Zootaxa, Redescription of Cirriformia crassicollis ...

Zootaxa 2625: 53–62 (2010) www.mapress.com / zootaxa/

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Redescription of Cirriformia crassicollis (Kinberg, 1866) and Timarete hawaiensis (Hartman, 1956) new combination, (Polychaeta: Cirratulidae), endemic polychaetes to the Hawaiian Islands WAGNER F. MAGALHÃES1 & JULIE H. BAILEY-BROCK1,2,3 1

Water Resources Research Center, University of Hawaii at Manoa, 2540 Dole Street, Honolulu, Hawaii 96822, USA Department of Zoology, University of Hawaii at Manoa, 2538 McCarthy Mall, Honolulu, Hawaii 96822, USA 3 Corresponding author. E-mail: [email protected] 2

Abstract Cirriformia crassicollis (Kinberg 1866) and Timarete hawaiensis (Hartman 1956), new combination are redescribed. Cirriformia hawaiensis is here referred to the genus Timarete Kinberg 1866, because the branchial filaments on chaetigers 10–18 are shifted dorsally forming a dorsolateral bulge over the notopodia. T. hawaiensis and C. crassicollis are presently known only from the Hawaiian Islands and are commonly associated with changing ecosystems such as harbors and dense populations of tube worms. Key words: taxonomy, endemism, Oahu Island, Hawaiian shallow waters

Introduction Three species of the family Cirratulidae have been described from the Hawaiian Islands: Cirriformia crassicollis (formerly Labranda crassicollis Kinberg 1866), Cirriformia semicincta (formerly Cirratulus semicinctus Ehlers 1905) and Cirriformia hawaiensis Hartman 1956 (formerly Audouinia branchiata Treadwell 1943). Cirriformia semicincta appears to have a circumtropical distribution (Hartman 1956) while C. crassicollis and C. hawaiensis are endemic to the Hawaiian Islands. The latter two species are poorly known taxonomically as the original and subsequent description (Kinberg 1866; Hartman 1948, 1966; BaileyBrock 1987) were based on highly variable morphological characters which change during ontogeny and accurate illustrations are lacking. In this paper we redescribe these two species following a review of the type material from museum collections and newly collected specimens. We transfer Cirriformia hawaiensis to the genus Timarete Kinberg 1866 and the highly variable ontogenetic characters (e.g. segmental origin of tentacular filaments, neuroacicular spines, and notoacicular spines) were measured to evaluate size-dependency.

Material and methods The type specimens were provided by the American Museum of Natural History, New York, USA (AMNH) and the Swedish Museum of Natural History, Stockholm, Sweden (SMNH); additional specimens were from the collections of the Wormlab, University of Hawai’i at Manoa, Hawaii, USA. These have been deposited at the United States National Museum of Natural History, Smithsonian Institution, Washington, D.C., USA (USNM) and SMNH. The characters analyzed were body length and width, total number of chaetigers, position of the tentacular filaments, first occurrence of acicular spines on notopodia and neuropodia, number of acicular spines on Accepted by P. Hutchings: 6 Aug. 2010; published: 24 Sep. 2010

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anterior and posterior parapodia, and where along the body the branchial filaments shift to a mid-dorsal location (only for T. hawaiensis). These characters were recorded from numerous complete specimens in order to determine how these characters vary with size and presumably age by correlation analyses. Individuals of both species were dehydrated through a series of increasing concentration of ethanol followed by two changes of absolute EtOH with subsequent critical point drying (in a SAMDRI-795) for Scanning Electron Microscopy (SEM) analysis. Specimens were mounted on stubs and coated with gold/ palladium. SEM observations were carried out using the Hitachi S-4800 at the Biological Electron Microscopy Facility (BEMF), University of Hawaii at Manoa.

Results Cirriformia crassicollis (Kinberg, 1866) Figures 1 (A, B, and E) and 2 (A–F) Labranda crassicollis Kinberg, 1866: p. 255. Cirriformia crassicollis; Hartman, 1948: p. 112; Hartman, 1966: p. 226; Bailey-Brock, 1987: p. 371.

