Pherecardia striata ( Kinberg, 1857 )

Salazar-Vallejo, Sergio I., Piotrowski, Christina N., Gustav, Leslie Harris & Paulay, Gustav, 2025, Revision of Pherecardia Horst 1886 (Annelida Amphinomidae), Zoosystema 47 (28), pp. 691-720 : 696-705

Pherecardia striata ( Kinberg, 1857) , restricted

( Figs 1A, B, F, G; 2A, B; 3; 4; 6-8)

Hermodice striata Kinberg, 1857: 13 ; 1910: 35 (diagn. repeated), pl. 12, fig. 8. — Ehlers 1920: 13-14. — Augener 1927: 122 ( partim); 1933: 188.

Pherecardia lobata Horst, 1886: 165-166 , pl. 7, figs 10-14; 1909: 300 (syn.); 1911: 17-21 (syn.); 1912: 32-33 (syn.) — Collin 1902: 741. — Fauvel 1919a: 336; 1919b: 349-350. — Day 1934: 27. — Bleeker & van der Spoel 1992: 152.

Amphinome sericata Fischli, 1903: 95-98 , pl. 4, figs 1, 2, pl. 7, figs 45-49, pl. 8. figs 79-80.

Hermodice pennata Treadwell, 1906: 1165 , fig. 41 (junior syn. P. striata fide Hartman 1956: 251 ).

Eucarunculata grubei Malaquin & Dehorne, 1907: 358-361 , pl. 51, fig. 2, pl. 53, figs 12-15, 17-20. — Potts 1909: 365-366. — McIntosh 1925: 13-14, pl. 1, figs 3-4.

Eucarunculata grubei var. gracilis Potts, 1909: 366-367 .

Eucarunculata grubei var. minuta Potts, 1909: 367 .

Pherecardia striata – Monro 1928: 77; 1933: 7. — Fauvel 1935: 292-293 ( partim); 1936: 257. — Okuda 1937: 265-266, fig. 5. — Monro 1939: 166-167 (epitokes). — Hartman 1948: 46; 1966: 181-182. — Day 1957: 67 (syn., partim); 1962: 636; 1967: 131, fig. 3.2p-t. — Fauchald 1977b: 13. — Blake 1991: 84. — Imajima 2005: 86-88, fig. 36A-G. — Cañete 2017: 199-201, Fig. 1.

DIAGNOSIS. — Pherecardia with dorsum bearing longitudinal stripes of similar width along body. Notopodia darker along anterior surface, posterior surface paler, not forming a complete blackish ring. Notochaetae abundant, longer than dorsal cirri along anterior chaetigers.

TYPE MATERIAL. — Holotype of Hermodice striata Kinberg, 1857 . French Polynesia • 1 specimen; Moorea; KSF Eugenies Expedition; Sta. 1245; coral gravel; SMNH 6080.

Holotype of Pherecardia lobata Horst, 1886 . Locality uncertain • 1 specimen; RMNH VER.1295 (locality given as Ambon Bay, III.1901, R. Semon leg. on label; the holotype specimen matches the original description, the label indicates that it was collected in 1901, 15 years after the original description. Horst (1886) noted that the provenance of the specimen was uncertain. Thus, the locality ( Ambon Bay) on the label is a later addition and the type locality must be considered uncertain).

Holotype of Hermodice pennata Treadwell, 1906 . Hawaiian Islands • 1 specimen; USFS Albatross; Sta. 4162; Nihoa Island; 38- 44 m; 8.VIII.1902; USNM 5209.

ADDITIONAL MATERIAL. — Kenya • 1 specimen; Mombasa; D. MacGregor leg., no further data; NHML 1961.8 .32 (complete, longitudinal stripes well defined dorsally, ventrally along anterior body half; notopodia without black rings; caruncle with eight pairs of lateral lobes [last one tiny]; median antenna without tip, 1/3 as long as caruncle; pygidium with anus terminal, anal plate truncate, probably in regeneration; body 62 mm long, 9 mm wide, 62 chaetigers). Madagascar • 1 specimen; Nossy Vorona ; off ENE side; 13°25’27”S, 48°21’15”E; 4 m depth; broken reef, extracted from large, dead Porites ; 15.V.2008; A. Anker, G. Bakary, E. Boissin, H. Bruggemann, T. Horeau, F. Michonneau, G. Paulay & T. Werner leg.; UF 750 (complete; dorsal longitudinal stripes of similar width, often branching; caruncle, interramal areas and venter maculate; anterior eyes two times as large as posterior ones, not protruded; caruncle with median ridge complete, with seven pairs of lateral smooth lobes, with venation; notopodia with anterior surface darker, without distal black rings; pygidium with anus terminal, anal plate truncate, barely notched; 83 mm long, 8 mm wide, 64 chaetigers) • 1 specimen; Nosy-Be; 7-26.V.2008; G. Paulay et al. leg.; UF 4230 (complete; dorsal longitudinal stripes of similar width, a few branching; caruncle interramal areas and venter maculate; anterior eyes three times as large as posterior ones, not protruded; caruncle with median ridge complete, with eight pairs of lateral smooth lobes, with venation; notopodia with anterior surface darker, without distal black rings; pygidium with anus terminal, anal plate truncate, barely notched; 108 mm long, 10 mm wide, 73 chaetigers) .

Mascarene Islands, La Réunion • 1 specimen; Saint Leu , Maison Verte, Cimetière; 21°11’36”S, 55°16’56”E; 8-9 m depth, fore reef, within dead digitate Acropora ; 13.VIII.2007; H. Bruggemann, N. Hubert, F. Michonneau & G. Paulay leg.; UF 654 (anterior fragment; dorsal longitudinal stripes of similar width, a few branching; not maculate; anterior eyes two times as large as posterior ones, not protruded; caruncle with median ridge complete, with 7-8 pairs of lateral lobes, barely pinnate; notopodia with anterior surface darker, without distal black rings; 26 mm long, 7 mm wide, 26 chaetigers) • 3 specimens; Saint Leu , Sec Jaune; 21°9’11”S, 55°16’51”E; 6-15 m depth; 17.VIII.2007; H. Bruggemann, N. Hubert, F. Michonneau & G. Paulay leg.; UF 664 (two complete, one without posterior region; dorsal longitudinal stripes of similar width, a few branching; not maculate; interramal belt and venter maculate; anterior eyes two times as large as posterior ones, not protruded; caruncle with median ridge complete, twisted, with seven pairs of lateral smooth lobes; notopodia with anterior surface darker, without distal black rings; pygidium with anus terminal, anal plate truncate, margin entire; 33-34 mm long, 7-8 mm wide, 50 chaetigers) .

