identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
1E1A4C68FF8FFFED512AFA65FEEADA80.text	1E1A4C68FF8FFFED512AFA65FEEADA80.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Dendronotus nordenskioeldi User 2021	<div><p>Dendronotus nordenskioeldi sp. nov.</p> <p>Figs 1–3, 5–7</p> <p>ZooBank: http:// urn:lsid:zoobank.org:act: 9FB63088-BF8A-4990-B292-7E26DEA0FDCF</p> <p>Etymology. In honour of Baron Nils Adolf Erik Nordenskiöld, outstanding Arctic explorer, geologist, and mineralogist. The Laptev Sea had been originally named “Nordenskiöld Sea”, after this Arctic explorer.</p> <p>Type locality. Laptev Sea.</p> <p>Type material. Holotype. ZMMU Op-665. Arctic region, Laptev Sea, sta. 100, 69 m depth, collector Olga Zimina.</p> <p>Description. Body relatively narrow, preserved holotype 19 mm length (fig. 5). Five to six main branched appendages on oral veil. Five appendages on rhinophoral stalks; 12–14 rhinophoral lamellae. Lateral papilla on rhinophoral sheath present. Five pairs of dorsolateral appendages. Circa 20 lip papillae. Dorsolateral appendages with relatively short primary stalk, moderately branched secondary branches and tertiary branches (fig. 5A, B). Basal colour is semitranslucent white. No additional pigment. Dorsal processes of jaws inclined posteriorly at approximately 53° to the longitudinal axis of the jaw body and 0.42 of its length (fig. 5D). Masticatory border of jaw with fine denticles (fig. 5E). Radular formula: 43 × 3–11.1.11–3. Central (rachidian) tooth bears very weak denticulation that are difficult to separate into individual denticles (up to 20 and more variously varied denticles at least) (fig. 5F, G). Denticles without deep furrows.Lateral teeth elongate, slightly curved, bearing up to seven denticles (+ 2–3 reduced denticles may present) (fig. 4F–H). Ampulla thickened, kidney-shaped. Prostate discoidal, consisting of up to 20 alveolar glands. Distal part of vas deferens moderate in length, transitioning to long, thick, curved copulatory organ. Bursa copulatrix large, distinctly elongated, with seminal receptaculum placed distally.</p> <p>Distribution. So far known only from the Laptev Sea.</p> <p>Remarks. Externally, Dendronotus nordenskioeldi sp. nov. resembles white forms of D. lacteus but differs by the absence of a distinct reticulation and peculiar ribs on the central teeth, which is a common characteristic of D. lacteus (Korshunova et al., 2017a). According to the molecular differences, a minimal Synopsis of the previously described valid intergroup distance for the COI gene (7.9%) taxa of the family Dendronotidae is found between D. nordenskioeldi sp. nov and D. europaeus. The mean intergroup Genus Dendronotus Alder &amp; Hancock, 1845 distances between D. nordenskioeldi sp. nov. Type species. Dendronotus frondosus and D. rufus, D. lacteus, and D. kamchaticus are (Ascanius, 1774)</p> <p>8.3%, 8.3%, and 8.8% respectively. A maximal intergroup distance (15.0%) is found between Diagnosis.Dorsolateralappendageswithdistinct</p> <p>D. nordenskioeldi sp. nov and D. patricki tertiary branches. Oral veil with branched (Appendix, table A2). appendages Downloaded. Jaws commonly from Brill.com with 12 set/12of /2023 tightly04:12:08PM 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/</p> <p>packed ridge-like semi-discoid structures and species, an extended diagnosis is provided also with denticles. Radula with at least more when possible. The fine-scale diagnostics of than eight lateral teeth in adult specimens. the genus Dendronotus is a complex task since Central teeth with cusp integrated within lateral variation patterns of habitus and anatomical denticles (or within teeth body if denticles characters overlap in various species. By a absent), the central cusp is not protruding. careful listing it is possible to step-by-step Denticle-bear part of central teeth separated fulfill the task and present a mutilelevel from lateral sides by usually distinct shoulder, organismal diversity, though not in a strictly thus central teeth with squarish base and defined form, as was targeted previously triangular top.Prostate with evident, conspicous (e.g., Odhner, 1936; Robilliard, 1970), but in alveolar glands, commonly represent by a a complex form that is best suited to the discoid structure (varies from a flattened disk evolutionary mosaic in character evolution. to a compressed spherical structure), or very There are also still uncertanties regarding rarely non-discoid, linear (in this case alveolar some particular characters as well as previous glands also present). Copulative organ basically overinterpretations. First of all, previously conical, with various ratios of length and width the pattern of branching in the dorsolateral and degrees of apical tapering. appendages was claimed to be of considerable taxonomic importance (Ekimova et al., Natural history. Representaives of the genus 2015: Fig. 8). In this respect, it is necessary to Dendronotus inhabit a broad bathymetric state, that while some trends exist in a few range, from intertidal to depths over 3000 m, species, importance of these characters were and a variety of substrates (including stony considerably overestimated partly due to the and soft benthic environments). A majority use of preserved specimens in the analysis. of the Dendronotus species feed on hydroids This can be illustrated by the D. kamchaticus (Robilliard, 1970; Thompson &amp; Brown, 1984; case, when according to Ekimova et al. (2015: Martynov &amp; Korshunova, 2011). There as a few Fig. 8D), D. kamchaticus has bulbous short notable exceptions, such as D. iris Cooper, 1863 branches that differ considerably from those from the northeastern Pacific,which is an active in any other Dendronotus species. This resulted predator of ceriantharian cnidarians (Wobber, from the use of preserved specimens with 1970; Shaw, 1991), whereas for the wide-bodied considerably contracted branches, whereas species D. robustus Verrill, 1870, D. velifer G.O. the living adult specimens of D. kamchaticus Sars, 1878 and D. bathyvela Martynov et al., have elongated branches of dorsolateral 2020a less specific prey spectrum (including appendages, which are not fundamentally polychaetes and hydroids) has been recorded different from the type species of the genus, (Roginskaya, 1990, 1997). D. frondosus (Korshunova et al., 2016a). The description of D. zakuro (similar to D. Remarks. The genus Dendronotus is restricted kamchaticus by radula morphology) is based here since some species have been moved to on recently collected specimens with both other genera. See discussions and remarks shorter “bulbous” (see Martynov et al., 2020a: under the genera Cabangus gen. nov. and Fig. 3A) and elongate dorsolateral appendages Pseudobornella. (Martynov et al., 2020a: Fig.3D, E), which is due to differences in physiological conditions. The Species composition. Diagnoses and remarks for preserved specimen of D.zakuro (see Martynov 27 included species are listed below. For each et al., 2020a Downloaded: Fig. 3C) from demonstrates Brill.com very 12/12/2023 similar 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/</p> <p>“bulbous” branches, which were incorrectly the genus Pseudobornella are mentioned after depicted as specific for D. kamchaticus in its the diagnoses of the species of the genus original description (Ekimova et al., 2015: Dendronotus.</p> <p>Fig. 8D) (see details in Synopsis under D. There are also various uncertainties and kamchaticus). Dendronotus lacteus apparently misinterpretations regarding internal charachas shorter tertiary branches, and a “curly” ters that need to be addressed before diagnoappearance of the dorsolateral appendages. ses can be provided. For example, Robilliard However, variation in D. lacteus does not only (1970) suggested to apply several coefficients concern ontogenetic stages, but also occurs of jaw proportions. However, it is easy to among different adult specimens. So, some incorrectly apply respective angle of measurelarge specimens of D. lacteus have quite long ments and receive very different results, as for elongate branches of dorsolateral appendages, example in Ekimova et al. (2015) compared similar to those in D. frondosus (Korshunova to the same species which jaws by Robilliard et al., 2017a). Most species of the genus (1970). As an example, for D. dalli in Ekimova Dendronotus possess considerably branched et al. (2015: 863) an angle of dorsal processes dorsolateral appendages with distinct tertiary to the longitudinal axis of 25° was indicated, branches (MacFarland, 1966; Robilliard, whereas in Robilliard (1970: 454) this was 60°.</p> <p>1970; Korshunova et al., 2016a, b, 2017a, In the present study we re-checked jaw pro- 2019b; Martynov et al., 2020a). Unbranched portions and presented updated values for the dorsolateralappendageswererecentlyreported majority of species with sufficient available only for D. claugei from the deep-sea (Valdes et information. This character still needs further al., 2018), but due to the most basal position testing, but some characteristic patterns for of this species (fig. 1) the generic position of different species are likely to occur. Similarly, D. claugei needs to be evaluated with more the number of the prostatic alveoles once data. For another deep-sea species, D. comteti, was thought to be a character that well charthe unbranched dorsolateral appendages acterizes different species (Robilliard, 1970; were also indicated (Valdes et al., 2018: 413), Ekimova et al., 2016). However, our previous however, according to the original description (Korshunova et al. 2016a, b) and present study of D. comteti, it possesses both branched and show that this character is much more variaunbranched dorsolateral appendages (Valdes &amp; ble, and while there are clear trends in some Bouchet, 1998: 320). Unbranched dorsolateral species, it should generally be used with care.</p> <p>appendages can occur in posterior parts of some In the species diagnoses below, differences other Dendronotus species (e.g., in D.robilliardi, in shape, length and proportions of the copsee Korshunova et al., 2016a). For some species, ulative organ (penis) are indicated. While in e.g., for D. dalli in Ekimova et al. (2015: 862), some species the differences in penial morthe indication of absent tertiary branches phology are clear, in other species this can be of dorsolateral appendages is incorrect. partly due to different degrees of contraction Dendronotus dalli definitely has tertiary after preservation. This character needs furbranches (Robilliard, 1970: 453; present study, ther investigation.</p> <p>fig. 7). Instead, in the genus Pseudobornella the In the present study we show that the shape general appearance of dorsolateral appendages and size of the bursa copulatrix, at least among does not show as branches, but it is ctenidium- several Dendronotus species (e.g., between like, because of very indistinct (or truly absent) D. kalikal and D. yrjargul), are rather contertiary branches (fig. 2). More details about stant and Downloaded can be used from Brill as.a com reliable 12/12/2023 diagnostic 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/ character. However, the exact position of the Martynov et al., 2020a), in the diagnoses commonly very small receptaculum seminis below we do not provide the number because can be very difficult to determine, but despite it can be misleading, because only a part of that, it will be given further attention in the the row was counted in various species. True diagnoses below. rodlets can occur in addition, as part of the Different terms were applied for the struc- thin labial cuticle, which is separated from tures on the masticatory edge within differ- the masticatory edges. Such true rodlets have ent species of the genus Dendronotus. For been well documented for the tropical repreexample, MacFarland (1966) used “denticles” sentatives of the family Dendronotidae (Pola or “ridge-like” denticles and Robilliard (1970) &amp; Stout, 2008), and possibly also occur in applied “denticles”, whereas Ekimova et al. many Dendronotus species, since the labial (2015) used “rodlets”. In this respect, the term cuticle with rodlets were already described for “rodlets” was used for rod-like structures that several representatives (see e.g., MacFarland, constitute the body of jaws in other nudi- 1966). However, these characters remain branch groups. In the genus Dendronotus, the largely untested by modern methods and we elements on the masticatory edges are usually do not include it in the diagnoses below. As termed as “denticles” when the bodies of jaws an additional complication, denticles and are entire. However, in the genus Dendronotus especially ridge-like structures on the mastithere is a special set of structures in addition catory edges can be easily discarded during to the denticles that appears as broad semi- preparation, and therefore in some previous disks or ridges, which can additionally be descriptions it was inappropriately indicated covered with smaller spine- or denticle-like that masticatory edges are devoid of any structures. It is likely that most species of the denticles or ridges. For example, the absence genus Dendronotus possess such structures. In of the denticles on the masticatory edges addition to the semi-disks/ridges at the masti- of D. frondosus