taxonID	type	description	language	source
396387E75F4BE01BFF50FE53FC62F9C6.taxon	materials_examined	Material examined. Locations adjacent to the Suez Canal, Mediterranean side: Israel: 10 samples, HUJ, depth ca. 0.3 m. Suez Canal material reported herein: 2 samples, 3 specs. Hebrew University-Smithsonian Expedition, Lake Timsah: SBE 7, 2 specs — Little Bitter Lake: SBE 1, 1 spec. Locations adjacent to the Suez Canal: Landlocked, Egypt: Lake Qârün, legit G. N. El Shabrawy, det. H. A. ten Hove 2000, ZMA V. Pol. 4981, 10 specs. Locations adjacent to the Suez Canal, Red Sea side: None. Suez Canal depth and substrates: 0.3 m, shallow; under rocks; encrustation on tin can. Colouration. Specimens from Lake Timsah with dark brown body; branchia of specimen from Bitter Lake with 6 rows of dark pigment.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F4BE01BFF50FE53FC62F9C6.taxon	distribution	Distribution. Worldwide in subtropical to temperate regions, northern and southern hemisphere; brackish. Mediterranean: Israel, Egypt.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F4BE01BFF50FE53FC62F9C6.taxon	discussion	Remarks. A new record for the Suez Canal dating from II. 1973. A mixo-hyperhaline brackish species (ten Hove & Weerdenburg 1978, ten Hove & van den Hurk 1993).	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F4BE01BFF50F9E6FB48F93B.taxon	materials_examined	Type locality. England, Weymouth. Not present in the Suez Canal, but see Salmacina.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F4BE019FF50F883FE3EFB2C.taxon	materials_examined	Material examined. More than 16 samples at the AMNH, USNM and YPM, with> 150 specimens, e. g., the following six samples: U. S. A., Connecticut, off New Haven, 11 – 15 m, syntypes YPM 2698, 2699, 8 specs — Massachusetts, New Bedford, from piles under wharf, legit R. Hall & A. Elwyn 2. VIII. 1909, det. W. G. van Name, redet. H. A. ten Hove; this H. dianthus is a “ reef ” (a solid aggregation of cemented tubes), AMNH 1122 (not counted, but at least 11 specimens). — New Jersey, Greater Egg Harbour, legit, det. H. E. Webster (1880: 128, 159), transferred by M. Pettibone to H. uncinata, redet. H. Zibrowius 1970 H. dianthus, USNM 381, 1 spec. — Florida, Alligator Harbour, Wilson’s Beach, Stn 22, Tall Timbers, legit J. Rudloe 25. III. 1966, det. H. A. ten Hove 1971 (unpubl.), USNM 51862, 2 specs. Mediterranean, Naples, Gulf, Zoological Station 1888, separated from Hydroides diramphus, RMNH 392, ZMA V. Pol. 3206, 1 spec. — without further data, det. K. J. Bush (1910: 498) Eupomatus uncinatus, redet. H. A. ten Hove 1985, YPM 2839, 10 dried specs. Locations adjacent to the Suez Canal, Mediterranean side: Cyprus, Limassol Harbour entrance, Cy- AI, ca. 0.3 m depth, scuba, from a ship propeller, legit G. Payiatas & P. Orfanou 27. X. 1997, det. M. N. Ben- Eliahu, HUJ-Poly- 80, 2 specs. Suez Canal proper: No previous records from the Suez Canal. Locations adjacent to the Suez Canal, Red Sea side: No records	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F4BE019FF50F883FE3EFB2C.taxon	distribution	Distribution. Temperate to subtropical coasts of northeast America and western Europe (not north of the English Channel). Atlantic Africa. Mediterranean: Italy, Naples; Cyprus (on ship propeller); Turkey, Aegean; Egypt, Alexandria, eastern harbour, Port Said, Suez Canal (Lake Timsah). Japan, Tokyo Bay.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F4BE019FF50F883FE3EFB2C.taxon	discussion	Remarks. Hydroides dianthus apparently originates from the Atlantic coast of North America and is common in biofouling along the U. S. western Atlantic seaboard (Zibrowius 1971: 704, 1973 a: 684). Contrary to the situation in the western Atlantic, there seemed to be no extensive populations in the eastern Atlantic and in the Mediterranean, except those in some Mediterranean lagoons (e. g., Thau lagoon). In the Mediterranean, Hydroides dianthus also occasionally occurs in harbours (H. Zibrowius, pers. comm.). The distribution pattern, conjunct in the western Atlantic and disjunct elsewhere, and the larger reservoir along the U. S. coast, supports the diagnosis of the western Atlantic as the provenance of this species (H. Zibrowius, pers. comm.). Zibrowius (1971) characterized it as tolerating a wide range of salinities and temperatures, between 28 – 50 ‰ and 5 – 30 ° C. Its first record in the eastern Mediterranean (Aegean, near Izmir) was in 1865 (Zibrowius 1973 a). Knight-Jones et al. (1991: 841) reported it from the only station in the Bay of Izmir with eutrophic conditions (turbid waters and rocks covered with Ulva and Mytilus). A first record from Cyprus (X. 1997) was that of two decaying individuals removed from a propeller of a ship in Limassol Harbour (Ben-Eliahu & Payiatas 1999), one an operculum without a body; this record does not provide a definitive proof of settlement in Cyprian waters. Hydroides dianthus has not been reported along the Mediterranean coast of Israel (M. N. Ben- Eliahu, unpubl. data); the specimen erroneously published long ago as H. cf. dianthus (Ben-Eliahu 1976: 106 – 107, fig. 1, and Ben-Eliahu & Safriel 1982: 387) is corrected herein to Hydroides operculatus, a Lessepsian migrant (see also Çinar 2006: 229 – 230, fig. 6). Moreover, Zibrowius & Bitar’s (1981) report of Hydroides dianthus from Lebanon was regarded as questionable (Ben-Eliahu & ten Hove 1992: 37); the two small Lebanese specimens, when re-examined by one of us (H. A. ten Hove) in 1998 appeared rather to belong to Hydroides cf. brachyacanthus Rioja, 1941, a Lessepsian migrant present on the coasts of Israel and Lebanon as well as the Levant coast of Turkey (Ben-Eliahu & ten Hove 1992: 43, Zibrowius & Bitar 2003, Çinar 2006: 225 – 226, fig. 2, respectively). Selim's (1988) first record of Hydroides dianthus from the southern Levant coast (Eastern Harbour, Alexandria), and a later record from Port Said, not within the canal proper, were documented by a well-illustrated operculum (Selim 1997 b fig. 2 a). However, Selim’s recent record (2009: 78) of Hydroides dianthus from within the canal, ca. 37 km from its northern mouth, from El Cap, referred to Ben-Eliahu's erroneous record of Hydroides cf. dianthus illustrated with a figure of the H. operculatus operculum (Ben-Eliahu 1976: 106, fig. 1), and the other references cited (Fauvel 1927: 357 – 358, Bellan 1964: 174 – 175 and Zibrowius 1968: 109 – 112) referred not to Hydroides dianthus, but rather to H. uncinatus, confused for it (reviewed in Zibrowius 1971). The verticil spines of Hydroides operculatus, a well known Lessepsian migrant, all point inwards (Çinar 2006: 229 – 230, fig. 6), whereas the spines in H. dianthus all point in the same (ventral) direction (Zibrowius, 1973 a: 683). Despite the uncertainty created by Selim’s use of these citations and our inability to review a voucher specimen, given Selim’s (1997) illustration, we tend to hypothesize that the identification is correct. If that is true, the record is also presumed to be a first record of Hydroides dianthus from within the Suez Canal. This could be indicative of a recent Atlantic-Mediterranean incursion into the canal, a finding with biogeographic significance. However, its disjunctive distribution — it is lacking along the northward Israeli coast but present in the Egyptian ports of Alexandria and Port Said — suggests it could rather be indicative of ship-transport as has been observed in Hydroides elegans and Hydroides diramphus. This is supported by a recent article by Link et al. (2009) that reported its first occurrence in East Asia, Tokyo Bay on artificial hard substrates, presumably due to ship transport, and a subsequent report from Osaka Bay by Otani & Yamanishi (2010). In the Discussion section below, we have enumerated Hydroides dianthus as a member of the Suez Canal serpulid fauna.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F49E006FF50FA95FDE1FE58.taxon	materials_examined	Material examined. Locations adjacent to the Suez Canal, Mediterranean side: None. Suez Canal proper: No records. Locations adjacent to the Suez Canal, Red Sea side: Egypt: 2 lots. Gulf of Suez, Suez, C. Crossland, details concerning substrate or habitat not available, det. H. L. M. Pixell Eupomatus spinosus, BM (NH) 1924: 6: 13: 135, 2 specs; syntypes of Eupomatus spinosus Pixell, 1913, donated J. S. Gardiner, BM (NH) 1938: 7: 25: 9 – 12, 4 specs. South Africa, legit W. Stephensen 11. VI. 1932, det. H. Zibrowius 1972, Hydroides spinosus, BM (NH) 1932.12.23.20, 1 spec.; Cape Peninsula, shore, Stn CP 138, legit, presented and det. J. H. Day H. dipoma, BM (NH) 1963.1.175, 1 spec.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F49E006FF50FA95FDE1FE58.taxon	distribution	Distribution. Atlantic Africa: Río de Oro, Morocco to the Cape of Good Hope; Suez.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F49E006FF50FA95FDE1FE58.taxon	discussion	Remarks. Pixell (1913: 78) reported Hydroides dipoma (as Eupomatus spinosus) from Suez. Zibrowius (1973 b: 36) commented on the species’ disjunctive distribution, from the length of the south east Atlantic coast, without additional records from the Mediterranean or from the Indian Ocean or Red Sea; and with a single Indo-West-Pacific location, Suez. We agree with him that it is likely that the record at Suez is due to a ship-borne introduction. We have not enumerated Hydroides dipoma among the species reported from the Canal. However, due to the proximity of Suez to the southern opening of the canal, we would not be surprised if it were to be found in the canal.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F56E004FF50FE61FC24FD1E.taxon	description	Figs 4, 5 A – C	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F56E004FF50FE61FC24FD1E.taxon	materials_examined	Material examined. Locations adjacent to the Suez Canal, Mediterranean side: Israel: 35 samples; one of these is the first sample from Israel, Haifa harbour, legit G. Haas, 22. III. 1937, det. M. N. Ben-Eliahu, HUJ- Poly- 867 (previously AN-II- 84) — Egypt, Sinai: 4 samples, 0.3 – 4 – 15 – [57 – 62] m, HUJ. — Egypt, Port Said, on buoy no. 723, Cambridge Expedition 13. XII. 1924, det. F. A. Potts Hydroides lunulifera, redet. H. Zibrowius (1971: 706), confirmed M. N. Ben-Eliahu 1986, CUZM-AN. I. 1930, 1 spec. Suez Canal material reported herein: 13 samples, 100 specs, Cambridge Expedition 6. X – 29. XII. 1924, 1 “ new ” BM (NH) sample: Toussoum: 1 spec. (see App. Table 2 A, and see just below). — Beets’ Great Bitter Lake samples: ca. 6 samples with empty tubes tentatively attributed to Hydroides diramphus (not enumerated, see App. Table 2 B). — Hebrew University-Smithsonian Expeditions, 1967 – 1973: 6 samples, 81 specs. Lake Timsah: SBE 7, 30 specs; SBE 8, 44 specs; Great Bitter Lake: SLC 67, 1 spec.; SLC 71, 1 spec.; SLC 3, 2 specs; SLC 117, 3 specs. — Great Bitter Lake “ Yellow Fleet ” Biofouling Samples, January 13 – 20, 1975: 3 subsamples, 3 specs; Bremerhaven dry dock, 18. VI. 1975, 7 m, 1 spec. with operculum still connected to partially decayed body — Lake Timsah, 1984, legit S. H. Shalla, det. H. A. ten Hove, ZMA V. Pol. 5001, 8 specs, tubes — El Tawan Beach, 10 / 19. XI. 1988, legit S. B. Shazly, det. H. A. ten Hove, ZMA V. Pol. 3818, 3 specs. Locations adjacent to the Suez Canal, Red Sea side, reported herein: Cambridge Expedition 6. X – 29. XII. 1924, 1 “ new ” BM (NH) sample from barge: Presumed Port Taufiq, 1 spec. (see App. Table 2 A). Suez Canal depth and substrates: 0.3 – 10 m; on algae, e. g., Digenea; on sponge, on molluscs; on a mollusc from a barge; on bivalves, Brachidontes pharaoni s, Crenatula picta, Pinctada radiata; on rocks; on artificial substrates: Encrusted tin can submerged in mud, rubber fenders and iron frames; ship fouling. Colouration. Radioles of specimens from Lake Timsah with alternating white and brown banding, widest bands in middle of radiole.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F56E004FF50FE61FC24FD1E.taxon	distribution	Distribution. Circum (sub) tropical; port fouling species. E. Mediterranean. Turkey, Lebanon, Israel, Egypt.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F56E004FF50FE61FC24FD1E.taxon	discussion	Remarks. Origin apparently tropical western Atlantic Coast; Zibrowius (1973 a) noted a first record of Hydroides diramphus from Naples in 1870 as its synonym, H. lunulifer (Claparède, 1870: 181 – 182 pl. 31, fig. 3). Its first record from the Levant Basin, from 1924, was from the sample collected by the Cambridge Expedition from a buoy in Port Said (Potts 1928: 701). Potts considered that Hydroides diramphus had not entered the Canal. However, in actuality, two additional samples of Hydroides diramphus had been collected, unknown to Potts, encrusted on the Cambridge Expedition molluscs that had been taken to the Natural History Museum in London and deposited in the Mollusc Section of the museum; they were found 62 years later during a visit to the Museum (by NBE) and are referred to above in the “ Material examined ” section as “ new ” Cambridge Expedition samples. One was from Toussoum (km 87) within the canal and the other from “ Barge 568 ”, presumably from Port Taufiq. [Potts referred only to “ barges ” without specifying their number but specifically mentioned only Port Taufiq as a location for them; the precise date of collection is unknown (J. Pickering, pers. comm., see App. Table 2 A)]. The first record of H. diramphus from within the Suez Canal is herein brought forward to 1924. It is likely that it was present among the dry Great Bitter Lake tubes collected later by C. Beets (1950); see Discussion and App. Table 2 B, but taphonomic residues, enabling confirmation of our tentative identification of tubes, were not found. In 1973, H. diramphus appeared to be present in larger numbers in Lake Timsah than in the Great Bitter Lake, and there have been consistent records of it there in subsequent Egyptian fouling research (e. g., Ghobashy 1984, Shalla 1985, Ghobashy et al. 1986, Ghobashy et al. 1990 and Mona 1992). Selim (1997 b) reported it in the five sites studied along the canal, i. e., in Port Said, El-Kab, Lake Timsah, Deversoir and Port Taufiq. Most of the present specimens had expanded T- or anchorshaped tips of the verticil spines and pointed chitinised tips of the funnel radii (Fig. 4). Bastida-Zavala & ten Hove (2003 b: 84) described tubes of Hydroides diramphus from California and Hawaii as variable in form “ having transversal ridges also with longitudinal ridges ” … and sometimes with peristomes. Variability in form is also confirmed for the present material; tubes with three ridges appeared to be rarer than those with two, although the median ridge may be very difficult to see. Fig. 5 A – C illustrates some variation in tube ornamentation in Hydroides diramphus. The tube in Fig. 5 A is less rugose than the one in Fig. 5 B that appears to be covered with a granular layer that could obscure surface sculpturing (i. e., could obscure a low median longitudinal ridge). Several tubes from the Mediterranean coast of Israel had three ridges so low that they could barely be seen with the water surface breaking at the level of the tube surface (these ridges were even lower than the median ridge figured in Fig. 5 A).	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F54E001FF50FCBBFD50F837.taxon	description	Figs 5 D, E, 6, 7	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F54E001FF50FCBBFD50F837.taxon	description	Suez Canal	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F54E001FF50FCBBFD50F837.taxon	description	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].	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F54E001FF50FCBBFD50F837.taxon	materials_examined	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. 6 B) and juveniles with only primary operculum (collar chaetae checked); km 72, north of Isma’iliya, 4. XII. 1924, Little Bitter Lake, K 2, (Kabrit), 17. X. 1924, with long central spine; K 9, (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 2 A]), 3 specs: Lake Timsah-Km 78, Isma’iliya: 1 spec.; T 9: 1 spec.; 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 2 B, Fig. 7). — Hebrew University-Smithsonian Expeditions, 1967 – 1973, 27 samples, 493 specs including 131 juveniles (see App. Table 2 C): 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 2 D). — 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 2 E). 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 2 A). — 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.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F54E001FF50FCBBFD50F837.taxon	distribution	Distribution. Worldwide in (sub) tropical to temperate regions; port fouling species. Mediterranean: Israel, Egypt.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F54E001FF50FCBBFD50F837.taxon	discussion	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, 1973 a: 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 6 C, 7 C), 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 2 A, 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 5 E, 6 D, E; 7 J – K). However, individuals with two prominent longitudinal ridges may also found (e. g., Fig. 5 D), with the part of the tube between them appearing to be sunken. This generally confirms the finding of Bastida-Zavala & ten Hove (2003 b: 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 7 J, K; see App. Table 2).	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F5FE00BFF50FF1BFB64FD0A.taxon	description	Figs 5 F, G, 8 – 10, presumably also Fig. 11	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F5FE00BFF50FF1BFB64FD0A.taxon	materials_examined	Material examined (Figs 8 – 10, presumably also Fig. 11): Locations adjacent to the Suez Canal, Mediterranean side: Israel: 20 samples, [1] – [8 – 10] – [18 – 22] – 33 m (830 m outlier record); INCNH unpublished data, first Israeli sample, Haifa Port, legit G. Haas, 22. III. 1937, det. M. N. Ben-Eliahu, HUJ-Poly- 878 (previously AN-II- 05). Suez Canal proper: Cambridge Expedition, El Qantara, det. F. A. Potts, CUZM-AN. I. 1930, 1 spec., verticil with taller dorsal spine + 6 spines with lateral spinules, processes with lateral spinules positioned rather high (subdistally); 32 radii in funnel, 21 radioles, ca. 22 mm. Suez Canal material reported herein: 35 samples with 106 +? 4 specs of which 4 belong to juvenile forms 1 and 2: One “ new ” Cambridge Expedition sample separated from a mollusc deposited at BM (NH); this single juvenile specimen preliminary named “ form no. 1 ” (Fig. 9) proved to be a Hydroides heterocerus juvenile (precise location in the canal not known [see App. Table 2 A]). — Hebrew University-Smithsonian Expeditions, 1967 – 1973: 4 samples, 4 specs of which two were the juvenile forms named nos. 1 and 2. Great Bitter Lake, east of Deversoir-Km 97: SLC 72, 1 spec.; SLC 74, 1 spec., juvenile form no. 1. Great Bitter Lake: SLC 117, 1 spec., juvenile form no. 2 (Fig. 10). SLC 123, 1 spec. — Great Bitter Lake “ Yellow Fleet ” Biofouling Samples, January 13 – 20, 1975: 25 +? 1 subsamples, 90 (1 juvenile) +? 