Iphione readi, Salazar-Vallejo & Piotrowski & Paulay, 2024

Salazar-Vallejo, Sergio I., Piotrowski, Christina N. & Paulay, Gustav, 2024, Revision of Iphione Kinberg, 1856 (Annelida, Aphroditiformia, Iphionidae), Zootaxa 5548 (1), pp. 1-84 : 61-69

Iphione readi sp. nov.

urn:lsid:zoobank.org:act:6FAE0FC9-1546-4ED0-BBC7-A187582753FB

Figs 1A–C , 27–30 , 36

Polynoe muricata : Savigny 1822: 21–22, Pl. 3, Fig. 1 (descr., partim). Eumolpe muricata : de Blainville 1828: 459, Pl. Chétopodes, Homocriciens Aphrodités, Fig. 1 (same figure from Savigny). Iphione muricata : Kinberg 1856: 383; Kinberg 1858: 8; Baird 1865: 181; de Quatrefages 1866: 266–268; Gravier 1901: 226–

231, Textfigs 232–239, Pl. 9, Figs 129–135 (redescr.); Potts 1910: 341; Fauvel 1932: 12–13; Fauvel 1933: 40–41; Fauvel

1935: 285–286; Day 1951: 14; Fauvel 1957: 3; Storch 1967: 148, pls. 1–4; Gibbs 1971: 123 (partim); Amoureux 1974: 430;

Amoureux et al. 1978: 68; Pettibone 1986: 9–16, Figs 1–5 (syn. partim, no figures made after Red Sea specimens); Wehe 2006: 62–67, Fig. 8 (partim); Goren et al. 2017: 134–136, Fig. 2 (non Savigny in Lamarck, 1818).

Type material. Red Sea, Saudi Arabia. Holotype ( UF 6343 ), Farasan Islands , Shuma Island (16°39’24.48” N, 41°33’16.92” E), 025 m, 25 Oct. 2014, R. Lasley, J. Moore & D. Uyeno, coll. One paratype ( UF 6340 ), Farasan Islands , Sarad Sarso (16°50’06.00” N ,, 41°36’48.24” E), 0.25 m, 25 Oct. 2014, D. Uyeno, R. Lasley & J. Moore, coll. (bent ventrally, elytra yellowish, with darker posterior margins and macrotubercles, some with white epibionts; macrotubercles in two rows, first one with largest tubercles; not measured nor dissected to avoid further damage). COI barcode: GenBank PQ 423939. One paratype ( UF 6360 ), Farasan Islands, Dumsuq Island (16°33’50.76” N, 42°03’30.6” E), 0–25 m, 26 Oct. 2014, R. Lasley, J. Moore & D. Uyeno, coll. (slightly bent ventrally, elytra with barely defined oblique dark bands, and darker areas along middorsal marginal portions, some with calcareous epibionts; elytra with two rows of macrotubercles, first row with largest tubercles; fimbriae often as long as macrotubercles along lateral margins; cirrigerous segments with basal tubercle of dorsal cirrophore projected; body 14 mm long, 9.5 mm wide, 29 chaetigers). COI barcode: GenBank PQ 423940.

Additional material

Red Sea, Gulf of Suez. Four specimens ( MNHN A-274 18), Mission Jousseaume, 1894 (no further data) (one straight, three specimens bent ventrally; elytra brownish, oblique longitudinal band well-defined, some with calcareous epibionts; macrotubercles in two rows, first row with markedly larger ones; body 15–29 mm long, 10–12 mm wide, 29 chaetigers) . One specimen ( MNHN A274-16 ), Mission Dolffus Egypte, Gulf of Suez, Sta. 17bis (28°14’ N, 33°22’ E), muddy sand and corals, 22 m, 25 Dec. 1928 (field data after Billard 1933), R.P. Dolfuss, coll. (slightly bent ventrally; elytra brownish without black spots; macrotubercles in two rows, first one with largest macrotubercles; body 19 mm long, 12.5 mm wide, 29 chaetigers) . Eleven specimens ( MNHN A274-28 ), Mission Jousseaume, Suez, 1894 (no further data; eight specimens bent ventrally; elytra brownish, oblique longitudinal band well-defined in most specimens, some with calcareous epibionts; macrotubercles in two rows, first row with largest ones; body 13.5–23 mm long, 8.0– 13.5 mm wide, 29 chaetigers) . One specimen ( MNHN A894 ), Mission A. Gruvel- P.G. Moazzo, Suez Bay , Jan.–Feb. 1934, entrance of canal (no further data; bent ventrally, some elytra detached, right parapodia of chaetiger 16 previously removed; elytra brownish, longitudinal bands barely visible; macrotubercles in two rows, first one with largest ones; body 31.5 mm long, 14 mm wide, 29 chaetigers) . One specimen ( ZMH PE 186 ), Gulf of Suez, 1914, F. Banarate, coll. (elytra yellowish, with an oblique darker band barely visible, with abundant sediment and epibionts; elytra with two rows of macrotubercles, first row with largest ones; cirrigerous segments with basal tubercle of dorsal cirrophore projected; body 25 mm long, 15 mm wide, 29 chaetigers) . One specimen ( ZMH V8365 ), Gulf of Suez, Bawmwarth, coll. (no further data; markedly bent ventrally; elytra yellowish without black bands, macrotubercles in two rows, first one with largest macrotubercles; not measured) .

