa copulatrix is large, rounded, and elongated (Fig. 5B, bc) with small seminal receptaculum placed distally (Fig. 5B, rs). Biology. Inhabits stony and rocky substrates, 7– 20 m. Distribution. Presently known from three remote locations in the North Western Pacific, from Hokkaido Island and Echizen coast (Honshu) in Japan and Starichkov Island, Kamchatka in Russia. Further intermediate findings on the Kurile Islands between these two remote points are therefore expected. Remarks. D. zakuro sp. nov. by combination of colouration, radular patterns and molecular phylogenetic data well differs from all previously described species of the genus Dendronotus. The almost smooth central teeth in adult radular morphology combined with the non-uniform, partly variegated external colouration of D. zakuro sp. nov. is only similar to D. kamchaticus. However, D. kamchaticus is phylogenetically distantly related to D. zakuro sp. nov. (Fig. 1). Furthermore, morphologically D. zakuro sp. nov. differs from D. kamchaticus by bright red to redbrownish colouration (the known range of colouration for D. kamchaticus is brownish to transparent greyish, see Korshunova et al., 2016a), larger number of rows of lateral teeth and patterns of their denticulations. D. subramosus with commonly brownish general colouration considerably differs from D. zakuro sp. nov. by the absence of lateral rhinophoral papillae and strongly denticulated central radular teeth. According to molecular phylogenetic analysis, evolutionary ties for D. zakuro sp. nov. are not fully clarified. Clade D. zakuro sp. nov. has the closest position to the clades D. arcticus, D. dalli, D. kamchaticus, D. niveus, D. lacteus, D. europaeus and D. rufus (Fig. 1), but none of these species (except D. kamchaticus) demonstrate any external or internal similarity to D. zakuro sp. nov. Maximum intraspecific and minimum interspecific genetic distances for the COI marker in the species of the genus Dendronotus including D. zakuro sp. nov. are presented in the Table 2.

Dendronotus bathyvela sp. nov. (Figures 4, 5C) http://zoobank.org/ 28A36F6D-C221-4C34-B40A-631E3129B943 Type material. Holotype, NSMT-Mo 94455, 45 mm long preserved, dissected, Japan, Tohoku, off Ofunato, Iwate Prefecture, 38° 53.5’ N 142° 2.8’ E, 18.10.2007, depth 303–307 m, collector K. Hasegawa, H. Komatsu. Three paratypes, NSMT-Mo 94450, 25, 26, 27 mm long (preserved), Japan, Tohoku, off Shimokita Peninsula, Aomori Prefecture, 41° 0.5’ N 141° 2.0’ E, 10.10.2006, depth 511– 510 m, collector T. Kuramochi, T. Fujita. Paratype, NSMT- Mo 94452, 35 mm long (preserved), Japan, Tohoku, off Kinkazan, Miyagi Prefecture, 38° 23.2’ N 141° 58.2’ E, 04.11.2006, depth 305–309 m, collector H. Komatsu. Paratype, NSMT-Mo 93078, 35 mm long (preserved), Japan, Tohoku, off Kesennuma, Miyagi Prefecture, 38° 4.6’ N 141° 55.3’ E, 19.11.2005, depth 249 m, collector T. Fujita, H. Saito. Paratype, NSMT-Mo 93080, 21 mm long (preserved), Japan, Tohoku, off Kesennuma, Miyagi Prefecture, 38° 4.2’ N 141° 58.9’ E, 19.11.2005, depth 306–309 m, collector T. Fujita, H. Saito. Paratype, NSMT-Mo 93061, 14 mm long (preserved), Japan, Tohoku, off Kinkazan, Miyagi Prefecture, 38° 2.3’ N 142° 2.1’ E, 17.11.2005, depth 382– 376 m, collector T. Fujita, H. Saito. Type locality. Japan, off the Pacific coast of Northern Honshu. Etymology. From Ancient Greek βάθος (deep sea) + velum (veil) in reference to the deepest record so far known of this wide-bodied Dendronotus species with a broad oral veil. Diagnosis. Body broad, 6–7 pairs dorsolateral appendages, colour dull reddish-brownish with numerous white spots, central tooth with well-defined denticles and furrows, vas deferens moderate in length, penis long, very thin. Description. Body broad, up to 45 mm in length (Fig. 4 A–D), 10–15 long branched appendages of oral veil, 5 appendages of rhinophoral stalks, 15–25 rhinophoral lamellae, branched rhinophoral lateral papilla absent, 6–7 pairs dorsolateral appendages (including smallest posterior ones), ca. 50–70 lip papillae. Dorsolateral appendages with moderate primary stalk, moderately branched secondary branches, and elongated tertiary branches (Fig. 4 A–D). Reproductive and anal openings placed laterally on right side. General colour dull reddish-brownish with numerous distinct opaque white dots on notum, tips of lateral appendages, oral appendages (Fig. 4 D). The jaws are ovoid with strong dorsal processes, denticles present (Fig. 4 E–F). Masticatory processes apparently bear ca 60 denticles. Radula formula is 36 × 1–13.1.13–1 (paratype 21 mm), 41 × 1–12.1.12–1 (paratype 27 mm), 37 × 1–13.1.13–1 (paratype 35 mm), 36 × 1–14.1.14–1 (holotype 45 mm). Central tooth with broad, relatively low cusp, strongly denticulated and bearing up to over 30–35 distinct to small denticles (Fig. 4G). Lateral teeth are long, slightly curved, commonly completely smooth or bearing few (up to 2) weak denticles (Fig. 4H). Reproductive system triaulic (Fig. 5C), ampulla folded several times (Fig. 5C, am), prostate consisting of 21–25 alveolar glands (Fig. 5C, pr), vas deferens long (Fig. 5C, vd) expanding to elongate penial sheath (Fig. 5C, psh), penis very long, thin (Fig. 5C, p), vagina very long and considerably twisted (Fig. 5C, vg), bursa copulatrix is large, rounded, and elongated (Fig. 5C, bc) with small seminal receptaculum placed distally (Fig. 5C, rs). Biology. Inhabits sand and muddy substrates with stones. Distribution. Presently known only off the Pacific coast of Northern Honshu (Japan), at depths of 249– 510 m. Remarks. According to the morphological data Dendronotus bathyvela sp. nov. clearly belongs to the group of wide-bodied Dendronotus species, which includes only a few species, i.e. D. patricki, D. robustus and D. velifer. Because all available D. bathyvela sp. nov. were formalin-fixed, this prevented us from including this species in the molecular phylogenetic analysis. However, the present morphological data are enough to distinguish D. bathyvela sp. nov. from all other wide-bodied species of the genus Dendronotus. The abyssal NE Pacific species, D. patricki, readily differs from the bathyal NW Pacific D. bathyvela sp. nov. by its uniform pinkish colouration (white pig- ment presents only on apices of dorsolateral and velar appendages, but not on the body) and details of the radula (Stout et al., 2011). Particularly, in D. patricki the number of lateral teeth is up to eight, whereas in D. bathyvela sp. nov. —up to 14. The number of lateral denticles on the central teeth is about 20 in D. patricki, whereas in D. bathyvela sp. nov. there are over 30 denticles. Previously, D. robustus had been recorded from the Sea of Japan and the Pacific side of Honshu (Roginskaya 1997; Hasegawa 2009). However, the true D. robustus from the shallow waters of the North Atlantic (see Lundin et al. 2017) considerably differs from the North Pacific material in the details of colouration and the radula. For a long time, D. velifer from Northern Europe and the Arctic was also confused with D. robustus, however recently it was shown that this is a distinct species, according to both morphological and molecular data (Lundin et al. 2017). The following differences between D. velifer and D. bathyvela sp. nov. are: 1) Appendages of the oral veil in living specimens of D. velifer are distinctly shorter than in living D. bathyvela sp. nov. (Fig. 4 A–D); 2) Large specimens of D. velifer commonly bear not more than four main dorsolateral appendages (an exceptionally large specimen was reported as having up to five main appendages plus a sixth smaller one), whereas even moderately sized D. bathyvela sp. nov. possess up to six main dorsolateral appendages (plus a seventh smaller one); 3) Colouration of living specimens of D. velifer is bright reddish, whereas D. bathyvela sp. nov. is dull reddish-brownish (Fig. 4D); 4) Number of denticles on the central teeth of D. velifer is commonly less than 30, whereas in D. bathyvela sp. nov. it reaches over 30; 5) Number of radular rows in D. velifer is up to 36, whereas in D. bathyvela sp. nov. the number of rows is up to 41 and more. It is remarkable that the North Atlantic shallowwater species D. robustus has up to seven dorsolateral appendages, like the NW Pacific shelf to bathyal new species D. bathyvela sp. nov., but not like the true predominantly Arctic shelf species D. velifer, which commonly has only four dorsolateral appendages. This feature readily distinguishes D. bathyvela sp. nov. from D. velifer. At the same time, D. bathyvela sp. nov. is well-distinguished from D. robustus by the shape of the central radula and external colouration. Bathymetrically D. velifer is known from relatively shallow waters with a range commonly ca. 15–230 m, whereas D. bathyvela sp. nov. inhabits the low part of the continental shelf (not shallower than 249 m) to upper bathyal (510 m) depths. True North Atlantic D. robustus is in turn exclusively a shallow water species and was not recorded deeper than 20–30 m (see Lundin et al. 2017). This set of several complex morphological and ecological features allow us to distinguish D. bathyvela sp. nov. from D. velifer. Recently a deep-sea (but not wide-bodied) species, D. claguei, was described (Valdés et al. 2018) off the NE Pacific coast. D. claguei fundamentally differs from D. bathyvela sp. nov. by its uniform translucent white colour, elongate body shape, radular details, and in addition D. claguei inhabits upper abyssal depths (2369 m) and not shelf to upper bathyal depths (249–510 m) as D. bathyvela sp. nov. does. In addition, the NE Pacific shallow-water species D. albopunctatus Robilliard, 1972 is somewhat similar to the wide-bodied species group of the genus Dendronotus, but has fewer, and longer, oral veil appendages, lacks lip papilla and considerably differs by the presence of distinct denticulation on the lateral teeth of the radula (Robilliard, 1972).

