Dendronotus primorjensis Martynov

User, GBIF. Org, 2021, A synoptic review of the family Dendronotidae (Mollusca: Nudibranchia): a multilevel organismal diversity approach, Contributions to Zoology 90 (1), pp. 93-153 : 126-129

publication ID	https://doi.org/ 10.1163/18759866-BJA10014
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scientific name	Dendronotus primorjensis Martynov
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Dendronotus primorjensis Martynov View in CoL ,

Sanamyan & Korshunova, 2015

Fig. 7 View FIGURE 7

Dendronotus primorjensis Martynov et al., 2015a: 60 , Fig. 5A–G View FIGURE 5 ; Martynov et al., 2015c: 75–76; Korshunova et al., 2016b: 15–28, Figs 1 View FIGURE 1 , 2 View FIGURE 2 ; Korshunova et al., 2016a: 20–32; Korshunova et al., 2019b: 6–8, Fig. 3A–F View FIGURE 3 .

Dendronotus dudkai Ekimova et al. (2016) : 33–38, Figs 9–11 (synonym of D. primorjensis ).

Extended diagnosis. Body narrow. Five to nine pairs of branched dorsolateral appendages (including 2–3 smaller pairs). Five to twelve oral veil appendages. Four to five appendages (internal middle and posterior ones longest) of rhinophoral stalks.Lateral papilla of rhinophoral sheaths present. Rhinophores with 8–12 lamellae.Five to 12lip papillae.Basal colour nonuniformly reddish brown with few opaque white stripes between dorsolateral processes to more uniformly olive almost without white pigment, or almost lacking general pigmentation. Dorsal processes of jaws inclined posteriorly at approximately 45–47° to the longitudinal axis of the jaw body and 0.48 of its length. Masticatory processes bear denticles and ridge-like structures. Radula with up to 41 rows of teeth. Central tooth with deep furrows and with up to 18 (common range 12–14) distinct denticles. Up to nine lateral teeth with up to eight denticles. Ampulla moderately narrow with several windings or folded loops. Bursa copulatrix large, rounded. Seminal receptaculum small placed distally at a moderately short distance from the vaginal opening. Prostate discoid with range 12– 19 alveolar glands. The vas deferens is moderate in length, penis relatvely long, curved. Reported body length up to 35 mm.

Bathymetry. 1819–1822 m depth. Distribution. NW Pacific, common in the northern part of the Sea of Japan, at both the Remarks. So far only known from three Russian coast and at Hokkaido on the Japanese specimens collected at the type locality. coast (Korshunova Downloaded et from al., 2016 Brill.com b, 2019 12/12/2023 04:12:08 PM

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Bathymetry. 3–20 m depth. teeth usually have more distinct denticles. We therefore retain the North Pacific D. purpureus

Remarks. Dendronotus dudkai Ekimova et al. as a separate species, potentially a sister species (2016)is a synonym of D.primorjensis according to D. lacteus , until further studies are done. to morphological, molecular and geographical data. For discussion see Korshunova et al. Dendronotus robilliardi Korshunova , (2016b, 2019b). Sanamyan, Zimina, Fletcher & Martynov, 2016

Fig. 7 View FIGURE 7

Dendronotus purpureus Bergh, 1879 Dendronotus robilliardi Korshunova , Dendronotus purpureus Bergh, 1879: 89–93 , Pl. Sanamyan, Zimina, Fletcher & Martynov, I, Figs 18–20; Pl. 3, Figs 7 View FIGURE 7 –12.; Bergh, 1904:15; 2016: 28–32, Figs 2 View FIGURE 2 , 3 View FIGURE 3 , 5 View FIGURE 5 . MacFarland, 1966: 254; Odhner, 1936:1108. Non Dendronotus purpureus var. auranti- Extended diagnosis. Body narrow. Five to eight aca Friele, 1879 (= D. frondosus or D. lacteus ). pairs (commonly more than six) of branched dorsolateral appendages, digestive gland

