Terebella cf. lapidaria Linnaeus, 1767

Lavesque, Nicolas, Daffe, Guillemine, Londoño-Mesa, Mario H. & Hutchings, Pat, 2021, Revision of the French Terebellidae sensu stricto (Annelida, Terebelliformia), with descriptions of nine new species, Zootaxa 5038 (1), pp. 1-63 : 53-55

Terebella cf. lapidaria Linnaeus, 1767

Figures 2A ; 25

Terebella lapidaria — Linnaeus 1767: 1092; Saint-Joseph 1894: 202–205, pl. VIII, figs 225–229, pl. IX, figs 230–231; Fauvel 1927: 254–255, fig. 87, f–l.

Synonyms. Terebella constrictor Montagu, 1819 ; Amphitrite neapolitana Delle Chiaje, 1828 ; Terebella misensis Costa, 1841 ; Terebella corallina Grube, 1855 ; Terebella pectinata Grube, 1855 ; Terebella rosea Grube, 1860 ; Terebella megalonema Schmarda, 1861 ; Heterophyselia bosci Quatrefages, 1866 ; Heteroterebella sanguinea Claparède, 1869 ; Terebella sulcigera Claparède, 1869 .

Material examined. MNHN-IA-PNT 131, complete, English Channel, Plougrescant, Pointe du Chateau, 48°52’11”N 3°13’30”W, intertidal, January 2019. SMA-LR-Tere-02, Northeastern Atlantic , Bay of Biscay, La Rochelle Harbour, 46°09’28”N 1°13’16”W, intertidal, September 2019 . SMA-Arc-Eupoly-GdBC, posteriorly incomplete, Northeastern Atlantic , Bay of Biscay, Arcachon Bay, Grand Banc, 44°40’27”N 01°11’46”W, intertidal, October 2017 , mount- ed for SEM. SMA-LR-Tere-03, posteriorly incomplete, Northeastern Atlantic , Bay of Biscay, La Rochelle Harbour, 46°09’28”N 1°13’16”W, intertidal, September 2019 .

Description. Longest complete specimen 36.7 mm long and 2.1 mm wide, for about 80 segments. Transverse prostomium attached to dorsal surface of upper lip; basal part with red eyespots more concentrated laterally, continuing dorso-laterally as a single band of eyespots, with eyespots progressively becoming more separated; distal part shelf-like. Buccal tentacles numerous, long, filiform, wrinkled and deeply grooved ( Fig. 25A–C ). Peristomium forming lips, upper lip hood-like convoluted, broader than high; lower lip swollen, cushion-like, broader than high ( Fig. 25B ).

Segment I narrow, forming low ventral lobe; lateral lobes absent on SG II–IV. Three pairs of dichotomous branchiae, on SG II–IV ( Fig. 25A–D ), with wide medial gap; second pair situated more laterally than first and third pairs; with numerous short filaments, arising from long stems and branches ( Fig. 25A–D ). Dorsum of anterior chaetigers tessellated ( Fig. 25C ). Ventral shields on SG II–XIII, rectangular, well defined; mid-ventral groove from SG XIV ( Fig. 25A–B ).

Notopodia from SG IV, extending posteriorly until last few chaetigers (n>80); notopodia short, rectangular; first pair slightly shorter than following pairs ( Fig. 25A–D ). Notochaetae from anterior chaetigers narrowly-winged from mid-length, with blade at an angle of about 45° with the shaft and curled serrated tip ( Fig. 25E–F ); notochaetae of posterior chaetigers with blade at an angle of about 90° with the shaft and curled serrated tip; presence of a spur at the base of the tip ( Fig. 25F ).

Neuropodia present from SG V, as low, almost sessile ridges throughout ( Fig. 25A–B, D ). Neurochaetae as short-handled avicular uncini, arranged in double rows from SG XI to posterior body, in a face-to-face arrangement. Uncini with triangular heel; long digitiform prow, pointing downwards and prolonged by a thin curved tendon, pointed dorsal button inserted halfway between base of main fang and prow, convex base, and main fang surmount- ed by crest with four rows of secondary teeth ( Figs 2A ; 25G–H ).

Nephridial and genital papillae on SG III and SG VI–X, globular, at base of branchiae on SG III, small, rounded above neuropodia and slightly displaced posteriorly on SG VI–X.

Pygidium discoidal and crenulated.

Type locality. Mediterranean Sea ( Gil 2011).

Type material. Cannot be traced, probably never deposited.

Distribution. Bay of Biscay, English Channel (this study; Fauvel 1927; Gil 2011), Mediterranean Sea ( Linnaeus 1767; Fauvel 1927), Adriatic ( Fauvel 1927, Gil 2011) and Aegean Sea ( Gil 2011).

Habitat. Under rocks, intertidal (this study), in muddy bottoms, shallow waters ( Gil 2011).