were incorrectly mentioned catory edges, more normally looking denticles in Ekimova et al. (2015), which was further can occur. Transitive denticles on masticatory used as a putative difference between D. fronedges were also occasionally reported for juve- dosus and D. primorjensis (Ekimova et al.,</p> <p>niles of D. frondosus s.l. (Colgan, 1914; Wägele 2016). However, in both species denticles on</p> <p>&amp; Willan, 2000). Therefore, in the present the masticatory edges are definitely present study we use two different terms “ridge-like (Korshunova et al., 2016a). The same is true structures” and “denticles” when character- also for other species, such as D. kamchaticus izing the masticatory edges of Dendronotus and D. kalikal. For D. kamchaticus the absence species. The few denticles seen in many cases of any structures on the mascticatory edges are likely a precursor that further transits into was indicated in the original description. For semi-disk structures on the masicatory edges D. kalikal the presence of merely “rodlets” was</p> <p>(see e.g., this clearly for D. kalikal, fig. 4Q). reported in the original description (Ekimova Because the full row of denticles and/or ridges et al., 2015). However in reality denticles and is not documented for all species due to diffi- ridge-like structures are present in both of culties during preparation, the exact number these species (Korshunova et al., 2016a, 2019b;</p> <p>for these structures can be uncertain. Though for D. kalikal see fig. 4Q). Therefore, when the whenever possible, we already presented the original descriptions denticles or ridge-like number of denticles/ridges, for example the structures was not indicated for species that recently described Dendronotus species (e.g., we could not Downloaded check, from we suppose Brill.com the 12/12presence /2023 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/ of at least one of these structures and indicate remarkable transformation during the develthat in the diagnoses. opment from early juvenile to adult. The cen- Though a pair of salivary glands indeed tral tooth in earlier juveniles is low and bears inserts into the pharynx on the sides of the few distinct lateral denticles. The lateral teeth oesophagus, the knowledge of these is limited in early juveniles are very few in number only to a few species (e.g., MacFarland, 1966; (1–3) or absent, showing an indistinct dentic- Valdes &amp; Bouchet, 1998; Pola &amp; Stout, 2008). ulation or they are almost smooth. In individ- The salivary glands are usually not figured uals approaching the adults stage, the central in the original descriptions (an exception: teeth become elongate-triangular with dis- Valdes and Bouchet, 1998) and were never tinct shoulders, lateral denticles become consistently used for taxonomic diagnostics. relatively smaller and numerous or com- We therefore do not include salivary glands pletely reduced. Meanwhile, the lateral teeth in the diagnoses of the species of the genus increase in number and the denticles become Dendronotus. more conspicuous. Since parts of juvenile The radular patterns within the genus radula may persist in some adults, this may Dendronotus still offer one of the best diag- create a considerable confusion regarding the nostic characters (Korshunova et al., 2017a). taxonomic importance of radular characters.</p> <p>However in several species groups quite For instance, D. dalli usually has a smooth similar patterns of the central tooth can be central teeth without denticles in adult stage, observed. These similarities occur between whereas for adults of D. dalli also denticles on groups/clades of closely related species (for central teeth were reported, which led to the example between species that closely related incorrect synonymyzation of D. dalli with D.</p> <p>to D. frondosus, e.g., D. primorjensis and D. frondosus (Odhner, 1936; MacFarland, 1966).</p> <p>venustus, between D. lacteus and D. rufus, This decision was later changed in favour of between D. dalli and D. elegans), but also the validity of D. dalli, but uncertanties over among apparently more distantly related spe- the taxonomic application of the putatively cies such as D. kamchaticus and D. zakuro, and variable radula remained (Robilliard, 1970; between D. albus and D. robilliardi (see fig. 1). Roginskaya, 1987). Such “variations” can Despite these similarities, in a majority of be overcome by a consistent application these cases fine-scale differences in morphol- of general ontogenetic principles in taxonogy of radula can be detected. At the same omy, using the attribution of a particular time, different species which are grouped in set of characters to a particular ontogenetic the same subclade (fig. 1) can demonstrate stage (Martynov et al., 2015b, 2020b). Thus, considerably different radular patterns, as for the application of a multilevel diagnostic example the clade consisting of D. jamsteci, within the genus Dendronotus (one that D. subramosus and D. albus (Robilliard, 1970; combines morphological characters of vari- Pola &amp; Stout, 2008; Korshunova et al., 2016a; ous levels and molecular data) instead of a Martynov et al., 2020a). Furthermore, the rad- search for immediate and clearly distinctive ulae of the morphologically very different D. differences, requires a fine-scale, complex robilliardi and D. gracilis demonstrate similar approach.</p> <p>patterns in the presence of a widened central tooth with numerous small lateral denticles Dendronotus albopunctatus Robilliard, 1972 without furrows (Baba, 1949; Korshunova Dendronotus albopunctatus Robilliard, 1972:</p> <p>et al. 2016a, 2019b). The radula undergoes 421–430, Downloaded Figs 1–6; from Behrens Brill.com, 1980 12/12: 72 /2023; Behrens 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/ and Hermosillo, 2005: 93; Lamb and Hanby, 2005: 264.</p> <p>Extended diagnosis. Body wide. Four to five (plus one–two posterior) pairs of dorsolateral appendages. Six to 12 appendages of oral veil. Three to seven appendages (equal in size) of rhinophoral stalks. Lateral papilla of rhinophoral sheaths absent. Rhinophores with 9–17 lamellae. No lip papillae reported in the first description. Basal colour yellowwhite, pale pink to red-brown, with red-brown spots and tiny opaque white spots scattered over whole body, solid white line along edge of foot and oral veil. Dorsal processes of jaws inclined posteriorly at approximately 50° to the longitudinal axis of the jaw body. Masticatory processes bear distinct denticles which increase in size towards the distal end of the process. Radula with up to 41 rows of teeth. Central tooth with up to 25 denticles. Up to 14 lateral teeth with up to 12 denticles. Ampulla voluminous, folded. Bursa copulatrix large, oval. Seminal receptaculum very small and placed distally at a short distance from the vaginal opening. Prostate discoid with about 11–16 alveolar glands. The vas deferens is long, penis short, conical. Body length up to 60 mm.</p> <p>Distribution. NE Pacific, Washington and Oregon, USA.</p> <p>Bathymetry. Depths of 18–40 m (soft substratum).</p> <p>Remarks. Externally, D. albopunctatus is somewhat similar to other species that also inhabit soft substrata, such as D. robustus and D. velifer. However, its radular characters are more similar to those of the majority of Dendronotus species.</p></div> 	https://treatment.plazi.org/id/1E1A4C68FF8FFFED512AFA65FEEADA80	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	User, GBIF. Org	User, GBIF. Org (2021): A synoptic review of the family Dendronotidae (Mollusca: Nudibranchia): a multilevel organismal diversity approach. Contributions to Zoology 90 (1): 93-153, DOI: 10.1163/18759866-BJA10014, URL: http://dx.doi.org/10.15468/dl.ph6kj8
1E1A4C68FF97FFEA53CDFF5AFEDEDF2D.text	1E1A4C68FF97FFEA53CDFF5AFEDEDF2D.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Dendronotus albus MacFarland 1966	<div><p>Dendronotus albus MacFarland, 1966</p> <p>Fig. 7</p> <p>Dendronotus albus MacFarland, 1966: 275– 279, Pl. 40, Fig. 1; Pl. 46, Figs 1–4; Pl. 47, Figs 8 –11; Pl. 48, Figs 7, 8; Pl. 49, Fig, 5; Pl. 50, Fig. 4; Pl. 51, Figs 6, 7.</p> <p>Dendronotus diversicolor Robilliard, 1970: 470–474, Figs 4–6, 25–28, Pl. 64, Figs 35, 36.</p> <p>Not D.albus sensu Robilliard, 1970:466–470; Behrens, 1980: 72; Behrens and Hermosillo, 2005: 93; McDonald, 1983: 172; McDonald, 2009: 463 = D. robilliardi Korshunova, Sanamyan, Zimina, Fletcher, Martynov, 2016.</p> <p>Extended diagnosis. Body relatively narrow. Four to five pairs of branched dorsolateral appendages, digestive gland penetrates only two pairs of dorsolateral appendages. Up to six appendages of oral veil. Three to seven appendages (external longest) of rhinophoral stalks. Lateral papilla of rhinophoral sheaths present. Rhinophores with 12–21 lamellae. Two to eight lip papillae. Basal colour translucent white to violet, dorsolateral appendage variable opaque orange or opaque white, including tip. Dorsal processes of jaws inclined posteriorly at approximately 50–60° to the longitudinal axis of the jaw body and 0.4–0.47 of its length. Masticatory borders apparently with ridge-like structures and denticles. Radula with up to 38 rows of teeth. Central tooth with up to 25 small distinct denticles without furrows. Up to nine lateral teeth with up to 14 denticles. Ampulla voluminous, folded. Bursa copulatrix large, oval. Seminal receptaculum very small and placed distally at a short distance from the vaginal opening. Prostate discoid with about 10–30 alveolar glands. The vas deferens is relatively short, bent, penis short, nearly straight, tapering to a blunt tip. Body length up to 73 mm.</p> <p>Distribution. NE Pacific: California to southern Alaska, Canada and US. Records from Baja California, Mexico, need verification.</p> <p>Bathymetry. Intertidal to at least 30 m depth.</p> <p>Remarks. Dendronotus albus has for almost 50 years persistently been confused with D. robilliardi. Despite this, D. albus is readily distinguished from D. robilliardi by the number of dorsolateral appendages (four to five in D. albus vs. five to eight in D. robilliardi) and the number of dorsolateral appendages into which the digestive gland branches penetrate (only in two anterior pairs in D. albus vs. three to six pairs including posterior ones in D. robilliardi). See Korshunova et al. (2016a) for discussion.</p> </div>	https://treatment.plazi.org/id/1E1A4C68FF97FFEA53CDFF5AFEDEDF2D	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	User, GBIF. Org	User, GBIF. Org (2021): A synoptic review of the family Dendronotidae (Mollusca: Nudibranchia): a multilevel organismal diversity approach. Contributions to Zoology 90 (1): 93-153, DOI: 10.1163/18759866-BJA10014, URL: http://dx.doi.org/10.15468/dl.ph6kj8
1E1A4C68FF90FFEA51C9FCA4FB87DF2D.text	1E1A4C68FF90FFEA51C9FCA4FB87DF2D.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Dendronotus arcticus Korshunova	<div><p>Dendronotus arcticus Korshunova,</p> <p>Sanamyan, Zimina, Fletcher &amp; Martynov, 2016</p> <p>Fig. 7</p> <p>Dendronotus arcticus Korshunova et al., 2016a: 26–28, Figs 1, 3A.</p> <p>large and rounded. Seminal receptaculum small placed distally at a moderately short distance from the vaginal opening. Prostate discoid with about 25–30 alveolar glands. The vas deferens is moderate in length, penis long and twisted. Body length up to 19 mm.</p> <p>Distribution. Arctic – Kara Sea to Laptev Sea, Russia.</p> <p>Bathymetry. 10–20 m depth, occasionally deeper.</p> <p>Remarks. This recently described species is easily distinguished from its congenerics (Korshunova et al., 2016a).This is a true Arctic species that so far has been recorded only from the Laptev Sea and Kara Sea.</p></div> 	https://treatment.plazi.org/id/1E1A4C68FF90FFEA51C9FCA4FB87DF2D	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	User, GBIF. Org	User, GBIF. Org (2021): A synoptic review of the family Dendronotidae (Mollusca: Nudibranchia): a multilevel organismal diversity approach. Contributions to Zoology 90 (1): 93-153, DOI: 10.1163/18759866-BJA10014, URL: http://dx.doi.org/10.15468/dl.ph6kj8
