Nephtys Caeca Class: Polychaeta, Errantia

Home , Errantia, Nephtyidae, Nephtys, Phyllodocida

Phylum: Annelida

Nephtys caeca Class: Polychaeta, Errantia

Order: Phyllodocida, Phyllodocida incertae sedis A sand worm Family: Nephtyidae

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

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. 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.

wart-like papillae (Fig. 1). center) of pigmentation on the lower end of Genitalia: the prostomium, a smooth proboscis without Nephridia: medial papilla, soft silky flowing setae and interramal cirri beginning on the third setiger. Possible Misidentifications Three other Nephtys species are not Worms of the family Nephtyidae can so easily confused with N. caeca. N. cornu- be distinguished by their anteriorly cylindri- ta, a small species (less than 15 mm in cal and posteriorly rectangular bodies (in length) that can be identified by its distinctive cross section), well-developed bi-lobed par- bifid ventral and posterior antennae. This apodia, interramal cirri, four small prostomi- species often retains larval eyes on the third al antennae, and eversible globular probos- setiger, a feature which is usually lost in oth- cis with terminal rows of papillae. They are er closely related species (Blake and Ruff strong and muscular worms that can be 2007). N. cornuta can also be differentiated good burrowers and strong swimmers because it has 18 distal paired papillae (Blake and Ruff 2007). Nephtyids superfi- (instead of 22 in N. caeca) (Lovell 1997). N. cially resemble the genus Nereis, however, punctata is much like N. caeca in size and they have no long anterior appendages form (Hartman 1938), but with interramal cirri (tentacular cirri) and their proboscis arma- beginning on setiger 8–10, and with incised ture is quite different (Kozloff 1993). The acicular lobes in the anterior parapodia. This distinctive taxonomic characters of N. cae- species is large and muscular with wide body ca include 22 distal paired papillae, 22 rows and short parapodia and is currently only of sub distal papillae with five papillae per reported in southern California (Hilbig 1997; row, no unpaired mid dorsal papilla and in- Blake and Ruff 2007) terramal cirri beginning on setigers five or N. parva, colorless except for a dark six (Blake and Ruff 2007). spot in the middle of its prostomium Some Nephtys species are distin- (Hartman 1968), a smooth proboscis proxi- guished from each other by very fine mor- mally, no medial papilla, eyespots on its third phological details. The species most close- setiger and interramal cirri beginning on the ly related to and difficult to differentiate from fourth setiger. The type material from this N. caeca include N. caecoides and N. cali- species is suspected to have been miscurat- forniensis. N. caecoides, is slightly smaller ed and the holotype appears to be that of N. (on average) than N. caeca, with dark cornuta, while the species description and bands of color on its anterior end, and a paratypes match N. caecoides more closely. smooth proboscis, not a rough one. N. cae- Thus, this species is not a currently valid tax- coides also has an unpaired medial papilla on (Lovell 1997; Blake and Ruff 2007). (not present in N. caeca) and interramal cir- N. ferruginea has the same number of ri beginning on the fourth setigers (rather paired distal and sub distal papillae, howev- than the fifth or sixth in N. caeca). It is er, the interramal cirri in this species begin probably the closest species, morphologi- on setiger three, rather than four in N. cae- cally, to N. caeca, and their two distribu- coides (Lovell 1997). N. ferruginea tions overlap in Coos Bay (Porch 1970). N. individuals have a distinct rust colored californiensis is found mostly on the outer pigment in a V-shape pattern on prostomium coast, or if in bays, only in very clean in addition to transverse bars mid dorsally on coarse sand. It has a distinctive V-shaped the first 20 setigers and oblique stripes pigment pattern (sometimes with red spot at dorsolaterally (Hilbig 1997).