Diagnosis (original description). Exact penetrates at least three pairs of dorolateral number of pairs of branched dorsolateral appendages. Four to six appendages of oral appendages unknown (at least, more than veil. Four to six appendages of rhinophoral three). Living colour purple, preserved reddish stalks. Lateral papilla of rhinophoral sheaths brown. Circa seven oral veil appendages. present. Rhinophores with 11–15 lamellae. Five appendages (described as equal in size, Five to ten lip papillae. Basal colour white, likely due to preservation) of rhinophoral dorsolateral appendage variable opaque stalks. Lateral papilla of rhinophoral sheaths orange or opaque white, including tip. Dorsal present. Rhinophores with circa 20 lamellae. processes of jaws inclined posteriorly at Several lip papillae (no exact count present in approximately 50–60° to the longitudinal original description). Masticatory processes axis of the jaw body and 0.43–0.45 of its of jaws with small denticles. Radula with up length. Radula with up to 43 rows of teeth. to 44 rows of teeth. Central tooth with a very Masticatory processes with ridge-like fine denticulation. Up to 14 lateral teeth with structures and smaller denticles. Central up to six denticles. Prostate and vas deferens tooth with up to 17 small, distinct denticles unknown, penis long, straight, conical. Body without furrows. Up to nine lateral teeth with length up to 20 mm. up to eight denticles. Ampulla is hook-shaped to swollen kidney-shaped, with various

Distribution. North Pacific, Bering Sea. Port degrees of folding. Bursa copulatrix large, Moller, Alaska, USA. rounded, with small seminal receptaculum placed distally at a short distance from the

Bathymetry. 31 m depth. vaginal opening. Prostate discoid with about

19–30 alveolar glands. The vas deferens is

Remarks. According to its original description short, penis relatively short, curved. Body (Bergh, 1879), D. purpureus has 14 lateral radula length up to 35 mm. teeth and is thus similar in this respect to the North Atlantic and Arctic species D. lacteus . Distribution. NW (from South Korea to However, D. lacteus of such a size has usually Commander Islands) and NE Pacific (from less than 14 lateral teeth, and also the central Alaska to Downloaded at least from Washington Brill.com State 12/12/2023, Canada 04:12:08PM

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Depth. circa 5–30 m. range does not extend further eastward than the Barents Sea.

Bathymetry. Intertidal to less than 20 m deep.

Remarks. See Remarks for D. albus . Remarks. T.A. Korshunova’s and A.V.

Martynov’s pre-2015 analysis has showed Dendronotus robustus Verrill, 1870 that there are at least two species in the “ D.

Fig. 7 View FIGURE 7 robustus complex”, namely D. robustus Verrill , Dendronotus robustus Verrill, 1870: 405 ; 1870 and D. velifer G.O. Sars, 1878 . However, Lundin et al., 2017: 303, 306, 309–315, Figs 3 View FIGURE 3 , 4 View FIGURE 4 . this analysis was unfortunately not included

Non auct., not Ekimova et al. (2015) (mix-up in the final version of the publication by I.A.

with D. velifer G.O. Sars, 1878 ). Ekimova (Ekimova et al., 2015). This detailed morphological and molecular analysis Extended diagnosis. Body wide. Six to seven has shown that D. robustus and D. velifer pairs of branched dorsolateral appendages. are well differentiated morphologically,

10 to 15 appendages of oral veil. Four to five bathymetrically and according to the appendages of rhinophoral stalks. Lateral molecular analysis (Lundin et al., 2017). In papilla of rhinophoral sheaths absent. the previous literature these two species have Rhinophores with 7–14 lamellae. More than been mixed.

20 lip papillae. Basal colour light grayish to yellowish with scattered opaque white Dendronotus rufus O’Donoghue, 1921

and yellow spots. Dorsal processes of jaws Fig. 7 View FIGURE 7

inclined posteriorly at approximately 47° Dendronotus rufus O’Donoghue, 1921:190 –

to the longitudinal axis of the jaw body and 192; Pl. 3, Figs 25–27; Pl. 4, Fig. 48; O’Donoghue,

0.55 of its length. Masticatory processes 1922:125; Robilliard, 1970: 456–460, Figs 4–6 View FIGURE 4 View FIGURE 5 View FIGURE 6 , apparently bear denticles (may possess ridge- 16–18, Pl. 63, Fig. 32.

like structures). Radula with up to 31 rows of Behrens, 1980:76; Behrens & Hermosillo, teeth. Central tooth with high cusp and up to 2005:95; Lamb & Hanby, 2005: 262.

20 small distinct denticles without furrows. Dendronotus frondosus sensu Odhner, 1936

Up to nine lateral teeth with up to seven (part.) non Ascanius, 1774.

denticles. Ampulla moderately narrow, with approximately three to four irregular loops. Extended diagnosis. Body relatively narrow.

Bursa copulatrix moderate in size, rounded to Six to nine pairs of branched dorsolateral pear-shaped,with small seminal receptaculum appendages. Five appendages of oral veil. Five placed distally near the vaginal opening. appendages of rhinophoral stalks. Lateral Prostate discoid with about 10 alveolar glands. papilla of rhinophoral sheaths present.

