<i>Ceuthonoe</i> <i>nezhai</i> gen. et sp. n. (Polynoidae: Polynoinae) commensal with sponges from Weijia Guyot, western Pacific

Ceuthonoe nezhai gen. et sp. n. (Polynoidae: Polynoinae) commensal with sponges from Weijia Guyot, western Pacific

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Yueyun Wang¹^,², Yadong Zhou¹^,², Chunsheng Wang¹^,²^,³^,⁴^,^*

Acta Oceanologica Sinica | 2021, 40(12) : 90 - 103

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Acta Oceanologica Sinica | 2021, 40(12): 90-103

• Environmental Study of Seamounts and Adjacent Waters in the Northwestern Pacific Ocean •

Ceuthonoe nezhai gen. et sp. n. (Polynoidae: Polynoinae) commensal with sponges from Weijia Guyot, western Pacific

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Yueyun Wang¹^,², Yadong Zhou¹^,², Chunsheng Wang¹^,²^,³^,⁴^,^*

Affiliations

¹ Second Institute of Oceanography, Ministry of Natural Resources, Hangzhou 310012, China

² Key Laboratory of Marine Ecosystem Dynamics, Ministry of Natural Resources, Hangzhou 310012, China

³ School of Oceanography, Shanghai Jiao Tong University, Shanghai 200030, China

⁴ Southern Marine Science and Engineering Guangdong Laboratory (Zhuhai), Zhuhai 519080, China

Published: 2021-11-25 doi: 10.1007/s13131-021-1885-0

Outline

Abstract

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Ceuthonoe, a new genus with type species C. nezhai sp. n. commensal with euplectellid glass sponge (Hexactinellida: Euplectellidae), is described and illustrated, based on the material collected from Weijia Guyot, western Pacific. The new genus is distinguished by a combination of characters: body with 32 segments; 14 pairs of elytra; prostomium with prominent cephalic peaks and facial tubercle; aciculae and chaetae on tentaculophores; biramous parapodium, each ramus with a long aciculum penetrating epidermis, without supra- or subacicular process; and noto- and neurochaetae with unidentate tips. Ceuthonoe gen. n. is mostly silimar to genera of Polynoinae in morphology. However the phylogenetic analysis based on cytochrome c oxidase subunit I, 16S, 18S, and 28S genes suggested that this new genus was separated from the main branch of Polynoinae. The study also discussed the genera related to the new genus.

Key words

new genus / new species / Polynoidae / taxonomy

Cite this Article

Yueyun Wang, Yadong Zhou, Chunsheng Wang. Ceuthonoe nezhai gen. et sp. n. (Polynoidae: Polynoinae) commensal with sponges from Weijia Guyot, western Pacific[J]. Acta Oceanologica Sinica, 2021 , 40 (12) : 90 -103 . DOI: 10.1007/s13131-021-1885-0

Full Text

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1 Introduction

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Polynoidae is one of the most diverse polychaete families, in the number of genera and species (Salazar-Vallejo et al., 2015; Bonifácio and Menot, 2019). It is widely distributed from the intertidal zone to the hadal zone (Paterson et al., 2009). Polynoids are characterized by having dorsoventrally flattened bodies, two palps, one to three antennae, elytrae on the dorsum and biramous parapodia with simple chaetae. Among approximately 900 polynoid species, more than 200 are known to be involved in about 550 associations with other marine invertebrates (Britayev et al., 2014; Martin and Britayev, 2018).

To date, about 58 genera have been included in the subfamily Polynoinae (Read and Fauchald, 2019). Most primitive species in Polynoinae are obligate symbionts with speciﬁc morphological adaptations (Serpetti et al., 2017; Taboada et al., 2020). Eight species (Harmothoe hyalonemae Martín, Rosell & Uriz, 1992; H. spongicola Hanley & Burke, 1991; Hermadion fauveli Gravier, 1918; Hololepidella nigropunctata (Horst, 1915); Intoshella euplectellae (McIntosh, 1885); I. dictyaulus Sui, Li & Kou, 2018; Malmgreniella murrayensis Pettibone, 1993a; Neopolynoe acanellae (Verril, 1881)) from six genera in Polynoinae are commensal with sponges (Martin and Britayev, 1998, 2018). The host-specificity of polynoids is not very high, and some species may also be commensal with other hosts, such as corals.

In May 2016, a sponge specimen (Hexactinellida: Lyssacinosida: Euplectellidae) was collected by the Chinese manned deep-sea submersible Jiaolong at a depth of 2 505 m from Weijia Guyot, western Pacific. A pair of polynoids were separated from the inner cavity of the sponge. Following a detailed examination of the morphology and sequence comparisons of mitochondrial cytochrome c oxidase subunit I (COI) using the BLAST search in GenBank, a new species of Polynoinae was fully described and illustrated, and a new genus was proposed after morphological comparision and phylogenetic analysis.

2 Materials and methods

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Four gene fragments (COI, 16S, 18S, and 28S) were used to infer the phylogenetic relationship of the new specimens with other polynoid scale worms. The non-coding 16S, 18S, and 28S gene sequences were aligned separately with MAFFT v.7.450 (Katoh and Standley, 2013); COI gene sequences were aligned using the MUSCLE algorithm. The alignments were manually concatenated in Geneirous R.11, resulting in a concatenated alignment of 4 054 bp: COI (552 bp), 16S (520 bp), 18S (1 953 bp), and 28S (1 029 bp). Nucleotide substitution models were selected based on the BIC criterion using jModelTest v. 2.1.10: GTR + F + I + G4 for COI and 16S, TN+F+R3 for 18S and TIM3+F+R3 for 28S. Sigalionidae and Acoetidae were selected as the outgroups based on previous studies (Zhang et al., 2018; Taboada et al., 2020). Maximum likelihood (ML) analyses were conducted in IQtree 1.6.10, with the bootstrap support values for each node determined using an ultrafast bootstrap algorithm for 100 000 replicates. Bayesian inference (BI) analyses were implemented in MrBayes v3.2 (Ronquist et al., 2012), with the substitution models replaced by GTR+G+I. Two parallel chain runs were performed each for 2 000 000 generations, with topologies sampled at every 1 000 generations. Tracer v1.6 was used to estimate the burn-in value, visualize, and ensure convergence.

Two specimens were removed from the body cavity of the sponge (Hexactinellida: Lyssacinosida: Euplectellidae: Placopegma sp.) collected by submersible Jiaolong, from Weijia Guyot, the western Pacific in May 2016. The specimens were preserved in 95% ethanol, subsequently dissected, and examined under a stereomicroscope (Zeiss Discovery V20). Notochaetae and neurochaetae were dissected carefully from the body, observed and photographed under a microscope (Zeiss Axio Imager A2). For barcoding purposes, COI, 18S, 28S, and 16S genes were ampliﬁed and sequenced. DNA was extracted using a DNAeasy Tissue Kit (Qiagen) following the manufacturer’s protocol. Primer pairs and PCR programmes used to amplify polynoid genetic markers were the same as those described by Zhou et al. (2019). GenBank accession numbers are presented in Table 1. The type specimens are deposited in the Sample Repository of Second Institute of Oceanography (RSIO), Ministry of Natural Resources, Hangzhou, China.

3 Results

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3.1 DNA barcoding and phylogenetics

Pairwise GTR-corrected distances of COI between Ceuthonoe and non-Ceuthonoe genera, which averaged 27.50% (range 21.46%–33.74%), fell in the range of inter-generic genetic distance among non-Ceuthonoe genera (range 6.90%–38.31%, average 24.53%; Appendix Table A1). Both ML and BI analyses obtained similar topology, in which Ceuthonoe nezhai sp. n. clustered with an undescribed polynoid commensal to a carnivorous sponge Chondrocladia verticillata Topsent, 1920, Polynoidae sp. 2 ST-2019 (Taboada et al., 2020), forming a discrete lineage outside a clade composed of species from Arctonoinae, Lepidastheniinae, Lepidonotopodinae, Macellicephalinae, and Polynoinae (Fig. 1).

3.2 Systematics

Family Polynoidae Kinberg, 1856

Genus Ceuthonoe gen. n.

Type species. Ceuthonoe nezhai gen. et sp. n.