Indonesia • 1 specimen; Lesser Sunda Islands; RV Siboga Expedition ; Sta. 60; Timor, Samau Island, Haingsisi; 23 m depth, reef, Lithothamnion in 3 m and less; 27-28.IV.1899; ZMA V.POL.1070.1 (complete, slightly twisted, breaking in two from chaetiger 17; some chaetae from chaetigers 10 and 25 removed for observation; dorsum with rather regular longitudinal stripes of similar width; caruncle with 6-8 lateral lobes with venation, not rugose; branchial filaments basally darker, paler distally; pygidium with anus terminal, anal plate truncate; body 53 mm long, 7 mm wide, 48 chaetigers) • 1 specimen; Maluku; RV Siboga Expedition ; Sta. 240; Banda anchorage; 9-45 m depth; trawl, black sand and coral; 22.XI–1.XII.1899; ZMA V.POL.1070.2 (bent ventrally; juvenile; some parapodia previously removed [lost?], breaking in two parts; dorsum with longitudinal stripes of similar width; caruncle with 5-6 lateral lobes with venation, not rugose; branchial filaments basally darker, tips paler; pygidium with anus terminal, anal plate wider than long, medially notched; body 19 mm long, 4 mm wide, 35 chaetigers) • 1 specimen; Lesser Sunda Islands, RV Siboga Expedition ; Sta. 300; 10°48.6’S, 123°23.1’E; 918 m; trawl, muddy bottom; 30.I.1900; ZMA V.POL.1070.4 (bent laterally, juvenile; dorsum with longitudinal stripes of similar width; caruncle with 5-6 lateral lobes with venation, not rugose; branchiae basally darker, tips paler; pygidium with anus terminal, anal plate slightly longer than wide, round; body 19 mm long, 4 mm wide, 35 chaetigers) • 1 specimen; Ambon Island; 3.II.1902; V. Kamper leg.; RMNH VER.1296 (partially dried, with salt spherules over chaetae and branchiae; dorsum with rather regular longitudinal stripes; venter with paler longitudinal stripes, less regular; caruncle in regeneration; eyes black, not protruded from prostomial margin; caruncle with lateral lobes with venation, not rugose; branchiae dark with tips pale; pygidium with anus terminal, anal plate truncate; body 93 mm long, 13 mm wide, 59 chaetigers) • 25 specimens; Ambon Island; 1887; H. Brock leg.; ZMH PE29 (17 complete; most grayish with purple to blackish longitudinal rather regular stripes, other brownish with more irregular streaking; caruncle lobes foliose, slightly indented, 6-10 pairs; eyes not protruded from prostomial margins; median antenna 1/4 to 2/3 as long as caruncle; pygidium with anus terminal, anal plate lobate, often slightly notched medially; complete with body 16-70 mm long, 6-13 mm wide, 32- 45 chaetigers • 1 specimen; Ambon Island; 1887; H. Brock leg.; ZMH PE30 (soft, complete, unpigmented [this has been retained in P. striata with hesitation after it was found in the type locality of P. lobata Horst, 1912 , which is a junior synonym of P. striata , but the fact that this was collected in the same locality as the type does not preclude it being something else]; left parapodia of chaetigers 24 and 25 previously removed [kept in container]; caruncle with eight pairs of lateral lobes; median antenna half as long as caruncle; pygidium with anus terminal, anal plate truncate; body 50 mm long, 10 mm wide, 52 chaetigers).

East Timor • 1 specimen; Timor , between Timor and Nusa Besi; anchorage; RV Siboga Expedition ; Sta. 282; 08°25.2’S, 127°18.4’E; 27- 54 m depth, trawl, sand and coral; 15-17.I.1900; ZMA V.POL. 1010.3 (complete, thinner medially, with progressively longer notochaetae from chaetiger 25 to end of body; some chaetae of chaetigers 25 and 50 removed for observation; dorsum with longitudinal stripes of similar width; caruncle with 5-6 lateral lobes with venation, not rugose; branchial filaments basally darker, paler distally; pygidium with anus terminal, anal plate with a median notch; body 60 mm long, 8 mm wide, 62 chaetigers) .