4 specs. — Bremerhaven dry dock, 18. VI. 1975, 3 subsamples, 11 specs, all decaying; one notably large tube, ca. 4 mm outer diameter; worm without operculum 4.7 cm (branchia 6 mm) / 3 mm width. — Lake Timsah, El Tawan Beach, legit S. B. Shazly 10 / 19. XI. 1988, ZMA V. Pol. 3819, 1 spec. Locations adjacent to the Suez Canal, Red Sea side: Gulf of Suez, J. K. Lord Expedition of 1870, det. H. A. ten Hove 1969 (separated from a mixed sample BM (NH) 1870: 12: 23: 31 by HAtH 1969, see Material Examined S. tetraceros), BM (NH) 1870: 12: 23: 61, specimen bioperculate, body rotted with epidermis abraded. — Gulf of Suez, Suez Quay, 3.7 – 18.3 m, legit J. S. Gardiner 1905 (the Percy Sladen Trust Expedition), det. H. L. M. Pixell (1913, Pl. 8, figs 2 a – c), confirmed M. N. Ben-Eliahu, several specs. —? Gulf of Suez, Suez, Stn R 5, Cambridge Expedition, 1924, CUZM AN. I. 1930, empty coiled tube, present in test-tube with Serpula vermicularis not Linnaeus, sensu Potts (1928: 700), presumably tube of Hydroides heterocerus (Fig. 11). Gulf of Aqaba: No records. Red Sea: Sudan, Suakin Harbour, legit C. Crossland 1904 – 1905, det. H. L. M. Pixell, H. Zibrowius, BM (NH) 1972.23, tube lacking, permanent mount dated 20. VII. 1904. — Eritrea, Dahlak Archipelago, Musseri Island, 15 ° 37 ’ N, 40 ° 43 ’ E, 27.5 m, Israel South Red Sea Expedition 1965, Stn 10, legit Ch. Lewinsohn 23. X. 1965, det. F. Rullier, TAU-NS- 1873, 1 spec. Gulf of Aden: Somalia, Ras Katib, outside harbour on a block of dead coral, legit, det. T. G. Pillai 29. III. 1982, ZMA V. Pol. 3851, 2 specs. Suez Canal depth and substrates: Shallow, 1 – 2.5 m, 10 m; on sponges; on bivalves, Brachidontes pharaonis and Spondylus spinosus; on barnacles, bryozoans, tunicates; on sandstone; on artificial substrates: Rubber fenders and iron frames.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F5FE00BFF50FF1BFB64FD0A.taxon	distribution	Distribution. Indo-West Pacific: Red Sea, Zanzibar, Madagascar, Persian (Arabian) Gulf, Sri Lanka,? New Caledonia. Lessepsian migrant to the Mediterranean: Israel, Lebanon, Cyprus, Turkey.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F5FE00BFF50FF1BFB64FD0A.taxon	discussion	Remarks. Potts (1928: 700 – 701) noted: “ In one specimen, lateral processes of spines were 1 / 3 length from the apical process, in the other, just under the apical process. Other authors have shown that there is a great deal of variation in their position. ” A non-quantitative survey of the present material showed that most of the lateral spinules were positioned close to the apical process (e. g., Fig. 8 A) and there appeared to be some variability in height of the spines in different individuals (a more thorough analysis of these characters is intended). Much more pronounced is the variability in the form of the marginal radii of the opercular funnel, ranging from “ bowling-pin-shaped ” radii with blunt or even stubby tips (Wesenberg-Lund 1949 and Ishaq & Mustaquim 1996, respectively; Fig. 8 E) to those with more developed side spinules as in Fig. 8 D, “ T ” or “ anchor-shape ” tips (Pixell 1913, Zibrowius 1971; Fig. 8 F), with intermediate forms between them. Specimens from the Sudanese Red Sea (Fig. 8 F) had much more developed side spinules than those in the Suez Canal population where the most developed of the side spinules were like those of Fig. 8 D; and less like Fig. 8 E. The number of verticil spines with lateral spinules in the both the Suez Canal population and the Lessepsian migrant population ranged from 5 – 7, 5.59 ± 0.53 spines, N = 59, Suez Canal and 5.78 ± 0.64, N = 27, the Lessepsian migrant population, respectively, the differences between them were not significant (T-test, p = 0.198). Çinar (2006 fig. 4 a, b) shows the tips of the marginal radii of the Lessepsian migrant from the Levant coast of Turkey to be like those of the Suez Canal population (Fig. 8 E). Specimens removed from the “ Münsterland ” in the Great Bitter Lake in January 1975 were large: The largest, 63 mm; verticil with 1 + 6 spines; funnel with anchor-tip to rounded marginal radii. However, the material sampled from the “ Münsterland ” in the Bremerhaven dry dock (in June 1975, 6 months later) gave the impression of even larger size (one tube had an outer diameter of 4 mm; the tubes were the largest of any taxon present in the assemblage) and many tubes were coiled. Most of the Hydroides heterocerus tubes from the Suez Canal were rounded (anterior end not attached to substrate) with marked transversal ridges; only 3 showed three longitudinal ridges, most marked in a small specimen (Fig. 5 G), less marked in the two others. We believe that the large coiled tube found in Potts’ (1928) nominal Serpula “ vermicularis ” sample from the Gulf of Suez rather belongs to H. heterocerus (Fig. 11). Lessepsian migrant H. heterocerus specimens had marginal teeth like Figs 8 D or E, or forms ranging between them. Juvenile specimens: The operculum of the juvenile form from the Cambridge Expedition (form we named, “ no. 1 ”, App. Table 2 A), had sharply pointed radii tips in the funnel (Figs 9 B, C), a marked deviation from the species description. A second specimen was obtained when searching through Brattström and Taasen’s biofouling material from the Great Bitter Lake. We attribute form “ no. 1 ” to Hydroides heterocerus, taking note of the variability in form that can prevail during Hydroides ontogeny (see ten Hove & Ben-Eliahu 2005). A single juvenile specimen, “ form no. 2 ”, was similarly found in the biofouling samples. In contrast, the funnel of this specimen had expanded T- or anchor-shaped marginal teeth; some of its verticil spines were characteristic for Hydroides heterocerus, while others appeared to be deviant, malformed (Figs 10 A, B). We attribute this specimen to Hydroides heterocerus as well. In studying the Hydroides heterocerus population from the Great Bitter Lake, we encountered additional specimens with malformations such as a lateral spinule on the dorsal verticil spine, or a stub of a verticil spine rather than a fully developed one.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F59E034FF50FF1BFAC7FC80.taxon	description	Figs 5 H, I, 12	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F59E034FF50FF1BFAC7FC80.taxon	description	(1968) Hydroides homocera: Ben-Eliahu 1991 b: 526, fig. 5 [Gulf of Suez, El Bilayim lagoon, first record from Gulf of Suez, collected 10. VI. 1968, sample SLR 1753 (Por et al. 1972); on Pectinidae. Marginal radii of funnel single anchor-shaped]; Selim 1997 b: 93 – 94, figs 6 a – e [Gulf of Suez, Port Taufiq, collected in 1988]. Red Sea, proper-Indo-West-Pacific (excluding citations from Gulf of Suez and Gulf of Aqaba already given above)	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F59E034FF50FF1BFAC7FC80.taxon	materials_examined	Material examined. Locations adjacent to the Suez Canal, Mediterranean side: Israel: 14 samples, 22 specs. First Mediterranean record of this Lessepsian migrant, INCNH data unpublished, Haifa Bay, 32 m, on Mimachlamys varia, legit E. Gilat (Gilat 91) 21. X. 1955, det. M. N. Ben-Eliahu ca. 1969, marginal radii singleanchor-shaped as figured by Wesenberg-Lund (1949, fig. 46 a), see Fig. 12 E, TAU-NS (no number), 1 spec. — Egypt, Sinai: 2 samples. Depth: 9 – [22 – 32] – 55 m. Suez Canal material reported herein: 8 samples with 38 specs. Beets’ Great Bitter Lake samples, no taphonomic residues found, tubes provisionally identified as those of H. homoceros in at least 4 samples — Hebrew University-Smithsonian Expeditions, 1967 – 1973, 3 samples, 33 specs: Great Bitter Lake, SLC 117, 31 specs; east of Deversoir, Km 97, SLC 50, 1 spec.; SLC 52, 1 spec., marginal radii of funnel single-anchorshaped, see Figs 12 A, D. — Great Bitter Lake “ Yellow Fleet ” Biofouling Samples, January 13 – 20, 1975: 4 subsamples, 4 specs. Bremerhaven dry dock, 1 spec. Locations adjacent to the Suez Canal, Red Sea side: Gulf of Suez, 1 sample. — Gulf of Aqaba: No records. Red Sea: South Red Sea: 2 samples (? m, 36.6 m). Sudan, Suakin Harbour, legit C. Crossland 7. II. 1905, BM (NH) 1924.6.13.145, 1 spec., syntype, marginal radii with 2 pairs of recurved lateral processes, i. e., “ double-anchor-shaped ” (Pixell, 1913, pl. 8 fig. 1 a as in Fig. 12 C). — Eritrea, Dahlak Archipelago, 15 ° 35 ’ N, 40 ° 44 ’, 36.6 m, on Malleidae, legit Ch. Lewinsohn, 23. X. 1965, Israel South Red Sea Expedition, 19085 Stn 12, det. M. N. Ben-Eliahu ca. 1985, marginal radii of funnel double-anchor-shaped, as in Fig. 12 C. Indian Ocean, Tanzania, off Zanzibar, scraped from the bottom of the “ Juba, ” legit C. Crossland, 1901 – 1902, 4 syntypes and slide of collar chaetae and uncini, BM (NH) 1924.6.13.147; although Pixell did not refer to variation in the form of the opercular spines, marginal radii of syntypes range from spur-tipped singleanchor to a more developed spur approaching double-anchor form (Pixell, 1913, pl. 8 fig. 1 a), i. e., a range in form between Figs 12 B to C (present paper). Arabian Sea: South Arabian Coast, 13.5 m, John Murray stn 53, 2. XI. 1933, det. C. C. A. Monro, BM (NH) 1937.9.2.540 – 1, 2 specs, marginal radii as in Pixell (1913), and as in Figs 12 B to C. — Oman, Gulf of Masirah, Masirah Island, Ras Al Ya; low tide, under stones, Stn 91 / 105, legit R. G. Moolenbeek & H. Dekker, 23. XI. 1991, ZMA V. Pol. 3838, marginal radii with spur-tipped single-anchor, see Fig. 12 B. Gulf of Oman, N. Oman, Khor al Quway, north-south running strait, gentle current, from east side of strait, 18.3 – 36.6 m, Royal Geographical Society (of Great Britain) Musandam Expedition, 1971 – 72, legit P. E. S. Cornelius; extracted and det. by H. Zibrowius 1972 from dead coral fragments from sandy bottom and limestone area, on coelenterates, BM (NH) 1972: 217, marginal radii double-anchor-shaped, as in Fig. 12 C. Strait of Hormuz, 5 miles S. E. of the Tunb light, 26 ° 12 ' N, 55 ° 22 ' E, 38 – 60 m, Petersen-grab, 7. IV. 1938, gravel and shells, Danish Expedition Stn 118, legit B. Løppenthin, det. E. Wesenberg-Lund (1949: 380 – 381), redet. H. A. ten Hove II. 2000, ZMUC, marginal radii of funnel single-anchor-shaped, see Figs 12 A, D, E. Persian (Arabian) Gulf. Sample details unknown, legit Ø. Støckland ca. 2004, det. M. N. Ben-Eliahu XI. 2004, marginal radii with single pair of lateral processes topped by a more or less expanded tip (cf., ten Hove 1970 a, figs 1 a – c), similarly from the Persian (Arabian) Gulf, Fig. 12 B. — Bahrain, 4 miles E. N. E. of Bahrain light-ship, Stn 87, m?, legit B. Løppenthin 20. III. 1928, ZMA V. Pol. 3002, marginal radii with one pair of recurved lateral pinnules, surmounted by spur (as in ten Hove 1970 a, figs 1 a – c), more recurved, i. e., more anchor-shaped than those in Fig. 12 B. Suez Canal depth and substrates: Shallow – 10 m, on algae: Digenea; sponges; on bivalve, Spondylus spinosus; on tunicate.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F59E034FF50FF1BFAC7FC80.taxon	distribution	Distribution. Lessepsian migrant to the Mediterranean: Israel, Turkey; Suez Canal; Red Sea: Gulf of Suez, Dahlak Archipelago; Indian Ocean: Zanzibar, Seychelles, South Arabian coast, Persian (Arabian) Gulf, Maldives. May occur in living corals (Millepora, Porites).	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F59E034FF50FF1BFAC7FC80.taxon	discussion	Remarks. Tube with 3 prominent longitudinal ridges of equal height; transversal ridges not prominent, giving the tube relatively smooth sides (ten Hove 1970 a: 55, figs 1 – 8; Figs 5 H – I). Very slight pink cast in some tubes. Of ca. 20 inhabited tubes, the longitudinal ridges were prominent in nearly all the tubes (although not equally prominent throughout the length of the tube, e. g., one of four fragments of the single tube in Fig. 5 H appears covered with a granular layer). Some of the three-ridged tubes on Beets’ shells were provisionally identified as Hydroides homoceros (see App. Table 2 B), however, due to the lack of taphonomic residues, presence in the Canal in 1950 is considered as likely but not conclusive. The first complete specimens (inhabited tubes), were collected in the Great Bitter Lake (east of Deversoir-Km 97 [SLC 50] in 1969, see App. Table 2 C). The operculum of Hydroides homoceros, and specifically the tips of the marginal radii of the funnels, shows an interesting variability in form (Figs 12 A – E) and all of the forms are present in the Persian Gulf. The “ double-anchor ” (Pixell 1913) and “ single-anchor ” (Wesenberg-Lund 1949) forms can be considered as the most extreme of these forms. The proximal paired lateral spinules may range in form from anchor-shaped to a more pointed “ T-shaped ” as in Figs 12 D – B. A distal pair of lateral spinules may be developed similar to the proximal pair (= “ double-anchor ”, Fig. 12 C), or the distal lateral spinules may be reduced to a more or less rounded spur, “ spur-tipped-T-shaped ” (Fig. 12 B); alternatively, distal lateral spinules may be lacking entirely (Fig. 12 D). The “ double-anchor-shape ” (Fig. 12 C) is similar to that figured in Pixell (1913, pl. 8 fig. 1 a) and in Mohammad (1981 fig. 2 c), but, in re-examining their material, we also found specimens with marginal radii ranging in form between Figs 12 B and C. This “ double-anchor ” form has been reported from the Indian Ocean, Zanzibar, Maldives (Pixell 1913), the Arabian coast, Oman (Monro 1937), from the Persian Gulf, Kuwait (Mohammad 1981, see references above), from the Seychelles (present paper), and the southern Red Sea - Dahlak Archipelago (Ben-Eliahu 1991 b, Fig. 12 C). Specimens from Oman showed some additional variation (present paper). It follows that in these populations there is more variability in the form of the marginal radii than given in their formal descriptions. The “ spur-tipped-anchor-shaped ” form is shown in Fig. 12 B; ten Hove (1970 a, figs 1 a – c) illustrated some of the variability in the tip of the spur; the population was from the Persian Gulf. The single-anchor form (Figs 12 D, E) reported from the Persian (Arabian) Gulf by Wesenberg-Lund (1949), has been found in the Gulf of Suez (present paper and Selim 1997 b: 87, 93 – 94, figs 6 a – e). It also characterizes both the Suez Canal population (Figs 12 A, D), and the Lessepsian migrant populations on the Levant coast of Egypt and Israel (ca. 80 individuals) (Figs 12 D, E). That only one of several forms present in the Persian (Arabian) Gulf has been found in the Gulf of Suez, colonized the Suez Canal and the Mediterranean coast of Israel, is an illustration of “ a founder effect ” (Mayr 1966, Ben-Eliahu 1991 b). Interestingly, the H. homoceros specimens removed from the aircraft carrier “ Foch ” in Toulon Port (Zibrowius 1979 b) belonged to the “ double-anchor ” Pixell type (H. Zibrowius, pers. comm.), thus, settlement on the “ Foch ” presumably occurred in the Indian Ocean or the Red Sea proper (Ben-Eliahu 1991 b). A single specimen recently collected from the Levant coast of Turkey (Iskenderun) by Çinar (2006, fig. 4 d – e), belongs to the “ spur-tipped-T-shaped ” form (in Figs 12 B, C, closer to B than to C). The distribution of the form (Persian (Arabian) Gulf, ten Hove 1970 a, and Turkey, Çinar 2006) provides a convincing illustration of a disjunctive population founded through ship-transport. Hydroides homoceros has a juvenile 2 - tier opercular ontogenetic stage belonging to the H. “ priscus ” type, one of several species in which this stage has been found (ten Hove & Ben-Eliahu 2005, fig. 2 a).	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F64E034FF50FC55FC3AFBC3.taxon	materials_examined	Type locality. Norway. Not present in the Suez Canal, but see H. elegans.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F64E033FF50FBF8FAFBF91A.taxon	description	Figs 13 – 15	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F64E033FF50FBF8FAFBF91A.taxon	materials_examined	Material examined. Locations adjacent to the Suez Canal, Mediterranean side: None. Suez Canal proper: 1 sample, Hebrew University-Smithsonian Expeditions, 1967 – 1973, Little Bitter Lake opp. opp. Kabrit-Km 120, SLC 85, 14. I. 1969, holotype and 2 paratypes. Suez Canal material reported herein: 17 samples, 28 specimens. Beets’ Great Bitter Lake samples, VIII / IX 1950, Stn 15, 10.4 m, from inside an empty Chama asperella shell, single tube with taphonomic residue of verticil, RMNH 18545 (Fig. 14). — Hebrew University-Smithsonian Expeditions, 1967 – 1973, 2 samples, 4 specs: Lake Timsah opp. Isma’iliya, Km 78: SBE 8, 3 specs; Great Bitter Lake: SLC 117, 1 spec. — Great Bitter Lake “ Yellow Fleet ” Biofouling Samples (January 13 – 20, 1975): 14 subsamples, 23 specs. Material deposited in HUJ; BM (NH) ZB. 1985.209, 1 spec.; Universidad del Mar UMAR-Poly 118, 3 specs; ZMA V. Pol. 3513, 1 spec.; SMF, 2 specs. Locations adjacent to the Suez Canal, Red Sea side: Gulf of Suez, legit J. K. Lord ca. 1870, det. H. Zibrowius 1969, Ben-Eliahu (1972 a), H. A. ten Hove (1990: 119, figs 16 – 18), 1 spec., BM (NH) 1870.12.23.62. Gulf of Aqaba: Egypt, Marsa Abu Samra, 68 m, dredge, on Malleidae TAU-MO- 19788, legit Ch. Lewinsohn 6. X. 1969, det. M. N. Ben-Eliahu ca. 1985 (Ben-Eliahu 1991 b: 525, fig. 4). Red Sea: Sudan, Sanganeb Atoll, 9 m, near Stn TQ 2, SAN 46, coral rubble with Serpula jukesii, legit D. Fiege 25. III. 1991, det. H. A. ten Hove 1999; 10 – 15 m, near Stn TQ 22 b, TQ 2, coral rubble with Vermiliopsis infundibulum / glandigera- complex, Protula spec., legit D. Fiege 19. IX. 1992, det. H. A. ten Hove 1992. — Eritrea: 6 samples (TAU-NS [no numbers], 4 samples, SMF 2 samples), depth range: 9 – [9 – 12] – 36.6 m). Dahlak Archipelago, 14 ° 58 ’ N, 40 ° 19 ’ E, 9 – 12 m, on Plicatulidae, Israel South Red Sea Expedition, ISRSE / 65, Stn 4, legit Ch. Lewinsohn 17. X. 1965, det. M. N. Ben-Eliahu ca. 1985 (Ben-Eliahu 1991: 525, fig. 4), TAU-MO- 1454, TAU-MO- 2186; Stn 12, 15 ° 35 ’ N, 40 ° 44 ’ N, 36.6 m, on Malleidae, legit Ch. Lewinsohn 17. X. 1965, TAU-MO- 1814; with H. homoceros, S. latiscapus, S. tetraceros, det. M. N. Ben-Eliahu ca. 1985 (Ben-Eliahu 1991 b: 525, fig. 4). Indo-West-Pacific: Philippines, Banacon Island, Danajon Bank, N. W. Bohol Island, reef and sand, on clams, legit C. Short, II – III. 1976; det. H. A. ten Hove 1986 (ten Hove 1990: 119), Australian Museum, 2 specs. Suez Canal depth and substrates: 0.4 – 10.4 m; on sponges, on molluscs: Brachidontes pharaonis, Chama asperella, Pinctada radiata and Spondylus spinosus; on bryozoans, tunicates, sandstone rock, and on a muddy tin can. Colouration. Lake Timsah field notes describe bodies with red-orange pigmentation.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F64E033FF50FBF8FAFBF91A.taxon	distribution	Distribution. Suez Canal: Lake Timsah, Great Bitter Lake, Little Bitter Lake; Red Sea: Gulfs of Suez and Aqaba, South Red Sea — Dahlak Archipelago; Philippines.