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Deepwater Tube Worms (Polychaeta, ) from the Hawaiian Islands1 JULIE H. BAILEy-BROCK2

THREE SERPULID TUBE WORMS have been dis­ (1906), but no serpulids were found. Hart­ covered on shells and coral fragments taken in man (1966a) reviewed the literature in an dredges from around the Hawaiian Islands. The extensive analysis of the Hawaiian two serpulines latiscapus Maren­ fauna. Straughan (1969), presented a more zeller and infundibulum Philippi recent survey of the littoral and upper sublit­ are new records for the islands. However, the toral Serpulidae. Other works by Vine (1972) small spirorbid Pileolaria (Duplicaria) dales­ and Vine, Bailey-Brock, and Straughan (1972) traughanae Vine has been described previously include ecological data collected from settle­ from within diving depths (Vine, 1972), but ment plates and by diving, but no records ex­ it is absent from shoal waters and intertidal re­ tend below 28 meters. Serpulids have been gions. 3 The occurrence of this species in the described from deepwater collections in other dredged collections indicates an extensive depth parts of the world. Southward (1963) found range in the Hawaiian Islands. 14 species of calcareous tube worms on hard The tube worms were obtained from col­ substrata dredged from depths as great as 1,755 lections taken during two separate oceano­ meters along the continental shelf off south­ graphic investigations in Hawaiian waters. western Britain. Antarctic collections yielded 14 Material consisting mostly of the pink serpuline genera and more than 23 species from depths Spirobranchus latiscapus was loaned by Dr. E. C. ranging from the littoral zone down to 4,930­ Jones of the National Marine Fisheries Service 4,963 meters in the South Sandwich Trench (N.M.F.S.) and was taken from an average (Hartman, 1966b, 1967). depth of 200 meters (lat 21 °9.6' N, long 157° 25.1' W). Examination of these samples, which DESCRIPTIONS OF SPECIES consisted of both living and preserved speci­ mens, proved very interesting. The other collec­ SpirobranchltS latiscapus has been included in tion contained all three serpulid species attached the genus Pomatostegus by von Marenzeller to the branches of the precious pink coral Coral­ (1885) and other authors, secondly in the lium secundum and on the solitary coral Cya­ genus Spirobranchus by Imajima and Hartman theras sp., both of which were collected by V. E. (1964) and other earlier authors, and possibly Brock during the 1966 Sange Pele survey for identified as Pomatocerus strigiceps by McIntosh locating areas of precious coral on Penguin (1885). H. A. ten Hove (1970) reviewed the Banks off Molokai (lat 20°51.6' N, long 157° literature on this species and retained the generic 19.3' W; depth, 220-600 meters). status, Spirobranchus. The specimens of S. la­ Previous literature on Hawaiian tiscapus from the N.M.F.S. were mainly associ- -,.,co""n"'st