South China Sea • 1 specimen; Macclesfield Bank ; no further data; NHML 1925.1 .128.143 (bent ventrally, without posterior end; anterior end damaged; caruncle with 5-6 pairs of lateral lobes; median antenna lost; longitudinal dorsal stripes barely visible; notopodial black rings not seen; body 39 mm long, 9 mm wide, 39 chaetigers) . Philippines • 1 specimen; Batangas Province, Tingloy, Maricaban Island, Sepok Point dive site; Hearst Philippine Biodiversity Expedition 2011; 13°41’23”N, 120°49’47”E; 6.V.2011; R. Van Syoc leg.; CASIZ 187243 (twisted dark specimen, with many longitudinal stripes, most branching laterally; notopodia with darker anterior surfaces, paler posteriorly; caruncle with median ridge complete, with 6 lateral lobes, smooth; median antenna 2/3 as long as caruncle; pygidium with anus terminal, anal plate truncate, margin incised; body 60 mm long, 10 mm wide, 59 chaetigers) • 5 specimens; Occidental Mindoro, Lubang Islands, Cabra Island, west of Cheeze Whiz Wall dive site; Verde Island Passage Expedition; Sta. LUB-12; 13°52’48”N, 120°1’11”E; 0-36 m depth; 24.V.2014; C. Piotrowski leg.; CASIZ 197440 (juveniles; two with pharynx exposed; smallest one broken in two pieces; smaller ones with paler pigmentation; all with darker areas over anterior notopodial surface; caruncle with 4-5 lateral lobes, smallest one with venation running along lobe, running across them in largest one; body 6.5-26.0 mm long, 1.5- 2.5 mm wide, 20-33 chaetigers) • 1 specimen; Lubang Island “Beyond Fish” dive site; 2014 Verde Island Passage Expedition; Sta. LUB-10; 13°47’36”N, 120°5’40”E; 7-30 m depth; 23.V.2014; C. Piotrowski leg.; CASIZ 198634 (complete juvenile specimen, straight; dorsal longitudinal stripes similar width throughout; notopodia darkly pigmented anteriorly only, lacking complete distal blackish rings; anterior eyes larger than posterior, not protruded from prostomial margin; caruncle complete, smooth, 4-5 lateral lobes with violet venation; median antenna 1/3 caruncle length, branchiae dark violet basally; anus terminal; 12 mm long, 3 mm wide, 28 chaetigers) • 1 specimen; Batangas Province, Tingloy, Maricaban Island, Sepok Point dive site; Hearst Philippine Biodiversity Expedition 2011; Sta. HEP-60; 13°41’17”N, 120°49’38”E; 0-24 m depth; coral rubble; 13.V.2011; E. Jessup & C. Piotrowski leg.; CASIZ 187263 (complete juvenile specimen, proboscis everted; faint longitudinal stripes of similar width throughout, in life dorsum with white spots; notopodia with anterior dark purple inverted triangular spots reddish in life, lacking blackish ring, in life ceratophores of dorsal cirri with yellow ring; red anterior eyes twice size of posterior eyes, not protruded from margin; caruncle with smooth complete median ridge, three lobate lateral lobes with simple venation; median antenna 2/3 length of caruncle, 2/3 length of lateral antennae; branchiae unpigmented; pygidium bulbous, in life with yellow ring; 5 mm long, 1 mm wide, 16 chaetigers) • 1 specimen; Tingloy, Maricaban Island, Bethlehem dive site; Hearst Philippine Biodiversity Expedition 2011; Sta. HEP-65; 13°40’23”N, 120°49’38”E; filamentous algae; 14.V.2011; T. M. Gosliner leg.; CASIZ 187288 (straight juvenile specimen; faint purple longitudinal stripes of similar width throughout dorsum; notopodia anterior with dark purple inverted triangular spot, lacking blackish ring; notochaetae dense, longer than dorsal cirri; red anterior eyes not protruded from margin; caruncle median ridge complete, smooth, 3-4 lateral lobes with specks of pigment, minimal venation; median antenna broken; branchiae pigmented basally; pygidium bulbous, bent ventrally, yellow in life; 6 mm long, 2 mm wide, 21 chaetigers) • 1 specimen; Tingloy, Maricaban Island, Sepok Point dive site; Hearst Philippine Biodiversity Expedition 2011; Sta. HEP-115; 13°41’16”N, 120°49’37”E; 0-26 m depth; 25.V.2011; C. Piotrowski leg.; CASIZ 187536 (posterior segments twisted, damaged; longitudinal stripes of similar width throughout dorsum, faint ventral longitudinal stripes; notopodia pigmented anteriorly only, lacking complete dark ring, dorsal cirri nearly as long as notochaetae; anterior eyes not protruded from prostomial margin; caruncle median ridge with pigmented transverse furrows, 5-6 lateral lobes with dark venation; median antenna 2-3 as long as caruncle, at least twice length of lateral antennae; branchiae pigmented basally; anus dorsal, pygidium elongate, bulbous; 70 mm long, 5 mm wide, 58 chaetigers) • 1 specimen; Verde Island, Coral Garden dive site; 2015 Verde Island Passage Expedition; Sta. VER-21; 13°33’37”N, 121°2’38”E; 6-15 m depth; coral rubble; 9.IV.2015; G. Paulay leg.; CASIZ 214577 (specimen twisted in S curve, internal organs prolapsed through anterior body wall from tissue sampling; longitudinal stripes similar width throughout dorsum, faint longitudinal stripes ventrally; notopodia pigmented anteriorly, lacking complete dark ring, notochaetae longer than dorsal cirri; eyes not protruded from prostomial margin; caruncle median ridge with pigmented transverse furrows, 5-6 lateral lobes with dark venation; median antenna 1/3 length caruncle, 3× length lateral antennae; branchiae pigmented basally; pygidium with terminal plate nearly twice longer than wide; 45 mm long, 6 mm wide, 51 chaetigers) .

Japan • 1 specimen; Okinawa Island, White Beach, Navy Pier ; 26°17’44”N, 127°54’22”E; semi-exposed, shaded artificial substrate; 0-1 m depth; 3.VII.2010; N. Evans, F. Michonneau, T. Schils & A. Devilliers leg.; UF 1723 (complete, twisted; dorsal longitudinal stripes of similar width, a few branched, integument maculate; notopodia without black distal rings; anterior end smashed, eyes of similar size, collapsed; caruncle with median ridge complete, 7-8 pairs of lateral lobes, barely pinnate; median antenna broken; pygidium with anus terminal, anal plate truncate, margin entire; body 101 mm long, 11 mm wide, 66 chaetigers) .

