Salmacina Claparède, 1870

Ben-Eliahu, M. Nechama & Ten Hove, Harry A., 2011, Serpulidae (Annelida: Polychaeta) from the Suez Canal- From a Lessepsian Migration Perspective (a Monograph) 2848, Zootaxa 2848 (1), pp. 1-147 : 62-65

Genus Salmacina Claparède, 1870

Preliminary remarks. Except for the presence / absence of an operculum on the first dorsal pair of radii in Filograna , the nominal genera Filograna and Salmacina are so similar that they have been classified together as indistinguishable by various authors. Moreover, different authors have dealt with the relationship of the operculate Filograna implexa to the non-operculate Salmacina dysteri ( Huxley, 1855) in different ways (e.g., Gee 1963; Day 1967). Until it is proven that both the operculate and non-operculate forms are the same genus and that the generic synonymy is warranted, valuable information may be lost by not separating these forms. The nominal taxa F. implexa , S. dysteri , S. incrustans , and even S. amphidentata have all been attributed a “world-wide” distribution. This of course may have been the result of ongoing ship-transport, but, in the past, several “world-wide” distributions have been split into more regionally distributed “sibling” taxa. In addition, errors in identification have also impacted on distributions cited in some of the literature (discussed below).

Most of the regional records are presumed to be a melange of the names, Filograna or Salmacina , with some later records cited more cautiously as Filograna / Salmacina – complex. Therefore we have refrained from giving lists of synonyms (including those from samples collected from Cyprus, Israel and Gulf of Suez in different Israeli collections that will eventually be dealt with separately). The Levant, Suez Canal and Red Sea specimens listed in the “Material examined” section below were all non-operculate; thus, we would now attribute them all to Salmacina sp. / spp., and not to Filograna . An updated generic description of both nominal genera is given in ten Hove & Kupriyanova (2009).

The genus Salmacina comprises very minute taxa requiring use of the SEM in order to elucidate the dentition of the collar chaetae and to count the number of teeth in the transverse rows of the uncini in frontal view (i.e., to determine the dental formula). Re-examination by SEM of some pre-SEM (i.e., pre-1965) voucher specimens showed that some of the regionally pertinent determinations were erroneous (e.g., pp. Pixell 1913, Fauvel 1933a, b, see “Material examined” below). To stabilize the taxa, the descriptions of published type material should be updated, and, when unavailable (as in Salmacina dysteri ), material from the type localities and habitats of the described taxa should be examined with the SEM and redescribed, e.g., Nogueira & ten Hove (2000). Similarly, voucher specimens of regional records published prior to the sixties should be verified and compared with this redescribed type or neotype material. To overcome the morphological variability found in these clonal species (Nishi & Nishihara 1997), new efforts at redescription should include comparative genetic analysis carried out on in conjunction with the morphological work. Unfortunately most museum specimens have been denatured due to fixation in formaldehyde, necessitating collection of fresh specimens from the type localities and making the revision of the Salmacina taxa a truly Sisyphean project. Although we were warned not to attempt to deal with this intractable genus (by H. Zibrowius, pers. comm.), we were determined to look for clues to the provenance of the “Yellow Fleet” Salmacina species from the Bitter Lake. We examined specimens from different biogeographical regions and sought to find robust characters to distinguish between the taxa. This effort resulted in the present identification of the “Yellow Fleet” taxon as Salmacina incrustans , provenance, Mediterranean Sea. We also succeeded in obtaining a more precise characterization of some Salmacina taxa in regions impacting the Suez Canal.

We introduce the term, “paucidentate-rasp-shaped” uncini (pru - type), to refer to thoracic uncini with few (2–3) teeth in the transverse row proximal to the fang (termed the “(F+1)” row), the number of teeth not increasing greatly in the transverse rows towards the apex, long and thin (rectangular) in frontal view (extreme form exemplified by the nominal Salmacina dysteri sensu Fauvel (1927 ; see Fig. 23A ). A contrasting term, “multidentate rasp-shaped” uncini (mru - type) refers to thoracic uncini that have the transverse row proximal to the fang, the (F+1) row, numbering 3 or more teeth, increasing greatly in the number of teeth (and in width of the uncinus) towards the apex, ovoid or wedge-shaped in frontal view (extreme form exemplified by Salmacina amphidentata not Jones, 1962 sensu Fiege & Sun, 1999; see Fig. 23B ). Frontal SEM views of uncini of these two forms are given in Figs 23A and B ; see also Figs 27B and E ). Data on thoracic uncini-type have been added to verified citations and the “Material examined” section, below.

