taxonID	type	description	language	source
B77C9C33CF30DC16FF61F8B1AE637BE9.taxon	description	(Figs 12 – 15) LSID urn: lsid: zoobank. org: act: AA 1775 A 1 - 6 C 41 - 40 B 1 - AC 54 - FB 9091 A 0 CD 99. Material examined: Holotype: CEAB. AP. 1000 A, February 2020, Gulf of Fos, France, approximate location 43 ° 25 ′ N 4 ° 56 ′ E, fixed in 4 % formalin seawater solution and preserved in 70 % ethanol, one specimen entire, dissected to extract jaws, collected by Creocean team. Paratypes: MNHNC MB 29: 000453 and MB 29: 000454, October 2023, Gulf of Fos, France, approximate location 43 ° 24 ′ 54 ′′ N 4 ° 50 ′ 42 ′′ E, fixed in 4 % formalin seawater solution and preserved in 70 % ethanol, one specimen entire, dissected to extract jaws, collected by Creocean team. CEAB. AP. 1000 B, November 2019, Port la Nouvelle, France, approximate location 43 ° 0 ′ 29 ′′ N 3 ° 4 ′ 35 ′′ E, fixed in 4 % formalin seawater solution and preserved in 70 % ethanol, one specimen (anterior fragment), collected by Creocean team. Description: Based on holotype. Body 49 mm long, 6.5 mm wide, width uniform along body, with 128 chaetigers (Table 2), rounded in cross section anteriorly, tapering and dorsoventrally flattened posteriorly, lacking most posterior segments (Fig. 12 A – F). Prostomium dorsoventrally flattened, with anterior end elevated, bilobate, with a conspicuous median sulcus (Fig. 12 A, D). One median and two lateral antennae, thin, distally tapering, folding back to middle of chaetiger 2; two palps, similar in shape to antennae, folding back until beginning of chaetiger 2 (Fig. 12 A – D). One pair of dark-brown reniform eyes, placed laterally to ceratophores of lateral antennae (Fig. 12 D). Calcareous cutting plates, 36 µm long and 67 µm wide, 1 / 4 shorter than sclerotized matrix, overall thick, with thin translucent borders, broadly B-shaped (Fig. 13 A, C); sclerotized matrix c. six times shorter than mandible carriers, D-shaped, distally straight, with smooth upper margin and growth ring-like marks seen though translucent calcareous cutting plates; left carrier with swollen distal end (Fig. 13 A, B). MxI c. three times as long as carrier, brown, with white, translucent tips; MxII c. four-fifths of MxI; MxIII arched, with anteriormost teeth more lateral than posteriormost ones, partly ventral to MxII, brownish, with teeth border whitish; attachment lamella of MxIII very small, straight, elongated, slightly sclerotized; left MxIV as wide as long, D-shaped; right MxIV four times longer than wider, arched; both Mx IV with concave side and posterior part of convex side brownish, anterior part of convex side and teeth border whitish; attachment lamellae of MxIV roughly C-shaped, ventral to Mx, right one with lower arm (pointed) c. three times shorter than right arm (rounded), left one with both arms rounded, similar in length; MxV roughly square, whitish; maxillary formula: I = 1 + 1, II = 4 + 4, III = 7 + 0, IV = 4 + 6, V = 1 + 1, VI absent (Fig. 13 D – F). Notopodial cirri triangular, tapering; longer (anterior), shorter (median and posterior), and longer (posteriormost) than chaetal lobes (Fig. 14 A – C). Branchial filaments starting at chaetiger 19, with a maximum of three filaments from chaetigers 50 – 73; longest filaments 4.8 times longer than notopodial cirri and 5.9 times longer than branchial stem (Fig. 14 B, C). Ventral cirri triangular, with rounded tips, with inflated bases from chaetiger 6 to posterior body end, as long as notopodial cirri in anteriormost chaetigers, one-third at midbody (Fig. 13 A – C). Notopodial aciculae inconspicuous. Neuropodial aciculae, brownish, with golden tips, up to three per