Remarks. The original description by Linnaeus (1767) is short, with very few taxonomic details and no precise type locality is given, which is not suprising as these taxonomic requirements were developed after Linnaeus. Specimens examined in this study match with descriptions of specimens from the French coasts studied by de Saint- Joseph (1894) and Fauvel (1927). However, all these specimens were sampled from the Atlantic coast while the type locality of this species is situated in the Mediterranean Sea. Until an accurate description, based on Mediterranean specimens, is available, we prefer to identify these specimens as Terebella cf. lapidaria .

Molecular data

During this study, 10 sequences of COI gene, belonging to eight species and 31 sequences of 16S gene, belonging to 15 species were obtained and deposited in Genbank ( Table 1, Figs 26 and 27 ). For COI, three sequences were attributed to two known species ( A. figulus , L. conchilega ), five sequences to three new species described during this study ( E. gili n. sp., P. sauriaui n. sp., T. banksyi n. sp.) and two sequences to two (probably) new species that could not be described here because sequences were obtained from a single damaged specimen ( Eupolymnia sp. C from the Gulf of Lion and Pista sp. A from the Bay of Biscay).

For 16S, eight sequences belonged to three species already known from Europe ( A. figulus , L. conchilega , T. lapidaria ), 17 sequences to seven new described species ( E. gili n. sp., E. lacazei n. sp., E. meissnerae n. sp., L. kellyslateri n. sp., P. miosseci n. sp., P. sauriaui n. sp., and T. banksyi n. sp.) and finally, as for COI, six sequences to six (probably) new species impossible to describe in this study ( Eupolymnia sp. A and Eupolymnia sp. D from Corsica, Eupolymnia sp. B from the Bay of Biscay, and finally Eupolymnia sp. C from the Gulf of Lion; Pista sp. A and Pista sp. B from the Bay of Biscay and Corsica respectively).

Obtaining these sequences was essential to separate species morphologically very similar. Indeed, several of the new species described here belong to complexes of cryptic or pseudo-cryptic species. Without molecular tools, it would have been impossible to differentiate and describe species of the P. cristata , L. conchilega or E. nebulosa complexes ( Figs 26 and 27 ). These data also permitted us to highlight that several sequences stored in Genbank do not correspond to the correct species (and probably belong to undescribed species). The first one is A. figulus . Based on COI molecular results ( Fig. 26 ), the French specimens differ from the A. figulus identified by Carr et al. (2011) from Canada, New Brunswick (GenBank accession number: HQ023982 ). However, as our specimens were sampled much closer to the type locality (East coast of Scotland) and are morphologically similar to the description of the species, we suggest that the Canadian specimens may belong to a different species, probably not new, as several species of Amphitrite have been described from that area, such as A. brunnea ( Stimpson, 1853) , A. ornata ( Leidy, 1855) , or A. attenuata Moore, 1907 . On the other hand, the French specimens match molecularly with specimens from the White Sea, Kandalaksha Bay (GenBank accession: HM417784 ) ( Hardy et al. 2011) ( Fig. 26 ). Lanice conchilega , identified by Stiller et al. (2020) from Scilly Islands ( England) is actually L. kellyslateri n. sp. and differs molecularly (but also morphologically, see L. kellyslateri n. sp. remarks) from specimens of L. conchilega sampled close from the type locality. These two species belong to a complex of cryptic species and are visually very difficult to separate. Finally, based on 16S molecular results ( Fig. 27 ), E. nebulosa sequenced by Stiller et al. (2020) and sampled from Banyuls-sur-Mer Bay is the same one as E. lacazei n. sp. described here from the same locality. This species is morphologically different from E. nebulosa described from UK (see E. lacazei n. sp. Remarks).

Figures

FIGURE 2. Schematic illustrations of different uncini morphologies, in lateral views (following Nogueira et al. 2010). A. Terebella cf lapidaria, MNHN-IA-PNT 131, SG X; B. Lomia ramzega, paratype MNHN-IA-TYPE 1791, SG XII; C. Eupolymnia meissnerae n. sp., holotype MNHN-IA-TYPE 2024, SG X. D. Pista sauriaui n. sp., paratype MNHN-IA-TYPE 2036, SG V. Abbreviations: Af, anterior filament; Ba, base; Cext, comma-shape extension; Cr, crest; DB, dorsal button; He, heel; LH, long handle; MF, main fang; Pr, prow. Scale bars: 25 µm.

FIGURE 25. Terebella lapidaria Linnaeus, 1767 MNHN-IA-PNT 131 (A–C, E–G), SMA-Arc-Eupoly-GdBC (D, H). A. Entire specimen, lateral view; B. Anterior end, lateral view, MG staining; C. Anterior end, dorsal view, MG staining; D. Anterior end, dorso-lateral view; E. Notochaetae, SG IX; F. Notochaetae, posterior segments; G. Uncinus, SG X; H. Uncini, SG V.Abbreviations: Bt, buccal tentacles; Ll, lower lip; Sp, spur. Numbers referring to segments.

FIGURE 26. Majority-rule consensus tree from Bayesian analysis using COI. Asterisk indicates posterior probability> 80%. Text in red refers to specimens sequenced during this study.

FIGURE 27. Majority-rule consensus tree from Bayesian analysis using 16S. Asterisk indicates posterior probability> 80%. Text in red refers to specimens sequenced during this study.