The vas deferens is moderate in length, penis Rhinophores with 19–24 lamellae. More massive, broad, slightly curved. Body length of than 30 lip papillae. Basal colour from red to verified specimens up to 50 mm. grayish white, white specimens commonly have remarkable differences in colour of Distribution. North Atlantic (both western and dorsolateral appendages (which can be brick eastern, Canada, Norway, Russia and USA), red to dark red). Dorsal processes of jaws very shallow waters only, in the Arctic, the inclined posteriorly Downloaded from at Brill approximately.com 12/12/2023 70 04:12 °:08PM via Open Access. This is an open access article distributed under the terms of the CC BY 4.0 License. https://creativecommons.org/licenses/by/4.0/ to the longitudinal axis of the jaw body and veil. Three to seven appendages (posterior 0.45 of its length. Masticatory processess with and medial ones longest) of rhinophoral ridge-like structures and smaller denticles. stalks. Lateral papilla of rhinophoral sheaths Radula with up to 35 rows of teeth. Central absent. Rhinophores with 3–14 lamellae. tooth with up to 28 small denticles (usually 0–4 lip papillae. Basal colour opaque white, smaller) with ribs or shallow furrows, never solid orange, brownish, ranges from pale completely smooth. Up to 16 lateral teeth yellow to dark brown, scattered small yellow with up to seven denticles. Ampulla relatively spots, often with four thin subparallel lines narrow, long, with several windings and (two on each side) next to the dorsolateral loops. Bursa copulatrix large, oval. Seminal appendages. Dorsal processes of jaws receptaculum small, placed distally at a inclined posteriorly at approximately 65° moderately short distance from the vaginal to the longitudinal axis of the jaw body and opening. Prostate discoid with numerous 0.33 of its length. Masticatory processess alveolar glands (at least more than 50). The with ridge-like structures. Radula with up vas deferens is very long, penis long, narrow, to 72 rows of teeth. Central tooth with up looped several times. Body length up to to 10 small distinct denticles with furrows. 170 mm. Up to seven lateral teeth with up to seven denticles. Ampulla voluminous, folded.Bursa Distribution. NE Pacific, from southern Alaska copulatrix large, oval. Seminal receptaculum to Washington state, Canada and USA. small placed distally at a moderately short distance from the vaginal opening. Prostate Bathymetry. 7–38 m depth. discoid with up to 12 alveolar glands. The vas deferens is moderate in length, penis thin, Remarks. This is among the largest long, winded. Body length up to 40 mm. Dendronotus species. According to the radular formula it is similar to D. lacteus and molecular Distribution. North Eastern Pacific, Canada data confirm D. rufus as a sister species to D. and USA. lacteus (fig. 1). In its biogeographical pattern it reveals quite narrow endemism, so far known Bathymetry. From intertidal to 20 m depth. only from southern Alaska to Washington state. Remarks. Dendronotus subramosus is unique among the clades of narrow-bodied Dendronotus subramosus MacFarland, 1966 species since it has no lateral papillae

Fig. 7 View FIGURE 7 on the rhinophoral sheath. However, Dendronotus subramosus MacFarland , phylogenetically D. subramosus is more 1966: 265–270, Pl. 40, Fig. 3 View FIGURE 3 ; Pl. 46, Figs 5–8 View FIGURE 5 View FIGURE 6 View FIGURE 7 View FIGURE 8 ; Pl. closely related to the externally similar- 47, Figs 3–7 View FIGURE 3 View FIGURE 4 View FIGURE 5 View FIGURE 6 View FIGURE 7 ; Pl. 49, Figs 1–3 View FIGURE 1 View FIGURE 2 View FIGURE 3 ; Pl. 50, Fig. 2 View FIGURE 2 ; Pl. 52, looking deep sea species D. jamsteci Figs 1 View FIGURE 1 , 2 View FIGURE 2 ; Robilliard, 1970: 462–466, Figs 3 View FIGURE 3 , 4 View FIGURE 4 , 6 View FIGURE 6 , (with the presence of rhinophoral sheath 19–21, Pl. 64, Fig. 33; Behrens, 1980:76; Behrens papillae), which was recently described & Hermosillo, 2005:95; McDonald, 1983: 177. off Pacific Honshu ( Japan) and also to the externally different-looking NE Pacific Extended diagnosis. Body narrow. Three shallow water species D. albus (figs 1, 2). Such to six pairs of branched dorsolateral a morphological and molecular evolutionary appendages. Four to six appendages of oral mosaic is Downloaded remarkable from and Brill.com contribute 12/12/2023 for 04:12 the:08PM