Material examined. Type material: Holotype (SMNH 498), from Honolulu, 21°19′ 157°52′ (Honolulu Harbor) Leg. Eugenie Exp. 1851–53 sta. 1074-77. Non-type specimens reported by Bailey-Brock (1984) and Dreyer et al. (2005) from samples collected at Niu Valley Beach Park, south coast of Oahu Island, 21°16′41′′ 157°44′40′′, coll., S. A. McCarthy, W. Estabrooks; #2 Jan 1979 (5, USNM 1145381), #3 Jan 1979 (1, USNM 1145380), #10 Sep 1992 (5, SMNH 110719), #2 Feb 1977 (7), #3 1978 (11), #3 Jan 1979 (2), #1 1979 (4), #2 (33); 11 Sep 1992 (pre Iniki hurricane) #1 (2), #3 (6), #5 (8), #7 (7), #9 (12); 5 Sep 1992 (post Iniki hurricane) #1 (3), #2 (14), #4 (2), #5 (10), #6 (4), #8 (4), #10 (5); Hanauma Bay, Keyhole, 21°16′14′′ 157°41′45′′, from hard substrate, 17 Aug 1999, coll. J.H. Bailey-Brock (1). Redescription of holotype. Holotype fragmented in several regions, containing body walls and body fragments without the body wall, with all in poor condition. Hartman (1948) previously described these fragments. Prostomium had a few dark spots (Hartman, 1948) but now, the holotype lacks the prostomium. Tentacular filaments arise above chaetiger 5 in two patches of filaments (5–7 each). Fragment 2.8 mm long, 2.0 mm wide (from left notopodium to right notopodium); six chaetigers remain on right side and ten chaetigers on left. Notopodium and neuropodium with yellow capillary chaetae number of 8–12 per fascicle. Larger piece of body wall (6.0 mm long with 4.6 mm wide) with only capillary chaetae in first chaetigers (eight chaetigers on neuropodium and 17 on notopodium only with capillaries). After that, 3 or 4 acicular spines alternate with 2–3 capillaries present. Distal part of this fragment with branchiae arising at a discrete distance from notopodial ridge (also reported by Hartman 1948). Largest body fragment lacks body wall and about 7 mm long and 2 mm wide with more than 30 chaetigers. Fragment rounded dorsally, ventral groove. Lateral projection of parapodium forms lateral shoulder. Yellow, slightly curved acicular spines present in fragment. Redescription. Specimens 2.5–50.0 mm long, 0.1–4.0 mm wide for 33–246 chaetigers. Body elongated, dorsally inflated, ventral surface with deep groove formed by projection of neuropodia ventrally. Smaller specimens with ventral surface flattened. First four chaetigers without distinct segmental markings, with dorsal surface resembling a dorsal crest. Crowded segments with well developed lateral shoulders. Color in alcohol yellow to tan, with branchial and tentacular filaments pale; color in life not observed. Prostomium triangular, pointed, almost as long as wide with pair of nuchal organs located near posterolateral border (Fig. 1A, B; 2A, B). Peristomium with two annuli and sub-annulations, the first annulus as large as four thoracic chaetigers and second one as large as two thoracic chaetigers (Fig. 1B; 2B). Junction between peristomial annulations deeply grooved (Fig. 1B; 2B). Tentacular filaments formed by two groups of filaments (8–12) organized in two rows each, dorsally located on chaetiger 4 or 5 on most specimens. Juveniles with tentacles on chaetiger 3 (Fig. 2F). Typically, two clusters positioned on only one chaetiger and this appears to be size-dependent (Fig. 1A and 3B). First

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MAGALHÃES & BAILEY-BROCK

pair of branchiae on chaetiger 1, one branchia per parapodium, well above notopodial ridge, more numerous on anterior end of body, present in posterior chaetigers. Largest specimen with branchial filaments continuing after chaetiger 50, inserted at discrete distance from notopodial ridge. However, branchial filaments do not become mid-dorsal as in Timarete species, but remain inserted laterally.

FIGURE 1. Cirriformia crassicollis. A, anterior end in dorsal view; B, anterior end in lateral view; E, Methyl green staining pattern of the anterior end. Timarete hawaiensis. C, anterior end in dorsal view; D, anterior end in lateral view; F, Methyl green staining pattern of the anterior end.

REDESCRIPTION OF TWO HAWAIIAN CIRRATULIDS

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Notopodia and neuropodia widely separated. On first five chaetigers, notopodia more dorsal and neuropodia more lateral; from chaetiger 6 notopodia become more lateral and neuropodia ventral (Fig. 2B). Anterior parapodia with large, yellow serrated capillary chaetae arranged in two rows (Fig. 2C). Notoacicular spines first present in neuropodia in all specimens examined from chaetigers 8–34 with an average gap of approximately 10 chaetigers until their appearance in notopodia (Fig. 3). Acicular spines amber in color, slightly curved on anterior end; 3–4 spines (rarely 5) alternating with 2–3 capillary chaetae throughout (Fig. 2D). Pygidium simple ventral lip with terminal anus (Fig. 2E).