parapodia, with blunt tips non-protruding from acicular lobe (Fig. 14 A – C). Chaetae in two distinct bundles: supracicular with limbates and pectinates at anterior edge, and subacicular with compound spinigers and one golden subacicular hook, with guards covering tip, bidentate, with short, rounded, upwards directed teeth separated by a narrow angle (Fig. 15 D, E), starting at chaetiger 40, present on all following chaetigers. Pectinate chaetae of four types, present in all chaetigers, except 1 – 4. Type 1 present from anterior parapodia, flat to little curved, isodont, slightly asymmetrical, with external teeth similar in length, with c. 25 narrow teeth about 4.3 µm long, similar in length on different chaetae, with pointed tips (Fig. 15 A). Type 2 thick, flat to little curved, isodont, slightly asymmetrical with short external teeth slightly differing in length, with 25 narrow teeth about 5 µm long, coarse, with pointed tips, slightly varying in length on different chaetae (Fig. 15 B, specimen from Port la Nouvelle). Type 3 thick, isodont, slightly asymmetrical, with external teeth of same length as internal ones, with 10 – 14 short coarse teeth with pointed tips (Fig. 15 B, specimen from Port la Nouvelle). Type 4 thick, non – curved, anodont, asymmetrical, with 7 – 11 coarse teeth with pointed tips, 18 – 25 µm long, c. eight times longer than wider (Fig. 15 C). Pygidium with four anal cirri, dorsal ones c. seven times longer than ventral. Variation: Specimens MNHNC MB 29: 000453 and MB 29: 000454 from Gulf of Fos fragmented in several parts, anterior one longest (2.4 – 5 cm long, 0.5 – 0.7 cm wide, for 72 – 83 chaetigers); branchiae starting at chaetigers 16 and 25, subacicular hooks starting at chaetigers 37 and 42; nondissected, jaws not seen. Specimen from Port La Nouvelle ripe female with intra-coelomic oocytes (195 – 210 µm in diameter, Fig. 14 F), body c. 40 mm long, 7.5 mm wide, with 67 segments, lacking middle and posterior segments (Table 2). Antennae folding back to middle of chaetiger 4 (Fig. 12 G, I; Table 2). Maxillary formula with MxIII 6 + 0 and MxIII 3 + 6 (Fig. 13 G; Table 2). Branchiae starting at chaetiger 24, with 2 (1) filaments to chaetiger 44, then three (Fig. 13 E) to chaetigers 67 (left side) or 64 (right side) (Fig. 13 H, 14 E); last three right-side segments with four filaments, longest filament 5.7 times longer than notopodial cirri (Fig. 14 F; Table 2). Aciculae 3 – 4 up to chaetiger 40, then 2 – 3 till last available body segment. Subacicular hooks starting from chaetiger 40 – 45 (Table 2), present to last available body segment, bidentate, with two guards, sometimes one broken (Fig. 15 F). Remarks: The specimens from southern France belong to the species of the ‘ sanguinea ’ group having bidentate subacicular hooks (Quatrefages 1866, Hansen 1882, Webster 1884, Gravier 1907, Treadwell 1917, Hutchings and Karageorgopoulos 2003, Liu, Hutchings and Sun 2017, Kara et al., 2020, Lavesque et al., 2020, Molina-Acevedo and Idris 2020, Abdulla et al., 2024). They differ from Marphysa birgeri Molina-Acevedo and Idris, 2020, M. haemasona Quatrefages, 1866, and M. multipectinata Liu et al., 2017, which have bidentate subacicular hooks from chaetigers 25 / 27, 60, and 20, respectively (40 / 45 in M. gili sp. nov.). Marphysa birgeri, which was described from the Adriatic Sea, resembles M. gili sp. nov. in having the teeth of bidentate subacicular hooks with blunt tips and upwards directed, but differs in having longer teeth (distal smaller than proximal) separated by a wide angle and occasionally bearing a second, unidentate subacicular hook (both teeth very short, separated by a narrow