Dendronotus jamsteci sp. nov. (Figures 1–2, 5A) http://zoobank.org/ B8DAC5AC-56F4-4C92-90EC-D48C5173772A Type material. Holotype, JAMSTEC No. 1160047475, 17.5 mm long preserved, dissected, Cruise KS-16-J03, on board ID HPD #1952, Japan, Northern Honshu, off Otsuchi, 39° 18.5378’ N 142° 17.82’ E, 08 March 2016, depth— 670 m, collector Shinji Tsuchida. One paratype, JAMSTEC No. 1160047461, Cruise KS-16-J03, on board ID HPD #1952-07, Dive No. HPD #1952, 21.5 mm long (preserved), Japan, Northern Honshu, off Otsuchi, 39° 18.4453’ N 142° 18.0498’ E, 08 March 2016, depth— 761 m, collector Shinji Tsuchida. Five paratypes, JAM- STEC No. 1160047464, Cruise KS-16-J03, on board ID HPD #1952-10, Dive No. HPD #1952, 2– 7 mm long (preserved), Japan, Northern Honshu, off Otsuchi, 39° 18.4453’ N 142° 18.0498’ E, 08 March 2016, depth— 761 m, collector Shinji Tsuchida. Ca. 60 paratypes, JAMSTEC No. 1160047463, Cruise KS-16-J03, on board ID HPD #1952-09, Dive No. HPD #1952, 1.5–15 mm long (preserved), Japan, Northern Honshu, off Otsuchi, 39°18.4453’ N 142°18.0498’ E, 08 March 2016, depth— 761 m, collector Shinji Tsuchida. Three paratypes, JAMSTEC No. 1160047392, Cruise KS-16-J03, on board ID HPD #1950-16, Dive No. HPD #1950, 11– 11.5 mm long (preserved), Japan, Northern Honshu, off Otsuchi, 39° 18.7059’ N 142° 18.0498’ E, 05 March 2016, depth— 684 m, collector Shinji Tsuchida. Nine paratypes, JAMSTEC No. 1160047475, Cruise KS-16-J03, on board ID HPD #1952, 2 – 21 mm long, Japan, Northern Honshu, off Otsuchi, 39° 18.5378’ N 142° 17.82’ E, 08 March 2016, depth— 670 m, collector Shinji Tsuchida. Three paratypes, JAMSTEC No. 1160047380, Cruise KS-16-J03, on board ID HPD #1950- 4, 7– 18 mm long (preserved), Japan, Northern Honshu, off Otsuchi, 39° 18.6579’ N 142° 18.0498’ E, 05 March 2016, depth— 704 m, collector Shinji Tsuchida. ......continued on the next page Type locality. Japan, off the Pacific coast of Northern Honshu. Etymology. In honour of the Japan Agency for Marine-Earth Science and Technology (JAMSTEC) which has made a considerable contribution to the understanding of deep-sea fauna in Japan and worldwide. Diagnosis. Body elongate, 6–7 pairs dorsolateral appendages, colour whitish with brownish spots and streaks, central tooth with well-defined denticles and furrows, vas deferens moderate in length, penis conical, moderately narrow. Description. Body elongate, up to 21.5 mm in length (Fig. 2A, B), 6–9 branched appendages of oral veil, 4–5 appendages of rhinophoral stalks, 12–15 rhinophoral lamellae, branched rhinophoral lateral papilla present, 6–7 pairs dorsolateral appendages, 20–25 lip papillae. Dorsolateral appendages with thickened primary stalk, moderately branched secondary branches, and blunted tertiary branches (Fig. 2A, C). Reproductive and anal openings placed laterally on right side. General colour whitish with scattered distinct opaque white dots on notum, tips of lateral appendages, oral appendages, lip papillae, and rhinophores (Fig. 2C). The jaws are ovoid with strong dorsal processes, denticles present. Masticatory processes bear ca. 60 denticles (holotype, including simple triangular and broadly lamellated ones (Fig. 2D, E). Radula formula is 33 × 1–6.1.6–1 (paratype 7 mm), 34 × 3–8.1.8–3 (holotype), 38 × 3–8.1.8–3 (paratype 20 mm), 29 × 3–8.1.8–3 (paratype 21.5 mm). Central tooth strongly denticulated and bearing up to 17 distinct denticles (Fig. 2F) with well-defined furrows. Lateral teeth are short, slightly curved, bearing up to 9 long denticles (Fig. 2G). Reproductive system triaulic (Fig. 5A), ampulla moderately thin, triple folded (Fig 5A, am), prostate consisting of 50–60 alveolar glands (Fig. 5A, pr), vas deferens moderate in length (Fig. 5A, vd) expanding to a narrow penial sheath (Fig. 5A, psh), penis short and conical (Fig. 5A, p), vagina long and twisted (Fig. 5A, vg), bursa copulatrix is medium-sized, rounded (Fig. 5A, bc) with small seminal receptaculum placed distally (Fig. 5A, rs). Biology. On hydroids within sand and muddy substrates. Distribution. Presently known only off the Pacific coast of Northern Honshu (Japan), at depths of 670– 761 m. Remarks. Dendronotus jamsteci sp. nov. is superficially similar to D. frondosus and D. primorjensis, but different in the details of 1) colouration: compared to D. frondosus and D. primorjensis, D. jamsteci sp. nov. has a weakly coloured whitish ground colour with dispersed brownish spots), 2) radula: D. jamsteci sp. nov. has different number of rows (up to 8 in D. jamsteci sp. nov., up to 9 in D. primorjensis, up to 10 in D. frondosus) and denticles on the central (up to 17 denticles in D. jamsteci sp. nov., up to 18 in D. primorjensis, up to 14 in D. frondosus) and lateral tooth (up to 9 denticles in D. jamsteci sp. nov., up to 8 in D. primorjensis, up to 7 in D. frondosus), and 3) reproductive system: D. jamsteci sp. nov. considerably differs in the number of alveolar glands (50–60 in D. jamsteci sp. nov., 12–19 in D. primorjensis, 16–30 in D. frondosus) as well as details in the patterns of the bursa and copulative organ arrangements. Importantly, according to the molecular phylogenetic data (Fig. 1), D. jamsteci sp. nov. doesn’t belong to the clade comprised of D. frondosus, D. primorjensis and D. venustus (this species readily differs externally from D. jamsteci sp. nov. by its bright yellow markings or large amounts of opaque white pigment) (MacFarland 1966; Stout et al. 2010) but is sister to the clade that includes D. subramosus and D. albus. However, both the latter species are considerably different morphologically from D. jamsteci sp. nov.. Particularly, although D. subramosus externally shares a similar brownish colouration with D. jamsteci sp. nov. it lacks lateral rhinophoral papilla common in most species of the genus Dendronotus including D. jamsteci sp. nov. By this feature, it readily differ from D. jamsteci sp. nov., whereas D. albus considerably differs from D. jamsteci sp. nov. by its uniform whitish or lilac colour and patterns of the radula and reproductive system. Such a discrepancy of molecular and morphological evolution is very interesting. From the NW Pacific species D. kalikal, D. jamsteci sp. nov. differs by the absence of subparallel brownish dorsal stripes, details of the radula and very significantly by the molecular data (Fig. 1). Two other NW Pacific species, D. kamchaticus and D. zakuro sp. nov. , are substantially different from D. jamsteci sp. nov. by their colouration and the smooth central teeth of their radula at the adult stage and in addition, they belong to a different molecular clade (Fig. 1). None of these species inhabit bathyal depths as D. jamsteci sp. nov. does, and their bathymetric distribution is restricted to shallow waters, not deeper than 60 m. Recently, a deep-sea species off the NE Pacific coast, D. claguei, was described (Valdés et al. 2018). For that species, only H3 gene data was provided in the cited publication, so the absence of COI and 16S data prevented its use here for molecular analysis because the H3 gene is almost identical for closely related species of the genus Dendronotus. D. claguei fundamentally differs from D. jamsteci sp. nov. by its uniform translucent white colour, radular details, and in addition D. claguei inhabits upper abyssal depths (2369 m) and not upper bathyal depths (670–761 m) as D. jamsteci sp. nov. does. Maximum intraspecific and minimum interspecific genetic distances for the COI marker in the species of the genus Dendronotus including D. jamsteci sp. nov. are presented in Table 2.

