Scolelepis Foliosa Has Many Variable Shape (Fig

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Phylum: Annelida Scolelep is foliosa Class: Polychaeta Order: Canalpalpita, Spionida A very large spionid polychaete Family: Spionidae

Taxonomy: The taxonomic history of this Anterior: Prostomium pointed and species is extensive and has yet to be fully tapered anteriorly (Scolelepis, Light resolved. The genus Scolelepis has many 1978; Pettibone 1963), but can also be synonyms including Aonis, Nerine, rounded (Berkeley and Berkeley 1952; Pseudomalacoceros, Nerinides, Scolecolepis Hobson and Banse 1981) (Fig. 2b). and Pseudonerine (Pettibone 1963). No frontal horns. Small occipital cirrus Furthermore, several subgenera have been present (Fig. 2a) and no caruncle (= proposed for Scolelepis. In 1963, Pettibone dorsal sense organ, Hobson and suggested the subgenera Scolelepis and Banse 1981) (Fig. 2b). Peristomium Nerinides based on neuropodial lamellae and enlarged and envelops prostomium ventral cirri. In 1987, Maciolek suggested the with two rolls (Light 1978) (Fig. 2b). subgenera Scolelepis and Parascolelepis Trunk: Fifth setiger not modified based on hooded hook morphology. These (Figs. 1, 2a) and no interparapodial subgenera were then erected to generic pouches (brackets, Fig. 1) (genus status by Williams (2007) based on the Scolelepis). phylogenetic analysis of Blake and Arnofsky Posterior: Anus and pygidium are (1999) (Blake 2006; Rocha and de Paiva dorsal (Fig. 3). Pygidium is a thick, 2012). The genus Scolelepis now includes 45 lobed pad with no cirri. species (Delgado-Blas 2006; Rocha and de Parapodia: Biramous, lamellar and with Paiva 2012). Scolelepis foliosa has many variable shape (Fig. 5). Podia become small synonyms including Aonis foliosa, Nerine toward worm posterior (Fig. 5d), but are not coniocephala, Aonis vittata, Nerine sarsiana glandular or thick. and Nerine foliosa occidentalis and may be Setae (chaetae): Fine spines (capillaries) in the same species as S. alaskensis and S. all noto- and neuropodia. Hooded hooks (Fig. bonnieri (Treadwell 1914; Pettibone 1963). 4a) begin in neuropodia after setiger 57 (sp. foliosa, Light 1978). Hooks also on notopodia Description on posterior-most setigers (Figs. 4, 5c, d). Size: The illustrated specimen (from Coos Hooks hooded, unidentate or multidentate Bay) is 500 mm in length and 15 mm in width, and worn (Fig. 4a). Capillary setae limbate with over 500 segments. The first 130 (no wings) and very finely striated, some with segments are 140 mm in length. Scolelepis horizontal ribs begin at setiger 39 (Fig. 4b). alaskensis, which may be the same species, Notosetae present in setiger one (Fig. 2a). has 130 segments and is 80 mm in length No setae with distal fringe. (Treadwell 1914). Eyes/Eyespots: Two pairs, anteriorly, in Color: The specimens we examined are some specimens (not the present specimen). golden tan with green palps that have white Anterior Appendages: Simple, massive and lines and spots, and red vessels. long (extending to setiger 20). Feeding palps General Morphology: A large and thick (Fig. 1) are easily broken off. worm that is rectangular in cross section and Branchiae: Branchiae are long, cirriform and bears many tightly arranged segments (Fig. partly fused to parapodial lamellae, begin on 1). setiger two (Figs. 2a, b) and continue almost Body: Not divided into distinct body regions to the worm posterior (genus Scolelepis, Light (Spionidae) (Blake 1996). 1978) (Fig. 1).

A publication of the University of Oregon Libraries and the Oregon Institute of Marine Biology Individual species: http://hdl.handle.net/1794/12682 and full 3rd edition: http://hdl.handle.net/1794/18839 Email corrections to [email protected]

