Hydroides elegans ( Haswell, 1883 )

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 : 19-24

Hydroides elegans ( Haswell, 1883)

Figs 5D, E, 6 , 7

Eupomatus elegans Haswell, 1883: 633 , pl. 12 fig. 1 [Type locality: Australia, Port Jackson ].

Hydroides elegans: Zibrowius 1971: 721–727 , figs 56–64 [synonymy of various older material determined as H. norvegica ; distribution: Florida; Senegal; Ghana; Angola; South Africa; France; Italy; Tunisia; Alexandria; Suez Canal, Isma’iliya; Suez; Mozambique; Indonesia, Java; western Australia; New South Wales; Queensland; Hawaii; California]; ten Hove 1974: 46–47, figs 1–3 [ Netherlands, U.K., France, Ghana, Tunisia, Malta, Italy, Mozambique, Sri Lanka, Argentina, the Netherlands Antilles]; Zibrowius 1979b: 133–134 [ France, Toulon Port, biofouling removed from the aircraft carrier “Foch” that had travelled in the Indo-West-Pacific, going and returning via the Suez Canal]; Bianchi 1981: 56–58, figs 18a–f [ Italy]; Ben-Eliahu & Fiege 1996: 29–30, 33, 38, fig. 10b [ Turkey, Cyprus, Israel, Egypt]; Zenetos et al. 2005: 73 [classified as an “established cryptogenic alien invasive species” in the Mediterranean].

Eastern Mediterranean

Hydroides norvegica not Gunnerus, sensu Potts 1928: 700 [ Egypt, Port Said from Buoy no. 723, collected in 1924, first record from the Levant Basin]; Fauvel 1937: 44 (synonymised in Zibrowius 1971: 722) [ Egypt, Alexandria, 4–10 m, on Caulerpa ]; Tebble 1959: 29 [ Israel, off Atlit, Stn 507, 16 m, legit A. Yashouv (= A. Wirszubski), Teb507, BM(NH) 1955.10.12.74, 8 specs (7 of them juveniles with primary opercula, material examined by us);? Banoub 1961: 8, fig. 4 Alexandria, material not examined by us, but in view of remark of Ghobashy & Ghobashy 2005 below most probably H. elegans ]; Laubier 1966: 9, synonymised in Zibrowius 1971: 722 [ Lebanon]; Ben-Eliahu 1970: fig. 2a [ Israel]; Ghobashy 1977: 214–218, tables 1–3, figs 3–5 [Alexandria ( Ghobashy & Ghobashy 2005: 90 implicitly synonymised this with H. elegans by naming H. elegans as the principle fouling organism in the harbour since 1976)]; Ghobashy & Selim 1976a: 287–295, figs 1–5 [Alexandria, eastern harbour (see note above)]; Ghobashy & Selim 1976b: 303–313 [Alexandria, harbour, (see note just above)]; Goren 1980: 278 [ Israel, Ashdod Port, panels, 0.2 m (synonymised in Ben-Eliahu & Fiege 1996: 7)].