1E1A4C68FF90FFE853ECFCA4FB86DE54.text	1E1A4C68FF90FFE853ECFCA4FB86DE54.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Dendronotus bathyvela Martynov, Fujiwara	<div><p>Dendronotus bathyvela Martynov, Fujiwara,</p> <p>Tsuchida, R. Nakano, N. Sanamyan, K. Sanamyan, Fletcher &amp; Korshunova, 2020</p> <p>Fig. 7</p> <p>Dendronotus bathyvela Martynov et al. 2020a: 507–510, Figs 4, 5.</p> <p>Extended diagnosis. Body relatively narrow. Extended diagnosis. Body wide. Six to seven Five to six pairs of dorsolateral appendages. pairs of branched dorsolateral appendages. 10 Six to eight appendages of oral veil. Five to to 15 appendages of oral veil. Five appendages six appendages (apparently equal in size) of rhinophoral stalks. Lateral papilla of of rhinophoral stalks. Lateral papilla of rhinophoral sheaths absent. Rhinophores rhinophoral sheaths present. Rhinophores with 15–25 lamellae. Lip papillae 50–70. Basal with 15–18 lamellae. Lip papillae 15–25. Basal colour dull reddish-brownish with numerous colour brownish with scattered distinct distinct opaque white dots on notum, tips opaque white dots. Dorsal processes of jaws of lateral appendages, oral appendages.</p> <p>inclined posteriorly at approximately 55° Dorsal processes of jaws inclined posteriorly to the longitudinal axis of the jaw body and at approximately 52° to the longitudinal 0.47 of its length. Masticatory processes axis of the jaw body and 0.56 of its length.</p> <p>apparently bear indistinct denticles (may Masticatory processes apparently bear possess ridge-like structures). Radula with up indistinct denticles (may possess ridge-like to 39 rows of teeth. Central tooth with up to structures). Radula with up to 41 rows of teeth.</p> <p>14 small denticles and reduced furrows. Up to Central tooth with high cusp and up to 35 nine lateral teeth with up to nine denticles. small distinct denticles without furrows. Up Ampulla folded twice. Bursa copulatrix is to 14 lateral Downloaded teeth from without Brill.com or 12 with /12/2023 few 04 weak: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/ lateral denticles. Ampulla voluminous, with approximately three loops. Bursa copulatrix large, rounded to oval. Seminal receptaculum small placed distally at a short distance from the vaginal opening. Prostate discoid with about 21–25 alveolar glands. The vas deferens is long, penis elongate, narrow, curved. Body length up to 45 mm.</p> <p>Distribution. Pacific coast of Northern Honshu, Japan, possibly may occur further in NW Pacific.</p> <p>jaws with irregular denticles and possibly with ridge-like structures. Radula with up to 32 rows of teeth. Central tooth with deep furrows and with up to 13 distinct denticles. Up to eight lateral teeth with up to ten denticles. Ampulla with three loops. Prostate compact with about nine alveolar glands. Bursa copulatrix large, rounded. Seminal receptaculum large placed distally at considerable distance from the vaginal opening. The vas deferens is moderate in length, penis relatively long, narrow. Body length up to 18 mm.</p> <p>Bathymetry. A shelf to deep-sea (bathyal) Distribution. Alarcón Rise, Gulf of California, species, so far recorded at depths of 249–510 Mexico, NE Pacific Ocean. m. Bathymetry. 2369 m depth. Remarks. This species has recently been described by Martynov et al. (2020a). Until Remarks. For this recently described deep-sea recently, only the single species D. robustus species there is no additional morphological was recognized among the wide-bodied, information and molecular data is very shallow-water dendronotids of the North limited. In particular, the dorsolateral Atlantic and the North Pacific shelf (Ekimova appendages need a further detailed study. et al.,2015).However, most recent data showed that “ D. robustus ” is a species complex, which Dendronotus comteti Valdés &amp; Bouchet, 1998 was already separated into D. robustus s.str., Dendronotus comteti Valdés and Bouchet, D. velifer and D. bathyvela (Lundin et al., 2017; 1998: 320–323, Figs 1–3. Martynov et al., 2020a). Diagnosis (original description). Body narrow. Dendronotus claguei Valdés, Lundsten &amp; Two or four pairs of partially branched, Wilson, 2018 partiallyunbranheddorsolateralappendages. Dendronotus claguei Valdés, Lundsten, Wilson, Four to six appendages of oral veil. Four 2018: 412–415, Figs 3B, 9. appendages (similar in size) of rhinophoral stalks. Rhinophores with seven to eight Diagnosis (original description). Body very lamellae.Lateral rhinophoral papilla possibly narrow. Four to six pairs of dorsolateral absent. Lip papillae probably absent. Basal appendages (reported as unbranched). Four colour uniformly pale cream (preserved). appendages of oral veil. Four appendages of Jaws reported as devoid denticles. Radula rhinophoral stalks (innermost appendages with up to 28 rows of teeth. Central tooth from each side longest). Lateral papilla of with deep furrows and with up to ten distinct rhinophoral sheaths absent. Rhinophores denticles. Up to six lateral teeth with up to with 11 lamellae. Lip papillae not reported in five denticles. The reproductive system was original description. Basal colour uniformly not studied in the original description. Body translucent white. Masticatory processes of length up to Downloaded 5. 5 mm from (preserved Brill.com). 12/12/2023 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/</p> <p>Distribution. Middle Atlantic, Lucky Strike area in the Mid-Atlantic Ridge.</p> <p>Bathymetry. 1685 m depth.</p> <p>Remarks. This is the only known species that originated from deep-sea hydrothermal habitats, and which was considered as belonging to the genus Dendronotus according to its external and internal characters. Molecular data is necessary to determine its phylogenetic position.</p> <p>denticles (may possess ridge-like structures). Radula with up to 51 rows of teeth. Central tooth completely smooth in adults. Up to 16 lateral teeth with up to 19 denticles. Ampulla relatively narrow with several (at least three) loops. Bursa copulatrix very large, oval. Seminal receptaculum small placed distally at a moderately short distance from the vaginal opening. Prostate discoid with up to 50 (and more) alveolar glands. The vas deferens is moderate in length, penis large, wide, with a blunt tip. Body length up to 14 cm.</p></div> 	https://treatment.plazi.org/id/1E1A4C68FF90FFE853ECFCA4FB86DE54	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	User, GBIF. Org	User, GBIF. Org (2021): A synoptic review of the family Dendronotidae (Mollusca: Nudibranchia): a multilevel organismal diversity approach. Contributions to Zoology 90 (1): 93-153, DOI: 10.1163/18759866-BJA10014, URL: http://dx.doi.org/10.15468/dl.ph6kj8
1E1A4C68FF92FFE351C9FD8FFE76D821.text	1E1A4C68FF92FFE351C9FD8FFE76D821.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Dendronotus dalli Bergh 1879	<div><p>Dendronotus dalli Bergh, 1879 Distribution. North Pacific (The Sea of Japan</p> <p>Fig. 7 in the west and at least Washington State, USA Dendronotus dalli Bergh, 1879: 94, Pl. 1, Fig. in the east to at least Chuckchii Sea; Russia, in 21; Pl. 2, Figs 9–12; Pl. 3, Figs 2–6; Robilliard, the Arctic (may possibly penetrate further to 1970: 452–455, Figs 13–15; McDonald, 2009: the west in the Arctic).</p> <p>463 (bibliography); Martynov &amp; Korshunova,</p> <p>2011: 155–157 (verified photographs of living Bathymetry. Verified records from 5 to 50 m specimens, description and biology in NW depth.</p> <p>Pacific); Ekimova et al., 2015: 860–863, Figs 6C,</p> <p>8G, 11, 12, 13A. Remarks. Until recently, the North Atlantic D. Dendronotus frondosus var. dalli Bergh, elegans (see below) has been confused with 1879 – Odhner, 1907: 19. the North Pacific D. dalli. The best external Not D.dalli sensu Bergh, 1886, Knipowitsch, feature to distinguish these two closely 1902, Roginskaya, 1987 (= D. elegans Verril, related species is the presence of an external 1880). opaque white pigment on the apical parts of the dorsolateral processes in D. dalli, whereas Extended diagnosis. Body relatively narrow. in D. elegans such a pigment, if present, is Four to eight pairs of branched dorsolateral internal. These species may co-occur in some appendages. Four to five appendages of oral Arctic localities, but this needs to be further veil. Four to 12 appendages (equal in size or investigated. For D.dalli an absence of tertiary few posterior ones longer) of rhinophoral branches of the dorsolateral appendages stalks. Lateral papilla of rhinophoral sheaths was incorrectly mentioned by Ekimova et al. present. Rhinophores with 16–33 lamellae. (2015).There is a possibility that light-coloured Lip papillae 15–40 (and more). Basal colour varieties of D. kamchaticus were depicted uniform translucent gray, light yellow to under D.dalli, judging from photographs in for orange, brownish or pinkish, with external example Behrens (1980) (see also Korshunova opaque white pigment on tips of dorsolateral et al., 2016a). In some studies, Dendronotus appendages. Dorsal processes of jaws inclined dalli was incorrectly considered a synonym of posteriorly at approximately 60° to the D. frondosus (Odhner, 1936; MacFarland, 1966) longitudinal axis of the jaw body and0.45 of its due to an underestimation of the importance length. Masticatory processes apparently bear of ontogenetic Downloaded radula from data Brill.com (central 12/12/2023 teeth 04:12 are: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/</p> <p>denticulated in juveniles and usually smooth the jaw body and 0.43of its length.Masticatory in adults). processes apparently bear denticles (which We have investigated a considerable num- may possess ridge-like structures). Radula ber of specimens and sequences of D. elegans with up to 42 rows of teeth. Central tooth from the North Atlantic and neighbouring completely smooth in adults. Up to at least subarctic regions. There are no confirmed 13 lateral teeth with up to eight denticles records of true D. dalli in the North Atlantic (can be more). Bursa copulatrix large, oval. and neighbouring regions. The indication of Seminal receptaculum small placed distally at D. dalli from the North Atlantic (Genbank a moderately short distance from the vaginal COI AF249800) in a table of the paper of opening. Prostate discoid with up to 60 and Wollscheid-Lengeling et al. (2001) is due to a more alveolar glands (their number was mix-up of localities, in this case with the North considerably underestimated in the original Pacific. The same can be said about Cadlina description of its new synonym D. niveus (see luteomarginata from the North Atlantic in remarks below). The vas deferens is moderate the same paper (Wollscheid-Lengeling et al., in length, penis strong, conical. Body length 2001) for which Johnson (2010: 142) explained up to 70 mm. that they originated from the North Pacific.</p> <p>Distribution. North Atlantic (known southern Dendronotus elegans Verrill, 1880 limit – Cape Cod, USA) to the Arctic</p> <p>Figs 7, 8 (including Barents Sea and White Sea, known Dendronotus elegans Verrill, 1880: 385–386. northeastern limit is the Laptev Sea, Russia). Dendronotus “elongatus”: Roginskaya, 1987: 175 (lapsus calami for D. elegans Verrill, 1880). Bathymetry. 10–25 m (White Sea) to 258 m Dendronotus niveus Ekimova, Korshunova, depth (western North Atlantic). Shepetov, Neretina, Sanamyan &amp; Martynov, 2015: 864–869, Figs 6D, 8F, 13B, 14, 15 syn. nov.; Remarks. Ekimova et al. (2015) overlooked Valdés et al., 2017: 6–7, 4B, 5B. important details in the original description Dendronotus dalli sensu Bergh, 1886 (non of D. elegans Verrill, 1880, for which a big Bergh, 1879); Robilliard, 1970: 452; Bleakney, animal with uniform salmon-coloured body 1996: 109 (D. elegans incorrectly mentioned as was used, and a special mentioning was synonym of D. dalli). made of the completely smooth central teeth</p> <p>(Verrill, 1880: 385–386). These characters are Extended diagnosis. Body relatively narrow. identical to those of the recently described Five to ten pairs of branched dorsolateral D. niveus Ekimova et al., 2015. The available appendages. Six to nine appendages of oral molecular data (Valdés et al., 2017; present veil. Four to five appendages (equal in size study, figs 1–3) also show that specimens or few posterior ones longer) of rhinophoral identified as D. niveus from off the east coast stalks. Lateral papilla of rhinophoral sheaths of North America (relatively close to the present. Rhinophores with 8–23 lamellae. Lip type locality of D. elegans) are identical to papillae 8–50. Basal colour uniform, pinkish specimens from the type locality of D. niveus to light orange, with internal opaque white in the White Sea. The holotype of D. elegans pigment of tips of dorsolateral appendages. in the Smithsonian Institution (NMNH, 2020) Dorsal processes of jaws inclined posteriorly at is not suitable for molecular study. However, approximately 40° to the longitudinal axis of a microslide Downloaded with radula from of Brill.com the holotype 12/12/2023 is 04:12:08PM</p> <p>via Open Access. This is an open access article distributed under the terms of the CC BY 4.0 License.</p> <p>https://creativecommons.org/licenses/by/4.0/ kept separately in the Yale Peabody Museum generally be described as highly branched,</p> <p>of Natural History (YPM IZ 010761.GP) and giving this species a fluffy appearance. Eight to available as online images (Yale Peabody 15appendages of oral veil.Five to six appendages Museum, 2020), which confirm the presence (posterior one very long) of rhinophoral stalks.</p> <p>of massive smooth central teeth. The date Lateral papilla of rhinophoral sheaths present.</p> <p>(6September1879), the depth (26fathoms =48 Rhinophores with 11–17 lamellae. Lip papillae meters) and locations (Stellwagen Bank, off 20–49.Basalcolourreddish-brownwithopaque Cape Cod) for collecting data of the holotype white stripes between dorsolateral processes to of D. elegans exactly match radula YPM IZ uniform light yellow with small brown spots.</p> <p>010761.GP, the preserved body USNM 842116 Dorsal processes of jaws inclined posteriorly and the original description of D. elegans at approximately 60° to the longitudinal in Verrill (1880). Additional morphological axis of the jaw body and 0.43 of its length.</p> <p>information of the molecularly investigated Masticatory processes apparently with fine specimens of D. elegans of the present study denticles. Radula with up to 37 rows of teeth.</p> <p>is given in fig. 8. It is evident that specimens Central tooth with up to 20 small denticles from the Atlantic North American coast without furrows, rarely completely smooth.