Burrow/Tube: Individuals make loose cirri. They also have branchiae only on the vertical burrows in shifting sands, but have no anterior segments – all characteristics that permanent tube (Pettibone 1963). are lacking in Scolelepis. Pharynx: Bears a large, eversible proboscis The genus Scolelepis is characterized which is saclike. by its pointed prostomium (which is blunt in Genitalia: S. foliosa), a caruncle (if present) to setiger Nephridia: two, a possible occipital cirrus, cirriform branchiae beginning on setiger two, an Possible Misidentifications unmodified fifth setiger, no interparapodial Spionidae can be distinguished by a pair pouches or saber setae and a pad-like of long prehensile grooved palps which pygidium (Light 1978). Other species that arise from the posterior peristomium (Blake could be found in our area include: 1996). Two other polychaete families have Scolelepis squamata, a species referred long palps: the Magelonidae (adhesive to as Nerinides acuta when it was found in palps, not long and flowing, and flattened San Francisco Bay (Jones 1961) and as spade-like prostomiums) and the Nerine cirratulus in South Slough, Coos Bay Chaetopteridae (have palps, but their (Hartman and Reish 1950) as well as other bodies are very obviously divided into three places (Berkeley and Berkeley 1952; quite different regions which is not the case Hartman 1969). This species also has in spionids). The similar family Cirratulidae, many synonyms (Spio acuta, Nerine minuta, may also have a large pair of palps, but they and Nerine agilis) (Light 1978) and is have tentacular filaments, which are lacking primarily distinguishable from S. foliosa by in the spionids. Spionids also have hooded its hooded hooks, which appear first on hooks in posterior segments, may or may setigers 25–40 and not on setiger 57. Other not have prostomial appendages or differences include its capillary setae, which branchiae, the prostomium is well are limbate and without ribs. The developed and fused with peristomium, the prostomium of S. squamata is pointed fore pharynx is without jaws and the setae are and aft and it usually possesses two pairs of mostly simple (Blake 1975). Often certain eyes. Scolelepis squamata can be up to 80 segments are highly modified and have mm in length and have up to 200 segments special setae, for example prostomial horns (Light 1978). Unlike S. foliosa, it has no are present in some genera (not occipital cirrus, and it does have a caruncle Scolelepis). Spionid parapodia are to setiger two. Like S. foliosa, it has long biramous, with acicula (see Leitoscoloplos cirriform branchiae from setiger two and the pugettensis) and sometimes have stout post-setal lamellae of setiger one are well saber setae (not Scolelepis). There are 19 developed. local spionid genera (Blake and Ruff 2007). Scolelepis tridentata has been found in Several other, locally occurring genera California and in Ireland. It is not known if it of spionids have well developed branchiae is present in Oregon. It has tridentate and pointed prostomia. Spio also has hooded hooks, each with a large central branchiae from setiger one, but lacks fang, which begin on setigers 15–16, not on accessary branchiae. In Pygospio, setiger 57. Capillary setae are limbate branchiae first appear on setiger 10. (Males (unlike in S. foliosa), but are similarly ribbed. have an additional pair on setiger two (Light Scolelepis texana is reported from 1978). In both genera Malacoceros and central CA and the eastern United States Laonice the branchiae begin on setiger two. including the Gulf of Mexico. This species Malacoceros species are distinct in having is recognizable by the lack of notosetae on ventral saber setae on their median and the first setiger and neuropodial hooded posterior segments and a prostomium with hooks from setigers 13–27 (Blake and Ruff horns (Light 1978). Laonice lack the horns 2007). and saber setae, but have a very long caruncle, interparapodial pouches, and anal

Hiebert, T.C. 2015. Scolelepis foliosa. In: Oregon Estuarine Invertebrates: Rudys' Illustrated Guide to Common Species, 3rd ed. T.C. Hiebert, B.A. Butler and A.L. Shanks (eds.). University of Oregon Libraries and Oregon Institute of Marine Biology, Charleston, OR. Ecological Information Bibliography Range: Scolelepis foliosa exhibits a cosmopolitan distribution, which may be due 1. AUDOUIN, J. F., and H. MILNE- in part to its convoluted taxonomic history EDWARDS. 1833. Classification des and extensive synonymies. Pacific Coast annelides et description de celles qui range includes British Columbia (Berkeley habitent les cotes de la France. and Berkeley 1952) and Washington Annales Des Sciences Naturelles. (Imajima and Hartman 1964). Individuals 80:411-425. are reported from Oregon and California (as 2. BERKELEY, E., and C. BERKELEY. S. f. occidentafis) by Audouin and Milne- 1952. Annelida. Polychaeta Edwards 1833, although this species is not sedentaria. Canadian Pacific Fauna. currently included in the most recent 9b:1-139. intertidal guide (Blake and Ruff 2007). 3. BLAKE, J. 2006. Spionida, p. 565-638. Local Distribution: Distribution in Coos Bay In: Reproductive biology and includes Clam Island, Pigeon Point, phylogeny of annelida. Enfield, Charleston mudflats (e.g., Portside) and bay Science Publishers. Vol. 4. Science mouth, subtidally. Publishers, Enfield. Habitat: Loose sandy mud and gravel in 4. BLAKE, J. A. 1975. Phylum Annelida: Coos Bay. Offshore in clean well-sorted sand Class Polychaeta, p. 151-243. In: (Hancock et al. 1984). Light's manual: intertidal invertebrates Salinity: Specimens collected at 30. of the central California coast. S. F. Temperature: Ranges from 9–15 °C. Light, R. I. Smith, and J. T. Carlton Tidal Level: Found at approximately 0.0 m at (eds.). University of California Press, Pigeon Point in Coos Bay. Offshore in about Berkeley. 5.4 m (Hancock et al. 1984). 5. —. 1996. Family Spionidae, p. 81-224. Associates: In: Taxonomic atlas of the benthic Abundance: Not common in Coos Bay. fauna of the Santa Maria Basin and Western Santa Barbara Channel. Vol. Life-History Information 6. J. A. Blake, B. Hilbig, and P. H. Reproduction: Pelagic development where Scott (eds.). Santa Barbara Museum larvae are mostly planktotrophic (Hannerz of Natural History, Santa Barbara, CA. 1956). 6. BLAKE, J. A., and P. L. ARNOFSKY. Larva: Larvae of Scolelepis foliosa are 1999. Reproduction and larval planktotrophic and can be recognized using development of the spioniform many adult characters including the presence polychaeta with application to of branchiae, setiger one and setiger five not systematics and phylogeny. modified and without specialized setae (as in Hydrobiologia. 402:57-106. adults), a pointed and triangular prostomium 7. BLAKE, J. A., and E. R. RUFF. 2007. that may or may not bear lateral horns and a Polychaeta, p. 309-410. In: Light and blackish brown gut (Crumrine 2001). Smith manual: intertidal invertebrates Juvenile: from central California to Oregon. J. Longevity: Carlton (ed.). University of California Growth Rate: Press, Berkeley, CA. Food: Selective surface deposit feeder 8. CRUMRINE, L. 2001. Polychaeta, p. (Fauchald and Jumars 1979). 39-77. In: Identification guide to larval Predators: marine invertebrates of the Pacific Behavior: Northwest. A. Shanks (ed.). Oregon State University Press, Corvallis, OR. 9. DELGADO-BIAS, V. H. 2006. Partial revision of Scolelepis (Polychaeta : Spionidae) from the Grand Caribbean

