Iphione muricata, sensu Savigny, 1822

Salazar-Vallejo, Sergio I., Piotrowski, Christina N. & Paulay, Gustav, 2024, Revision of Iphione Kinberg, 1856 (Annelida, Aphroditiformia, Iphionidae), Zootaxa 5548 (1), pp. 1-84 : 16-25

Iphione muricata (Savigny in Lamarck, 1818) restricted

Figs 1D–F , 3D–F , 4–6 , 36

Polynoe muricata Savigny in Lamarck, 1818: 308–309 (diagn.). Iphione glabra de Quatrefages, 1866: 268–269 ; Solís-Weiss et al. 2004: S14. Iphione muricata : Potts 1910: 341; Fauvel 1919: 334; Fauvel 1932: 12–13 (partim); Fauvel 1935: 285–286 (partim); Fauvel

1939: 509 (partim); Monro 1939: 168 (partim); Day 1951: 14 (partim); Gibbs 1971: 123 (partim); Amoureux 1974: 430

(partim); Pettibone 1986: 9–16, Figs 1–5 (syn. partim); Hartmann-Schröder 1991: 19 (partim).

Type material. MNHN TYPE 335: Mauritius (Île de France) M. Mathieu, coll. no further data; holotype of Polynoe muricata and syntype of Iphione glabra (see Remarks) (bent ventrally, most elytra previously removed; body and elytra with a massive accumulation of precipitates, probably salts; two elytra cleaned, one median and one posterior elytra removed; body 24 mm long, 16 mm wide, 29 chaetigers). MNHN TYPE 334: Mauritius (Île de France), Péron & Lesueur, coll. no further data; syntype of Iphione glabra only; better preserved, used for redescription)

Additional material

Egypt, Red Sea, Gulf of Suez. Four specimens ( MNHN A274-19 ), Mission Jousseaume, 1894 (no further data; three specimens bent ventrally; elytra brownish, oblique longitudinal band barely defined, some with calcareous epibionts; macrotubercles in two rows, progressively smaller posteriorly; body 19–28 mm long, 10–15 mm wide, 29 chaetigers). Ten specimens ( MNHN A274-27 ), Mission Jousseaume, 1894 (no further data; four specimens bent ventrally; elytra brownish, oblique longitudinal band barely defined, most with calcareous epibionts; macrotubercles in two rows, progressively smaller posteriorly; body 14–23 mm long, 7.5–14.0 mm wide, 29 chaetigers) .

Sultanate of Oman. Two specimens ( UF 411 ), Muscat Governorate, Qurm Beach (23°37’33.6” N, 58°28’51.6” E), 0–1 m, sandstone, under rocks, mid-low water intertidal, 26 Jan. 2005, V. Bonito, M. Claereboudt & G. Paulay, coll. (markedly bent ventrally; pharynx exposed; elytra golden to brownish, oblique longitudinal blackish line better defined in largest specimen; macrotubercles in two rows, progressively smaller posteriorly; cirrigerous segments with basal tubercle of dorsal cirrophores projected, barely in smallest specimen; body length not measured, 0.8–1.6 mm wide, 28–29 chaetigers) . One specimen ( UF 9313 ), Muscat Governorate, Haramel Village cove (23°35’42.0” N, 058°36’03.6” E), 14 Jan. 2020, G. Paulay, S. Maslakova & M. Cherneva, coll. (markedly bent ventrally; body wall reddish; elytra brownish with an oblique, slightly darker longitudinal band; macrotubercles in three rows, progressively smaller posteriorly; basal tubercle of dorsal cirrophores projected; body not measured) . One specimen ( UF 9318 ), Muscat Governorate, Haramel Village cove (23°35’42.0” N, 58°36’03.6” E), 14 Jan. 2020, G. Paulay, S. Maslakova & M. Cherneva, coll. (complete, bent ventrally; body wall reddish; elytra brownish with an oblique, slightly darker longitudinal band; macrotubercles in two rows, progressively smaller posteriorly; basal tubercle of dorsal cirrophores projected; body 27 mm long, 16 mm wide, 29 chaetigers) . One specimen ( UF 9319 ), Muscat Governorate, Haramel Village cove (23°35’42.0” N, 58°36’03.6” E), 14 Jan. 2020, G. Paulay, S. Maslakova & M. Cherneva, coll. (complete, slightly bent ventrally; body wall reddish; elytra brownish with an oblique, slightly darker longitudinal band; macrotubercles in three rows, progressively smaller posteriorly; basal tubercle of dorsal cirrophores projected; body 27 mm long, 17 mm wide, 29 chaetigers) . One specimen ( UF 9320 ), Muscat Governorate, Haramel Village cove (23°35’42.0” N, 58°36’03.6” E), 14 Jan. 2020, G. Paulay, S. Maslakova & M. Cherneva, coll. (complete, slightly bent ventrally; body wall reddish; elytra brownish with an oblique, slightly darker longitudinal band; macrotubercles in three rows, progressively smaller posteriorly; basal tubercle of dorsal cirrophores projected; body 25 mm long, 15 mm wide, 29 chaetigers) . One specimen ( UF 9368 ), Muscat Governorate, Bandar Al Jissah (23°33’25.2” N, 58°39’03.6” E), 16 Jan. 2020, G. Paulay, M. Claereboudt, S. Maslakova & G. Kolbasova, coll. (complete, slightly bent ventrally; body wall reddish; elytra golden, without black spots; macrotubercles in two rows, progressively smaller posteriorly; basal tubercle of dorsal cirrophores projected; body 19 mm long, 13 mm wide, 29 chaetigers) . One specimen ( UF 9454 ), Muscat Governorate, West side of Fahal Island (23°40’55.2” N, 58°30’00.0” E), 19 Jan. 2020, G. Paulay, A. Bemis, G. Kolbasova & S. Brown, coll. (juvenile; right parapodia of chaetigers 13 and 14 previously removed; elytra yellowish, without black spots, macrotubercles in two rows, progressively smaller posteriorly; cirrigerous segments with basal tubercle not projected; other data in variation: body 11 mm long, 6 mm wide, 28 chaetigers) . One specimen ( UF 10504 ), Dhofar Governate, Mirbat, Chinese Wreck (16°57’57.6” N, 54°42’28.8” E), 26 Ja. 2022, G. Paulay, R. Lasley, A. Uehling & E. Ciareboudt, coll. (bent ventrally; right parapodia of chaetigers 11–16 previously removed; right elytra 5 and 6 removed for observation (kept in container); elytra brownish, with an oblique longitudinal wide band, and white epibionts; macrotubercles in 2—three rows, progressively smaller posteriorly; cirrigerous segments with basal tubercle projected; body mm 26 long, 15 mm wide, 29 chaetigers) .