Red Sea. Egypt. One specimen ( ZMH V6120 ), Khalij al Jimshan, Hartmeyer, coll. (no further data; elytra brownish with abundant white epibionts; elytra with two rows of macrotubercles, first row with largest ones, sometimes duplicate; body 21 mm long, 13 mm wide, 29 chaetigers) .

Red Sea. Saudi Arabia. One specimen ( CAS 192038 ), Red Sea Biodiversity Cruise 2013, Makkah Province, Thuwal, Kaust Beach , intertidal, 15 Mar. 2013, T.M. Gosliner, coll. (complete; two right parapodia dissected for observation, one midbody elytron removed for observing details; body 12.5 mm long, 8 mm wide, 28 chaetigers) . One specimen ( UF 2847 ), Thuwal (22°16’26.4” N, 39°03’03.6” E), 2–20 m, seaward reef slope and front of shelf reef, 10 Oct. 2012, M. Berumen & G. Paulay, coll. (bent ventrally; right parapodial and elytral fragments removed for molecular studies; 1 row of macrotubercles; basal tubercle of dorsal cirrophore projected; body 11 mm long, 6.5 mm wide, 29 chaetigers) . One specimen ( UF 3498 ), Farasan Islands, Naf Shuma (16°45’10.8” N, 41°36’18.0” E), 1–5 m, silty reef flat to slope, 9 Mar. 2013, A. Anker, P. Norby & G. Paulay, coll. (slightly bent ventrally; oblique brownish longitudinal bands, and posterior elytral margins barely pigmented; left parapodia of chaetigers 13 and 14 removed for observation; body 23 mm long, 12 mm wide, 29 chaetigers) . One specimen ( UF 3500 ), Farasan Islands, Mahama Island (16°29’20.4” N, 41°56’38.4” E), 4–17 m, reef rubble, fringing slope, 9 Mar. 2013, A. Anker, P. Norby & G. Paulay, coll. (elytra brownish with wide oblique blackish bands, and darker posterior elytral margins in posterior segments, many elytra with white epibionts; two rows of macrotubercles; basal tubercle of dorsal cirri clearly projected; body 15 mm long, 8.5 mm wide, 29 chaetigers) . One specimen ( UF 3505 ), Farasan Islands, Naf Shuma (16°45’10.8” N, 41°36’18.0” E), 1–5 m, silty reef flat to slope, 9 Mar. 2013, A. Anker, P. Norby & G. Paulay, coll. (slightly bent ventrally; elytra brownish with wide oblique blackish band and darker posterior elytral margins in median and posterior elytra; basal tubercle of dorsal cirrophore barely projected; body 18 mm long, 9 mm wide, 29 chaetigers) . One specimen ( UF 3514 ), Farasan Islands, Zahrat Durakah (16°50’09.6” N, 42°18’21.6” E), 4 m, fringing reef slope around sand canal, in rubble, 11 Mar. 2013, A. Anker, P. Norby & G. Paulay, coll. (bent ventrally; elytra brownish with wide oblique longitudinal band; body 17 mm long, 9 mm wide, 29 chaetigers) . One specimen ( UF 3583 ), off Thuwal, Abu Shosha Reef (22°12’10.8” N, 39°02’52.8” E), 5 m, sheltered side, 24 Mar. 2013, J. Moore & J. Bouwmeester, coll. (slightly bent ventrally; elytra yellowish with a barely continuous, longitudinal black band; two rows of macrotubercles; basal tubercle of dorsal cirri barely projected; body 11.5 mm long, 7.5 mm wide, 29 chaetigers) . One specimen ( UF 4488 ), Amr’s Dropoff (25°39’24.5916” N, 36°42’43.4592” E), 3–12 m, silty fringing reef, 1 Feb. 2016, J. Moore, coll. (juvenile; brownish longitudinal bands discontinuous in same elytron, made by darker areas; middorsal elytral margins darker than other areas; some elytra removed for observation; macrotubercles in two rows, first one with largest tubercles; body 17 mm long, 8.5 mm wide, 29 chaetigers) . One specimen ( UF 4489 ), Amr’s Dropoff (25°39’24.5916” N, 36°42’43.4592” E), 3–12 m, silty fringing reef, 1 Feb. 2016, J Moore, coll. (juvenile; right elytron 4 and parapodium previously removed, left parapodium of chaetiger 10 removed for observation (kept in container); macrotubercles in two rows, each with a few tubercles, first row with largest tubercles; body 5.5 mm long, 2.5 mm wide, 23 chaetigers; 11 elytral pairs) . One specimen ( UF 6287 ), Wasaliyat Island (17°46’58.08” N, 41°26’08.88” E), 0–25 m, patch reef, 17 Oct. 2014, D. Uyeno, R. Lasley & J. Moore, coll. (markedly bent ventrally, not measured to avoid any damage; elytra with discontinuous oblique dark bands, posterior elytral margins darker in posterior elytra; macrotubercles in two rows, first row with tubercles markedly larger than the others) . One specimen ( UF 6320 ), Farasan Islands, North Ghorab Island (17°06’30.96” N, 42°04’07.68” E), 5–7 m, fringing reef, silty bottom, 22 Oct. 2014, D. Uyeno, R. Lasley & J. Moore, coll. (elytra yellowish with brown spots forming irregular longitudinal band, some with white epibionts; macrotubercles darker, in two rows, size markedly different; median cirrigerous segments with dorsal cirrophore with basal tubercle barely bilobate; body 10.5 mm long, 6 mm wide, 29 chaetigers) . One specimen ( UF 6338 ), Farasan Islands, Dhi Dahaya (16°52’22.44” N, 41°26’27.6” E), 0.25 m, 24 Oct. 2014, D. Uyeno, R. Lasley, & J. Moore, coll. (juvenile; left parapodium of chaetiger 14 removed for observation; elytra maculate, dark areas small, spread apart, not forming distinct longitudinal bands; macrotubercles in two rows; basal tubercle indistinct; body 13 mm long, 7.5 mm wide, 29 chaetigers) .

