Spiophanes norrisi sp. nov. (Polychaeta: Spionidae) — a new species from the NE Pacific coast, separated from the Spiophanes bombyx complex based on both morphological and genetic studies Author Blank, Miriam text Zootaxa 2009 2278 1 25 journal article 10.5281/zenodo.191147 e46b99bc-4894-4e3b-877e-2c06869dad00 1175-5326 191147 Spiophanes bombyx ( Claparède, 1870 ) ( Figs 2–4 ) Spio bombyx Claparède, 1870 : 485 –487, plate XII, fig. 2. Spiophanes bombyx . – Mesnil, 1896 : 249 –257, plate XV, figs 1–22. – Mesnil, 1897 : 91 –92, plate III, figs 17, 20, 21. – Söderström, 1920 : 243 –244, figs 135. – Fauvel, 1927 : 41 , figs 14a–i. –Hartmann– Schröder, 1971 : 327 –328, fig. 112. –Meißner, 2005: 54–58, figs 33–35 (in part).– Not Okuda, 1937 : 222 –223, figs 3–4. –Not Light, 1978 : 60 –62, figs 60–61. –Not Blake, 1996 : 146 –147, figs. 4.18.a–e. –Not Imajima, 1991 : 128 –132, figs 8–9. –Not Blake & Kudenov, 1978 : 224 . ? Spiophanes verrilli Webster & Benedict, 1884 : 728 –729, pl. VI, figs 65–72. FIGURE 2. Spiophanes bombyx (Claparède, 1870) : A. Anterior end, dorsal view. B. Chaetigers 9–15 with dorsal ciliated organs and dorsal ciliated crests, dorsal view. — Specimens from the North Sea, German Bight, depth 24 m, leg. K. Meißner, Jul 2008. Scale: in µm. Non– type material. NE Atlantic Ocean: Norwegian Sea : 60° 36.990' N 2° 46.070 E , 110 m , 6- V-1998, 1 specimen (ZSRO-P 715). North Sea : German Bight: 54°19.728' N 6°59.700' E , 39.6 m , fine sand (d50 = 0.1 mm ), 24- VII-2008, 18 specimens ( ZMH P- 24840); 54°01.810' N 7° 01.732 E , 32.1 m , fine sand (d50 = 0.175 mm ), 24- VII-2008, 1 specimen ( ZMH P- 24841); 53°41.186' N 6° 29.870 E , 19.3 m , medium sand (d50 = 0.409 mm ), 23 VII-2008, 1 specimen ( ZMH P- 24842); 53°42.741' N 6° 26.494 E , 22.2 m , medium sand (d50 = 0.422 mm ), 23- VII-2008, 1 specimen ( ZMH P- 24843); 53°41.609' N 6° 29.839 E , 23.9 m , medium sand (d50 = 0.313 mm ), 23- VII-2008, 4 specimens ( ZMH P- 24844). Mediterranean Sea : Gulf of Naples: 40°45.987' N 14° 22.445 E , 20 m , silty sand, 22- III-2005, 2 specimens ( ZMH P- 24832); Litorale Domitio: 40°57.337' N 13° 59.505 E , 5 m , fine sand, VIII-2003, 9 specimens ( ZMH P- 24833), Costa del Azahar: between cabo de San Antonio and Puerto de Valencia [ 39°27.513' N 0° 18.846 W ], 1 specimen ( MNCN 16.01/ 2653), 1 specimen ( MNCN 16.01/2620), 1 specimen ( MNCN 16.01/2891); Costa del Sol: Nerja, [ 36°44.759' N 3° 43.856 W ], 14- VI-1983, 1 specimen ( MNCN 16.01/9855); Punta de Torrox [ 36°42.834' N 3° 57.229 W ], II-1995, 1 specimen ( MNCN 16.01/8754), Asturias: [ 36°42.834' N 3° 57.229 W ], 1 specimen ( MNCN 16.01/ 9918); Aegean Sea : Izmir Bay [ 38°25.935' N 27° 03.271 E ], 24 m , muddy sand, 13- VIII-2001, 1 specimen ( ZMH P- 24836), Mersin Bay [ 36°46.983' N 34° 36.817 E ], 5 m , mud, 17- IX-2005, 3 specimens ( ZMH P- 24834), Iskenderun Bay: [ 36°44.561' N 35° 42.783 E ], 25 m , muddy sand, 10- IX-2005, 2 specimens ( ZMH P- 24835), [ 36°44.561' N 35° 42.783 E ], 10 m , muddy sand, 10- IX-2005, 6 specimens ( ZMH P- 24837). FIGURE 3. Spiophanes bombyx (Claparède, 1870) : Chaetiger 10 – 18, dorsal view, methyl green stained nuchal organs. — Specimen from Turkey, Iskenderum Bay, sublittoral, leg. E. Dagli, Sep 2005. Scale: 0.5 mm. Additional material. Spiophanes verrilli Webster & Benedict, 1884 . North Atlantic Ocean, Massachusetts : Provincetown, Wellfleet, intertidal, 5 slides with several chaetigers, 1 middle and 1 posterior fragment ( USNM 480). Description. Examined specimens between 0.2 and 1.5 mm in width. Prostomium broad anteriorly, subtriangular, with long, digitiform anterolateral horns ( Fig. 2 A). Occipital antenna absent. Usually two pairs of black or red eyes present. Dorsal ciliated organs starting posterior to the prostomium as continuous ciliated bands to the end of chaetiger 2, thereafter as pair of segmental ciliated patches increasing in size, eventually becoming pairs of almost