Martynov, Alexander, Fujiwara, Yoshihiro, Tsuchida, Shinji, Nakano, Rie, Sanamyan, Nadezhda, Sanamyan, Karen, Fletcher, Karin, Korshunova, Tatiana (2020): Three new species of the genus Dendronotus from Japan and Russia (Mollusca, Nudibranchia). Zootaxa 4747 (3): 495-513, DOI: https://doi.org/10.11646/zootaxa.4747.3.4

Onchidoris macropompa sp. nov. (Figures 6 I, J; 7 D; 13; 14 A���F; 15 A, B. Table 3) Onchidoris sp.: Martynov, 1997: 235 Material. Holotype, ZMMU Lc- 37463, NW Pacific near Kamchatka peninsula, Starichkov Id., 6���7 m depth, collected by T.A. Korshunova and A.V. Martynov, 25.07. 2008. Paratypes, Lc- 37464, two specimens (one dissected), same locality and collectors as holotype, 25.07. 2008. Paratype, ZMMU Lc- 37465, one specimen (dissected), 21���26 m depth, same locality and collectors as holotype, 14.08. 2008. Paratypes, ZMMU Lc- 37466, two specimens (dissected), 12���15 m depth, same locality and collectors as holotype, 19.08.2008; Paratype, ZMMU Lc- 37467, one specimen (dissected), Commander Ids., Medny Id., Cape Drovyanye Stolby, sample 101, 9��� 12 m depth, collector Oshurkov V.V., 17.07. 1992. Paratype, ZMMU Lc- 37468, one specimen (dissected), Commander Islands, Beringa Id., Ariy Kamen Id., sample 225, 25 m depth, rock, collector V.I. Shalukhanov, 0 2.08. 1991. Paratypes, Lc- 37469, two specimens (one dissected), Commander Islands, Mednyi Id., Kekur Korabelny Stolb Id., 18���19 m depth, rock, collector V.I. Shalukhanov, 11.07. 1992. Type locality. NW Pacific, SE Kamchatka, Starichkov Id., 6���26 m depth. Etymology. From macros (Greek, = large) and pompa (Russian, Italian, = pump) refers to the largest relative size of the buccal pump of the present species within Onchidorididae. Description. External morphology. The length of holotype is 7 mm, the width is 4 mm (Fig. 13 A). The length of 5 living specimens ranged from 6 to 15 mm, the width ranged from 4 to 6.5 mm. The consistency of the living animals is hard. The notum is moderately broad, rounded in front and posteriorly. The rhinophores are long and retract into sheaths with smooth edges, except for 3���4 tubercles of various sizes that are connected with the edges of the sheaths (Fig. 13 F). The rhinophoral sheath edges are capable of some contraction in living specimens. There are 7���13 rhinophoral lamellae. The notum is densely covered with mushroom-shaped or club-shaped tubercles on a short stalk. The larger tubercles dominate all over the notum, but a single irregular zigzag row of smaller tubercles appears in the mid-notal line. Some smaller tubercles are also scattered around various notal areas and at the edge of the notum. The rays of the spicules that extend from the bases of tubercles form a dense network that shines through the notum surface (Figs. 6 I; 13 A, D). Each tubercle contains dense bundles of spicules, characteristically slightly protruding from the tubercle surface. The strongly calcified spicules are of various sizes; most have a narrow channel inside, some are solid (Fig. 6 J). A gill cavity is absent. Inside the gill circlet there are several narrow tubercles of variable height. Ten to twelve unipinnate gills form a semicircle around the anus (Fig. 13 D). The oral veil is semicircular (Figs. 13 B, C). The foot is broad, anteriorly rounded and thickened, and posteriorly not projecting beyond the notum, forming a rounded tail (Fig. 13 A). Colour. The living specimens are off-white, semitransparent. Rhinophores (including lamellae) and gills are similar to the ground colour. Dull white gill glands are placed at the gills base. A postbranchial gland is indicated by a slightly conspicuous flattened area behind the gills. Anatomy. Digestive system. The anterior part of the buccal bulb is modified to a prominent, large, very broad, buccal pump that sits on a short, narrow, conspicuous stalk (Fig. 7 D). The buccal pump is fully banded by the broad peripheral muscle (Fig. 7 D). Lateral sides of the buccal pump are provided with thin muscular fibres. The rounded labial disk is covered by colourless cuticle with indistinct labial elements. The radular formula in five specimens (8���15 mm length) is 35��� 38 x 1.1. 1.1. 1, radular teeth are almost colourless. The central tooth is distinct, relatively large, elongated, rectangular, and folded (Figs. 14 A���C, E). The first lateral tooth is provided with a long, wide base and a strong, almost straight beak-shaped cusp. All teeth are entirely devoid of any denticles (Figs. 14 A���F). The second lateral teeth are rectangular plates, with a downward directed cusp on its lower outside corner (Figs. 14 B���C; E���F). The stomach is relatively small and narrow. The stomach caecum is absent. Circulatory system. In the pericardial sac there is a rather wide triangular posterior auricle and a smaller sized and also triangular ventricle. The massive blood gland forms a single piece that is located above the central nervous system and projects slightly anteriorly and posteriorly. Central nervous system. The cerebral and pleural ganglia are well separated, the latter being somewhat smaller in size. The optic nerve is very short. The eyes are relatively large, with black pigment in all studied specimens. The pedal ganglia are smaller than the cerebrals, lay below them and are connected to them by very short connectives. The rhinophoral ganglia are rather irregular, globular or elongate. The buccal ganglia are slightly oval. Gastro-esophageal ganglia are present. Five pairs of cerebral nerves, two pleural and three pedal ones are detected. Reproductive system. (Figs. 15 A, B). The ampulla is wide and swollen, somewhat kidney-shaped, and filled with sperm (Figs. 15 A, a). A long post-ampullar duct is placed along a shallow groove in between the seminal receptacle and vagina and then bifurcates into a long vas deferens (Fig. 15 A, pr) and the proximal oviduct (Fig. 15 A, pov). The prostatic part of the vas deferens is a relatively long loop adjacent to, but not encircling the bursa copulatrix (Fig. 15 A, pr). The prostate is narrow, not granulated; it narrows and then rapidly widens into a long swollen penial sheath; this forms ca. 2.5 loops, and contains several folds of the ejaculatory duct. (Fig. 15 A, psh). The everted penial sheath and ejaculatory duct (penis) have two short terminal processes, and a third, very long one, that are united into a trifurcate penis (Fig. 15 B, tp). The globular bursa copulatrix (Fig. 15 A, bc) contains a dark brown substance; it enters the proximal part of the vagina via a narrow, relatively long stalk. At its base, a duct, which is similar in diameter to the vagina, exits the vagina and leads to the ovoid seminal receptacle (Fig. 15 A, rs). The vagina is wide and long (Fig. 15 A, v). The small distal oviduct (Fig. 15 A, dov) exits the vagina (Fig. 15 A, v) close to the vaginal opening. Biology. Specimens were found predominantly under small stones covered with several species of encrusting bryozoa, at 6���15 m depth, rarely down to 18���25 m depth. Distribution. Presently known only from the type locality in Kamchatka waters and from Commander Ids. (Martynov 1997, as Onchidoris sp.; present study). Remarks. Onchidoris macropompa sp. nov. is externally most similar to Onchidoris muricata but can be readily distinguished from the latter by the smooth cusp of the first lateral teeth, completely devoid of any traces of denticles and in all studied specimens from different, distantly placed localities (i.e. from Kamchatka and from Commander Islands) (Figs. 14 A���F). In contrast, numerous specimens of Onchidoris muricata from both North Atlantic and North Pacific waters always have revealed the presence of various numbers of cusp denticles (Thompson & Brown 1984; Millen 1985; present study, Figs. 14 G���I). In addition, the cusps of the first lateral teeth of Onchidoris macropompa are straight (Figs. 14 B���D, F), whereas in Onchidoris muricata it is typically distinctly curved (Figs. 14 H, I). In addition, Onchidoris macropompa differs from most other species of the genus Onchidoris by presence of a central tooth in the radula (see Table 3 for comparison). The single other congener with radula formula of 1.1. 1.1. 1 and having smooth first lateral teeth, O. bilamellata, differs from Onchidoris macropompa by its colouration, the shape of the buccal pump, the shape of the first lateral teeth, and the pattern of the reproductive system., Published as part of Martynov, Alexander, Korshunova, Tatiana, Sanamyan, Nadezhda & Sanamyan, Karen, 2009, Description of the first cryptobranch onchidoridid Onchimira cavifera gen. et sp. nov., and of three new species of the genera Adalaria Bergh, 1879 and Onchidoris Blainville, 1816 (Nudibranchia: Onchidorididae) from Kamchatka waters, pp. 1-43 in Zootaxa 2159 on pages 35-37, DOI: 10.5281/zenodo.188931, {"references":["Martynov, A. V. (1997) Opistobranch gastropods at the Commander Islands with remarks on their fauna of the Russian Far Eastern Seas. In: Benthic flora and fauna of the shelf zone of the Commander Islands. 1997. Vladivostok, Dalnauka Press., ed. A. V. Rzhavsky, 230 - 241.","Thompson, T. E. & Brown, G. H. (1984) Biology of opisthobranch molluscs, vol. 2, Ray Society, no. 156., 229 pp.","Millen, S. V. (1985) The nudibranch genera Onchidoris and Diaphorodoris (Mollusca, Opisthobranchia) in the northeastern Pacific. The Veliger 28, 80 - 93."]}

