taxonID	type	description	language	source
563E8793D242FFDFFF26F8EEFA79CC9B.taxon	description	(Figs 3 – 5)	en	Hookabe, Natsumi, Yamamori, Luna, Ogawa, Akito, Jimi, Naoto, Kohtsuka, Hisanori, Rouse, Greg W. (2025): An updated phylogeny of Iphitime (Annelida: Dorvilleidae) revealing multiple host switching, with description of I. nubila sp. nov. Zoological Journal of the Linnean Society 204 (4): 1-16, DOI: 10.1093/zoolinnean/zlaf101, URL: https://doi.org/10.1093/zoolinnean/zlaf101
563E8793D242FFDFFF26F8EEFA79CC9B.taxon	vernacular_names	Japanese name: unryû-mino-isome.	en	Hookabe, Natsumi, Yamamori, Luna, Ogawa, Akito, Jimi, Naoto, Kohtsuka, Hisanori, Rouse, Greg W. (2025): An updated phylogeny of Iphitime (Annelida: Dorvilleidae) revealing multiple host switching, with description of I. nubila sp. nov. Zoological Journal of the Linnean Society 204 (4): 1-16, DOI: 10.1093/zoolinnean/zlaf101, URL: https://doi.org/10.1093/zoolinnean/zlaf101
563E8793D242FFDFFF26F8EEFA79CC9B.taxon	description	ZooBank: urn: lsid: zoobank. org: act: 55 B 4 F 2 D 2 - FD 69 - 43 E 9 - 96 DE- 3 DCAD 7 BE 9 FAD.	en	Hookabe, Natsumi, Yamamori, Luna, Ogawa, Akito, Jimi, Naoto, Kohtsuka, Hisanori, Rouse, Greg W. (2025): An updated phylogeny of Iphitime (Annelida: Dorvilleidae) revealing multiple host switching, with description of I. nubila sp. nov. Zoological Journal of the Linnean Society 204 (4): 1-16, DOI: 10.1093/zoolinnean/zlaf101, URL: https://doi.org/10.1093/zoolinnean/zlaf101
563E8793D242FFDFFF26F8EEFA79CC9B.taxon	materials_examined	Materials examined: Holotype (NSMT-Pol H- 986): female, fixed / preserved in 5 % formalin in seawater, collected off Minabe, Wakayama, Japan, on 2 December 2021 from Dardanus crassimanus with a shell of Tutufa bufo (Röding, 1798) (Fig. 3 A, B) caught with a gill net at depths of 5 – 50 m; several branchiae were fixed and preserved in 99 % ethanol. Paratype 1 (NSMT-Pol P- 987): male, fixed / preserved in 10 % formalin in seawater, collection date, location, and host individual same with the holotype. Paratype 2 (NSMT-Pol P- 988), female, fixed / preserved in 70 % ethanol, collected off Minabe, Wakayama, Japan, on 9 December, from D. crassimanus with a shell of T. bufo caught with a gill net at depths of 5 – 50 m; Paratype 3 (NSMT-Pol P- 989), female, fixed / preserved in 70 % ethanol, collected off Minabe, Wakayama, Japan, on 9 December, from Dardanus pedunculatus with a shell of Charonia lampas (Linnaeus, 1758) caught with a gill net at depths of 5 – 50 m; Paratype 4 (NSMT-Pol P- 990), female, fixed / preserved in 70 % ethanol, collected off Minabe, Wakayama, Japan, on 9 December, collected from D. pedunculatus with a shell of Tonna luteostoma (Küster, 1857) caught with a gill net at depths of 5 – 50 m, head region dissected for observation of jaw morphology. Exact coordinates of sampling localities unavailable.	en	Hookabe, Natsumi, Yamamori, Luna, Ogawa, Akito, Jimi, Naoto, Kohtsuka, Hisanori, Rouse, Greg W. (2025): An updated phylogeny of Iphitime (Annelida: Dorvilleidae) revealing multiple host switching, with description of I. nubila sp. nov. Zoological Journal of the Linnean Society 204 (4): 1-16, DOI: 10.1093/zoolinnean/zlaf101, URL: https://doi.org/10.1093/zoolinnean/zlaf101