</p> <p>and from the Arctic Eurasian regions are Up to 11 (usually 9–10) lateral teeth with up morphologically similar according to their to seven denticles. Ampulla with at least two external and internal features. We therefore loops. Bursa copulatrix large, rounded. Seminal synonymize Dendronotus niveus Ekimova et receptaculum small placed distally at a short al., 2015 with D. elegans Verrill, 1880. Another distance from the vaginal opening. Prostate species with smooth central teeth at the discoid with a range of 35–40 alveolar glands.</p> <p>adult stage is D. dalli Bergh, 1879 (see above, The vas deferens is very long, penis relatively figs 1, 2), but this one is found predominantly narrow, bent. Body length up to 100 mm.</p> <p>in the North Pacific and differs from D.</p> <p>elegans in habitus morphology (apical parts Distribution. NE Atlantic, so far known from of dorsolateral appendages with external the Netherlands, Norway, Sweden and the UK.</p> <p>opaque white pigment vs. with internal or absent white pigment in D. elegans) and Bathymetry. 15–150 m depth.</p> <p>molecular data.</p> <p>Remarks. This species has been separated Dendronotus europaeus Korshunova, from from D. frondosus and D. lacteus based on</p> <p>Martynov, Bakken &amp; Picton, 2017 both morphological and molecular evidence</p> <p>Fig. 7 (Korshunova et al., 2017b).</p> <p>Dendronotus europaeus Korshunova,</p> <p>Martynov, Bakken, Picton, 2017: 1–8, Figs. 1–2; Dendronotus frondosus (Ascanius, 1774) Supplementary information, text and figure. Fig. 7</p> <p>Dendronotus frondosus sensu Thompson Amphitrite frondosa Ascanius, 1774: 155–158,</p> <p>and Brown, 1984 (partim.); not Ascanius, 1774. Pl. 5, Fig. 2 (neotype selected in Ekimova et al.</p> <p>2015).</p> <p>Extended diagnosis. Body relatively narrow. Dendronotus frondosus (Ascanius, 1774) –</p> <p>Six to nine pairs of branched dorsolateral Odhner, 1936: 1105–1109, Fig. 39 (mixture of appendages, including two to three posterior several species); Robilliard, 1970: 441–446, ones. The dorsolateral appendages can Pl. 63, Fig. Downloaded 29, text and from Figs Brill.com 4, 7, 8 12,/9 12/ (2023 partim04:12,: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/ mostly referred to D. venustus); Ekimova et al. eight) lateral teeth with up to seven denticles.</p> <p>2015: 848–857, Figs 1–5, 6A, 7A, 8A. Ampulla voluminous, folded. Bursa copulatrix Doris arborescens Müller, 1776:229 (mixture large, oval to rounded. Seminal receptaculum of several species). small placed distally at a moderately short Doris cervina Gmelin, 1791: 3105, no. 12. distance from the vaginal opening. Prostate Tritonia reynoldsii Couthouy, 1838: 74–80, discoid with range 16–30 alveolar glands.</p> <p>Pl. 2, Figs 1, 2, 3, 4. The vas deferens is moderate in length, penis Tritonia felina Alder &amp; Hancock, 1842: 33. relatively long, curved. Common body length</p> <p>Tritonia pulchella Alder &amp; Hancock, 1842: no more than 50 mm.</p> <p>33–34.</p> <p>Tritonia ascanii MØller, 1842: 5. Distribution. North Atlantic, both east and Amphitridea fabricii Kröyer, 1847: 114. west parts, not distributed to the Arctic Campaspe pusilla Bergh, 1863: 471–478, Pl. further than the easternmost border of the 12, Figs 28–35. Barents Sea.</p> <p>Campaspe major Bergh, 1886: 21–24; Pl. 1,</p> <p>Figs. 23–26; Pl. 2, Figs 1 –11. Bathymetry. Intertidal to 20–30 m depth.</p> <p>Dendronotus luteolus Lafont, 1871: 267.</p> <p>Dendronotus junior Mörch, 1875: 125. Remarks. Three common North Atlantic Dendronotus arborescens var. aurantiaca shallow water species, namely D. europaeus Friele, 1879: 284. Korshunova et al., 2017b, D. frondosus NB: Mixture of several species under the (Ascanius, 1774), and D. lacteus, show an name Dendronotus frondosus in the literature extremely similar range of habitus variation before 2015. (see Korshunova et al., 2017b) and have been confused in older literature. Therefore, the Extended diagnosis. Body narrow. Five to six verified distribution of D. frondosus can only pairs of branched dorsolateral appendages. be assessed using recent data (Korshunova Four to seven appendages of oral veil. Four to et al., 2017b) that covers at least the North East five appendages (middle and posterior ones Atlantic from Norway and also the Barents can be longer) of rhinophoral stalks. Lateral Sea and the White Sea to possibly France / papilla of rhinophoral sheaths present. northern Spain. The range of D. frondosus Rhinophores with six to 12 lamellae. Lip does not extend further than the the true papillae four to 12. Basal colour brownish to Arctic. Also, several deep-water records of reddish-brown, often with small white and D. frondosus (e.g., Odhner, 1939; Swennen, yellow specks, but usually without opaque 1961; Thompson &amp; Brown, 1984) most likely white stripes between dorsolateral processes refer to D. lacteus, since according to our data to completely white translucent specimens. D. frondosus was usually found shallower Dorsal processes of jaws inclined posteriorly than 20–30 m depth. We have examined an at approximately 60° to the longitudinal extensive collection of various Dendronotus axis of the jaw body and 0.4 of its length. from the NW Pacific and NE Pacific and Masticatory processes apparently bear ridge- have so far not found any specimen from the like structures and denticles. Radula with up North Pacific for which the identity could to 42 rows of teeth. Central tooth with deep be confirmed as D. frondosus despite the furrows and with up to 14 (common range presence of a few published records (e.g., 8–12) distinct denticles. Up to 10 (usually up to Ekimova Downloaded et al., 2016 from Brill). In.com this12/12 respect /2023 04,:12 the: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/ common occurrence of true D. frondosus in 1–3) with stable distinct differences in exter-</p> <p>the North Atlantic and the extreme rarity nal and internal morphology (fig. 4) and of D. frondosus in the North Pacific, with without any gene flow between each other.</p> <p>the simultaneous common presence in Dendronotus kalikal thus does not occur the North Pacific of other species formerly naturally in the North Atlantic. A potential considered as “ D. frondosus ”, (e.g., D. kalikal, future introduction of some North Pacific D. kamchaticus, D. primorjensis, D. venustus), species from the “ D. frondosus megacomclearly suggests that true D. frondosus does plex” may take place, but so far there are not occur naturally in the North Pacific. no reliable records of any North Pacific spe- Here, it is largely substituted by other species cies in the North Atlantic. At least recently,</p> <p>from the D. frondosus species complex. In a case of evident anthropogenic transporparticularly, D. primorjensis endemic to the tation of the dendronotid Pseudobornella</p> <p>Sea of Japan, is morphologically similar to orientalis was reported from Japan or China D. frondosus and sister to it according to our to NE Pacific (Agarwal et al., 2017) (for taxmolecular phylogenetic analysis (fig. 1). D. onomy of Pseudobornella see below). Thus, primorjensis has several traceable diagnostic most previous records of this species from features in radula and reproductive system the Northwest Pacific refer to one of three</p> <p>(see Korshunova 2016a and Synopsis below). recently described species, D. kamchaticus</p> <p>By such features and molecular data D. Ekimova et al., 2015, D. kalikal Ekimova et primorjensis could be excellent candidate al., 2015, or D. primorjensis Martynov et al.,</p> <p>for an example of a relict species, with 2015, and in the Northeast Pacific, previous close relatives with ranges extending to the records refer to D. venustus Stout et al., 2010 Arctic, which afterwards formed a separate or D. kamchaticus Ekimova et al., 2015 (see species, D. frondosus, in the North Atlantic. also Korshunova et al., 2016a, b).</p> <p>As a result of such an evolutionary history, D.</p> <p>primorjensis and D. frondosus are evidently Dendronotus gracilis Baba, 1949</p> <p>naturally separated from from closely related Dendronotus gracilis Baba, 1949: 167, Pl. 35, North Pacific and North Atlantic species, Fig. 127, text-fig. 109; Robilliard, 1970: 461–462; because true D. frondosus does not occur in Nakano, 2018: 385.</p> <p>the Arctic. On the contrary, the potentially disrupted populations of D. frondosus in Diagnosis (original description). Body the North Atlantic and the North Pacific do narrow. Four pairs of branched dorsolateral not reveal morphological and molecular appendages. Four veil appendages. Five to differences. Most reliable explanations for six processeses of the rhinophoral stalks.</p> <p>such an occurrence of few of so far known Rhinophores with 18 lamellae. Basal colour specimens of D. frondosus in the North bluish-white, with numerous scattered Pacific can be an anthropogenic introduction yellow spots, various processes opaque white.</p> <p>or a mistake in the sorting or processing of Masticatory processes of jaws with denticles.</p> <p>collection material. Dendronotus frondosus Radula with up to 41 rows of teeth. Central does not occur naturally in the North Pacific. tooth with up to 20 small denticles without</p> <p>In this study we also evidently show that furrows, rarely completely smooth. Up to the North Pacific D. kalikal and the North eight lateral teeth with up to nine denticles.</p> <p>Atlantic and Arctic D. yrjargul sp. nov. are Reproductive system unknown. Body length invariably placed in two distinct clades (figs 25 mm (living Downloaded, original from description Brill.com 12) /. 12/2023 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/ Distribution. Japan, Pacific side of Honshu, Japan.</p> <p>transportation, which is hard to explain in this case and needs verification.</p> <p>Bathymetry. 160 m depth(original description). Dendronotus iris Cooper, 1863</p> <p>Fig. 7</p> <p>Remarks. This species is insufficiently known. Dendronotus iris Cooper, 1863: 59; Information on the internal morphology of MacFarland, 1966: 257–265, Pl. 47, Figs 12– specimens from the type locality is restricted 18; Plate 48, Figs 1–6; Pl. 49, Fig. 4; Pl. 50, Fig.</p> <p>to the original description (Baba, 1949). 1; Pl. 51, Figs 1–5; Robilliard, 1970: 446–450, Additional data in Pola and Stout (2008) Figs 4–6, 10–12, Pl. 63, Fig. 30; Behrens, 1980: referring to D. gracilis specimens are from 74; Behrens &amp; Hermosillo, 2005: 94.</p> <p>considerably remote locations off the tropical Dendronotus giganteus O’Donoghue, 1921: island of Okinawa, whereas the type locality 187–190; Pl. 4, Fig. 47; Pl. 5, Figs 57–59.</p> <p>of D. gracilis is in the temperate Sagami Bay.</p> <p>Their study lacks molecular data and may Extended diagnosis. Body relatively represent a separate species. The Okinawan narrow. Four to eight pairs of branched specimen indicated in Pola and Stout (2008) dorsolateral appendages. Two to four oral has eight pairs of dorsolateral processes, six veil appendages. Two to five appendages processess of the oral veil, and 12 rhinophoral (longest internal) of rhinophoral stalks.</p> <p>lamellae, whereas according to the original Lateral papilla of rhinophoral sheaths description in Baba (1949) there are four pairs present. Rhinophores with 15–31 lamellae.</p> <p>of dorsolateral processes, four processesses 20–40 lip papillae. Basal colour grayish to of the oral veil, and 18 rhinophoral lamellae, reddish-orange, with white line bordered respectively. Taking into consideration that foot edge. Dorsal processes of jaws inclined details of colour patterms between the original posteriorly at approximately 40° to the description of D. gracilis and Okinawan longitudinal axis of the jaw body and 0. 35 specimens are also different as indicated by of its length. Masticatory processes bear Pola and Stout (2008: 65), we cannot confirm denticles (possibly including ridge-like the conspecificity of the Okinawan specimens structures and denticles). Radula with up with the original D.gracilis, even if we consider to 61 rows of teeth. Central tooth quite the potentially cooler waters in Okinawa at 69 narrow, with deep furrows and with up to</p> <p>m depth and that only a few specimens were 18 distinct denticles. Up to 21 lateral teeth involved in the comparison. Furthermore, Pola commonly smooth, occasionally with up to and Stout (2008) considered specimens from eight denticles. Ampulla relatively narrow New Zealand as belonging to D. gracilis as well, with numerous loops. Bursa copulatrix which also show differences in colouration. moderate in size, oval, bent. Seminal However, in the light of a modern emerging receptaculum small, placed distally at a paradigm of multilevel organismal diversity moderately short distance from the vaginal that challenges the previously concept of opening. Prostate very long, non-discoid, widely claimed polytypic species with broad linear, with numerous alveolar glands. The ranges in favour of considerably smaller, often vas deferens is short, penis moderately geographically restricted species, a potential short, straight, with blunt tip. Body length presence of true D. gracilis in New Zealand of live specimens may reach more than can only be the result of an anthropogenic 20 cm. Downloaded from Brill.com 12/12/2023 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/ Distribution. NE Pacific, from southern Alaska to California (but records from Baja California need to be confirmed). As an adult it feeds on cerianthid anthozoans (Wobber, 1970; Shaw, 1991), quite different to other Dendronotus species.</p> <p>Bathymetry. 10–30 m depth.</p> <p>Remarks. This species is clearly distinguished by a combination of external characters (uniform grayish to reddish-orange colour, massive dorsal appendages) and internal characters (linear instead of discoid prostate – unique for the genus), and also by adult behavioral patterns (special “rearing attack” feeding on species of Ceriantharia, e.g., Shaw, 1991). Dendronotus nanus Marcus et Marcus, 1967 was considered a synonym of D. iris (Stout et al., 2010), but in the absence of molecular data and presence of morphological differences (see below) we prefer to retain D. nanus as a separate species (see details below). Dendronotus iris possibly is the largest species of its genus.</p> </div>	https://treatment.plazi.org/id/1E1A4C68FF92FFE351C9FD8FFE76D821	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	User, GBIF. Org	User, GBIF. Org (2021): A synoptic review of the family Dendronotidae (Mollusca: Nudibranchia): a multilevel organismal diversity approach. Contributions to Zoology 90 (1): 93-153, DOI: 10.1163/18759866-BJA10014, URL: http://dx.doi.org/10.15468/dl.ph6kj8