angle and no additional unidentate subacicular hook in M. gili sp. nov.). Marphysa tribranchiata Liu et al., 2017 and M. schmardai Gravier, 1907 have up to three branchial filaments (up to six in M. gili sp. nov.). Marphysa brasiliensis (Hansen, 1882) and M. mullawa Hutchings and Karageorgopolous, 2003 have branchiae from chaetigers 28 – 33, M. acicularum Webster, 1884 from chaetigers 27 – 35, and M. victori from 29 to 33 (19 – 24 in M. gili sp. nov.). Marphysa victori also has three types of pectinate chaetae and M. merchangensis Abdullah et al., 2024 has five types (four in M. gili sp. nov.). Marphysa viridis Treadwell, 1917 has one type of isodont pectinate chaetae (four in M. gili sp. nov.) with fewer teeth in middle body chaetigers (14 vs. 25 in M. gili sp. nov.). Marphysa gili sp. nov. also resembles M. gaditana and, particularly, M. sanguinea (Table 2). However, M. gaditana may have two subacicular hooks, the second one being either unidentate (Cádiz) or bidentate (Tunis), with the bidentate ones with triangular and laterally directed teeth (only one hook, with teeth short, round, upwards directed in M. gili sp. nov.) and their Type 4 pectinate chaetae have very long teeth tapering to very long filiform tips (not particularly long having pointed, non-filiform tips in M. gili sp. nov.). In turn, M. sanguinea has subacicular hooks with long, triangular, upwards directed teeth and start from chaetigers 21 – 25 (short, rounded, upwards directed teeth, starting from chaetiger 40 – 45 in M. gili sp. nov.), Type I pectinate chaetae with 14 – 15 short filiform teeth and Type 4 with 8 – 10 teeth with pointed tips eight times longer than wider (Type 1: 20 – 22, with pointed tips; Type 4: 4 – 13, with short filiform tips up to 10 times longer than wider in M. gili sp. nov.).	en	Martin, Daniel, Chaibi, Marwa, Lavesque, Nicolas, Daffe, Guillemine, Daramy, Flore, Hutchings, Pat, Jourde, Jérôme, Romano, Chiara (2025): Integrative taxonomy reveals further hidden diversity of Marphysa (Polychaeta: Eunicidae) in European Atlantic and Mediterranean waters. Zoological Journal of the Linnean Society 204 (3), DOI: 10.1093/zoolinnean/zlaf018, URL: https://doi.org/10.1093/zoolinnean/zlaf018
B77C9C33CF30DC16FF61F8B1AE637BE9.taxon	etymology	Etymology: The specific epithet ‘ gili’ is dedicated to Dr João Gil, worldwide renowned polychaetologist, indefatigable advocate of meticulous taxonomic work, and tireless editor of the World Polychaete Database. Distribution: Western Mediterranean: Bay of Fos and off Port La Nouvelle (France). Chaetiger 205 216 – 233 175 128 (67) 138 – 270 number Body length 122 89 – 105 60 49 (40, incomplete) 48.4 – 163.1 (mm) Body width 6.5 7 – 8 5.2 6.5 (7.5) 3.7 – 6.6 (mm) Chaetiger lenght up to 15 up to 13 13 up to 12 (18) 10 – 15 vs. width Antenae central / 2 / 3 3 / 5 1 / 1 2 (4) 1 / 2 lateral (up to chaetiger) Palps (up to 2 / 3 2 / 3 peristomium 2 peristomium chaetiger) Mx I 1 + 1, brown with 1 + 1, brown with white 1 + 1, brown 1 + 1, brown with 1 + 1 dark tips tips with white white tips tips Mx II 5 + 6 brown with 8 + 7, brown with some 4 + 4, brown 4 + 4 (4 - 4), brown 3 – 4 + 5 dark tips white tips with dark tips with white tips Mx III 5 + 0 brown with 7 + 0, brown with some 7 + 0, brown 7 + 0 (6 + 0), brown 6 – 7 + 0 dark tips white tips with some with slightly whitish white tips upper side Mx IV 3 + 5 brown with 3 + 8, brown with some 4 + 7, brown 4 + 6 (3 + 6), brown 4 + 5 – 6 dark tips white tips with whitish with whitish upper upper side side Mx V 1 + 1, brown 1 + 1, pale brownish 1 + 1, whitish 1 + 1, whitish 1 + 1 Notopodial cirri triangular, tapering; triangular, tapering; longer