in color from pale pink to a deep salmon orange. to the tube opening (Fig. Ie). Among seven There is an obvious median, longitudinal ridge specimens from the Sange Pele dredge haul with blunt, triangular to rectangular teeth, (depth between 220-600 meters), both adult which are most well developed toward the and juvenile stages were found. Hollow, cup­ mouth (Fig. la). Where the tube has grown shaped, calcareous brood chambers develop prior up off the substratum, as seen in a few speci­ to breeding and one specimen was incubating mens, there are three or more finely toothed two embryos. This is perhaps the most open ridges on either side of the median ridge (Fig. type of brood chamber that has been described la). Erect portions of tube are circular in cross in the Spirorbinae literature. Vine (1972) said section; attached portions are typically triangular that one of the specimens he found had six in section. developing embryos, but only two could fit in The opercula are composed of 1 to 7 cal­ the confines of the opercular chamber (Fig. If). careous tiers, which get smaller toward the This would seem to be a hazardous method for apex (Fig. Ib). These tiers comprising the incubating the developing trochophore larvae, operculum can be dislodged and some of the except under the most ideal circumstances. largest specimens were lacking one or more of the older tiers. The opercular stalk has broad DEPTH AND ZOOGEOGRAPHICAL RANGE lateral wings. There is an incomplete collar, seven thoracic segments, and a large abdomen H. A. ten Hove (1970) reviewed the geo- with as many as 48 setigerous segments. graphical range of Spirobranehus latiseapus, a The color of living specimens is an overall range which extends throughout the Indo-West pink-red. The opercular stalk has red patches Pacific. Station records from Moore and Bush extending over the wings; the opercular plates (1904) and Takahasi (1938) showed that this vary from shades of orange to yellow. The serpulid occurs between 80-150 meters. Hawaii are red at the bases and have red bands represents the most easterly record of this on an orange background extending to the tips. species in the Pacific. My observations agree with those of Takahasi The type locality of Vermiliopsis infundibu­ (1938), who described this serpulid from Japan lum is the Mediterranean Sea, but it is found in as Pomatostegus latiseapus. the Atlantic, Indian, and Pacific oceans (Zibro- The Hawaiian specimens of Vermiliopsis in- wius, 1968). The shallowest record is by Hart­ fundibulum Philippi agree with the descriptions man (1969), who found the species from by Fauvel (1927), Zibrowius (1968), and Hart- within diving depths off Santa Catalina Island, man (1969: 779-780). The whitish tubes have California. Southward (1963) found three spe­ a characteristic fluted appearance with an ex- cies of Vermiliopsis, including V. infundibu­ panded flute forming the mouth (Fig. Ie). The lum, taken from depths between 205-1,430 operculum is a ringed, chitinous dome, golden meters on the continental shelf. This cosmopoli­ brown in color, and the stalk lacks lateral wings tan distribution may be a result of water tem­ (Fig. Id). perature and current systems, as well as available Pileolaria (Dupliearia) dalestraughanae, an substrata for larval settlement. The vertical opercular-brooding spirorbid, occurred in both range of Pileolaria (Dupliearia) dalestraugha­ San e Pele and N.M.F.S. collections. This spe- nae is extensive. The species has a known dis------=c"'"":ie--=s"--w-a=-=s--:r--=e-=-ce:-:n:-;tl1:y-=-d'e-=-s=-=c"='ri'Lb-=-edTLb--:y'V."..-in=-=e-:-T(1;-;9""7"2")...---r"r!'oC::mC:------"'tr"'1L:u--,-;t,...-lo,-n-m--:~Hr:a:=w=-ai'l from 8 meters (Vme,T97T)c------1 Maili Point and Pokai Bay, Oahu, from depths to between 220-600 meters off Penguin Banks. of 8 and 15 meters, respectively. The tube coils The upper limit of this considerable range may sinistrally (clockwise) in a flat spiral and has be determined by the degree of water movement four distinct longitudinal ridges which extend and possibly by the amount of suspended ma-

FIG. 1. Spirobranchus latiscapus: (a), three tubes attached to a carrier shell; (b) tiered operculum with winged stalk. Vel'miliopsis infundibulum: (c), tube; (d), ringed operculum, Pileolaria (Duplica.·ia) dale­ stl'aughanae: (e), sinistral tube; (f), adult operculum containing eggs. NOTE: e and f after Vine (1972). Deepwater Tube Worms-BAILEy-BROCK 407

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10.25 mm 0·125 mm E 408 PACIFIC SCIENCE, Volume 26, October 1972