Sudan. Six specimens ( NHM 1941.4.4.168-172), Sudan Pearl Fisheries Investigations, Dongonab, 1905, C. Crossland, coll. (variably bent; most elytra on site, smaller ones yellowish, larger ones with brownish elytra, oblique longitudinal black band indistinct; left elytra 7 removed from 3 specimens for observation (kept in container), macrotubercles of first row very large; body 9–23 mm long, 6.5–14 mm wide, 29 chaetigers) .

Djibouti. Two specimens ( MNHN A274-8 ), Djibouti, 1897, No. 29, M. Coutière, coll. (bent ventrally; elytra brownish, smallest one with a black spot, largest one with an oblique, longitudinal black band; macrotubercles in two rows, first row with largest macrotubercles; body 15–24 mm long, 8–11 mm wide, 29 chaetigers) . Four specimens ( MNHN A274-11 ), Mission Gravier, Pingodiu & Météor reefs, 18–20 m, 17 Jan. 1906, C. Gravier, coll. (dehydrated, one bent ventrally; elytra brownish with a black spot to an oblique longitudinal black band; macrotubercles in two rows, first one with markedly larger macrotubercles; no further dissected; body 10–16 mm long, 4.5–9.0 mm wide, 29 chaetigers) . One specimen ( MNHN A274-12 ), Mission Gravier, Île Moucha, Gulf of Tadjourah, coral, 23 Jan. 1904, C. Gravier, coll. (slightly bent ventrally; elytra brownish without black spot; macrotubercles in two rows, first row with markedly larger ones; body 10.5 mm long, 7.5 mm wide, 29 chaetigers) . One specimen ( MNHN A274-24 ), Obock, Obock, Mission Jousseume , 1895, M. Jousseume, coll. (body damaged; most elytra detached, some in container; elytra brownish with a darker area over outer elytral half; macrotubercles in two rows, first row with largest macrotubercles; body 19 mm long, 11.5 mm wide, 29 chaetigers) . One specimen ( MNHN A274-30 ), Mission Gravier Djibouti, Djibouti Bay , in base of coral, 13 Jan. 1904, C. Gravier, coll. (elytra brownish, outer elytral area darker; macrotubercles in two rows, first row with largest ones; body 17 mm long, 9.5 mm wide, 29 chaetigers) . One specimen ( MNHN A274-31 ), Mission Gravier Djibouti, Île Moucha, Gulf of Tadjourah, in rocky shore, 23 Jan. 1904, C. Gravier, coll. (juvenile, markedly bent ventrally; elytra brownish with dispersed black spots; macrotubercles in 1 row, very large and bent dorsally, at least the largest ones; not measured) .