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F64E033FF50FBF8FAFBF91A.taxon	discussion	Remarks. There have not been many records of this species. Hydroides steinitzi was described from 3 specimens collected from the Little Bitter Lake in 1969 (Ben-Eliahu 1972 a). Its verticil is unusual in being comprised of a single element with out-pocketing, somewhat resembling a peeled orange when seen in apical view. The verticil of the holotype had 6 out-pockets (bulges), and the funnel 12 pointed radii (Ben-Eliahu 1972 a). Note variability in the shape as well as in the number and the degree of chitinisation of the funnel radii (Figs 13 A – G). Hydroides steinitzi was first collected from the Gulf of Suez by the J. K. Lord Expedition in 1870 (specimen BM (NH) 1870: 12: 23: 62), presumably from a coral reef habitat (the sample largely consisted of Spirobranchus species that inhabit living coral [ten Hove 1970 b: 49 – 50]). The original sample, BM (NH) 1870: 12: 23: 31, was subdivided into 5 different taxa in 1969, during a visit by one of us (HAtH) to the Natural History Museum, London; two of the taxa were Spirobranchus spp., three taxa, Hydroides spp. In 1971, H. Zibrowius re-identified the provisionally identified specimens of Hydroides, giving them temporary BM (NH) codes. On receiving the Ben-Eliahu manuscript describing Hydroides steinitzi, H. Zibrowius identified one of these Gulf of Suez specimens (BM (NH) 1971: 21), as H. steinitzi, and the newly identified specimen was then assigned a permanent code by the museum, BM (NH) 1870.12.23.62. Unfortunately, Zibrowius’ reply to Ben- Eliahu (letter, 26. XI. 1971) reporting the existence of the Gulf of Suez specimen came when the issue with the description was already in production, but a “ note added in press ” was permitted, mentioning the record (Ben- Eliahu 1972 a). The Gulf of Suez specimen measured ca. 11 mm in length, body 65 chaetigers, with ca. 10 radioles per lobe; the operculum had 6 out-pockets in the verticil and 10 radii in the funnel (H. Zibrowius, pers. comm.; ten Hove 1990: figs 16 – 18). The operculum resembled Figs 13 C 1 and C 2 in form. The first record of Hydroides steinitzi from within the Suez Canal, from 1950, is from the Great Bitter Lake, from a tube within a shell collected by Beets that contained a taphonomic residue of a verticil (RMNH 18545 [Fig. 14]). We have no way of knowing the date when the species settled in the canal. Altogether, it has been collected on four separate occasions (see above). Presently, the total number of H. steinitzi specimens from the Suez Canal comes to 30, with most of the specimens obtained from the Great Bitter Lake “ Yellow Fleet ” biofouling aggregation. These later specimens enabled providing a first description of the tube (the tubes of the type specimens were unwittingly damaged). Hydroides steinitzi was considered to be a potential Lessepsian migrant as it had been found both in the Bitter Lake and in Lake Timsah (Ben-Eliahu 1991 b). Subsequent reports from the Suez Canal or from the Mediterranean areas adjacent to it (e. g., Alexandria) have not included it, but this may be due to not sampling in its preferred microhabitats or on preferred substrate taxa (e. g., molluscs, bryozoans, etc.). H. Zibrowius (1979 b) reported it among ship-transported biofouling taxa removed from the aircraft carrier “ Foch ” that arrived in Toulon Port (via the Suez Canal) after voyaging for 7 months in the western Indian Ocean. A first description of the Hydroides steinitzi tube (Figs 14, 15): The tube has three longitudinal ridges close together on the upper surface and transversal ridges, which give it a generally rugose (not delicate) appearance. The median longitudinal ridge may be as prominent as the lateral longitudinal ridges as seen in three of the four tubes figured from above (Figs 15 A, B and E) and in some cross-sections (Figs 15 D, E); in Figs 14 F 1 and F 2, the median ridge is lower, but it can be discerned. Note that in Figs 14 A, B, a median longitudinal ridge cannot be perceived; the tube surface appears to be covered by a granular overlay — as in many of the H. steinitzi specimens — obscuring the surface sculpturing (Fig. 15 B); it is not clear whether the overlay is made by the worm. In some cross-sections (Figs 15 C, D), the tube appears generally rounded, with a flattened upper surface, but in others (e. g., Figs 15 E, F) the tube is sub-trapezoidal, expanded basally; the upper surface may appear flattened. The tube may be somewhat coiled or looped (Figs 14 A, B, 15 B).	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F61E03EFF50FF1BFD22FE4B.taxon	materials_examined	Material examined. Suez Canal material reported herein: Hebrew University-Smithsonian Expeditions, 1967 – 1973: Little Bitter Lake: SBE 1, juvenile, tube rather square in cross-section with lateral ridges, 1 spec.; SLC 117, some of the 109 juveniles with collar chaetae spot-checked for presence of characteristic Hydroides elegans chaetae belonged to a different taxon or taxa. — Great Bitter Lake “ Yellow Fleet ” Biofouling Samples, January 13 – 20.1975: 6 subsamples, 7 specs. Suez Canal depth and substrates: On bivalves, Brachidontes pharaonis and Spondylus spinosus; on tunicate; under rock. (2) Hydroides cf. “ priscus ” (Pillai, 1971) - type juveniles with a secondary opercular form found in several Hydroides taxa (ten Hove & Ben-Eliahu 2005), see Fig. 16.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F61E03EFF50FF1BFD22FE4B.taxon	description	Eastern Mediterranean	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F61E03EFF50FF1BFD22FE4B.taxon	materials_examined	Material examined. Hydroides grubei not Pillai, sensu Zibrowius & Bitar 1981: 159 – 160 [Lebanon, Beirut, Zaitouné, 5 m, on bivalve, 23. IX. 1978]. Suez Canal material reported herein: 2 samples, 3 specimens. Cambridge Expedition “ new ” sample, from BM (NH) mollusc sample, Lake Timsah, Km 78, Isma’iliya, 1 spec. (App. Table 2 A; ten Hove & Ben- Eliahu 2005, fig. 4 c). — Hebrew University-Smithsonian Expeditions, 1967 – 1973: Great Bitter Lake: SLC 117 (Fig. 16 B), 2 specs (ten Hove & Ben-Eliahu 2005, fig. 4 d). Suez Canal depth and substrates: 10 m, on bivalve, Pinctada radiata; on barnacle.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F61E03EFF50FF1BFD22FE4B.taxon	discussion	Remarks. The “ Hydroides priscus ” ontogenetic stage with a juvenile secondary operculum containing a funnel and verticil was mentioned in Ben-Eliahu & ten Hove (1989: 393), and again as “ Hydroides novaepommeraniae in Ben-Eliahu (1991 b: 527) and Ben-Eliahu & ten Hove (1992: 42, syn. H. grubei). It is a juvenile ontogenetic stage, and has been documented in several individuals with different mature opercula on the opposite lobe, e. g., in Hydroides homoceros, H. cf. brachyacanthus (Ben-Eliahu 1991 b: 527), as well as in H. minax, H. cf. albiceps and H. trivesiculosus (ten Hove & Ben-Eliahu 2005). The first three are Lessepsian migrants (Ben-Eliahu & ten Hove 1992: 42). Hydroides grubei not Pillai sensu Zibrowius (1979 b) could be a juvenile stage of these or other taxa, and thus we excluded it from Zibrowius (1979 b) ’ s list of species present in biofouling scraped from the aircraft carrier “ Foch ” (see Discussion Section below). The Great Bitter Lake SLC 117 “ H. priscus ” specimen had sharp pointed teeth in the funnel radii (ten Hove & Ben-Eliahu 2005, fig. 4 d) and its bayonet collar chaetae had sharp teeth.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F6EE03FFF50FD81FDFBFE00.taxon	description	Fig. 3 C	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F6EE03FFF50FD81FDFBFE00.taxon	materials_examined	Material examined. Locations adjacent to the Suez Canal, Mediterranean side: Crete: 1 sample (intertidal). — Turkey: 3 samples (intertidal). Cyprus: 24 samples (6 samples included the “ thick-rim ” form described in Ben-Eliahu & Payiatas (1999) along with the typical form; 2 samples comprised only the thickrim form, [0 – 0.5] – [0 – 8] – [10 – 18] – 92 m]). — Lebanon, Beirut, 1 spec., permanent mount, det. K. J. Bush 1905 Josephella humilis, and several tubes on a bivalve, collectively catalogued as YPM 2969 though not specified as a type, synonymised with J. marenzelleri by Fauvel (1927: 380 – 381, fig. 129 m – t), confirmed M. N. Ben- Eliahu 1998. — Israel: 7 samples (3 samples with both thick-rim and typical forms, 1 with only thick-rim form, [intertidal – [4 – 6] – [8 – 10] – 18 m]). Suez Canal material reported herein (Fig. 3 C): Beets’ Great Bitter Lake samples, VIII / IX 1950, residues of minute tubes on shells, 2 subsamples, presumed to be Josephella. — Great Bitter Lake “ Yellow Fleet ” Biofouling Samples, January 13 – 20 1975, 18 subsamples, ca. 119 specimens. Locations adjacent to the Suez Canal, Red Sea side: Gulf of Suez: None. — Gulf of Aqaba: Egypt, Israel, typical form, but population from North Beach, Elat included individuals lacking opercula: 6 samples, intertidal – 3 – 8 m, HUJ. Suez Canal depth and substrates: On sponges; on the bivalves, Brachidontes pharaoni s and Spondylus spinosus; on tube of Spirobranchus tetraceros, on barnacles; in crevices of bryozoans; on tunicates; in crevices of biofouling conglomerate.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F6EE03FFF50FD81FDFBFE00.taxon	distribution	Distribution. Worldwide in (sub) tropical to temperate regions. Mediterranean: Israel; Gulf of Aqaba: Israel, Egypt.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F6EE03FFF50FD81FDFBFE00.taxon	discussion	Remarks. A cryptofaunal species frequently found in calcareous concretions, e. g., coralligène, calcareous algae. The new record is most likely due to the stable reef-like substrate provided by the biofouling aggregation on the “ Yellow Fleet ” ships. Erect tubes were also found on Spondylus spinosus between the spines (Fig. 3 C) and on tunicates in folds near the siphons. Some tubes were coiled. The Great Bitter Lake individuals seemed typical. A thick-rimmed opercular form, described in Ben-Eliahu & Payiatas (1999), is present along with the typical form in populations from both Cyprus and Israel. The Red Sea population from the Gulf of Aqaba (Sinai Peninsula) only had typical opercula, suggesting that the Suez Canal population originated from the that population.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F6FE03FFF50FDDEFF26FD20.taxon	materials_examined	Type locality. Arctic Ocean, Greenland. Presumed not present in the Suez Canal, but see Placostegus sp., below.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F6CE03CFF50FD7EFF13FC16.taxon	description	Pillai (2009: 146 – 148) makes a good case for synonymising these three genera on morphological criteria, uniting about 30 taxa that have been kept apart for almost a century in a single genus. A full discussion of this synonymy is not within the province of the present paper. Given the genetic tools available today, however, this synonymy should be backed by genetic data. For the time being, we prefer to use the traditional taxonomy.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F6CE03CFF50FBB3FE05FAE8.taxon	materials_examined	Type localities: Cape of Good Hope and New Zealand. This nominal taxon is a species inquirendam, already evident from the two “ type localities ”, the material of which refers to two different taxa, Pomatoleios kraussii and Spirobranchus carinifer. For records under the name Pomatoceros caeruleus from the Suez Canal, see Spirobranchus tetraceros.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F6CE03CFF50FAFDFCD1F990.taxon	materials_examined	Type locality. Not precisely known, but North Atlantic. Not present in the Suez Canal, but see Spirobranchus tetraceros, below. On ecological grounds, it is inconceivable that Pomatoceros triqueter has replaced Spirobranchus tetraceros in the intertidal and splash zone around Alexandria, Egypt as recently reported by El-Rashidy et al. (2009), a finding that contradicts Abd-Elnaby (2005) [quoted by Ghobashy & Ghobashy (2005: 90 – 93); see Spirobranchus tetraceros, below]).	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F6CE03AFF50F945FBBDFF2A.taxon	description	Eastern Mediterranean	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F6CE03AFF50F945FBBDFF2A.taxon	materials_examined	Material examined. Locations adjacent to the Suez Canal, Mediterranean side: Israel: 11 samples, with one, at the most, two, individuals [0 – 0.3] – [0.1 – 1] – [0.5 - 0.75] – [4 – 6] m. Substrates along Israeli coast: On gastropod Stramonita haemastoma (TAU-MO- 16366), legit J. Krystal 27. X. 1958, det. M. N. Ben-Eliahu, 4 specs, first record from Israeli coast; on asbestos plates at Electric Corporation, Haifa; on tide-pool snails; under stones; on Stramonita haemastoma gastropods (TAU-MO – 16356; 16370, 16371, 20717), on muricid gastropod; on breakwater, 0.5 – 0.75 m, and 4 – 6 m; Tel Aviv, Reading power plant, presumed 0 – 2 m. Suez Canal proper: Lake Timsah, legit, det. S. H. Shalla ca. 1984 Pomatoceros triqueter, redet. H. A. ten Hove Pomatoleios kraussii, ZMA V. Pol. 4999, 5 specs. Locations adjacent to the Suez Canal, Red Sea side: 0 samples.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F6CE03AFF50F945FBBDFF2A.taxon	distribution	Distribution. Widely distributed in the Indo-West-Pacific, from the Cape Province in S. Africa to Hawaii. Lessepsian migrant to the Mediterranean: Suez Canal, Israel, Lebanon, Turkey.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F6CE03AFF50F945FBBDFF2A.taxon	discussion	Remarks. The species forms mid-littoral “ belts ” or “ zones ” in the Pacific parts of its range [e. g., northern Australia, Straughan (1967 a: 224)]. According to Miura & Kajihara (1984), Pomatoleios kraussii prefers sheltered areas. Pillai (1960: 17) reported that the larvae prefer aerated and well-lit water. In the Suez Canal, isolated individuals were reported by S. H. Shalla (1985) and by Ghobashy et al. (1986, 1990) only from the western side of Lake Timsah (most abundant in their “ clear-water station ”, no. I). By April 1997, Shalla & Holt (1999) reported well-formed intertidal reefs on the western shores of both Lake Timsah and the Great Bitter Lake. As predicted by Shalla & Holt (1999: 135), this same pattern of pioneer settlement of individuals eventually promoting gregarious settlement and developing into reefs has been repeated in the Lessepsian migrant population along the Lebanese coast (Zibrowius & Bitar 2003). The species is common along the Lebanese coast, locally in large aggregations (H. Zibrowius, pers. comm.). Within the past two years, aggregations have similarly developed in Haifa Bay, Israel, near the Kishon Port (G. Rilov, pers. comm.; Rilov et al., in prep.). Various aspects relating to gregarious settlement are discussed in ten Hove & van den Hurk (1993: 35). Characters distinguishing this species (the solitary form) from two other shallow-water serpulids with triangular tubes are discussed in the section on Spirobranchus tetraceros, below.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F6AE026FF50FE8BFC5EFBF0.taxon	description	For a generic diagnosis, see ten Hove & Kupriyanova (2009: 81 – 83). Preliminary remarks. In their recent review of the taxonomy of serpulid genera, ten Hove & Kupriyanova (2009: 81) classified Protula as the most problematic serpulid taxon. That is because the phylogenetic basis for this genus is ill-defined and based on negative characters, such as lack of an operculum, lack of special collar chaetae and lack of any characteristic ornamentation of the tubes (mostly). Furthermore, characters for species distinction are scarce and poorly understood (ten Hove 1984: 187; 1994: 109). Not only were the initial descriptions scanty (Table 1), but nearly all were based on only one or, at the most, a few specimens (ten Hove & Pantus 1985: 421), so that variability in taxonomic characters could not be taken into consideration. Many taxa have been described on at least partly-presumed differences in chaetation (ten Hove 1994: 109), such as, the characters, presence and location of Apomatus chaetae, that figure so importantly throughout the Protula literature, e. g., Uchida (1978) and in Table 1. To evaluate the extent of variability of diagnostic characters in Protula, ten Hove & Pantus (1985) carried out a study of live Mediterranean individuals considered to be Protula tubularia (Montagu, 1803). Within the same population, they found individuals that possessed Apomatus chaetae and others that lacked them (ten Hove & Pantus 1985: 425; ten Hove & Kupriyanova 2009: 81). Apomatus - chaetae, extremely transparent, are very difficult to discern within the thick bundles of limbate chaetae, so that while presence data can be accepted as valid, absence data might be evaluated as provisional, particularly when the data derive from examination of whole animals under the compound microscope. On the other hand, examining dissected parapodia under high magnification should enable observing Apomatus chaetae if present (Fig. 22 F), as well as examining whole specimens with the SEM, where transparency has no relevance. In the studied Mediterranean Protula tubularia population, the location of rows of thoracic uncini also showed variability, with uncinigerous tori typically present from chaetigers 2 – 7. However, uncini might also be completely lacking, and, particularly in larger specimens, there was an impression that uncini might disappear secondarily (ten Hove & Pantus 1985: 429). The variability in both these characters illustrates why inferences based on few specimens should be viewed with the greatest caution, including inferences from literature. The urgently needed revision of this genus must therefore be based both on a comparison of all available types, and of sufficient topotypical material assembled for each taxon to produce statistically evaluated morphological criteria. The proposed review should include modern molecular techniques, providing supplementary evidence as to whether morphologically similar (allopatric) populations belong to one or more taxa (e. g., Golani & Ritte 1999, Bucciarelli et al. 2002 and Bonhomme et al. 2003). In our objective of dealing with the Suez Canal as a pathway and / or as a habitat, our first priority is elucidation of the source of the population inhabiting it. A first step entails assembling information on the taxa of the biogeographic regions adjoining the canal, and then excluding those taxa that do not correspond with the present material. (This approach should be carried out with other problematic taxa, as well, e. g., with Vermiliopsis sp. / spp. [see below]). Two Protula taxa from the Mediterranean must be taken into account: A larger taxon, Protula intestinum (Lamarck, 1818), type locality, “ European seas ”, 8 – 12 cm in length / 8 – 12 mm in width, and a smaller one, Protula tubularia (Montagu, 1803), type locality, England, 2 – 5 cm in length, 3 – 8 mm in width (Fauvel 1927: 382 – 384). In surveying the tropical Indo-West-Pacific Protula literature, the records can roughly be brought together into two taxa: (a) a larger taxon with branchial radioles arranged in a spire of 3 – 8 whorls, with a tube-diameter of up to 2 cm (ten Hove 1994: 109; Smith 1985: 92 – 97, 433, fig. 9 e), named Protula bispiralis (Savigny, 1822), type locality, “ les mers des Indes ” (it was collected by F. Péron, thus, probably Australia [ten Hove 1994: 109; Weinberg 1996]) and, (b) a smaller one, Protula palliata, type locality Sri Lanka, described from a single specimen and characterised by a semicircular arrangement of the branchial radioles, at most in a 3 / 4 