straight short ciliated bands until about chaetiger 15; after chaetiger 15 ciliated bands often slightly shorter, sometimes comma-shaped or slightly oblique; application of Shirlastain A reveals additional thin transverse ciliary bands between metameric ciliated patches; metameric dorsal ciliated organs extending well into the abdominal region but not present on posteriormost chaetigers ( Figs. 2 A–B, 3, 4D, Tab. 2). Peristomium moderately developed. First parapodium oriented dorsally; postchaetal lamellae subulate, of about same length in neuro- and notopodium ( Fig. 2 A). Parapodia of chaetigers 2–4 in dorsolateral to lateral position; postchaetal notopodial lamellae subulate; neuropodial postchaetal lamellae subtriangular to rounded. Chaetigers 5–8 with short, subtriangular notopodial and reduced neuropodial postchaetal lamellae ( Fig. 4 A). From chaetiger 9, notopodial postchaetal lamellae subtriangular to cirriform with broad base; neuropodial lamellae reduced ( Figs 2 B, 3). From chaetiger 15, notopodial postchaetal lamellae cirriform with broad base ( Figs 3 , 4 D). FIGURE 4. Spiophanes bombyx (Claparède, 1870) : A. Chaetigers 4–9 with openings of glandular organs on chaetigers 5, 7, 8, 9, lateral view. B. Neuropodial hooks from chaetiger 16, dorsal apical view. C. Chaetigers 9–11, lateral view. D. Posterior end with stout curved notochaeta, dorsal view. — A, B, D specimens from the North Sea, German Bight, depth 24 m, leg. K. Meißner, Jul 2008; C specimen from the Gulf of Naples, depth 20 m, leg. P. Lanera, March 2005. Scale: in µm. Chaetal spreader “0+1 type ” usually with undulate opening well developed on chaetigers 5, 7 and 8; on chaetiger 6 opening of glandular organ reduced to a short slit; on chaetigers 9–14 (or sometimes 9–13 in small juvenile specimens) openings as lateral vertical slits ( Fig. 4 A, C). Ventrolateral intersegmental pouches absent. Dorsal ciliated crests present from chaetiger 3, increasing in size in further posterior segments. Chaetiger 1 usually with one stout, crook-like chaeta in each neuropodium; remainder of chaetae all capillaries; hirsute under SEM and appearing granulated when viewed with light microscopy; notochaetae arranged in a tuft; neurochaetae in two rows. Chaetigers 2–4 with capillaries with narrow but distinct sheath in both neuro- and notopodia; hirsute (SEM) or appearing granulated (light microscopy); notochaetae arranged in a tuft; neurochaetae arranged in two rows. Notochaetae of chaetigers 5–14 sheathed capillaries, arranged in three rows; neuropodia with stout sheathed capillaries, distally pointed, in one or two rows. Neuropodial hooks with partially reduced hood first present from chaetiger 15, sometimes small juvenile specimens with neuropodial hooks from chaetiger 14 ( Fig. 8 left); hooks quadridentate, with main fang surmounted by single unpaired tooth and pair of smaller distal teeth, additional small teeth sometimes present ( Fig. 4 B); 4–11 hooks arranged in a single row, number depending on specimen size ( Fig. 9 ); notopodia with narrowly sheathed capillaries. Ventral sabre chaetae first present in hook-bearing chaetigers. Stout, curved notochaetae in far posterior parapodia present ( Fig. 4 D). TABLE 2. Comparison of important morphological characters of Spiophanes species similar to S. bombyx , additional thin transverse cilia bands observed in S. bombyx , S. norrisi sp. nov. and S. cf. uschakowi are not drawn; S. verilli Webster & Benedict, 1884 not included since status unresolved; *= observations based on limited number of specimens. S. bombyx S. norrisi S. cf. uschakowi S. aucklandicus S. anoculata 1 1 1 1 1 2 2 2 2 2 3 3 3 3 3 4 4 4 4 4 5 5 5 5 5 6 6 6 6 6 Dorsal ciliated 7 7 7 7 7 organs 8 8 8 8 8 9 9 9 9 9 10 10 10 10 10 11 11 11 11 11 12 12 12 12 12 13 13 13 13 13 14 14 14 14 14 15 15 15 15 15 16 16 16 16 16 17 17 17 17 17 18 18 18 * 18 * 18 * Pygidium usually with two thin, cirriform anal cirri.

