Nephtys Caeca Class: Polychaeta Order: Phyllodocida a Sand Worm Family: Nephtyidae

Phylum: Annelida Nephtys caeca Class: Polychaeta Order: Phyllodocida A sand worm Family: Nephtyidae

Taxonomy: Nephtys caeca is the name used Setae (chaetae): All nephtyid setae are in current local intertidal guides (e.g., Blake simple and the setae of both rami are of and Ruff 2007). One can find several similar morphology. Overall, there are four synonyms, however, including variants of the main types of nephtyid setae including generic spelling (Nephthys), subjective capillary (e.g., spinose), barred (which are synonyms (e.g., N. margaritacea, N. oerstedii pre-acicular), lyrate and setae with spines and N. bononensis) and species described (Dnestrovskaya and Jirkov 2011). Nephtys and later determined to be an earlier caeca has fan-like bunches of neuro- and developmental stage of N. caeca (e.g., notosetae on the parapodial lobes. Post- Nephthys nudipes) (Rainer 1991). acicular setae (Fig. 5) are long and fine, with single lateral barbs (Fig. 4a) and pre-acicular Description setae short and with transverse bars (Figs. Size: Individuals to 20 cm in length and 10– 4b, 5). 15 mm in width (Hartman 1968). 90–150 total Eyes/Eyespots: None (Fig. 2). body segments. Anterior Appendages: Four small, simple Color: Body color is pale pink and can be (unforked) antennae (Fig. 2). light to dark green or brown. No prominent Branchiae: The interramal cirri, which are external pigment patterns. The proboscis is inserted just beneath each dorsal cirrus, are iridescent. sometimes called branchiae (Blake and Ruff General Morphology: Anterior cylindrical in 2007). cross-section and becomes slender and Burrow/Tube: Nephtys caeca can move rectangular posteriorly (Nephtyidae, Blake rapidly through loose sand and makes and Ruff 2007). temporary burrows (MacGinitie 1935). Body: Individuals long, slender and Pharynx: Bears short and wide proboscis quadrangular in cross-section (Hartman with a variety of papillae, their number and 1968). arrangement is of taxonomic significance Anterior: Prostomium pentagonal, (Blake and Ruff 2007). The proboscis in flattened and no pigment pattern (Fig. Nephtys species can be divided into three 2). distinct regions including the proximal, sub Trunk: Thick with widely separated distal and distal (Lovell 1997) (Fig. 1). The parapodial rami (Fig. 1, 5). proboscis, when fully everted, is globular, with Posterior: Pygidium with very small 22 rows of paired distal papillae forming a and hairlike posterior cirrus (Fig. 1). crown-like structure. Twenty-two rows of sub Parapodia: Fleshy flaps extending laterally distal papillae with five small papillae in each from each segment, are biramous and rami row (Fig. 1). The proximal surface of the are widely separated, densely packed and proboscis is rough and covered with minute their setae are fan-shaped (Nephtyidae, Blake wart-like papillae (Fig. 1). and Ruff 2007). Each lobe with a broad and Genitalia: rounded notopodium and a neuropodium. Nephridia: Post-acicular lobes become foliaceous posteriorly (Hartman 1968) (Fig. 5). Bears Possible Misidentifications interramal cirri that are long and recurved Worms of the family Nephtyidae can be between the two parapodial lobes (Figs. 3, 5). distinguished by their anteriorly cylindrical