Adalaria slavi sp. nov. (Figures 3 C, G; 6 D���F; 7 B; 9; 11 A���D; 12 A, B. Table 2) Type Material. Holotype, ZMMU Lc- 37456, (23 mm length), NW Pacific near Kamchatka peninsula, Starichkov Id., 20���26 m, collected by T.A. Korshunova and A.V. Martynov, 14.08. 2008. Paratypes, ZMMU Lc-37457, 28 specimens, same locality and collectors as holotype 19.08. 2008. Paratypes, ZMMU Lc- 37458, seven specimens (one dissected) same locality and collectors as holotype, at 18���24 m depth, large boulders, 19.08. 2008. Paratypes, ZMMU Lc- 37459, five specimens (three dissected), same locality and collectors as holotype, 14.08. 2008. Paratypes, ZMMU Lc-37460, 10 specimens, same locality and collectors as holotype, 19.08. 2008. Paratypes, ZMMU Lc-37461, 9 specimens, same locality and collectors as holotype, 19.08. 2008. Paratype, ZMMU Lc- 37462, one dissected specimen, same locality and collectors as holotype, 19.08. 2008. Type locality. NW Pacific, SE Kamchatka, Starichkov Id., 18��� 26 m. Etymology. This species is named in honour of Vyacheslav G. Shipilov, captain of the boat ���Chaika��� in recognition of his generous help in organizing scuba diving. Description. External morphology. The length of the holotype is 23 mm and width 11.5 mm (Fig. 9 A���B). The length of 20 living specimens ranged from 8 to 23 mm, the width ranged from 4.5 to 11.5 mm. The consistency of living animals is rather soft. The notum is moderately broad, rounded in front and posteriorly. The rhinophores are long and retracted into sheaths with smooth edges, except for 5���6 tubercles of various size that are connected with edge of each sheath (Fig. 9 D). The rhinophoral sheath edges are capable of some contraction in living specimens. There are 11���14 rhinophoral lamellae. The clavus of the rhinophore has a low ridge posteriorly. The notum is densely covered with inflated cylindrical or almost globular tubercles on a short stalk. Tubercles in the central notal area are somewhat wider and more globular than those at the notal edge. Larger tubercles are regularly intermingled with smaller ones. The rays of spicules radiating from the bases of tubercles form a sort of network under the surface of the apparently soft notum (Figs. 6 D���E). The spicules are not conspicuous externally. Each tubercle contains dense bundles of spicules, not protruding through the tubercle surface. The strongly calcified spicules are of various sizes, most of them are solid (Fig. 6 F). The gill cavity is absent. Six to eight bi- and tripinnate gills form an almost complete semicircle around the anus, and one tubercle may be present just behind the anus. (Fig. 9 E). Three gills were detected in a juvenile of 5���6 mm length. The oral veil is large, since it consists of two pairs of processes: a single, broad trapezoid upper triangular projection that is not medially fused with the hyponotum, and two flattened lobes below (Fig. 9 B). The foot is broad, anteriorly rounded, and posteriorly slightly projecting beyond the notum in crawling animals forming a rounded tail (Fig. 9 A). Colour. The living specimens are milky white, slightly transparent, with a brownish intestine scarcely visible in the middle of the notum. The integument of the notum (including hyponotum) and rhinophores are densely covered with small, faint opaque white dots (Figs. 9 A, C). On the tubercle tops and rhinophoral lamellae edges the white pigment is almost entirely absent. The gill edges are covered with white dots. The white gonad shines through mature animals (including the holotype), and ventrally the reddish digestive gland can be seen. Anatomy. Digestive system. The anterior part of the buccal bulb is modified into the prominent, sessile buccal pump (Fig. 7 B). The buccal pump is fully banded by a relatively narrow peripheral muscle (Fig. 7 B). The lateral sides of the buccal pump are provided with thin muscular fibres. The salivary glands are massive triangular lobes (Fig. 7 B, left figure). The rounded labial disk is covered by yellowish cuticle bearing fine, knob-like labial elements (Figs. 3 C, G). The radular formula in six studied specimens (15���21 mm length) is 27��� 32 x 6 ���9.1.1.1.9��� 6. Radular teeth are slightly yellowish. The central tooth is small, elongated, rectangular and folded (Figs. 11 A, D). The first lateral tooth is provided with a long, wide base and a strong slightly curved beak-shaped cusp, bearing 10���15 small denticles (Fig. 11 C). The outer denticles gradually reduce in size towards the internal ones. Outer lateral teeth have slightly elongated bases, with a curved, hooked cusp on its lateral corner; all are similar in size and shape (Figs. 11 A, C). In smaller specimens (8���9 mm length) the denticles of the innermost lateral teeth are relatively larger and fewer (8���10) (Fig. 11 D) and the cusp itself is straighter. This condition is somewhat similar to the condition in juveniles of Adalaria proxima (Fig. 11 L) and adult specimens of Adalaria olgae sp. nov. (Figs. 11 F, H). The stomach is relatively small and narrow. A stomach caecum is absent. Circulatory system. In the pericardial sac a triangular posterior auricle and a smaller sized oval ventricle are present. The blood gland is rather large in relation to the central nervous system, lies above it and comprises from both posterior and anterior lobes. Central nervous system. The cerebral and pleural ganglia are well separated, the latter being somewhat larger in size. The optic nerve is very short. The eyes are not large, with black pigment in all studied specimens. The pedal ganglia are similar in size to the cerebrals, lay below them and are connected to them by very short connectives. The rhinophoral ganglia are spherical. The buccal ganglia are slightly oval. Gastroesophageal ganglia are not differentiated. Six pairs of cerebral nerves, three pleural and three pedal ones are detected. Reproductive system. (Figs. 12 A, B). The ampulla is moderately short and narrow (Figs. 12 A, B, a). The post-ampullar duct bifurcates into a long vas deferens and a short proximal oviduct (Fig. 12 B, pr and pov). The prostate has two distinct parts; a proximal, narrow, rather long convoluted duct partially encircles the bursa copulatrix, a distal, short but greatly swollen part is wrapped within a thin sheath and forms a few lobes (Fig. 12 A, pr). The prostate transits to a long single-looped penial sheath, which contains several loops of the ejaculatory duct (Fig. 12 A, psh). The inverted penial sheath and the ejaculatory duct (penis) is long and rather thick, without spines and additional terminal processes (Fig. 9 F). The moderately sized, globular bursa copulatrix contains some pinkish-red substance; it enters into the vagina via a short narrow stalk (Fig. 12 B, bc). The proximal oviduct (Fig. 12 B, pov) is short and rather straight; it extends from near to the junction of ampulla and prostate to the vagina at the bases of the seminal receptacle and bursal stalk. The seminal receptacle is wide, swollen, similar in diameter to the vagina and appears as its prolongation rather than as a separate structure (Fig. 12 B, rs). The vagina is a long, wide and convoluted duct (Fig. 12 A, v); near its opening, it has an additional pouch, the vaginal bursa (Fig. 12 A, vb), and then it opens via a short distal descending part (Fig. 12 A, pv) and also transits to the off-white nidamental glands by a short wide indistinct distal oviduct (Fig 12 A, dov). Biology. Specimens were found predominantly on large boulders covered with several species of encrusting bryozoa, at 18���26 m depth, where it is a very common species. Distribution. Presently known only from the type locality. Remarks. Adalaria slavi sp. nov. is well distinguished from other species of the genus by a number of characters. The present species is similar to A. proxima (Fig. 11 G) and A. loveni in the number of outer laterals (up to 9) but markedly differs regarding the shape of the first lateral tooth (beak shaped covered with small denticles instead of smooth straight cusp), shape of the prostate comprising two parts, different shape of the notal tubercles and characteristic opaque white small dots densely scattered all over the dorsal body side. Adalaria tschuktschica Krause, 1885 (Figs. 11 I, M) and the poorly described Lamellidoris spiculoides Volodchenko, 1941 were considered as nomina dubia by Martynov (2005) and Millen (2006); they differ from Adalaria slavi in having spiniform elongated notal tubercles, by the shape of the first lateral teeth and by a considerably fewer number of outer lateral teeth (5���6 instead of 6���9). The recently described NE Pacific A. evincta Millen, 2006 significantly differs from Adalaria slavi by the presence of globular tubercles on a very narrow stalk with protruding long spicules, by the differently shaped lateral teeth, the smaller number of outer laterals (3���6 instead of 6���9), the long, convoluted, narrow prostate, and by the colouration. Specimens of Adalaria jannae Millen, 1987, have been found in course of the present study from Kamchatka waters, but