Djibouti

Two specimens ( MNHN A274-7 ), Djibouti , 1897, No. 29, M. Coutière, coll. (largest specimens with all left elytra previously removed; elytra brownish without black spots; macrotubercles in two rows, progressively smaller posteriorly; body 19–27 mm long, 11–13 mm wide, 29 chaetigers) . One specimen ( MNHN A274-12 ), Mission Gravier Djibouti , Île Moucha, Gulf of Tadjourah, coral, 23 Jan. 1904, C. Gravier, coll. (bent ventrally; elytra brownish; macrotubercles in two rows, decreasing in size posteriorly; body 17.5 mm long, 9.5 mm wide, 29 chaetigers) .

Seven specimens ( MNHN A274-23 ), Obock, Obock, Mission Jousseume , 1895, M. Jousseume, coll. (body damaged; two with most elytra detached, some in container; elytra brownish without black spots; macrotubercles in two rows progressively smaller posteriorly; body 14–27 mm long, 9–15 mm wide, 29 chaetigers) . One specimen ( MNHN A274-32 ), Mission Gravier Djibouti, Îlot Residence , in mixed bottoms (sand and rock), 1904, C. Gravier, coll. (markedly bent ventrally; elytra brownish without black spots; macrotubercles in two rows progressively smaller posteriorly; not measured) .

Kenya. Seven specimens ( NHM 1924.6.18.187–191), F.A. Potts, coll. (no further data; elytra yellowish to brownish, most with a barely pigmented longitudinal blackish band, macrotubercles in two rows, first row with macrotubercles slightly larger than those in second row; body 10–21 mm long, 9.0– 9.5 mm wide, 29 chaetigers) .

Mozambique. One specimen ( NHM 1963.1.6), Delagoa Bay , Inhaca Island (25°50’ S, 32°50’ E), J.H. Day, coll. (elytra brownish, blackish band barely visible, macrotubercles slightly smaller posteriorly, in 2 irregular rows; body 23 mm long, 14 mm wide, 29 chaetigers) .

Tanzania. One specimen ( CAS 227559 ), complete, Zanzibar Expedition 2018, Zanzibar, Mangapwani (05°59’24.0” S, 39°11’24.0” S), 0.2–3.0. m, T.M. Gosliner, coll. (data used for variation) . Three specimens ( NHM 1941.4.4.165–167), Pemba Island, sandbanks, 18 Nov. 1901, C. Crossland, coll. (one completely bent, another one almost without elytra; elytra brownish with a longitudinal blackish band barely visible, macrotubercles in two rows of similar size; body 17–25 mm long, 11–14 mm wide, 29 chaetigers) . One specimen ( ZMH PE 187 b), Bawe , Zanzibar, 15 Jul. 1989, Votzkow, coll. (no further data; slightly dehydrated; first few pairs of anterior and posterior elytra, and left parapodium of chaetiger 16 previously removed (parapodium lost); elytra brownish with two rows of macrotubercles progressively smaller posteriorly; cirrigerous segments with basal tubercle of dorsal cirrophores projected; body 28.5 mm long, 11 mm wide, 29 chaetigers) . One specimen ( ZMH V4575 ), Bani Island , Zanzibar, Stuhlmann, coll. (no futher data; markedly bent ventrally, partially dehydrated; elytra brownish, without black bands, with two rows of macrotubercles decreasing in size posteriorly; not measured) . One specimen ( ZMH V4577 ), Kokotoni , Zanzibar, Stuhlmann, coll. (no further data; bent ventrally; elytra brownish with high amount of sediment particles mostly on fimbriae; darker middorsal bands indistinct; elytra with two rows of macrotubercles heavily eroded, progressively smaller posteriorly; body 20.5 mm long, 14 mm wide, 29 chaetigers) .

Mayotte. One specimen ( MNHN A274-9 ), 1883, A. Vinnout, coll. (no further data; markedly bent ventrally; left elytron 10 removed for observation (kept in container); elytra with an oblique, longitudinal black band, expanded anteriorly; macrotubercles in two rows, progressively smaller posteriorly; not measured) .