Hydroides elegans: Ben-Eliahu 1976: 107 [ Israel, intertidal cryptofauna]; Zibrowius & Bitar 1981: 159–160 [ Lebanon, Beirut, Zaitouné, 5 m, 23.IX.1978, on bivalve]; Ghobashy 1984: 41 [ Egypt, Alexandria, harbour]; Ben-Eliahu 1991b: 518 [ Cyprus, Israel]; El-Komi 1991a: 6, 7, tables 1–4, fig. 2 [Alexandria]; Ben-Eliahu & ten Hove 1992: 40 [ Israel, entire coast from north to south, 0–24 m]; El-Komi 1992a: 123, 130, 133, tables 2, 4, 5, fig. 3, pls. 1, 2 [Alexandria harbour]; El-Komi & El-Sherif 1992: 259–260, 263, tables 3, 4, fig. 4 [Alexandria, Eastern Harbour]; Ben- Eliahu & Fiege 1996: 29–30, 33, 38 [ Cyprus, Levant coast of Turkey, Israel, Egypt]; Ergen & Çinar 1997: 237 [ Turkey, Antalya Bay]; Selim 1997b: 91–92, figs 4a–f [Alexandria, Eastern Harbour (citing Selim 1978, see App. Table 4)]; El-Komi 1998: 259, 262, 263 [buoys in Alexandria harbour]; Zibrowius & Bitar 2003: 71 [ Lebanon, Beirut]; Ghobashy & Ghobashy 2005: 90 [Alexandria, Eastern Harbour: Review, refers to Ghobashy & Selim 1976 a, b, Ghobashy 1977, Mona 1978, Selim 1978, 1996, El-Komi 1991, 1992a, b and 1998; the western harbour review quotes Ramadan (1986, see App. Table 4) on settlement]; Çinar 2006: 226, fig. 3 d, e [Levant coast of Turkey, Iskenderun Harbour, Yumurtalık Harbour, Mersin Bay, 0.1– 5 m, on rope and tyre, IX.2005]; Ramadan et al. 2006: 26, table 1 [Alexandria, Eastern Harbour; also reviewing Banoub 1961, Megally 1970, Ghobashy 1977, El-Komi 1991a, b, 1992a, 1998]; Abd-Elnaby 2009: 10 [Alexandria]; Selim 2009: 76 [Egyptian waters].

Hydroides sp. : Fishelson & Haran 1987: 122 [ Israel, Mikhmoret; specimens examined in 1999 by Ben-Eliahu all proved to be Hydroides elegans ].

Serpula concharum not Langerhans, sensu Fishelson & Haran 1987: 122 [ Israel, Mikhmoret; redet. Ben-Eliahu & ten Hove in 1999 as Hydroides elegans specimens with primary opercular stages (see “Remarks”, below)].

? Serpula vermicularis not Linnaeus, sensu El-Komi 1991a: 7, table 2 [ Egypt, Alexandria, fouling study, presumably specimens with primary opercula]; El-Komi 1991b: 290–291, table 1, [ Egypt, Alexandria (see preceding remarks)].

Hydriodes [sic] elegans: El-Komi 1991b: 290–291 , tables 1, 3 [ Egypt, Alexandria].

? Hydroides sp. : El-Komi 1997: 109 [ Egypt, Lake Manzalah, with Ficopomatus enigmaticus ].

Surpula [sic] sp.: El-Komi 1998: 262–263 [ Egypt, Alexandria, buoys in harbour; presumably specimens with primary opercula (see “Remarks”, below)].

Suez Canal

Hydroides elegans: Zibrowius 1971: 721–727 , figs 56–64 [synonymy of H. norvegica from “Pola” Expedition 1895, Isma’iliya, first record from the Suez Canal (see “Material examined”, below, and Fig. 6 ) and synonymy of various older specimens determined as H. norvegica ; Ben-Eliahu 1972a: 77 [El Qantara, Kabrit, Tis’a]; Ghobashy et al. 1980: 84, 86–91, table 2, figs 4, 5 [major fouling component throughout the canal]; Ghobashy & El-Komi 1981a: 170–177 [Lake Timsah, 4.5– 10 m], 1981b: 180–183 [Toussoum, Deversoir, Kabrit, Geneva, el Shalloufa]; Ghobashy 1984: 39, 45–47 [throughout Suez Canal]; Ghobashy et al. 1986: 319–326, fig. 3 [Lake Timsah], 1990: 677– 686 [Lake Timsah]; Selim 1997b: 91–92, figs 4a–f [El-Kab, Lake Timsah, Deversoir, ranked “first serpulid” in canal by abundance of individuals]; Wehe & Fiege 2002: 126 [Suez Canal; list of references)]; Emara & Belal 2004: 192– 204 [pooled Lake Timsah and Bitter Lakes]; Ghobashy & Ghobashy 2005: 90 [a review of the Suez Canal fouling research showed dominance of H. elegans on fouling panels throughout the canal—all authors quoted by them are referred to above, with exception of Barbary (1992, see App. Table 4)]; Selim 2009: 73 [northern part of the canal to Toussoum]; Abd-Elnaby 2009: 10 [southern part of the canal].