whorl, with a tube diameter of a few millimetres (Table 1). In Willey’s type specimen, the body measured 19 mm in length (without branchial lobes), 3 mm in width; 30 – 40 radioles per lobe. Willey named it for its most exceptional character, the collar, with very long lateral lobes “ rolled on themselves ” and “ probably able to follow the branchial spire to its termination. … ” and “ ventral border was slightly concave ” (Willey 1905: 316; Table 1). Other characters: Thorax with 7 chaetigers with limbate chaetae, and with Apomatus chaetae and uncini from the 4 th chaetiger; thoracic uncini, P + ~ 20 teeth; Willey only illustrated an Apomatus chaeta, a sickle-shaped abdominal chaeta, and a profile view of the peg of a thoracic uncinus (Willey 1905, pl. 7, figs 183 – 185). The taxon, Protula palliata, as recorded in the literature, is possibly a complex of species (ten Hove 1994: 109). Twenty years before Protula palliata was described, McIntosh (1885: 511) published an Indo-West- Pacific Protula taxon, P. arafurensis, collected in the Arafura Sea by the Challenger Expedition. Unfortunately, the description was based on a single specimen lacking its branchial crown. As the species cannot be identified with certainty (ten Hove 1994; Table 1), P. arafurensis can only be regarded as a " species inquirendam ", one that cannot even be taken into consideration for species identification (Table 1). The single later record, Treadwell’s (1906) Protula? arafurensis, is erroneous, the material was re-examined by one of us (HAtH) and belongs to the Vermiliopsis infundibulum / glandigera – complex. Protula procera Ehlers, 1918, type locality Aru Islands, Indonesia, differs from P. palliata in its short collar, larger size, larger number of radioles in a single lobe (56), and in a pectinate (rather than circular) arrangement of the radioles; the abdominal chaetae appear sickle-shaped (Ehlers 1918 pl. 17, fig. 9). Protula anomala Day, 1955, type locality South Africa, and also reported from E. Australia (Smith 1985: 87 – 92), also can be excluded since it is clearly differentiated by its compound radiolar eyes and retrogeniculate abdominal chaetae (for a definition of retrogeniculate, see ten Hove & Kupriyanova 2009: 26). The present Suez Canal specimens from the Bitter Lakes are small, with a tube diameter of <4 mm (Figs 21 A, D), see below, and the branchiae are arranged in two semi-circles (Figs 17 A, B and 19 A – C). Thus, we can confidently exclude the Indo-Pacific taxon, Protula bispiralis, based on its spiralled branchial structure and large size, as well as the Mediterranean taxon, Protula intestinum, on account of its large size. (We have studied material of both nominal taxa from areas more or less adjacent to the Suez Canal). With much greater caution, we need to approach the identification of the smaller taxa from the Indo-Pacific and the Mediterranean that appear to be rather similar morphologically, as the mixed synonymies below suggest. Here the problem of allocation to species is exacerbated by Fauvel’s having included the Persian (Arabian) Gulf and the Indian Ocean as part of the range of the Mediterranean species, Protula tubularia, in his “ Faune de France, ” with a citation of the Indo-Pacific taxon, P. palliata, as a questionable synonym of P. tubularia (Fauvel 1927: 382 – 383). In his earlier Persian Gulf paper, Fauvel (1911: 433), the Protula included was P. palliata, and there Fauvel questioned whether P. tubularia was its synonym. However, in the report of the “ Siboga ” Expedition to the Malay Archipelago, Mesnil & Fauvel (1939: 35) listed a Protula species as P. tubularia, with P. palliata as its synonym, and in that paper the authors noted that, “ as concerns P. tubularia and P. palliata, neither the tubes, nor the worms showed differences sufficient to distinguish between them. ” Fauvel’s (1953) “ Fauna of India ” again referred to the nominal species, Protula tubularia, and cited Protula palliata as its synonym. The illustrations were copied from the “ Faune de France ” (1927 fig. 257). We will never know whether, in arriving at this synonymy, Fauvel took into consideration the collar structure described by Willey, or whether he assumed the character to be of no particular importance, aberrant, or just described in an exaggerated way. In the later Indo-Pacific literature, Wesenberg-Lund (1949) listed Protula palliata and remarked on its very large thoracic membranes; Dew (1959) also listed P. palliata (citing Fauvel’s 1911 publication); as did Pillai (1960: 5 – 7). Fauvel’s taxonomic (biogeographic) philosophy, especially on the occurrence of wide-spread “ cosmopolitan ” species, impacted not only on his own work but on an entire generation of polychaete researchers as well as on the present generation of non-specialists. In the following section on the genus Salmacina, we quote from Fauvel’s remarks on his predisposition to regard similar taxa from different biogeographic regions as belonging to the same taxon (Fauvel 1933 b: 144). In mitigation, Fauvel, who died in 1958 (age 92), seven years before the Scanning Electron Microscope became commercially available (Breton 2006), may have been unaware that he lacked an adequate tool for comparison of minute morphological structures. Moreover, in 1933, when he published the report on the Dollfus Expedition polychaetes, he would have been unfamiliar with the concepts of invertebrate sibling species, first introduced by E. Mayr in 1942 (see also Mayr 1963; Grassle & Grassle 1976), and of cryptic species (e. g., Bucciarelli et al. 2002; Agapow et al. 2004). 1) It is not possible to be absolutely certain that the 5 records of Protula palliata all refer to the same taxon. — 2) McIntosh’s description of “ Protula arafurensis ” is too incomplete to use for identification; P. arafurensis should be regarded as a “ species inquirendam ” (see p. 46, above). — 3) Although the details of McIntosh’s (1885, fig. 18) thoracic uncinus are not clear, the number of horizontal rows of teeth above the peg may come to 23 / 24 rows. In order to minimize the potential for error due to synonymising similar taxa from different biogeographical provinces, we propose, in contradiction to the philosophy expressed in Fauvel (1933 b), to adopt the most conservative biogeographical approach possible, treating Protula taxa from the Atlantic- Mediterranean and from the Indo-West-Pacific biogeographical provinces as separate species unless / until synonymy can be proved for them. It then follows that the name, Protula tubularia, should be reserved for the Atlantic-Mediterranean forms — as noted, the type locality is England (Montagu 1803) — and P. tubularia - like taxa occurring in the Indo-West-Pacific region should provisionally be assigned to P. palliata or to P. cf. palliata including the two unidentified Protula taxa with un-spiralled branchiae from the Seychelles and Amirantes Islands (ten Hove 1994: 109). We intend to continue looking for robust characters to distinguish between these taxa, particularly as the Suez Canal joins both the Atlantic-Mediterranean and Indo-West- Pacific regions, in order to establish the provenance of the Suez Canal Protula species that colonized it. Molecular techniques are not presently applicable to denatured formaldehyde-fixed specimens; thus for comparing the older specimens found in most museum collections, we can still only employ morphological techniques. We consider that some characters used in the past to distinguish between species of the genus Protula are not sufficiently robust to sufficiently discriminate between similar species. For example, the question of type of abdominal chaetae, whether sickle-shaped or geniculate, that Fauvel (1927: 382) used to discriminate between the two Protula species from the Mediterranean, may exist in reality, and may prove to be a workable character to differentiate between taxa. However, ten Hove & Pantus (1985, fig. 2 i – k) showed that the differences between the character states “ sickle ” and “ geniculate ” are not absolute but gradual, e. g., form of the abdominal chaetae varied within a single bundle from straight to geniculate, also with more or less sickle-shaped ones, and suggested that this character should be reserved for populations rather than for single specimens. In this regard, note the variability of shapes in abdominal chaetae of Protula specimens collected from underneath the same small rock in the Little Bitter Lake that ranged from sickle-shaped to geniculate (Figs 18 G – J). We have even observed these different forms in a single individual from the Little Bitter Lake. Fauvel himself noted a reversal of the contour of abdominal chaetae produced by exposure to different chemicals (Fauvel 1927: 384, legend of fig. 130 h, i), putting in question the differences in contour of abdominal chaetae as a robust taxonomic criterion. Differences in the collar structure also may not be a reliable criterion (ten Hove & Pantus 1985: 427). A character that may prove useful in fresh Protula material (ten Hove & Pantus 1985: 429 – 430), but not in preserved specimens, is the presence / absence of radiolar and prostomial eyespots, because eyespots observed in freshly described specimens may fade in preservative (Ben-Eliahu & Fiege 1996; Table 2), and their durability in preservative is insufficiently well known (e. g., paired crimson eyespots densely positioned on the rachides of Spirobranchus tetraceros radioles faded within three years of formalin-fixation and preservation in alcohol [Ben-Eliahu et al. 2003]). Presently, the best and most reliable morphological comparisons appear to be of the ultrastructure of the thoracic and abdominal uncini, using SEM micrography as a standard tool to determine, in frontal view, the number of rows of teeth and the number of teeth in the horizontal rows, the shape of the anterior tooth (in Protula, the “ peg ”), whether pointed, rounded, more or less swollen, truncated, “ square-ended ” or bilobed (e. g., Figs 18 C, D, K, L; Fig. 20 A, P:? 4: 4: 4: 3: 3: 1: 1: 1: 1: etc.; Table 3), while the light microscope should be used as well for squash preparations to provide a profile view (Fig. 20 B). Mediterranean material of the nominal Protula tubularia has been studied with SEM by ten Hove & Kupriyanova (2009) and again by the present authors. However, even with the SEM, due to the very close packing of the uncini in the torus, adjacent structures may be difficult to distinguish clearly. Thus, to better visualise the structure, uncini of Protula tubularia have been “ cut ” out of the photograph of the torus (Fig. 18 K [thoracic uncinus, cut from ten Hove & Kupriyanova (2009 fig. 39 c), see detailed description in the legend]; and Fig. 18 L — abdominal uncinus of specimen from the same sample) to compare with the abdominal uncini from specimens from the Little Bitter Lake (Figs 18 C, D). The following key presents some differences noted between these uncini: 1 a. In the Suez Canal Protula taxon, both thoracic and abdominal uncini are of “ saw-to-rasp-shaped ” type, i. e., the teeth in the posterior apical (saw ”) - part are in single file (Figs 18 C, D); the peg tip is bilobed, with rounded lobes (Figs 20 D – G); the thoracic uncini have ca. P + 19 rows with ca. 3 –? 4 multiple rows, grading to? 4 minute teeth in the row proximal to the peg (Fig. 20 A). 1 b. In the Mediterranean Protula tubularia (Figs 18 K, L), in both the thoracic and abdominal uncini, the teeth in the apical part form an angled double row, the uncini are thus “ rasp ” - shaped type in both apical (posterior) and proximal (anterior) parts and the tip of the peg appears more blunt (possibly abraded); moreover, the number of teeth is much greater; the thoracic uncini have ca. 21 apical angled double rows and ca. 17 multiple anterior rows grading to ca. 5 – 7 minute teeth in the rows proximal to the peg, with ca. 7 teeth in the proximal row, ca. 38 rows of minute teeth (counted from enlarged figs 39 a, c in ten Hove & Kupriyanova (2009) and Fig. 18 K). These differences in ultrastructure of the uncini point to the Mediterranean and Suez Canal Protula taxa as being different. Although recognizing that the ultrastructure may be variable and that the small number of specimens examined lacks a statistical basis of confidence, these differences presumably justify excluding the Mediterranean taxon, P. tubularia, as a founder population for the Bitter Lake specimens and support a Red Sea derivation for the population. That inference is bolstered by the fact that Beets’ Protula tubes were found on molluscs dredged from the floor of the Great Bitter Lake, thus, not connected directly to ship fouling, more so, as foulers of this genus have not yet been recorded from the Indo-Pacific (ten Hove 1994: 109). Another factor supporting the inference of a Red Sea derivation is the lecithotrophic nature of the Protula larva that dictates a short-term larval life and rapid settlement (Kupriyanova et al. 2001; See Discussion, Section 6 a). Taking these factors and the taxonomic approach discussed above into consideration, we designate the Protula species from the Suez Canal as P. cf. palliata, subject to confirmation by comparison with topotypical P. palliata material. (Meanwhile, efforts to obtain topotypical specimens have not yet proved successful. Moreover, the National Museum Colombo, where A. Willey had been a director, does not have any specimens [M. Goonatilake, NMC and Dr. T. G. Pillai, NHM, pers. comms.]).	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F76E026FF50FBEEFA5EF8BB.taxon	description	Figs 17 – 22, Table 2 Records from the Red Sea proper-Indo-West-Pacific that (most probably) can be attributed to the nominal taxon Protula palliata are:	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F7BE056FF50FE4EFEDAF873.taxon	materials_examined	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 1933 a, 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. 23 A). 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. 23 B). Frontal SEM views of uncini of these two forms are given in Figs 23 A and B; see also Figs 27 B 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.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F7BE056FF50FE4EFEDAF873.taxon	materials_examined	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. 26 A, 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., “ tu 1 ” or “ tu 2 ” 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 ZMA 11 - tu 1: (F + 1) = 3,2,3,3,3 (five adjacent uncini counted in SEM micrograph of first thoracic torus). ZMA 11 - tu 2: (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). ZMA 11 - tu 2: (F + 1) = 2,3,3,3,3,3,3,3,3,2. ZMA- 11 - tu 3: (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 - tu 2: (F + 1) = 2,3,2,2,3,2. S- 14 - 1 - tu 3: (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 ATW 1 - 200 - w 2 - 007: (F + 1) = 3,2,2,3,2,3,2,2,3 (sequence of nine uncini). SEM ATW 2 - 1 - 209 - w 1 - 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 ATW 1 - 219 - w 3 - 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 - tu 2: (F + 1) = 2,2,3,2,3,3,3. SEM Paris-Vaast- 1 - 3 - 018 - tu 4: (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 - tu 5: (F + 1) = 2,2,3,3,3,3,2. 61 - 72 - 2 - tu 6: (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 K 2 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, 32 º 44 ' – 32 º 47 ' E, 28 º 49 – 28 º 54 ' N, 25 – 30 m, coralligenous sand, det. P. Fauvel (1933 a: 80) Salmacina dysteri, MNHN. SEMs: Paris-GS- 1933 - 004 - tu 1: (F + 1) = 5,5,5,4,4,?, 4,5 (counted seven of eight uncini in first torus). Paris-GS- 1933 - 003 - tu 2: (F + 1) = 4,5,?, 4,4,4,4. Paris-GS- 1933 - 002 - tu 3: (F + 1) = 4,5,5,?, 4,4,4,4,?, 4,4 (see Fig. 27 B, 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 - tu 1: (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 - tu 3: (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, tu 1: (F + 1) = 3,3,3,? 4,?,?,?. Second torus, tu 2: (F + 1) =?, 4, 3, 3,4,5; redetermined herein as Salmacina sp. of mru - type). — Hawaii: Biofouling, legit J. H. Bailey-Brock. tu 2: (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. 27 E). SEM-A-Elat- 003 - tu 1: (F + 1) = 5,4,5,5,4,5,4,?, 4,4 (counted nine of 10 uncini of first torus). SEM-B-Elat-tu 1: (F + 1) =?, 5,5,4,4,5,?. SEM-D- Elat- 009 - tu 1: (F + 1) = 4,4,3,3,3,3,5,3,4,3,4,4. Identified as Salmacina sp. of mru – type (Fig. 27 E). 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]).	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F07E050FF50FF1BFD7EF849.taxon	description	Figs 3 B, 24 – 26, 27 A	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F07E050FF50FF1BFD7EF849.taxon	materials_examined	Suez Canal material reported herein: 37 samples, total number estimated as between ca. 200 – 500 individuals. Beets’ Great Bitter Lake samples, VIII / IX 1950, Stns 8 – 10, 33,? 42, B 3, 5 +? 1 samples with tentatively identified (empty) tube residues (see Appendix Table 2 B). — Great Bitter Lake “ Yellow Fleet ” Ship Biofouling Samples (January 13 – 20, 1975), 37 subsamples, 200 – 500 specs (Appendix Table 2 D). Suez Canal depth and substrates: Beets’ samples: Residues of empty tubes on shells between 2.5 – 10.6 m, on Chicoreus erythraeus [as Murex anguliferus], * Murex forskoehlii; * Brachidontes pharaonis and * Gastrochaena cymbium. — Brattström & Taasen Great Bitter Lake “ Yellow Fleet ” ship-biofouling; on sponges; bivalves, * Brachidontes pharaonis (Figs 3 B, 24), * Malvufundus regulus, * Pinctada radiata and * Spondylus spinosus; as well as on barnacles, crabs, bryozoans, and tunicates. (* Denotes an Indo-Pacific mollusc taxon that has migrated and established populations in the Levant Basin of the Mediterranean, i. e., a Lessepsian migrant mollusc [H. K. Mienis, pers. comm.]).	