References for the Filograna / Salmacina- complex names used for material from the Suez Canal or its vicinity.

Protula dysteri Huxley, 1855: 113 , figs 1–11 [Type locality: Wales, Caermarthen Bay, Tenby]; referred to Salmacina dysteri by Claparède (1870: 176).

Salmacina incrustans Claparède, 1870: 519 [Type locality: Italy, Naples]; Fauvel 1927: 378–380, fig. 129l [Atlantic: Madeira, Cape Verde, Mediterranean: Malaga, Valencia, Marseilles, Naples].

Salmacina aedificatrix Claparède, 1870: 519 [Mediterranean, Tyrrhenian Sea, Naples; “Usually synonymised with S. dysteri ” ( Nogueira & ten Hove 2000: 158) ].

Filograna / Salmacina species complex: Wehe & Fiege 2002: 124 [Suez Canal].

Material examined for comparison. The number of teeth in the horizontal rows of the rasp-shaped uncini is a comparatively easy morphological character to quantify and to use as a parameter of variability. In the SEM micrographs, we have been able to observe variability between uncini of the same torus; even in the number of teeth in the proximal row to the fang—in the (F+1) rows of adjacent uncini. In Fig. 26A , for example, the teeth in the (F+1) row proximal to the fang of the seven adjacent uncini counted from right to left numbered 3,3,2,3,3,2,2 teeth (the remaining uncini in the torus were not distinct). Regretfully, the number of individuals available from different locations or that we could SEM was in most cases very small, thus the observations given below are carried out without performing statistical comparisons. Nonetheless, some differences found between uncini of specimens from different locations were clearly distinct. The abbreviation, “tu” refers to which torus was examined, e.g., “tu1” or “tu2” refer to the first or the second thoracic torus.

Presumed Salmacina dysteri s. str. based on North Atlantic location and description. North Sea, Scotland: Orkney Islands, Mainland, Sound, Point of Hellia, Stn 20, 15.5 m, legit M.H. de Kluyver 1990, det. H.A. ten Hove 1990 Salmacina sp. , ZMA V.Pol. 3768. SEM ZMA11 -tu1: (F+1) = 3,2,3,3,3 (five adjacent uncini counted in SEM micrograph of first thoracic torus) . ZMA11 -tu2: (F+1) = 3,2,4 s,2,3,4 s (six adjacent uncini counted from second torus; s denotes an apparently aberrant splitting of the apical end observed in several of these uncini, never remarked until now) . ZMA11 -tu2: (F+1) = 2,3,3,3,3,3,3,3,3,2 . ZMA-11 -tu3: (F+1) = 2,?,2,3,3,2,2,2,2.— Hebrides, Isle of Muck, Ruadh leam laranche, possibly legit Mrs. E.A.T. MacEwen 30.VII.1938, Salmacina dysteri , determinator unknown, Royal Scottish Museum, Edinburgh; interlacing tube aggregate, ca. 20 specs with (F+1) = 2 or 3. SEM S-14-1-tu2: (F+1) = 2,3,2,2,3,2. S-14-1-tu3: (F+1) = 3,3,3,2,2,2,2. All thoracic uncini of pru - type .

Irish Sea, Wales: Holyhead, Anglesey, Menai Bridge Marine Biological Station , 11.XI.1957, Salmacina dysteri , determinator unknown, NMWZ 1985.042 .1957. SEM ATW1-200-w2-007: (F+1) = 3,2,2,3,2,3,2,2,3 (sequence of nine uncini). SEM ATW2-1-209-w1-tu: (F+1) = 3,3,3,3,2,3,3,3,2,2,2,3,2; thoracic uncini of pru - type ).— Menai Bridge, Menai Bridge Marine Biological Station , VIII.1953, Salmacina dysteri , det. unknown, NMWZ 1985.042 .672, tube aggregations, several small, presumably net-like, clusters. SEM ATW1-219-w3- tu: (F+1) = 4,3,3,3,3,2,3,2,3,3,2 (thoracic uncini of pru - type) .