Kenya. One specimen ( NHM unnumb.), Nyal , Mombasa, under coral boulders, D. McGregor, coll. (no further data; bent ventrally, some elytra detached; elytra yellowish, with two rows of macrotubercles, first row with largest ones; body 16 mm long, 8 mm wide, 29 chaetigers) . Two specimens ( USNM 97380 ) Mombasa, off South coast, removed from NHM 1924.6.18.187–191 (Elytra without blackish bands, with two rows of macrotubercles; body 16–33 mm long, 9–14 mm wide, 29 chaetigers; smallest specimen with longer macrotubercles; neurochaetae unidentate) .

Tanzania. One specimen ( NHM 1937.9.2.44–45), John Murray Expedition , Sta. 111 (05°04’18” S, 39°14’12” E), Agassiz trawl, 73–165 m, 14 Jan. 1934 (juvenile; partially dissected, 9 pairs of elytra, some elytra previously removed; elytra yellowish, with 1–2 fimbriae, and two large macrotubercles, each representing one row, largest macrotubercle towards external margin, additional one close to posterior margin; no further dissections to avoid further damage; body 4 mm long, 2.5 mm wide, 23 chaetigers) .

South Africa. One specimen ( USNM 81940 ), Delagoa Bay , near Johaca Island, Jul. 1935, C.J. van der Horst, coll. (all elytra previously removed, and two right parapodia dissected from chaetigers 14 and 15, all kept in container; body 25 mm long, 15 mm wide, 29 chaetigers) .

Sri Lanka. Two specimens ( NHM 1928.4.26.889/890), Gulf of Manaar (no further data; one markedly bent ventrally; elytra brownish, with 2–3 rows of macrotubercles, first row with the largest ones; less bent specimen 19.5 mm long, 11.5 mm wide, 29 chaetigers) .

Andaman Islands. Four specimens ( MNHN A399 ), RV Investigator, Sta. 665, 5 Feb. 1924 (no further data; slightly bent ventrally; elytra golden to brownish, with a wide longitudinal blackish band, most with 2–3 rows of macrotubercles, largest ones lateral in first row; cirrigerous parapodia with large basal tubercle to dorsal cirrophore; body 14.5–34 mm long, 8.5–14.0 mm wide, 29 chaetigers) . One specimen ( NHM 1938.5.7.3), Sta. 665, South Point , Outram Island, no further data (slightly bent ventrally; some elytra previously removed (kept in container), each with two rows of macrotubercles, the first row with largest ones; body 27.5 mm long, 17 mm wide, 29 chaetigers) .

Indonesia. One specimen ( ZMA V475 ) Yakarta Bay , Java, C.P.H. Sluiter, coll. (no further data; markedly bent ventrally; elytra brownish, darker along external elytral half; macrotubercles in two rows, first row with macrotubercles markedly larger than those present in second row; cirrigerous segments with basal tubercle of dorsal cirrophore projected; not measured) . One specimen ( ZMH PE 187 ), Java (no further data; markedly bent ventrally; brownish, elytra without dark bands, and with abundant sediment and epibionts; elytra with two rows of macrotubercles, first row with largest ones, some broken; not measured to avoid further damage) .

Hong Kong. One specimen ( AM 19943 ), Breaker Reef, 6 Apr. 1986, P. Hutchings, coll. (juvenile, markedly bent ventrally; elytra yellowish with an oblique longitudinal black band; macrotubercles in two rows, first one with largest ones; not measured) . One specimen ( AM 20044 ), Chek Chau, W side of Hong Kong, 5 m, 14 Apr. 1986, P.A. Hutchings, coll. (bent ventrally; elytra with an oblique longitudinal, ill-defined black band; macrotubercles in two rows, first row with largest ones; cirrigerous segments with basal tubercle of dorsal cirrophore projected; body 12 mm long, 8 mm wide, 29 chaetigers) .

