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Phylum:

Cerebratulus marginatus Class: ,

A ribbon worm Order:

Family:

Taxonomy: Cerebratulus marginatus was Mouth: Ventral and behind the brain and dis- described by Renier (1804) from Naples, Ita- tinct from proboscis pore (order Heteronemer- ly. This now has a worldwide distri- tea) (Kozloff 1974). bution and an extensive list of synonyms Proboscis: Eversible (phylum Nemertea) (see Gibson 1995). Thus it is very likely that and, when not everted, coiled inside rhyncho- there are several different species currently coel (cavity). Proboscis of moderately size referred to as C. marginatus from differing with sticky glandular surface is everted less regions. readily than in C. californiensis. Proboscis bears no stylet (class Anopla) (Kozloff 1974). Description Tube/Burrow: An excellent swimmer and Size: Among the largest of local nemerte- strong burrower, C. marginatus does not in- ans, where sizes range from 50 cm to 1 m in habit a permanent tube or burrow. length and up to 15 mm in width (Coe 1901, 1905). Possible Misidentifications Color: Slate brown, dark grey or grayish Eight Cerebratulus species are report- green (Coe 1901). Individuals vary in color; ed from central CA to OR (Roe et al. 2007). locally they can be very dark brown. Some- Species in this have firm, non- times more pale ventrally, thin lateral mar- contractile and often ribbon-like bodies. One gins often colorless or white (Coe 1901, species that is easily mistaken for C. margina- 1905). tus is C. californiensis. Both are slate colored General Morphology: Large, thick and and possess thin lateral margins that are col- round worm anteriorly but very flat and rib- orless or white. Cerebratulus californiensis bon like in mid-body. Non-segmented can be identified by a head that is smaller (phylum Nemertea). than the body width (compare Figs. 1a and Body: Rounded anteriorly and very dorso- 1b) and by thin wide margins that span anteri- ventrally flattened posteriorly with thin lateral orly farther than in C. marginatus (T. Hiebert margins in intestinal region, ribbon-like (Coe and Maslakova, pers obs). 1943) (Fig 2). Not very contractile, individu- Other NE Pacific Cerebratulus species als fragment easily when handled (Roe et al. include C. albifrons, a dark brown species 2007). with white head, up to 30 cm in length and Anterior: Head spade-shaped with found intertidal and subtidal in Alaska to San pointed anterior tip (Fig. 1a), widening to just Diego, CA (Coe 1901; Roe et al. 2007); C. wider than or equal to body width. Cephalic montgomeryi with red body and head with grooves large and deep (Coe 1905), often white tip, occurs intertidal and subtidally from flaring when swimming. Alaska to Monterey Bay (Coe 1901); C. occi- Posterior: Caudal cirrus (tail) pre- dentalis is a subtidal species, up to 30 cm in sent, thin (Coe 1943) and easily lost when length, reddish brown dorsally with lighter collecting. ventral pigment, from Alaska to Puget Sound Eyes/Eyespots: No ocelli. and San Francisco Bay (Coe 1901); C. longi-

A publication of the University of Oregon Libraries and the Oregon Institute of Marine Biology Individual species: https://oimb.uoregon.edu/oregon-estuarine-invertebrates and full 3rd edition: http://hdl.handle.net/1794/18839 Email corrections to: [email protected]

Hiebert, T.C. 2015. Cerebratulus marginatus. 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, Charles- ton, OR.