1E1A4C68FF9AFFDF51C9FA0EFED5DD7C.text	1E1A4C68FF9AFFDF51C9FA0EFED5DD7C.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Dendronotus kamchaticus Ekimova	<div><p>Dendronotus kamchaticus Ekimova, Remarks. Though this species previously was</p> <p>Korshunova, Shepetov, Neretina, Sanamyan confused with D. frondosus, D. albus and D.</p> <p>&amp; Martynov, 2015 venustus, D. kamchaticus is well delineated</p> <p>Fig. 7 from those species according to the</p> <p>Dendronotus kamchaticus Ekimova, morphological features (semitransparent</p> <p>Korshunova, Shepetov, Neretina, Sanamyan grayish Downloaded with from white-tipped Brill.com 12/12dorsolateral /2023 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/</p> <p>appendages and digestive gland often Dendronotus lacteus (Thompson, 1840) penetrating all branches of all dorsolateral Fig. 7</p> <p>appendages to body brownish colouration Tritonia lactea Thompson, 1840: 88–89, Pl.</p> <p>without dorsal subparallel stripes), radular 2, Fig. 3.</p> <p>patterns (usually almost smooth central teeth Dendronotus lacteus Eliot, 1910: 161;</p> <p>in adults) and molecular phylogenetic data Nordsieck, 1972: 68; Pruvot-Fol, 1954: 356;</p> <p>(figs1,2).In the original description branches of Thollesson, 1998: 191–193, Figs 1–3; Evertsen the dorsolateral appendages were incorrectly and Bakken, 2005: 18; Korshunova et al., 2017: described as bulbous and short (see Ekimova 2–7, Figs 1, 2.</p> <p>et al., 2015: Fig. 8D), despite the presence of not Becher, 1886: 14.</p> <p>this species in the same paper (Ekimova et al., not Eliot, 1910:112 (= D. dalli).</p> <p>2015: Fig. 16A, B) represented by photos of live = D. frondosus auct. non Ascanius, 1774.</p> <p>paratypes with more elongated, not exactly bulbous, though still shortened branches. Extended diagnosis. Body relatively narrow.</p> <p>Further, in a specimen of D. kamchaticus from Four to eight pairs of branched dorsolateral the Sea of Japan the branches were described appendages which in adults commonly show as elongated and also very different from the specific “curly” pattern though other specimens original description and Fig. 8D in Ekimova with more “typical” for genus Dendronotus (2015). This significant discrepancy compared elongated appendages may occur. Four to ten with the original description was explained: oral veil appendages. Four to six appendages “because different ontogenetic stages were (inetrnal middle and posterior ones longest)</p> <p>used in the description and this study, as of rhinophoral stalks. Lateral papilla of well as cerata morphology may be altered in rhinophoral sheaths present. Rhinophores preserved specimens” (Ekimova et al., 2016: with 10–14 lamellae. At least more than 15 lip</p> <p>38). However, this discrepancy was not due papillae.Basalcolourfromuniformlydarkredto to the different ontogenetic stages. In the uniform white, often with small brownish dots.</p> <p>original description of D. kamchaticus, the Redddish and brownish specimens may have type specimens were represented by an almost white stripes between cerata, white specimens adult holotype (12 mm preserved length) milky, non-transparent in appearance. Dorsal with a mature reproductive system whereas processes of jaws inclined posteriorly at seven paratypes were subadults (2–10 mm approximately 55–60° to the longitudinal of preserved length). Most importantly, the axis of the jaw body and 0.52 of its length.</p> <p>drawing in Ekimova(2015: Fig.8D)depicted the Masticatory processes possibly bear denticles largest holotype, with an adult reproductive and ridge-like structures. Radula with up to 43 system and the artificially bulbous branches rows of teeth. Central tooth with up to 40–50</p> <p>were only seen in the preserved specimen, small denticles (commonly smaller), often without any considerable ontogenetic with peculiar ribs, never completely smooth.</p> <p>modifications. The largest specimen so far Up to 16 lateral teeth with up to five denticles.</p> <p>known of D. kamchatcius (adult, 30 mm) Ampulla moderately narrow with several from the NE Pacific, also has considerably windings or folded loops. Bursa copulatrix elongated branches (Korshunova et al., large, rounded to oval. Seminal receptaculum 2016a). This fully confirms that the branches small placed distally at considerable distance were described and figured incorrectly in the from the vaginal opening. Prostate discoid with original description. a range from Downloaded 12 to more from than Brill30.com alveolar 12/12/2023 glands04:12.: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/</p> <p>The vas deferens is relatively long in length, Ampulla was not described in the original penis relatively long, bent. Body length up to description. Bursa copulatrix large, rounded 150 mm. to oval. Seminal receptaculum small placed distally at considerable distance from the</p> <p>Distribution. North Atlantic and Arctic vaginal opening. Glandular part of prostate (eastern limit at least Laptev Sea, Russia). was reported in the original description as in early development of differentiation.</p> <p>Bathymetry. From circa 5 to 427 m depth; The vas deferens is extremely long, with common at shallow, subtidal depths at around numerous loops, penis moderately short, 10–20 m. straight, with blunt tip. Body length up to</p> <p>13 mm.</p> <p>Remarks. Dendronotus lacteus together with</p> <p>D. frondosus and D. europaeus form a species Distribution. NE Pacific, Baja California, complex that is difficult to distinguish Puerto Peñasco, Sonora, Mexico.</p> <p>based on external characteristics, but in which it is still possible to find fine-scale Bathymetry. Floating on seawater surface. morphological differences. In contrast, the radulae of these three species demonstrate Remarks. Dendronotus nanus was considered distinctly different patterns (Korshunova et a synonym of D. iris by Stout et al. (2010), but al., 2017b). in the absence of molecular data we prefer to keep them separate. Furthermore, despite the</p> <p>Dendronotus nanus Marcus &amp; Marcus, 1967 fact that Stout et al. (2010) mentioned only</p> <p>Dendronotus nanus Marcus &amp; Marcus, minor morphological differences between</p> <p>1967: 210–214, Figs 64–65; Farmer, 1980: 72; D. iris and D. nanus, Marcus and Marcus Robilliard, 1972: 428–430. (1967) in their original description specially highlighted that D. nanus differs from D. iris</p> <p>Diagnosis (original description). Body by a considerably longer vas deferens in narrow. Six pairs of branched dorsolateral animals of only 13 mm length (an considered appendages. Circa six oral veil appendages. to be not fully mature), whereas D. iris that Five appendages (posterior ones the longest) was studied anatomically by Robilliard of rhinophoral stalks. Lateral papilla of (1970) had a considerably larger average body rhinophoral sheaths present. Rhinophores length 65–120 mm but a significantly shorter with about 15 lamellae. Circa 20 lip papillae. vas deferens. In the BOLD database there is a Basal colour semitransparent grayish with sequence of D. nanus (in GenBank identified a slight reddish hue and a little brown as D. iris), but from a very remote location in pigment, with white line bordered foot British Columbia whereas the type locality edge, white tipped dorsolateral appendages of D. nanus is Baja California, Sonora, thus with blackish brown subapical rings and an this available sequence does not represent orange base. Masticatory processeses with a true D. nanus. Even if taken into account crested pointed denticles. Radula with up to that the glandular part of vas deferens 26 rows of teeth. Central tooth quite narrow, likely is underdeveloped in D. nanus (see with deep furrows and with up to 10 distinct Marcus &amp; Marcus, 1967), the non-glandular denticles. Up to 14 lateral teeth commonly muscular part of vas deference of D. nanus is smooth, occasionally with up to 12 denticles. considerably Downloaded longer from than Brill.com in D. iris. 12/12/2023 Robilliard 04:12:08PM via Open Access. This is an open access article distributed under the terms of the CC BY 4.0 License.</p> <p>https://creativecommons.org/licenses/by/4.0/</p> <p>(1972) compared D. iris and D. nanus in detail and mentioned similarity of both species but did not synonymize them. All these characters need to be re-analysed before any further attempts at synonymizing these species should be done.</p> </div>	https://treatment.plazi.org/id/1E1A4C68FF9AFFDF51C9FA0EFED5DD7C	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	User, GBIF. Org	User, GBIF. Org (2021): A synoptic review of the family Dendronotidae (Mollusca: Nudibranchia): a multilevel organismal diversity approach. Contributions to Zoology 90 (1): 93-153, DOI: 10.1163/18759866-BJA10014, URL: http://dx.doi.org/10.15468/dl.ph6kj8
1E1A4C68FFA5FFDF512AFE57FF3DDA1F.text	1E1A4C68FFA5FFDF512AFE57FF3DDA1F.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Dendronotus patricki Stout, Wilson	<div><p>Dendronotus patricki Stout, Wilson &amp;</p> <p>Valdés, 2011</p> <p>Dendronotus patricki Stout et al., 2011: 63–67, Figs 2–4.</p> <p>Diagnosis (original description). Body broad. Three to four pairs of branched dorsolateral appendages. Six appendages of oral veil. Four appendages (posterior ones long) of rhinophoral stalks. Lateral papilla of rhinophoral sheaths absent. Number of rhinophoral lamellae and lip papillae not indicated in the original description. Colour uniformly reddish brown (appears in original description as rather pinkish) with a semitransparent body. Masticatory processeses with ridge-like structures. Radula with up to 31 rows of teeth. Central tooth with up to 20 small denticles without furrows. Up to eight lateral teeth without denticles. Ampulla moderately voluminous with two folded loops. Bursa copulatrix very large, rounded to oval. Seminal receptaculum moderate in size placed distally near the vaginal opening. Prostate discoid with range at least 10 alveolar glands. The vas deferens is very long, penis relatively long, straight, conical. Body length up to 25 mm.</p> <p>Distribution. Whalefall ‘Patrick’ area (36º42’ 30.31’’ N, 122º 06’ 18.72’’W), Monterey Canyon, USA.</p></div> 	https://treatment.plazi.org/id/1E1A4C68FFA5FFDF512AFE57FF3DDA1F	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	User, GBIF. Org	User, GBIF. Org (2021): A synoptic review of the family Dendronotidae (Mollusca: Nudibranchia): a multilevel organismal diversity approach. Contributions to Zoology 90 (1): 93-153, DOI: 10.1163/18759866-BJA10014, URL: http://dx.doi.org/10.15468/dl.ph6kj8