shallowly and never together with the new species (Fig. 8 A). They are well distinguished externally from Adalaria slavi in having smaller and more slender notal tubercles, a very hard dorsal notum with a strong network of spicules shining through, the presence of a well defined postbranchial gland, and the semitransparent white or yellowish colour without opaque white dots. Internally A. jannae (Figs. 11 J, K) also clearly differs from Adalaria slavi by its radula that is entirely devoid of central teeth, sharper denticles on the first lateral tooth cusp, fewer outer laterals (4���6 instead of 6���9), and a shorter ejaculatory duct in the reproductive system. Finally, the present species markedly differs from the sympatric Adalaria olgae sp. nov., which inhabits the same depth, by its white colour, bi- and tripinnate gills instead of unipinnate ones, distinct tentacle lobes on the oral veil, a sessile buccal pump, the shape of the first lateral teeth, and by a larger number of outer lateral teeth which differ in their shape. Adalaria slavi is readily distinguished from all known Adalaria species by having a large trilobed oral veil with paired lower lobes and an entire upper lobe, in combination with other features such as body size, colour, and radular features that are summarized in Table 2. The present species, like at least most other Adalaria and Onchidoris species (see Millen 1987; Schmekel & Portmann 1982; present study, Figs. 12 B; 15 A, C), shows an arrangement of reproductive organs that differs from usual doridoidean systems with oocytes and allosperm entering the female gland mass via separate ducts (i.e., oviduct and uterine duct): the proximal oviduct does not enter the female gland mass but connects to the vaginal system; oocytes and allosperm enter the female gland mass via a distally situated, combined duct. Contrary information on Adalaria jannae, A. proxima (Alder & Hancock, 1854), Onchidoris bilamellata (L., 1767) and O. muricata (M��ller, 1776) by Fahey & Vald��s (2005) likely are due to observational errors (Thompson 1966; Millen, 1987; own reexaminations of A. jannae, O. bilamellata and O. muricata)., Published as part of Martynov, Alexander, Korshunova, Tatiana, Sanamyan, Nadezhda & Sanamyan, Karen, 2009, Description of the first cryptobranch onchidoridid Onchimira cavifera gen. et sp. nov., and of three new species of the genera Adalaria Bergh, 1879 and Onchidoris Blainville, 1816 (Nudibranchia: Onchidorididae) from Kamchatka waters, pp. 1-43 in Zootaxa 2159 on pages 26-29, DOI: 10.5281/zenodo.188931, {"references":["Krause, A. (1885) Ein Beitrag zur Kenntniss der Mollusken-Fauna des Beringsmeeres. II. Gastropoda und Pteropoda. Archiv fur Naturgeschichte 51, 256 - 302.","Volodchenko, N. I. (1941) New species of the nudibranchiate molluscs from the Far-Eastern seas of USSR. Issledovaniya dalnevostochnykh morei USSR, 1, 53 - 68.","Millen, S. V. (2006) A new species of Adalaria (Nudibranchia: Onchidorididae) from the Northeastern Pacific. Proceedings of the California Academy of Sciences, 57, 357 - 364.","Millen, S. V. (1987) The nudibranch genus Adalaria, with a description of a new species from the Northeastern Pacific. Canadian Journal of Zoology, 65, 2696 - 2702.","Schmekel, R. L. & Portmann A. (1982) Opisthobranchia des Mittelmeeres, Nudibranchia und Saccoglossa. Fauna e flora del Golfo di Napoli 40, Monografia della Stazione Zoologica di Napoli, pp. i - viii, 1 - 410, Springer-Verlag.","Muller, O. F. (1776) Zoologiae Danicae. Prodromus seu animalium Daniae et Norvegiae indigenarum characteres, nomina, et synonyma imprimis popularium. Heineck et Faber, Hafnia et Lipsia, 282 pp.","Fahey, S. J. & Valdes, A. (2005) Review of Acanthodoris Gray, 1850 with a phylogenetic analysis of Onchidorididae Alder and Hancock, 1845 (Mollusca, Nudibranchia). Proceedings of the California Academy of Sciences, Ser. 4., 56, 213 - 273."]}

Genus Adalaria Bergh, 1879 Adalaria: Bergh, 1879: 360 Synonyms: Arctadalaria Roginskaya, 1971 Type species: Doris loveni Alder & Hancock, 1862, by monotypy Diagnosis. The notum is covered with well-defined spiculose tubercles of variable shape. The integument contains a dense network of spicules. The rhinophores are lamellate. The rhinophoral pockets have poorly defined, contractile smooth sheaths adjoined by several tubercles. The contractile gills are bi-tri to unipinnate, arranged separately in an almost complete circle around the anus. A common gill cavity or separated gill pits are completely absent. The oral veil is well defined, semi-circular, in one species distinctly trilobed. The foot is broad, not bilobed anteriorly, slightly narrowed posteriorly. The labial cuticle contains weakly defined polygonal elements. The buccal pump is large, prominent, sessile or on a short broad stalk, fully banded medially by a broad peripheral muscle. The salivary glands are short. The radula formula is 3 ���13.1.1.1.13��� 3. The central teeth are small and rectangular; present in most species. The first lateral tooth is large, beakshaped, in most species bearing well defined denticles. Further lateral teeth are small elongate and characteristically excavated. The stomach is relatively small, a stomach caecum is absent. The male part of the reproductive system is comprised of a looped, moderately narrow prostate, and a long musculary sheathened ejaculatory duct. The penis is not armed. The post-ampullar gonoduct bifurcates into a vas deferens and a proximal oviduct that connects to the vagina. The combined distal oviduct / uterine duct starts separately from the vaginal duct, close to the external genital opening. The bursa copulatrix is small and rounded. The seminal receptacle is distinct but always buried within nidamental glands near bursa base. The 7 valid species (including 2 described in the present paper) are listed in Table 2. The North Pacific is the center of the genus diversity., Published as part of Martynov, Alexander, Korshunova, Tatiana, Sanamyan, Nadezhda & Sanamyan, Karen, 2009, Description of the first cryptobranch onchidoridid Onchimira cavifera gen. et sp. nov., and of three new species of the genera Adalaria Bergh, 1879 and Onchidoris Blainville, 1816 (Nudibranchia: Onchidorididae) from Kamchatka waters, pp. 1-43 in Zootaxa 2159 on page 25, DOI: 10.5281/zenodo.188931, {"references":["Bergh, R. (1879) Gattungen nordischer Doriden. Archiv fur Naturgeschichte 45, 340 - 369.","Roginskaya, I. S. (1971) Arctadalaria septemtrionalis gen. n., sp. n. (Onchidorididae) a new nudibranchiate mollusc from the Laptev sea. Zoologicheskii Zhurnal, 50, 1154 - 1157.","Alder, J. & Hancock, A. (1862) Description of a new genus and some new species of naked Mollusca. Annals and Magazine of Natural History, series 3, 10, 102 - 105."]}

Adalaria olgae sp. nov. (Figures 3 D, H; 6 G–H; 7 C; 10; 11 E, F, H; 12 C–D. Table 2) Type Material. Holotype, ZMMU Lc- 37451, NW Pacific near Kamchatka peninsula, Starichkov Id., 20–26 m depth, collected by T.A. Korshunova and A.V. Martynov, 14.08. 2008. Paratypes, ZMMU Lc- 37452, two dissected specimens, same locality and collectors as holotype, 20–25 m depth, 14.08. 2008. Paratype, ZMMU Lc- 37453, one dissected specimen, same locality and collectors as holotype, 19.08. 2008. Paratypes, ZMMU Lc- 37454, three specimens, same locality and collectors as holotype, 14.08. 2008. Paratypes, ZMMU Lc- 37455, ten specimens, same locality and collectors as holotype, 14.08. 2008. Type locality. NW Pacific, SE Kamchatka, Starichkov Id., 18–26 m depth. Etymology. Adalaria olgae sp. nov. is named in honour of the daughter of two authors (AM and TK). Description. External morphology. The dimensions of the holotype are 13.5 mm x 7 mm (Fig. 10). The length of 10 living specimens ranged from 5.5 to 13.5 mm, the width ranged from 3.5 to 7 mm. The consistency of the living animals is rather soft. The notum is moderately broad, rounded in front and posteriorly. The rhinophores are long and retract into sheaths with smooth edges, except for 3 tubercles of variable size at the edges of the sheaths (Fig. 10 D). The rhinophoral sheath edges are capable of some contraction in living specimens. There are 6–9 rhinophoral lamellae. The clavus of each rhinophore lacks a ridge posteriorly. The notum is densely covered with club-shaped or almost globular tubercles on a short stalk (Fig. 10 A). The top of the tubercles often has a peculiar wrinkled appearance, somewhat tulip-shaped (Fig. 10 C). The tubercles of the mid-notal area are somewhat wider and more globular than those at the notal edge. Larger tubercles are regularly intermingled with smaller ones. Rays of spicules radiating from the bases of tubercles form a network on the surface of the apparently soft notum (Fig. 6 G). The spicules are not conspicuous externally. Each tubercle contains dense bundles of spicules, which do not protrude through the tubercle surface. The strongly calcified spicules are of various sizes, some with a narrow channel inside, some solid (Fig. 6 H). A gill cavity is absent. Ten to thirteen unipinnate gills form a semicircle around the anus (Figs. 10 A, C). Within the gill circlet, one long and four shorter elongate tubercles are situated just before the anus. The oral