563E8793D242FFDFFF26F8EEFA79CC9B.taxon	diagnosis	Diagnosis: Prostomium with two antennae protruding forward. Branchiae cirriform without branching, present from chaetiger 1, and inserted dorsally. Parapodia uniramous with 3 – 5 supraacicular simple chaetae, 6 – 10 subacicular compound chaetae, and 1 or 2 slender capillary chaetae. Pygidium with two cirri. Sex dimorphism present; female jaws P-type; male jaws K-type.	en	Hookabe, Natsumi, Yamamori, Luna, Ogawa, Akito, Jimi, Naoto, Kohtsuka, Hisanori, Rouse, Greg W. (2025): An updated phylogeny of Iphitime (Annelida: Dorvilleidae) revealing multiple host switching, with description of I. nubila sp. nov. Zoological Journal of the Linnean Society 204 (4): 1-16, DOI: 10.1093/zoolinnean/zlaf101, URL: https://doi.org/10.1093/zoolinnean/zlaf101
563E8793D242FFDFFF26F8EEFA79CC9B.taxon	description	Description: Body flattened, tapered in posterior region (Fig. 3 C, D). Prostomium rounded without eyespots. Two digitiform antennae dorsally protruding forward on prostomium; mature female antennae long, extending beyond prostomium (Fig. 3 E); male antennae short, not extending beyond prostomium (Fig. 3 D). Peristomium represented by two apodous achaetous rings, 1 st ring slightly longer than 2 nd ring (Fig. 3 D, E). Palps absent. Chaetae present at first segment after peristomium (Fig. 3 E). Branchiae cirriform inserted dorsally (Fig. 3 F) and present from chaetiger 1 (Fig. 3 E). Parapodia uniramous with a rounded acicular lobe and papilliform dorsal cirri (Fig. 4 A, B). Three kinds of chaetae present; 3 – 5 supraacicular simple chaetae with hook-shaped tips (Fig. 4 D, F); 7 – 10 subacicular compound chaetae slightly serrated (Fig. 4 E, G); at ventralmost, 1 – 3 slender capillary chaetae curved up dorsally or just extending laterally (Fig. 4 A, B). Paired pygidial cirri digitiform shorter than antennae (Fig. 3 G). Female (holotype): 10.6 cm in length, 3.0 mm in maximum width for 403 chaetigers (Fig. 3 C). Two long digitiform antennae 2.1 mm in length, extending beyond prostomium (Fig. 3 E). Branchiae first digitiform, tapering and becoming longer in mid-body but shorter in the posterior 20 segments (Fig. 3 C). Chaetae comprising of 3 supraacicular simple, 9 or 10 subacicular compound, and 1 – 3 slender capillary chaetae (Fig. 4 A, C – E). Subacicular compound chaetae more dorsally socketed in posterior part of body. Paired short pygidial cirri about 0.9 mm in length (Fig. 3 G), shorter than antennae (Fig. 3 E). Mandibles lost during preparation. P-type maxillae 490 μm in length with six pairs of denticles fused with each other; maxilla I with at least eight teeth, maxilla II with at least six teeth, and maxilla III with at least six coarse teeth variously developed (Fig. 5 A, B). The posteriormost denticles (maxillae III – VI) fused with each other (Fig. 5 A, B). Eggs about 120 μm diameter found between the branchiae (Fig. 3 C). Male (paratype 1): 4.0 mm in length, 1.2 mm in maximum width for 79 chaetigers (Fig. 3 D). Two short digitiform antennae 0.5 mm in length, without extending beyond prostomium (Fig. 3 D). Branchiae first digitiform, tapering and becoming longer in mid-body but shorter in the posterior 20 segments (Fig. 3 D). Chaetae comprising of 5 supraacicular simple, 6 subacicular compound, and 1 or 2 slender capillary chaetae (Fig. 4 B, F, G). Pygidial cirri lost. Rod-like mandibles 350 μm in length, strongly sclerotized, comprising of two shafts distally widening and smooth; distal cutting edge without serration (Fig. 5 C, D). K-type maxillae 600 μm in length, with smooth ice tong-shaped forceps and at least one free-denticle (Fig. 5 C, D).	en	Hookabe, Natsumi, Yamamori, Luna, Ogawa, Akito, Jimi, Naoto, Kohtsuka, Hisanori, Rouse, Greg W. (2025): An updated phylogeny of Iphitime (Annelida: Dorvilleidae) revealing multiple host switching, with description of I. nubila sp. nov. Zoological Journal of the Linnean Society 204 (4): 1-16, DOI: 10.1093/zoolinnean/zlaf101, URL: https://doi.org/10.1093/zoolinnean/zlaf101