Vietnam. Four specimens ( MNHN A398 ), Bay of Nhatrang. 1931, C. Dawydoff, coll. (no further data; three markedly bent specimens, elytra yellowish to brownish, with a wide, longitudinal black band, and darker posterior margins; each elytron with 1–3 rows of macrotubercles, smallest specimen with a single row; body 9–29 mm long, 4–15 mm wide, 28–29 chaetigers) .

Australia. One specimen ( AM 17861 ), Christmas Island , 1980, R. Hicks, coll. (no further data; juvenile, markedly bent ventrally; elytra yellowish, without black spots; macrotubercles in two rows, first one with largest macrotubercles; not measured) . One specimen ( MAGNT W17 ), Burford Island , NT, Sta. CP 14, reef flat, under rocks, low subtidal, 13 Oct. 1981, J.R. Hanley, coll. (markedly bent ventrally; all elytra previously detached; elytra brownish with an oblique longitudinal black band; macrotubercles in two rows, some broken, first row with macrotubercles markedly larger than those present in second row; not measured) . One specimen ( MAGNT W18 ), Burford Island , NT, Sta. CP 14, reef flat, under rocks, low subtidal, 13 Oct. 1981, J.R. Hanley, coll. (markedly bent ventrally; many elytra previously detached; elytra brownish with an oblique longitudinal black band, and two rows of macrotubercles, many broken, first row with macrotubercles markedly larger than those present in second row; not measured) . One specimen ( MAGNT W2816 ), East Point , NT, Sta. EP 11, rock embedded in mud, subtidal, 18 May 1985, J.R. Hanley, coll. (juvenile; elytra brownish, with two rows of macrotubercles, first row with largest ones; body 10 mm long, 7 mm wide, 28 chaetigers) . One specimen ( MAGNT W2817 ), East Point , NT, Sta. EP 11, rock embedded in mud, subtidal, 18 May 1985, J.R. Hanley, coll. (juvenile; elytra brownish, with two rows of macrotubercles, first row with largest ones; body 11.5 mm long, 7 mm wide, 29 chaetigers) . One specimen ( MAGNT W6883 ), Scorpion Island , WA, Sta. RH 91-55, fringing shore reef, low subtidal, 15 Aug 1991, J.R. Hanley, coll. (bent ventrally; first right elytron detached; elytra with a oblique longitudinal black band, macrotubercles in two rows, many broken, first row with largest macrotubercles; body 12 mm long, 8 mm wide, 29 chaetigers) . Five specimens ( NHM 1925.1.28.36–40), Christmas Island , no date or depth data, C.W. Andrews, coll. (elytra yellowish to brownish, at least external elytral half, with abundant sediment particles and some epibionts; some elytra previously removed, each with two rows of macrotubercles, first one with macrotubercles markedly larger than those in second row; body 10–23 mm long, 6.5–11.0 mm wide, 29 chaetigers) .

Papua New Guinea. One specimen ( MNHN IA 2021-686 ), Papua Niugini Expedition, Sta. PS 31 (5°08.2’ S, 145°49.4’ E), 10–37 m, 27 Nov. 2012, S. Hourdez et al., coll. (markedly bent ventrally; elytra dirty yellowish, oblique longitudinal black band present; macrotubercles in two rows, first row with tubercles markedly larger than those present in following row; cirrigerous segments with basal tubercle of dorsal cirrophore projected; not measured) .

Vanuatu. Two specimens ( MNHN A274-6 ), Malekula (Mallicolo), Malampa, Nov. 1894, M. Francois, coll. (tied up, poorly preserved; elytra brownish with a median oval black spot (smaller specimen), or an oblique longitudinal black band; not further dissected; body 17–19 mm long, 9–11 mm wide, 29 chaetigers) .

American Samoa. Two specimens ( USNM 81937 ), Tutuila Island , Pago Pago, without further data (body 28–30 mm long, 13–14 mm wide, 29 chaetigers) .

Diagnosis. Iphione with median antenna reduced to nuchal papilla; elytra fimbriate; macrotubercles spine-like to digitate with distal spines, in 2–3 rows, first row with macrotubercles markedly larger than those in other rows; cirrigerous segments with dorsal cirrophores with basal tubercle projected; neurochaetae unidentate.