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F07E050FF50FF1BFD7EF849.taxon	discussion	Remarks. Salmacina dysteri, originally described from boreal temperate waters (South Wales), was listed in the Cambridge Expedition Report on the Polychaeta Sedentaria (Potts 1928) as present in the southern part of the Suez Canal and reported by Pixell (1913) and Potts (1928) from the Gulf of Suez end of the canal. Regrettably, Potts (1928) provided no details concerning his nominal S. dysteri material other than that there were especially large aggregations at the bathing place at the sites Port Taufiq 3, and Port Taufiq outside the Canal. That description is insufficient even to conjecture whether the Beets’ and “ Yellow Fleet ” Salmacina specimens sampled ca. 50 years later belong to the same taxon as that of Pixell (1913) or Potts (1928). Fauvel (1933 b: 144) reported Salmacina dysteri from the Gulf of Suez, citing Pixell’s (1913) and Potts’ (1928) reports, and discussed the philosophical basis for his taxonomic work: “ Many species considered for a long time as endemic to the Indo-Pacific region appear absolutely identical with European species ... One risks, therefore, considering as an important new species a taxon simply designated by a different name although in reality it is synonymous. ” Fauvel was, very likely, the most redoubtable polychaetologist of his day, and this biogeographical philosophy presumably influenced H. L. M. Pixell (1913) and F. A. Potts (1928) as seen in their “ cosmopolitan attribution ” of S. dysteri (or they may all have influenced each other). Together, these authors provided an unlikely mix of distributions including those of nominal S. dysteri: The Atlantic, North Sea, English Channel, Cape Verde, Mediterranean, Red Sea, Suez Canal, Gulf of Suez, Zanzibar and Australia (Fauvel 1927). Regrettably, Pixell’s (1913) and Potts’s (1928) important voucher specimens from the Suez Canal could not be found, however, we were able to obtain for SEM examination Pixell’s nominal Salmacina dysteri specimens from Zanzibar and Wasin Island; one of Fauvel’s (1933 a, b) samples from the Gulf of Suez; a nominal specimen of S. dysteri from Hawaii; as well as live Salmacina specimens from the Gulf of Aqaba (from Elat). Our findings did not support the determination of S. dysteri for these specimens or the ascription of S. dysteri as a “ cosmopolitan species ” (see “ Material examined for comparison ”, above). (Interestingly, Fauvel himself had some doubts as to the identification of the Gulf of Suez specimen as he put [three!] question marks after the “ Salmacina dysteri ” in his handwritten label of the voucher sample although the question marks were not cited in his two Gulf of Suez publications [Fauvel 1933 a: 80, 1933 b: 1431]). On the contrary, the new findings support our cautious approach of generalising unverified determinations of Salmacina when the identifications cannot be validated by re-examining the specimens (see also Noguiera & ten Hove 2000). Here it is important to stress that generalising the species determination applies only to the specific sample examined, as in none of the given areas was the density of sampling sufficiently great as to definitively exclude the possibility of more than one Salmacina taxon being present, particularly in the port areas, such as Port Taufiq, and even in the Suez Canal itself. As the late Prof. Heinz Steinitz noted (pers. comm.), finding a taxon in a given area is meaningful, but not having found it may not be indicative of its true absence unless high density and thorough sampling is carried out. This truism is particularly relevant concerning Salmacina and other taxa that can be transported by ship. The type locality of Salmacina dysteri is Tenby, Carmarthen Bay, South Wales. Scanning electron microscope examination of nominal S. dysteri specimens from nearby biogeographic locations, i. e., the North Sea, the Irish Sea, as well voucher specimens from the English Channel identified by P. Fauvel (see Material examined for comparison, above), establishes that Fauvel’s particulars of the thoracic uncini of S. dysteri from its biogeographic home region — “ 7 rows with 2 – 3 teeth in a row ” — were accurate (Fauvel 1927; see also Rioja 1931 and Nogueira & ten Hove 2000), although in his figure, 129 i on p. 379, Fauvel showed only a lateral view of the uncinus. Precise detail was naturally missing in his lateral illustration of the collar chaetae reported as having numerous fine teeth on the fin. Fauvel’s illustration of the S. dysteri aggregate (Fauvel 1927, fig. 129 k, a copy of Huxley 1855, fig. 1) is variously described as “ a bundle of intertwining networks of tubes ” (Fauvel 1927), a “ rope-like twisted mesh ” (Knight-Jones et al. 1996: 256) or an “ arborescent ” aggregate (Nishi & Nishihira 1997). Examination of the nominal S. dysteri tube aggregations from the Hebrides, from Menai Bridge and Fauvel-identified Saint Vaast-la-Hougue (La Manche) material bears that out. The contrasting description of Salmacina incrustans referred to collar chaetae with 4 – 6 large teeth on the fin (Fauvel 1927: 377 – 380, fig. 129 a – l). Other than “ rectangular with several rows of teeth above a larger tooth at the base, ” Fauvel (1927: 378) did not provide any details of the thoracic uncini of S. incrustans. Although we were not able to obtain specimens of published S. incrustans for SEMming, we SEMmed material from traditional locations of S. incrustans from Spain and Croatia. The micrograph of the collar chaetae showed there was a prominent gap between the fin and the blade, as well as a perceptible difference in size between the prominent teeth of the fin and the other more lateral and proximal teeth. Distinct frontal views of a few thoracic uncini in a SEM micrograph of the 4 th torus of the S. incrustans from the Costa Brava specimen (SEM no. 0016), show three teeth in the F + 1 row, with the number of teeth increasing to ca. 5 in the horizontal rows towards the apex. A nominal S. incrustans ’ uncinus from the Adriatic had F + 1 with two teeth. For the tube aggregate of S. incrustans (type locality, Italy), Fauvel (1927: 378 – 379) reported two forms, either, “ Tubes more or less aggregated in colonies encrusting algae, shells and stones, ” or, “ More rarely, forming structures analogous to those of Salmacina dysteri. ” Fauvel reported prostomial ocelli as present in both Salmacina dysteri and S. incrustans (Fauvel, 1927: 377, 378, respectively). However, in their study of taxonomic characters in Salmacina sp. / spp., ten Hove & Pantus (1985) found that newly-collected Mediterranean specimens (not determined to species) lacked prostomial ocelli altogether. In live or freshly collected Salmacina specimens from the Gulf of Aqaba, the character, brilliant crimson prostomial ocelli was very distinctive, but we noted that the ocelli faded shortly after alcohol-preservation (Ben-Eliahu et al. 2007). Our conclusion was that the lack of prostomial or branchial ocelli is a robust character only in live or fresh material, not to be taken into consideration when studying preserved museum specimens (Ben-Eliahu et al. 2007). We can provide no explanation for the discrepancy between Fauvel’s finding and that of ten Hove & Pantus (1985). Nonetheless, we have confidence that the chaetal structures described above and the form of the tube aggregate are robust and durable characters despite some variability in structure. Both of our present Salmacina samples come from the Great Bitter Lake (Appendix Table 2 B, D). The empty tube residues on C. Beets’ shells, collected in 1950, can be identified as Salmacina sp. due to their size and aggregate form, similar to those in Figs 3 B, 24 A, B, thus, presumably, belonging to the same taxon as the Salmacina specimens collected by Brattström & Taasen in 1975 from the “ Yellow Fleet ” ships. The flat- encrusting form of the tube aggregation corresponds to Fauvel’s (1927) description of Salmacina incrustans and excludes a determination of S. dysteri. The collar chaetae and uncini also conform to Fauvel’s characterisation of S. incrustans, taking into account his presumed magnification limitations. Fig. 25 shows the collar chaetae of seven SEM “ Yellow Fleet ” specimens. Although there was considerable variability in their structure, even within a single fascicle (e. g., Figs 25 D 1 – 3, F 1 – 2), there were nonetheless consistent characteristics: (1) the chaeta has a distinct non-denticulate gap separating the basal fin from the blade; (2) ca. 4 particularly large teeth on the fin, the distal-most tooth being the largest, the other three teeth grading in size proximally in the midline; the teeth grading laterally and proximally into smaller teeth, and then even smaller teeth laterally and more proximally reaching their smallest size in the most lateral and most proximal parts of the fin. Tori of thoracic uncini of two specimens are shown in Figs 26 A, C; two uncini were counted in each torus (see legend, Fig. 26 A, uncini designated 1 and 2, and in Fig. 26 C, designated 5 and 6, respectively). The number of teeth in the proximal row to the fang (F + 1) position was 2 – 3, with the number increasing to 6 and 5 teeth and 4 and 5 teeth in the widest rows, respectively, altogether 6 – 7 rows of teeth above the fang (for counts of the abdominal uncini, see Figs 26 B and E). There was considerable variability in structure between the uncini even within the same torus. Both the uncini and the flat-encrusting tube structure of our Salmacina from the Bitter Lake differ from Fauvel’s specimen from the Gulf of Suez (Fauvel 1933 a, b) and from that in the Gulf of Aqaba (Figs 27 B, E, respectively), as well as from a nominal S. dysteri from Hawaii. With less trepidation than before attempting this comparative study, we feel able to identify our Bitter Lake specimens as S. incrustans sensu Fauvel (1927). Salmacina incrustans had been recorded from Alexandria (101 m), ca. 225 km west of the Mediterranean opening of the Suez Canal (Fauvel 1937: 47 [again, voucher specimens not available]). Presuming that our determination as S. incrustans from the “ Yellow Fleet ” population in the Bitter Lake was correct, it is a reasonable inference that this taxon was transported by the ships from the Atlantic and Mediterranean into the Suez Canal, and eventually “ infected ” the Great Bitter Lake, as most probably did two other biofouling taxa, Hydroides elegans and H. diramphus (see above). As regards the Salmacina dysteri sample misidentified by Fauvel from the Gulf of Suez (1933 a, b; Figs 27 B – D), it is worth mentioning that the tube aggregation resembled that of the S. dysteri, e. g., from the English Channel (Manche, St. Vaast-la-Hougue (see Comparative material studied). Fauvel’s sample contained only a single worm, the one we SEMmed, however, the micrograph clearly showed thoracic uncini of the multidentate-rasp-shaped type, with four teeth in the row proximal to the fang (F + 1) = 4 (Fig. 27 B), rather than the 2 – 3 teeth characteristic of S. dysteri; thus, it is certain that this Gulf of Suez specimen was not S. dysteri s. str. despite the similar tube aggregation structure. An important implication of this finding is that the presence of an aggregation of netlike intertwining tubes (presumably the basis for Fauvel’s identification), is not exclusively diagnostic for S. dysteri. Our results indicate that the character is present in additional Salmacina taxa. In the Salmacina populations from the Gulf of Aqaba, the thoracic uncini similarly have multidentate rasp-like uncini with 4 – 5 teeth above the fang: (F + 1) = 4 – 5, Fig. 27 E, and net-like aggregations. This was also true for nominal Salmacina dysteri specimens from Hawaii (Bailey-Brock 1976). Pixell’s Salmacina dysteri from Zanzibar, Tanzania, Wasin Harbour (BM (NH) 1924.6.13.152); from the bottom of S. S. “ Juba ” (BM (NH) 1938.7.25, 26 – 40); off the coast of Zanzibar, Jembani (BM (NH) 1938.7.25.41 – 52), and presumably also Zanzibar 1938.7.25.13 – 25, uncini partly covered (Pixell 1913). In the present paper, we are not attempting to deal in detail with all these taxa, except for generalising them to Salmacina sp. — there may be more than one taxon involved. Multidentate rasp-like uncini are also present in Salmacina amphidentata not Jones, 1962 sensu Fiege & Sun (1999) from Hainan Island as well as in the Caribbean species, S. amphidentata Jones, 1962. Finally, as concerns tube-aggregate form, if both flat-encrusting and net-like forms of aggregate are characteristic of Salmacina incrustans as Fauvel (1927: 379) suggested, this raises the question whether the form of the aggregation can be related to ecological adaptation, for example, to hydrographic factors, i. e., to a less rather than a more protected environment?	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F01E051FF50FF1BFEACFC33.taxon	materials_examined	Type locality. Atlantic Ocean, Madeira. Not present in the Suez Canal; however, for literature records under this name from the Suez Canal, see Hydroides elegans, Hydroides spp. and Serpula hartmanae.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F01E051FF50FF1BFEACFC33.taxon	discussion	Remarks. The name has frequently been used for the juvenile stage of Hydroides that lacks an upper verticil (ten Hove & Ben-Eliahu 2005). As concerns Serpula concharum s. str., with its more than 150 literature records, this Atlantic- Mediterranean species should be a well-known taxon. Its tube is rounded trapezoidal in cross-section, with five more or less similar longitudinal ridges (e. g., Langerhans 1880: 118 – 119; Bianchi 1981: 47 – 49, fig. 14). However, according to Zibrowius (1968: 98 – 100), the tube may have 3 – 5 ridges, and intermediate forms exist. Fauvel (1927: 352 – 353, fig. 121) and Rioja (1931: 404 – 406, pl. 128) explicitly mention 5 ridges in the text, but figure only three. Material collected by us along the Israeli coast up to depths of 20 meters only showed 3 - ridge tubes, with a rather square trapezoidal cross-section, as opposed to the material from deeper localities with 5 ridges and more circular cross-sections. Therefore, Ben-Eliahu & ten Hove (1992: 41) attributed the shallow water population from Israel to Serpula cf. concharum; subsequently Ben-Eliahu & Fiege (1996: 6) used S. concharum “ type B ” to indicate this form with 3 longitudinal ridges while Bianchi & Morri (2000: 260, fig. 2) used Serpula sp. Probably S. concharum sensu auct. from the Atlantic- Mediterranean is a complex of a complex of — at least — two species and much work remains to be done on the variability of taxonomic characters, particularly in the shallow water material. The revision of the taxon should be based both on the available morphological specimens and on genetic studies. Records under this name from outside the Atlantic-Mediterranean probably belong to other taxa (see Remarks in section of Serpula hartmanae, below). A recent report of Serpula concharum from shallow water off Alexandria in Egypt by El-Rashidy et al. (2009) may well be the same taxon as the material present in shallow depth from the coast of Israel.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F01E044FF50FB91FBB2FBCB.taxon	description	Figs 28 – 32, Table 4	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F01E044FF50FB91FBB2FBCB.taxon	description	No previous records, but see Remarks. Suez Canal No previous records. Gulf of Suez and Gulf of Aqaba Serpula concharum not Langerhans, sensu Amoureux et al. 1978: 143, fig. 11, as S. c. [part; Gulf of Aqaba, Elat, Stn 6846, underside of a buoy, legit L. Fishelson, 18. IX. 1970, on dead corals, 2 specs, synonymised herein (Fig. 29). Red Sea proper – Indo-West-Pacific (excluding citations from Gulf of Suez and Gulf of Aqaba already given above) Serpula concharum not Langerhans, sensu Imajima 1982: 38 – 39, figs 1 a – i [fide Imajima & ten Hove (1984), Micronesia, Palau and Yap Islands].	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F01E044FF50FB91FBB2FBCB.taxon	materials_examined	Material examined. Locations adjacent to the Suez Canal, Mediterranean side: Questionable (somewhat similar) material examined (see below). Suez Canal material reported herein: Great Bitter Lake “ Yellow Fleet ” Biofouling Samples, January 13 – 20, 1975: 4 subsamples, 4 specs, two specimens with normally developed opercula (larger, B & T- 02, HUJ- Poly- 4388 (Figs 30, 31); smaller, B & T- 01, HUJ-Poly- 4387 (Fig. 32); and two with small, regenerating opercula, B & T- 03, HUJ-Poly- 4390, respectively B & T- 04, HUJ-Poly- 4389). Locations adjacent to the Suez Canal, Red Sea side: Gulf of Suez: None. — Gulf of Aqaba: Israel, Elat, Serpula concharum not Langerhans, sensu Amoureux et al.: 1978: 143, fig. 11, as S. c. [part; Gulf of Aqaba, Elat, Stn 6846, underside of a buoy, legit L. Fishelson, 18. IX. 1970, on dead corals, 2 specs (Fig. 29) — Egypt, Taba Beach, under rock, 11. V. 1988, 3 questionable specs, see remarks below. Indo-West-Pacific: Marshall Islands, Bikini Atoll, Ocean Side of Enyu Island, legit, det. D. Reish 6. IX. 1956, holotype USNM 38400, paratype 38401 (Fig. 28). — Solomon Islands, Mamara Point and Komimbo Bay, British Royal Society Expedition to the Solomon Islands, 1965, attached to coral on reef platform, legit, det. P. E. Gibbs, BM (NH) 1970.833 – 835, 3 specs. — Palau & Yap Islands, off Arumonogui, Japanese Expedition to the Palau and Yap Islands, VII. 1980, M. Imajima (1982) det. S. concharum, redet. Imajima & ten Hove (1984) S. hartmanae, ZMA V. Pol. 3452, 1 spec. (of the 15 specs collected), tube. Suez Canal depth and substrates: On Brachidontes pharaonis bivalves; on a barnacle; on a bryozoan.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F01E044FF50FB91FBB2FBCB.taxon	distribution	Distribution. Red Sea: Gulf of Aqaba; Indo-West Pacific:? Seychelles;? India, Micronesia.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F01E044FF50FB91FBB2FBCB.taxon	description	Description, based on the largest individual from the Great Bitter Lake with the most differentiated operculum, B & T- 02 (Figs 30, 31, Table 3). Length, 26.5 mm plus (some posterior chaetigers lacking); ca. 120 chaetigers (7 thoracic); operculum with 15 blunt radii without tubercles, apical grooves nearly reaching the centre, surface, concave, cup-like, depth ca. ½ the length of the vesicle. Profile bell-shaped with a waist, basal bulb slightly expanded, distal diameter 3.5 times diameter of waist which is positioned 41 % from the base (Fig. 30 D); length of external grooves 42.5 %. Peduncle with a slight but distinct ventral asymmetrical boss proximal to the marked constriction (more marked in glycerine mount than in alcohol, Figs 30 D, F, respectively). Number of branchial radioles per lobe, ca. 11. Hyaline bodies in the branchial radioles present in all four Suez Canal specimens (Figs 31 F, G and 32 E) and in a Palau Island individual as well. Apron present. Meristic characters are detailed for each of the four Suez Canal specimens in Table 3: Length, 12.7 – 26.5 + mm; width, 0.8 – 1.2 mm; number of chaetigers, 80 – 121 + (7 thoracic), with opercula radii numbering 12 – 16 (mature opercula with 12 and 15 radii and the regenerating opercula with 15 and 16 radii). The number of radii in Amoureux et al. ’ s (1978) synonymised nominal Serpula concharum specimen was also 12; the paratype has 18 radii (Fig. 28 A). The number of branchial radioles in the lobe ~ 8 – 11 (~ 8, 10, 10, 11), respectively (largest individual underlined). Length of external grooves, ~ 37 – 44 %; the maximum opercular diameter 1.8 – 3.5 times that of waist; waist positioned 28.9 – 50 % from the base. Contour bell-shaped, with waist less developed in small regenerating opercula. Characteristic asymmetrical peduncular boss present in B & T- 02, much less distinct in B & T- 01, not present in the two regenerating opercula. Collar chaetae: Bayonet chaetae with 2 conical teeth; in some of the chaetae, a small (fine), very easily overlooked tooth, positioned behind and between the two large teeth and the blade (Figs 28 E 1, 29 I, J; 31 A, second chaeta from left; see also Fig. 32 F). Collar capillary chaetae present. Other thoracic chaetae “ limbate ” and capillary chaetae (Fig. 31 B). Thoracic uncini saw-shaped, F + 5 (Figs 29 M, 31 D), (F + 4, three smaller individuals). Anterior abdominal uncini saw-shaped, F + 4, two individuals; both F + 4 and F + 3 uncini in one individual; F + 3 in one individual (Figs 28 G, 31 E). Abdominal uncini becoming rasp-shaped posteriorly and increasing both in numbers of teeth in vertical row and in numbers of teeth in transversal row (e. g., F: 2: 3: 4: 4: plus several rows of smaller teeth towards apex, e. g., Fig. 29 L). Abdominal chaetae asymmetric flat-trumpet chaetae with ca. 25 teeth (Figs 28 F, 29 K; 31 C, 32 I replaced by capillary chaetae posteriorly (B & T- 02 - capillary chaetae in last 17 abdominal chaetigers + a few missing chaetigers. Hyaline bodies were present in the branchial radioles; becoming infrequent towards the proximal part of the radiole, noted in all four individuals from the Suez Canal (Figs 31 F, G, 32 E). Tube: White with 3 rounded longitudinal ridges on flat upper surface, trapezoidal in cross-section, with transversal ridges; in more anterior parts of the tube the 3 longitudinal ridges are less pronounced, tube appears less rugose (Figs 28 C, D; 29 A – C; 30 A, B; 32 A, B). Thin granular hyaline surface overlay present. 