Mariana Islands • 1 specimen; Guam , West Cocos Barrier Reef; 13°14’00”N, 144°38’28”E; 10-15 m depth; fore reef, rubble; 19.V.1997; J. Starmer & G. Paulay leg.; UF 19 (regenerating posterior end; dorsal longitudinal stripes of different width, about four wider dorsal and three on each side; integument without maculae; notopodia without distal blackish rings; anterior eyes two times as large as posterior ones, not protruded from prostomial margins; caruncle pale, with median ridge complete, with 8-9 pairs of lateral smooth lobes; median antenna without tip, 2/3 as long as caruncle; pygidium with anus terminal, anal plate truncate; 135 mm long, 12 mm wide, 75 chaetigers) • 1 specimen; Guam , Pago Bay; 13°25’11”N, 144°47’36”E; 7 m depth; fore reef, under rocks; 20.II.1996; G. Paulay leg.; UF 49 (complete; dorsal longitudinal stripes of similar width, some branching; integument without maculae; notopodia without distal blackish rings; anterior eyes two times as large as posterior ones, not protruded from prostomial margins; caruncle pale, with median ridge complete, with 10 pairs of lateral pinnate lobes; median antenna entire, half as long as caruncle; pygidium with anus terminal, prolapsed, anal plate truncate; 64 mm long, 10 mm wide, 60 chaetigers) • 1 specimen; Guam , Orote Peninsula; 13°26’19”N, 144°37’39”E; 20-25 m depth; under rocks; 6.II.1998; G. Paulay leg.; UF 173 (complete; dorsal longitudinal stripes of similar width, some branching; integument without maculae; notopodia without distal rings; anterior eyes two times as large as posterior ones, not protruded from prostomial margins; caruncle pale, with median ridge complete, with seven pairs of lateral smooth lobes; median antenna without tip, 1/3 as long as caruncle; pygidium with anus terminal, anal plate notched; 58 mm long, 10 mm wide, 61 chaetigers) • 1 specimen; Asuncion Island , NE of island; 19°41’59”N, 145°25’0”E; 10-16 m depth; 5.IX.2003; M. Malay leg.; UF 2876 (pharynx partially exposed; without pygdium; dorsal longitudinal stripes of similar width, a few branching; integument without maculae; notopodia without distal rings; anterior eyes two times as large as posterior ones, not protruded from prostomial margins; caruncle pale, with median ridge complete, with 6-7 pairs of lateral smooth lobes; median antenna without tip, half as long as caruncle; 33 mm long, 10 mm wide, 51 chaetigers) • 1 specimen; Maug Islands ; 20°02’N, 145°13’E; 12 m depth; reef slope, in holes along crevices; 2.VI.1992; P. Schupp leg.; UF 163 (complete; dorsal longitudinal stripes of similar width, a few branching; integument without maculae; notopodia without distal blackish rings; anterior eyes slightly larger than posterior ones, not protruded from prostomial margins; caruncle pale, with median ridge complete, with eight pairs of lateral smooth lobes; median antenna broken, 1/5 as long as caruncle; pygidium with anus terminal, anal plate truncate, margin entire; 44 mm long, 7 mm wide, 47 chaetigers) .

Line Islands • 1 specimen; Kiritimati Atoll , West side of atoll, north of passage, outer reef slope; 2°0’30”N, 157°29’22”E; 10-14 m depth; from dead Pocillopora head; 4.VIII.2005; G. Paulay & N. Knowlton leg.; UF 572 (without posterior region, anterior region with gut exposed in chaetiger 9; dorsal longitudinal stripes of similar width, rarely branching; notopodia with an anterior darker area, without distal black rings; caruncle with median ridge complete, five pairs of lateral lobes with venation, not pinnate, distal pair smooth; median antenna without tip, half as long as caruncle; 22 mm long, 8 mm wide, 31 chaetigers) • 1 specimen; Palmyra Atoll, southern outer reef at North-South Causeway intersection; CAS Palmyra Expedition 2006; Sta. CP-18; 5°52.069’N, 162°4.156’W; 9-18 m depth; 11.IX.2006; M. Roth & T. Konotchick leg.; CASIZ 174407 (complete, twisted, small specimen; dorsal longitudinal stripes of similar width; notopodia dark basally, distally unpigmented, lacking distal blackish rings; anterior eyes slightly protruded from prostomial margin; caruncle median ridge complete, smooth, 5-6 lateral lobes with purple venation; median antenna 2/3 as long as caruncle; branchiae basally dark, distally unpigmented; pygidium with anus terminal, ringed in purple distally; 17-20 mm long, 3 mm wide, 35 chaetigers) .

Hawaii • 1 specimen; Hawaii Island, Kona coast, in front of Hotel Kona Surf ; 3-10 m depth; VI.1973; A. J. Ferreira leg.; CASIZ 4899 (complete; regular longitudinal stripes distinct; anterior surfaces of notopodia darker than adjacent areas, without black ring; body 29 mm long, 4 mm wide, 45 chaetigers) • 4 specimens; Oahu Island, Honakalu Harbor, near Kailua ; 7-8 m depth; no date; J. Bailey leg.; LACM-AHF 14775 (complete; regular longitudinal stripes distinct, anterior surfaces of notopodia darker than adjacent areas, without black rings; body 30-57 mm long, 5-10 mm wide, 47- 54 chaetigers) • 1 specimen; Oahu Island, off Kewalo ; 21°17’20”N, 157°51’54”W; 9-12 m depth; outer reef slope, settling plates, ARMS 1; 1.XII.2008; R. Moffitt leg.; UF 822 (without pygidium, bent laterally; dorsal longitudinal stripes of similar width, a few branching; notopodia without distal rings; anterior eyes two times as large as posterior ones, not protruded from prostomial margins; caruncle brownish, median ridge complete, with seven pairs of lateral lobes; median antenna half as long as caruncle; 34 mm long, 6 mm wide, 41 chaetigers) • 1 specimen; Oahu Island, Kaneohe Bay , N of Kekepa Island; 21°28’37”N, 157°46’33”W; 15-18 m; 26.V.2017; IZ team leg.; UF 5438 (anterior fragment, markedly contracted, bent ventrally; dorsal longitudinal stripes of similar width, some branching; notopodia without distal rings; anterior eyes larger than posterior ones; caruncle whitish with median ridge distinct, and seven pairs of lateral lobes, mostly smooth; median antenna without tip, 1/3 as long as caruncle; 21 mm long, 8 mm wide, 25 chaetigers) • 1 specimen; Oahu Island, Kaneohe Bay , N of Kapapa Island; 21°29’23”N, 157°47’20”W; 16-19 m depth; 27.V.2017; IZ team leg.; UF 5461 (without pygidium, twisted; dorsal longitudinal stripes of similar width, some branching; notopodia without distal rings; abundant white spots along notopodial bases, continued ventrally, abundant ventrally; anterior eyes slightly larger than posterior ones; caruncle brownish, maculate, with median ridge distinct, and seven pairs of lateral pinnate lobes; median antenna without tip, 4/5 as long as caruncle; 121 mm long, 12 mm wide, 70 chaetigers) • 1 specimen; Oahu Island, Kaneohe Bay , outside main reef pass; 21°28’37”N, 157°46’15”W; 14-17 m depth; 28.V.2017; IZ team leg.; UF 5499 (complete, twisted, some anterior left parapodia removed for molecular studies; dorsal longitudinal stripes of similar width, a few branching; notopodia without distal rings; anterior eyes two times as large as posterior ones, not protruded from prostomial margins; caruncle pinkish with median ridge distinct, 7-8 pairs of lateral lobes, mostly smooth; median antenna almost complete, 1/2 as long as caruncle; pygidum terminal, anal plate truncate; 70 mm long, 6 mm wide, 62 chaetigers) .