563E8793D242FFDFFF26F8EEFA79CC9B.taxon	distribution	Distribution and habitat: The species is found inside the apex of shells carried by hermit crabs (D. pedunculatus and D. crassimanus) with small amounts of mud possibly host faeces (Fig. 3 A, B). They are only known from the type locality, off Minabe, Wakayama, Japan, at depths of 5 – 50 m. Two to four worms were found in a single shell, including a large female and multiple smaller males.	en	Hookabe, Natsumi, Yamamori, Luna, Ogawa, Akito, Jimi, Naoto, Kohtsuka, Hisanori, Rouse, Greg W. (2025): An updated phylogeny of Iphitime (Annelida: Dorvilleidae) revealing multiple host switching, with description of I. nubila sp. nov. Zoological Journal of the Linnean Society 204 (4): 1-16, DOI: 10.1093/zoolinnean/zlaf101, URL: https://doi.org/10.1093/zoolinnean/zlaf101
563E8793D242FFDFFF26F8EEFA79CC9B.taxon	etymology	Etymology: The specific epithet originates from the Latin word for ‘ cloudy’ or ‘ foggy’, inspired by the Cloud Dragon painting ‘ Unryû-zu’ on the ceiling of Tenryu-ji Temple in Kyoto, Japan. It describes the species’ prominent dorsal branchiae, reminiscent of dragon scales, and their attachment to the ventral surface inside their host hermit crab shells, reminiscent of the painting.	en	Hookabe, Natsumi, Yamamori, Luna, Ogawa, Akito, Jimi, Naoto, Kohtsuka, Hisanori, Rouse, Greg W. (2025): An updated phylogeny of Iphitime (Annelida: Dorvilleidae) revealing multiple host switching, with description of I. nubila sp. nov. Zoological Journal of the Linnean Society 204 (4): 1-16, DOI: 10.1093/zoolinnean/zlaf101, URL: https://doi.org/10.1093/zoolinnean/zlaf101
563E8793D242FFDFFF26F8EEFA79CC9B.taxon	discussion	Remarks The new species can be distinguished from other described species by combination of several morphological characters. The prostomium has two antennae protruding forward, in contrast to several dorvilleids. The branchiae are cirriform and lack branching, unlike I. cuenoti, I. doederleini, and I. loxorhynchi, which have branched branchiae. Additionally, the branchiae are present from chaetiger 1, whereas in I. paguri, I. loxorhynchi, I. holobranchiata (possible junior synonym of I. loxorhynchi; see below), Iphitime sartorae de Paiva & Nonato, 1991, and I. hartmanae, branchiae appear chaetigers 2 – 5. The branchiae are inserted dorsally, distinguishing the new species from I. cuenoti, I. doederleini, I. loxorhynchi, and I. holobranchiata, where the branchiae are inserted dorsolaterally. The number of chaetae, pygidium cirri, and the presence of sex dimorphism further differentiated the new species from the congeneric species. Iphitime nubila resembles I. paguri from Europe, though they are not closely related suggesting that having hermit crab hosts has evolved twice (Figs 1, 2). Both species have cirriform unbranched branchiae, exhibit sexual dimorphisms in jaw morphology, and utilize hermit crabs as hosts. However, I. nubila can be distinguished from I. paguri by the insertion position of the branchiae (Fig. 3 E); in I. nubila, they insert from chaetiger 1 (Fig. 3 E), whereas in I. paguri, they are present from chaetiger 4 – 5 (HØisaeter and Samuelsen 2006). The ML tree topology inferred that I. nubila was sister to I. loxorhynchi fromsouthernCalifornia (Fig. 1), althoughthesupportvalue was quite low. The two species can be distinguished by the insertion position of the branchiae (chaetiger 1 in I. nubila vs. 2 in I. loxorhynchi), and the numbers of supraacicular simple chaetae (3 – 5 in I. nubila vs. 20 in I. loxorhynchi) and subacicular compound chaetae (6 – 10 in I. nubila vs. 20 or more in I. loxorhynchi) (Hartman 1952). Furthermore, the branchiae of I. nubila are entirely cirriform without distal branching as reported in I. loxorhynchi (Hartman 1952).	