1) For characterizing the opercula, the following contour types are given: A – Funnel shaped (rather straight walls), B – Bell-shaped without flare or waist, C – Bell- (trumpet) - shaped with flare, without waist, D – Bell-shaped with flare and waist, without markedly expanded basal “ bulb ”, E – Bell-shaped with flare and waist, with markedly expanded basal “ bulb ”, — 2) Constriction of peduncle at the base of the opercular vesicle, — 3) The asymmetric boss at the distal end of the opercular peduncle, characteristic for Serpula hartmanae Reish: +++ = very prominent asymmetric boss; ++ prominent (see Fig. 29 F); + visible; – lacking, — 4) Dentition of the collar chaetae, designated as follows: 2 L = 2 large conical teeth; 2 L, 1 m = 2 large conical teeth with 1 smaller median tooth positioned between the teeth and the blade (Fig. 29 J); 2 L, 1 M denotes 3 large conical teeth positioned as above; note that Serpula hartmanae bayonet chaetae lack proximal rasps or additional dentition, — 5) For radiolar hyaline bodies see Figs 31 F, G, 32 E. Brattström & Taasen specimen no. 2 (removed from Bryozoa [Figs 30 – 31]), — 6) Missing several pygidial segments (Fig. 30 E), — 7) Figs 30 D, F, — 8) Fig. 30 C, — 9) Figs 30 D – G, — 10) Fig. 30 G, — 11) 1 – 2 collar chaetae with small median tooth (Fig. 31 A, 2 nd chaeta from left), — 12) Fig. 31 D, F denotes the sharply pointed anterior tooth in Serpula uncini, here seen in profile view, — 13) Fig. 31 E, — 14) see Figs 31 F, G; thinning out of radiolar hyaline bodies towards proximal part (base of radiole). — 15) Fig. 30 B, — 16) Abbreviations, “ tu / t ”, thoracic uncini in torus; “ ac ”, abdominal chaetae; “ au / t ” abdominal uncini per torus; “ caps ”, capillary chaetae in abdomen; number of uncini in thoracic torus, 49; number of abdominal chaetae in fascicle, 13; number of abdominal uncini in torus, 74; last 17 chaetigers with capillary chaetae. Brattström & Taasen specimen no. 1 (removed from Mollusca [Fig. 32]), — 17) Specimen lost after photographing, — 18) Counted on digital photo, — 19) Small operculum, not fully developed and not fully chitinised, without waist (Fig. 32 D), — 20) Fig. 32 C, — 21) Peduncular boss barely developed (Fig. 32 D), — 22) Fig. 32 F, frontal view, left chaeta, shows 2 teeth (intermediate area is not in focus), right chaeta clearly shows 2 large teeth and small median tooth between the large teeth and the blade, — 23) Fig. 32 G, — 24) Fig. 32 H, — 25) Hyaline bodies present, — 26) Fig. 32 A, B. Brattström & Taasen specimen no. 4 (removed from Mollusca; specimen with regenerating branchial crown and operculum): 27) Branchial crown of small size, — 28) Constriction present, not marked, not chitinised, — 29) Basal boss not developed, — 30) Most of the bayonet chaetae with 2 pointed teeth, somewhat abraded; the most basal chaeta appears to have an intermediate tooth between these teeth and the blade, — 31) In one torus, dorsal uncini F + 5, ventral uncini F + 4. Brattström & Taasen specimen no. 3, removed from Cirripedia, specimen with regenerating branchial crown and operculum, — 32) Minute intermediate tooth present. — 33) The number of transverse rows increases posteriorly; frontally, the uncini are rounded, thus; only some of the teeth were in focus; mau F + 5, seemingly saw-shaped; more posterior uncini F + 7 / 3 – 4 sawto-rasp-shaped (compiled from two foci). Red Sea specimen TAU-NS- 6188 (Amoureux et al. 1978: 143) (Fig. 29): 34) Posterior abdominal part missing, — 35) Fig. 29 E — 36) Fig. 29 D, — 37) Single unbroken chaeta, dissected from base of fascicle (Figs 29 I, J) — 38) Fig. 29 M lateral view, 39) Fig. 29 L, — 40) Lateral longitudinal ridges higher than median ridge (Figs 29 A – C). Paratype from Marshall Islands, USNM 38401: 41) Returned to USNM before the difficult-to-observe minute middle posterior tooth was noted in other specimens (the chaetae are abraded but the left-most chaeta seems to show middle posterior tooth). — 42) Thoracic uncini from enlarged fig. 15 of Reish (1968), — 43) Returned to USNM prior to studying character, — 44) Tube with granular layer, covering ridges. IWP Palau, ZMA V. Pol. 3452 (Imajima & ten Hove 1984: 36 – 38, figs 1 a – d). IWP Seychelles, ZMA V. Pol. 4353, S. cf. hartmanae: 45) Very abraded chaetae with 2 - teeth; figure of basal chaeta seems to show tip of small medial-tooth.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F01E044FF50FB91FBB2FBCB.taxon	discussion	Remarks. Narrowing down the identification of our Suez Canal material of the genus Serpula, we considered the species neighbouring the Canal at both sides (and further away as well); see extensive discussion of this approach under Protula. The name Serpula concharum has erroneously been given to Indo- West-Pacific taxa, e. g., Amoureux et al. (1978, see above). Identifying “ Serpula concharum ” - like taxa with a similar opercular contour and a low number of blunt marginal radii from the Suez Canal area, we can discount both Mediterranean taxa of this nominal species (see remarks on S. concharum, above) by the fact that they lack the main diagnostic characters for S. hartmanae: The asymmetric ventral peduncular boss (swelling) proximal to the constriction of the operculum, termed by Reish (1968) “ 2 knobs ”, and a granular surface overlay on the white tube (ten Hove 1994). However, in the Solomon Island specimens of P. E. Gibbs (1971), a sketch of spec. no. 833 showed a prominent asymmetric boss just below the opercular constriction while that of no. 835 was not prominent (notes, Ben-Eliahu, 1986), an indication as to the variability of this character. Considering Serpula taxa distributed in the Mediterranean, we excluded S. cavernicola Fassari & Mòllica, 1991, S. israelitica Amoureux, 1976 and S. vermicularis, all with more (40 +) radii than our present material. It should be noted that we consider S. vermicularis strictly as an Atlantic / Mediterranean taxon, not cosmopolitan, cf. ten Hove & Jansen-Jacobs (1984). Serpula lobiancoi Rioja, 1917 has a pronouncedly zygomorphic operculum. In considering described Indo-West-Pacific (IWP) taxa, we excluded several IWP taxa with more radii such as Serpula indica Parab & Gaikwad, 1989 (maybe including the very similar S. nudiradiata Pillai, 2009, see further down), or S. jukesii, and taxa with a similar number of radii but with more teeth on the collar chaetae such as S. oshimae Imajima & ten Hove, 1984 that also has a predominantly orange tube and 7 – 9 thoracic chaetigers (Imajima 1978, as S. cf. kaempferi) or S. vittata Augener, 1914 (syn. S. palauensis Imajima, 1982) that has Hydroides elegans - like collar chaetae and a tube dotted with brownish speckles. Another Indo-West-Pacific species, S. rubens Straughan, 1967 b (p. 209 – 211, fig. 4 a – i), with a similar opercular contour and number of radii (16 – 18), and collar chaetae with a large and several small teeth, is a much smaller species (to 8 mm in length), with fewer pairs of radioles (a maximum of 7 pairs), and with a variable number of thoracic segments, 8 – 11 (Imajima & ten Hove 1984: 38). The Suez Canal material comprised only 4 individuals and, regrettably, two of them, B & T- 03 and - 04, had regenerating opercula. Revising Serpula hartmanae parameters from Imajima (1982, as S. concharum) and Imajima & ten Hove (1984) provides a reported length of up to 24 mm, with up to 105 chaetigers; thorax with typical constant seven chaetigers; 15 – 16 radioles, and 11 – 25 blunt opercular radii. Meristic differences between their S. hartmanae material and that of Reish (1968) appeared to these authors insufficient for taxonomic distinction (Imajima & ten Hove 1984: 38). Ten Hove (1994) reported Seychelles material with an even greater number of opercular radii (24 – 35) as Serpula cf. hartmanae due to the presence of an asymmetric peduncular boss (“ peculiar swelling ”) and a granular tube surface. Moreover, some of the Seychelles specimens’ opercular grooves were wavy, not smooth, thus it might well be a separate species. The Red Sea S. hartmanae individual had 12 opercular radii (Fig. 29 D). As Table 3 shows, the largest individual from the Suez Canal (B & T- 02) slightly exceeds the compiled length described for Serpula hartmanae. Its operculum shows an asymmetric peduncular boss (Figs 30 D – F), though less prominent than that of the Red Sea specimen (Fig. 29 E, F) which is similar in size to that of the paratype (Fig. 28 B) — it is smaller than that figured for the holotype by Reish (1968, fig. 11). A Palau Island individual had an even larger boss than the paratype or the Red Sea individual (Imajima & ten Hove 1984: 37, fig. 1 a). The number of chaetigers in these Suez Canal specimens is also slightly greater than that described for Serpula hartmanae, but that is also true for other S. hartmanae individuals, e. g., from Palau Island (material of Imajima & ten Hove 1984); that of the Red Sea specimen with a missing posterior abdomen was 67 +. The number of radioles in the Suez Canal material was less than that described for Serpula hartmanae, while the Red Sea individual had 15 radioles in the non-opercular lobe, which conforms to the description. In the several descriptions, differences in the chaetal dentition of the bayonet collar chaetae also emerge. Reish (1968) figured them with 2 teeth, rather abraded in the paratype (see Figs 28 E 1, E 2); ten Hove & Jansen- Jacobs (1984: 149, fig. 2 q) similarly noted 2 teeth without accessory teeth, however, Imajima (1982) and Imajima & ten Hove (1984: 36) referred to “ 2 – 3 heavy conical teeth and a variable number of small accessory teeth at base. ” The collar chaetae of one of their Palau specimens (Imajima & ten Hove 1984) were abraded, except for a basal chaeta with 2 large conical teeth; whether or not there is an intermediate tooth positioned between these teeth and the blade could not be seen; there were no accessory teeth at the base. The Red Sea specimen had only one unbroken bayonet chaeta at the base of the fascicle; it had a distinct small median tooth positioned between the conical teeth and the blade (Figs 29 I, J) similar to some of the Suez Canal bayonet chaetae, described above (Figs 31 A, 32 F) — the remaining collar chaetae were broken. In a Seychelles specimen (of Serpula cf. hartmanae), the median tooth was larger than that in the Red Sea and Suez Canal chaetae (Table 3). Thoracic uncini: An enlargement of Reish’s (1968) figure of the thoracic uncini (fig. 15) shows a dentition of F + 6. The thoracic uncinus given by Imajima (1982: 39, fig. 1 f) was F + 7 teeth, and Imajima & ten Hove (1984) reported F + 7 thoracic uncini. However, a single torus from a Palau Island individual, showed several uncini of F + 6 dentition, while most were F + 5. The Red Sea specimen showed F + 5 dentition (Fig. 29 M). As noted, the largest Suez Canal individual had F + 5 uncini (Fig. 31 D), while the thoracic uncini of the three others was F + 4, a lower number than previously described for Serpula hartmanae. Abdominal uncini: Reish (1968) did not specify dentition of the saw-shaped anterior abdominal uncini; however that of the paratype was F + 4 (Fig. 28 G). The abdominal uncini described by Imajima (1982), and Imajima & ten Hove (1984: 36) are F + 7 anterior and posterior uncini, with up to 4 horizontal rows of teeth in the transversal rows of the rasp-shaped posterior uncini. The Red Sea specimen’s anterior abdominal uncini were F + 4 while the not quite posterior-most uncini were F + 7 / 4. All Suez Canal individuals show a trend of increasing the number of teeth in the vertical rows posteriorly, and the increase in number of teeth in the horizontal rows continued, detailed for some individuals in Table 3, which also shows the variability of dentition within a single torus (most individuals have a posterior dentition of F + 7 / 4 [see “ Treatment of the specimens in “ Material and Methods ” section]). For example, dentition of a posterior abdominal uncinus of specimen B & T – 2 was F + 7 / 4 (or 5) (detailed F: 1: 2: 3: 3: 2: [4 or 5]: 4), while in specimen B & T – 4, in the same torus, the number of teeth were: F + 8 / 4 (detailed F: 1: 1: 1: 2: 3: 4: 4: 2) in one uncinus, and F + 7 / 4 (= F: 1: 1: 2: 2: 3: 4: 3) in another (seen under compound microscope). Abdominal chaetae: Abdominal chaetae cannot be properly described without SEM due to their transparent nature (ten Hove & Jansen-Jacobs 1984: 145); photographs (Figs 28 F, 29 K, 31 C and 32 I) show mostly the fibrils and not the transparent distal teeth; thus, counts of the teeth done under the compound microscope are not reliable. Tube: The Micronesian material from Palau Island (Imajima & ten Hove 1984: 36 – 38, figs 1 c, d) appears more granular than that of the paratype, the Red Sea or the Suez Canal individuals, but all of the tubes appeared to have a thin granular layer. In the Gulf of Aqaba specimens from Taba Beach, the seeming lack of a granular surface on the tubes, despite a three-ridged cross-section, raises a question regarding the identification of the three specimens taken from under a rock (11. V. 1988 [cited above]); the identification as Serpula hartmanae is supported by the low number of opercular radii, 17, 16 opercular radii, 11 radii in a juvenile with marked constriction present at the base of the operculum. Meristic differences found by Imajima & ten Hove (1984) did not seem to them sufficient for taxonomic distinction. Similarly, the Suez Canal material, though showing some variation, should be attributed to Serpula hartmanae. Our results suggest that the degree of waistline in the profile of the operculum is a function of the opercular development (size) and this appears to be true also for the development of the characteristic asymmetric peduncular boss. It is not known whether the closely packed hyaline bodies observed in the branchial radioles have disappeared in some preserved specimens. What is clear is that variability in various characters in the taxon S. hartmanae needs still to be further explored, and particularly, variability in development of the peduncular boss (bosses), in structure of bayonet collar chaetae, dentition of the uncini as well as the effect of different environmental conditions on the formation and degree of tube granulation, and the thickness of the lateral keels. In this exploration, S. amplilobata Pillai, 2009 (pp. 134 – 135, figs 27 a – k, 18 A – E), recently described upon a single specimen only, should be taken into account as well. Apparently Pillai was not aware of the existence of S. hartmanae, since not referring at all to this taxon in his paper, while it is at least very similar — if not synonymous — with his newly described taxon regarding operculum, granular tube, collar chaetae, etc. On reviewing the present paper, H. Zibrowius contacted H. A. ten Hove concerning a new finding by Zibrowius & Bitar that Serpula hartmanae is abundant on the Lebanese coast (Zibrowius & Bitar, in prep.). This finding should cause it to be classified as a Lessepsian migrant proper, sensu Por (see Introduction). The report galvanized us to examine Serpula material encrusted on recently obtained molluscs (samples of the Tel Aviv University mollusc collection), as well as our own collections from 1990, diagnosed as S. cf. concharum (by Ben-Eliahu & ten Hove 1992: 41). The specimens are clearly different from Serpula concharum Langerhans, 1880 s. str., from deeper waters. All 22 specimens surveyed have a three-keeled tube. However, in some tubes, the two dorso-lateral keels are much more prominent, more rounded, and more blunt than the thinner medial one (as in Serpula hartmanae [Fig. 29 B]), but in others (perhaps half of the tubes), the keels are all narrow and rather similar — very likely this is related to size / age, but that still needs verification. Tubes are mostly covered by a thin granular layer. Some of the opercula show a small but distinct ventral peduncular boss proximal to the marked constriction (n = 3) as in Serpula hartmanae (Figs. 29 E, F, 30 D, F, and 31 D), but, in most specimens, the boss is not pronounced. The number of radii range between 10 – 13.3 – 19 (n = 22, SD 2.3). Perhaps the variability observed in the tube structure and the peduncular boss falls within the range of a single taxon, but we cannot presently be sufficiently certain to make a conclusive determination of that. We assume that Zibrowius & Bitar’s Lebanese record belongs to the same taxon, which, in the past, we had identified as S. cf. concharum. This raises the question whether this material belongs to a taxon indigenous to the Mediterranean, as forms with a 3 - keeled tube have been mentioned under various names, the oldest probably being Serpula sulfurata Milne Edwards, 1836. Alternatively, there is a reasonable similarity with our specimen from the Gulf of Aqaba (Fig. 29), presumably identical with S. hartmanae, which indeed might make the taxon yet another Lessepsian migrant. In view of the fact that the taxonomy of the genus Serpula is exceedingly problematic (see Remarks under Serpula jukesii, this paper), we have refrained from reaching a decision at the present time. A revision of S. concharum sensu lato, and all material of Indo-West-Pacific records of similar taxa, including S. hartmanae, is necessary (see remark on variability of this nominal taxon one paragraph above as well), but falls outside the scope of the present paper. Serpula cf. concharum, recently reported in shallow Egyptian waters (Alexandria) by El-Rashidy et al. (2009), is very possibly the same taxon as that found along the Israeli coast (see remarks under Serpula concharum, above).	