veil, as in Adalaria slavi sp. nov., consists of a single, broad trapezoid anterior triangular projection which is medially not fused with the hyponotum, and two ventro-lateral flattened lobes (Fig. 10 B). The tentacular lobes are less defined than in Adalaria slavi, so the general appearance of the oral veil is more similar to other Adalaria and Onchidoris species (Figs. 8 A–C). The foot is broad, anteriorly rounded and thickened, and posteriorly slightly projecting beyond the notum in crawling animals forming a rounded tail (Fig. 10 A). Colour. The living specimens have a remarkable, bright lemon yellow ground colour, which is constant in all studied specimens (Fig. 10). Under magnification the yellow pigment appears as numerous small dots. The rhinophores (including lamellae) are similar in colour to the ground colour. The upper part of tubercles is lighter. The gills are semitransparent-white, without traces of the yellow pigment (Fig. 10). Bright white gill glands are in the notum at the gill bases. Anatomy. Digestive system. The anterior part of the buccal bulb is modified into the prominent buccal pump having a short, broad, but conspicuous stalk (Fig. 7 C). The buccal pump is fully banded by a relatively broad peripheral muscle (Fig. 7 C). The lateral sides of the buccal pump have thin muscular fibres. The rounded labial disk is covered by yellowish cuticle bearing fine distinct regular polygonal elements (Figs. 3 D, H). The radular formula in four specimens (11–13 mm length) is 30– 31 x 3 –4.1.1.1.4– 3. A few anterior radular rows have only two or three outer lateral teeth, whereas most of the further radula rows possess four laterals. Radular teeth are slightly yellowish. The central tooth is small, elongated, rectangular, and folded (Figs. 11 E, H). The first lateral tooth is provided with a long, wide base and a strong, slightly curved cusp. The cusp bears 4–8 denticles that are placed in a characteristic pattern. The outermost 1–3 denticles are conspicuously larger than the rest (Figs. 11 F, H). There is a prominent triangular knob medial to the denticular ridge and a poorly defined medial wing. Outer lateral teeth are slightly elongated plates, with a downward directed cusp on its lower outside corner, and all are similar in size and shape (Figs. 11 E–F). The stomach is relatively small and narrow. A stomach caecum is absent. Circulatory system. In the pericardial sac the thin-walled, triangular posterior auricle and a smaller sized, oval ventricle are present. A well-defined blood gland lies above the central nervous system. Central nervous system. The cerebral and pleural ganglia are well separated, the latter being somewhat smaller in size. The optic nerve is very short. The eyes are relatively large, with black pigment in all studied specimens. The pedal ganglia are smaller than the cerebrals, lay below them, and are connected to them by very short connectives. The rhinophoral ganglia are rather irregular, round or elongate. The buccal ganglia are roundish-oval. Gastro-esophageal ganglia are present. Five pairs of cerebral nerves, two pleural and three pedal ones are detected. Reproductive system. (Figs. 12 C, D). The ampulla is long and wide, but not filled by sperm in all studied specimens and thus generally appeared inconspicuous (Figs. 12 C, D, a). The post-ampullar duct bifurcates into a thick prostatic loop and a proximal oviduct (Fig. 12 D). The prostatic part of vas deferens is a relatively long thickened loop (Figs. 12 C, D, pr), which does not encircle the bursa copulatrix. The prostate is wide, swollen, filled with sperm and not granulated. The prostate transitions into a long swollen single-looped penial sheath, which contains several loops of the ejaculatory part of the vas deferens (Fig. 12 C, psh). The inverted penial sheath and ejaculatory duct (penis) are long and rather thick, without spines, but have two additional short terminal knobs, not united into a singular structure (Figs. 10 E; 12 D, p). The vagina is very short and somewhat widened (Fig. 12 D, v); it forms a short stalk to the relatively large globular bursa copulatrix (Fig. 12 D, bc) containing a pinkish-brown substance. The proximal oviduct is very short and indistinct, entering near the base stalk of the bursa copulatrix. The seminal receptacle is hardly distinguishable and just represented by an elevation of the bursa stalk (12 D, rs). The combined distal oviduct and uterine duct emerges near the junction of the ampulla and seminal receptacle. This duct is long and wide, narrowing and diminishing within the nidamental glands (Figs. 12 C, D, dov). Biology. Specimens were found predominantly on large boulders covered with several species of encrusting bryozoa, at 18–26 m depth, where the species is considerably less common than Adalaria slavi. Distribution. Presently known only from the type locality. Remarks. Adalaria olgae sp. nov. is well distinguished from other species of the genus by a number of characters. From A. proxima (Figs. 8 B; 11 G) and A. loveni the present species markedly differs in having considerably fewer outer lateral teeth (3–4 instead of 9–13), by the shape of the first lateral tooth (curved beak shaped with small denticles instead of almost straight smooth cusp) by its relatively short and markedly swollen prostate, a penis with two short terminal knobs, by the different shape of the notal tubercles, the bright yellow-lemon ground colour with off-white gills, and also by its smaller body length. Adalaria tschuktschica (Figs. 11 I, M) and Lamellidoris spiculoides differ from Adalaria olgae in having spiniform, elongate notal tubercles, a semicircular oral veil without evident tentacle lobes and a larger number of distinctly shaped outer lateral teeth. Adalaria evincta significantly differs from Adalaria olgae by its globular tubercles on a very narrow stalk bearing distinctly protruding long spicules, and a couple of internal features that are summarized in Table 2. Adalaria jannae is found in Kamchatka waters, but more shallowly and never together with the new species (Fig. 8 A); it is well distinguished externally from Adalaria olgae in having smaller and more slender notal tubercles, a very hard notum with a translucent strong spicule network (Figs. 6 K; 8 A), and the presence of a well defined postbranchial gland. Internally A. jannae also clearly differs from Adalaria olgae by its radula that is entirely devoid of central teeth, among other features (Table 2). Finally, the present species differs considerably from the sympatric and syntopic Adalaria slavi sp. nov. (Figs. 9; 11 A–D) regarding colour, unipinnate versus bi- and tripinnate gills, less defined tentacle lobes on the oral veil, a stalked buccal pump, fewer outer lateral teeth and their shape, a swollen prostate, and the tiny knob-shaped seminal receptaculum that is integrated at the base of the bursal duct. The coloration of Adalaria olgae (intense lemon yellow with semitransparent-white gills) is especially remarkable and diagnostic. Whereas for some predominantly white onchidoridid species yellowish colour variations have been reported (e.g. for Adalaria proxima (Fig. 8 B) and Onchidoris muricata; Thompson & Brown 1984; Millen 1985) all collected specimens of Adalaria olgae showed a homogenous invariable intense lemon yellow colour, which markedly differs this new species from any yellowish onchidoridid colour varieties (including yellow and orange variants of A. jannae recorded from NE Pacific only). Its colour pattern also readily distinguishes Adalaria olgae from all other onchidoridids of similar size of the Kamchatka waters. In the present study radulae of Adalaria olgae were compared with those of juvenile Adalaria proxima (Fig. 11 E–F and 11 L respectively). Adalaria proxima at the length of ca. 10 mm transforms the denticulate first lateral teeth (Fig. 11 L) into a smooth one (Fig. 11 G) (Thompson 1958; Thompson & Brown 1984; present study). Adalaria olgae has denticulate first lateral teeth, and, at this size, already has a mature reproductive system. Thus, the first lateral teeth of the juvenile type persist in adult Adalaria olgae. Denticulation pattern, however, differs between A. proxima juveniles and Adalaria olgae: the former posess 2–3 large similar-sized denticles (Fig. 11 L), whereas the new species has usually more than 5 denticles, which are differentiated into larger and smaller ones (Fig. 11 F, H). All studied A. proxima specimens of 10–15 mm length (Barents Sea, Martynov et al. 2006) had immature, poorly developed reproductive systems but already entirely smooth first lateral teeth, whereas all investigated Adalaria olgae (8–13 mm) possess mature, well differentiated reproductive system and denticulated first laterals. Details of the reproductive system (for instance the very short vagina and distinct swollen prostate) and dorsal tubercles shape also significantly differ between Adalaria olgae and A. proxima (Table 2). Comparisions of the external shape of the adults and juveniles of A. proxima (Fig. 8 B and 8 C respectively) and adult specimens of Adalaria olgae (Fig. 10) highlight these differences. Distinguishing features of Adalaria olgae sp. nov. are summarized in Table 2.