1E1A4C68FFA5FFDB53CDFF5AFEFBDCEC.text	1E1A4C68FFA5FFDB53CDFF5AFEFBDCEC.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Dendronotus primorjensis Martynov	<div><p>Dendronotus primorjensis Martynov,</p> <p>Sanamyan &amp; Korshunova, 2015</p> <p>Fig. 7</p> <p>Dendronotus primorjensis Martynov et al., 2015a: 60, Fig. 5A–G; Martynov et al., 2015c: 75–76; Korshunova et al., 2016b: 15–28, Figs 1, 2; Korshunova et al., 2016a: 20–32; Korshunova et al., 2019b: 6–8, Fig. 3A–F.</p> <p>Dendronotus dudkai Ekimova et al. (2016): 33–38, Figs 9–11 (synonym of D. primorjensis).</p> <p>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.</p> <p>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</p> <p>b). via Open Access. This is an open access article distributed under the terms of the CC BY 4.0 License.</p> <p>https://creativecommons.org/licenses/by/4.0/</p> <p>Bathymetry. 3–20 m depth. teeth usually have more distinct denticles. We therefore retain the North Pacific D. purpureus</p> <p>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 &amp; Martynov, 2016</p> <p>Fig. 7</p> <p>Dendronotus purpureus Bergh, 1879 Dendronotus robilliardi Korshunova, Dendronotus purpureus Bergh, 1879: 89–93, Pl. Sanamyan, Zimina, Fletcher &amp; Martynov, I, Figs 18–20; Pl. 3, Figs 7 –12.; Bergh, 1904:15; 2016: 28–32, Figs 2, 3, 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</p> <p>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</p> <p>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</p> <p>Bathymetry. 31 m depth. vaginal opening. Prostate discoid with about</p> <p>19–30 alveolar glands. The vas deferens is</p> <p>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</p> <p>via Open Access. This is an open access article distributed under the terms of the CC BY 4.0 License.</p> <p>https://creativecommons.org/licenses/by/4.0/ and USA). Northern Japan (photographic record only, Korshunova et al., 2019b).</p> <p>Depth. circa 5–30 m. range does not extend further eastward than the Barents Sea.</p> <p>Bathymetry. Intertidal to less than 20 m deep.</p> <p>Remarks. See Remarks for D. albus. Remarks. T.A. Korshunova’s and A.V.</p> <p>Martynov’s pre-2015 analysis has showed Dendronotus robustus Verrill, 1870 that there are at least two species in the “ D.</p> <p>Fig. 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, 4. this analysis was unfortunately not included</p> <p>Non auct., not Ekimova et al. (2015) (mix-up in the final version of the publication by I.A.</p> <p>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,</p> <p>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.</p> <p>20 lip papillae. Basal colour light grayish to yellowish with scattered opaque white Dendronotus rufus O’Donoghue, 1921</p> <p>and yellow spots. Dorsal processes of jaws Fig. 7</p> <p>inclined posteriorly at approximately 47° Dendronotus rufus O’Donoghue, 1921:190 –</p> <p>to the longitudinal axis of the jaw body and 192; Pl. 3, Figs 25–27; Pl. 4, Fig. 48; O’Donoghue,</p> <p>0.55 of its length. Masticatory processes 1922:125; Robilliard, 1970: 456–460, Figs 4–6, apparently bear denticles (may possess ridge- 16–18, Pl. 63, Fig. 32.</p> <p>like structures). Radula with up to 31 rows of Behrens, 1980:76; Behrens &amp; Hermosillo, teeth. Central tooth with high cusp and up to 2005:95; Lamb &amp; Hanby, 2005: 262.</p> <p>20 small distinct denticles without furrows. Dendronotus frondosus sensu Odhner, 1936</p> <p>Up to nine lateral teeth with up to seven (part.) non Ascanius, 1774.</p> <p>denticles. Ampulla moderately narrow, with approximately three to four irregular loops. Extended diagnosis. Body relatively narrow.</p> <p>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.</p> <p>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</p> <p>Fig. 7 on the rhinophoral sheath. However, Dendronotus subramosus MacFarland, phylogenetically D. subramosus is more 1966: 265–270, Pl. 40, Fig. 3; Pl. 46, Figs 5–8; Pl. closely related to the externally similar- 47, Figs 3–7; Pl. 49, Figs 1–3; Pl. 50, Fig. 2; Pl. 52, looking deep sea species D. jamsteci Figs 1, 2; Robilliard, 1970: 462–466, Figs 3, 4, 6, (with the presence of rhinophoral sheath 19–21, Pl. 64, Fig. 33; Behrens, 1980:76; Behrens papillae), which was recently described &amp; 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</p> <p>via Open Access. This is an open access article distributed under the terms of the CC BY 4.0 License.</p> <p>https://creativecommons.org/licenses/by/4.0/ fulfilling of the term “mutlilevel fine-scale organismal diversity”.</p></div> 	https://treatment.plazi.org/id/1E1A4C68FFA5FFDB53CDFF5AFEFBDCEC	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	User, GBIF. Org	User, GBIF. Org (2021): A synoptic review of the family Dendronotidae (Mollusca: Nudibranchia): a multilevel organismal diversity approach. Contributions to Zoology 90 (1): 93-153, DOI: 10.1163/18759866-BJA10014, URL: http://dx.doi.org/10.15468/dl.ph6kj8
1E1A4C68FFA1FFDB512AFEE6FDA0DAAF.text	1E1A4C68FFA1FFDB512AFEE6FDA0DAAF.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Dendronotus velifer G. O. Sars 1878	<div><p>Dendronotus velifer G.O. Sars, 1878</p> <p>Fig. 7</p> <p>Dendronotus velifer G.O. Sars, 1878:238–239, Pl. 15, Fig. 4, Pl. 28, Fig. 2; Lundin et al., 2017: 303–309, 310–318, Figs 1, 2, 4.</p> <p>Non D. velifer auct. (mix-up with D.robustus and D. bathyvela).</p> <p>Extended diagnosis. Body wide. Four to six pairs of branched dorsolateral appendages. Six to twelve main appendages on oral veil. Four to five appendages on rhinophoral stalks. Lateral papilla of rhinophoral sheaths absent or very small.Rhinophores with 12–15lamellae. 30–35 lip papillae. Basal colour commonly light-to bright red with scattered white dots. Dorsal processes of jaws inclined posteriorly at approximately 65° to the longitudinal axis of the jaw body and 0.55 of its length. Masticatory processes with denticles. Radula with up to 36 rows of teeth. Central tooth with low to moderate cusp and up to 26 small distinct denticles without furrows. Up to 15 lateral teeth with which is smooth or with up to four rudimentary denticles. Ampulla twolooped. Bursa copulatrix large, rounded, with small seminal receptaculum placed distally at some distance from the vaginal opening. Prostate discoid with more than 50 alveolar glands. The vas deferens is moderate in length, penis thin, long. Body length up to 90 mm.</p> <p>Distribution. North Atlantic (southern limit – Gullmarfjord, Sweden) and Arctic (at least to the Laptev Sea), Norway and Russia.</p> <p>Bathymetry. Commonly found below 50 m depth, in the Arctic not shallower than 10 m.</p> <p>Remarks. See Remarks for D. robustus.</p> </div>	https://treatment.plazi.org/id/1E1A4C68FFA1FFDB512AFEE6FDA0DAAF	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	User, GBIF. Org	User, GBIF. Org (2021): A synoptic review of the family Dendronotidae (Mollusca: Nudibranchia): a multilevel organismal diversity approach. Contributions to Zoology 90 (1): 93-153, DOI: 10.1163/18759866-BJA10014, URL: http://dx.doi.org/10.15468/dl.ph6kj8
1E1A4C68FFA1FFD853CDFF5AFC8DD91B.text	1E1A4C68FFA1FFD853CDFF5AFC8DD91B.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Dendronotus venustus MacFarland 1966	<div><p>Dendronotus venustus MacFarland, 1966</p> <p>Fig. 7</p> <p>Dendronotus venustus MacFarland, 1966: 271– 275, Pl. 40, Fig. 2; Pl. 46, Figs 9–12; Pl 47, Figs 1, 2; Pl. 49, Fig. 6; Pl. 50, Fig. 3; Pl. 52; Figs 3–6; Stout et al. 2010: 7.</p> <p>D. frondosus sensu Robilliard, 1970 and auctorum non Ascanius, 1774.</p> <p>Diagnosis. Body narrow. Four to eight pairs of branched dorsolateral appendages. Four to eight appendages of oral veil. Four to six appendages (posterior and medial ones longest) of rhinophoral stalks. Lateral papilla of rhinophoral sheaths present. Rhinophores with 8–14 lamellae. Lip papillae absent or up to eight. Colour forms include white specimens with yellow spots and brownish or reddish specimens with yellow dots and/or tubercles or large amount of white pigment, but with no white-tipped appendages. Dorsal processes of jaws inclined posteriorly at approximately 55–60° to the longitudinal axis of the jaw body and 0.41 of its length. Masticatory processess with ridge-like structures and smaller denticles. Radula with up to 48 rows of teeth. Central tooth with up to 12 small distinct denticles with furrows. Up to eight lateral teeth with up to five denticles. Ampulla voluminous, folded. Bursa copulatrix large, pear-shaped. Seminal receptaculum small placed distally at considerable distance from the vaginal opening. Prostate discoid with few (5–12) alveolar glands. The vas deferens is long, possibly also partly prostatic since thickened on considerable length and then distally narrowed, penis thin, long. Body length up to 42 mm.</p> <p>Distribution. NE Pacific, Canada and USA.</p> <p>Bathymetry. Common in shallow waters, at depths of circa 5–20 m.</p> <p>Remarks. Despite MacFarland’s (1966) detailed description of D. venustus, it was almost universally considered a synonym of D. frondosus (Robilliard, 1970; McDonald, 1980; Behrens &amp; Hermosillo, 2005) until molecular data showed that it is distinct from D. frondosus (Stout et al., 2010; Korshunova et al., 2019b). Robilliard (1970) did not doubt the synonymy of D. venustus, although he rarely observed more than 12 of the prostatic alveolar glands in the NE Pacific specimens, whereas in true D. frondosus their number is usually up to at least 30. There are also minor differences in the radular patterns and some details of colouration between D. frondosus and D. venustus (e.g., common presence of white forms with small yellow spots in D. venustus). There is a possibility of the presence of hidden diversity within D. venustus. However, a specimen from Alaska included in our present study (fig. 7) matches the original description of D.venustus very well by external and internal features (MacFarland, 1966: 271–275, pl. 40 fig. 2, pl. 46, fig. 9, pl. 50 fig. 3). These characters include the presence of yellow-greenish spots on a greyish to whitish body, a specifically broadened, somewhat cheliform first lateral tooth, and a long, broadened, apparently partly prostatic vas deferens in addition to a small discoid alveolar prostate. We therefore surely refer to this specimen as a true D. venustus in our present phylogenetic analysis (fig. 1).</p> <p>Five to seven appendages of oral veil. Circa five appendages (posterior ones the longest) of rhinophoral stalks. Lateral papilla of rhinophoral sheaths present. Rhinophores with 10–11 lamellae. Lip papillae 20–30. Basal 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. Dorsal processes of jaws inclined posteriorly at approximately 70° to the longitudinal axis of the jaw body and 0.5 of its length. Masticatory processess with ridge-like structures. Radula with up to 36 rows of teeth. Central tooth completely devoid of denticles (except for the anteriormost juvenile radula in some specimens), only sometimes faint traces of furrows occur. Up to 12 lateral teeth with up to seven denticles. Ampulla moderately narrow, sligthly bent. Bursa copulatrix very large, oval. Seminal receptaculum small placed distally at considerable distance from the vaginal opening. Prostate discoid with circa 25 alveolar glands. The vas deferens is moderate in length, penis conical, curved. Body length up to 30 mm.</p> <p>Distribution. NW Pacific, so far recorded from Kamchatka, Russia, southern part of Hokkaido and Honshu, Japan (the Sea of Japan side).</p></div> 	https://treatment.plazi.org/id/1E1A4C68FFA1FFD853CDFF5AFC8DD91B	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	User, GBIF. Org	User, GBIF. Org (2021): A synoptic review of the family Dendronotidae (Mollusca: Nudibranchia): a multilevel organismal diversity approach. Contributions to Zoology 90 (1): 93-153, DOI: 10.1163/18759866-BJA10014, URL: http://dx.doi.org/10.15468/dl.ph6kj8
1E1A4C68FFA2FFCB51C9FABDFC35DE9D.text	1E1A4C68FFA2FFCB51C9FABDFC35DE9D.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Dendronotus zakuro Martynov, Fujiwara, Bathymetry. Shallow	<div><p>Dendronotus zakuro Martynov, Fujiwara, Bathymetry. Shallow waters, at depths of circa</p> <p>Tsuchida, R. Nakano, N. Sanamyan, 7–20 m.</p> <p>K. Sanamyan, Fletcher &amp; Korshunova, 2020</p> <p>Fig. 7 Remarks. This species has recently been</p> <p>Dendronotus zakuro Martynov, Fujiwara, described in detail (Martynov et al., 2020a).</p> <p>Tsuchida, Nakano, Sanamyan, Sanamyan,</p> <p>Fletcher, Korshunova, 2020a: 505–507, Figs 3, Genus Cabangus gen. nov.</p> <p>5B. ZooBank: http:// urn:lsid:zoobank.org:act: 3B779CBF-A310-4B47-85B1-488104FAD541</p> <p>Extended diagnosis. Body narrow. Six to seven Type species. Dendronotus regius Pola &amp;</p> <p>pairs of branched dorsolateral appendages. Stout, 2008 Downloaded from Brill.com 12/12/2023 04:12:08 PM 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 /</p> <p>Etymology. From the Indonesian word denticle-bearing part of the central teeth is not “cabang” meaning “branch” in reference to separated from the lateral sides by a distinct this genus as “dendronotids of the tropics” shoulder.Thus the central teeth have a squarish and to respect the great contribution of base and a triangular top. This makes the the Indonesian fauna to global marine radula of both species more similar (though biodiversity (e.g., Hoeksema, 2007). not identical) to that of the family Bornellidae, whose species are also partly similar in external Diagnosis. Dorsolateral appendages with morphology. However, details of the stomach distinct tertiary branches. Oral veil with and the reproductive systems are different.</p> <p>branched appendages. Radula with at least Therefore, taking molecular and morphological up to nine lateral teeth in adult specimens. evidence into account, we propose here the Central teeth with cusp integrated within new genus Cabangus gen. nov. with as type lateral denticles. Denticle-bearing part of species Dendronotus regius Pola &amp; Stout, 2008.</p> <p>central teeth not separated from lateral sides, The holotype of the only other tropical species distinct shoulders absent, thus central teeth described so far, D. noahi, is likely a juvenile triangular in outline. Prostate represented by since it possesses only four rows of lateral teeth a thickened structure without evident alveolar and immature reproductive system. Although glands and disk. Copulative organ conical. this species is now included in the genus Cabangus gen. nov., this needs further study.