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F14E040FF50FB03FB94FAEE.taxon	materials_examined	Type locality. East coast of Australia. Most probably not present in the Suez Canal proper, see Remarks, below. Preliminary remarks. Since the presence of Serpula jukesii Baird (1865: 20, pl. 2 fig. 6) in the Suez Canal has not (yet?) been authoritatively demonstrated, we have refrained from a full treatment of the taxon here. For comparison, however, we reviewed the literature of larger Serpula taxa from Indo-West-Pacific origin and studied dozens of samples, as S. jukesii (including its holotype BM (NH) ZB 1982: 84), vermicularis sensu auct., vermicularis granulosa, magna, vasifera, originating from Kuwait to Japan, from India to Australia. Some of this has been summarized in Table 4. Below we also comment upon relevant records from the nearby Gulfs of Suez and Aqaba. Material examined. Gulf of Suez: Cambridge Expedition, stn R 5, 5. XI. 1924, det. F. A. Potts (1928: 700) Serpula vermicularis, fragments of a specimen discussed in detail below. — 33 ° 23 ’ E., 28 ° 14 ’ N, 22 m, muddy sand, St. 17 bis, R. Ph. Dollfus Exp., 25. XII. 1928, det. P. Fauvel (1933 a: 76) S. vermicularis, redet. S. jukesii herein, CUZM, 1 spec. Operculum of specimen symmetrical with blunt teeth, its “ waist ” at mid-point; expanded basal bulb; constriction, deep funnel, 54 marginal radii (teeth); 40 radioles per lobe; tube white, pink inside, tubes without ridges; collar trilobed; apron; collar chaetae with 2 teeth; thoracic uncini F + 4; anterior abdominal uncini F + 5 and F + 6. Red Sea, legit G. von Frauenfeld, det. Grube (1868: 640) Serpula gervaisii, redet. H. A. ten Hove 1982 S. jukesii (Table 4), Muzeum Przyrodnicze Wroclaw, 1 spec. Operculum showing tubercles on inner radii as opposed to type of S. jukesii. (*) Revised numbers of opercular radii and number of radioles per branchial lobe by H. A. ten Hove following his observation of specimen. — 1) The number of opercular radii in Atlantic-Mediterranean Serpula vermicularis counted from different illustrations. — 2) It cannot be excluded that Mediterranean records earlier than 1991 included measurements of S. cavernicola, with 45 – 80 radii (Fassari & Mòllica 1991). — 3) ten Hove & Jansen-Jacobs (1984: 150). — 4) Marenzeller (1885: 216) compared Serpula princeps Grube, 1877 described from N. Japan with his Serpula granulosa. However, Grube (1877) as referred to by Marenzeller [Naturh. Ber. d. Schles. Ges. f. vaterl. Cultur, 1877: 62] cannot be traced in the Dutch libraries, neither in the available bibliographies, nor is the name used again by Grube (1878 a: 104 – 105), where the same diagnosis as cited by Marenzeller (1885: 216) has been attributed to Serpula? jukesii by Grube. Serpula princeps should be regarded to be a nomen nudum. — 5) Correction of observations of Straughan (1967 a) in ten Hove & Jansen-Jacobs (1984), taxon synonymised with S. jukesii. — 6) Not given in Marenzeller, 1885.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F14E040FF50FB03FB94FAEE.taxon	discussion	Remarks. The taxonomy of the genus Serpula is exceedingly problematic. Without comparison of specimens to be identified with type specimens, or at least with topotypical material, identification is difficult under the best of circumstances. This is exacerbated when a specimen collected from within a transoceanic canal is unavailable for direct examination. The only record under the name Serpula jukesii was listed in a table (Abd- Elnaby 2009: 10), without any additional details. Regretfully, it was not possible to forward the material to Amsterdam for direct examination, however, the author generously responded to ten Hove’s request to count the number of opercular radii. On the basis of its having too few opercular radii (19 radii after damage, F. A. Abd-Elnaby, pers. comm. to H. A. ten Hove), we excluded the identification as Serpula jukesii for this Suez Canal specimen. In contrast, the taxon is excluded from an identification as Serpula hartmanae due to its having too great a number of radii (see Table 4), bayonet chaetae with 2 teeth, and by lacking an asymmetric boss on the peduncle, proximal to the constriction separating the peduncle from the operculum (F. A. Abd-Elnaby, pers. comm. to H. A. ten Hove). Given the present constraints, the most plausible conclusion is that the specimen is a juvenile Hydroides with a primary operculum (for a discussion of opercular ontogeny in Hydroides, see ten Hove & Ben-Eliahu 2005). As it lacks the characteristic collar chaetae of Hydroides elegans, we are provisionally identifying it as Hydroides sp. (not enumerated among the verified taxa of the canal, and listing it among the non-determined species below). A nominal Serpula vermicularis sensu auct. had been collected in the Gulf of Suez and was included in Potts’ 1928 article on the Cambridge Expedition Sedentaria although not having been collected from within the Suez Canal proper. Potts (1928) referred to Pixell’s (1913) report of S. vermicularis in the Red Sea, which included citations from Antarctica, Patagonia and the Mediterranean Sea, “ clearly a very unlikely distribution ” (ten Hove & Jansen-Jacobs 1984). Potts’ (1928) Serpula was presumably of Indo-West-Pacific or / and Red Sea affinity, whereas the type locality of Serpula vermicularis is Sussex, England (Heppell 1963: 445). By comparison with the distributions of other boreal-temperate taxa such as Pomatoceros triqueter, it may be safely assumed that Serpula vermicularis also occurs throughout the Atlantic-Mediterranean region. However, even this Atlantic-Mediterranean taxon might not be a single species. Ben-Eliahu & ten Hove (1992) and Ben-Eliahu & Fiege (1996) mentioned differences in tube structure and colour, which may have taxonomic importance. Preliminary work done by us (in 1990 – unpublished) indicates that there may be statistical differences in number of opercular radii, length of interradial grooves between opercula of the “ echinate ” form and Serpula vermicularis “ s. str. ”. Earlier Mediterranean records too may have included material of S. cavernicola. The cosmopolitan status of Serpula vermicularis has been questioned by ten Hove & Jansen-Jacobs (1984); as a consequence Kupriyanova & Rzhavsky (1993) and Kupriyanova (1999) separated 2 valid North Pacific taxa, S. columbiana Johnson, 1901 and S. uschakovi Kupriyanova, 1999, from the previously “ cosmopolitan ” S. vermicularis. Potts (1928: 700) nominal Serpula vermicularis reported an operculum with 70 opercular radii (“ radiating ridges ”), radially symmetrical (“ insertion not as in S. lobiancoi ”), borne on a lobe with 22 branchial radioles. Potts gave no details on body structure or chaetae. Regretfully, the vial with Potts’ nominal S. vermicularis specimen, comprised only a tube and a posterior fragment when reviewed in 1986 at the CUZM by M. N. Ben- Eliahu; the branchial crown and anterior were already missing. However, when rechecked in IX. 2002, only the tube was present, along with some abdominal epidermal tissue adhering to it (Fig. 11). As Fig. 11 A – C shows, Potts’ description of the tube was both Painstaking and accurate: A massive tube, “ coiled in a plane spiral ” and “ the tube is not free anteriorly over a great part of its length ”, as Fauvel states is the case in Serpula vermicularis (Fauvel 1927: 352). The number of longitudinal ridges on the tube was difficult to make out ” (Potts 1928). The tube resembles an enormous Spirorbis, and has a rather flattened upper surface with two blunt longitudinal ridges. Although the mouth of the tube appears rounded in cross-section, there are pronounced transversal ridges (i. e., former peristomes), and it has a thick granular overlay (Fig. 11 C, see discussion below). Other Indo-West-Pacific records of nominal Serpula vermicularis should be checked individually against the voucher specimens. For example, Serpula jukesii sensu Grube (1878), from Japan should be referred to S. uschakovi (fide Kupriyanova 1999). Potts’ (1928) questionable Gulf of Suez specimen would fit into the biogeographical pattern of Serpula jukesii. Dealing with possible synonymies of nominal Serpula taxa, ten Hove & Jansen-Jacobs (1984) wrote: Of the small number of useful taxonomic characters in Serpula, the “ most important is the number of opercular radii ”. According to these authors, Serpula vermicularis from Australian waters generally should be named S. jukesii (including S. magna). Serpula vermicularis granulosa sensu Mohammad (1971, Kuwait, 2 specs) and S. gervaisii not Quatrefages, 1866, sensu Grube (1868), Red Sea, 1 spec., were considered to be possible synonyms of S. jukesii. They are characterised by similar form in tubes, thoracic membranes not forming an apron on the ventrum, 3 - toothed collar chaetae, and a very large number of opercular radii (i. e., marginal teeth) with and sometimes without distinct surface tubercles. Data on the collar chaetae and the thoracic membranes of Potts’ Serpula would have provided important support for identifying the specimen. Nonetheless, the described operculum falls within the Serpula jukesii / S. granulosa complex (sensu ten Hove & Jansen-Jacobs 1984: 143 – 152). In a later paper, ten Hove (1994: 111) suggested that both taxa might be valid; ten Hove & Kupriyanova (2009: 94) treated both as separate taxa. The number of opercular radii of these nominal taxa ranges from 40 – 100 (Table 4). For perspective, Table 4 also gives the range of number of marginal radii for Atlantic-Mediterranean Serpula vermicularis s. str., ranging from 25 – 61, with an illustration of local differences in this taxon in numbers of radii, as opercula from a purely Norwegian population were of larger size than those compiled from various areas including the Mediterranean. Size differences for warmer and colder-water populations of invertebrate species are a phenomenon described by Gunter (1957: 174) and Mayr (1966: 327) and might be found in S. vermicularis as well. Alternatively, the Atlantic-Mediterranean S. vermicularis sensu auct. might prove to be a complex of two (cryptic) species. Attributing the operculum described by Potts (1928) to the Serpula jukesii / granulosa- complex creates a dilemma, as the tube, with its regular flat tight spiral coiling, is not characteristic for Serpula. It appears certain that the tube and operculum are “ mismatched ”. Flat coiled spiralling is, however, known for an occasional Hydroides (H. Zibrowius, pers. comm. and our own unpublished observations and illustrated by Kupriyanova & Jirkov 1997), for Spiraserpula (Pillai & ten Hove, 1994), as well as for the bathyal Nogrobs (cf. ten Hove & Kupriyanova 2009: 68). A Hydroides identification for the tube is not contraindicated by the abdominal chaetae from the epidermis removed from the tube lumen (Fig. 11 D – F). The most plausible solution to this dilemma is that the opercular crown and the tube were fragments of two taxa collected together and placed in the same vial (H. Zibrowius, pers. comm.), so that Potts’ record of this sample should be revised to Serpula? jukesii and to Hydroides sp. Moreover, once the possibility of Hydroides sp. is accepted, we suggest the tube could belong to H. heterocerus, the largest of the Hydroides species known from this area, and one with several records from the Gulf of Suez, indeed from the same general location, e. g., Suez quay (Pixell 1913: 75), Gulf of Suez (Fauvel 1933 a, b) [see above section dealing with Hydroides heterocerus]). Potts (1928) did not indicate whether he had examined a “ loose ” branchial crown — one separated from a worm body floating in the sample material — or seen the body belonging to the operculum. Serpula? jukesii as understood by ten Hove & Jansen-Jacobs (1984: 149) shows considerable intraspecific variability in the collar chaetae: One heavy tooth and two accessory ones or three heavy teeth and two accessory ones, or four nearly equal teeth. However, Serpula nudiradiata Pillai 2009 (pp. 136 – 139) may have been included in S. jukesii by ten Hove & Jansen-Jacobs, possibly throwing a different light upon this variation, their material should be restudied. It should be noted that Pillai apparently did not see the holotype of Serpula jukesii in the BM (NH), at least it was not mentioned by him, and he might have observed that the holotype of S. jukesii does not show tubercles, as opposed to Pillai’s material of this nominal taxon (and that of S. gervaisii sensu Grube (1868 )) which both do show tubercles. A further difference between the holotype and Pillai’s specimens is the presence of an apron, as the ample thoracic membranes are not joined to form an apron in the holotype of S. jukesii. In fact, this was one of the characters upon which ten Hove & Jansen-Jacobs (1984) determined synonymy of their S. jukesii and of S. magna (by comparison of types); however, this characterstate easily may be overlooked if not specifically studied and in some specimens it may be damaged (e. g., in S. gervaisii sensu Grube (1868 )). In conclusion, Serpula jukesii too should be revised. Other Indo-West- Pacific Serpula taxa listed in ten Hove (1994) or described subsequently have fewer radii or opercula of different shapes than in the S. jukesii / granulosa- complex (the following list mentions only opercula for comparison with the Potts (1928) description, without taking into consideration tube, collar chaetae and other Serpula pacifica (Uchida, 1978), a juvenile specimen, lacks an operculum (and was regarded as of uncertain identity by Pillai & ten Hove (1994: 103); Serpula japonica Imajima, 1979, 21 – 24 radii, operculum zygomorphic; Serpula rubens has a bell-shaped operculum with or without a waist, with greater or lesser flare, with only 15 – 18 radii (types restudied by us). Two taxa with opercula with very long external grooves; with hardly any distal flair; elongate, with rather straight sides (basal bulb not expanded, waist not well-defined: Serpula watsoni Willey, 1905, ampulla deeply elongated; 33 – 55 radii, and Serpula sp. of Ishaq & Mustaquim (1996) 16 – 33 radii, ampulla shorter, profile rather squat; 9 – 31 radioles. Two taxa with opercula with very long external grooves, but distally flared, tapering to the constriction, basal bulb not expanded: Serpula vittata 18 – 23 radii and S. vasifera, 31 radii, respectively (according to their written descriptions, the illustrations are contradictory; other material identified by ten Hove had 25 – 34 radii). Three opercula with short external grooves, distally flared, tapering to the constriction, basal bulb not expanded: Serpula tetratropia Imajima & ten Hove, 1984 with 18 – 20 radii, raised central disc, somewhat pointed radii; S. zelandica Baird, 1865, 13 – 21 radii (incompletely described) and S. longituba (Imajima, 1979), 31 radii, short external grooves, with a markedly tapered elongated ampulla. Four taxa with distally flared, bell-shaped opercula with rather short external grooves, and with a well-defined waist, basal bulb more or less expanded: Serpula willeyi Pillai, 1971, 10 radii; S. hartmanae, 16 – 25 radii, with an asymmetric boss proximal to the constriction; S. oshimae 19 – 26 radii, lacking the boss, differs from S. hartmanae in collar chaetae. Similar, but larger, greatly flaring distally, Serpula indica 48 – 56 radii. Two taxa with hexagonal tusk-like tubes, Serpula crenata (Ehlers, 1908) and S. sinica Wu, Sun & Chen, 1979 with 17 – 22 radii and 23 radii, respectively. Three gregarious Indo-West Pacific and Mediterranean taxa were transferred to the genus Spiraserpula because they have internal tube structures (ITS) by Pillai & ten Hove, (1994): Spiraserpula minuta (Straughan, 1967 b: 216, fig. 6 h – m) with only filiform pseudopercula (Australia); Spiraserpula lineatuba Straughan, 1967 b: 216, fig. 6 h – m) (Australia, Queensland and New South Wales), operculum slightly zygomorphic with 22 – 25 radii; outer grooves ca. midway and Spiraserpula massiliensis (Zibrowius, 1968: 102 – 105, pl. 1 figs 24 – 37, pl. 14 fig.), operculum with elongate funnel and rather flat distal cup with 10 – 23 radii and short outer grooves. The nominal Indo-West- Pacific Serpula philippensis McIntosh, 1885, from abyssal depths, is poorly described (questionable, fide ten Hove & Kupriyanova 2009: 95), lacking a description for an operculum and the collar chaetae appear to be atypical. The present, redetermined Serpula jukesii records of Potts (1928) and Fauvel (1933) from the Gulf of Suez are not enumerated among the ca. 16 species reported from within the canal.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F10E040FF50FAEBFAD9F856.taxon	materials_examined	Type locality. European Seas. Not present in the Suez Canal, but see Hydroides elegans, Hydroides spp., Serpula hartmanae and S. jukesii sections, above.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F10E040FF50FAEBFAD9F856.taxon	discussion	Remarks. As discussed above, the taxon Serpula is difficult to classify. The record of S. vermicularis by Selim (2009: 73) from the Suez Canal is unlikely to belong to this boreal-temperate taxon. Given as a listing in a table only, not even with the number of opercular radii, presumably it should be attributed to an unidentified Hydroides species juvenile stage with a primary operculum, or possibly even to Serpula hartmanae. Without verification from direct examination of the specimen, it is presently only possible to regard this record from the northern canal as questionable. Comparative material examined. Atlantic: North Atlantic: Lervik, Stordoe, Hardanger Fjord, Norway, 183 – 238 m, VI. 1892, det. S. F. Harmer Serpula vermicularis, CUZM, 5 specs. Largest, length 58 mm; opercular diameter, 3 mm; 60 radii (36 – 46 – 60, s. d. = 9.08, N = 5, radii). — Mediterranean: Much Levant Mediterranean material in the National Natural History Collections of the Hebrew University of Jerusalem presently determined as Serpula vermicularis (see Ben-Eliahu & ten Hove 1992; Ben-Eliahu & Fiege 1996).	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F11E041FF50FF1BFB12FF0F.taxon	materials_examined	Type locality. West Indies. Not present in the Suez Canal, but see Spirobranchus tetraceros.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F11E041FF50FECEFAA5FE62.taxon	materials_examined	Type locality. Mediterranean Sea, Italy. Not present in the Suez Canal, but see Spirobranchus tetraceros.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F11E048FF50FE61FC63FBF0.taxon	description	Fig. 33, Table 5 Preliminary remarks. The nominal taxon, “ Pomatoceros triqueter ”, was reported from shallow fouling panels from Alexandria harbour and from various locations within the Suez Canal (see Mediterranean and Suez Canal Synonymy Sections below). That contradicts what is known of the ecology and depth preferences of Pomatoceros triqueter, a boreal-temperate taxon with an Atlantic-Mediterranean distribution in sublittoral depths (e. g., in the Levant Basin, samples from 38 – 67 m from Israel (N = 3), and from 119 m from Cyprus (N = 1) (authors’ unpubl. database). Along the coast of Israel and presumably also along the Mediterranean coast of Egypt, there are three rather similar species found in shallow habitats. Like Pomatoceros triqueter, all three have triangular tubes and calcareous opercular plates borne on winged opercular peduncles: Pomatoceros lamarckii (Quatrefages, 1866), an Atlantic-Mediterranean species (Bianchi 1981), Spirobranchus tetraceros and Pomatoleios kraussii, Lessepsian migrant species (Ben-Eliahu & ten Hove 1992) that may occur on biofouling panels, both of them present in the Suez Canal. Spirobranchus tetraceros is distinguished from the others by its more elaborate opercular spines and its fringed opercular wings (in most specimens) and special collar chaetae (Figs 33 A, D and E). The opercular peduncles of both Pomatoceros lamarckii and Pomatoleios have wings with smooth margins. Pomatoleios kraussii has a somewhat concave opercular plate, frequently with a diagnostic sunken (internal) talon (ten Hove 1973 fig. 43); best seen when cleared in glycerine, and it lacks collar chaetae. Pomatoceros lamarckii has fine small limbate collar chaetae (Ben-Eliahu & ten Hove 1992). Ghobashy et al. (1990) retroactively corrected citations of “ Pomatoceros triqueter ” for the Suez Canal to Spirobranchus tetraceros. However, given the presence of the similar taxon, Pomatoleios kraussii, we have placed question marks before the problematic citations.