Hydroides norvegica not Gunnerus, sensu Potts 1928: 700 [collected in 1924, “the most common serpulid throughout the canal”; redet. Zibrowius 1971, see “Material examined”, below]; Fauvel 1933a: 76 [R.Ph. Dollfus Expedition, 7.I.1928, Isma’iliya, on boat hull (synonymised Zibrowius 1971)].

? Serpula vermicularis not Linnaeus, sensu Ghobashy et al. 1980: 84; Ghobashy 1984: 45 [Lake Timsah)]; Ghobashy & Ghobashy 2005: 93 [found in Suez Canal on fouling plates in few numbers, and referring to Ghobashy et al. 1980]; Emara & Belal 2004: 192–199 [pooled Lake Timsah and Bitter Lakes data]; Selim 2009: 73 [Ras el Esh]; all these records presumably specimens with primary opercular stages of H. elegans (or other Hydroides taxa, see Remarks below).

? Serpula concharum not (Langerhans), sensu Selim 2009: 73 [northern part of canal, Ras el Esh]; Abd-Elnaby 2009: 10 [southern part of the canal], presumably specimens with primary opercula (see “Remarks”, below).

? Serpula sp. : Ghobashy et al. 1980: 84, 87, table 2; Ghobashy & El-Komi 1981a: 172 [Lake Timsah, 4.5– 10 m], presumably specimens with primary opercula (see “Remarks”, below).

Gulf of Suez and Gulf of Aqaba Hydroides norwegica [sic] not Gunnerus, sensu Pixell 1913: 74 [Gulf of Suez, Suez, dry dock, bottoms of S.S. “Thira ”

and S.S. “Slide ” that had come from Alexandria, BM(NH) 1924.6.13.148 (synonymised by Zibrowius 1971: 722)]. Hydroides norvegica not Gunnerus, sensu Potts 1928: 700 [Port Taufiq, on barges]; Amoureux et al. 1978: 144–145

[Gulf of Aqaba, Elat, Venezia lagoon, biofouling on floats].? Serpula concharum not Langerhans, sensu Amoureux et al. 1978: 143 [Gulf of Aqaba, Elat, Venezia lagoon, biofouling on same floats as those referred to just above, presumably specimens with primary opercular stages of H. elegans

(abundant in the lagoon [1999, M.N. Ben-Eliahu & H.A. ten Hove, unpubl. data]); also pro parte S. hartmanae (see

“Remarks”, below)]; El-Komi, et al. 1998: 16–24 [Suez Bay]; Ghobashy & Ghobashy 2005: 93 [same].? Serpula vermicularis not Linnaeus, sensu Amoureux et al. 1978: 143 [Gulf of Aqaba, Elat, Venezia lagoon, presumably specimens with primary opercular stages of H. elegans [part], see above, and “Remarks”, below]; El-Komi et al. 1998: 16, 17, 20 [Suez Bay]; Ghobashy & Ghobashy 2005: 93 [same].

Hydroides elegans: Ghobashy & El-Komi 1981b: 180–183 [Gulf of Suez, Suez]; Amoureux 1983: 369 [Gulf of Aqaba, sand]; Selim 1997b: 91–92, figs 4a–f [Gulf of Suez, collected in 1988, Port Taufiq]; El-Komi et al. 1998: 11–20 [Suez Bay, collected 1992–1993 on polystyrene fouling panels and buoys]; Abd-Elnaby 2009: 10 [Gulf of Suez].

Hydroides sp. : El-Komi 1996: 7 [Suez Bay, presumably H. elegans ].

Red Sea proper-Indo-West-Pacific (excluding citations from Gulf of Suez and Gulf of Aqaba already given above)

Hydroides norvegica not Gunnerus, sensu Dew 1959: 24–25, figs a–i [various locations in western Australia, Queensland, New South Wales, Victoria, South Australia, New Zealand, New Britain].