Generic diagnosis. Body short, with about 32 segments. Elytra 14 pairs, on segments 2, 4, 5, and 7, alternate segments to 23, 26, and 29. Elytra without fringed papillae and with conical microtubercles.

Prostomium bilobed, with two distinct cephalic peaks, with paired palps, and three antennae. Lateral antennae with ceratophores inserted ventrally; a prominent conical facial tubercle under the median antenna. Eyes two pairs. Pharynx with two pairs of hooked jaws without denticulation on the inner border. Tentaculophore with an aciculum and few chaetae. Second segment with first pair of elytra and biramous parapodia.

Parapodia biramous; noto- and neuropodia well-developed with elongate acicular lobes; tips of noto- and neuroacicula penetrating the epidermis. Notochaetae thicker than neurochaetae. Nephridial papillae bulbous.

Etymology. The genus name is a combination of the Greek words “Ceuth” meaning “hidden” and “noe” from “Polynoe”, a sea nymph of Greek mythology. It refers to the type species commensal with sponges. Gender: feminine.

Remarks. This new genus is unique in having a low number of segments, prominent cephalic peaks, facial tubercle, aciculae, and chaetae on the tentacular segment, long aciculae spines penetrating the epidermis.

The new genus may be confused with Harmothoe, which has neurochaetae with bidentate tips whereas Ceuthonoe gen. n. has neurochaetae with unidentate tips. The new genus is also superficially similar to Antarctinoe Barnich et al. 2006, Bylgides Chamberlin 1919, Eunoe Malmgren 1866, and Gattyana McIntosh, 1897. They all have a low number of segments, cephalic peaks, and a facial tubercle. Antarctinoe and Bylgides are distinguishable from the new genus by having a digitiform supra-acicular process on neuropodia. The new genus differs from Eunoe and Gattyana in the form of chaetae. The distal tips of notochaetae of Eunoe are pointed and glabrous or blunt and split. Notochaetae of the new species have slightly curved, smooth tips. Neurochaetae of Eunoe are sturdy and have somewhat expanded and serrated subterminal parts, while that of the new genus are slender with tapering ends. Additionally, the neuropodia of Eunoe have a digitiform supra-acicular process (Wehe, 2006). Notochaetae of Gattyana are thinner than neurochaetae, while notochaetae of the new genus are thicker than neurochaetae.

Ceuthonoe nezhai gen. et sp. n.

(Figs 2 and 3)

Type material. Holotype: B6317500002, complete, Weijia Guyot, western Pacific Ocean, JL-Dive106, Station S3-1, 13.018 45°N, 156.948 00°E, 2 505 m depth, May 1, 2016. Paratype: B6317500004, one complete specimen, same collecting data as holotype.

Description. Holotype about 19.8 mm from the prostomium to the end of the body (without anal cirri), 2.7 mm wide at the widest part excluding parapodia (Fig. 2a). Paratype about 15.2 mm in length and 1.8 mm wide at the widest part excluding parapodia (Fig. 2b). Parapodia long and subequal to the width of corresponding segments. The following description is based on holotype and paratype.

Body short, with 32 segments, tapering posteriorly; the ventral side flattened and the dorsal side slightly convex. A deep longitudinal groove on the ventral side. Elytra 14 pairs, on segments 2, 4, 5, and 7, alternate segments to 23, 26, and 29. Elytra large, covering the dorsum except the middle part. Elytra oval in shape, without fringed papillae, with many conical microtubercles on surfaces (Figs 2c, 3c and d). Microtubercles near anterior and inner regions increasing in number and becoming larger. Dorsal cirri and dorsal tubercles near each other, present on non-elytrigerous segments. Dorsal cirri composed of short cylindrical cirrophores and long styles, up to three times the length of the parapodia (Figs 2j and 3b).

Prostomium bilobed, wider than long, with distinct cephalic peaks and three antennae; ceratophore of median antenna in the anterior notch, lateral antennae with ceratophores inserted ventrally to median antenna (Figs 2d, e and 3a). Median antenna missing in both specimens. Styles of lateral antennae covered with short soft papillae (Fig. 2k). A pair of stout palps with smooth surface. A distinct conical facial tubercle under the ceratophore of the median antenna. Two pairs of eyes arranged in a reverse trapezium, posterolaterally on the widest part of the prostomium (Fig. 2f). Pharynx with two pairs of hooked jaws without denticulation on the inner border (Fig. 2l). Tentaculophores lateral to prostomium, each with a long projecting aciculum and many chaetae projecting anteriorly (Fig. 3a). Notochaetae similar to those in the following segments. Tentacular cirri shorter and thinner than palps.

Second segment with first elytra, biramous parapodia, and coiled ventral cirri longer than the following ones, without nuchal lobe. Parapodia biramous, with yellowish noto- and neuroaciculae and numerous chaetae (Figs 2m and 3b). Notopodium shorter than neuropodium, with a slender acicular lobe, and tip of aciculum penetrating epidermis (Fig. 3f). Notochaetae and neurochaetae the same from the anterior segments to the posterior end. Notochaetae stout, about 1.5 times as wide as neurochaetae. Notochaetae projecting dorsally, with numerous rows of spinules having slightly curved, smooth tips (Figs 2m and 3e). Neuropodium well developed, with obtuse postchaetal lobe and longer prechaetal acicular lobe, and tip of aciculum penetrating epidermis. The upper part of neurochaetae extending, wider than the shaft, with spinous regions. The tips of neurochaetae slightly curved and smooth (Fig. 3g). Ventral cirri short and less than half length of parapodia.

Nephridial papillae short bulbous, beginning on Segment 5. Anal cirri lost.

Pigmentation. Signiﬁcant brown pigmentation on the ventral side of parapodia and body, especially on sides of the ventral groove. Tips of ventral cirri dark-brown. Pigmentation stable in alcohol-preserved specimens.

Type locality. Weijia Guyot, western Pacific.

Distribution. Only known from the type locality. Commensal with sponges.

Etymology. The specific name is dedicated to Nezha, which is a Chinese mythological figure.

Remarks. Polynoids are often found to be associated with other invertebrates, such as sponges, corals, echinoderms, mollusks, and other polychaetes. Eight Polynoinae species (Harmothoe hyalonemae from Western Mediterranean, 1 083–2 256 m; H. spongicola from Fairway Reefs, Coral Sea, 720 m; Hermadion fauveli from Cape Verde Islands, Atlantic Ocean; Hololepidella nigropunctata, widely distributed in Indo-west and central Pacific Ocean, 1–76 m; Intoshella euplectellae from Philippines, 184 m; I. dictyaulus from near the Mariana Trench, West Pacific, 890 m; Malmgreniella murrayensis from the South Arabian Coast, 38 m; Neopolynoe acanellae from east coast of the MA, USA) are reported commensal with sponges. Ceuthonoe nezhai sp. n. is similar to Harmothoe species in having distinct cephalic peaks, a facial tubercle, and lateral antennae inserted ventrally on the prostomium. However, the new species differs from the latter by possessing neurochaetae all with unidentate tips. Hololepidella, Intoshella and Malmgreniella species can also be easily distinguished from the new species by the number of segments, morphology of elytra, neurochaetae and prostomium. Hololepidella nigropunctata has 55 or more segments, with mostly smooth elytra, smooth dorsal cirri, and bidentate neurochaetae (Barnich et al., 2004; Wehe, 2006). Intoshella species has smooth elytra and no cephalic peaks. Malmgreniella murrayensis has 36 segments, no cephalic peaks, and bidentate neurochaetae. The new species is also closely related to species of Bylgides Chamberbin, 1919. They all have a small number of segments and elytrae, and aciculae on tentaculophores. They can be distinguished by the form of the chaetae and acicular lobes. Neurochaetae of Bylgides have long spinous regions and capillary tips, but the neurochaetae of Ceuthonoe nezhai sp. n. have blunt tips. Long conical neuroacicular lobes of Bylgides have a supraacicular digitiform process, but the acicular lobes of Ceuthonoe nezhai sp. n. have no supraacicular or subacicular process. The new species is also very similar to genus Neopolynoe species in anterior body. The species of Neopolynoe are distinguishable from the new species by having numerous segments and naked dorsum of the posterior body. Comparisons of diagnostic morphological characteristics of closely related genera with those of the new genus are presented in Table 2.