Papua New Guinea • 1 specimen; Louisiades Archipelago, Misima Island , 1.6 km North of Point Ebola; 10°36’43”N, 152°32’23”E); north coast; 31.V.1998; G. Paulay leg.; UF 31 (complete; dorsal longitudinal stripes of similar width; eyes of similar size, not protruded, caruncle with median ridge complete, with six pairs of lateral lobes, smooth; median antenna 1/3 as long as caruncle; notopodia without distal black rings; pygidium with anus terminal, anal plate truncate cleft; 34 mm long, 9 mm wide, 54 chaetigers) • 1 specimen; East New Britain, Ralum ; dead coral; 4.X.1896; H. Dahl leg.; ZMH V10312 (complete, regenerating posterior end; anterior end compressed; longitudinal stripes of different width, reaching intersegmental spaces, well-defined dorsally and ventrally, distinct laterally; notopodia with black rings; caruncle asymmetric, with 11 left lobes and six right lobes; median antenna without tip, surpassing anterior caruncle half; right parapodium of chaetiger 25 removed for observation (kept in container); dorsal cirrophore 2-3 times wider and about three times longer than cirrostyle; pygidium with anus terminal, anal plate small, truncate; body 100 mm long, 18 mm wide, 58 chaetigers) .

Australia • 1 specimen; Harry’s Canyon, Heron Island , Queensland; 15 m depth; 18.XI.2009; Creefs Heron 2009 team leg.; MAGNT 28828 (twisted, without posterior end, broken in two pieces, dorsal longitudinal stripes of similar width along body; eyes nor protruded from prostomial margin; fragment 15 + 36 mm long, 11 mm wide, 17 + 35 chaetigers) • 1 specimen; fore reef north of Tantabiddi , Ningaloo Reef, WA; 23 m depth; 14.VI.2008; G. M. Dally leg.; MAGNT 29300 (juvenile, bent laterally; longitudinal stripes of similar width along body; notopodia with black band anteriorly; branchiae blackish with tips pale; caruncle with 6 lateral foliose lobes, all with blackish venation; median antenna bent posteriorly; 1/4 as long as caruncle, as long as palps; lateral antenna slightly shorter than palps; eyes not protruded from prostomial margin, anterior eyes almost 2× as large as posterior ones; pygidium with anus terminal, anal plate slightly notched; body 11.5 mm long, 5 mm wide, 32 chaetigers) .

New Caledonia region: Matthew Island • 1 specimen; Northwest side; 22°20’29”S, 171°21’1”E; 10 m depth; 1.VIII.2017; S. Hannam, A. Reid, C. Bedford & I. Middleton leg.; AM 51660 (body twisted, broken in two fragments; longitudinal anastomosing stripes blackish dorsally and ventrally, paler ventrally, present throughout body; caruncle with pale ridges, darker blades, with eight lateral lobes foliose, rugose; median antenna 1/3 as long as caruncle; a few noto- and neurochaetae removed from chaetigers 9 and 25; harpoon notochaetae present, with more denticles in median chaetigers; neurochaetae with denticles better defined in anterior chaetigers, some with sharp tips and over 20 subdistal denticles; anus dorsal between last two chaetigers, anal plate round, blunt; body 23 + 44 mm long, 11 mm wide, 21 + 40 chaetigers) • 1 specimen; Northwest side; 22°20’29”S, 171°21’1”E; 10 m depth; 1.VIII.2017; S. Hannam, A. Reid, C. Bedford & I. Middleton leg.; AM 51662 (anterior fragment, bent dorsally; longitudinal anastomosing stripes blackish dorsally and ventrally, paler ventrally, present throughout fragment; caruncle with thin blackish anastomosing stripes, 1/2 as wide as anterior end; with eight lateral foliose lobes; median antenna 1/4 as long as caruncle; chaetae no removed; fragment 19 mm long, 8 mm wide, 27 chaetigers) .

Samoa • 1 specimen; Upolu ; from Museum Godeffroy; no further data; ZMH V694 (colorless; caruncle asymmetrical with four left and six right lobes, median ridge base smooth; median antenna lost; pygidium with anus terminal, anal plate barely bifid; body 40 mm long, 7 mm wide, 46 chaetigers) .

Cook Islands • 1 specimen; Aitutaki , Rapae Canyon; 18°50.868’S, 159°48.041’W; 20 m depth; coral rubble; 30.VI.2001; R. Wetzer & N. D. Pentcheff leg.; LACM-AHF 14777 (complete; regular longitudinal stripes, anterior surface of notopodial lobes darker, without black ring; body 61 mm long, 10 mm wide, 59 chaetigers) .