Hydroides elegans: Mohammad 1976: 133 [Persian (Arabian) Gulf, Kuwait]; Vine & Bailey-Brock 1984: 139–140, fig. 2a [Red Sea, Sudan, Port Sudan, settlement panels at 20 m, coral slabs 30–35 m, undersides of ships (part; probably only ship-fouling material, deep material probably different species)]; Bailey-Brock 1985: 209–210, figs 11a–d [“common fouling species with a cosmopolitan distribution in warm waters of the Atlantic, Pacific and Indian oceans (see Zibrowius 1971)]; Ishaq & Mustaquim 1996: 170, figs 5a–h [ Pakistan, intertidal rocks, boat hull]; Wehe & Fiege 2002: 126 [Red Sea, Persian (Arabian) Gulf; list of references]; Abd-Elnaby 2009: 10 [Red Sea].

Hydriodes [sic] sp.: El-Komi 1992b: 638 [fouling plates, Egypt, Red Sea, Ghardaqa].

? Serpula concharum not Langerhans, sensu El-Komi 1992b: 638 [fouling plates, Red Sea, Ghardaqa, Egypt]; Ghobashy & Ghobashy 2005: 93 [citing El-Komi 1992b: 638)]; Abd-Elnaby 2009: 10 [Red Sea]. All presumably specimens with primary opercular stages of H. elegans (see Remarks below).

Material examined. Locations adjacent to the Suez Canal, Mediterranean side: Turkey: 4 samples, intertidal— Cyprus: 14 samples, <0.5 –[0–8]–[10–18]– 26 m — Israel: 106 samples, [intertidal–0.3]– [2–3]– 37 m ; one of these is the first sample from Israel, Haifa Port , 27.V.1934, leg. G. Haas, det. M.N. Ben-Eliahu, HUJ- Poly-722 — Egypt, Sinai, 11 samples, shallow–[10–15] – 46 m.

Suez Canal proper: 13 samples, no. specs not enumerated for each sample, including dry tubes: “Pola” Expedition 1895–1898, Isma’iliya, 17.X.1895 (fide Sturany 1899), on two Potamides conicus gastropods, det. E. von Marenzeller H. norvegicus (fide Stagl et al. 1996), redet. H. Zibrowius 1971 ( NHMW, Evert. Varia 2034, alte invert. no. 17199), 2 specs ( Fig. 6 ).—Cambridge Expedition: Ras-el-Esh, Km 14, 20.XII.1924 ( Fox 1926: 50); Ferry Post, Isma’iliya, 27–28.II.1924: 3 specs, 1 with long central spine (as in Fig. 6B ) and juveniles with only primary operculum (collar chaetae checked); km 72, north of Isma’iliya, 4.XII.1924, Little Bitter Lake, K2, (Kabrit), 17.X.1924, with long central spine; K9, (Kabrit), 25.X.1924, on “Bollard” and on houseboats, various dates, 1 spec. with 2 opercula, with central spine well-developed; and 1 sample, Gulf of Suez, Port Taufiq, “Barge 678,” 10.X.1924, mass of empty Hydroides tubes, det. F.A. Potts (1928: 700) H. norvegicus , redet. H. Zibrowius (1971: 722) H. elegans , confirmed M.N. Ben-Eliahu 1986; several specs; samples catalogued together as CUZM-AN.1.1930.