Madagascar. One specimen ( MNHN A274-10 ), Baie St. Augustin, No. 23, 1909, M. Bastard, coll. (no further data; some elytra previously removed; elytra brownish without black spots; macrotubercles in two rows, progressively smaller posteriorly; cirrigerous segments with basal tubercle of dorsal cirrophores barely projected; body 20 mm long, 12 mm wide, 29 chaetigers) . Four specimens ( MNHN A274-29 ), Mission F. Geay, Tuléar, reefs, No. 4978 (no further data; partially dehydrated, one markedly bent ventrally with pharynx fully exposed, jaws with one additional denticle; elytra brownish without black spots, outer elytral surface darker than inner one; macrotubercles in two rows, progressively smaller posteriorly; body 14.0– 19.5 mm long, 8.0– 10.5 mm wide, 29 chaetigers) . Five specimens ( MNHN A274-35 ), 1919, M. Decary, coll. (no further data; very soft specimens, elytra detaching or in site; elytra with an oblique longitudinal blackish band in two specimens, the others with elytra brownish; macrotubercles in two rows, progressively smaller posteriorly; not measured) . One specimen ( NHM 1961.8.3), Diego Suarez & Nossy-Bé (no further data; elytra covered with fine particles, with a wide oblique blackish band, two rows of macrotubercles, first one with tubercles slightly larger than those in second row, probably after the sedimentary environment; body 18 mm long, 11 mm wide, 28 chaetigers) .

Mascarene Islands. One specimen ( ZMH V499 ), Mauritius. Robillard, coll. (no further data; markedly bent ventrally; no other dissections to avoid further damage; elytra brownish with two rows of macrotubercles progressively smaller posteriorly; not measured) . One specimen ( UF 635 ), Réunion Island, La Saline , Trou d’eau (21°06’03.6” S, 55°14’38.4”E 0–2 m, 27 Jul. 2007, N. Hubert & F. Michonneau, coll. (markedly bent ventrally, pharynx exposed; left parapodia of chaetigers 15–20 previously removed; elytra brownish, oblique longitudinal black band barely defined, with three rows of macrotubercles progressively smaller towards margins; pharynx exposed, jaws with 2 mostly eroded accessory denticles; body length not measured, 13 mm wide, 29 chaetigers) . One specimen ( UF 636 ), Réunion Island, Saint-Leu , Sec Jaune (21°09’19.8» S, 55°16’52.02» E), 0–2 m, 17 Aug. 2007, H. Bruggemann, N. Hubert & F. Michonneau, coll. (bent laterally after removal of left parapodia of chaetigers 12–17; elytra with wide blackish band, with three rows of macrotubercles progressively smaller towards margins; pharynx exposed, jaws with 3 accesory denticles; body 17 mm long, 8 mm wide, 29 chaetigers) . One specimen ( UF 638 ), Réunion Island, La Saline , Trou d’eau (21°06’03.6” S, 55°14’38.4” E), 0–2 m, 30 Jul. 2007, N. Hubert & F. Michonneau, coll. (bent laterally after removal of right parapodia of chaetigers 15–19; elytra with three rows of macrotubercles progressively smaller towards margins; pharynx exposed, upper jaws with 3 accessory denticles, lower ones with 4; body 25 mm long, 13 mm wide, 29 chaetigers) . One specimen ( UF 662 ), Réunion Island, La Posses , Banc des Lataniers (20°55’21.0» S, 55°20’29.04» E), boulder and cobble beach, under rocks, 0–2 m, 7 Aug. 2007, H. Bruggemann, N. Hubert, F. Michonneau & G. Paulay, coll. (slightly bent ventraly; elytra with three rows of macrotubercles progressively smaller towards margins; body 31 mm long, 16 mm wide, 29 chaetigers) .

Seychelles. One specimen ( ZMH 5758 ), Brauer, coll. (no further data; bent ventrally, pharynx fully exposed; elytra brownish with an oblique, longitudinal darker band barely distinct; elytra with two rows of macrotubercles progressively smaller posteriorly; body 20 mm long, 12 mm wide, 29 chaetigers) .

Nicobar Islands. One specimen ( NHMD 1184777 ), RV Galathea Expedition, Unspecified locality and date (fixed in ethanol, bent ventrally; elytra with a barely pigmented dark band; 3–4 rows of macrotubercles, progressively smaller towards margins; basal tubercle of dorsal cirrophores bilobed; neurochaetae unidentate; body 27 mm long, 13 mm wide, 29 chaetigers) .

Vietnam. Two specimens ( MNHN A274-1 ), Tonkin, Mar. 1897, M. Lichtenfelder, coll. (no further data; one bent ventrally; some elytra previously removed; elytra brownish with an oblique longitudinal blackish band; macrotubercles in two rows, decreasing in size posteriorly; basal tubercle to dorsal cirrophore bilobed; body 19–22 mm long, 11.5–12 mm wide, 29 chaetigers) . One specimen ( MNHN A274-2 ), Cap St. Jacques, Capt. Modest, Sep. 1908 (no further data; bent ventrally, some elytra previously removed; elytra brownish with an oblique, longitudinal black band; macrotubercles in two rows, decreasing in size posteriorly; cirrigerous segments with basal tubercle of dorsal cirrophore projected; neurochaetae slightly falcate, unidentate; body 22 mm long, 10 mm wide, 29 chaetigers) . Two specimens ( MNHN A274-5 ), Cap St. Jacques, Capt. Modest, No. 37, Sep. 1908 (poorly preserved; elytra brownish with an oblique, longitudinal black band, better defined in a few elytra; macrotubercles in two rows, decreasing in size posteriorly; most fimbriae detached; body 20–28 mm long, 12.0– 13.5 mm wide, 29 chaetigers) .