Region, with the description of two 19. MACIOLEK, N. 1987. New species new species and a key to species and records of Scolelepis (Polychaeta: recorded in the area. Contributions to Spionidae) from the east coast of Zoology. 75:75-97. North America, with a review of the 10. FAUCHALD, K., and P. A. JUMARS. subgenera. Proceedings of the 1979. Diet of worms: a study of Biological Society of Washington. polychaete feeding guilds. 7:16-40. Oceanography and Marine Biology. 20. PETTIBONE, M. H. 1963. Revision of 17:193-284. some genera of polychaete worms of 11. HANCOCK, D. R., P. O. NELSON, C. the family Spionidae, including the K. SOLLITT, and K. J. WILLIAMSON. description of a new species of 1984. Coos Bay offshore disposal site Scolelepis. Proceedings of the investigation: interim report phase I, Biological Society of Washington. February, 1979-March, 1980, 76:89-103. Portland, OR. 21. ROCHA, M. B., and P. C. DE PAIVA. 12. HANNERZ, L. 1956. Larval 2012. Scolelepis (Polychaeta: development of the polychaete Spionidae) from the Brazilian coast families Spionidae Sars, Disomidae with a diagnosis of the genus. Mesnil, and Poecilochaetidae N. fam: Zoologia. 29:385-393. in the Gullmar Fjord (Sweden), p. 204. 22. TREADWELL, A. L. 1914. In: Zoologiska bidrag fran Uppsala. Polychaetous annelids of the Pacific Uppsala University, Uppsala, Sweden. Coast in the collections of the 13. HARTMAN, O. 1969. Atlas of the Zoological museum of the University sedentariate polychaetous Annelids of California. University of California from California. Allan Hancock Publications in Zoology. 13. Foundation, University of Southern 23. WILLIAMS, J. D. 2007. New records California, Los Angeles, CA. and description of four new species of 14. HARTMAN, O., and D. J. REISH. Spionids (Annelida: Polychaeta: 1950. The Marine annelids of Oregon. Spionidae) from the Philippines: the Oregon State College, Corvallis, genera Dispio, Malacoceros, Polydora, Oregon. and Scolelepis, with notes on palp 15. HOBSON, K. D., and K. BANSE. ciliation patterns of the genus 1981. Sedentariate and archiannelid Scolelepis. Zootaxa. 1-35. polychaetes of British Columbia and Washington. Canadian Bulletin of Fisheries and Aquatic Sciences. 209:1-144. 16. IMAJIMA, M., and O. HARTMAN. 1964. The polychaetous annelids of Japan. Part II. Allan Hancock Foundation Publications Occasional Papers. 26:239-452. 17. JONES, M. L. 1961. A quantitative evaluation of the benthic fauna off Point Richmond, California. University of California. Publications in Zoology. 67:219-320. 18. LIGHT, W. J. 1978. Spionidae: Polychaeta, Annelida. Boxwood Press, Pacific Grove, CA.