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F11E048FF50FE61FC63FBF0.taxon	description	677 – 685 [Lake Timsah]; Ben-Eliahu 1991 b: 519, fig. 2 [el Qantara-Km 45, Lake Timsah, Bitter Lakes]; Selim 1997 b: 94 – 96, figs 7 a – c, 8 a – e [Suez Canal, “ very abundant in all sites investigated ”, i. e., El-Kab, Lake Timsah, Deversoir, collected in 1988; for Lake Timsah, also citing Shalla (1985) and Mostafa (1992) (see App. Table 4)]; Emara & Belal 2004: 192 – 199 [pooled Lake Timsah and Bitter Lakes data]; Ghobashy & Ghobashy 2005: 91, 93 [referring to Ghobashy et al. (1986) and to Barbary (1992, see App. Table 4); Selim 2009: 74 [northern part of the canal]; Abd-Elnaby 2009: 10 [southern part of the canal].? Pomatoceros triqueter not (Linnaeus), sensu Ghobashy et al. 1980: 84, table 2 [El Tina, Isma’iliya]; Ghobashy & El- Komi 1981 a: 170 [Lake Timsah]; Ghobashy & El-Komi 1981 b: 181 [southern canal]; Ghobashy 1984: 45 [Lake Timsah]; Ghobashy & Ghobashy 2005: 91; 93 and referring to Ghobashy et al., 1980. In 1986, Spirobranchus tetraceros was first cited by this research group and all previous records of P. triqueter presumably refer to S. tetraceros. Spirobranchus sp.: Ghobashy 1984: 45 [Lake Timsah]; presumably S. tetraceros.? Spirobranchus giganteus not Pallas, sensu Selim 2009: 73 [Selim cited Fauvel (1933 a, b) that referred to S. coutieri (Gravier, 1908) and S. multicornis (Grube, 1862), both taxa synonymised with S. tetraceros, and thus Selim’s record is attributable to what we presently regard to be the S. tetraceros - complex]. Gulf of Suez and Gulf of Aqaba	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F11E048FF50FE61FC63FBF0.taxon	materials_examined	Material examined. Locations adjacent to the Suez Canal, Mediterranean side: Rhodes: 1 sample, littoral. — Cyprus: 1 empty tube, [3 – 5] m (H. Zibrowius, pers. comm.). — Lebanon: common (H. Zibrowius, pers. comm.). — Israel: 37 samples, 0.2 – [3 – 18] – 31 m; first Israeli sample from Haifa Bay, 5. XI. 1967, SLM 272, 27 m, det. M. N. Ben-Eliahu, INCNH unpublished data. Suez Canal material reported herein: 57 samples, 137 specimens. Beets’ Great Bitter Lake samples: No triangular Spirobranchus tetraceros tubes appeared to be present on any of the shells. — Hebrew University- Smithsonian Expeditions, 1967 – 1973, 8 samples, 12 specs: el Qantara, Km 45: SLC 6, 1 spec.; SLC 31, 1 spec.; Lake Timsah, Km 78, opposite Isma’iliya: SBE 7, 1 spec.; SBE 8, 3 specs; Great Bitter Lake, east of Deversoir, Km 97, SLC 50, 1 spec.; SLC 61, 1 spec.; SLC 71, 1 spec.; Great Bitter Lake, SLC 123, 3 specs — Great Bitter Lake “ Yellow Fleet ” Biofouling Samples, January 13 – 20, 1975: 45 subsamples, 115 specs; Bremerhaven dry dock, 18. VI. 1975, 2 subsamples, 2 specs, both individuals large, decayed. — Lake Timsah, legit, det. S. H. Shalla ca. 1984 Spirobranchus sp., Conopomatus sp., redet. H. A. ten Hove, ZMA V. Pol. 4997, 4998, 6 specs — El Tawan Beach, 10 / 19. XI. 1988, legit S. B. Shazly, det. H. A. ten Hove, ZMA V. Pol. 3820, 2 specs. Locations adjacent to the Suez Canal, Red Sea side: 7 samples, shallow. Gulf of Suez, J. K. Lord Expedition 1870, det. H. A. ten Hove 1969 S. tetraceros (3 specs left under the original BM (NH) number by HAtH, from a mixed sample of tubes, one Spirobranchus corniculatus (now BM (NH) 1870.12.23.60), three S. tetraceros and 3 epibiontic species of Hydroides (now BM (NH) 1870.12.23.61, 62, 63), first listed together under an unpublished ms. name by W. Baird), BM (NH) 1870.12.23.31. Since both Spirobranchus taxa inhabit living coral, the sample was presumably from a coralligenous habitat; same, redet. H. A. ten Hove 1969 S. tetraceros, BM (NH) 1870.12.23.30, 2 specs — Gulf of Suez, legit R. MacAndrew, listed under an unpublished ms. name by W. Baird, redet. H. A. ten Hove 1969 S. tetraceros, BM (NH) 1869.7.8.42. — Gulf of Suez, Cambridge Expedition, R 6, 6. XI. 1924, det. F. A. Potts (1928: 701) Spirobranchus jousseaumi; in 1986, Ben- Eliahu noted opercular wings reduced without crenellations, interradiolar membranes with unilobed processes, characteristic antler-like opercular spines; subsequent re-examination (in 2002) showed the operculum had been cut off at the base of the peduncle, and was missing, 1 spec., CUZM AN. I. 1930. Gulf of Aqaba: 10 samples, [0.2 – 1] – [6 – 8] – [20 – 25] – 25. Elat, shallow up to 2 m, 1955, E 55 / 441, legit H. Steinitz, det. P. Fauvel Pomatoceros coeruleus [sic], specimen lacked tube, operculum and collar chaetae; “ branchia and collar characterized by dark indigo blue colour ” (Fauvel 1957: 10); redet. herein due to presence of interradiolar processes]. Red Sea: South Red Sea, 8 samples, 1 – 27.4 – 36.6 m, HUJ. Indian Ocean: Seychelles, 11 samples, [1 – 7] – [4 – 8] – [12 – 15] – 45 m, ZMA, RMNH. Suez Canal depth and substrates: shallow, 0.4 – [3 – 4] m, on alga: Laurencia; on sponge; on bivalves: Brachidontes pharaonis, Malvufundus regulus, Pinctada radiata and Spondylus spinosus; barnacles; bryozoans; tunicates; rocks; artificial substrates: Canal walls, tin can submerged in mud; rubber fenders and iron frames.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F11E048FF50FE61FC63FBF0.taxon	distribution	Distribution. Circumtropical, but see Remarks. Red Sea: Gulfs of Suez and Aqaba. Lessepsian migrant to the Mediterranean: Israel, Lebanon, Cyprus, Turkey and Rhodes. Colouration (Field notes for Lake Timsah specimen): Ventral side of operculum pink, dorsal side bluebrown. Radioles, from distal to proximal part, with white band, short blue band, white band, brown band, white band at base. Thorax dark red-brown. Noted very pink tube (Great Bitter Lake subsample Biv 11 [App. Table 2 D]). One of us (HAtH) has the impression that individuals with bluish tubes that are buried in living corals and attached to dead Pocillopora damicornis (Linnaeus) skeletons may belong to a different taxon than the pink tube specimens found on various hard substrates, which might lend some support to Pillai’s (2009) distinction between S. tetraceros and S. semperi.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F11E048FF50FE61FC63FBF0.taxon	discussion	Remarks. The taxon Spirobranchus tetraceros as redefined by ten Hove (1970 b) is presently considered to be a complex of species, with some taxa in the Indo-West-Pacific and another in the Caribbean (Smith 1985; ten Hove & Kupriyanova 2009). Some of the Indo-West-Pacific species have been defined as “ S. tetraceros s. str., S. coronatus and S. decoratus ”, but, according to Smith (1985) there may be further subdivision that should be done within the taxon S. tetraceros “ s. str. ” based on characters such as the number of the main spines, colouration of the tube and the tube lumen, the extent of serration of filiform processes on the peduncular wings, and the presence and number of pairs of radiolar ocelli on the radioles. When we were finalizing the present paper, Pillai (2009) published his discussion on Spirobranchus. As he did with other taxa, Pillai split S. tetraceros sensu ten Hove (1970) (i. e., the S. tetraceros- complex) into about 8 different taxa, based on different character combinations (Pillai 2009: 158, notably the discussion on p. 162). In order to evaluate this attempt to clarify the Spirobranchus tetraceros - complex, we compared the main characters used by Pillai with our material from the Suez Canal (see Table 5). By Pillai’s standards, the Suez Canal material is characterized by a unique combination of characters meriting the description of yet another species, intermediate between his definition of Spirobranchus tetraceros and S. semperi. Using Pillai’s criteria, ten Hove's (1970) figures, from a single Bahrain sample, should be named S. tetraceros (figs 23 – 24), S. semperi (figs 25 – 26, 28 – 29) and S. arabicus (figs 1 – 2, 15 – 16). We are not prepared to accept this approach since most of Pillai’s taxa are based upon single samples with 1 – 6 specimens only and variability of various character states has not been studied by Pillai. For instance, Pillai’s (2009) description of S. elatensis, from the Gulf of Aqaba, was distinguished from S. tetraceros mainly by it’s conical operculum, based only on a single specimen. Our work is based on hundreds of samples, with several hundreds of specimens (admittedly not consistently enumerating all character states dealt with recently by Pillai). As regards S. tetraceros, we have found both conical and fully branched opercula and intermediate forms in the same Suez Canal samples, and consider them to belong to a single species (see Figs 33 C – E; see also ten Hove 1970, figs 6, 30 for a Caribbean population of what we nowadays would identify as the Caribbean S. dendropoma Mörch, 1863; cf. Smith 1985: 66). We have noted that the presence / absence of papillae, and the shape of these papillae / fringes, seems to be dependent on their position, dorsal or more ventrally, in the radiolar circles. Without population studies, and without genetic support, we find Pillai’s splitting difficult to accept. Thus, we prefer not to adopt the classification given by Pillai (2009), but to retain the more “ traditional ” concept of this taxon as both in the present paper and Selim (1997 b), thus synonymising S. elatensis with S. tetraceros s. str. A review of all the literature citations, taking into consideration Smith’s (1985) and Pillai’s (2009) hypotheses is not within the province of the present paper. However, when the review of this taxon will finally be carried out, it should comprise both morphological and genetic evidence from populations. 1) No. of spines sensu ten Hove 1970; in ten Hove & Kupriyanova (2009) these structures were referred to as horns, — 2) See Bailey-Brock (1985, fig. 8 e) or Pillai (2009, fig. 37 c) and Fig. 33 D. The character was never looked for consistently, but ten Hove’s unpublished notes mention it was present in material from New South Wales to Northern Territory, Australia, — 3) Information missing in Pillai (2009), but present in Bailey-Brock (1985: 204 – 207, fig. 8), — 4) Data from ten Hove's sketches of the types of S. coronatus, — 5) From ten Hove's notes on the type material; the 4 th specimen from Heron Island, Queensland, Australia, det. S. coronatus by D. Straughan is a specimen exactly like S. " baileybrockae ”, — 6) Pillai (2009, figs 48 b, c) suggest 1 medioventral (split in two) and two laterodorsal split in two; thus 6 horn tips, — 7) A double fringe was only figured before by Selim (1997 b, fig. 7 b); see Fig. 33 D, — 8) According to Pillai (2009: 178) four spines: 2 ventral spines and 2 dorsal ones with short mediodorsal tine. We interpret his figures 57 c, d slightly otherwise, — 9) One also might argue that there are 4 opercular spines: 2 ventral ones split twice and two laterodorsal ones split twice of which the 1 st branch is a short mediodorsal tine. Regional citations given above are for the species complex as known subsequent to ten Hove (1970 b). We should note, however, that fig. 16 S. j. of Amoureux et al. (1978: 150) from the Gulf of Aqaba, Elat, Venezia lagoon appears to be that of Spirobranchus coronatus; thus, both S. tetraceros and S. coronatus are present in the northern Gulf of Aqaba. Spirobranchus polytrema not Philippi, sensu Amoureux et al. (1978: 150) is corrected to S. tetraceros based on the illustration marked S. p. in his fig. 16; thus the nominal taxon S. polytrema, was not a Mediterranean migrant to the Red Sea via the Suez Canal as stated by these authors. The Suez Canal population appears to belong to a single taxon (Figs 27 A – E), a fouling species characterised by small peduncular wings with a pointed tip, generally fringed (Figs 27 A, D, E), sometimes with a double fringe (clearly visible in Fig. 33 D, right wing; see note Table 5); interbranchial membranes usually with unilobed processes and the specimens examined from the Levant Basin of the Mediterranean appear to belong to the same taxon, known also from the Gulf of Aqaba and the Gulf of Suez (Ben-Eliahu 1991, fig. 2). Examination of live material from Elat showed two dense rows of bright crimson ocelli with minute lenses on each side of the rachis of the radioles (base of prostomium not looked for in this live specimen), suggesting the Elat population might be the same as Smith’s “ Spirobranchus tetraceros species C ” (Smith 1985, Table 5). Selim (1997 b fig. 7) remarked and illustrated the polymorphism of the operculum of S. tetraceros, and we have also included examples in Figs 33 A – E. Ten Hove (1970 b: 4, 48) noted that Pixell (1913) had dealt with the variability of Spirobranchus tetraceros opercula of the C. Crossland collection in the Natural History Museum, London by giving them a variety of names (see synonymy above). In their ranking of the Lessepsian migrants within the category, “ Worst Invasive Species ”, Zenetos et al. (2005) appear to underestimate the importance of Spirobranchus tetraceros, known from Beirut from 1965 (Laubier 1966, as Spirobranchus giganteus coutierei), and as far as Rhodes already in 1970 (Ben-Eliahu 1991, collected by the “ Biota ” project). Selim et al. (2005) reported that S. tetraceros had replaced Hydroides elegans as the dominant fouling species in Alexandria Mediterranean waters, such that it amply fulfils the criterion “ having an serious impact on diversity ” (Zenetos et al. 2005).	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F18E049FF50FBEEFE02FE20.taxon	description	In view of these taxonomical problems, morphology cannot be used to determine the provenance of Vermiliopsis infundibulum / glandigera / pygidialis in the Suez Canal [contrary to the somewhat formalistic distinction by Pillai (2009: 108, fig. 8), made on literature comparison only]. In contrast, the Mediterranean / Atlantic species, Vermilopsis striaticeps (Grube, 1862), can be distinguished from V. infundibulum (e. g., Zibrowius 1979 a) by marked taxonomic characters, although it has been confused with this taxon in the past. For example, Monro (1930: 209 – 211, fig. 88), figured four opercula in describing his “ Vermiliopsis glandigerus ” from the Atlantic Gulf of Guinea, two of them with clear longitudinal striations belonging to Vermiliopsis striaticeps, the other two belonging to the Vermiliopsis infundibulum / glandigera complex (material studied by HAtH). It appears that the Vermiliopsis infundibulum / V. glandigera - complex and V. striaticeps are both present in the Mediterranean and also in the Suez Canal. Vermilopsis striaticeps was reported from Deversoir [as V. infundibulum in Selim (1997 b), citing Ghobashy et al. (1980) and Ghobashy & Ghobashy (2005), who referred to the same single specimen]. The excellent illustration in Selim (1997 b, fig. 9 a) enables confirming this specimen confidently as Vermiliopsis striaticeps. However, that finding called into question other citations based on Selim’s identifications, e. g., those of Vermiliopsis infundibulum from the Mediterranean. For example, Selim (1997 b: 97) cited a record of Ghobashy et al. (1980) from the Suez Canal for Vermiliopsis infundibulum without providing morphological detail. Similarly, it is not possible to know whether the record from Alexandria harbour (Ghobashy 1984) belongs to the Mediterranean Vermiliopsis infundibulum or V. striaticeps. Determinations of Vermiliopsis infundibulum based on empty tubes must also be considered as questionable and are therefore cited with a question mark before the date of publication in the synonymy section below.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F19E076FF50FDD9FAECFAEB.taxon	description	figs 25 a – h. Vermiliopsis infundibulum glandigera - group: Imajima 1976: 139 – 141, fig. 11. Eastern Mediterranean	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F19E076FF50FDD9FAECFAEB.taxon	description	? Vermi [o] liopsis infundibulum: Emara & Belal 2004: 196, 199 [pooled Lake Timsah and Bitter Lakes data]. Vermiliopsis glandigera: Abd-Elnaby 2009: 10 [reported from southern part of the canal and Red Sea]. Gulf of Suez and Gulf of Aqaba	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F19E076FF50FDD9FAECFAEB.taxon	description	Cheval Paar].	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F19E076FF50FDD9FAECFAEB.taxon	materials_examined	Material examined. Locations adjacent to the Suez Canal, Mediterranean side: Vermiliopsis infundibulum: Rhodes 1 sample, shallow — Turkey 2 samples, [108 – 135] m — Cyprus 59 samples, [3 – 5] – [18 – 78] – 2432 m — Israel 30 samples, 1 – [18 – 66] – 1435 m — Egypt (incl. Sinai) 6 samples, 18 – [137 – 198] – [1993 – 2025] m, HUJ and SMF. Suez Canal material reported herein: 2 samples, 2 specimens. Hebrew University-Smithsonian Expeditions, 1967 – 1973: Exp. 11. VII. 1968: SLC 45, 1 spec. — East of Deversoir – Km 97: SLC 71, 1 spec. Locations adjacent to the Suez Canal, Red Sea side: As Vermiliopsis infundibulum / glandigera (see introductory note): Gulf of Suez, Suez, det. H. L. M. Pixell (1913: 86 – 87) V. pygidialis, redet. ten Hove (1975: 68) Vermiliopsis infundibulum / glandigera, BM (NH) 1924.6.13.140, 2 specs — Gulf of Aqaba: 14 samples, [intertidal – 3] – [6 – 9] – [22 – 25] m. Red Sea proper 9 samples, 1 – [5 – 15] – 50 m, HUJ. Indian Ocean, Seychelles 18 samples, [1 – 7] – [4 – 11] – [30] – [45 – 55] m, ZMA. Suez Canal depth and substrates:? from pool and “ sublittoral ”, on alga; on Pectinidae.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F19E076FF50FDD9FAECFAEB.taxon	discussion	Remarks. Particularly the non-Atlantic-Mediterranean Vermiliopsis taxa are in need of revision.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F26E077FF50FAE1FB55FD9E.taxon	materials_examined	Material examined. Locations adjacent to the Suez Canal, Mediterranean side: Turkey, 2 samples, 0.5 m — Rhodes, 1 sample — Cyprus, 26 samples [0.5 – 0.8] – [3 – 5] – 46 m — Israel, 36 samples, intertidal – 10 – 67 m. No additional specimens from the Suez Canal in addition to those reported by Selim (1997 b) as Vermiliopsis infundibulum.	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
396387E75F26E077FF50FAE1FB55FD9E.taxon	distribution	Distribution. Mediterranean, Atlantic (from the Gulf of Guinea to Cornwall, U. K.)	en	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): 1-147, DOI: 10.11646/zootaxa.2848.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2848.1.1