French Polynesia • 1 specimen; Austral Islands, Rurutu , near the village; 22°28’48”S, 151°21’17”W; 5 m depth; sandy bottom next to reef, hard bottom outside lagoon; 10.VIII.1991; J. K. Lowry & J. M. Poupin leg.; AM 35512 (body twisted, without posterior end; longitudinal stripes visible, some anastomosing, better retained along a few anterior segments; caruncle pale, 1/6-1/5 as wide as anterior end, with 8 lateral foliose lobes; a few noto- and neurochaetae removed from chaetigers 9 and 25; harpoon notochaetae present; neurochaetae with tip blunt, slightly falcate, emerging from a short basal hood; margin denticulate with about 10 subdistal denticles; body 135 mm long, 17 mm wide, 83 chaetigers) • 1 specimen; Society Islands; Moorea, Vai’are Pass, north side; Sta. BIZ 713; 17°31.338’S, 149°45.7308’W; 20 m depth; 10.XII.2010; J. Moore leg.; MAGNT 29885 (juvenile, twisted; longitudinal stripes of similar width along body; notopodia without black ring; branchiae blackish with tips pale; caruncle with five lateral foliose lobes, all with brownish venation; median antenna bent laterally, 1/4 as long as caruncle, slightly longer than palps and lateral antennae; eyes not protruded from prostomial margin, anterior eyes almost 2 × as large as posterior ones; pygidium with anus terminal, anal plate slightly notched; body 12 mm long, 3 mm wide, 32 chaetigers) • 1 specimen; Society Islands, Moorea; barrier reef between Cook’s and Opunohu Bays ; 17°28’37”N, 149°49’50”W; 0-2 m depth; fore reef, under rocks; VII.2006, K. McKeon & G. Paulay leg.; UF 523 (complete, some parapodia removed for molecular studies; dorsal longitudinal stripes of similar width, some branching; notopodia with anterior surfaces dark, without distal rings; anterior eyes two times as large as posterior ones, not protruded; caruncle with median ridge complete, with 6-7 pairs of lateral smooth lobes, with venation; median antenna 1/2 as long as caruncle; pygidium with anus terminal, anal plate truncate, margin entire; 77 mm long, 8 mm wide, 60 chaetigers) • 1 specimen; Society Islands, Moorea; Price’s settling plates, CTD1; 17°28’28”S, 149°48’46”W; ex Nicole, no depth or date data; N. Price leg.; UF 545 (without posterior end, removed for molecular studies; dorsal longitudinal stripes of similar width, a few branching; notopodia with anterior surfaces darker, without distal rings; anterior eyes three times as large as posterior ones, not protruded; caruncle with median ridge complete, with 6-7 pairs of lateral crenulate lobes, with venation; median antenna erect, as long as caruncle; 48 mm long, 4 mm wide, 46 chaetigers) • 1 specimen; Society Islands, Moorea; lagoon pinnacle, south of Afareaitu ; 17°33’50”S, 149°47’21”W; 15-18 m depth; silty patch reef with overhangs; 12.XI.2009; S. McPherson leg.; UF 1330 (juvenile, complete; without longitudinal stripes; anterior stomach greenish; notopodia with anterior surfaces black, without distal rings; anterior and posterior eyes of similar size, not protruded; caruncle with median ridge complete, with two pairs of lateral smooth lobes, without venation; lateral antennae and palps biarticulate, ceratophores and ceratostyles, and palpophores and palpostyles of similar size; median antenna directed forward, 1/2 as long as caruncle; branchiae single filaments along chaetigers 1-2, other segments without branchiae; pygidium with anus terminal, anal plate truncate, notched; 9.4 mm long, 0.8 mm wide, 9 chaetigers, last segment achaetous) • 1 specimen; Society Islands, Moorea; in front of Hilton ; 17°28’37”S, 149°50’41”W; 13-24 m depth; fore reef, rubble; 8.XII.2009; K. White, W. El- Tourky & J. Thomas leg.; UF 1434 (juvenile, complete, some parapodia removed for molecular studies; dorsal longitudinal stripes of similar width, not branching; notopodia with anterior surfaces darker, without distal rings; anterior and posterior eyes of similar size, not protruded; caruncle with median ridge complete, with 4-5 pairs of lateral smooth lobes, with venation; median antenna 1/4 as long as caruncle; pygidium with anus terminal, anal plate truncate, notched; 13 mm long, 3 mm wide, 27 chaetigers) • 1 specimen; Society Islands, Moorea, off NW Motus ; 17°29’2”S, 149°54’59”W; 19-20 m depth; outer reef, under rocks; 3.XI.2009; G. Paulay, S. McKeon & T. Lotufo leg.; UF 1269 (small juvenile, cut in two fragments, a few segments removed for molecular studies; dorsal longitudinal stripes of similar width, none branched; integument not maculate; notopodia with anterior surfaces blackish, without distal rings; anterior eyes slightly larger than posterior ones, not protruded; caruncle with median ridge complete, 3-4 pairs of smooth lobes, with minimal venation; branchiae few, single filaments throughout body; pygidum fleshy, tubular, anus terminal, anal plate barely developed; 3 + 2 mm long, 2 mm wide, 9 + 7 chaetigers) • 1 specimen; Society Islands, Moorea; between Opunohu Bay and Motus ; 17°28’56”S, 149°53’27”W; 18-22 m depth; outer reef, rubble; 3.XI.2010; J. Moore leg.; UF 2125 (without posterior region; bent in three; dorsal longitudinal stripes of similar width, a few branched; caruncle, dorsum, interramal belt and venter maculate; notopodia without distal black rings; anterior eyes three times as large as posterior ones, not protruded; caruncle with median ridge complete, 6-7 pairs of smooth lobes; 27 mm long, 7 mm wide, 37 chaetigers) • 1 specimen; Society Islands, Moorea; between Opunohu Bay and Motus ; 17°28’56”S, 149°53’27”W; 18-22 m; outer reef, rubble; 3.XI.2010; J. Moore leg.; UF 2312 (without posterior region; soft, colorless, dorsum and venter maculate; notopodia without distal black rings; prostomium broken, eyes indistinct; caruncle distorted, median ridge indistinct, four pairs of smooth lobes; 64 mm long, 7 mm wide, 58 chaetigers) • 1 specimen; Marquesas Islands; Nuku Hiva, Les 4 Grottes ; 8°56’13”S, 140°7’14”W; 20-23 m depth; I.2012; J. Starmer leg.; UF 3804 (markedly contracted, anterior and posterior fragments; dorsal longitudinal stripes of similar width; notopodia with darker anterior surfaces, without distal black rings; anterior eyes barely visible, posterior eyes hidden after body contraction; caruncle with median ridge complete, 6-7 pairs of lateral smooth lobes, with venation; pygidium with anus terminal, anal plate truncate; 37 + 33 mm long, 10 mm wide, 36 + 30 chaetigers) .