Description. Holotype (UF 6343) complete, slightly bent ventrally, a few elytra previously detached (in container); body 19.5 mm long, 11 mm wide, 29 chaetigers. Elytra brownish, with oblique longitudinal black bands, darker areas along inner posterior margins, fimbriae with abundant fine calcareous particles, and calcareous epibionts ( Fig. 27A ); notochaetae whitish, neurochaetae golden. Body wall pale, venter pale.

Elytra with fimbriae ( Fig. 27C , inset 2), better developed along lateral margins throughout body, peduncle thin, fimbriae incorporate abundant sediment particles, fimbriae about 5 times longer than wide; macrotubercles digitate, about 3 times longer than wide, arranged in two rows, first one with largest macrotubercles, the largest one usually in the borderline between hyaline and opaque elytral surfaces, bent middorsally ( Fig. 27C , inset 1), macrotubercles in irregular second row markedly smaller, digitate; microtubercles along posterior margin, some with long filaments ( Fig. 27C , 3 ).

Prostomium roughly hexagonal ( Fig. 27B ), wider than long, with anterior incision running along anterior prostomial third. Anterior lobes projected into ceratophores, slightly darker, longer than prostomium, as long as ceratostyles, subdistally swollen, tips tapered. Eyes black, positioned in posterior prostomial half, anterior eyes slightly larger than posterior ones, placed in lateral protuberances, posterior eyes dorsal, in prostomial posterior margins. Palps thick, bent ventrally, twice longer than antennae, with longitudinal rows of papillae, tips tapered, elongate. Nuchal papillae distinct, pear-shaped, under nuchal flap.

Tentacular segment dorsally reduced, without chaetae, tentacular cirri about as long as palps. Facial tubercle visible dorsally, pale.

Segments 2–4 directed anteriorly. Second segment visible dorsally, with a semicircular nuchal lappet, expanded laterally, slightly wider than long, covering nuchal papilla. Ventral buccal segment inserted ventrally, almost three times longer than following cirri, directed ventrally, projected beyond chaetal tips. Dorsal nodules well-developed in segment 3, markedly smaller in segments 4–5.

Median cirrigerous segments with dorsal cirri projected beyond chaetal tips; basal tubercle of dorsal cirrophore projected as a single lobe ( Fig. 27D ). Notochaetae very abundant, dirty white, with series of transverse funnel-shaped spinose rows, tips bare. Neuropodia with neuracicular lobe truncate, with small globular papillae. Neurochaetal lobe with marginal papillae, neurochaetae abundant, basally smooth, subdistally swollen, with many rows of fine denticulations along swollen region, tips barely falcate, unidentate, sharp ( Fig. 4D , inset).

Anus dorsal, visible between elytral pairs 12 and 13.

Etymology. The specific epithet is after Dr. Geoffrey Read, editor of Annelida in the World Register of Marine Species (WoRMS), in recognition for his long-term editorial efforts for updating the website. The specific epithet is a noun in the genitive case ( ICZN 1999, Art. 31.1.2).

Variation. Examined specimens were 4–34 mm long, 2.5–15.0 mm wide. The first row of macrotubercles includes the largest ones, however, their size depends on the number of macrotubercles such that when there are only 3–4, they can be markedly larger than when there are 6–7 ( Fig. 1A–C ), but even then, they are markedly larger than those present in the following row.

Some specimens (MNHN A 274 18) are bent ventrally, and elytral features were observed in their right elytron 6. The elytra have an oblique longitudinal black band, macrotubercles in two rows but first one had the largest macrotubercles, and the largest macrotubercle is the outermost, usually rising from the borderline between the hyaline and opaque elytral surfaces; the second row is more irregular and its macrotubercles are markedly smaller than those present in first row, although the outermost can be slightly smaller than the largest one, the others are markedly smaller. The filaments of the marginal microtubercles are delicate and are not as long as those present in the neotype.

One Red Sea specimen (CAS 192038) has a longitudinal wide rust-red band along the elytra ( Fig. 28A ); elytra with fimbriae and a row of very large conical macrotubercles ( Fig. 28B ), and 1–2 additional ones, size decrement exponential, not gradual, with smaller tubercles along margin ( Fig. 28B , insets); cirrigerous segments with dorsal cirri markedly surpassing chaetae ( Fig. 28C ), cirrophore with basal tubercle conical; neurochaetae unidentate, tips falcate.