Indonesia. One specimen ( ZMA V471.7 ), Sulawesi, RV Siboga Exped., Stat. 127 (Great Sangir Island, Taruna Bay ), 45 m, trawl and reef-exploration, fine sand, 20–21 Jul. 1899 (markedly bent; elytra yellowish, without black spots, with fine sediment particles; macrotubercles in two rows, progressively smaller posteriorly; not measured) . One specimen ( ZMA V471.9 ), Maluku, RV Siboga Exped., Sta. 142 (Obi Major, anchorage off Laiwui), 23 m, dredge + reef exploration, muddy bottom, 5–7 Aug. 1899 (markedly bent ventrally; elytra yellowish, oblique longitudinal darker band barely visible; macrotubercles in two rows, progressively smaller posteriorly; not measured) . One specimen ( ZMA V471.12 ), Irian Jaya, RV Siboga Exped., Stat. 164 (01°42.5´S, 130°47.5´E), 32 m, dredge, sand, stones, shells, 20 Ago. 1899 (bent ventrally; right elytra 6 and 7 removed for observation (kept in container); elytra yellowish, without black spots, with some areas in regeneration, with different epibionts, without fine sediment particles; macrotubercles in two rows, progressively smaller posteriorly; body 17 mm long, 11 mm wide, 29 chaetigers) . One specimen ( ZMA V471.13 ), Maluku, RV Siboga Exped., Sta. 174 (Ceram (Seram), N coast, Waru Bay ), 18 m, dredge, reef exploration, muddy bottom, 28–29 Aug. 1899 (elytra yellowish, without black spots; macrotubercles in two rows, progressively smaller posteriorly; body 9.5 mm long, 6 mm wide, 29 chaetigers) . One specimen ( ZMA V471.24 ), Lesser Sunda Islands, Rotti (or Rodti) Island, E coast, Pepela Bay , RV Siboga Exped., Sta. 301 (10°38´S, 123°25.2´E), 22 m, dredge, muddy bottom, coral, 30 Jan. 1900 (markedly bent; elytra brownish with white epibionts towards posterior margins, without black spots; macrotubercles in two rows, progressively smaller posteriorly; not measured) . One specimen ( ZMA V471.30 ), Maluku, RV Siboga Exped., Sta. 250 ( Kur Island , anchorage off Kilsuin), 20–45 m, coral, 6–7 Dec. 1899 (bent ventrally; elytra yellowish, without black spots, macrotubercles in two rows, progressively smaller posteriorly; body 8 mm long, 6 mm wide, 28 chaetigers) . One specimen ( ZMA V476 ), Sumatra, Nias Island , southern coast, Telok Dalam (Teluk Dalam), de Zwaan, coll. (no further data; complete, right elytra 6 and 7 previously removed; elytra with oblique longitudinal black band, and darker posterior margins, with white and filamentous epibionts; macrotubercles in two rows, progressively smaller posteriorly; body 23 mm long, 13 mm wide, 29 chaetigers) .

Philippines. Two specimens ( ZMH V500 ), Mactan Island , Cebu, Ringe, coll. (no further data; bent ventrally, one without elytra; the other with elytra brownish; elytra with oblique longitudinal blackish band, and darker posterior elytral area, with two rows of macrotubercles progressively smaller posteriorly; cirrigerous segments with basal tubercle of dorsal cirrophores projected; body 19 mm long, 11 mm wide, 29 chaetigers) .

Hong Kong. One specimen ( AM 20043 ), Chek Chau, W side of Hong Kong, 5 m, 14 Apr. 1986, P.A. Hutchings, coll. (bent ventrally; elytra brownish with an oblique longitudinal black band; macrotubercles in two rows, progressively smaller posteriorly; cirrigerous segments with basal tubercle of dorsal cirrophore projected; body 22 mm long, 10 mm wide, 29 chaetigers) .