Pacific Panama • 1 specimen; Gulf of Chiriqui, Coiba National Park; off Islas Uva ; 7°48’57”N, 81°45’36”E; coral reef; 13.III.2006; P. W. Glynn leg.; LACM-AHF 14778 (complete, regenerating posterior body half; regular longitudinal stripes, anterior surface of notopodia darker, without black ring; body 43 mm long, 8 mm wide, 48 chaetigers) • 4 specimens; Taboga; S. Y. St. George Expedition , 1923-1924; from colony of Pocillopora ; C. Crossland leg.; NHML 1932.12.24.271-275 (none complete, two with posterior end in regeneration; longitudinal stripes better retained in largest specimen; all with notopodia with black rings; two with duplicate parapodia between chaetigers 10/11; median antenna 2/3-4/5 as long as caruncle; caruncle with 7-8 pairs of lateral lobes; body 40- 53 mm long, 5-10 mm wide, 39-42 chaetigers) .

DISTRIBUTION. — East Africa to West America, in shallow water, coral or rocky bottoms.

DESCRIPTION OF TYPE SPECIMENS

Holotype of Hermodice striata (SMNH 6080) consisting of an anterior fragment and two dissected chaetigers, both compressed by labels and partially dehydrated; anterior fragment 8 mm long, 5.5 mm wide, six chaetigers ( Fig. 4A). Anterior end damaged, median antenna without tip, lateral antennae complete, left palp complete, right palp broken; caruncle eroded or removed ( Fig. 4B); first two chaetigers with notochaetae longer than dorsal cirri. Dissected segments depressed, one notopodium separate from remains, with dorsal cirrophore about as long as cirrostyle, notochaetae abundant, longer than dorsal cirrus, mostly capillaries and shorter harpoon-chaetae ( Fig. 4C, inset); branchiae with approximately 10 filaments. Following details after original illustration: Dorsum with homogeneous longitudinal stripes ( Fig. 4D). Median antenna slightly longer than laterals or palps, 1/3 as long as caruncle; caruncle with five pairs of lateral lobes ( Fig. 4E). Eyes black, of similar size ( Fig. 4F). Mouth between chaetigers 2 and 3 ( Fig. 4G). Dorsal cirri longer than ventral cirri ( Fig. 4H), branchiae arborescent, shorter than dorsal cirri. Notochaetae including harpoon chaetae ( Fig. 4I); neurochaetae including smooth capillaries ( Fig. 4J), and acicular neurochaetae, tips entire, slightly bent ( Fig. 4K).

Holotype of Pherecardia lobata (RMNH VER.1295) complete, anterior end damaged from compression in container, dissections, and pinning for illustrations; posterior region bent ventrally, last six chaetigers partially detached. Dorsum brownish, longitudinal stripes faded, with regular, longitudinal thin furrows along each segment ( Fig. 5A); venter brownish with a longitudinal midventral blackish thin line. Dorsal and ventral cirri and branchiae markedly contracted. Many notochaetae broken. Body 87 mm long, 9 mm wide, 66 chaetigers.

Prostomium bent dorsally, ovoid, approximately as long as wide, anterior margin and longitudinal lip broken, mouth ventrally restricted to first chaetiger. Lateral antennae short, right one without tip, palps lost ( Fig. 5B). Median antenna with ceratophore partially surrounded by base of caruncle median ridge, ceratostyle lost. Eyes black, of similar size, not protruded from prostomial margins. Mouth opens between chaetigers 2 to 4.

Caruncle pale, reaching chaetiger 3, 1/4 as wide as anterior end; median ridge transversely sliced previously, cut extended throughout caruncle length, with 6-7 lateral foliose lobes, some cut off, blades pinnate.

Parapodia not dissected to avoid further damage, all segments with arborescent branchiae with many filaments as long as dorsal cirrophores. Notopodia cylindrical with dorsal cirrophores twice as wide and 2/3 as long as cirrostyles; notochaetae surrounding dorsal cirrophores, abundant, soft. Neuropodial lobes subcylindrical, neurochaetae arise from middle. Ventral cirri with cirrophore short, subcylindrical, as long as wide, cirrostyle 3-4 times longer than cirrophores. Neurochaetae with distinct distal tooth, subdistal denticles eroded, barely visible in smaller neurochaetae ( Fig. 5C).

Posterior region tapered; anus terminal, broken; anal plate indistinct.

Holotype of Hermodice pennata Treadwell, 1906 (USNM 5209) including two fragments, without pigmentation pattern; anterior fragment 11 mm long, 6 mm wide, 15 chaetigers, and median fragment 9 mm long, 9 mm wide, 10 chaetigers.

VARIATION

Other specimens of P.striata exhibit interesting modifications in ocular and caruncle development. Eyes are protruded from the prostomial surface in most specimens, but not in all of them and this is not a size-related modification ( Fig. 6 A-D). The caruncle may be modified due to the loss of the median ridge such that the lateral lobes become distorted, and symmetry is difficult to assess ( Fig. 6E, F), yet pinnate development remains visible.

The caruncle lobes are smooth with venation barely protruded [ 28 mm specimen, Fig. 6A, B], but caruncle lobes are rugose due to venation from transverse low projections on each lobe, resulting in a pinnate appearance, especially in larger specimens ( Fig. 6C, D).