Onchimira gen.nov. Type Species: Onchimira cavifera gen. et sp. nov. Diagnosis. The notum is soft and almost smooth, relatively broad, usually covered with very short tubercles. The integument contains a sparse network of spicules. The rhinophores are lamellate. The rhinophoral pockets have well defined contractile sheaths with smooth edges. The gills are unipinnate united by a common membrane into a circle around the anus, and are retractable into a common true gill cavity. The gill cavity border is moderately raised and has a smooth edge, which is capable of closing entirely over the gills. The oral veil is well defined, trapezoid, with oblique lateral sides and a convex anterior edge. The foot is broad, not bilobed anteriorly, slightly narrowed posteriorly. The labial cuticle contains dark rods with bent tips. The buccal pump is large, sessile, fully banded medially by a broad peripheral muscle. The salivary glands are elongate. The radula formula is 7 ���9.1.1.1.9��� 7. The central teeth are small and rectangular. The first lateral tooth is large, beak-shaped, bearing small faint denticles. Further lateral teeth are small and elongate. The stomach is relatively large, a caecum is absent. The male part of the reproductive system comprises a small single looped prostate, a moderately sized muscular vas deferens and a large, distinctly swollen penis, which contains a smaller evertable part. The penis is not armed. The ampulla bifurcates into a vas deferens, uterine duct and oviduct. The bursa copulatrix is small and rounded. The seminal receptacle does not represent a separate structure; instead it is formed by a swollen terminal part of the vagina, which is wide, massive and moderately convoluted. Etymology. From onchi in referring to the family Onchidorididae and mira (Latin), = remarkable, extraordinary. Onchimira is a noun of feminine gender. Remarks. See under species description., Published as part of Martynov, Alexander, Korshunova, Tatiana, Sanamyan, Nadezhda & Sanamyan, Karen, 2009, Description of the first cryptobranch onchidoridid Onchimira cavifera gen. et sp. nov., and of three new species of the genera Adalaria Bergh, 1879 and Onchidoris Blainville, 1816 (Nudibranchia: Onchidorididae) from Kamchatka waters, pp. 1-43 in Zootaxa 2159 on pages 17-18, DOI: 10.5281/zenodo.188931

Genus Onchidoris Blainville, 1816 Onchidoris: Blainville, 1816: 96 Synonyms: Ancylodoris Dybowski, 1900 Atalodoris Iredale & O'Donoghue, 1923 Lamellidoris Alder & Hancock, 1855 Oicodespina Gistl, 1848 Onchidiorus F��russac, 1822 Onchidora Cuvier, 1830 Onchiodora Desmarest, 1858 Oncidoris Herrmannsen, 1847 Onchidorus Blainville, 1816 Oncidiodoris Gray, 1847 Oncodoris Agassiz 1846 Proctaporia M��rch, 1857 Villiersia Orbigny, 1837 Type species: Onchidorus leachii Blainville, 1816, by monotypy Diagnosis. The notum is covered with well-defined spiculose tubercles of various shapes in most species. The integument contains a dense network of spicules. The rhinophores are lamellate. The rhinophoral pockets have poorly defined contractile smooth sheaths adjoined by several tubercles. The contractile gills are unipinnate, inserted separately in an almost complete circle around the anus. A common gill cavity or separated gill pits are completely absent. The oral veil is well defined, semi-circular. The foot is broad, not bilobed anteriorly, slightly narrowed posteriorly. The labial cuticle contains weakly defined polygonal elements. The buccal pump is large, prominent, on a narrow stalk, fully banded medially by a broad peripheral muscle. The salivary glands are short. The radula formula is 1 ���0.1.0���1.1.0��� 1 (but two outer laterals were reported from O. muricata by Thompson & Brown (1984)). The central teeth are small and rectangular; absent in most species. The first lateral tooth posesses a variably beak-shaped cusp, bearing small denticles in most species. The outer lateral teeth are squarish or elongate and usually posess a posterior or anterior denticle. The stomach is relatively small, a caecum is absent. The male part of the reproductive system shows a single short or looped prostate, and a long ejaculatory duct within a muscular sheath. The penis is not armed. The post-ampullar gonoduct bifurcates into a vas deferens and a proximal oviduct, which connects to the vagina. The combined distal oviduct and uterine duct starts separately from the vaginal duct, close to the external genital opening. The bursa copulatrix is small and rounded. The seminal receptacle is distinct but always buried within nidamental glands near to the base of the bursa. The genus includes 15 valid species (one of them is described in the present paper, Table 3). The North Atlantic and the Mediterranean Sea is the center of the genus diversity. Most of the Atlantic species of the genus still are poorly known anatomically. Some species, e.g. O. tridactila Ortea & Ballesteros, 1982 were described only from few specimens and their deliniation from for instance O. neapolitana (Delle Chiaje, 1841) is not completely satisfactory; both O. neapolitana and O. tridactila share very similar first lateral teeh with a strongly folded flange., Published as part of Martynov, Alexander, Korshunova, Tatiana, Sanamyan, Nadezhda & Sanamyan, Karen, 2009, Description of the first cryptobranch onchidoridid Onchimira cavifera gen. et sp. nov., and of three new species of the genera Adalaria Bergh, 1879 and Onchidoris Blainville, 1816 (Nudibranchia: Onchidorididae) from Kamchatka waters, pp. 1-43 in Zootaxa 2159 on pages 34-35, DOI: 10.5281/zenodo.188931, {"references":["Blainville, H. (1816) Quartrieme memoire sur les mollusques, de l'ordre des cyclobranches. Bulletin des Sciences par la Societe Philomathique, Paris 13, 93 - 97.","Thompson, T. E. & Brown, G. H. (1984) Biology of opisthobranch molluscs, vol. 2, Ray Society, no. 156., 229 pp.","Ortea, J. & Ballesteros, M. (1982) Sobre algunos Onchidoris Blainville, 1816 (Mollusca, Opistobranchia, Doridacea) del litoral iberico. Investigacion Pesquera 46, 239 - 254.","Delle Chiaje, S. (1841) Descrizione e notomia degli animali invertebrati della Sicilia citeriore osservati vivi negli anni 1822 - 1830, Napoli, 7, plates 86 - 173."]}

Onchimira cavifera gen. et sp. nov. (Figures 2; 3 A, E; 4 A���C, F, J���K; 5 A, B; 6 A, B; 7 A. Table 1) Type material. Holotype, ZMMU Lc- 37446, NW Pacific near Kamchatka peninsula, Starichkov Id., 20���26 m, large boulders, collected by T.A. Korshunova and A.V. Martynov. 14.08. 2008. Paratypes, ZMMU Lc- 37447, 11 specimens (three dissected), same locality and collectors as holotype, 14.08. 2008. Paratypes, ZMMU Lc- 37448, three specimens (one dissected), same locality and collectors as holotype, 18���24 m, 19.08. 2008. Paratypes, ZMMU Lc- 37449, three specimens (one dissected), same locality and collectors as holotype, 19.08. 2008. Paratypes, ZMMU Lc- 37450, three specimens (one dissected), same locality and collectors as holotype, 19.08. 2008. Type locality. NW Pacific, SE Kamchatka, Starichkov Id., 18���26 m depth. Etymology. The species epithet from the Latin cavi (= cavity) and fero (= to bear) refers to the presence of a well-defined gill cavity. Description. External morphology. The dimensions of the holotype are 22 mm x 12 mm (Fig. 2). The length of fifteen measured living specimens ranged from 8.5 to 25 mm, the width ranged from 4.5 to 14 mm. The consistency of the living animals is soft. The notum is rather broad, rounded in front and posteriorly. The rhinophores are long and retracted into raised sheaths with smooth, soft, sometimes slightly crenulate edges, not bearing tubercles (Fig. 2 C). The rhinophoral sheath edges are capable of considerable contraction in living specimens. There are 5���9 rhinophoral lamellae. The rhinophore clavus lacks a posterior ridge. The notum is almost smooth, sparsely covered with wrinkled low elevations, sometimes raised to very low tubercles (Fig. 2 A). Rays of spicules radiate from the bases of such elevations and form a sparse network in the notum (Fig. 6 A), but spicules are not conspicuous externally (Fig. 2 A). Each elevation contains sparsely placed spicules, which do not protrude from the tubercles. The strongly calcified spicules are of various size, most with a narrow channel inside (Fig. 6 B). 