Suez Canal material reported herein: 60 +?1 samples, 565 +?1 specs (of which 145 were juveniles) plus some additional non-enumerated Lake Timsah juveniles, as well as three of Beets dry samples with 14 determinable taphonomic specs (Appendix Table 2): 4 “new” Cambridge Expedition samples (samples separated from BM ( NH) molluscs [see App. Table 2A]), 3 specs: Lake Timsah-Km 78, Isma’iliya : 1 spec.; T9: 1spec.; Toussoum-Km 87: 1 spec. — Beets’ Great Bitter Lake samples, VIII / IX 1950, taphonomic material found in 3 samples, Stns 1, 4 and 21, RMNH 18528–18535 , 18538 , 18546–18549 , 14 specs, plus 24 samples with empty tubes (not enumerated) tentatively attributed to Hydroides elegans (see App. Table 2B, Fig. 7 ).— Hebrew University-Smithsonian Expeditions, 1967–1973, 27 samples, 493 specs including 131 juveniles (see App. Table 2C): Port Fouad-Km 12: SLC 45, 1 spec.; el Qantara-Km 45: SLC 6, 197 specs (1 juvenile); SLC 20, 5 specs; SLC 31, 9 specs (3 of them juveniles); SLC 32, 1 spec.; SLC 38, 1 spec.; Lake Timsah , Km 78, opposite Isma’iliya: SLC 4, 2 specs (juveniles); SBE 5, 2 specs; SBE 7, 1 spec.; SBE 8, 39 specs; Great Bitter Lake , east of Deversoir-Km 97: SLC 50, 18 specs (3 of them juveniles); SLC 52, 1 spec. (juvenile); SLC 55, 2 specs (juveniles); SLC 60, 6 specs; SLC 61, 6 specs (5 juvenile); SLC 64, 4 specs; SLC 67, 1 spec. (juvenile); SLC 71, 4 specs; SLC 74, 2 specs (1 of them juvenile); SLC 76, 1 spec.; Km 98: SLC 3; 6 specs; SLC 117, 175 specs (109 juveniles, spot-checked); SLC 123, 2 specs (juveniles); Little Bitter Lake opp. Kabrit-Km 120: SLC 77, 1 spec.; SLC 85, 4 specs (1 juvenile); eastern shore rocky promontory: SBE 1, 1 spec.; Tis’a-Km 149: SLC 2, 1 spec. — Great Bitter Lake “Yellow Fleet” Biofouling Samples, January 13–20, 1975: 24 +?1 subsamples, 33 +?1 specs (13 of them juveniles, see App. Table 2D).— Lake Timsah , legit S.H. Shalla ca. 1984, det. H.A. ten Hove, ZMA V.Pol. 4997 (juveniles), 5000, 5001, 5002, altogether 17 specs, and juveniles (not enumerated).— El Tawan Beach, legit S.B. Shazly 10 / 19.XI.1988, det. H.A. ten Hove, ZMA V.Pol. 3817, 4 specs (App. Table 2E) .

Locations adjacent to the Suez Canal, Red Sea side: Gulf of Suez , Suez, in crevice of stone on shore, legit C. Crossland 1904–1905, det. H. Zibrowius, confirmed H.A. ten Hove, BM ( NH) ZB 1972 : 22, 1 spec. — Cambridge Expedition 6.X–29.XII.1924, 1 “new” BM ( NH) sample from barge: Presumed Port Taufiq , 1 spec. (see App. Table 2A). — Gulf of Aqaba: Israel, Elat, 13 samples, 0.5– [6–8]–[40–46]– [81–90] m, HUJ .

Suez Canal Depth and Substrates: 0.2– 10 m; on algae: Cystoseira myrica , Digenea , Laurencia , Sargassum ; on sponge; on the gastropod, Murex forskoehli ; on bivalves: Brachidontes pharaonis , Chama gryphoides , Chicoreus erythraeus , Crenatula picta , Fulvia fragilis , Fusinus verrucosus , Malvufundus normalis , M. regulus , Pinctada radiata , Spondylus spinosus , Pectinidae (not designated); on bryozoans; barnacles; crabs; tunicates; under rocks; on canal walls; on artificial substrates, e.g., on tin can submerged in mud, rubber fenders and iron frames.

Colouration. Lake Timsah field notes describe radioles with 5 red-orange stripes.

Distribution. Worldwide in (sub)tropical to temperate regions; port fouling species. Mediterranean: Israel, Egypt.

Remarks. Hydroides elegans ’ area of origin is believed to be Australia, its type locality ( Zibrowius 1994). It was present in an 1888 harbour-fouling sample from Naples (RMNH 932; ZMA V.Pol. 3205). Its first published Mediterranean record, also from Naples, was by LoBianco (1892: 85) under the synonymous name H. pectinatus ( Philippi, 1844) . The first Levant Basin reference for Hydroides elegans (as H. norvegica ), from 1924, was the specimen collected by the Cambridge Expedition from a buoy in Port Said ( Potts 1928). All publications from the Suez Canal report it as the dominant fouling serpulid (see synonymy section, above). Hydroides elegans occurs worldwide in subtropical to temperate regions (northern and southern hemisphere), as opposed to the superficially similar congener, H. norvegicus Gunnerus, 1768 that has a mainly boreal and deeper Mediterranean distribution (see Zibrowius 1971: 717–721, 1973a: 683; ten Hove 1974: 46).