Australia. One specimen ( AM 2650 ), Lindeman Island, Whitsunday Pass , Queensland, 16 m, Jan. 1929, M. Ward, coll. (elytra brownish, oblique longitudinal black band better defined in posterior chaetigers; macrotubercles in two rows, progressively smaller posteriorly; basal tubercle of dorsal cirrophore projected); body 22.5 mm long, 14.5 mm wide, 29 chaetigers ). One specimen ( AM 2737 ), Northwest Island, Capricorn Group , Queensland, 1929, Ward & Boardman, coll. (no further data; elytra dark brown; elytra with a few sediment particles and epibionts; macrotubercles in two rows, decreasing in size posteriorly; basal tubercle of dorsal cirrophore projected; body 21 mm long, 12.5 mm wide, 29 chaetigers) . Three specimens ( AM 2750 ), Rat Island, Port Curtis , Queensland, Jul. 1929, Ward & Boardman, coll. (no further data; elytra brownish with an oblique longitudinal black band; macrotubercles in 2–three rows, progressively smaller posteriorly; cirrigerous segments with basal tubercle of dorsal cirrophore projected; body 28–29 mm long, 14.0– 14.5 mm wide, 29 chaetigers) . One specimen ( AM 2845 ), Ninety Mile Beach , between Broome and Wallal, 13 m, 1931, R. Bourne, coll. (markedly bent ventrally; elytra yellowish with an oblique longitudinal black band; macrotubercles in two rows, progressively smaller posteriorly; basal tubercle of dorsal cirrophore projected; not measured) . One specimen ( AM 2934 ), Great Barrier Expedition, Queensland, N of North Direction Island, Great Barrier Reef , Sta. 17 (14°43’48.0” S, 145°30’00.0” E), 35 m, 9 Mar. 1929 (bent ventrally; elytra yellowish with an oblique longitudinal ill-defined band; macrotubercles in two rows, progressively smaller posteriorly; basal tubercle of dorsal cirrophore projected; body 20 mm long, 10.5 mm wide, 29 chaetigers) . One specimen ( AM 5432 ), Mandorah , Darwin, NT, under rock, low tide, 22 Nov. 1972, N. Coleman, coll. (slightly bent ventrally; elytra with an oblique black longitudinal band and several black spots; macrotubercles in two rows, progressively smaller posteriorly; body 15.5 mm long, 10 mm wide, 29 chaetigers) . One specimen ( AM 5437 ), Kendrew Island , Dampier, WA, under rocks, low tide, 14 Oct. 1972, N. Coleman, coll. (slightly bent ventrally, some elytra previously removed (kept in container); elytra yellowish with an oblique longitudinal black band; macrotubercles in two rows, decreasing in size posteriorly; body 22 mm long, 15 mm wide, 29 chaetigers) . One specimen ( AM 5452 ), Blow Holes , Quobba, WA, 2 m, 22 Jun 1972, N. Coleman, coll. (markedly bent ventrally; elytra brownish with an oblique longitudinal black band; macrotubercles in two rows, progressively smaller posteriorly; not measured) . One specimen ( AM 38418 ), Dampier Archipelago, Georgeff Reef , intertidal, 28 Aug. 1999, R.A. Peart, coll. (bent ventrally; some elytra previously removed; elytra yellowish with an oblique longitudinal black band and some calcareous epibionts; macrotubercles in two rows, progressively smaller posteriorly; body 16 mm long, 11 mm wide, 29 chaetigers) . One specimen ( AM 38419 ), Dampier Archipelago, W side Malus Island , 2.6 m, sand and algae bottom, 27 Aug. 1999, R.A. Peart, coll. (juvenile, posterior end bent ventrally; all elytra previously removed; elytra yellowish without black spots; macrotubercles in two rows, second row with a few macrotubercles, progressively smaller posteriorly; body 14 mm long, 6 mm wide, 28 chaetigers) . One specimen ( MAGNT W252 ), Oxley Island, NT, Sta. OX 2, Acropora , low subtidal, 26 Oct. 1982, J.R. Hanley, coll. (markedly bent ventrally; elytra with abundant sediment particles, and a tapered, oblique longitudinal dark band; macrotubercles in two rows, anterior row with macrotubercles slightly larger than those of second, more irregular row; not measured) . One specimen ( MAGNT W4894 ), Lee Point, Darwin, NT, Sta. RH 87-48, coral rubble, subtidal, 23 Oct. 1987, J.R. Hanley, coll. (juvenile, slightly bent ventrally, posterior end damaged; elytra yellowish with spread black spots, 2–3 submarginal fimbriae, macrotubercles in two rows, decreasing in size posteriorly; cirrigerous segments with basal tubercle of dorsal cirrophore projected; neurochaetae unidentate; body 6 mm long, 4 mm wide, 28 chaetigers) . One specimen ( MAGNT W4897 ), Lee Point, Darwin, NT, Sta. RH 87-48, coral rubble, subtidal, 23 Oct. 1987, J.R. Hanley, coll. (bent ventrally; elytra yellowish with oblique longitudinal black band, some fimbriae between macrotubercles, macrotubercles in two rows, decreasing in size posteriorly; body 14 mm long, 9.5 mm wide, 29 chaetigers) . One specimen ( MAGNT W4939 ), Lee Point, Darwin, NT, Sta. RH 87-48, exposed rocky reef, subtidal, 25 Sep. 1987, J.R. Hanley, coll. (bent ventrally, some elytra previously detached; elytra yellowish with oblique longitudinal wide black band, macrotubercles in two rows, decreasing in size posteriorly; body 11 mm long, 7.5 mm wide, 28 chaetigers) .

One specimen ( MAGNT W5139 ), Point Samson , near Roebourne, WA, Sta. RH 88-35, coral rubble, low tide, 9 Feb. 1988, J.R. Hanley, coll. (bent ventrally; many elytra and left parapodium of chaetiger 12 previously removed (all kept in container); elytra covered with orange silt; prostomium paler than adjacent body wall regions; elytra yellowish with an irregular median black spot; macrotubercles in two rows, gradually smaller posteriorly; cirrigerous segments with basal tubercle of dorsal cirrophore projected; body 10 mm long, 7 mm wide, 28 chaetigers) . One specimen ( MAGNT W5140 ), Point Samson , near Roebourne, WA, Sta. RH 88-35, coral rubble, low tide, 9 Feb. 1988, J.R. Hanley, coll. (some elytra previously removed (kept in container), left parapodium of chaetiger 10 previously removed (lost); elytra covered with orange silt; prostomium white, contrasting with body wall pigmentation; pharynx partially exposed, slightly damaged by compression, partially dissected for observing jaws and its 2 accessory denticles, elytra yellowish with an oblique longitudinal black band; macrotubercles in two rows, gradually smaller posteriorly; body 15 mm long, 10 mm wide, 29 chaetigers) . One specimen ( MAGNT W5141 ), Point Samson , near Roebourne, WA, Sta. RH 88-35, coral rubble, low tide, 9 Feb. 1988, J.R. Hanley, coll. (markedly bent ventrally; many elytra, and right parapodium of chaetiger 10 previously removed (all kept in container); elytra covered with orange silt; prostomium white; elytra yellowish, median black spots barely visible; macrotubercles in two rows, gradually smaller posteriorly; cirrigerous segments with basal tubercle of dorsal cirrophore projected; not measured) . One specimen ( MAGNT W6882 ), Scorpion Island , WA, Sta. RH 91-54, fringing shore reef, low tide, 14 Aug. 1991, J.R. Hanley, coll. (bent ventrally; elytra yellowish with a wide irregular dark band; macrotubercles in two rows, decreasing in size posteriorly; body 14 mm long, 7 mm wide, 29 chaetigers) . One specimen ( MAGNT W5142 ), Point Samson , near Roebourne, WA, Sta. RH 88-35, coral rubble, low tide, 9 Feb. 1988, J.R. Hanley, coll. (markedly bent ventrally; many elytra previously removed, left parapodium of chaetiger 13 previously removed (all kept in container); elytra covered with orange silt; prostomium white, contrasting with body wall pigmentation; pharynx partially exposed, slightly damaged by compression; elytra yellowish with an irregular medial spot; macrotubercles in two rows, gradually smaller posteriorly; not measured) . One specimen ( MAGNT W28949 ), East Point , NT, Sta. EP 11, rocks imbedded in mud, low tide, 18 May 1985, J.R. Hanley, coll. (bent ventrally; pharynx fully exposed, jaws with 2 accessory denticles; right elytra 5 and 7 removed for observatioin; elytra with a wide, ill-defined black band; macrotubercles in two rows decreasing in size posteriorly; body 11.5 mm long, 7.5 mm wide, 28 chaetigers) . Eleven specimens ( NHM 1970.113 a), Grahams Point (no further data; elytra yellowish in smaller specimens, progressively darker in larger ones; many elytra eroded, with white epibionts; two rows of macrotubercles, first one with macrotubercles slightly larger than those present in second row; body 10–28 mm long, 6.0– 13.5 mm wide, 29 chaetigers) . Two specimens ( ZMH P 20976 ), Heron Island , coral reef, under rocks, intertidal, 3 Feb. 1976, G. Hartmann-Schröder, coll. (smallest specimen broken by compression, largest one bent ventrally, with elytra yellowish, without black spots, two rows of macrotubercles progressively smaller posteriorly; body 25 mm long, 14 mm wide, 29 chaetigers) .