Ecological Information µm (Caron et al. 1995) and ripe adults spawn Range: Type localities include Greenland into their temporary burrows (Bently et al. and the Arctic (Hartman 1968). Known dis- 1984). However, in the River Tyne Estuary tribution is Alaska to northern California and (United Kingdom), individuals are known to circumboreal. Possibly introduced from the spawn in late spring or early summer (Olive eastern United States (Blake and Ruff 1977). 2007). Larva: Nephtyid trochophore larvae have a Local Distribution: Coos Bay distribution pair of eyes, dome-shaped prostomium and includes many stations, especially those barrel-shaped body. They have well devel- within South Slough. The distribution of N. oped prototrochs and telotrochs, with neu- caeca is much like that of the polychaete rotrochs present in young larvae. They are Scoleteoma zonata. common in plankton samples and are recog- Habitat: Sand, mud or mixed sediments. nized by their shape and species-specific Individuals also occur with eelgrass and pre- bright body colors (Lacalli 1980; Fernald et al. fer more mud than Scoleteoma zonata 1987). Early trochophore larvae of Nephtys (Porch 1970). caeca have been described (Thorson 1946; Salinity: Collected at salinities of 30, but Lacalli 1980). They have dull red to brown can tolerate lower salinities (i.e., freshwater pigmentation on the episphere, prostomium of stream beds) (Porch 1970). and pygidium. The posterior pigmentation is Temperature: A cold water species, N. arranged in two bands, one anterior to and caeca does not extend far southward to the other posterior to the pygidium (Lacalli California. 1980). They also have an olive colored gut Tidal Level: Intertidal (+ 0.15 m) to lower and no blue pigmentation, a common charac- intertidal and depths of 1000 m (Rainer teristic of other nephtyid larvae (e.g., http:// 1991). invert-embryo.blogspot.com/2012/12/ Associates: Known associates include bar- confirmed-identity-of-wild-caught.html). Eight- nacles and the large polychaete, Pista paci- setiger stages measure approximately 670 fica. µm in length and have simple capillary setae Abundance: Not common, locally (Blake (Lacalli 1980). Nephtyid trochophore and and Ruff 2007). In the St. Lawrence Estuary metatrochophore larvae are predatory (Québec, Canada), most individuals were (Fernald et al. 1987; Crumrine 2001). observed in the lowest sampling sites Juvenile: The prostomium transitions from (specific sampling heights not indicated, Ca- rounded to angular in newly metamorphosed ron et al. 1995). individuals (Fig. 5, Lacalli 1980). Juveniles may possess eyes on one of the first three Life-History Information setigers that are usually, although not always Reproduction: Nephtys caeca, as is the (e.g. N. cornuta), lost in adults (Nephtys, Hil- case for other Nephtys species, are free- big 1997). Advanced larval or juvenile stages spawning with pelagic larval development were collected benthically from the River Tyne that proceeds via a trochophore larva Estuary and described by Olive in 1977. (Fernald et al. 1987; Crumrine 2001; Pleijel Unique features included yellow-brown pig- and Rouse 2006). In the St. Lawrence Estu- mented prostomium with rusty brown pigment ary (Québec, Canada), oocyte maturation granules dorsally, green intestine becoming occurred in the late summer to autumn, oo- deep blue posteriorly, spade-shaped pygidium cyte diameters were approximately 140–160 and both smooth and striated setae. The four

pairs of anterior antennae only developed in brates from central California to Oregon. J. advanced stages (Olive 1977). No larval T. Carlton (ed.). University of California settlement was observed intertidally in the Press, Berkeley, CA. St. Lawrence Estuary, suggesting sublittoral 4. CARON, A., L. BOUCHER, G. larval recruitment. Instead, juveniles were DESROSIERS, and C. RETIERE. 1995. seen intertidally in June–July (1986) (Caron Population-dynamics of the polychaete et al. 1995). Nephtys caeca in an intertidal estuarine Longevity: Seven or more year life-span environment (Quebec, Canada). Journal of (United Kingdom, Olive 1977). the Marine Biological Association of the Growth Rate: Sexual maturity is reached in United Kingdom. 75:871-884. two years (United Kingdom, Olive 1977). 5. CARON, A., G. DESROSIERS, P. J. W. Food: Nephtys caeca adults are carnivorous OLIVE, C. RETIERE, and C. NOZAIS. and predation by them is known to regulate 2004. Comparison of diet and feeding ac- other infaunal populations within a communi- tivity of two polychaetes, Nephtys caeca ty. They are a primary predator of Macoma (Fabricius) and Nereis virens (Sars), in an balthica (St. Lawrence Estuary) and are estuarine intertidal environment in Que- known to feed on smaller conspecifics bec, Canada. Journal of Experimental Ma- (Ambrose 1984; Caron et al. 2004). Juve- rine Biology and Ecology. 304:225-242. nile N. caeca, however, are herbivores 6. CRUMRINE, L. 2001. Polychaeta, p. 39- (Caron et al. 2004). 77. In: Identification guide to larval marine Predators: invertebrates of the Pacific Northwest. A. Behavior: Individuals are active, and are Shanks (ed.). Oregon State University good swimmers and burrowers (MacGinitie Press, Corvallis, OR. 1935). Bioturbation from N. caeca has been 7. DNESTROVSKAYA, N. Y., and I. A. shown to homogenize particles in the first JIRKOV. 2011. Microscopical studies of few centimeters of sediment while burrowing Nephtyid chaetae (Annelida: Polychaeta: or moving (Piot et al. 2008). Nephtyidae) from Northern Europe and Arctic. Italian Journal of Zoology. 78:219- Bibliography 228. 1. AMBROSE, W. G. 1984. Increased emi- 8. FERNALD, R. L., C. O. HERMANS, T. C. gration of the amphipod Rhepoxynius LACALLI, W. H. WILSON, JR, and S. A. abronius (Barnard) and the polychaete WOODIN. 1987. Phylum Annelida, Class Nephtys caeca (Fabricius) in the pres- Polychaeta, p. 138-195. In: Reproduction ence of invertebrate predators. Journal of and development of marine invertebrates Experimental Marine Biology and Ecolo- of the northern Pacific coast. M. F. Strath- gy. 80:67-75. mann (ed.). University of Washington 2. BENTLEY, M. G., P. J. W. OLIVE, P. R. Press, Seattle, WA. GARWOOD, and N. H. WRIGHT. 1984. 9. HARTMAN, O. 1938. Review of the anne- The spawning and spawning mechanism lid worms of the family nephtyidae from of Nephtys caeca (Fabricius, 1780) and the northeast pacific, with descriptions of Nephtys hombergi Savigny, 1818 five new species. Proceedings of the Unit- (Annelida, Polychaeta). Sarsia. 69:63-68. ed States National Museum. 85:143-158. 3. BLAKE, J. A., and R. E. RUFF. 2007. 10. HARTMAN, O. 1968. Atlas of the erranti- Polychaeta, p. 309-410. In: The Light ate polychaetous annelids from California. and Smith manual: intertidal inverte- Allan Hancock Foundation, University of