A publication of the University of Oregon Libraries and the Oregon Institute of Marine Biology Individual species: http://hdl.handle.net/1794/12672 and full 3rd edition: http://hdl.handle.net/1794/18839 Email corrections to: [email protected] and posteriorly rectangular bodies (in cross (Blake and Ruff 2007). N. cornuta can also section), well-developed bi-lobed parapodia, be differentiated because it has 18 distal interramal cirri, four small prostomial paired papillae (instead of 22 in N. caeca) antennae, and eversible globular proboscis (Lovell 1997). N. punctata is much like N. with terminal rows of papillae. They are caeca in size and form (Hartman 1938), but strong and muscular worms that can be good with interramal cirri beginning on setiger 8– burrowers and strong swimmers (Blake and 10, and with incised acicular lobes in the Ruff 2007). Nephtyids superficially resemble anterior parapodia. This species is large the genus Nereis, however, they have no and muscular with wide body and short long anterior appendages (tentacular cirri) parapodia and is currently only reported in and their proboscis armature is quite southern California (Hilbig 1997; Blake and different (Kozloff 1993). The distinctive Ruff 2007) taxonomic characters of N. caeca include 22 N. parva, colorless except for a dark distal paired papillae, 22 rows of sub distal spot in the middle of its prostomium papillae with five papillae per row, no (Hartman 1968), a smooth proboscis unpaired mid dorsal papilla and interramal proximally, no medial papilla, eyespots on its cirri beginning on setigers five or six (Blake third setiger and interramal cirri beginning on and Ruff 2007). the fourth setiger. The type material from Some Nephtys species are this species is suspected to have been distinguished from each other by very fine miscurated and the holotype appears to be morphological details. The species most that of N. cornuta, while the species closely related to and difficult to differentiate description and paratypes match N. from N. caeca include N. caecoides and N. caecoides more closely. Thus, this species californiensis. N. caecoides, is slightly is not a currently valid taxon (Lovell 1997; smaller (on average) than N. caeca, with Blake and Ruff 2007). dark bands of color on its anterior end, and N. ferruginea has the same number a smooth proboscis, not a rough one. N. of paired distal and sub distal papillae, caecoides also has an unpaired medial however, the interramal cirri in this species papilla (not present in N. caeca) and begin on setiger three, rather than four in N. interramal cirri beginning on the fourth caecoides (Lovell 1997). N. ferruginea setigers (rather than the fifth or sixth in N. individuals have a distinct rust colored caeca). It is probably the closest species, pigment in a V-shape pattern on prostomium morphologically, to N. caeca, and their two in addition to transverse bars mid dorsally on distributions overlap in Coos Bay (Porch the first 20 setigers and oblique stripes 1970). N. californiensis is found mostly on dorsolaterally (Hilbig 1997). the outer coast, or if in bays, only in very clean coarse sand. It has a distinctive V- Ecological Information shaped pigment pattern (sometimes with Range: Type localities include Greenland red spot at center) of pigmentation on the and the Arctic (Hartman 1968). Known lower end of the prostomium, a smooth distribution is Alaska to northern California proboscis without medial papilla, soft silky and circumboreal. Possibly introduced from flowing setae and interramal cirri beginning the eastern United States (Blake and Ruff on the third setiger. 2007). Three other Nephtys species are not Local Distribution: Coos Bay distribution so easily confused with N. caeca. N. includes many stations, especially those cornuta, a small species (less than 15 mm within South Slough. The distribution of N. in length) that can be identified by its caeca is much like that of the polychaete distinctive bifid ventral and posterior Scoleteoma zonata. antennae. This species often retains larval Habitat: Sand, mud or mixed sediments. eyes on the third setiger, a feature which is Individuals also occur with eelgrass and usually lost in other closely related species

Hiebert, T.C. 2015. Nephtys caeca. 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. prefer more mud than Scoleteoma zonata common characteristic of other nephtyid (Porch 1970). larvae (e.g., http://invert- Salinity: Collected at salinities of 30, but can embryo.blogspot.com/2012/12/confirmed- tolerate lower salinities (i.e., freshwater of identity-of-wild-caught.html). Eight-setiger stream beds) (Porch 1970). stages measure approximately 670 µm in Temperature: A cold water species, N. length and have simple capillary setae (Lacalli caeca does not extend far southward to 1980). Nephtyid trochophore and California. metatrochophore larvae are predatory Tidal Level: Intertidal (+ 0.15 m) to lower (Fernald et al. 1987; Crumrine 2001). intertidal and depths of 1000 m (Rainer 1991). Juvenile: The prostomium transitions from Associates: Known associates include rounded to angular in newly metamorphosed barnacles and the large polychaete, Pista individuals (Fig. 5, Lacalli 1980). Juveniles pacifica. may possess eyes on one of the first three Abundance: Not common, locally (Blake and setigers that are usually, although not always Ruff 2007). In the St. Lawrence Estuary (e.g. N. cornuta), lost in adults (Nephtys, (Québec, Canada), most individuals were Hilbig 1997). Advanced larval or juvenile observed in the lowest sampling sites stages were collected benthically from the (specific sampling heights not indicated, River Tyne Estuary and described by Olive in Caron et al. 1995). 1977. Unique features included yellow-brown pigmented prostomium with rusty brown Life-History Information pigment granules dorsally, green intestine Reproduction: Nephtys caeca, as is the becoming deep blue posteriorly, spade- case for other Nephtys species, are free- shaped pygidium and both smooth and spawning with pelagic larval development that striated setae. The four pairs of anterior proceeds via a trochophore larva (Fernald et antennae only developed in advanced stages al. 1987; Crumrine 2001; Pleijel and Rouse (Olive 1977). No larval settlement was 2006). In the St. Lawrence Estuary (Québec, observed intertidally in the St. Lawrence Canada), oocyte maturation occurred in the Estuary, suggesting sublittoral larval late summer to autumn, oocyte diameters recruitment. Instead, juveniles were seen were approximately 140–160 µm (Caron et al. intertidally in June–July (1986) (Caron et al. 1995) and ripe adults spawn into their 1995). temporary burrows (Bently et al. 1984). Longevity: Seven or more year life-span However, in the River Tyne Estuary (United (United Kingdom, Olive 1977). Kingdom), individuals are known to spawn in Growth Rate: Sexual maturity is reached in late spring or early summer (Olive 1977). two years (United Kingdom, Olive 1977). Larva: Nephtyid trochophore larvae have a Food: Nephtys caeca adults are carnivorous pair of eyes, dome-shaped prostomium and and predation by them is known to regulate barrel-shaped body. They have well other infaunal populations within a developed prototrochs and telotrochs, with community. They are a primary predator of neurotrochs present in young larvae. They Macoma balthica (St. Lawrence Estuary) and are common in plankton samples and are are known to feed on smaller conspecifics recognized by their shape and species- (Ambrose 1984; Caron et al. 2004). Juvenile specific bright body colors (Lacalli 1980; N. caeca, however, are herbivores (Caron et Fernald et al. 1987). Early trochophore larvae al. 2004). of Nephtys caeca have been described Predators: (Thorson 1946; Lacalli 1980). They have dull Behavior: Individuals are active, and are red to brown pigmentation on the episphere, good swimmers and burrowers (MacGinitie prostomium and pygidium. The posterior 1935). Bioturbation from N. caeca has been pigmentation is arranged in two bands, one shown to homogenize particles in the first few anterior to and the other posterior to the centimeters of sediment while burrowing or pygidium (Lacalli 1980). They also have an moving (Piot et al. 2008). olive colored gut and no blue pigmentation, a