A juvenile (5.5 mm long) (UF 4489) has cephalic appendages and dorsal cirri tapered, not subdistally swollen, as is the case in larger specimens; elytra transparent with irregular brownish spots ( Fig. 29A, B ). The elytra have a single row with 2–3 large conical macrotubercles, and fimbriae are very short, 8-shaped structures bordering elytra along their posterior margin ( Fig. 29C ). The cirrigerous segments have dorsal tubercles without basal tubercle ( Fig. 29D ); neurochaetae falcate, sharp.

Remarks. Iphione readi sp. nov. corresponds to I. muricata sensu Savigny (1822) . Iphione readi resembles I. muricata (Savigny in Lamarck, 1818) (see above) by having elytra with fimbriae well-developed, and neurochaetae unidentate. These two species differ in the relative size of macrotubercles and in the presence of fimbriae along posterior elytral margins. In I. readi , the macrotubercles of the first row are markedly larger than those present in the following row, and the posterior margin has microtubercles with distal spines, whereas I. muricata has macrotubercles progressively smaller posteriorly, and the microtubercles along the posterior margin have very few or no spines.

Savigny (1822: 21) indicated that the plate for his I. muricata was made after a specimen from the Gulf of Suez. The original figure by Savigny has been simplified ( Fig. 30 ) to illustrate the main diagnostic features. The body is oval, almost completely covered by elytra ( Fig. 30A ), and cephalic appendages and dorsal cirri are exposed beyond the elytral margins, they are all subdistally swollen. The anterior end was observed from an oblique frontal perspective; palps have distinct rows of papillae, there is no median antenna and only one pair of eyes is visible after the first elytra were removed, leaving the basal elytral plate fixed on elytrophores ( Fig. 30B ). Elytra are areolate and have a thin longitudinal dark band along body, and each elytron has two rows of macrotubercles, those present along the first row are markedly larger than the others, but their arrangement was unclear because it looks like a row with alternating long and short, instead of being shown as arranged in two rows, and lateral margins are fimbriate ( Fig. 30C ). The pharynx was dissected, as indicated by the collapsed marginal papillae ( Fig. 30D ), and the jaws have two accessory denticles ( Fig. 30E ). The cirrigerous segments show a dorsal cirrophore with a round, projected basal tubercle ( Fig. 30F ).

The dissection damaged the specimen that Savigny illustrated, and it appears to have been lost soon after his publication. However, de Quatrefages (1866: 266–268) provided additional information of the specimen, but it was based upon the illustrations, not on the specimen that Savigny had studied and illustrated, though de Quatrefages did not include any illustration. An earlier repetition of the original Savigny’s illustration was made by de Blainville (1828: 459), who made slight modifications to the original illustration, because he indicated a wider longitudinal black band. Grube (1870: 281) noted that Savigny’s collection was not transferred to the Paris Museum. The specimen that Savigny had illustrated was not found in the Paris collections.

Gravier (1901: 226) studied and illustrated several Red Sea specimens of I. muricata sensu Savigny (1822) . His close-up of the marginal portion of the elytra (his Pl. 9, Fig. 133) shows the large, finger-shaped, slightly bent macrotubercles and fimbriae. The macrotubercles in the first row are 3–5 times larger than those present in the second row.

Pettibone (1986) did not illustrate specimens from the Red Sea, but used specimens from other localities including type specimens of other species she regarded as junior synonyms. None of her figures show the striking size difference in macrotubercles characterizing I. readi sp. nov. and none may represent this species. Wehe (2008) redescribed the species with Red Sea specimens; he illustrated the larger macrotubercles in the first row.