The number of longitudinal stripes decreases with size in preserved specimens, being more abundant and thinner in smaller specimens ( Fig. 7A). They remain of similar width and length in each segment in larger specimens ( Fig. 7D), although the branching of each stripe tends to be reduced in larger individuals. The prostomium is difficult to observe due to contraction of the anterior end, especially in the smaller specimen. The caruncle has a distinct median ridge with an expanded, roughly heart-shaped anterior region, but the lateral foliose lobes become more developed with maturity; small specimens have fewer and almost smooth lobes ( Fig. 7C), whereas larger specimens have more lobes and each is pinnate ( Fig. 7F). Along the body, there is a lateral longitudinal rugose belt, usually paler than in adjacent parapodial areas ( Fig. 7B, E), which becomes mottled in larger specimens. Branchiae are abundant with reddish filaments and pale tips; in larger specimens the paler area is relatively smaller than observed in small specimens. Notopodia are protruded muscular cylinders, slightly darker in their anterior aspects than in adjacent areas ( Fig. 7G), and notochaetae are very abundant and surround the dorsal cirri, with dorsal cirri having cirrophores about as long as, or up to twice longer than cirrostyles; whereas the neuropodia are lobate with chaetae emerging above the ventral cirri ( Fig. 7G). Staining with Methyl green pigment failed to reveal a glandular role for the maculate interramal belt ( Fig. 7H). A cross section of this area reveals a thick furrowed integument and melanophores concentrated mostly in furrows, with the paler spots corresponding to non-acidic glands ( Fig. 7I). Neurochaetae include denticulate capillaries and aciculars with a small distal tooth, and often a series of subdistal denticles which are better defined in upper than in lower chaetae of the same chaetiger ( Fig. 7J).

REMARKS

Hartman (1948: 46) indicated the holotype of Hermodice striata Kinberg, 1857 was an anterior fragment 8 mm long with seven segments, with branchiae from chaetiger 1, each with five filaments, and mouth in chaetiger 3. We observed some minor differences between her description and the type, but its condition is rather precarious, such that manipulation was kept as minimal as possible.

De Quatrefages (1866) introduced Amphinome formosa for a specimen collected in Hawaii (Sandwich Islands) that was 65 mm long, 7 mm wide, and 90 segments. The caruncle was described as ‘very developed, reaching segment 4. Its shape is oval, with 7-8 foliose lobes fused medially by a ridge with a free margin.’ These features match Pherecardia ; however, because there is no type specimen the status of the species cannot be determined (see below).

Horst (1886) proposed Pherecardia , with P. lobata as its type species, based on a damaged specimen with most of its pigmentation faded away, but he noted that it was ‘marked on its dorsal side with faint longitudinal folds’ which may have corresponded with longitudinal stripes. Details of the prostomium were not provided because it was damaged.

Fischli (1903) described Amphinome sericata in detail with several illustrations, and although prostomial details were not given, the color plates portray the dorsum with rather regular longitudinal stripes of similar width, and this explains why Horst (1911) may have synonymized it with P. striata . Fischli lived in Linthal, Switzerland, but whether his type specimens are extent and if so where they are located, could not be clarified. Inquiries to nearby natural history museums about potential types were not answered.

Treadwell (1906) proposed Hermodice pennata based on a 15-segment anterior fragment collected in Hawaii. He indicated that pigmentation included numerous longitudinal stripes extending across segments, and that the eyes were ‘prominent on dorsal view’, and his illustration shows them not protruded from the prostomial margin. The caruncle was described and illustrated as having seven smooth lobes per side. The holotype is in two fragments. The caruncle is as illustrated and while the pigmentation is now gone it is indicated in the original description. Based on examination of the type and Treadwell’s description we confirm that this species is a synonym of P. striata , as proposed by Augener (1927).

Malaquin & Dehorne (1907) proposed Eucarunculata with E. grubei as its type species without citing any of the previous literature on the above species. They indicated that anterior eyes were slightly larger than posterior ones, and that the animal had 10-15 ‘dark longitudinal stripes running from segment to segment along the whole body.’ Their illustrations show the anterior end with longitudinal stripes of similar width. Horst (1909) indicated that E. grubei was a junior synonym of his P. lobata . We did not locate the type of E. grubei . Based on the description and illustration we agree with Horst that E. grubei is a synonym of P. stricta .

Potts (1909) proposed two varieties for E. grubei based on two smaller specimens: E. g. var. gracilis and E. g. var. minuta , both with regular longitudinal stripes, but no type specimens were deposited. He noted in a footnote (page 365) that ‘the differences in form of caruncle and chaetae are probably due to varying age.’ His specimens have not been located, and it is likely that they were not deposited in a museum. The typical form in his treatment included a specimen 104 mm long, 7 mm wide, and with 73 segments; its caruncle had 6-7 lateral lobes, notochaetae included smooth capillaries and non-spurred harpoon chaetae, neurochaetae were distally falcate, either with subdistal denticles or smooth. His variety gracilis included one specimen 24 mm long, and 4 mm wide with 41 segments, with 5-6 lateral lobes in the caruncle, and the neurochaetae had ‘more distinct and numerous serrations’ than in the typical form. His variety minuta was based on a specimen 16 mm long, 2 mm wide, and with 34 segments. He noted that neurochaetae had distinct, variable serrations. Hartman (1959: 133) regarded both as junior synonyms of Pherecardia lobata . We agree with her conclusions.

Differences between Pherecardia striata and P. distincta , are mostly in pigmentation pattern, as indicated in the key above. In P. striata the longitudinal stripes are more or less regular, having similar width and length, and the notopodia have a darker pigmentation along anterior surface than on posterior one, whereas in P. distincta the longitudinal stripes are irregular, of different width and length, and the notopodia have a blackish ring. Further, these two species are separated by 22.4-23.5 % K2P distance in COI ( Fig. 8). Intraspecific variation between the two sequenced species is 0.3-2% in P. striata , and 1.2% in P. distincta . The records by Amoureux (1977: 1095) and Rajasekaran & Fernando (2012: 3) could not be confirmed.