Southern California, Los Angeles. G. W. Rouse and F. Pleijel (eds.). Science 11. HILBIG, B. 1997. Family Nereididae, p. Publications, Enfield, NH. 291-316. In: Taxonomic atlas of the ben- 19. PORCH, L. L. 1970. Polychaetes of Coos thic fauna of the Santa Maria Basin and Bay. In: OIMB Unpublished Student Re- Western Santa Barbara Channel. Vol. 4. port, Summer 1970. J. A. Blake, B. Hilbig, and P. H. Scott 20. RAINER, S. F. 1991. The genus Nephtys (eds.). Santa Barbara Museum of Natural (Polychaeta, Phyllodocida) of Northern Eu- History, Santa Barbara, CA. rope: a review of species, including the 12. KOZLOFF, E. N. 1993. Seashore life of description of N. pulchra sp-n and a key to the northern Pacific coast: an illustrated the Nephtyidae. Helgolander Meeresunter- guide to Northern California, Oregon, suchungen. 45:65-96. Washington, and British Columbia. Uni- 21. THORSON, G. 1946. Reproduction and versity of Washington Press, Seattle, larval development of Danish marine bot- WA. tom invertebrates, with special reference 13. LACALLI, T. C. 1980. A guide to the ma- to the planktonic larvae in the Sound rine flora and fauna of the Bay of Fundy: (Oresund). Medd fra Komm Danmarks polychaete larvae from Passamaquoddy Fisk Havunder-sogelser Ser Plankton. 4:1- Bay. Canadian Technical Report of Fish- 523. eries and Aquatic Sciences. 940:1-27. 14. LOVELL, L. L. 1997. A review of six spe- Updated 2015 Nephtys cies of (Cuvier, 1817) T.C. Hiebert (Nephtyidae: Polychaeta) described from the eastern Pacific. Bulletin of Marine Science. 60:350-363. 15. MACGINITIE, G. E. 1935. Ecological as- pects of a California marine estuary. American Midland Naturlist. 16:629-765. 16. OLIVE, P. J. W. 1977. Life-history and population-structure of polychaetes Nephtys caeca and Nephtys hombergii with special reference to growth rings in teeth. Journal of the Marine Biological Association of the United Kingdom. 57:133-150. 17. PIOT, A., A. ROCHON, G. STORA, and G. DESROSIERS. 2008. Experimental study on the influence of bioturbation performed by Nephtys caeca (Fabricius) and Nereis virens (Sars) annelidae on the distribution of dinoflagellate cysts in the sediment. Journal of Experimental Marine Biology and Ecology. 359:92-101. 18. PLEIJEL, F., and G. W. ROUSE. 2006. Phyllodocida, p. 431-496. In: Reproduc- tive biology and phylogeny of annelida.