en	Hookabe, Natsumi, Yamamori, Luna, Ogawa, Akito, Jimi, Naoto, Kohtsuka, Hisanori, Rouse, Greg W. (2025): An updated phylogeny of Iphitime (Annelida: Dorvilleidae) revealing multiple host switching, with description of I. nubila sp. nov. Zoological Journal of the Linnean Society 204 (4): 1-16, DOI: 10.1093/zoolinnean/zlaf101, URL: https://doi.org/10.1093/zoolinnean/zlaf101
563E8793D24DFFDCFCCDFC43FE86C995.taxon	description	(Figs 6 – 8)	en	Hookabe, Natsumi, Yamamori, Luna, Ogawa, Akito, Jimi, Naoto, Kohtsuka, Hisanori, Rouse, Greg W. (2025): An updated phylogeny of Iphitime (Annelida: Dorvilleidae) revealing multiple host switching, with description of I. nubila sp. nov. Zoological Journal of the Linnean Society 204 (4): 1-16, DOI: 10.1093/zoolinnean/zlaf101, URL: https://doi.org/10.1093/zoolinnean/zlaf101
563E8793D24DFFDCFCCDFC43FE86C995.taxon	vernacular_names	Japanese name: mino-isome.	en	Hookabe, Natsumi, Yamamori, Luna, Ogawa, Akito, Jimi, Naoto, Kohtsuka, Hisanori, Rouse, Greg W. (2025): An updated phylogeny of Iphitime (Annelida: Dorvilleidae) revealing multiple host switching, with description of I. nubila sp. nov. Zoological Journal of the Linnean Society 204 (4): 1-16, DOI: 10.1093/zoolinnean/zlaf101, URL: https://doi.org/10.1093/zoolinnean/zlaf101
563E8793D24DFFDCFCCDFC43FE86C995.taxon	materials_examined	Materials examined: NSMT-Pol 113618: fixed and preserved in 10 % formalin in seawater, collected in Sagami Bay, Kanagawa, Japan, on 2 December 2021 from Macrocheira kaempferi (Fig. 4 A, B) caught with a gill net set at depths of 250 – 300 m; NSMT-Pol 113619, fixed and preserved in 70 % ethanol, collection date and locality same above, jaws dissected. NSMT-Pol 113620, fixed and preserved in 70 % ethanol, on 22 September 2022, from M. kaempferi collected off Shirahama. Coordination for sampling localities unavailable.	en	Hookabe, Natsumi, Yamamori, Luna, Ogawa, Akito, Jimi, Naoto, Kohtsuka, Hisanori, Rouse, Greg W. (2025): An updated phylogeny of Iphitime (Annelida: Dorvilleidae) revealing multiple host switching, with description of I. nubila sp. nov. Zoological Journal of the Linnean Society 204 (4): 1-16, DOI: 10.1093/zoolinnean/zlaf101, URL: https://doi.org/10.1093/zoolinnean/zlaf101
563E8793D24DFFDCFCCDFC43FE86C995.taxon	description	Description Body flattened, tapered in posterior region; 15 – 21 mm in length and 1.6 mm in maximum width (Fig. 6 C). Prostomium rounded without eyespots (Fig. 6 D). Two short digitiform antennae 250 μm in length, dorsally protruding forward on prostomium, without extending beyond prostomium (Figs 6 D, 7 A, 8 A). Peristomium represented by two apodous achaetous rings, 1 st ring as long as 2 nd ring (Figs 6 D, 7 A). Chaetae present at first segment after peristomium (Figs 7 A, 8 A). Branchiae palmate inserted dorsolaterally, present from chaetiger 1 (Figs 6 D, E, 7 C, 8 A). A pair of pygidial cirri digitiform, 320 μm in length (Fig. 6 F), slightly longer than antennae (Fig. 6 D). Chaetae comprising of 2 – 4 supraacicular simple and 7 subacicular compound chaetae (Figs 7 C – E, 8 B – D). Subacicular compound chaetae with serration (Figs 7 E, 8 D). Jaws of K-type present (Fig. 6 B). Sex unknown in the specimen examined. Rod-like mandibles 240 μm in length, strongly sclerotized, comprising of two shafts distally widening and smooth; distal cutting edge without serration (Fig. 7 B). K-type maxillae 340 μm in length, with smooth ice tong-shaped forceps and paired free-denticles (Fig. 7 B).	