As apparent from the above list of regional synonyms, in recent years, the citations in the fouling literature of this area show increasing awareness that the principle fouling species is Hydroides elegans rather than H. norvegicus and many of the erroneous citations of H. “ norvegicus ” have been corrected retroactively. However, the references to Serpula sp. from these same fouling samples / same habitat (or of “ S. concharum ”, or “ S. vermicularis , mainly boreal species) suggest that individuals with primary (immature) opercula have been and are probably still being confused for species of the genus Serpula rather than being recognized as immature ontogenetic stages of Hydroides , presumably of H. elegans (see Moran 1984; ten Hove & Ben-Eliahu 2005 for a review of the relevant ontogenetic history). Fortunately, H. elegans has rather distinctive collar chaetae ( Figs 6C , 7C ), and by spot-checking the collar chaetae, e.g., in sample SLC 117 and in the biofouling samples, it was possible to confirm that most of the juveniles in the present samples belonged to this species ( H. elegans specimens listed as juvenile in App. Table 2A, C–E lack the upper verticil, but the characteristic collar chaetae showed them to belong to this species). The tube of H. elegans is characteristically smooth, without peristomes; flanges or prominent transversal ridges, usually with a narrow flattened area on the dorsal (upper) surface bordered by low smooth, not pronounced longitudinal ridges with a rounded transition to the sides of the tube (see Figs 5E, 6D, E ; 7J–K ). However, individuals with two prominent longitudinal ridges may also found (e.g., Fig. 5D), with the part of the tube between them appearing to be sunken. This generally confirms the finding of Bastida-Zavala & ten Hove (2003b: 86). In minute tubes, a barely perceptible median “line” can be distinguished. Many tubes are sufficiently characteristic as to enable confident attribution to this species (e.g., Figs 7J, K ; see App. Table 2).

Figures

FIGURE 6. The very first serpulid collected from the Suez Canal—Hydroides elegans—from a boat hull, collected by the “Pola” Expedition (Isma’iliya, 17.X.1895, coll. Mus. Vindobonensis, Vienna, Evertebr. Varia, inv. no. 2034 [alte invert. no. 17199]), det. H. Zibrowius. A—Operculum photographed in methylene-blue-glycerine; focus on spines of upper verticil, B—Same, focusing on long tooth arising from the central disk of the verticil, C—Collar chaetae showing characteristic fine teeth and proximal rasp, D, E—Aggregates of tubes on gastropod, Potamides conicus; note the slightly flattened upper surface of the tube. Scales: A, B—100 µm, C—50 µm, D, E—1 mm.

FIGURE 7. Taphonomic residues of Hydroides elegans from mollusc shells collected in the Great Bitter Lake in 1950 by C. Beets. The chitinous residues “survived” more than 50 years in the dried serpulid tubes. A—Body of dried worm, B—Thoracic chaetae, C—Broken but distinctive collar chaetae, D—Residue of verticil, E—Fragment of tube with adhering fascicles of thoracic chaetae (putative identification of species based on tube), F—Thoracic uncini (putative identification to species based on tube), G—Operculum, apical view of verticil, H—Operculum, ¾ view. I. Tube with projecting verticil, lower side of tube. J. Hydroides elegans tube fragment on shell of Gafrarium pectinatum, K—Fusinus polygonoides shell (ca. 11 cm in length), bearing tubes of H. elegans. Scales: A, B, D, E, I, J—1 mm; G, H—100 µm; C, F—50 µm. Figs A, B, D–F, I, K—from Beets’ Stn 4, 0.2–0.8 m, C, G —from Stn 21, 1.5–3 m, H—from Stn 1: 1–2.7 m, J—from Stn 19: 5–5.5 m (for stations, see Beets (1953), Appendix Table 2B).