terial that could obstruct the adult brood HOVE, H. A. TEN. 1970. Serpulinae (Poly­ chamber. As the operculum is merely a modified chaeta) from the Caribbean. I, The genus it is extruded from the protection of the Spirobranchus. Stud. Fauna Curac;ao. 32 tube every time the extends the branchial (117) :1-57. crown during feeding. Spirorbinae that brood IMA]IMA, M., and O. HARTMAN. 1964. The within the confines of the tube in an egg string polychaetous of Japan. ace. Pap. or thoracic pouch are not so vulnerable to loss Allan Hancock Fdn. 26:1-452. of embryos (Bailey, 1969). The relatively calm MARENZELLER, E. VON. 1885. Siidjapanische water off Oahu's leeward coast appears to per­ Anneliden. 2. Ampharetea, Terebellacea, mit an unusually shallow distribution of this Sabellacea, Serpulacea. Denkschr. Akad. spirorbid. P. (D.) dalestraughanae has been Wiss. Wien 49 (2) :197-224. recorded from 20-27 meters off southern Africa McINTOSH, W. C. 1885. Annelida Polychaeta. and is tentatively identified in collections from In Report on the Scientific Results of the the Cape Verde Islands by Phyllis Knight-Jones Voyage of H.M.S. Challenger during the (personal communication). These records give Years 1872-1876. Vol. 12, 689 p., 94 pI., some idea of the geographical range of this 1 map. Reprint, 1966. Johnson Reprint species. Corp., New York. I am grateful to Dr. E. C. Jones (N.M.F.S.) MOORE, J. P., and KATHARINE J. BUSH. 1904. for both preserved and live and for and Serpulidae from Japan, with reading the manuscript; to Drs. H. A. ten Hove, descriptions of new species of . Proe. Zoologisch Laboratorium, Utrecht, and J. H. Acad. Nat. Sci. Philad. 36:157-179. Day, University of Cape Town, for their as­ SOUTHWARD, EVE C. 1963. Some new and little­ sistance with the identification of Spirobranchus known serpulid polychaetes from the conti­ latiscapus; and to Dr. Olga Hartman, Allan nental slope. J. Mar. BioI. Ass. U.K. 43:537­ Hancock Foundation, for identifying Vermili­ 587. opsis infundibulum. The author also wishes STRAUGHAN, D. 1969. Serpulidae (Annelida: to acknowledge a University of Hawaii Biomed­ Polychaeta) from Oahu, Hawaii. Bull. S. ical Support Grant, 1970/71. Calif. Acad. Sci. 68 (4) :229-240. TAKAHASI, K. 1938. Polychaetous of LITERATURE CITED Izu Peninsula. 1. Polychaeta, collected by the "Misago" during the Zoological survey BAILEY, JULIE H. 1969. Methods of brood pro- around the Izu Peninsula. Sci. Rep. Tokyo teetion as a basis for the reclassification of the Bunrika Daig. B, 3 (57) :193-220. Spirorbinae (Serpulidae). J. Linn. Soc. TREADWELL, A. 1906. Polychaetous annelids of (Zool.) 48:387-407. the Hawaiian Islands, collected by the Steamer FAUVEL, P. 1927. Polychetes sedentaires. Faune Albatross in 1902. Bull. U.S. Fish Comm. Fr. 16:1-494. 23:1145-1181. HARTMAN, O. 1966a. Polychaetous annelids of the Hawaiian Islands. ace. Pap. Bishop Mus. VINE, P. J. 1972. Spirorbinae (Polychaeta, Ser- 23 (11) :163-252. pulidae) of the Hawaiian chain. Part 1, new ___. 1966b. Polychaeta Myzostomidae and species. Pacif. Sci. 26 (2) :140-149. ----'S'e'd-en-'-;t,-a--.ri~a-o'f,---"A,-n--ft~ar::-:c-;-.ti-=-ca-.-A-.-'-n~ta'-r'ct~1Cc-TR'-e-"-se""'a'-rc"CTh-VINn,-P;-J:;-JtJIoI-E-H-;-B:AItE-Y~BR0eK-,--a:nd-9-;-.-- Series, vol. 7. Amer. Geophys. Union, Wash- STRAUGHAN. 1972. Spirorbinae (Polychaeta, ington, D.C., publ. 1414. 158 p. Serpulidae) of the Hawaiian chain. Part 2, ---. 1967. Polychaetous annelids collected Hawaiian Spirorbinae. Pacif. Sci. 26 (2): by the USNS Eltanin and Staten Island 150-182. cruises, chiefly from Antarctic seas. Allan ZIBROWIUS, H. 1968. Etude morphologique, Hancock Monogr. Marine BioI. 2 :1-387. systematique et ecologique des Serpulidae ---. 1969. Atlas of the sedentariate poly- (Annelida: Polychaeta) de la region de Mar- chaetous annelids from California. Allan seille. Ree. Trav. St. Mar. End. Bull. 43 (59) Hancock Fdn., Los Angeles. 812 p. 81-252.