4 Discussion

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Polynoidae is one of the most diverse polychaetes family having the highest number of species involved in symbiotic relationships (Britayev et al., 2014). To date, phylogenetic relationships within Polynoidae have not been fully resolved. Phylogenetic analyses conducted on a limited number of polynoid sequences indicate that Polynoinae is not a monophyletic group (Bonifácio and Menot, 2019; Taboada et al., 2020). The morphological characteristics of the new genus suggest that it should be a member of Polynoinae based on the diagnosis given by Wehe (2006). However, the results obtained from molecular phylogenetic analyses reveal that Polynoinae is a polyphyletic group. The new genus and Paralepidonotus are separated from the major clade of Polynoinae, which is intermingled with species of Arctonoinae, Lepidastheniinae, Lepidonotopodinae, and Macellicephalinae in the phylogenetic tree, indicating that a wider range of taxa and genetic marker sampling is needed for further revision of this clade. Members belonging to Arctonoinae have subbiramous parapodia and no chaetae on tentaculophores, while those belonging to the Polynoinae usually have chaetae on tentaculophores and biramous parapodia (Barnich and Fiege, 2001). This new genus is more similar to a member of Polynoinae. Members of Lepidastheniinae are characterized by lateral antennae terminal, tentaculophores without chaetae, and reduced notopodia; notochaetae are usually missing (Salazar-Vallejo et al., 2015). The new genus is similar to Parahalosydna Horst, 1915 (Lepidastheniinae) in the number of segments and elytra. However, lateral antennae are in the terminal position in Parahalosydna and in the ventral position in the new genus. Subfamily Lepidonotopodinae was erected for genus Lepidonotopodium Pettibone (1983), characterized by having no lateral antennae. Bonifácio and Menot (2019) synonymized Lepidonotopodinae with Macellicephalinae based on the the phylogenetic analysis of morphological and molecular datasets. Both subfamilies were characterized by the synapomorphic absence of lateral antennae. Hatch et al. (2020) analyzed the phylogenetic relationships of Branchinotogluminae, Branchiplicatinae, Branchipolynoinae, Lepidonotopodinae, Bathykurila guaymasensis, and Levensteiniella (described as Macellicephalinae). The results suggested that these taxa formed a well-supported clade. Therefore, the oldest subfamily name Lepidonotopodinae Pettibone 1983 was reinstated, including Branchipolynoe, Branchinotogluma, Bathykurila, Branchiplicatus, Lepidonotopodium, Levensteiniella, Thermopolynoe, and Peinaleopolynoe. Our result showed that these two subfamilies were sister groups with robust support (Fig. 1). Lepidonotinae has a fixed number of segments (26) or more (up to 70 and more), with lateral antennae inserted terminally or subterminally on the prostomium. This new genus is closely related to the short-bodied genus Hermenia belonging to Lepidonotinae. Both of them have chaetae on the tentacular segment and a bulbous facial tubercle. Parapodia of Hermenia are subbiramous with reduced notopodia forming small lobes on the anterodorsal faces of stout neuropodia, while those of the new genus are biramous with normal-sized notopodia. Neurochaetae of Hermenia are stout, with falcate tips and 1 or 2 stout teeth. Neurochaetae of the new genus have unidentate tips. According to the taxonomic revision of Wehe (2006), the new genus is similar to the genera Eunoe and Harmothoe; they all have cephalic peaks, ventrally inserted lateral antennae, and chaetae on tentaculophores. Their differences are discussed in remarks on the new genus.

Symbiotic relationships between annelid polychaetes and other marine invertebrates are common; approximately 45% of all known commensal species belong to Polynoidae Kinberg, 1856 (Taboada et al., 2020). The species Polynoidae sp. 2 ST-2019 reported by Taboada et al. (2020) may also be included in this new genus owing to its close molecular phylogenetic relationship with Ceuthonoe nezhai gen. et sp. n. Both of them are similar in morphological characteristics. Polynoidae sp. 2 ST-2019 was collected in association with carnivorous sponge Chondrocladia verticillata from the Gulf of Mexico, while Ceuthonoe nezhai gen. et sp. n. was collected in association with sponge Placopegma sp. More findings are needed to clarify the aforementioned relationships.

Acknowledgements

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We are very grateful to all the scientists and crew of the R/V Xiangyanghong 9 and the team of the submersible Jiaolong for their help: n collecting deep-sea materials. We thank Ruiyan Zhang for molecular data collection. We also thank Vlaams Instituut voor de Zee Library for their help in providing important references.

Funding

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The National Natural Science Foundation of China under contract No. 41806179; the China Ocean Mineral Resources Research and Development Association Program under contract No. DY135-E2-2-03.

References

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Barnich R, Fiege D. 2001. The Mediterranean species of Malmgreniella Hartman, 1967 (Polychaeta: Polynoidae: Polynoinae), including the description of a new species. Journal of Natural History, 35(8): 1119–1142, doi: 10.1080/00222930152434463

Barnich R, Fiege D, Micaletto G, et al. 2006. Redescription of Harmothoe spinosa Kinberg, 1856 (Polychaeta: Polynoidae) and related species from Subantarctic and Antarctic waters, with the erection of a new genus. Journal of Natural History, 40(1–2): 33–75, doi: 10.1080/00222930500445044

Barnich R, Fiege D, Sun Ruiping. 2004. Polychaeta (Annelida) of Hainan Island, South China Sea: Part III. Aphroditoidea. Species Diversity, 9(4): 285–329, doi: 10.12782/specdiv.9.285

Bock G, Fiege D, Barnich R. 2010. Revision of Hermadion Kinberg, 1856, with a redescription of Hermadion magalhaensi Kinberg, 1856, Adyte hyalina (G. O. Sars, 1873) n. comb. and Neopolynoe acanellae (Verrill, 1881) n. comb. (Polychaeta: Polynoidae). Zootaxa, 2554: 45–61, doi: 10.11646/zootaxa.2554.1.4

Bonifácio P, Menot L. 2019. New genera and species from the Equatorial Pacific provide phylogenetic insights into deep-sea Polynoidae (Annelida). Zoological Journal of the Linnean Society, 185(3): 555–635, doi: 10.1093/zoolinnean/zly063

Britayev T A, Gil J, Altuna Á, et al. 2014. New symbiotic associations involving polynoids (Polychaeta, Polynoidae) from Atlantic waters, with redescriptions of Parahololepidella greeffi (Augener, 1918) and Gorgoniapolynoe caeciliae (Fauvel, 1913). Memoirs of Museum Victoria, 71: 27–43, doi: 10.24199/j.mmv.2014.71.04

Fiege D, Barnich R. 2009. Polynoidae (Annelida: Polychaeta) associated with cold-water coral reefs of the northeast Atlantic and the Mediterranean Sea. Zoosymposia, 2: 149–164, doi: 10.11646/zoosymposia.2.1.13

Jirkov I A. 2001. Polikhety severnogo Ledovitogo Okeana [Polychaeta of the Arctic Ocean] (in Russian). Moscow, Russia: Yanus-K Press, 126–127

Hanley J R. 1991. Revision of the genus Parlalepidonotus Horst, 1915 (Polychaeta: Polynoidae). Invertebrate Taxonomy, 4(5): 1053–1075

Hanley J R. 1993. Scaleworms (Polychaeta: Polynoidae) of Rottnest Island, Western Australia. In: Wells F E, Walker D I, Kirkman H, et al., eds. Proceedings of the Fifth International Marine Biological Workshop: The Marine Flora and Fauna of Rottnest Island, Western Australia. Perth: Western Australian Museum, 305–320

Hanley J R, Burke M. 1991. Polychaeta Polynoidae: Scaleworms of the Chesterfield Islands and Fairway Reefs, Coral Sea. In: Crosnier A, ed. Résultats des Campagnes MUSORSTOM, Volume 8. Mémoires du Muséum national d'histoire naturelle, 151(1): 9–82

Hatch A S, Liew H, Hourdez S, et al. 2020. Hungry scale worms: Phylogenetics of Peinaleopolynoe (Polynoidae, Annelida), with four new species. ZooKeys, 932: 27–74, doi: 10.3897/zookeys.932.48532