triangular, triangular, tapering; Elongate, triangular, longer (anterior (anterior and median), tapering; longer (anterior and digitiform in last and median), shorter (posterior) and longer (an- median), shorter chaetigers, longer shorter (pos- longer (posterior-most) terior and me- (posterior) and than chaetal lobes terior) and longer than chaetal lobes dian), shorter longer (posterior- (posterior-most) (posterior) most) than chaetal than chaetal and longer lobes lobes (posteriormost) than chaetal lobes Branchiae 20 / 25 – 195 19 / 24 – 174 / 223 22 / 24 – 126 19 – 112 (24 –?) 13 / 21 – 133 / 255 (chaetigers) Branchial filaments up to 4 up to 5 up to 3 up to 3 (4) up to 4 – 6 Maximum number 39 - 160 49 – 66 35 / 35 – 50 – 73 (65, right? from chaetiger 103 / 110 side –?) Neuropodial up to 6, golden up to 4 (5), golden brown up to 4, golden up to 3 (4), brownish, up to 4, dark brown aciculae brown brown with golden tips Table 2 (cont.) Cádiz Arcachon (NL 126) Tunis Fos (PLN) M. sanguinea Subacicular hook from chaetiger from chaetigers 44 / 72, from chaetiger from chaetiger 40 (45), from chaetiger 40 / 55, absent absent form many para- 42 / 65, light present in all para- 21 / 25, absent from from some podia (completely absent yellow, podia, dark yellow, some chaetigers, parapodia, dark in two specimens), dark bidentate, slightly bidentate, darkish basally and yellow, bidentate yellow, bidentate, with covered by with round tips translucent distally, with round tips, round tips laterally dir- two guards. upwards directed, bidentate with covered by two ected, covered by two covered by two thick short round tips guards; some- guards, in some para- guards; second hook upwards directed, times a second podia of one specimen, in some parapodia, covered by two one, unidentate, a second bidentate hook similar to main one. guards. withouth guards. covered by two guards.	en	Martin, Daniel, Chaibi, Marwa, Lavesque, Nicolas, Daffe, Guillemine, Daramy, Flore, Hutchings, Pat, Jourde, Jérôme, Romano, Chiara (2025): Integrative taxonomy reveals further hidden diversity of Marphysa (Polychaeta: Eunicidae) in European Atlantic and Mediterranean waters. Zoological Journal of the Linnean Society 204 (3), DOI: 10.1093/zoolinnean/zlaf018, URL: https://doi.org/10.1093/zoolinnean/zlaf018
B77C9C33CF34DC1CFC45FD5FAF6B7AAC.taxon	materials_examined	Material examined: Tunisia: CEAB. AP. 995 A – 995 D, 29 March 2022, Gulf of Tunis, 36.804722 ° N, 10.294444 ° E, collected by a professional fisherman in muddy sand bottoms, fixed and preserved in 95 % ethanol, four specimens; MNHNC MB 29: 000455 and MB 29: 000456, 29 March 2023, Gulf of Tunis, 36 ° 48 ′ 16.99 ′′ N, 1017 ′ 39.99 ′′ E, collected by a professional fisherman in muddy sand bottoms, fixed and preserved in 95 % ethanol, two specimens. France: CEAB. AP. 995 E – 995 F, 9 November 2022, Arcachon Bay, collected by NL in mud close to oyster reefs, fixed and preserved in 80 % ethanol, three specimens. MNHN-IA- 2017 - 2210, 22 June 2020, Oléron Island, approximate location 45 ° 54 ′ 17 ′′ N, 1 ° 11 ′ 50 ′′ W, collected by Stéphane Guenneteau and Jacques Pigeot on muddy sediments close to a rocky reef, fixed and preserved in 95 % ethanol, one specimen. Description: Based on Tunisian specimens: CEAB. AP. 995 A for all characters except mandibular apparatus, CEAB. AP. 995 B for mandibular apparatus, and CEAB. AP. 995 A – D to provide ranges. Body up to 233 cm long, up to 8 mm wide (Table 2); rounded in cross-section anteriorly, tapering and dorsoventrally flattening posteriorly. Prostomium dorsoventrally flattened, with anterior end elevated and bilobate, with a conspicuous median sulcus (Fig. 4 A, D). One median and two lateral antennae, folding back to middle of chaetigers 2 – 3; two palps, folding back until beginning of chaetigers 2 – 3 (Fig. 4 A, C, D). One pair of dark eyes, inserted laterally to ceratophores of lateral antennae (Fig. 4 A, D). Calcareous cutting plates 42 µm long and 79 µm wide, 1 / 4 longer than sclerotized matrix, overall thick, with thin translucent borders, broadly D-shaped (Fig. 5 A – C); sclerotized matrix c. 13 times shorter than mandible carriers, D – shaped, distally straight, with smooth upper margin and growth ring-like marks seen through translucent calcareous cutting plates (Fig. 5 A – C). MxI c. twice as long as carrier, brown, with white, translucent tips; MxII c. four-fifths of MxI; MxIII arched, with anteriormost teeth more lateral than posteriormost ones, partly ventral to MxII, concave side and posterior part of convex side brownish, anterior part of convex side and teeth border whitish; attachment lamella of MxIII very small, slightly sclerotized; left MxIV as wide as long, D-shaped; right MxIV four times longer than wider, arched; colour of both Mx IV as in MxIII; attachment lamellae of MxIV roughly C-shaped, ventral to MxIII, right one with lower arm (pointed) c. three times shorter than right arm (rounded), left one with both arms rounded, similar in length; MxV roughly square, pale brownish to yellowish; maxillary formula: I = 1 + 1, II = 8 + 7, III = 7 + 0, IV = 3 + 8, V = 1 + 1, VI absent (Fig. 5 D – F). Branchiae starting at chaetigers 19 – 24, with a maximum of three filaments from chaetigers 50 – 73, longest filaments 5.3 times longer than notopodial cirri and 4.1 times longer than branchial stem (Fig. 6 A – D). Notopodial cirri triangular, tapering; longer (anterior and median), shorter (posterior), and longer (posteriormost) than chaetal lobes (Fig. 6 A – D). Ventral cirri triangular, with rounded tips, with inflated bases from chaetiger 6 to posterior body end (except about last 6), about three-quarters as long as notopodial cirri in anteriormost chaetigers, two-thirds at midbody and as long as in posterior chaetigers (Fig. 6 A – D). Notopodial aciculae inconspicuous. Neuropodial aciculae, brownish, with golden tips, up to 2 – 4 (5) per parapodia, with blunt tips non-protruding from acicular lobe (Table 2; Fig. 6 A – D). Chaetae in two distinct bundles. Supracicular bundle with limbate and pectinate chaetae at anterior edge; pectinate chaetae of four types (Table 2), present from chaetiger 5 till body end; Type 1 present from anterior parapodia, thin, flat to slightly curved, slightly asymmetrical, heterodont with very long external teeth, with c. 25 teeth c. 7 – 10 µm long, slightly varying in length in different chaetae, evenly tapering to short filiform tips (Fig. 7 A, B); Type 2 thin, flat to slightly curved, heterodont with short external teeth, slightly asymmetrical, with 22 – 25 teeth c. 5 µm long, narrow, with short filiform tips slightly varying in length in different chaetae (Fig. 7 C); Type 3 thick, flat to slightly curved, isodont with short external teeth, slightly asymmetrical, with 10 – 14 teeth c. 20 µm long, coarse, with short filiform tips slightly varying in length on different chaetae (Fig. 7 D); Type 4 thick, flat, anodont, asymmetrical, with 4 – 6 teeth with short filiform tips, c. 55 – 70 µm long, coarse, c. 7 – 10 times longer than wider (Fig. 7 E, F). Subacicular bundle with compound spiniger chaetae and one (two in some parapodia) golden subacicular hook, with two guards covering tip, bidentate, with rounded teeth upwards directed, distal one c. half-size of proximal one (Fig. 7 G), starting at chaetiger 