One specimen ( ZMH V7883 ), Turtle Island , Western Australia, Gale, coll. (no further data; elytra brownish with an oblique longitudinal black band; two rows of macrotubercles, progressively smaller posteriorly; basal tubercle of dorsal cirrophore projected as a single lobe; body 25 mm long, 14 mm wide, 29 chaetigers) . One specimen ( ZMH V9333 ), Cape Jaubert , Western Australia (no further data; body brownish, slightly bent ventrally; elytra with two rows of macrotubercles, progressively smaller posteriorly; basal tubercle of dorsal cirrophore with a projected lobe; body 21 mm long, 12 mm wide, 29 chaetigers) .

Papua New Guinea. One specimen ( MNHN IA 2022-929 ), Papua Niugini Expedition, Sta. PR 149 (5°09.4’ S, 145°48’ E), 19 m, 1 Dec. 2012, S. Hourdez et al., coll. (markedly bent ventrally; elytra brownish, with oblique longitudinal black band, macrotubercles in two rows, progressively smaller posteriorly; cirrigerous segments with basal tubercle of dorsal cirrophore projected; body 9 mm wide) .

Solomon Islands. Two specimens ( NHM 1970.11 ), Komimbo Bay , Guadalcanal (no further data; smaller specimen with tiny black dots on elytra, larger specimen with brownish elytra, each with 2—three rows of macrotubercles; basal tubercle of dorsal cirrophores projected; neurochaetae unidentate, slightly falcate; body 7–16 mm long, 4.5–10 mm wide, 28–29 chaetigers) . One specimen ( NMH 1970.107 ), Tetel Island , 8 Jul. 1975, P.E. Gibbs, coll. (elytra yellowish with a darker band over an area larger than external elytral half, macrotubercles in two rows; left elytra 1–3 previously removed (not in container); body 13.5 mm long, 7 mm wide, chaetigers) .

New Caledonia. One specimen ( MNHN A274-11 ), Grand Terre , Kuto Bay, Île des Pins (no further data; bent ventrally, partially dehydrated; pharynx fully exposed, jaws with 3 accessory denticles, elytra brownish with a small black spot; macrotubercles arranged in two rows, progressively smaller posteriorly; body 22 mm long, 9.5 mm wide, 29 chaetigers) .

Samoa. One specimen ( MNHN A274-22 ), Moris, Sta. 56, 1889 (no further data; bent ventrally; right elytron 6 removed for observation (kept in container); elytra brownish, without black spots; macrotubercles in two rows, progressively smaller posteriorly; body 20 mm long, 13 mm wide, 29 chaetigers) . One specimen ( NHM 1921.5.1.533), Upolu, W.C. McIntosh, coll. (no further data; bent ventrally; elytra brownish, darker along posterior margins; macrotubercles in two rows, first row with macrotubercles slightly larger; two elytra previously removed (kept in container); body 25 mm long, 15 mm wide, chaetigers) .

Diagnosis. Iphione with median antenna reduced to nuchal papilla; elytra fimbriate; macrotubercles spine-like, barely longer than wide, with distal spines, in 2—three rows, first row with macrotubercles slightly larger than those in other rows; cirrigerous segments with dorsal cirrophores with basal tubercle projected; neurochaetae unidentate.

Description. Syntype of Iphione glabra de Quatrefages, 1866 (MNHN TYPE 334) bent ventrally, first eight pairs or elytra previously removed ( Fig. 4A ), some left in container; right parapodia of chaetigers 15 and 16 previously removed (lost), left parapodium of chaetiger 14 removed for observation (kept in container); body 17 mm long, 9 mm wide, 29 chaetigers). Elytra brownish, with sediment particles and an oblique dark band visible in most elytra; notochaetae whitish, neurochaetae golden. Body wall pale, venter pale.

Elytra with fimbriae, eroded from most detached elytra ( Fig. 4B, C ), better preserved in the other syntype (MNHN TYPE 335) ( Fig. 4F–H ), with variable amounts of sediment particles, especially along posterior segments. Fimbriae 6–10 times longer than wide, with a few spines along stem in syntype 334 ( Fig. 4B , inset), or markedly spinous, probably after contraction in syntype 335 ( Fig. 11I ); macrotubercles conical, most eroded, about twice longer than wide, arranged in two rows, decreasing in size posteriorly and laterally.

Prostomium roughly hexagonal, wider than long, with anterior incision running along anterior prostomial third. Anterior lobes projected into ceratophores, longer than prostomium, shorter than ceratostyles, subdistally swollen, tips tapered. Eyes colorless, indistinct. Palps thick, bent, left one bent to the right, right one bent dorsally, slightly longer than antennae, with longitudinal rows of papillae, tips mucronate. Nuchal papillae not visible dorsally, covered by nuchal flap.