FIGURE 2. SEM of Cirriformia crassicollis showing: A, anterior end in dorsal view; B, anterior end in lateral view; C, anterior serrated capillary chaeta; D, mid-body neuroacicular spines alternated with capillary chaeta; E, posterior end and pygidium in dorsal view; F, a juvenile specimen.

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Methyl green staining pattern. Prostomium with anterior tip not staining; dorsal junction with peristomium intensely stained (Fig. 1E). Second peristomial annulation and anterior chaetigers before dorsal tentacles staining intensely showing dorsal crest (Fig. 1E). Branchial and tentacular filaments staining weakly. Mid-segmental areas staining intensely while inter-segmental areas not staining. Notopodial and neuropodial ridges not staining while parapodia stain intensely. Ventral groove weakly stained.

FIGURE 3. Variation of size-dependent characters in Cirriformia crassicollis. A—Relationships among the commencement of notoacicular and neuroacicular spines and the total number of chaetigers in complete specimens. B— Relationships among the body length (in mm), the number of chaetigers, and the position where the tentacular filaments begin (chaetiger 3, 4, 5 or 6) in complete specimens.

Remarks. None of the current descriptions of C. crassicollis (i.e. Kinberg 1866, Hartman 1948, 1966, Bailey-Brock 1987) are sufficiently detailed to correctly identify this species. Cirriformia crassicollis differs from other congeners by the general shape of the adult body including the well developed parapodial ridge forming “shoulders”, and a deep ventral groove along the body; the tentacular filaments occur in only one chaetiger (chaetiger 4 or 5 in adult specimens), there is a gap of approximately ten chaetigers between the commencement of neuro- and notoacicular spines, and a distinct methyl green staining pattern that shows a strong reaction at the junction between the prostomium and peristomium and on the second peristomial annulation. Juvenile individuals of this species presented only capillary chaetae throughout, less than 40 chaetigers, and tentacular filaments located on chaetiger 3 (Fig. 2F). Additionally, the branchial filaments of juveniles had a few black spots sparsely distributed over their length. Blake (1996) reported different bifid spines in juveniles of Cirriformia tentaculata and C. moorei that were not present in adults. Cirriformia crassicollis differs from the other Hawaiian species C. semicincta (Ehlers 1905) on the position of the tentacular filaments. In C. semicincta the tentacular filaments arise over chaetigers 3 and 4 (Abbott 1946, Okuda 1937) while in C. crassicollis they arise above only one chaetiger, generally 4 or 5. Biology. Cirriformia crassicollis were collected in Diopatra dexiognatha Paxton & Bailey-Brock, 1986 mounds; dense aggregations were found in a specific intertidal area of Honolulu. Cirriformia crassicollis (as Cirriformia sp. – Bailey-Brock 1984 and as Cirratulidae – Dreyer et al. 2005) was one of the most numerous polychaetes in these communities. Some specimens of C. crassicollis were found actually inhabiting Diopatra tubes (Bailey-Brock, pers. obs.). Bailey-Brock (1979) reported this species as a component of chaetopterid mounds on a fringing reef, south shore of Oahu, reaching densities of 200 ind./m². Distribution. The type locality for C. crassicollis is Honolulu, Oahu, Hawaii. A new record from Halape, Big Island, Hawaii, was reported by Hartman (1966) and Bailey-Brock (1979) reported C. crassicollis from Fort Kamehameha Ahua reef near Pearl Harbor entrance channel, Oahu, Hawaii. All the specimens herein analyzed were from Niu Valley Beach Park, south coast of Oahu, and Hanauma Bay, east coast of Oahu, Hawaii. Therefore, C. crassicollis is endemic to the Hawaiian Islands.



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Timarete hawaiensis (Hartman, 1956), new combination Figures 1 (C, D, and F) and 4 (A–F) Audouinia branchiata Treadwell, 1943: p. 1, Figs. 1–3. Cirriformia hawaiensis Hartman, 1956: p. 293. Cirriformia hawaiensis; Hartman, 1966: p. 226; Bailey-Brock, 1987: p. 371, plate 3.II.3.c.