Katoh K, Standley D M. 2013. MAFFT multiple sequence alignment software version 7: Improvements in performance and usability. Molecular Biology and Evolution, 30(4): 772–780, doi: 10.1093/molbev/mst010

Levenstein R Y. 1982. New genera of the subfamily macellicephalinae (Polychaeta, Polynoidae) from the Tasman hollow. Zoologicheskii Zhurnal, 61(9): 1291–1296

Martin D, Britayev T A. 1998. Symbiotic polychaetes: review of known species. Oceanography and Marine Biology: An Annual Review, 36: 217–340

Martin D, Britayev T A. 2018. Symbiotic polychaetes revisited: An update of the known species and relationships (1998–2017). Oceanography and Marine Biology, 56: 371–448

Martín D, Rosell D, Uriz M J. 1992. Harmothöe hyalonemae sp. nov. (Polychaeta, Polynoidae), an exclusive inhabitant of different Atlanto-Mediterranean species of Hyalonema (Porifera, Hexactinellida). Ophelia, 35(3): 169–185, doi: 10.1080/00785326.1992.10429925

McIntosh W C. 1885. Report on the Annelida Polychaeta collected by H. M. S. Challenger during the years 1873–76. Report on the Scientific Results of the Voyage of H. M. S. Challenger during the years 1873–76. Zoology, 12: 1–554

Paterson G L J, Glover A G, Barrio Froján C R S, et al. 2009. A census of abyssal polychaetes. Deep Sea Research Part II: Topical Studies in Oceanography, 56(19–20): 1739–1746, doi: 10.1016/j.dsr2.2009.05.018

Pettibone M H. 1975. Review of the genus Hermenia, with a description of a new species (Polychaeta: Polynoidae: Lepidonotinae). Proceedings of the Biological Society of Washington, 88(22): 233–248

Pettibone M H. 1983. A new scale worm (Polychaeta: Polynoidae) from the hydrothermal rift-area off western Mexico at 21°N. Proceedings of the Biological Society of Washington, 96(3): 392–399

Pettibone M H. 1993a. Scaled polychaetes (Polynoidae) associated with ophiuroids and other invertebrates and review of species referred to Malmgrenia McIntosh and replaced by Malmgreniella Hartman, with descriptions of new taxa. Smithsonian Contributions to Zoology, 538: 1–92

Pettibone M H. 1993b. Revision of some species referred to Antinoe, Antinoella, Antinoana, Bylgides, and Harmothoe (Polychaeta: Polynoidae: Harmothoinae). Smithsonian Contributions to Zoology, 545: 1–41

Read G, Fauchald K. 2019. World Polychaeta database. Polynoidae Kinberg, 1856. World Register of Marine Species. http://www.marinespecies.org/aphia.php?p=taxdetails&id=939 [2019-12-30/2021-04-01]

Ronquist F, Teslenko M, Van der Mark P, et al. 2012. MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology, 61(3): 539–542, doi: 10.1093/sysbio/sys029

Salazar-Vallejo S I, González N E, Salazar-Silva P. 2015. Lepidasthenia loboi sp. n. from Puerto Madryn, Argentina (Polychaeta, Polynoidae). ZooKeys, 546: 21–37, doi: 10.3897/zookeys.546.6175

Serpetti N, Taylor M L, Brennan D, et al. 2017. Ecological adaptations and commensal evolution of the Polynoidae (Polychaeta) in the Southwest Indian Ocean Ridge: A phylogenetic approach. Deep Sea Research Part II: Topical Studies in Oceanography, 137: 273–281, doi: 10.1016/j.dsr2.2016.06.004

Sui Jixing, Li Xinzheng, Kou Qi. 2019. A new species of the genus Intoshella Darboux, 1899 (Polychaeta: Polynoidae) commensal with a deep-sea sponge from a seamount near the Mariana Trench. Marine Biodiversity, 49(3): 1479–1488, doi: 10.1007/s12526-018-0922-5

Taboada S, Silva A S, Neal L, et al. 2020. Insights into the symbiotic relationship between scale worms and carnivorous sponges (Cladorhizidae, Chondrocladia). Deep Sea Research Part I: Oceanographic Research Papers, 156: 103191, doi: 10.1016/j.dsr.2019.103191

Uschakov P V. 1982. Polychaetes of the Suborder Aphroditiformia of the Arctic Ocean and the Northwestern Part of the Pacific, Families Aphroditidae and Polynoidae. Fauna of the USSR, Polychaeta, 2: 1–272

Wehe T. 2006. Revision of the scale worms (Polychaeta: Aphroditoidea) occurring in the seas surrounding the Arabian Peninsula: Part I. Polynoidae. Fauna of Arabia, 22: 23–197

Zhang Yanjie, Sun Jin, Rouse G W, et al. 2018. Phylogeny, evolution and mitochondrial gene order rearrangement in scale worms (Aphroditiformia, Annelida). Molecular Phylogenetics and Evolution, 125: 220–231, doi: 10.1016/j.ympev.2018.04.002

Zhou Yadong, Chen Chong, Sun Yanan, et al. 2019. Amphisamytha (Annelida: Ampharetidae) from Indian Ocean hydrothermal vents: Biogeographic implications. Deep Sea Research Part I: Oceanographic Research Papers, 154: 103148, doi: 10.1016/j.dsr.2019.103148

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doi: 10.1007/s13131-021-1885-0

Receive Date：2021-05-06
Online Date：2026-03-06
Published：2021-11-25

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Received：2021-05-06
Accepted：2021-06-21

Funding

The National Natural Science Foundation of China under contract No. 41806179; the China Ocean Mineral Resources Research and Development Association Program under contract No. DY135-E2-2-03.

Affiliations

¹ Second Institute of Oceanography, Ministry of Natural Resources, Hangzhou 310012, China

² Key Laboratory of Marine Ecosystem Dynamics, Ministry of Natural Resources, Hangzhou 310012, China

³ School of Oceanography, Shanghai Jiao Tong University, Shanghai 200030, China

⁴ Southern Marine Science and Engineering Guangdong Laboratory (Zhuhai), Zhuhai 519080, China

Corresponding:

*E-mail: wangsio@sio.org.cn

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表12种不同金属材料的力学参数

科 Family	属数 Number of genus	种数 Number of species	占总种数比例 Percentage of total species (%)	属 Genus	种数 Number of species	占总种数比例 Percentage of total species (%)
鹅膏菌科Amanitaceae	2	11	5.26	鹅膏菌属 Amanita	10	4.78
小菇科 Mycenaceae	2	12	5.74	丝盖伞属 Inocybe	5	2.39
多孔菌科 Polyporaceae	8	14	6.70	蜡蘑属 Laccaria	5	2.39
红菇科 Russulaceae	3	23	11.00	小皮伞属 Marasmius	6	2.87
				小菇属 Mycena	11	5.26
				光柄菇属 Pluteus	5	2.39
				红菇属 Russula	17	8.13
				栓菌属 Trametes	5	2.39

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Table 1. List of taxa included in the phylogenetic analysis including the NCBI accession number