A publication of the University of Oregon Libraries and the Oregon Institute of Marine Biology Individual species: http://hdl.handle.net/1794/12672 and full 3rd edition: http://hdl.handle.net/1794/18839 Email corrections to: [email protected] Bibliography Northern Europe and Arctic. Italian Journal of Zoology. 78:219-228. 1. AMBROSE, W. G. 1984. Increased 8. FERNALD, R. L., C. O. HERMANS, T. emigration of the amphipod C. LACALLI, W. H. WILSON, JR, and Rhepoxynius abronius (Barnard) and S. A. WOODIN. 1987. Phylum the polychaete Nephtys caeca Annelida, Class Polychaeta, p. 138- (Fabricius) in the presence of 195. In: Reproduction and invertebrate predators. Journal of development of marine invertebrates Experimental Marine Biology and of the northern Pacific coast. M. F. Ecology. 80:67-75. Strathmann (ed.). University of 2. BENTLEY, M. G., P. J. W. OLIVE, P. Washington Press, Seattle, WA. R. GARWOOD, and N. H. WRIGHT. 9. HARTMAN, O. 1938. Review of the 1984. The spawning and spawning annelid worms of the family mechanism of Nephtys caeca nephtyidae from the northeast pacific, (Fabricius, 1780) and Nephtys with descriptions of five new species. hombergi Savigny, 1818 (Annelida, Proceedings of the United States Polychaeta). Sarsia. 69:63-68. National Museum. 85:143-158. 3. BLAKE, J. A., and R. E. RUFF. 2007. 10. HARTMAN, O. 1968. Atlas of the Polychaeta, p. 309-410. In: The Light errantiate polychaetous annelids from and Smith manual: intertidal California. Allan Hancock Foundation, invertebrates from central California to University of Southern California, Los Oregon. J. T. Carlton (ed.). University Angeles. of California Press, Berkeley, CA. 11. HILBIG, B. 1997. Family Nereididae, 4. CARON, A., L. BOUCHER, G. p. 291-316. In: Taxonomic atlas of the DESROSIERS, and C. RETIERE. benthic fauna of the Santa Maria 1995. Population-dynamics of the Basin and Western Santa Barbara polychaete Nephtys caeca in an Channel. Vol. 4. J. A. Blake, B. Hilbig, intertidal estuarine environment and P. H. Scott (eds.). Santa Barbara (Quebec, Canada). Journal of the Museum of Natural History, Santa Marine Biological Association of the Barbara, CA. United Kingdom. 75:871-884. 12. KOZLOFF, E. N. 1993. Seashore life 5. CARON, A., G. DESROSIERS, P. J. of the northern Pacific coast: an W. OLIVE, C. RETIERE, and C. illustrated guide to Northern California, NOZAIS. 2004. Comparison of diet Oregon, Washington, and British and feeding activity of two Columbia. University of Washington polychaetes, Nephtys caeca Press, Seattle, WA. (Fabricius) and Nereis virens (Sars), in 13. LACALLI, T. C. 1980. A guide to the an estuarine intertidal environment in marine flora and fauna of the Bay of Quebec, Canada. Journal of Fundy: polychaete larvae from Experimental Marine Biology and Passamaquoddy Bay. Canadian Ecology. 304:225-242. Technical Report of Fisheries and 6. CRUMRINE, L. 2001. Polychaeta, p. Aquatic Sciences. 940:1-27. 39-77. In: Identification guide to larval 14. LOVELL, L. L. 1997. A review of six marine invertebrates of the Pacific species of Nephtys (Cuvier, 1817) Northwest. A. Shanks (ed.). Oregon (Nephtyidae: Polychaeta) described State University Press, Corvallis, OR. from the eastern Pacific. Bulletin of 7. DNESTROVSKAYA, N. Y., and I. A. Marine Science. 60:350-363. JIRKOV. 2011. Microscopical studies 15. MACGINITIE, G. E. 1935. Ecological of Nephtyid chaetae (Annelida: aspects of a California marine estuary. Polychaeta: Nephtyidae) from American Midland Naturlist. 16:629- 765.