</p> <p>Remarks. According to the present molecular Externally, Cabangus spp. are different from the phylogenetic analysis, the tropical Dendronotus common external appearance of Dendronotus regius Pola &amp; Stout, 2008 comes as a most basal spp. by a combination of a very narrow clade, distinct from all presently known species body and short dorsolateral appendages. By of the genera Dendronotus and Pseudobornella. these characters, Cabangus gen. nov. is also Morphologically, D. regius differs from any somewhat similar to the family Bornellidae, known species of the genus by the presence which is phylogenetically distant from the of a homogenous, fine prostate, which is Dendronotidae. Cabangus gen. nov. represents externally smooth and does not show any a basalmost clade within the Dendronotidae alveoles. Pola and Stout (2008: Fig. 4) evidently and hence may retain some features of the did not indicate any alveols in the prostate common ancestors with phylogenetically more of their D. regius, but in the text they (Pola &amp; distantly related families. Cabangus gen. nov.</p> <p>Stout, 2008: 48) mentioned a “large prostate is also well supported by biogeographic data with the proximal limit being marked by a because the majority of its species inhabits closely set ring of alveolar glands”. To clear this Arctic and temperate waters,whereas Cabangus contradiction between this figure and the text gen. nov. represents a distinct tropical lineage.</p> <p>in the original description, we checked several adult specimens of D. regius and confirm Cabangus noahi (Pola &amp; Stout, 2008) comb.</p> <p>that there are no externally or internally nov.</p> <p>conspicous alveols, otherwise so characteristic Dendronotus noahi Pola &amp; Stout, 2008: 55–63,</p> <p>for a majority of true Dendronotus species. figs 6A, B.</p> <p>There is another striking difference between both strictly tropical species of Dendronotus Diagnosis (original description). Body very</p> <p>(D. regius plus D. noahi) and any other narrow. Four pairs of branched dorsolateral known species of this genus, namely that the appendages Downloaded. Six appendages from Brill.com of oral 12/12 veil /2023. Four 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/ appendages(posterior longest) of rhinophoral stalks. Small lateral papilla of rhinophoral sheaths possibly present. Rhinophores with nine lamellae. Lip papillae not indicated in the original description. Colour semitransparent white with brownish branches of digestive gland. Masticatory processes with denticles. Radula with up to 18 rows of teeth. Central tooth quite narrow, with shallow furrows and with up to 15 distinct denticles. Up to four lateral teeth with up to eight denticles. Reproductive system reported as immature in the first description. Body length up to 4 mm (juvenile specimen).</p> <p>seven denticles. Ampulla voluminous, folded. Bursa copulatrix large, pear-shaped to oval. Seminal receptaculum small placed distally at a short distance from the vaginal opening (in the original description entangled duct from ampulla was incorrectly identified as “receptaculum seminis”). Prostate non discoid, thickened, without evident alveolar glands.The vas deferens is moderate in length, penis tapered. Body length up to 15 mm.</p> <p>Distribution. Tropical Indo-west Pacific.</p> <p>Bathymetry. Up to 45 m deep.</p> <p>Distribution. Papua New Guinea, north coast, Remarks. See under Cabangus gen. nov. outer barrier reef, Bagabag Island, Bismarck Sea. Genus Pseudobornella Baba, 1932</p> <p>Type species. P. orientalis Baba, 1932 (fig. 2) Bathymetry. 30. 5 m depth.</p> <p>Diagnosis. Dorsolateral appendages without Remarks. Molecular data unavailable, as well distinct (or with few very short) tertiary as additional specimens. Species is in need of branches. One of the rhinophoral stalk further study. appendages from both sides is extremely long,</p> <p>usually exceeding body length. Oral veil with Cabangus regius (Pola &amp; Stout, 2008) comb. long simple unbranched appendages. Radula nov. with very small number of lateral teeth Fig. 2 (presently known to be no more than two rows Dendronotus regius Pola &amp; Stout, 2008: 46 – of lateral teeth). Central teeth with protruding</p> <p>54, Figs 1–5. cusp distinctly separate from lateral denticles.</p> <p>Prostate represents by narrow tube, without Extended diagnosis. Body very narrow. alveols and disk. Copulative organ partly Three to four pairs of branched dorsolateral flattened and with a widened subcircular appendages. Circa six appendages of oral apical part. veil. Four to five appendages (posterior ones longest) of rhinophoral stalks. Rhinophores Remarks. The genus Pseudobornella differs from with 10–11 lamellae. Lateral papilla of both Dendronotus and Cabangus gen. nov. by rhinophoral sheaths present. Lip papillae a unique combination of external and internal possibly absent. Basal colour white or yellow, characters.These characters include a very long with blue, brown and reddish patches. appendage of the rhinophoral sheath, absence Masticatory processes with denticles. Radula of distinct tertiary branches of the dorsolateral with up to 36 rows of teeth. Central tooth appendages, resulting in the general appearance with up to 20 small distinct denticles with of dorsolateral appendages of Pseudobornella as furrows. Up to nine lateral teeth with up to ctenidium-like Downloaded. Instead from, all Brill.com species of 12/12/2023 the genus 04:12:08PM</p> <p>via Open Access. This is an open access article distributed under the terms of the CC BY 4.0 License.</p> <p>https://creativecommons.org/licenses/by/4.0/ Dendronotus possess well-defined tertiary coupled with the molecular data (including branchesonthedorsolateralappendages,andno presenting for the first time data on COI gene species has such a exceedingly long appendages for the type species Pseudobornella orientalis,</p> <p>of the rhinophoral sheaths. There is a deep-sea figs 1, 2) show that the genus Pseudobornella species, Dendronotus claugei (see above), for represents an early offshoot of the family which unbranched dorsolateral appendages are Dendronotidae. This evolution was fuelled reported, but this species is only known from by the paedomorphic reduction of the lateral a single specimen and needs further studies. teeth and juvenilization of the central teeth According to the present phylogeny D. claugei (for criteria of paedomorphosis (Korshunova may represent a separate genus, but more data et al., 2020a; Martynov et al., 2020b). A pre-</p> <p>are needed to support this. Furthermore, so far vious morphological cladistic study (Pola et no single species of the genus Dendronotus and al., 2009) placed the genus Pseudobornella Cabangus gen. nov. have smooth unbranched outside the family Dendronotidae, thus appendages in the oral veil, whereas numerous highligthing the morphological peculiarities reported Pseudobornella specimens invariably of this genus as inconsistent with those of show strong unbranched appendages of the other dendronotid genera. However, a solely oral veil. Ultimately, the radular pattern of molecular study suggested to synonymize the genus Pseudobornella differs from that in Pseudobornella with Dendronotus (Pola &amp;</p> <p>the adult stage of species of Dendronotus and Gosliner, 2010). We confirm here the validity Cabangus gen. nov. by a very small number of of the genus Pseudobornella Baba, 1932 using lateral teeth (so far reported no more than two). integrative evidences. Specimens of the type Adult specimens of the genera Dendronotus species Pseudobornella orientalis from the Sea and Cabangus gen. nov. possess at least eight of Japan (fig. 2) in the present study matched lateral teeth, and only very rarely in poorly morphologically well with those in the origknown deep-sea lineages, the number of the inal description of P. orientalis (Baba, 1932), lateral teeth rows can reach six in number. The including the characteristic small chocotropical C. noahi was reported to have only four late-brown spots and yellow lines on the dor-</p> <p>rows of lateral teeth, but only a single juvenile sal side, and the shape of the radular teeth.</p> <p>specimen is known (Pola &amp; Stout, 2008). All investigated species of the genus Dendronotus Species composition. This presently monoshowed the presence of 1–3 rows of lateral teeth specific genus only contains P.orientalis Baba, during the early stages of their ontogeny (e.g., 1932.</p> <p>Martynov et al., 2020a), while only in the genus</p> <p>Pseudobornella such feature became apparent in adults stage. Furthermore, the shape of the Discussion central teeth of Pseudobornella is also different from that in any adult Dendronotus and General taxonomic and biogeographic</p> <p>Cabangus gen. nov. species by the presence of a overview of the family Dendronotidae:</p> <p>strongly protruding central cusp on the central a revolution of fine-scale defining of species teeth, which is distinctly separated from the The family Dendronotidae and its type genus lateral denticles. Dendronotus represent an emerging model ASEMimageoftheradulaof P.orientalis from for studying speciation, and ultimately for the Sea of Japan was presented in Martynov investigating general problems related to et al. (2015a: 60, fig. 5G). Radular characters the “species Downloaded concept from ”. The Brill.com total12number /12/2023 04of: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/ species belonging to the genus Dendronotus Dendronotus europaeus inhabits the north-</p> <p>in restricted sense is 27 (a review of all the erneastern Atlantic in European waters only.</p> <p>previously described and species consid- Dendronotus arcticus is reported only from ered valid with diagnoses is presented in the the true Arctic, e.g., Kara and Laptev seas.</p> <p>Synopsis section). Though some new spe- Dendronotus lacteus occurs in northeastern cies were recognized using morphological Atlantic and Arctic with the Laptev Sea as methods (e.g., MacFarland, 1966; Robilliard, easternmost limit. Dendronotus yrjargul sp.</p> <p>1970, 1972), the species composition of the nov. inhabits mostly the Arctic (with the Kara genus Dendronotus remained quite conserv- Sea as easternmost limit) and the neighbourative (Thompson &amp; Brown, 1984; Roginskaya ing northern European regions with middle 1987, 1997). The genus Dendronotus was Norway as southernmost limit. Dendronotus represented by just about 10 recognised nordenskioeldi sp. nov. is so far only reported shallow-water species worldwide (plus one from the Arctic Laptev sea. Dendronotus deep-sea and two or three species of unclear kalikal is distributed in the northwestern status) until a considerable amount of hid- Pacific until the Bering Strait. Dendronotus den diversity was discovered recently in the kamchaticus is limited to the Northern Pacific shallow waters of both the North Pacific and only, with scattered records from the norththe North Atlantic (e.g., Korshunova et al., western and northeastern parts. Dendronotus 2017a; Martynov, 2020a). zakuro is reported from the northwestern The conservative approach in the tax- Pacific only, ranging from middle Honshu onomy of the family Dendronotidae relied to Kamchatka Peninsula. Dendronotus priconsiderably on Mayr’s (1942, 1969) polytypic morjensis is restricted merely to the Sea of species concept and allowed the presence Japan and neighbouring localities in Japan.</p> <p>of broadly defined “species” distributed over Dendronotus venustus is restricted to the very large geographic distances. For example, northeastern Pacific. This considerable until about the year 2000 only D. frondosus regionalisation in a majority of the species is was recognised among the North Atlantic remarkable since all of them have planctonic narrow-bodied Dendronotus spp. (Thompson veliger larvae and may potentially disperse</p> <p>&amp; Brown, 1984). The wide geographic range of very widely. But in reality instead of a single D. frondosus at that time also encompassed “infinitely variable” species around whole the entire North Pacific up to subtropical Eurasia we have an enormously complicated regions of the middle of Honshu (Japan) and radiation of evidently separate species. From China (e.g., Robilliard, 1970; Baba, 1993; Lin these species, so far only D. lacteus shows a et al., 1986). Later this has showed not to be maximum range from the northern European the case, and the supposedly super-polytypic seas to the Arctic. However, it also shows a and pan-geographic “species” D. frondosus considerable morphological variation comhas been split into at least 11 narrow-defined pared to other species, which may lead to this species (see below and figs 1, 7). Each of these species becoming further subdivided. A very species occupies a particular geographic broad geographic range (the North Atlantic, region. The natural range of true D. frondo- Arctic and North Pacific) was previously also sus sensu stricto is limited solely to the east- indicated for Dendronotus dalli (Robilliard, ern and western parts of the North Atlantic 1970; Roginskaya, 1987). Mutilevel data with the subarctic Barents Sea as easternmost for the putative North Atlantic and Arctic limit, while it is absent in the Arctic region. “ D. dalli Downloaded ” show instead from that Brill.com it represents a 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/ separate species D. elegans (as “ D. niveus ”). Notably, Robilliard (1970) noted differences in Dendronotus elegans is a predominatly Arctic the reproductive system between European species, which reaches cold water masses at D. frondosus and his NE Pacific specimens of the Canadian and US North Atlantic coasts. D. venustus that he at that time identified as Dendronotus dalli instead is a predominantly “ D. frondosus ”. At the same time, apparently North Pacific species, which reach Arctic key reproductive features (e.g., number of seas into the Bering Strait and neighbouring prostatic alveols and shape of ampulla) can waters, but it is absent in the subarctic North vary significantly, as in the cases of D. albus or Atlantic regions. D. elegans. The separate characters should be Importantly, many of these species could therefore used with considerable care to dis-</p> <p>12/12/2023</p> <p>be separated by a careful application of mor- tinguish particular species.