en	Hookabe, Natsumi, Yamamori, Luna, Ogawa, Akito, Jimi, Naoto, Kohtsuka, Hisanori, Rouse, Greg W. (2025): An updated phylogeny of Iphitime (Annelida: Dorvilleidae) revealing multiple host switching, with description of I. nubila sp. nov. Zoological Journal of the Linnean Society 204 (4): 1-16, DOI: 10.1093/zoolinnean/zlaf101, URL: https://doi.org/10.1093/zoolinnean/zlaf101
563E8793D24DFFDCFCCDFC43FE86C995.taxon	distribution	Distribution and habitat: The species is found in the branchial cavity of Macrocheira kaempferi (Fig. 6 A, B). Since its first discovery byMarenzeller (1902) in Japanese waters (specific locality not documented), this species has been reported at depths of 548 – 640 m in Sagami Bay, under the junior synonym of Coelobranchus papillosus Izuka, 1912 (Izuka 1912, Imajima and Hartman 1964). It has also been reported from Seto and Minabe, Wakayama, Japan (Japan Initiative for Biodiversity Information; https: // gbif. jp / gbif _ search / dataset? datasetId = snet- 1096 - 105).	en	Hookabe, Natsumi, Yamamori, Luna, Ogawa, Akito, Jimi, Naoto, Kohtsuka, Hisanori, Rouse, Greg W. (2025): An updated phylogeny of Iphitime (Annelida: Dorvilleidae) revealing multiple host switching, with description of I. nubila sp. nov. Zoological Journal of the Linnean Society 204 (4): 1-16, DOI: 10.1093/zoolinnean/zlaf101, URL: https://doi.org/10.1093/zoolinnean/zlaf101
563E8793D24DFFDCFCCDFC43FE86C995.taxon	discussion	Remarks: Specimens collected from Sagami Bay conform to the original description of I. doderleini as well as C. papillosus (Marenzeller 1902, Izuka 1912).	en	Hookabe, Natsumi, Yamamori, Luna, Ogawa, Akito, Jimi, Naoto, Kohtsuka, Hisanori, Rouse, Greg W. (2025): An updated phylogeny of Iphitime (Annelida: Dorvilleidae) revealing multiple host switching, with description of I. nubila sp. nov. Zoological Journal of the Linnean Society 204 (4): 1-16, DOI: 10.1093/zoolinnean/zlaf101, URL: https://doi.org/10.1093/zoolinnean/zlaf101
563E8793D24EFFDCFEC0F948FB21CBDC.taxon	discussion	We obtained the mitogenome of I. doederleini (15980 bp in total, 26.4 % GC content) and I. nubila (15081 bp in total, 26.9 % of GC content) (Fig. 9). In I. doederleini from Sagami Bay, 18 S was 3128 bp long, with 14411 reads and an average coverage of 659.02 ×; in I. doederleini from Wakayama, 18 S was 3233 bp long, with 23803 reads and an average coverage of 1018.31 × and the mitochondrial genome was 15980 bp long, with 12734 reads and an average coverage of 115.27 ×; in I. nubila, 18 S was 2407 bp long, with 886 reads and an average coverage of 69.00 × and the mitochondrial genome was 15081 bp long, with 13587 reads and an average coverage of 133.94 ×. For both species, 13 protein-coding genes (PCGs) including cox 1, cox 2, cox 3, nad 1, nad 2, nad 3, nad 4, nad 4 l, nad 5, nad 6, cytb, atp 6, and atp 8, 12 S rRNA, 16 S rRNA, and 22 tRNAs were identified (Fig. 9). The gene order of each species was identical: cox 1 - cox 2 - cox 3 - nad 6 - nad 5 - nad 4 l-nad 4 - rrnS-rrnR-cytb-nad 1 - nad 3 - nad 2 (Fig. 9).	en	Hookabe, Natsumi, Yamamori, Luna, Ogawa, Akito, Jimi, Naoto, Kohtsuka, Hisanori, Rouse, Greg W. (2025): An updated phylogeny of Iphitime (Annelida: Dorvilleidae) revealing multiple host switching, with description of I. nubila sp. nov. Zoological Journal of the Linnean Society 204 (4): 1-16, DOI: 10.1093/zoolinnean/zlaf101, URL: https://doi.org/10.1093/zoolinnean/zlaf101