Material examined. Type material: Holotype (AMNH 3260), Pearl Harbor, Oahu, Hawaii, 1925, coll. A.E. Verrill. Non-type material: Pearl Harbor, Oahu, Hawaii, 1925, coll. A.E. Verrill (47, AMNH 3280); Hospital Point, Pearl Harbor, Oahu, Hawaii, 21°20′ 56′′ 157°58′ 07′ ′ , 2m, Oct 1996, coll. R. DeFelice (2, USNM 1145382); Hull of Stoddart, Pearl Harbor, Oahu, Hawaii, 21°21′20′′ 157°58′11′′, Nov 2000, coll. R. Brock (3); Kaimana Beach on Gracilaria salicornia mats, Oahu, Hawaii, 21°15′49′′ 157°49′21′′, 9/04/09, coll. C. Moody (2), Kahuku, North shore of Oahu, Hawaii, removed from oyster shells, 12/27/97, 21°41′22′′ 157°56′35′′, coll. J.H. Bailey-Brock (4); Hanauma Bay, Keyhole, Oahu, Hawaii, 21°16′11′′ 157°41′41′′, from hard substrate, 8/ 17/99, coll. J.H. Bailey-Brock (3). Redescription. Holotype 40 mm long, 2.2 mm wide for about 186 chaetigers. Other specimens 5–43 mm long, 0.8–2.8 mm wide for 66–182 chaetigers. Body rounded dorsally, flattened ventrally, narrower on posterior one-third in comparison with anterior end. Some specimens with shallow ventral groove on last chaetigers. Anterior and posterior ends with segments concertined. Distinct segments throughout. Color in alcohol yellow to tan; black spots on prostomium, peristomium and mid-body segments. Branchial and tentacular filaments pale yellow. Some specimens with darker area over dorsal surface including branchial filaments from posterior end of the body. Color in life not known. Prostomium rounded, wider than long, without eyes (Fig. 4A). Some specimens with row of pigment spots along anterior-ventral margin of peristomium and chaetigers 1–3. Peristomium formed by three annulations with first one dorsally inflated resembling dorsal crest. Tentacular filaments in two widely separated groups, each dorsally located on chaetigers 3–4 in all specimens analyzed (Figs. 1C and 4A, D), not size-dependent. Each patch with about 7–9 filaments arranged in two rows oblique to chaetigers 3 and 4, with more filaments on chaetiger 4. Branchial filaments first present on third peristomial annulation, one pair per segment; on chaetiger 1 positioned above notopodial ridge (Fig. 4D). On chaetigers 10–18, branchial filaments shifted from near notopodium to mid-dorsum, with a larger distance than that between noto- and neuropodia from notopodium (Figs. 1D and 4B). Dorsolateral bulge over notopodium on chaetigers 1–10 shifted to mid-dorsum on chaetiger 10–18. Branchial filaments numerous on anterior end and few to absent on posterior body. Shifting of branchial filaments not a sizedependent character (Fig. 5C). Parapodia well developed on anterior chaetigers forming distinct shoulder; this elongated forming lateral bulge where branchial filaments shift to mid-dorsum (Figs. 1D and 4B). Noto- and neuropodium distinctly separated; notopodium lateral and neuropodium ventral throughout. Serrated capillary chaetae more abundant on anterior parapodia, with 6–8 capillaries (Fig. 4F). Neuropodial acicular spines from chaetiger 8–19 while notoacicular spines from chaetigers 21–78; appearance of acicular spines size-dependent (Fig. 5A). In several specimens, neuroacicular spines begin where branchial filaments shift dorsally above notopodia. Anterior acicular spines 3–4 per neuropodium, accompanied by 1–2 capillaries; fewer aciculars posteriorly, with 2–3 spines per neuropodium, rarely accompanied by capillaries. Anterior notoacicular spines 2–3 (rarely 4) per notopodium, with 1–2 acicular spines and rarely one capillary on far posterior chaetigers. Number of acicular spines on anterior and posterior noto- and neuropodia constant in relation to number of chaetigers. Neuroacicular spines curve distally, more robust than notoacicular spines (Fig. 4C). Notoacicular spines pale yellow, straight, fragile, and mostly broken. Pygidium with terminal anus (Fig. 4E). Methyl green staining pattern. Entire body including branchial and tentacular filaments staining slightly. Prostomium not staining while first peristomial annulation stained intensely. The second peristomial annulation not stained dorsally (Fig. 1F). Notopodial and neuropodial ridges not stained.