Tentacular segment dorsally reduced, with chaetae; tentacular cirri slightly shorter than palps. Facial tubercle not visible dorsally, pale.

Segments 2–4 directed anteriorly. Second segment visible dorsally, with a semicircular nuchal lappet, wider than long, covering nuchal papilla. Ventral buccal cirri inserted ventrally, almost three times longer than following cirri, left lost, right one not projected beyond chaetal tips. Dorsal nodules barely projected, more elevated in segment 2, progressively shorter.

Median cirrigerous segments with dorsal cirri markedly projected beyond chaetal tips, cirrophore smooth, with a projected, single barely projected lobe ( Fig. 4D ). Cirrostyle subdistally swollen, surface papillate. Notochaetae very abundant, dirty white, with series of transverse funnel-shaped spinose rows, tips bare. Neuropodia with neuracicular lobe blunt, with globular papillae. Neurochaetal lobe with marginal papillae, neurochaetae abundant, basally smooth, subdistally swollen with many rows of fine denticulations along swollen region, tips slightly falcate, unidentate, sharp ( Fig. 11E ).

The other syntype (MNHN 335) brownish, with large amounts of adsorbed materials on body and elytra ( Fig. 4F ), and a few elytra attached; body 24 mm long, 16 mm wide, 29 chaetigers. After cleaning, both median ( Fig. 4G ) and posterior elytra ( Fig. 4H ) with two irregular rows of macrotubercles, mostly eroded to their base; oblique darker longitudinal band barely visible; fimbria subcylindrical, with abundant large spines along stem, and with long blunt spines distally ( Fig. 4I ). Left parapodium of chaetiger 12 removed for observation (kept in container); cirrigerous parapodia with a barely developed basal tubercle to dorsal cirrophore, cirrostyles slightly swollen subdistally ( Fig. 4J ).

Variation. Some features of the exposed pharynx ( Fig. 3D–F ) were referred to above. One specimen (CAS 227559) complete, brownish, slightly bent dorsally, pharynx fully exposed, jaws with three accessory denticles; body 27 mm long, 13 mm wide, 29 chaetigers ( Fig. 5A ); left elytron 7 removed for observation, and left parapodium of chaetiger 12 removed for observation (kept in container); elytra brownish, larger macrotubercles arranged in two rows, macrotubercles about 2–3 times longer than wide ( Fig. 5B ); fimbriae arranged along lateral and posterior margins, longer in lateral ( Fig. 5C ) than in posterior region ( Fig. 5D ); macrotubercles blunt conical, each with 2–3 distal spines ( Fig. 5E ). Median cirrigerous segments with dorsal cirri reaching neurochaetal tips, basal tubercle of dorsal cirri swollen, bilobed, not projected as a cone ( Fig. 5F ); ventral cirri blackish basally. Neurochaetae bronze to brownish, smooth basally, denticulate along swollen subdistal region, tips slightly falcate, unidentate, smooth ( Fig. 5G ).

A juvenile 11 mm long specimen (UF 9454) has elytra without black spots ( Fig. 6A ). The elytra have two rows of macrotubercles and their size decreases posteriorly ( Fig. 6B , insets 1, 2), whereas the posterior microtubercles have short filaments ( Fig. 6B , inset 3). The cirrigerous parapodia have long cirrophores and tapered cirrostyles ( Fig. 6C ), and its neurochaetae are unidentate.

Additional specimens were 6–29 mm long, 4–16 mm wide. There are two rows of macrotubercles, the first row has slightly larger ones than those present in the second row, and this does not change with body size, although the second row can be more irregular and have additional tubercles.

Remarks. Nomenclatural problems regarding the type material of Polynoe muricata have confounded identification of this species. Savigny in Lamarck (1818: 308–309) provided a short diagnosis for the species and recorded it only from Mauritius (Île-de-France), based on one specimen collected by M. Mathieu (MNHN TYPE 335). Later, Savigny (1822: 21–22, Pl. 3, Fig. 1 ) described and illustrated a different specimen, from the Gulf of Suez, as P. muricata that actually belonged to a different species (here described as I. readi sp. nov.). Most subsequent authors followed Savigny’s (1822) text and illustration to identify I. muricata , which together with a poor understanding of species limits led to substantial confusion about the identity of I. muricata .

De Quatrefages (1866: 268) recognized Savigny’s specimen from Suez and Lamarck’s specimen from Mauritius as different species, incorrectly applied I. muricata to the former, and redescribed the latter specimen (including the holotype of P. muricata ), as I. glabra , together with a second syntype (MNHN TYPE 334), collected from the same locality ( Mauritius) by Péron & Lesueur. His selection of Lamarck’s type for his I. glabra , makes P. muricata and I. glabra objective synonyms. De Quatrefages provided some additional measurements and details for the illustrated specimen from Suez and noted differences between the two species. The main differences noted for I. glabra were that antennae, tentacular and dorsal cirri were tapered, not subdistally swollen, antennae were tiny, elytral margins were barely fimbriate, posteriorly smooth (hence the name), dorsal cirri were short, not surpassing notochaetae, and ventral cirri were minute. The slight difference in size of macrotubercles in rows 1 and 2 was indicated as “not resembling the spines illustrated by the naturalist (Savigny)” ( de Quatrefages 1866: 269).

We are thus restricting I. muricata (Savigny in Lamarck, 1818) to specimens with elytra having fimbriae and macrotubercles progressively smaller posteriorly. Day (1951: 14) noted that “there is great variation in the tubercles of the elytra.” However, elytra have mostly two rows of macrotubercles, and even though they can be eroded in older specimens, the striking size difference between those of the first row against the second one in I. readi sp. nov. (= I. muricata sensu Savigny 1822 ) separates it from those being gradually smaller posteriorly as in I. muricata . After the study of juvenile specimens of both species, we conclude this difference is present once they have two rows of macrotubercles; in I. muricata they are progressively smaller posteriorly, whereas in I. readi the first row has markedly larger macrotubercles.