Family	Subfamily	Taxon	18S	28S	16SrRNA	COI
Note: — means the sequence was not reported.
Acoetidae		Panthalis oerstedi	AY839572	JN852845	JN852881	AY839584
Iphionidae		Iphione sp.	JN852819	—	JN852886	JN852921
		Thermiphione sp.	JN852820	JN852849	JN852887	JN852922
Polynoidae	Polynoinae	Ceuthonoe nezhai gen. et sp. n.	—	MW063140	MW063141	MW064265
	Polynoinae	Acholoe astericola	AY839567	JN852850	JN852888	AY839576
	Lepidastheniinae	Alentia gelatinosa	AY839566	—	—	AY839577
	Polynoinae	Antarctinoe ferox	MG905039	—	MG905033	KJ676611.1
	Polynoinae	Antinoe aequiseta	—	—	—	MH337356.1
	Polynoinae	Antipathipolyeunoa sp.	KU738169	KU738184	KU738149	KU738202.1
	Polynoinae	Austrolaenilla antarctica	—	—	KJ676606	KJ676627.1
	Macellicephalinae	Bathykurila guaymasensis	DQ074765	—	MG905034	DQ074766
	Macellicephalinae	Branchinotogluma sandersi	JN852821	JN852851	JN852889	JN852923
	Macellicephalinae	Branchipolynoe symmytilida	—	—	AF315055	AY646021
	Polynoinae	Brychionoe sp.	KU738182	KU738200	KU738167	—
	Polynoinae	Bylgides elegans	JN852822	JN852852	JN852890	JN852924
	Polynoinae	Bylgides sarsi	JN852823	JN852853	JN852891	JN852925
	Arctonoinae	Capitulatinoe cf. cupisetis	KF919301	KF919302	KF919303	—
	Polynoinae	Enipo torelli	—	HQ023831	—	HQ023480.1
	Polynoinae	Eunoe nodosa	JN852824	JN852854	JN852892	JN852926
	Polynoinae	Gastrolepidia clavigera	JN852825	JN852855	JN852893	JN852927
	Polynoinae	Gattyana ciliata	AY894297	DQ790035	—	AY894312
	Polynoinae	Gattyana cirrhosa	JN852826	JN852856	JN852894	JN852928
	Macellicephalinae	Gesiella jameensis	Ky454403	Ky823476	Ky454412	Ky454429
	Polynoinae	Gorgoniapolynoe corralophila	KU738173	KU738189	KU738154	KU738208.1
	Polynoinae	Grubeopolynoe tuta	—	—	—	MF121411.1
	Lepidonotinae	Halosydnella australis	KY823449	KY823463	KY823480	KY823495
	Lepidonotinae	Halosydna brevisetosa	JN852827	JN852857	JN852895	AY894313
	Polynoinae	Harmothoe glabra	JN852828	JN852858	JN852896	JN852929
	Polynoinae	Harmothoe imbricata	AY340434	AY340400	AY340463	AY839580
	Polynoinae	Harmothoe impar	JN852829	JN852859	JN852897	JN852930
	Polynoinae	Harmothoe oculinarium	AY894299	JN852860	JN852898	AY894314
	Polynoinae	Hermadion truncata	—	—	—	MH242794.1
	Lepidonotinae	Hermenia verruculosa	JN852830	JN852861	JN852899	JN852931
	Lepidastheniinae	Hyperhalosydna striata	JN852831	JN852862	JN852900	JN852932
	Polynoinae	Intoshella dictyaulus	MG519807	—	—	MG519808.1
	Lepidastheniinae	Lepidastenia elegans	JN852832	JN852863	JN852901	JN852933
	Lepidonotinae	Lepidonotus clava	JN852833	JN852864	JN852902	JN852934
	Lepidonotinae	Lepidonotus squamatus	AY176290	JN852865	JN852903	AY894316
	Polynoinae	Leucia violacea	—	—	—	KX946860.1
	Polynoinae	Malmgreniella mcintoshi	JN852834	JN852866	JN852904	JN852935
	Polynoinae	Melaneis loveni	JN852835	JN852867	JN852905	JN852936
	Polynoinae	Neopolynoe acanellae	MN653050	MN653123	MN653064	MN656076
	Polynoinae	Neopolynoe paradoxa	JN852836	JN852868	JN852906	JN852937
	Polynoinae	Neopolynoe chondrocladiae	MN653051	MN653124	MN653092	MN656104
	Polynoinae	Paradyte crinoidicola	JN852837	JN852869	JN852907	JN852938
	Polynoinae	Paralepidonotus ampulliferus	JN852838	AF185164	JN852908	JN852939
	Macellicephalinae	Pelagomacellicephala cf. iliffei 2	KY454408	KY823474	KY454420	KY454435
	Macellicephalinae	Pelagomacellicephala cf. iliffei 3	KY454411	KY823475	KY454428	KY454443
	Macellicephalinae	Pelagomacellicephala cf. iliffei 4	KY454405	—	KY454416	KY454431
	Polynoinae	Polyeunoa laevis	KF713424	KU738193	KF713464	KU738213.1
	Polynoinae	Polynoe scolopendrina	JN852839	JN852870	JN852909	JN852940
	Polynoinae	Robertianella synophthalma	MN653053	MN653126	MN653122	MN656132
	Lepidonotinae	Thormora jukesii	JN852840	JN852871	JN852910	JN852941
	Polynoinae ?	Polynoidae sp. 2 ST 2019	MN653052	MN653125	MN653121	MN656131
	Polynoinae	Subadyte pellucida	—	—	—	KX946868.1
Sigalionidae		Neoleanira tetragona	AY839570	JN852872	JN852911	AY839582
		Psammolyce sp.	JN852843	JN852877	JN852916	—
		Sigalion bandaensis	AB106254	AF185165	—	AY583699
		Sigalion spinosus	AY894304	DQ790062	—	AY894319
		Sthenelais boa	AY839575	DQ779711	DQ779635	AY839587
		Sthenelais limicola	JN852844	JN852878	—	—
		Sthenelanella uniformis	AY894306	DQ790064	—	AY894322

2 Morphological comparison of Genera related to Ceuthonoe gen. n. Unless otherwise indicated, character information is from original descriptions and illustrations

No.	Genus	Num. of segments— elytrae pairs	Facial tubercle	Chaetae on tentaculophore	Cephalic peaks	Bidentate tip of neurochaetae	supraacicular process	neuroacicula penetrating epidermis	Symbiotic relations	Ref.
Note: + indicates with, − indicates without, ? indicates unknown information, and * indicates monotypic genera. Symbiotic relations referencing to Martin and Britayev (1998, 2018).
1	Antarctinoe	<50; 15	+	+	+	−	+	+		Barnich et al. (2006)
2	Australonoe*	up to 39; 16	+	+	+	+	+	?		Hanley (1993)
3	Bathynotalia*	21; 11	?	−	+	−	−	−		Levenstein (1982)
4	Bylgides	up to 38; 14/15	?	+/−	+	−	+	?		Pettibone (1993b)
5	Eucranta	38–40; 15	?	?	+	+	?	?		Uschakov (1982)
6	Eunoe	34–40; 15	?	+/−	+/−	−	?	?	E. opalina commensal with Synallactida	Uschakov (1982)
7	Gattyana	34–40; 15	?	+/−	+/−	−	?	?	G. spp. commensal with other polychaetes	Uschakov (1982)
8	Harmothoe	up to 50; 15	+	+	+	+	+	+	H. spp. commensal with Alcyonacea, polychaete, decapod, Bivalve, Sponge	Barnich et al. (2006)
9	Hermadion	up to 50; 15	?	+	−	−	−	+		Bock et al. (2010)
10	Hermenia	26; 12	+	+	−	+	−	−		Pettibone (1975)
11	Hololepidella	>55; 20–55; less	?	+/−	+	+	?	?	commensal with Ochrophyta and Scleractinia	Wehe (2006)
12	Intoshella	up to 50; up to 23	−	+	−	+	+	?	commensal with Holothuroid, Sponge	Sui et al. (2019)
13	Leucia	up to 50; 16	−	+	+	+	+	+	L. violacea commensal with Scleractinia	Fiege and Barnich (2009)
14	Malmgreniella	up to 46; 14/15	?	+/−	−	+/−	+	?	M. spp. commensal with Holothuroid, Ophiuroid, Polychaete, Sipunculid, Echinoid, Asteroid, Cnidarian, Balanoglossid, Decapod and Sponge	Pettibone (1993a)
15	Neopolynoe	up to 59; 15	−	+/−	+	−	+	+	N. acanellae commensal with Cnidarian and Sponge; N. paradoxa commensal with Cnidarian	Bock et al. (2010)
16	Paralepidonotus	36–41; 15	+	+	−	+/−	+/−	+/−		Hanley (1991)
17	Parahalosydna	reduced number of segments; 15	−	−	−	−	−	?		Jirkov (2001)
18	Ceuthonoe gen. n.	32; 14	+	+	+	−	−	+	commensal with glass sponge Placopegma sp.	this study