40 – 55 to body end. Two pairs of pygidial cirri, dorsal pygidial cirri c. 5 – 10 times longer than ventral ones (Fig. 4 E – G). Remarks: The Tunisian specimens of M. gaditana do not differ molecularly from those found in the Bay of Cádiz (Figs 1 – 3; Table 1). However, they are 17 – 33 longer, and have 11 – 28 more segments and 0.3 – 0.4 times longer antennae (Table 2). Their maxillary formula differs in the number of teeth at MXII, III, and IV, and the maxillae show whitish tips (dark in the Iberian specimens) (Table 2). The branchiae have one more filament, start one segment before, may end about 20 chaetigers before or 30 chaetigers after (depending on the specimens), and show the maximum number of filaments in 10 – 60 more posterior segments and over 100 less segments the Iberian specimens, while some branchial filaments are ramified in one specimen (might be the result of a malformation) (Table 2). The maximum number of aciculae is lower than in the Iberian specimens (Table 2). In some specimens, very few parapodia have subacicular hooks, while two completely lack them, and a single specimen has two subacicular hooks in some parapodia, both bidentate — although one of them could appear eroded — (Fig. 7 G). Finally, there is a slight variability in the number of teeth of some pectinate chaetae, with Type 1 having 3 – 4 fewer, Type 2 having 3 – 5 more, and Type 4 having 1 – 4 fewer (Table 2). The French specimens from the Bay of Arcachon occur in sympatry with the introduced M. victori, being initially confused with its juveniles. However, as for the Tunisian specimens, our molecular analyses reveal that they cannot be distinguished molecularly from the specimens of M. gaditana from Cádiz, while showing some morphological differences. Among them, they are smaller, with shorter antennae and palps (Fig. 8 A – E; Table 2). Maxillae I, II, III, and IV show different number of teeth and MIV and MV have some whitish colour (Fig. 9 A – E) that is absent in the specimens of Cádiz (Table 2). The aciculae are less Hidden diversity of Marphysa • 7 numerous (Fig. 10 A – E; Table 2), the subacicular hooks start in more posterior chaetigers, and the pectinate chaetae show teeth with shorter filiform tips (Figs 11 A – E; Table 2). While similarly lacking molecular differences (Figs 1 – 3), some morphological variability occurs also between the Tunisian and Arcachon specimens. The latter are smaller, with shorter antennae and palps (Fig. 8 A – E; Table 2) and have MxII and MxIV with fewer teeth (Fig. 9 A – E; Table 2). Branchiae have fewer filaments and the maximum number of filaments starts earlier (Fig. 10 A – C; Table 2). Subacicular hooks are present in all chaetigers, a second hook has not been seen, the pectinate chaetae show slight variability in number of teeth, with some types having teeth with shorter filiform tips (Figs 11 A – E; Table 2). Distribution: Atlantic Ocean: Cádiz Bay (Spain), Sado Estuary (Portugal, only sequenced), Arcachon Bay (Biscay Bay, France), Oléron Island (Biscay Bay, France, only sequenced), Cap de la Hague (English Channel, France, only sequenced), Florida and Virginia (USA, only sequenced); Western Mediterranean: Gulf of Tunis (Tunisia). Habitat: Shallow sandy mud bottoms.	en	Martin, Daniel, Chaibi, Marwa, Lavesque, Nicolas, Daffe, Guillemine, Daramy, Flore, Hutchings, Pat, Jourde, Jérôme, Romano, Chiara (2025): Integrative taxonomy reveals further hidden diversity of Marphysa (Polychaeta: Eunicidae) in European Atlantic and Mediterranean waters. Zoological Journal of the Linnean Society 204 (3), DOI: 10.1093/zoolinnean/zlaf018, URL: https://doi.org/10.1093/zoolinnean/zlaf018