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Remarks. The transfer of Cirriformia hawaiensis to the genus Timarete was made based on the dorsal position of the branchial filaments on mid-body segments. This characteristic was previously reported in the descriptions by Treadwell (1943) and Hartman (1956, 1966). Timarete punctata (Grube 1859) is a common species in Hawaii and differs from T. hawaiensis, new comb., by the body color. Timarete punctata is dark brown with black irregular spots throughout and tentacular filaments with black lateral stripes (Çinar 2007). Whereas, T. hawaiensis has dusky dark brown pigment only on the ventral surface of the peristomium; the branchial and tentacular filaments of alcohol preserved specimens are pale yellow. The tentacular filaments arise on chaetigers 3 and 4 on both species.

FIGURE 4. SEM of Timarete hawaiensis showing: A, anterior end in dorsal view; B, anterior end in lateral view; C, mid-body neuropodium and notopodium; D, anterior end showing branchial and tentacular insertion, abbreviation: br— branchia; E, Pygidium; F, anterior serrated capillary chaetae.



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FIGURE 5. Variation of size-dependent characters in Timarete hawaiensis. A—Relationships among the commencement of notoacicular and neuroacicular spines and the total number of chaetigers in complete specimens. B— Relationships between the body length (in mm) and the number of chaetigers. C—The position in chaetigers where the branchial filaments shift to mid-dorsum.

Treadwell (1943) and Hartman (1956) reported five or six neuroacicular spines in the holotype; however, none of the specimens examined, including the holotype, have more than four aciculars in the neuropodium. Timarete hawaiensis differs from its congeners by the presence of branchial filaments on the third peristomial annulation. Biology. The specimens from the type-locality inhabit sponges and are found with Haplosyllis spongicola Grube (1855) and Ophiactis cf. modesta Brock 1888. Timarete hawaiensis reproduces both sexually and asexually by fission. Some specimens were found regenerating anterior and/or posterior ends. Several specimens had completely regenerated anterior ends that were pale in color that contrasted with the darker pigmentation of the rest of the body. In addition, mature females were found with eggs. Petersen (1999) also stated that other Timarete species reproduce by architomic fragmentation. Some specimens from Paiko Lagoon, south coast of Oahu Island, Hawaii, 21°16′58′′ 157°43′29′′, identified by Dr. Olga Hartman as Cirriformia hawaiensis were observed in aquaria by Thomas (1973). He pointed out that this species does not construct burrows but occupies a vertical subsurface position in the sediment with feeding tentacles and a few branchiae protruding above the surface. Timarete hawaiensis inhabits the sediment surface layers to about 10 cm in depth but is more abundant in the upper 5 cm. The long branchiae protrude into the overlying water for respiration; the tentacles picking up detritus at the sediment water interface that is presumably ingested. Distribution. The type-locality is Pearl Harbor, south shore of Oahu, Hawaii (Treadwell 1943). Additional specimens were found at Paiko Lagoon (Thomas 1973; Bailey-Brock 1987), Hanauma Bay, and Kaimana beach, south shore, and at Kahuku, north shore of Oahu, Hawaii. This distribution indicates that T. hawaiensis is endemic to Oahu, Hawaii.

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Discussion Both species redescribed here are endemic to the Hawaiian Islands and inhabit fragile habitats such as Diopatra and chaetopterid mounds and massive growth of sponges that are readily disturbed by boat traffic, fishermen, swimmers, and physical disturbances such as hurricanes and storms. Some species-level characters for cirratulids are strongly related to the growth (Blake 1996). The appearance of acicular spines on the notopodium and neuropodium in Cirriformia crassicollis are clearly sizedependent. However, a gap of approximately ten chaetigers is consistently found on all specimens regardless of size. On the other hand, for Timarete hawaiensis, the appearance of notoacicular spines are more related to growth than the appearance of neuroacicular spines. The gap between noto- and neuroacicular spines appearance is greater on larger specimens than on smaller ones. This gap between the appearance of noto- and neuroacicular spines may be a good species-level character but needs further investigation. The previous descriptions of both species were based on one or few specimens. Based on previous studies and on the results shown here, the description of polychaetes belonging to the family Cirratulidae should take into account the variability of some characters during ontogeny. The position of where the acicular spines begin on the noto- and neuropodia should be measured on specimens over a range of sizes to allow for accurate descriptions and correct identifications in the future. However, cirratulid species are rarely dominants in benthic communities (although very common), making the sampling of a full range of sizes very difficult. Juveniles would need to be identified and measured but they may show different chaetae patterns from adults, as seen in some species of Cirriformia (Blake, 1996) and juveniles and adults may not be present at a study site at the same time.