Pettibone (1986: 3, 9–10) studied the syntype of I. glabra, MNHN 335 that was also the holotype of I. muricata . She did not illustrate it and correctly regarded it as a junior synonym of I. muricata (Savigny in Lamarck, 1818).

Iphione muricata resembles I. ankeri sp. nov. from Guam because both have two or up to three rows of macrotubercles; they differ because in I. muricata the posterior elytral margin is almost smooth, whereas in I. ankeri the microtubercles have long filaments. Omani specimens of I. muricata appears to be closely related to I. fimbriata based on COI sequence data ( Fig. 36 ).

Distribution. Red Sea to Mozambique, east to Samoa, on coral reefs and in rocky or mixed bottoms in shallow water (0–45 m).

Figures

FIGURE 1. Iphione readi sp. nov. (incl. I. muricata sensu Savigny (1822)), non-type specimens, left elytra 7 of non-type specimens (NHM 1941.4.4.168–172), arranged vertically. A. 9 mm long specimen. B. 16 mm long specimen. C. 23 mm long specimen. D. Iphione muricata (Savigny in Lamarck, 1818) non-type small specimen (11.5 mm long, MAGNT W2894), elytron 6 (E, F indicate areas enlarged in E and F, and asterisk indicates the enlarged area in G). E. Close-up of macrotubercle (Mt) and fimbriae. F. Same. G. Close-up of microtubercles along posterior margin. Scale bars: A, 0.4 mm; B, D, 0.5 mm; C, 0.8 mm; E, F, 100 µm; G, 70 µm.

FIGURE 2. Iphione ovata Kinberg, 1856, non-type specimens (2 ECOSUR 0000, Sta. 30, 13 mm long). A) Ventral view. B. Ventral body wall and most muscle bundles removed, oval masses are ovaries. C. Pharynx and gut ventral wall removed (*: enteric caeca). D. Pharynx, upper jaws separated medially. E. Pharynx, lower jaws separated medially (Pa: papilla, Ph: pharynx, St: stomach). Scale bars: A, 1.38 mm; B, 1.43 mm; C, 1.48 mm; D, 0.32 mm; E, 0.27 mm.

FIGURE 3. Pharynx features. A. Iphione henshawi Pettibone, 1986, non-type specimen (BPBM R481), anterior end, pharynx exposed, dorsal view. B. Same, ventral view. C. Same, frontal view of pharynx (numbers indicate papillae). D. Same, left lateral view. E. Iphione muricata (Savigny in Lamarck, 1818), restricted, non-type specimens (UF 411), smaller specimen (9 mm wide), pharynx, frontal view (numbers indicate upper papillae). F. Another, larger specimen (16 mm wide), pharynx, frontal view (numbers indicate upper papillae). Scale bars: A, B, 1.3 mm; C, 0.9 mm; D, 0.7 mm; E, 0.3 mm; F, 0.6 mm.

FIGURE 4. Iphione muricata (Savigny in Lamarck, 1818), restricted. A. Syntype MNHN 334, dorsal view. B. Same, anterior elytron (inset: fimbriae from another elytron). C. Same, median elytron. D. Chaetiger 14, left parapodium, posterior view after removal of notochaetae. E. Same, median neurochaetal tips. F. Syntype MNHN 335, two detached elytra before cleaning. G. Same, median elytron. H. Same, posterior elytron. I. Fimbriae of posterior elytron. J. Chaetiger 12, left parapodium, posterior view, after removal of notochaetae. Scale bars: A, 2 mm; B, 0.7 mm; C, 1 mm; D, 0.4 mm; E, 30 µm; F, 1.3 mm; G, 1.2 mm; H, 1.6 mm; I, 30 µm; J, 0.6 mm.

FIGURE 5. Iphione muricata (Savigny in Lamarck, 1818), restricted, non-type specimen (CAS 227559). A. Dorsal view, depressed by glass slide. B. Left elytron 7, dorsal view (C–E: sections enlarged in following pictures). C. Close-up of fimbriae. D. Close-up of smaller, marginal spines. E. Close-up of larger, inner spines. F. Chaetiger 12, left parapodium, posterior view, notochaetae partially trimmed off (after Methyl green staining). G. Same, basal neurochaetal tips. Scale bars: A, 2.5 mm; B, 1.5 mm; F, 0.9 mm; G, 0.3 mm.

FIGURE 6. Iphione muricata (Savigny in Lamarck, 1818), restricted, non-type specimen (UF 9454), juvenile. A. Dorsal view, after Methyl green staining. B. Right elytron 6, dorsal view (insets: enlarged areas indicated by numbers). C. Chaetiger 12, left parapodium, posterior view after removal of notochaetae. Scale bars: A, 0.8 mm; B, 0.4 mm; C, 0.2 mm.

FIGURE 11. Iphione henshawi Pettibone, 1986, non-type specimen (BPBM R481). A. Anterior end, dorsal view (elytra 1–3 removed). B. Left elytra 1–3, dorsal view. C. Elytron 3, close-up of fimbriae, some macrotubercles in the lower area. D. Same, close-up of macrotubercles. E. Cirrigerous left parapodium of median segment, posterior view. Scale bars: A, 0.8 mm; B, 1.4 mm; C, D, 0.3 mm; E, 0.5 mm.

FIGURE 36. Maximum Likelihood phylogeny of Iphione species with localities and catalog numbers. Numbers above branches are bootstrap support (>70 only) (PNG: Papua New Guinea) (modified after Piotrowski et al. 2024).