A1 GTR_corrrected distance between genera

	Acholoe astericola	Alentia gelatinosa	Antarctinoe ferox	Antinoe aequiseta	Antipathipoly-eunoa sp.	Austrolaenilla antarctica	Bathykurila guaymasensis	Branchinotog-luma sandersi	Branchipolynoe symmytilida	Bylgides elegans	Enipo torelli
Acholoe astericola
Alentia gelatinosa	0.269 511 76
Antarctinoe ferox	0.231 803 84	0.240 436 13
Antinoe aequiseta	0.230 661 56	0.240 084 59	0.238 238 74
Antipathipolyeunoa sp.	0.223 309 43	0.224 370 71	0.196 251 31	0.200 569 18
Austrolaenilla antarctica	0.202 646 68	0.236 284 87	0.185 219 68	0.194 404 15	0.195 290 08
Bathykurila guaymasensis	0.274 921 18	0.337 522 95	0.287 268 84	0.287 884 17	0.294 589 25	0.270 288 77
Branchinotogluma sandersi	0.281 914 74	0.271 519 12	0.270 206 93	0.260 584 28	0.270 202 27	0.244 822 91	0.252 109 06
Branchipolynoe symmytilida	0.309 160 94	0.284 317 45	0.271 797 33	0.315 925 49	0.289 288 93	0.252 788 07	0.227 130 61	0.283 901 42
Bylgides elegans	0.226 497 14	0.226 630 04	0.211 128 53	0.222 389 21	0.220 205 22	0.186 782 59	0.299 272 87	0.246 542 57	0.319 554 30
Enipo torelli	0.214 481 27	0.267 762 17	0.221 269 04	0.177 320 96	0.237 277 14	0.201 776 54	0.300 720 36	0.255 073 55	0.347 967 32	0.198 563 52
Eunoe nodosa	0.197 076 00	0.266 600 16	0.199 355 95	0.167 388 09	0.179 691 39	0.183 210 39	0.277 403 25	0.236 569 86	0.298 012 57	0.219 793 65	0.173 788 99
Gastrolepidia clavigera	0.242 957 88	0.290 416 73	0.259 805 33	0.237 094 81	0.244 147 70	0.215 785 44	0.287 618 36	0.291 316 93	0.329 487 16	0.246 373 89	0.240 090 99
Gattyana cirrhosa	0.212 864 51	0.244 646 38	0.239 538 42	0.122 068 74	0.176 761 36	0.184 642 70	0.269 115 61	0.267 278 54	0.287 962 53	0.200 045 79	0.204 698 22
Gorgoniapolynoe corralophila	0.193 333 76	0.260 437 08	0.192 422 22	0.188 417 29	0.135 026 24	0.166 976 60	0.268 311 74	0.282 361 87	0.280 199 88	0.223 725 86	0.173 564 55
Grubeopolynoe tuta	0.220 858 89	0.267 683 01	0.198 286 67	0.195 345 06	0.195 528 71	0.175 008 95	0.288 466 02	0.241 374 59	0.303 052 61	0.188 509 19	0.165 281 12
Halosydna brevisetosa	0.246 788 18	0.238 669 65	0.242 028 98	0.233 554 11	0.245 726 78	0.230 254 20	0.301 610 36	0.249 249 98	0.296 928 61	0.227 365 33	0.234 313 35
Harmothoe glabra	0.192 352 03	0.244 530 07	0.216 255 57	0.191 693 62	0.209 978 24	0.184 001 30	0.280 242 77	0.275 991 52	0.294 245 75	0.170 236 82	0.214 601 17
Hermadion truncata	0.229 504 67	0.251 499 76	0.219 228 82	0.159 241 33	0.208 486 08	0.190 918 68	0.308 311 79	0.293 185 09	0.322 352 42	0.232 435 32	0.156 596 02
Hermenia verruculosa	0.305 168 84	0.269 749 68	0.256 179 24	0.295 978 15	0.281 563 30	0.282 018 94	0.344 430 19	0.280 998 35	0.337 009 88	0.291 506 20	0.284 984 95
Hyperhalosydna striata	0.273 750 83	0.231 817 83	0.235 834 70	0.278 617 54	0.244 322 85	0.241 693 28	0.307 256 22	0.264 947 19	0.309 317 45	0.235 659 74	0.268 375 32
Intoshella dictyaulus	0.194 629 57	0.265 175 37	0.216 771 35	0.221 247 99	0.189 103 76	0.196 123 93	0.279 661 45	0.281 273 84	0.268 856 26	0.215 814 17	0.197 426 94
Iphione sp.	0.316 665 68	0.361 590 51	0.331 609 20	0.334 787 72	0.317 127 13	0.306 283 23	0.299 539 01	0.309 792 58	0.294 255 56	0.383 103 97	0.344 926 36
Lepidasthenia elegans	0.258 976 51	0.227 797 27	0.207 000 92	0.243 207 28	0.212 362 65	0.178 575 62	0.285 072 42	0.277 965 74	0.300 226 39	0.237 199 40	0.259 237 35
Lepidonotus clava	0.247 554 50	0.242 824 25	0.208 851 81	0.243 305 83	0.213 591 09	0.187 816 39	0.288 998 41	0.277 081 70	0.305 522 14	0.233 925 37	0.261 132 73
Leucia violacea	0.194 010 48	0.243 055 26	0.226 959 85	0.194 385 23	0.166 102 50	0.175 617 13	0.260 462 13	0.246 952 91	0.297 629 16	0.209 694 44	0.186 552 32
Malmgreniella mcintoshi	0.232 223 17	0.246 995 25	0.202 821 73	0.185 027 50	0.193 855 30	0.158 548 82	0.285 923 59	0.275 636 02	0.318 913 51	0.201 501 20	0.198 394 53
Melaenis loveni	0.244 679 84	0.246 257 21	0.232 654 72	0.179 744 00	0.195 361 77	0.189 717 60	0.274 822 25	0.242 988 83	0.296 000 19	0.218 078 70	0.190 774 79
Neopolynoe paradoxa	0.211 068 22	0.287 160 42	0.187 489 18	0.213 484 61	0.204 421 34	0.170 350 44	0.290 913 03	0.258 700 07	0.277 083 64	0.210 487 41	0.177 050 14
Paradyte crinoidicola	0.207 096 61	0.276 264 54	0.216 611 69	0.187 294 12	0.169 536 94	0.185 016 15	0.258 660 71	0.243 733 42	0.281 107 32	0.185 766 25	0.213 930 20
Paralepidonotus ampulliferus	0.274 653 67	0.298 324 79	0.259 376 15	0.271 419 99	0.235 694 85	0.258 323 07	0.294 102 04	0.288 761 42	0.303 365 44	0.244 609 84	0.286 540 02
Polyeunoa laevis	0.258 812 32	0.261 462 18	0.186 313 11	0.190 165 03	0.171 548 02	0.221 625 88	0.287 715 35	0.264 763 76	0.312 900 01	0.221 275 26	0.240 161 40
Polynoe scolopendrina	0.230 075 01	0.232 860 97	0.227 610 52	0.245 412 48	0.246 152 12	0.216 098 89	0.304 536 93	0.288 410 09	0.294 613 49	0.234 203 21	0.233 852 75
Subadyte pellucida	0.204 346 96	0.260 655 32	0.240 490 63	0.235 773 84	0.219 236 64	0.217 392 50	0.301 267 81	0.313 934 90	0.305 682 76	0.236 602 34	0.243 015 01
Thermiphione sp.	0.283 893 21	0.249 393 81	0.255 527 20	0.294 418 02	0.277 039 43	0.248 392 38	0.300 535 92	0.291 855 58	0.288 677 19	0.285 407 60	0.272 284 18
Thormora jukesii	0.266 083 22	0.256 615 03	0.256 609 22	0.283 712 09	0.258 401 10	0.279 393 90	0.308 337 30	0.282 862 82	0.333 650 04	0.263 473 57	0.275 364 88
Ceuthonoe nezhai gen. et sp. n.	0.277 990 39	0.235 979 10	0.249 207 93	0.300 987 09	0.281 188 34	0.243 551 99	0.337 392 74	0.274 951 94	0.324 302 20	0.247 913 46	0.300 167 41
Gesiella jameensis	0.252 124 85	0.300 859 60	0.272 240 16	0.239 164 15	0.279 897 28	0.221 936 89	0.272 362 16	0.308 544 64	0.290 703 32	0.279 877 94	0.249 943 21
Pelagomacellicephala cf. iliffei 2	0.264 930 78	0.258 735 24	0.276 788 20	0.287 890 34	0.248 698 64	0.278 013 05	0.330 923 79	0.310 294 94	0.345 141 98	0.299 767 44	0.284 886 32
Halosydnella australis	0.252 941 22	0.234 158 89	0.244 374 41	0.236 259 02	0.242 185 79	0.256 159 53	0.288 168 04	0.237 212 48	0.267 194 41	0.237 171 24	0.251 627 76
Polynoidae sp. 2 ST 2019	0.253 565 13	0.247 879 10	0.231 397 79	0.287 037 04	0.257 491 59	0.256 217 50	0.302 787 53	0.285 703 69	0.284 474 27	0.253 523 10	0.271 653 55
Robertianella synophthalma	0.210 912 34	0.220 457 62	0.203 697 34	0.194 457 67	0.172 252 91	0.182 272 99	0.272 772 77	0.254 813 47	0.286 825 3	0.208 351 71	0.169 457 35