Acknowledgments The first author is indebted to Drs Rodolfo Elías and Maria Silvia Rivero (Universidad Nacional de Mar del Plata) for kindly introducing him to cirratulid taxonomy. We thank the University of Hawaii Water Resources Resource Center for SEM funds. Tina Carvalho (BEMF) assisted with SEM procedure. Specimens would not have been available for study without the research interests and collecting efforts of S. A. McCarthy, W. Estabrooks, R. Brock, V. Brock, C. Moody, the City and County of Honolulu, and collection permits. We are grateful for Dr Elín Sigvaldadóttir (The Swedish Museum of Natural History) and Dr Mark Siddall (The American Museum of Natural History) for graciously loaning the holotypes of C. crassicollis and T. hawaiensis, respectively. Linda Ward and Cheryl Bright helped cataloguing specimens in the USNM. Drs James Blake, Pat Hutchings and one anonymous reviewer made helpful comments that improved this manuscript.

Literature cited Abbott, D.P. (1946) Some polychaetous annelids from a Hawaiian fish pond. University of Hawaii Research Publication, 23, 4–24. Bailey-Brock, J.H. (1979) Sediment trapping by chaetopterid polychaetes on a Hawaiian fringing reef. Journal of Marine Research, 37(4), 643–656. Bailey-Brock, J.H. (1984) Ecology of the tube dwelling polychaete Diopatra leuckarti Kinberg, 1865 (Onuphidae) in Hawaii: community structure, and sediment stabilizing properties. Zoological Journal of the Linnean Society, 80, 191–199. Bailey-Brock, J.H. (1987) Phylum Annelida. In: Devaney, D.M., Eldredge, L.G. (Eds.), Reef and Shore Fauna of Hawaii. Bishop Museum Press, Hawaii, pp. 213–454. Blake, J.A. (1996) 8. Family Cirratulidae Ryckholdt, 1851. Including a revision of the genera and species from the eastern North Pacific. In: J. A. Blake, B. Hilbig and P.H. Scott (eds.), Taxonomic Atlas of the Benthic Fauna of the Santa Maria Basin and the Western Santa Barbara Channel. Volume 6 – The Annelida Part 3, Polychaeta: Orbiniidae to Cossuridae. Santa Barbara Museum of Natural History, pp. 263–384. Santa Barbara, California.



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Çinar, M.E. (2007) Re-description of Timarete punctata (Polychaeta: Cirratulidae) and its occurrence in the Mediterranean Sea. Scientia Marina, 71(4), 755–764. Dreyer, J., Bailey-Brock, J.H. & McCarthy, S.A. (2005) The immediate effects of Hurricane Iniki on intertidal fauna on the south shore of O’ahu. Marine Environmental Research, 59, 367–380. Hartman, O. (1948) The marine annelids erected by Kinberg with some notes on some other types in the Swedish State Museum. Arkiv för Zoologi, 42A(1), 1–137. Hartman, O. (1956) Polychaetous annelids erected by Treadwell, 1891 to 1948, together with a brief chronology. Bulletin of the American Museum of Natural History, 109, 239–310. Hartman, O. (1966) Polychaetous annelids of the Hawaiian Islands. Occasional Papers of the Bernice P. Bishop Museum, 23, 163–252. Kinberg, J.G.H. (1866) Annulata nova. Öfversigt af Förhandlingar Konglia Vetenskaps Akadamiens, 22, 239–258. Okuda, S. (1937) Polychaetous annelids from the Palau Islands and adjacent waters, the South Sea Islands. Bulletin of the Biogeographical Society of Japan, 7(12), 257–315. Petersen, M.E. (1999) Reproduction and development in Cirratulidae (Annelida: Polychaeta). Hydrobiologia, 402, 107– 128. Thomas, K. (1973) A contribution to the Zoology and distribution of annelids in Paiko Lagoon, Oahu. Master Thesis, University of Hawai‘i at Mānoa, Honolulu, 109pp. Treadwell, A.B. (1943) New species of polychaetous annelids from Hawaii. American Museum Novitates, 1233, 1–4.

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