Although many records for I. muricata followed Savigny illustrations, once the presence of fimbriae or the size of macrotubercles were disregarded as diagnostic features, several records of I. muricata do not match I. muricata (Savigny in Lamarck, 1818), or I. readi . For example, Gibson (1886: 150) recorded it from Liverpool, but he noted all scales were lost, which is rarely the case in Iphione ; it was later shown by Hornell (1891: 224) that this record belonged to Gattyana cirrhosa ( Pallas, 1766) . Potts (1910: 341) noted one of his many specimens was found on a white-brown holothurian in the Addu Atoll, Maldives. Horst (1917b: 65) reported 50 specimens from Indonesia. Pruvot (1930: 3) recorded I. muricata from New Caledonia and described and illustrated the elytra as marginally smooth, which is not the case for true I. muricata . Monro (1931: 7) recorded the same species from several Southeastern Australian reefs. Fauvel (1932: 12) recorded it from several localities in the Indian Ocean. Day (1934: 25; 1962: 628) recorded I. muricata for South Africa and Madagascar, but no diagnosis or illustration were given. Fauvel (1937: 51) recorded I. muricata as a senior synonym of I. hirotai Izuka, 1912 from Japan, even though the latter has smooth elytral margins. Okuda (1937: 267) reached the same conclusion and recorded it from Koror (or Kororu), Palau Islands. Monro (1939: 168) also disregarded the difference in elytral margins and recorded smooth and papillose specimens under the same name. For his paper on New Caledonian polychaetes, Fauvel (1947: 13) used Schmarda (1861) illustration for his I. peronea . Fauvel (1953a: 173) recorded this species for Tahiti without diagnosis or description. Hartman (1966: 175) recorded it from subtidal substrates in Hawaii. The record by Amoureux (1974: 430) from 15 m off Tanikely, Madagascar is challenging to interpret because it was based upon a 4 mm long specimen with 20 pairs of elytra. He indicated he was following Rullier (1972: 34), but the latter indicated there were only 13 pairs of elytra, although the latter recorded one specimen with marginal papillae and five others without them, just as recorded by Pruvot (1930: 4) for New Caledonia. Bailey-Brock & Hartman (1987: 235, Fig. 3.2.4) used Gravier (1901) illustrations, made after Red Sea specimens, for their report on Hawaiian polychaetes. The record by Hanley (1992: 364) for Hong Kong did not include diagnosis or illustrations. This series of records emphasizes the historical confusion regarding the delineation of I. muricata (Savigny in Lamarck, 1818).

Distribution. Red Sea, South Africa and east to Samoa, on reefs and rocky or mixed bottoms (0–165 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 8. Iphione fimbriata de Quatrefages, 1866, reinstated, paratype of I. fustis Hoagland, 1920 (USNM 18969).A.Anterior end, dorsal view, after Methyl green staining. B. Anterior elytra, above: right elytron from chaetiger 2; below: left elytron from chaetiger 4. C. Left elytron from median chaetiger, dorsal view (insets: fimbriae, and smaller marginal spines). D. Chaetiger 14, left parapodium, posterior view after Methyl green staining. E. Same, upper and medial neurochaetae. F. Same, medial and lower neurochaetae. Scale bars: A, 1.1 mm, B, 1 mm; C, 2 mm; D, 0.6 mm; E, 100 μm; F, 80 μm.

FIGURE 27. Iphione readi sp. nov. (incl. I. muricata sensu Savigny (1822)) holotype (UF 6343). A. Dorsal view. B. Anterior end, dorsal view, after Shirlastain-A. C. Left elytron 6, insets indicated by number (1, macrotubercle; 2, smallest fimbria; 3, microtubercle). D. Chaetiger 12, left parapodium, posterior view after removal of notochaetae (inset: close up of upper neurochaetal tips). Scale bars: A, 2 mm; B, D, 0.5 mm; C, 1.2 mm.

FIGURE 28. Iphione readi sp. nov. (incl. I. muricata sensu Savigny (1822)), non-type specimen (CAS 192038). A. Dorsal view. B. Left elytron 6, seen from above (insets, from left to right: fimbriae, large macrotubercle, marginal tubercles). C. Chaetiger 12, left parapodium, posterior view, after Methyl green staining (inset: median neurochaetae). Scale bars: A, 1 mm; b, 0.7 mm; C, 0.3 mm.

FIGURE 29. Iphione readi sp. nov. (Incl. I. muricata sensu Savigny (1822)), non-type juvenile specimen (UF 4489).A. Dorsal view. B. Same, after Methyl green staining. C. Right elytron 4, inner margin slightly bent. D. Chaetiger 10, left parapodium, posterior view. Scale bars: A, B, 0.6 mm; C, 0.4 mm; D, 0.1 mm.

FIGURE 30. Iphione readi sp. nov. (Incl. I. muricata sensu Savigny (1822)), original illustrations after explanation by Audouin (1825: 66). A. Dorsal view. B. Anterior end, oblique frontal view. C. Median elytron, schematic. D. Pharynx with jaws and papillae, lateral view. E. Jaws in lateral, frontal and posterior views. F. Median cirrigerous chaetiger, right parapodium, posterior view (Abbreviations: LA, lateral antennae; Pa, palp; TC, tentacular cirri; VC2, ventral cirrus segment 2; 3, segment 3).

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).