	Eunoe nodosa	Gastrolepidia clavigera	Gattyana cirrhosa	Gorgonia-polynoe corralophila	Grubeopolynoe tuta	Halosydna brevisetosa	Harmothoe glabra	Hermadion truncata	Hermenia verruculosa	Hyperhalosydna striata	Intoshella dictyaulus
Acholoe astericola
Alentia gelatinosa
Antarctinoe ferox
Antinoe aequiseta
Antipathipolyeunoa sp.
Austrolaenilla antarctica
Bathykurila guaymasensis
Branchinotogluma sandersi
Branchipolynoe symmytilida
Bylgides elegans
Enipo torelli
Eunoe nodosa
Gastrolepidia clavigera	0.230 553 42
Gattyana cirrhosa	0.176 298 19	0.234 438 00
Gorgoniapolynoe corralophila	0.182 612 24	0.198 853 27	0.177 961 08
Grubeopolynoe tuta	0.184 155 78	0.221 042 24	0.182 638 65	0.204 819 49
Halosydna brevisetosa	0.252 711 79	0.286 497 41	0.253 979 65	0.236 245 97	0.247 410 81
Harmothoe glabra	0.191 495 70	0.248 724 62	0.187 712 73	0.175 291 04	0.203 279 96	0.253 316 97
Hermadion truncata	0.191 894 05	0.223 757 95	0.185 233 86	0.208 363 62	0.157 801 96	0.248 028 91	0.211 556 72
Hermenia verruculosa	0.286 394 57	0.313 060 00	0.296 280 09	0.276 232 50	0.250 622 43	0.267 142 73	0.315 345 25	0.269 289 63
Hyperhalosydna striata	0.260 873 72	0.277 651 58	0.249 057 91	0.260 622 67	0.223 436 46	0.221 559 62	0.270 354 62	0.256 691 90	0.251 614 99
Intoshella dictyaulus	0.183 429 46	0.225 124 80	0.197 763 21	0.190 982 56	0.172 953 66	0.243 018 05	0.202 968 68	0.200 585 24	0.275 599 46	0.242 327 43
Iphione sp.	0.324 520 94	0.331 518 98	0.315 205 02	0.313 046 18	0.294 201 05	0.310 234 10	0.337 045 01	0.297 278 36	0.263 715 40	0.332 025 36	0.291 977 33
Lepidasthenia elegans	0.223 113 99	0.256 619 95	0.203 475 14	0.211 745 18	0.209 645 53	0.197 876 74	0.237 512 87	0.251 528 72	0.258 954 31	0.202 483 54	0.215 940 27
Lepidonotus clava	0.204 499 44	0.254 313 36	0.224 049 89	0.213 950 69	0.207 991 33	0.231 825 58	0.235 439 42	0.256 479 15	0.257 271 52	0.199 815 05	0.226 808 59
Leucia violacea	0.194 010 48	0.243 055 26	0.226 959 85	0.194 385 23	0.166 102 50	0.175 617 13	0.260 462 13	0.246 952 91	0.297 629 16	0.209 694 44	0.186 552 32
Malmgreniella mcintoshi	0.196 958 40	0.242 155 33	0.207 399 27	0.159 617 90	0.198 844 16	0.211 095 15	0.197 814 93	0.206 854 23	0.304 734 97	0.259 651 52	0.195 079 98
Melaenis loveni	0.199 703 28	0.224 598 57	0.188 333 16	0.173 354 66	0.185 409 05	0.235 414 95	0.195 684 58	0.195 908 72	0.287 408 57	0.258 211 50	0.218 006 00
Neopolynoe paradoxa	0.217 263 64	0.240 369 26	0.214 145 35	0.146 993 96	0.183 532 45	0.213 418 75	0.203 498 14	0.191 350 88	0.279 434 65	0.245 411 80	0.222 117 69
Paradyte crinoidicola	0.175 448 28	0.221 621 43	0.165 450 59	0.164 970 01	0.198 606 51	0.208 852 84	0.184 042 66	0.228 521 74	0.315 447 07	0.224 258 82	0.210 535 58
Paralepidonotus ampulliferus	0.249 283 83	0.318 035 72	0.242 480 56	0.268 899 62	0.190 062 62	0.297 300 20	0.265 164 31	0.240 878 53	0.293 344 58	0.277 046 47	0.234 892 15
Polyeunoa laevis	0.184 910 02	0.244 840 93	0.176 401 85	0.167 527 27	0.200 012 62	0.233 991 51	0.189 429 18	0.218 635 03	0.273 161 74	0.239 704 14	0.210 475 10
Polynoe scolopendrina	0.244 755 21	0.268 659 78	0.247 842 24	0.211 480 47	0.250 635 47	0.245 013 12	0.257 187 26	0.246 196 07	0.324 827 68	0.260 268 20	0.237 975 66
Subadyte pellucida	0.233 765 57	0.250 280 36	0.193 058 37	0.167 634 48	0.206 039 06	0.275 731 28	0.198 133 89	0.253 302 23	0.285 443 58	0.285 458 40	0.217 177 49
Thermiphione sp.	0.276 234 08	0.263 181 65	0.265 361 85	0.275 164 57	0.272 733 23	0.255 222 52	0.298 458 60	0.264 910 80	0.224 641 27	0.246 763 16	0.256 312 08
Thormora jukesii	0.268 978 33	0.217 538 43	0.270 850 39	0.266 804 65	0.266 917 31	0.238 240 83	0.293 402 84	0.278 678 68	0.218 003 77	0.218 974 20	0.262 411 77
Ceuthonoe nezhai gen. et sp. n.	0.276 396 21	0.305 799 82	0.286 475 96	0.284 129 43	0.270 907 13	0.244 206 38	0.242 056 05	0.277 236 82	0.263 989 56	0.283 160 18	0.290 976 75
Gesiella jameensis	0.254 264 96	0.262 184 39	0.227 858 62	0.238 921 91	0.242 303 75	0.294 418 97	0.234 671 26	0.234 262 95	0.378 684 02	0.278 753 18	0.225 928 92
Pelagomacellicephala cf. iliffei 2	0.290 277 05	0.299 494 26	0.276 459 00	0.226 452 54	0.314 473 22	0.261 203 35	0.281 603 88	0.288 750 13	0.312 755 78	0.273 650 22	0.306 710 16
Halosydnella australis	0.232 425 03	0.286 432 11	0.249 430 97	0.273 323 00	0.220 579 49	0.196 589 56	0.282 000 77	0.249 248 59	0.240 995 10	0.221 562 33	0.260 490 71
Polynoidae sp. 2 ST 2019	0.276 219 07	0.264 289 79	0.262 096 38	0.270 379 22	0.242 365 35	0.241 605 94	0.237 146 06	0.216 446 34	0.256 813 00	0.255 874 27	0.268 627 51
Robertianella synophthalma	0.174 658 16	0.221 806 28	0.181 413 18	0.141 881 76	0.190 976 75	0.223 334 57	0.188 236 68	0.196 780 63	0.271 056 61	0.246 077 83	0.182 762 73