</p> <p>phological characters without aid of molec- All these cases clearly confirm that not ular data. The possibility of morphological only molecular data, per se, changed the separation is evident in the case of D. venustus modern landscape of biodiversity research.</p> <p>from the NE Pacific. Dendronotus venustus was The fundamental changes have occurred described in considerable detail and already in the dominated paradigm of the species became delineated from the resembling D. description. A major modern feature of this frondosus in the mid-20th century (published is the general denial of huge polytypic and in MacFarland, 1966) without use of molecular pan-geographic species in favour of fine-scale techniques and scanning electron microscopy interspecific morphological differences in (SEM). SEM has been available to taxonomists combination with limited geographic ranges.</p> <p>since the 1970s and its use would enable dif- Within restricted “narrow species” a polymorferentiation of some of the Dendronotus spe- phism (a basis for the polytypic concept) is cies that were described in the 21st Century. often manifested in the occurrence of parallel However, this did not happen because all these morphs, which may obscure species diversity characters could be considered as too subtle (see e.g., Korshunova et al., 2020b). Though from the perspective of the past dominant Mayr was among the major developers of the paradigm. For instance, Thompson and Brown population-based, variable, “non-typological” (1984) published an SEM image of radula of a species concept, he also admitted the possispecimen identified as D. frondosus but which ble existence of a hardly detectable diversity actually belongs to the recently described D. of “sibling species” (Mayr, 1942; Yoder et al., europaeus (Korshunova et al., 2017b). Even 2005). Despite this, the gravity of “a morpho-</p> <p>such morphologically well distinguished logically super-variable species concept” was species as D. albopunctatus (Robilliard, 1972) clearly unfavourable for seeking of fine-scale was commented as very similar to D. robus- differences. This was one of the reasons why tus (see Rudman, 2007). While data on the researchers for a long time omitted evidence multilevel diversity within Dendronotidae for morphological differences in the “ D. fron-</p> <p>are currently accumulating, more and finer dosus megacomplex”. The polytypic concept cases will be discovered in the future among became popular more than five decades already restricted “species”. Nevertheless, a ago and at that time Mayr (1942) suggested majority of the recently separated species can a potential solution for how to unfold the either be differentiated by SEM radula data or apparent “chaotic species” of “typologists”</p> <p>by a combination of the habitus data, radular into a polymorphic “biological” species.</p> <p>data and features of the reproductive system. However this Downloaded concept from can Brill now.com be12considered /12/2023 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/ as rather an obstacle than an aid in the dis- not yet described diversity within the genus covery of real patterns of biodiversity. As a Dendronotus was discovered in the temperate clear evidence for that, until very recently waters of Southern Hemisphere (e.g., Burn, only drastically different species within the 2015).</p> <p>genus Dendronotus have been recognised Most of the previously recognised without doubts (Robilliard, 1970; McDonald, Dendronotus species (seven out of nine, see 1983), despite uncertanties in their delimita- Robilliard, 1970, 1972) occur in the NE Pacific, tion (Robilliard, 1975). An especially remarka- while in the North Atlantic and NW Pacific ble example is the notorious problem of how only one or two species were commonly rec-</p> <p>to distinguish “ D. albus ” from “ D. diversicolor ”, ognized. This caused a distorted picture of a which has persisted since Robilliard (1970). biogeographic asymmetry in the Dendronotus This problem was in a profound confusion of diversity between the North Atlantic, norththe morphological characters rather than in western Pacific and northeastern Pacific. One the absence of molecular data and was solved of the important results of the fine-scale speonly recently by the distinction of D. robil- cies definition is a considerable reduction liardi (Korshunova et al., 2016a). of the “biogeographic asymmetry” among The majority of the described species of the Dendronotus spp. between NE Pacific, NW genus Dendronotus inhabit cold and temper- Pacific, Arctic and North Atlantic localities.</p> <p>ate regions of the Northern Hemisphere and The resulting numbers of shallow-water and their diversity clearly declines towards trop- shelf species in such regions, as the North ical localities. Pola and Stout (2008) assigned Atlantic and the NW Pacific, are impressive.</p> <p>two tropical dendronotids first to the genus I.e., in the North Atlantic and western subarc- Dendronotus. However, the morphology of the tic regions (Barents Sea as limit) there are curmale part of the reproductive system of the rently seven species, in the true Arctic (Kara, these tropical representatives is different from Laptev and East Siberian seas) there are also those in the genus Dendronotus, and available seven species so far described and recorded, molecular data for“ D.regius ”show it to be a sep- while in the NW Pacific (Bering Strait as northarate with a more basal placement compared ernmost limit) there are eigth species and in to any other dendronotid (figs 1, 2). Therefore NE Pacific there are 10 species. In all regions “ D. regius ” has become the type species of the the number of species will likely increase furnew genus Cabangus gen. nov., which encom- ther. This analysis clearly suggests that not passes exclusively tropical dendronotids. This only geographical proximity to the rich warm corroborates well with the new general agenda water fauna (as in the case of NE Pacific) is of the multilevel organismal diversity, which important, but also that ecological and other proposes narrowly defined taxa not only on a factors (including the history of faunal forspecies level, but also at the genus and family mation) have contributed substantially to levels (Korshunova et al., 2017a, b, c; 2019a). an active speciation. The presence of at least This prevents to produce non-diagnosable seven species of Dendronotus in the Artic as huge taxa that not only lack support from mor- one of the world’s coldest regions is a clear eviphology but also from various other data, such dence for this. All these data and implications as biogeography. The taxonomic diversity of have contributed to rapid and revolutionary the family Dendronotidae undergoes a process advancements in the taxonomy of the family of understanding and more taxa in this family Dendronotidae (Thollesson, 1998; Stout et al., are expected to be discovered. For example, a 2010; Martynov Downloaded et from al. 2015 Brill.com a, c, 2020 12/12 /a 2023; Ekimova 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/ et al. 2015; Korshunova et al. 2016a, b, 2017a, genus Dendronotus implies the presence of 2019b; Lundin et al., 2017; present study). many taxa that are incomparably more close to D. frondosus as the type species of the genus Multilevel Dendronotidae diversity and (e.g., D. primorjensis, D. venustus) than to the diFferent degrees of evolutionary separation much different wide-bodied D. robustus (and</p> <p>The majority of dendronotids are putatively its closely related species, e.g., D. velifer, D.</p> <p>united under the genus Dendronotus, which bathyvela). Therefore, although a direct comrepresents a morphologically heterogenous parison between D. robustus sensus strictus assemblage of several distinct clades with a and D. frondosus sensus strictus is possible, significant molecular divergence (see figs 1, 2; this is not taxonomically meaningful any- Synopsis; Appendix, table A2). The case of the more considering the presence of such hardly family Dendronotidae is therefore particularly distinguishable complexes. The similarities illustrative of multilevel differences among between the “ Dendronotus robustus complex” apparently taxonomically equally recognized and the “ D. frondosus megacomplex” are thus “species”. These apparently natural units are similarities at genus or higher level.This needs not equal by their morphological and molecu- to be adressed in a further study.</p> <p>lar properties, and broader, ontogenetic prop- The common formulation of a procedure erties. Hence, the term “species” is not equal to find differences within such multilevel over large phylogenetic distances but also organismal diversity, and hence the degree of among relatively small levels, such as genus separation between different groups of bio-</p> <p>and family. To make taxonomy consistent logical organisms is a “species delimitation”</p> <p>with the multilevel diversity among the fam- despite the species concept itself having no ily Dendronotidae, we separate a new genus universal fundamental agreement even in Cabangus gen. nov. and confirm a separate recent publications (Stanton et al., 2019). In status of the genus Pseudobornella. the present study we found a remarkable case Notably, even among the “core species” when the mean COI difference of a group is that is still constitute the genus Dendronotus slightly above 2% (normally indicative of</p> <p>(fig. 1) our present study demonstrates various a single species) but when morphological, degrees of separation from species in a state of ontogenetic and biogeographic evidence being morphologically well distinguished but are examined the two Dendronotus groups molecularly low delineated (the D. yrjargul deserve to be taxonomically marked as dis-</p> <p>sp. nov. – D. kalikal pair) to morphologically tinct species, namely D. kalikal and D. yrjarsubtle but molecularly well distinguished gul sp. nov. (figs 1–4, 6). At the same time species (D. frondosus – D. primorjensis, D. dal- D. nordenskioeldi sp. nov., despite its habitus li– D. elegans, D. lacteus – D. europaeus and similarity with D. lacteus, differs substantially D. nordenskioeldi sp. nov.) (figs. 1–8). We can from that species by its central teeth with unambiguously present distinguishing char- very weak denticles devoid of rib-like strucacters only when these apparent groups tures (fig. 5) and intriguingly, (fig. 1) it demoncurrently known as species really belong to strates significant molecular distance from sufficiently distantly related subclades, as in all know species of the genus Dendronotus easily recognizable species such as D. iris or (Appendix, table A2). Despite the lack of D. subramosus, or D. frondosus and D. robus- consensus about the meaning of “species” we tus (figs 1, 7). A very complicated mutilevel can observe some “groups of genetically and morphological and molecular diversity of the ecologically Downloaded similar individuals from Brill.com” (12 e./ g 12.,/2023 Shapiro04:12:08PM 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/ et al., 2016) in nature, even though there are of live specimens. Our warm gratitude goes to no clear criteria about how to separate one Karin Fletcher for improvement of English in similar group from another similar group. an earlier version of the manuscript and for</p> <p>A conclusion was reached that the “popula- providing us with the wonderful pictures of tion-to-species” level represents a continuum D. rufus and D. venustus. Electron Microscopy (Mallet et al., 2009; Coates et al., 2018). Using Laboratory MSU is thanked for support these numerous available data on different with electron microscopy. Reviewers are groups and our present data, we therefore dis- thanked for providing comments. The study agree with the most recent suggestion to save was supported by the Norwegian Taxonomy “species” as a universal concept using a “weak Initiative project #sneglebuss Barents Sea (19- realism” approach (Reydon &amp; Kunz, 2019). 18_70184240). The work of AM was supported Because even in its weaker form, “species” as by the research project of MSU Zoological a reality still implies the universal application Museum (AAAA-A16-116021660077-3). The of this concept despite that it encompasses work of TK was conducted under the IDB RAS groups that have different degrees of separa- Government basic research program in 2020 tion, and hence different morphological and № 0108-2019-0002.</p> <p>molecular properties. 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