10���15 (usually 12) unipinnate gills are united by a common membrane into a circle around the anus. Gills are retractable into a common true gill cavity, which is capable of closing over the gills completely (Figs. 2 E���H). The border of the gill cavity is moderately raised and has a smooth edge (Figs. 2 E, F). The oral veil is well defined, trapezoid, with oblique lateral sides and convex anterior edge (Fig. 2 B). The foot is broad, anteriorly rounded and not thickened; posteriorly it projects slightly from the notum in crawling animals, forming a rounded tail. Colour. The living specimens are grayish with creamy tinge (Fig. 2). The rhinophores (including lamellae) are similar to the ground colour but more intensively cream. The gills are semitransparent white, similar to the ground colour. The pinkish digestive gland is slightly visible through the notum dorsally and shines more clearly through the foot ventrally. A purple-blackish female gland mass is visible through the anterior part of the right side in some specimens. Anatomy. Digestive system. The anterior part of the buccal bulb is modified into the sessile, large, buccal pump which is medially fully banded by a broad peripheral muscle (Fig. 7 A). The lateral sides of the buccal pump are provided with thin muscular fibres. The rounded labial disk is covered by a brown cuticle bearing distinct, rod-shaped labial elements with bent tips (Figs. 3 A, E). The radular formula in four specimens (17���23 mm length) is 25- 28 x 7 ���9.1.1.1.9��� 7. The radular teeth are slightly yellowish. The central tooth is small, elongate, rectangular and folded on the lateral edges (Fig. 4 A). The first lateral tooth is large and provided with a long, wide base and a strong, slightly curved beak-shaped cusp (Figs. 4 A���C, F). The cusp sometimes bears small faint denticles (Fig. 4 C). Outer lateral teeth are elongated small plates without cusps, all similar in size and shape (Figs. 4 A, B, F). The salivary glands are relatively long and narrow (Fig. 7 A). The stomach is relatively large and broad, then rapidly narrowing to the intestine. The stomach caecum is absent. Circulatory system. In the pericardial sac the broadly triangular posterior auricle and the smaller sized oval ventricle are present. The rather massive, irregularly rectangular blood gland is located above the central nervous system. Central nervous system. The cerebral and pleural ganglia are well separated, the latter being somewhat smaller in size. The optic nerve is very short. The eyes are relative large, with black pigment in all studied specimens. The pedal ganglia are smaller than the cerebrals, lay below them and are connected to them by very short connectives. The rhinophoral ganglia are rather irregular, rounded or elongate. The buccal ganglia are rounded-oval (Fig. 7 A). Gastro-esophageal ganglia are present. Five pairs of cerebral nerves, two pleural and three pedal ones are detected. Reproductive system. (Figs. 5 A, B). The ampulla is relatively short and narrow, not filled by sperm in all studied specimens (Fig. 5 B, a). The ampulla trifurcates into the moderately long vas deferens, uterine duct and oviduct (Fig. 5 B, pr, ud and ov). The prostatic part of the vas deferens is a very short, slightly swollen and bending duct, which does not encircle the bursa copulatrix (Figs. 5 A, B, pr). The prostate transits to a moderately long and narrow single-looped vas deferens (Figs. 5 A, B, vd), which rapidly widens and enters a common genital atrium (Fig. 5 A, at), terminating into the large, wide, and prominent penis, which contains a smaller evertable part (Figs. 4 J, K; 5 A, B, p). The vagina is relatively wide, moderately convoluted (Figs. 5 A, B, v), and enters a rather small, flattened bursa copulatrix (Figs. 5 A, B, bc). The uterine duct is long and narrow (Fig. 5 B, ud); it begins at the ampulla bifurcation and then enters the terminal part of the vagina forming a small pointed elevation. A separate seminal receptacle is absent, but the terminal part of the vagina forms a large swollen area (Fig. 5 B, tv+rs), which may serve as a receptacle. In freshly dissected living specimens, a tiny, almost inconspicuous knob-shaped structure was found on the vagina at the bursa base. This structure possibly is a vestige of a seminal receptacle, but in the ethanol-fixed specimens it was no longer detectable. The oviduct is well defined, wide (Figs. 5 A, B, ov). It starts at the ampulla, and is an irregularly convoluted duct. The capsular gland is unusually purplish-blackish in colour, having an alveolar surface (Figs. 5 A, B, cgl). Biology. Specimens were found predominantly on large boulders covered with several species of encrusting bryozoa, at 18���26 m depth, considerably less commonly than Adalaria slavi sp. nov. Distribution. Presently known only from the type locality. Remarks. Onchimira cavifera gen. et sp. nov. possesses all the usual onchidoridid characters, e.g. a welldefined buccal pump which is fully banded by the peripheral muscle, a rectangular rachidian tooth (when present), a distinct, hooked, first lateral teeth, and a number of elongate, reduced, outer laterals. The new species differs from all known onchidorids by possessing a true gill cavity of a cryptobranch dorid type. The gills are capable of complete retraction into the gill cavity, the edges of which may fully contract over the gills (Figs. 2 E-H). These features clearly delineate Onchimira gen. nov. from other onchidoridid genera (Table 1). The only genus of the family Onchidorididae, which also demonstrates the presence of a well defined gill cavity, is Calycidoris Abraham, 1876. In the present study, numerous specimens of the single known species of this genus, Calycidoris guentheri Abraham, 1876, were examined for comparison (Fig. 8 E). It is confirmed that C. guentheri possesses a well defined gill cavity, which even can contract to some degree. However, no specimens were found with a completely closed gill pocket, i.e. with edges of the cavity fully contracted over the gills. The general external appearance of Onchimira cavifera is similar to that of cryptobranch dorids, e.g. Cadlina, in having an elevated body, and markedly differs from Calycidoris guentheri, which has a flattened notum (Fig. 8 E). The notum spicule pattern of the genera Onchimira and Calycidoris is also different ― the former has a soft notum that is sparsely filled with spicules (Figs. 2 A, 6 A) whereas in the latter genus the notum is hard and contains a dense spicule network (Fig. 6 C). Internally, Onchimira gen. nov. differs from Calycidoris by the presence of a large distinct penis (Figs. 2 D, 5 A, B, p) instead of a long, narrow ejaculatory duct (Fig. 5 C, psh). An especially distinctive feature is the seminal receptacle that is apparently fused with the terminal part of the vagina forming a large swollen area (Fig. 5 B, tv+rs). This part of the vagina is similar to the pattern found in the genera Adalaria and Onchidoris, where possibly all known species have a wide, swollen seminal receptacle that directly transits into the vagina (e.g. Figs. 12 B, rs and 15 A, C, rs). In contrast, members of the genera Calycidoris and Acanthodoris possess a well-defined, long-stalked seminal receptacle (Figs. 5 C, D, rs), which is distinct from the vagina, as is also found in many cryptobranch dorids. The radula of Onchimira gen. nov. has central teeth (Fig. 4 A), while in Calycidoris central teeth are absent (Figs. 4 H, I). The lateral teeth of Calycidoris are very massive (Figs. 4 G���I), whereas those of Onchimira gen. nov. are considerably thinner (Figs. 4 A���C). The general radular appearance of Onchimira gen. nov. is rather similar to the genus Acanthodoris (Figs. 4 D, E), except for the presence of the central tooth. The poorly described Lamellidoris beringi Volodchenko, 1941 was indicated as having a common gill sheath (Volodchenko 1941). However, a single specimen personally identified by N.I. Volodchenko as Onchidoris beringi stored in the Zoological Institute, St. Petersburg, showed an external morphology that is typical for the genus Onchidoris (including the presence of numerous mushroom-shaped notal tubercles) and lacking a common gill sheath. Other features of Lamellidoris beringi indicated by Volodchenko (1941), i.e. long cylindrical notal tubercles, smooth rhinophores, narrow conical oral tentacles, short thick cusp of the first lateral tooth and 5 outer laterals, significantly differ from Onchimira cavifera. Another genus traditionally placed within family Onchidorididae, Diaphorodoris Iredale & O���Donoghue, 1923, also possesses a small gill cavity (Millen 1985; present study), but other external and internal characters are very different from the genera Onchimira, Calycidoris, and from any other onchidoridid taxa (Table 1). Earlier it was suggested (Martynov 1999 a, b) that Diaphorodoris is closely related to the phanerobranch family Anculidae. Onchimira cavifera thus is a member of Onchidorididae but cannot be incorporated into any existing genus (Table 1). There are a number of important differences regarding external features as well as digestive and reproductive organ systems. Rather than widening and confusing the current generic diagnoses we establish the new genus Onchimira. Phylogenetic analyses including characters of the newly described taxa will support or reject this hypothesis., Published as part of Martynov, Alexander, Korshunova, Tatiana, Sanamyan, Nadezhda & Sanamyan, Karen, 2009, Description of the first cryptobranch onchidoridid Onchimira cavifera gen. et sp. nov., and of three new species of the genera Adalaria Bergh, 1879 and Onchidoris Blainville, 1816 (Nudibranchia: Onchidorididae) from Kamchatka waters, pp. 1-43 in Zootaxa 2159 on pages 18-25, DOI: 10.5281/zenodo.188931, {"references":["Abraham, P. S. (1876) Notes on some genera of nudibranchiate Mollusca, with notices of a new genus and some hitherto undescribed species, in the collection of the British Museum. Annals and Magazine of Natural History, series 4, 18, 132 - 146.","Volodchenko, N. I. (1941) New species of the nudibranchiate molluscs from the Far-Eastern seas of USSR. Issledovaniya dalnevostochnykh morei USSR, 1, 53 - 68.","Millen, S. V. (1985) The nudibranch genera Onchidoris and Diaphorodoris (Mollusca, Opisthobranchia) in the northeastern Pacific. The Veliger 28, 80 - 93.","Martynov, A. V. (1999 a) Buccal pumps, gills pockets and new understanding of suctorial phanerobranchial dorids. In Systematic, Phylogeny and Biology of Opisthobranch Molluscs, (2 nd International Workshop of Malacology) Menfi, Italy, June 10 - 14 1999, 13 - 14.","Martynov, A. V. (1999 b) Nudibranch molluscs of North-West part of the Sea of Japan with discussion on taxonomy and phylogeny of the order Nudibranchia. Ph. D. thesis. Zoological Institute RAS, St. Petersburg. 424 p."]}