English Channel, coast of France, Saint-Vaast-La Hougue, legit, det. J.L.A. de Quatrefages Filograna ; redet. P. Fauvel S. dysteri, MNHN ; subsamples deposited in BM (NH): BM (NH) 1928.4.26.106 and 1928.4.26.886, agglomerated tubes. SEM Paris-Vaast-1-3-016-tu2: (F+1) = 2,2,3,2,3,3,3. SEM Paris-Vaast-1- 3-018-tu4: (F+1) = 3,3,3,3,3,3,2,2,3,3,2 (thoracic uncini of pru- type).

Salmacina dysteri confirmed based on comparison of specimens with above North Atlantic material. Central Atlantic Ocean , Cape Verde Islands, São Vicente, legit C. Crossland 11.IX.1904, presented by J.S. Gardiner, det. H.L.M. Pixell Salmacina dysteri, BM(NH) 1924.6.13.131. SEM 131-017, left part of torus: (F+1) =?,?,3,3,2,3,3,2,2,3 (counted eight of the ten uncini in the micrograph). SEM 131-018, central part (F+1) = 2,3,2,3,2,3,2 (maximum no. of teeth in widest horizontal row 4; thoracic uncini of pru - type).— Additional sample from same locality, det. H.L.M. Pixell 25.IX.1912, BM (NH) 1938.7.25.61–75 including permanent mount. SEM 61-72-2-tu5: (F+1) = 2,2,3,3,3,3,2. 61-72-2-tu6: (F+1) = 3,3,3,3,2,3,3; thoracic uncini of pru - type ).

Questionable nominal Salmacina dysteri records of which voucher specimens are missing, determination generalised to Salmacina sp.

Suez Canal proper: Potts (1928: 701–702) reported Salmacina sp. as S. dysteri from two sites in the southern part of the canal, Stn K2 at km 124 in the Little Bitter Lake from buoys between 0–4 m, and from km 152, presumably scraped from the pier or from bivalves encrusted on the pier.

Gulf of Suez: Two samples from the entrance to the Suez Canal: (1) The Percy Sladen Trust Indian Ocean Expedition of 1905, sample from a small buoy, det. H.L.M. Pixell (1913: 87) Salmacina dysteri . —(2) The Cambridge Expedition, two sites at Port Taufiq: Stn PT 0, scraped from piers along the avenue; Stn PT 3, scraped from a wooden fence along the bathing place, det. F.A. Potts (1928: 701–702) Salmacina dysteri (sample details from Fox 1926).

Nominal Salmacina dysteri records that proved to belong to taxa with multidentate rasp-shaped uncini (i.e., erroneous determinations) generalised to Salmacina sp. / spp.

Gulf of Suez: R. Ph. Dollfus Expedition. stn XI, 32º44'– 32º47'E, 28º49– 28º54' N, 25–30 m, coralligenous sand, det. P. Fauvel (1933a: 80) Salmacina dysteri, MNHN. SEMs: Paris-GS-1933-004-tu1: (F+1) = 5,5,5,4,4,?,4,5 (counted seven of eight uncini in first torus). Paris-GS-1933-003-tu2: (F+1) = 4,5,?,4,4,4,4. Paris-GS-1933-002-tu3: (F+1) = 4,5,5,?,4,4,4,4,?,4,4 (see Fig. 27B , second thoracic uncinus) .