	Iphione sp.	Lepidasthenia elegans	Lepidonotus clava	Leucia violacea	Malmgreniella mcintoshi	Melaenis loveni	Neopolynoe paradoxa	Paradyte crinoidicola	Paralepidonotus ampulliferus	Polyeunoa laevis	Polynoe scolopendrina
Acholoe astericola
Alentia gelatinosa
Antarctinoe ferox
Antinoe aequiseta
Antipathipolyeunoa sp.
Austrolaenilla antarctica
Bathykurila guaymasensis
Branchinotogluma sandersi
Branchipolynoe symmytilida
Bylgides elegans
Enipo torelli
Eunoe nodosa
Gastrolepidia clavigera
Gattyana cirrhosa
Gorgoniapolynoe corralophila
Grubeopolynoe tuta
Halosydna brevisetosa
Harmothoe glabra
Hermadion truncata
Hermenia verruculosa
Hyperhalosydna striata
Intoshella dictyaulus
Iphione sp.
Lepidasthenia elegans	0.305 133 86
Lepidonotus clava	0.327 080 51	0.069 029 20
Leucia violacea	0.325 422 76	0.220 475 25	0.227 334 56
Malmgreniella mcintoshi	0.335 880 20	0.224 442 19	0.243 446 83	0.166 212 01
Melaenis loveni	0.306 347 04	0.217 783 64	0.213 898 80	0.156 596 54	0.201 151 53
Neopolynoe paradoxa	0.317 490 91	0.223 078 62	0.234 389 84	0.167 223 87	0.169 209 05	0.183 982 47
Paradyte crinoidicola	0.352 818 84	0.210 381 11	0.226 220 33	0.197 551 57	0.218 904 31	0.204 414 55	0.183 302 31
Paralepidonotus ampulliferus	0.350 436 40	0.276 609 39	0.267 248 55	0.241 540 63	0.286 180 13	0.261 012 11	0.242 734 69	0.254 090 31
Polyeunoa laevis	0.327 543 47	0.212 394 33	0.224 644 00	0.188 466 18	0.208 599 90	0.192 117 86	0.201 721 11	0.208 898 45	0.266 239 14
Polynoe scolopendrina	0.379 922 59	0.240 515 68	0.250 207 72	0.226 628 55	0.200 161 20	0.263 861 77	0.218 945 65	0.218 053 34	0.331 403 56	0.256 973 49
Subadyte pellucida	0.343 507 42	0.228 415 03	0.214 772 70	0.219 804 12	0.225 842 28	0.237 269 98	0.255 722 32	0.208 584 10	0.260 675 34	0.219 229 36	0.234 952 52
Thermiphione sp.	0.263 006 49	0.258 821 49	0.266 638 56	0.275 063 22	0.288 452 76	0.259 538 45	0.265 448 72	0.258 043 64	0.302 945 23	0.255 540 32	0.316 058 18
Thormora jukesii	0.299 040 69	0.226 966 15	0.245 154 99	0.259 031 84	0.275 049 80	0.256 666 12	0.293 642 37	0.268 555 73	0.314 272 24	0.248 903 11	0.289 059 08
Ceuthonoe nezhai gen. et sp. n.	0.309 143 32	0.238 719 40	0.262 519 18	0.227 254 55	0.264 798 05	0.261 822 39	0.283 928 11	0.282 577 95	0.333 408 55	0.277 422 08	0.271 520 25
Gesiella jameensis	0.324 869 84	0.255 000 27	0.246 016 98	0.229 823 80	0.241 013 36	0.248 550 36	0.229 394 51	0.225 900 92	0.276 710 05	0.284 447 17	0.278 292 97
Pelagomacellicephala cf. iliffei 2	0.280 079 28	0.232 513 74	0.242 258 70	0.268 277 13	0.305 783 34	0.246 061 90	0.277 972 64	0.277 033 53	0.334 771 16	0.264 822 34	0.299 314 67
Halosydnella australis	0.296 441 67	0.234 982 59	0.229 825 03	0.230 775 49	0.267 737 29	0.249 162 84	0.275 053 16	0.262 852 90	0.270 092 62	0.261 288 93	0.249 387 30
Polynoidae sp. 2 ST 2019	0.307 291 97	0.258 075 09	0.260 563 01	0.256 598 64	0.266 367 21	0.276 010 20	0.266 913 52	0.278 854 36	0.290 574 60	0.245 405 36	0.262 331 06
Robertianella synophthalma	0.301 457 84	0.231 979 14	0.234 923 17	0.172 299 41	0.184 401 70	0.168 897 56	0.167 438 32	0.187 523 51	0.215 383 11	0.184 582 03	0.228 736 51

	Subadyte pellucida	Thermiphione sp.	Thormora jukesii	Ceuthonoe nezhai gen. et sp. n.	Gesiella jameensis	Pelagomacellic-ephala cf. iliffei 2	Halosydnella australis	Polynoidae sp. 2 ST 2019
Acholoe astericola
Alentia gelatinosa
Antarctinoe ferox
Antinoe aequiseta
Antipathipolyeunoa sp.
Austrolaenilla antarctica
Bathykurila guaymasensis
Branchinotogluma sandersi
Branchipolynoe symmytilida
Bylgides elegans
Enipo torelli
Eunoe nodosa
Gastrolepidia clavigera
Gattyana cirrhosa
Gorgoniapolynoe corralophila
Grubeopolynoe tuta
Halosydna brevisetosa
Harmothoe glabra
Hermadion truncata
Hermenia verruculosa
Hyperhalosydna striata
Intoshella dictyaulus
Iphione sp.
Lepidasthenia elegans
Lepidonotus clava
Leucia violacea
Malmgreniella mcintoshi
Melaenis loveni
Neopolynoe paradoxa
Paradyte crinoidicola
Paralepidonotus ampulliferus
Polyeunoa laevis
Polynoe scolopendrina
Subadyte pellucida
Thermiphione sp.	0.304 207 13
Thormora jukesii	0.279 639 16	0.214 375 32
Ceuthonoe nezhai gen. et sp. n.	0.274 900 97	0.253 297 56	0.291 380 26
Gesiella jameensis	0.252 228 82	0.308 232 42	0.360 050 13	0.316 815 13
Pelagomacellicephala cf. iliffei 2	0.299 009 24	0.291 921 97	0.284 222 39	0.279 183 68	0.286 840 27
Halosydnella australis	0.270 208 81	0.231 993 28	0.238 275 86	0.260 840 91	0.315 887 72	0.311 794 26
Polynoidae sp. 2 ST 2019	0.267 175 30	0.235 863 10	0.269 466 07	0.214 625 26	0.241 997 03	0.292 326 13	0.216 593 34
Robertianella synophthalma	0.192 200 27	0.251 141 20	0.263 141 64	0.272 085 73	0.246 861 48	0.255 376 93	0.244 626 42	0.232 877 43

Fig. 1. Phylogeny of polynoids based on the concatenated alignments of four genes using BI and ML analyses, imposed on the consensus ML tree. Node supports are shown as ML bootstrap values (BP, above) and Bayesian posterior probabilities (PP, below). Nodes not recovered in the BI analysis are marked with --.

Fig. 2. Ceuthonoe nezhai gen. et sp. n.: Holotype, dorsal view (a), Paratype, ventral view (b), Elytra from Segment 2 (c), Appendages of prostomium from holotype and paratype (d and e), Dorsal view of prostomium (first pair of elytra removed) (f) Median segments, ventral view (g), anal segment, dorsal and ventral view (h and i), Dorsal cirrus (j), Lateral antenna (k), Dorsal view of jaws from paratype (l), and Parapodium of Segment 8, end view (m). dc: dorsal cirrus, dtc: dorsal tentacular cirrus, ep: eltrophore, ft: facial tubercle, laa: lateral antenna, lac: lateral antenna ceratophore, mac: median antenna ceratophore, np: nephridial papilla, pal: palp, vc: ventral cirrus, vtc: ventral tentacular cirrus, 2vc: ventral cirrus on Segment 2, and 3vc: ventral cirrus on Segment 3. Scale bars: 2.0 mm (a, b, k), 0.2 mm (c–e), 0.5 mm (f–j), and 1.0 mm (m).

Fig. 3. Ceuthonoe nezhai gen. et sp. n. Holotype. a. Prostomium, b. Parapodium of Segment 8 (chaetae omitted), end view, c. Tubercles on elytra, d. Elytra, e. Notochaeta and detailed view, f. Notoacicula, and g. Neurochaeta and detailed view. e–g are views under light microscope. Scale bars: 0.2 mm (a–d), 0.1 mm (e–g).

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