Chaetal spreader fully developed	on chaetigers 5,7,8	on chaetigers 5,7,8	on chaetigers 5,7,8 *	on chaetigers 5,7,8	on chaetigers 5-8
Sabre chaetae	only in hook-bearing chaetigers	from chaetiger 10 (rarely from 9 th)	from chaetiger 9 or 10 *	only in hook- bearing chaetigers	from chaetiger 4
Neuropodial hooks	from chaetiger 15 (14) (depending on specimen size)	from chaetiger (13), 14 or 15 (depending on specimen size)	from chaetiger 15 *	from chaetiger 14	from chaetiger 15
Number of neuropodial hooks	4-11	4-10	6-7*	6-13	6-8 (Blake 1996) 8-12 (Hartman 1969)

Pigmentation. Reddish pigment in neuropodia often visible in chaetigers of the middle body region; sometimes also present in neuropodia of chaetigers 15–18. Some specimens with brownish pigment on the prostomium close to the eyes. Methyl green staining pattern. Inconspicuous. Biology. Hannerz (1956) in a paper about the larval development of S. bombyx with detailed descriptions of the morphology of the different larval stages reports planktonic larvae to occur between April to December in Swedish coastal waters. Remarks. S. bombyx can be distinguished from all other Spiophanes species in the North and Mediterranean Seas by its strikingly long anterolateral horns, the chaetal spreaders of the “0+1 type ” with undulate opening fully developed on chaetigers 5, 7, and 8, the dorsal ciliated organs starting as two short continuous bands extending to the end of chaetiger 2 followed by metameric patches in a species-specific pattern (see species description above), the exclusive presence of sabre chaetae in hook-bearing chaetigers, and the presence of partially hooded hooks in neuropodia of the posterior body region. Differences between morphologically similar species from other geographic regions most importantly concern the orientation of the dorsal ciliated patches on chaetigers of the middle and posterior body region ( Tab. 2 ), the start of neuropodial hooks on chaetiger 15 (on chaetiger 14 only in small juvenile specimens) as well as the restriction of sabre chaetae to hook-bearing chaetigers. In addition, the number of neuropodial hooks is lower in S. bombyx compared to S. aucklandicus if comparing specimens of same size (see Meißner 2005); no differences in regard to this character were found between S. bombyx and S. norrisi sp. nov. ( Fig. 9 ). S. bombyx can no longer be regarded as a species with a world-wide distribution. The distribution might be wider than reflected by the records from the North and Mediterranean Seas confirmed here, but the status of S. bombyx as a cosmopolitan species has to be dismissed. The synonymy of S. verrilli Webster & Benedict, 1884 from Wellfleet, Massachusetts with S. bombyx has to be verified in future studies based on the investigation of material from the type locality (also see chapter Discussion in this paper). The type material of S. verrilli was investigated by the first author but its study did not allow final conclusions since important characters are not observable any longer. It consists of five permanent slides with several chaetigers, plus one middle and one posterior fragment. Geographical distribution. North Atlantic Ocean: along European coasts, Mediterranean Sea. In shallow waters up to subtidal depths.