Red Sea proper-Indo-West-Pacific (excluding citations from Gulf of Suez and Gulf of Aqaba already given above) Indian Ocean, Tanzania, Zanzibar, Wasin Harbour, legit C. Crossland, 1902, presented by J.S. Gardiner, det. H.L.M. Pixell (1913: 38) Salmacina dysteri, BM(NH) 1924.6.13.152 [SEM not permitted]. In Pixell’s permanent mount, studied with oil immersion, the single uncinus in focus showed (F+1) = 4 teeth; the uncini are F+8, i.e., with eight horizontal rows of teeth “above” the fang (see Fig. 23 ), and with a maximum number of six teeth in the widest row (formula F=8/6).— Zanzibar area, legit C. Crossland, 22.VIII.1901, presented by J.S. Gardiner, det. H.L.M. Pixell (1913: 38) Salmacina dysteri, BM(NH) 1938.7.25.13–25 (material previously dried up, specimen extracted from tubes by HAtH). SEM 13-25-09-tu1: (F+1) = 3,3,3,3,3,3 (counted sequence of six uncini in first torus); the maximum number of teeth in the transverse rows of both the 1 st and 2 nd uncini was 7). SEM 13-25-tu3: (F+1) = 3,3,3,3,3,3,3,3 (in the eight uncini in third torus, only the three anterior horizontal rows exposed, others covered); presumed to be a Salmacina sp. of mrutype.—From Zanzibar area, from bottom of S.S. “Juba”, legit C. Crossland 29.VII.1901, presented by J.S. Gardiner, det. H.L.M. Pixell (1913: 38) Salmacina dysteri, BM(NH) 1938.7.25, 26–40. SEM 26-40-023-tu: (F+1) =?,?,4,4,? (counted only two uncini in torus, other uncini not in focus or covered over). The glycerine mounts of wet material studied with oil immersion showed (F+1) = 4 teeth in proximal row to fang, thus, uncini of mru - type. Pixell’s slide A, seen with oil immersion, seems to show (F+1) = 3 or 4 teeth (none of the details were seen clearly).—Off coast of Zanzibar, Jembani, 22.VIII.1901, legit C. Crossland, presented by J.S. Gardiner, det. H.L.M. Pixell (1913: 38) Salmacina dysteri, BM(NH) 1938.7.25.41–52. SEMs (HAtH) show truncated tips of fang, frontal view of these thoracic uncini more rectangular than others: First torus, tu1: (F+1) = 3,3,3,?4,?,?,?. Second torus, tu2: (F+1) =?, 4, 3, 3,4,5; redetermined herein as Salmacina sp. of mru - type). — Hawaii: Biofouling, legit J.H. Bailey-Brock. tu2: (F+1) = 5,5,4,4,4,5,4,4 (counting the last uncinus showed that the number of horizontal rows was 9 and its detailed formula was F: 4:5:6:7:8:10:?9:9:?6 (formula F+9 / 10), thus it is characterised as a Salmacina sp. of mru - type).

Salmacina sp. of multidentate rasp-shaped uncini - type. Gulf of Aqaba, Elat, North Beach, 5–10 m, legit N. Shashar 24.06.2002, det. M.N. Ben-Eliahu Salmacina sp. , on / in crevices of red Bryozoa on rope from fish cages, few; live material with red clusters of prostomial ocelli ( Fig. 27E ). SEM-A-Elat-003-tu1: (F+1) = 5,4,5,5,4,5,4,?,4,4 (counted nine of 10 uncini of first torus). SEM-B-Elat-tu1: (F+1) =?,5,5,4,4,5,?. SEM-D- Elat-009-tu1: (F+1) = 4,4,3,3,3,3,5,3,4,3,4,4. Identified as Salmacina sp. of mru – type ( Fig. 27E ).

Comparative specimens identified as Salmacina incrustans Claparède, 1870 from Atlantic, Mediterranean (consistent with the Fauvel, 1927 description). Spain, Costa Brava near Playa de Aro, Playa de San Pol, S, stones and debris in rock-crevices, 0.5– 1 m, legit, det. H.A. ten Hove 11.VI.1971, S. incrustans, ZMA V.Pol. 3814, ZMA-SEM 0032 (number on the ZMA micrograph H 46), uncini with 7 horizontal rows of teeth, proximal row to fang (F+1) = 3,3,3,3,2 or 3,3,3 (counted seven uncini in sequence); the only uncinus positioned in full frontal view that could be counted showed maximum no. of teeth, 5–?6, in fifth horizontal row.—Ligurian Sea, Monaco, coll. Prince of Monaco 22.XII.1910, on hull of “Eider”, det. P. Fauvel S. incrustans, MNHN , tubes only.— Monaco harbour, det. P. Fauvel S. incrustans, BM(NH) 1928.4.26.144 / 154 ( SEM not allowed).—Tyrrhenian Sea, Naples, det. P. Fauvel S. incrustans, BM(NH) 1928.4.26.954.— Croatia, Istria, S. Pomer, outside harbour pier next to Pomer camping ground, boulders, legit, det. H.A. ten Hove VII.2003 S. incrustans, HUJ-SEM (only one uncinus in focus, F:2:3:3:3:3:3:4:4:?2 [covered]).