ceps is found in the low intertidal, dredged, mon, rendering this species useful for experi- is up to 30 cm in length, dark reddish brown mental research (e.g. Bianchi 1969; Bianchi et with pale anterior and occurs from Alaska al. 1972; Voogt 1973). and Tamales Bay (Coe 1901; Corrêa 1964); Life-History Information C. herculeus is enormous in size (up to 2 m Reproduction: Males and females sexually long and 25 mm wide), burrows in soft sedi- mature in summer, gametes can be seen ment inter tidally and subtidally from Alaska through body wall in serially arranged trans- to southern California (Coe 1901); C. lineo- verse lines. latus is pale grey with olive longitudinal lines Larva: A classic species for embryological extending the entire body length (up to 20 work, the development of this species was cm) and is found intertidally up to 70 m from documented in 1899 (Coe) and fully described southern California to Mexico, and also in in 1930 (Schmidt) and proceeds indirectly via Miami, Florida (Coe 1905; Corrêa 1964). a planktotrophic pilidium larva (Coe 1899; Because of the many identifying char- Schmidt 1930; Coe 1905, 1940). acteristics, which are internal and not visible, Juvenile: it is sometimes very difficult to distinguish Longevity: among Nemerteans without dissecting them. Growth Rate: Ways in which the worms flatten, contract, Food: Predatory and feeds on polychaetes and coil are useful as aids to identification of and clams. live specimens. Predators: Ecological Information Behavior: Excellent swimmers and burrowers Range: Described from specimens in Na- (Coe 1901) (Fig 3), individuals sometimes col- ples, Italy (Renier 1804). The distribution of lected by net swimming at night (Coe 1943; C. marginatus is not known with certainty as Gibson 1995). the synonymy of this species is complicated. Bibliography Widespread in the northern hemisphere, Pa- cific coast of North America, Japan, western 1. BIANCHI, S. 1969. Histochemistry of neu- North Atlantic, Arctic, northern Europe in- rosecretory system in Cerebratulus cluding the Mediterranean (the type region). marcinatus (Heteronemertini). General Southern distribution reaches Madeira and Comparative Endocrinology. 13:206- (Gibson 1995; Roe et al. 2007). 213. Local Distribution: Coos Bay and South 2. BIANCHI, S., V. ESPOSITO, and N. Slough, at many sites. GRANATA. 1972. Cerebral organs of Cer- Habitat: Sand, mud, or fine gravel sedi- ebratulus marginatus (Heteronemertini). ments (Gibson 1995). General and Comparative Endocrinology. Salinity: 18:5-20. Temperature: The distribution of this spe- 3. COE, W. R. 1899. On the development of cies suggests a wide temperature tolerance. the Pilidium of certain Nemerteans. Trans- Tidal Level: Intertidal to sublittoral, dredged actions of the Connecticut Academy. x:235 at 50 m (Coe 1905, 1940) to 150 m (Gibson -262. 1995). 4. —. 1901. Papers from the Harriman Alas- Associates: ka Expedition. The Nemerteans. Proceed- Abundance: Frequently encountered in es- ings of the Washington Academy:1-110. tuarine mudflats in Charleston, OR. Com- 5. —. 1905. Nemerteans of the west and

A publication of the University of Oregon Libraries and the Oregon Institute of Marine Biology Individual species: https://oimb.uoregon.edu/oregon-estuarine-invertebrates and full 3rd edition: http://hdl.handle.net/1794/18839 Email corrections to: [email protected]

northwest coasts of North America. Bul- ebratulus marginatus. Archives Interna- letin of the Museum at Harvard College. tionales De Physiologie De Biochimie Et xlvii:1-318. De Biophysique. 81:871-880. 6. —. 1940. Revision of the nemertean fau- Updated 2014 na of the Pacific coasts of north, central T.C. Hiebert and northern South America. Allan Han- cock Foundation of Scientific Research. Allan Hancock Pacific Expeditions. Re- ports. 2:247-323. 7. —. 1943. Biology of the nemerteans of the Atlantic coast of North America. Transactions of the Connecticut Acade- my of Arts and Sciences. 35:129-328. 8. CORRÊA, D. D. 1964. Nemerteans from California and Oregon. Proceedings of the California Academy of Sciences (series 4). 31:515-558. 9. GIBSON, R. 1995. Nemertean genera and species of the world: an annotated checklist of original names and descrip- tion citation, synonyms, current taxonom- ic status, habitats and recorded zoogeo- graphic distribution. Journal of Natural History. 29:271-562. 10. KOZLOFF, E. N. 1974. Keys to the ma- rine invertebrates of Puget Sound, the San Juan Archipelago, and adjacent re- gions. University of Washington Press, Seattle. 11. ROE, P., J. L. NORENBURG, and S. MASLAKOVA. 2007. Nemertea, p. 221- 233. In: Light and Smith manual: intertid- al invertebrates from central California to Oregon. J. T. Carlton (ed.). University of California Press, Berkeley, CA. 12. SCHMIDT, G. A. 1937. Bau und Entwick- lung der pilidien von Cerebratulus pan- therinus und marginatus und die frage der morphologischen merkmale der hauptformen der pilidien. Zoologische Jahrbuecher Jena Anatomie. 62:423- 448. 13. VOOGT, P. A. 1973. Biosynthesis and composition of sterols in nemertean Cer-

Hiebert, T.C. 2015. Cerebratulus marginatus. 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, Charles- ton, OR.