Figures

FIGURE 23. Frontal SEM views of the two extreme forms (types) of thoracic uncini in Salmacina spp. A—Paucidentate rasp-shaped uncini (pru-type) of Salmacina dysteri: Uncini from third thoracic torus of specimen from the Isle of Muck, Hebrides, Scotland, 30.VII.1938 (Scottish Museum, Edinburgh); note “above” the fang (the enlarged anterior-most tooth, designated in the figure as F), 7 transverse rows of teeth, with two teeth in the row proximal to the fang (rows in figure numbered and designated by a line); that proximal to fang designated as (F+1); apical (posterior) row, here, the 7th row, designated as (F+7); the detailed dental formula of the left uncinus in A is then (F+1) = 2: (F+2) = 2: (F+3) = 3: (F+4) = 3: (F+5) = 3: (F+6) = 3 and (F+7) = 3, or, briefly, F:2:2:3:3:3:3:3 (as noted above, in the detailed dental formula, the different transverse rows of the uncinus are separated by a colon), B—Multidentate rasp-shaped thoracic uncini (mru-type) present in Salmacina amphidentata not Jones, 1962 sensu Fiege & Sun (1999) from Hainan Island, near China (from Fiege & Sun 1999: 135, fig. 22c), “above” the fang, 9 transverse rows of teeth, with six teeth in the (F+1) row proximal to the fang, and with the number of teeth in the transverse rows increasing to 10–11 towards the apex (transverse rows 5– 7); the dental formula of the right uncinus is F:6:7:?9:10:11:11:11:?8:9.

FIGURE 26. Salmacina incrustans: Chaetae (other than collar chaetae) and uncini of specimens shown in Figs 25. A, B—Thoracic and abdominal uncini of spec. 13-3: A: Thoracic uncini (numbers refer to marked uncini in the figures): (1) F:3:4:4:5:6:6:3 = F+7 rows. (2) F:2:3:3:4:5:4 = F+6 rows, B—Abdominal uncini: (3) F:5:7:7:7:8:8:7:6:4 = F+9 rows; (4) F:5:5:6:6:3:5:6:5:5:5 = F+10 rows, C—Thoracic uncini of spec. 10-3, thoracic uncini teeth count: (5) F:3:4:4:4:4:4 = F+6 rows. (6) F:3:4:3:4:5:5:3 = F+7 rows, D—Apomatus chaeta and limbate chaetae (specimen 10-2), E—Abdominal uncini of spec. 13-2, (7) F:4:5:6:7:6:7:7:5:5 = F+9; (8) F:5:5:6:8:7:7:6:7:4 = F+9, F, G—Abdominal chaetae of two specs (15-1 and 8-1. Magnifications: A—13,000 x, B—16,000 x, C—12,000x, D—6,000 x, E—10,000 x, F—12,000 x, G— 16,000 x.

FIGURE 27. Comparison of Salmacina spp. from the Suez Canal, Gulf of Suez, and Gulf of Aqaba. A—Salmacina incrustans from Great Bitter Lake, spec. 10-3, thoracic uncini both (F+1) = 3, left uncinus detailed F:3:3:3:3:4:4 (compare Fig. 23), B—Salmacina not dysteri (Huxley) from Gulf of Suez (32°44’–32°47’ E., 28°49’–28°54 N., 25–30 m, legit R. Ph. Dollfus, Stn 11, 8.XII.1928, coralligenous sand, det. P. Fauvel (1933a: 80, 1933b: 143), generalised herein to Salmacina sp.; thoracic uncini from second torus of single (minute) specimen, apical part of uncini covered by a flap; F:4:5:5:? (left uncinus) and F:4:6:6:6:? (right uncinus), C—Collar chaeta fin, blade broken off, D—Intertwining, netlike tube aggregation from which minute specimen was removed, E—Thoracic mru-type uncini of Salmacina specimen from Gulf of Aqaba (from Elat). Magnifications: A—12,000 x, B, C—10,000 x, E—8,000 x.