Pseudopolydora Kempi Class: Polychaeta, Sedentaria, Canalipalpata

Phylum: Annelida

Pseudopolydora kempi Class: Polychaeta, Sedentaria, Canalipalpata

Order: Spionida, Spioniformia A tube-dwelling sedentary polychaete worm Family: Spionidae

Taxonomy: Pseudopolydora kempi was de- Anterior: Prostomium rather blunt, scribed by Southern in 1921 from a brackish with small bi-lobed lateral horns (Fig. 2). No water lake in India and subsequently found caruncle, but with occipital cirrus between in Japan, for which the subspecies P. kempi palps (Fig. 2). japonica was later designated by Imajima Trunk: and Hartman (1964). When this species Posterior: Pygidium cup shaped, flar- was found in California, another subspecies ing and with two dorsal projections or pro- was designated (P. kempi californica) (Light cesses (Fig. 4). 1969). However, after re-examining the type Parapodia: Biramous. Anterior noto- and specimens of P. kempi californica, Blake neurosetae include several kinds of capillary and Woodwick (1975) determined that the and limbate spines (Figs. 5a and b). Notopo- subspecific designations were not necessary dial post-setal lobes on setigers 2–5 (Fig. 3). and, instead, P. kempi, was likely introduced Neuropodial lobes reduced at setiger eight, to California from Japan (Carlton 1975; when they become tori, with hooded hooks. Blake and Woodwick 1975; Light 1978; Co- Setae (chaetae): Modification on setiger five hen and Carlton 1995). Although the spe- consists of a special J-shaped double row of cies which occurs in Oregon is currently re- falcigers (Fig. 5a) (sp. kempi, Light 1978), in ferred to as P. kempi, developmental differ- addition to typical bilimbate setae (Fig. 5b). ences suggest that this species is not the Setiger one with neurosetal fascicle only, no same as those from India and Japan (Blake notosetae (Figs. 2, 3). Posterior neurosetae and Ruff 2007). (from setiger eight) are bidentate hooded hooks in row of 18–20 (Fig. 5c) (genus Description Pseudopolydora, Light 1978). Size: Individuals up to 28 mm in length Eyes/Eyespots: Four small eyes, an outer (Blake 1975). Our specimens (from Coos pair anterior and darker with inner pair sub- Bay) are 16 mm in length and 1.5 mm in dermal, close together and between palps width, with nearly 40 segments. Average (Fig. 2). specimens are 12 mm in length, with 50 Anterior Appendages: Anterior with small bi- body segments (Light 1978). lobed lateral horns and two conspicuous Color: Pigmentation is variable (Light 1978), palps each about 1/3 body length (Fig. 1). but typically pale, with transverse interseg- Branchiae: Present on 15–25 segments, be- mental rows of black spots anteriorly on ginning on setiger seven (Light 1978) (Figs. 1, most specimens (sp. kempi, Blake 1975) 3). (Fig. 3). Burrow/Tube: Tube mucoid and animal with- General Morphology: Body thickened ante- in is completely hidden, except for extended riorly, becoming narrow posteriorly. No divi- palps. sion of body into distinct sections. Fifth se- Pharynx: tiger only slightly modified (Fig. 4). Body:

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. Pseudopolydora kempi. 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.

Genitalia: only 10–12 pairs (Light 1978). The major Nephridia: setae on setiger five are U- or J-shaped (Light 1978), but its pygidium is narrow and cup-like, Possible Misidentifications lacking dorsal projections seen in P. kempi. Spionidae can be distinguished by a Pseudopolydora paucibranchiata is small (4–6 pair of long prehensile grooved palps which mm, rarely more than 12 mm in length (Light arise from the posterior peristomium (Blake 1978)) and its palps have yellow reflective 1996). Two other polychaete families have spots (Blake 1975). long palps: the Magelonidae, with adhesive Other genera in this common estuarine palps (not long and flowing) and with flat- family include Boccardia, Polydora and Py- tened spade-like prostomiums and the gospio. Boccardia have branchiae from Chaetopteridae, which have palps, but their setiger two and a strongly modified setiger bodies are very obviously divided into three five. Polydora also have a strongly modified quite different regions, which is not the case fifth setiger, their branchiae begin on setiger in spionids. The similar family Cirratulidae, six and they lack post-setal parapodial lobes may also have a large pair of palps, but they (Hartman 1969). Pygospio (see P. elegans) have tentacular filaments, which are lacking have branchiae beginning posterior to setiger in the spionids. Spionids also have hooded 10 and the fifth setiger is unmodified. Their hooks in posterior segments, may or may tubes are papery and clear, to which fine sand not have prostomial appendages or grains adhere. branchiae, the prostomium is well developed and fused with peristomium, the pharynx is Ecological Information without jaws and the setae are mostly simple Range: Type locality is a brackish water lake (Blake 1975). Often certain segments are in India (Blake and Woodwick 1975). Alt- highly modified and have special setae, for hough it is currently unknown if this species is example prostomial horns are present in widely distributed or consists of many cryptic some genera. Spionid parapodia are species from distant locations, the current biramous, with acicula (see Leitoscoloplos range includes locations in India, South Afri- pugettensis) and sometimes have stout ca, Kurile Islands and the Pacific coast. Cali- saber setae. There are 19 local spionid fornia populations were likely introduced with genera (Blake and Ruff 2007). oysters (Crassostrea) from Japan in the Pseudopolydora spionids can be dis- 1960s (Light 1978). tinguished from other genera by their unusu- Local Distribution: Collection sites in Coos al J-shaped row of hooks on setiger five Bay include South Slough. Individuals also (Fig. 3), and by their neuropodial hooded collected in the Columbia River estuary. hooks, which begin on setiger eight. Pseu- Habitat: Individuals inhabit mucoid tubes in dopolydora branchiae begin on setiger sev- sandy mud of bays and are often found out- en (Fauchald 1977). side beds of the mud shrimp Callianassa. Pseudopolydora paucibranchiata is Salinity: Brackish to nearly fresh water with the only other common Pacific Coast salinity ranging from 6.3–31.9. Collected at species in the genus Pseudopolydora (Blake salinities of 30 in Coos Bay. and Ruff 2007). Unlike P. kempi, it has a Temperature: 10–15 °C. rounded prostomium with no pigment stripes Tidal Level: Intertidal to shallow depths on the anterior segments. Branchiae are (Hartman 1969) and high intertidal (Coos Bay, present from setiger seven, but there are South Slough in Callianassa beds).

Associates: The amphipod, Eobrolgus spi- ulations from California and India and the Sea nosus is often found within the tubes of P. of Japan (Blake and Ruff 2007). kempi. Juvenile: Metamorphosis is marked by the Abundance: South Slough, June abun- transition of larval organs to juvenile struc- dance was measured in cores (15cm diame- tures. Anterior feeding palps, for example, ter x 13 cm depth) at three tidal heights. which do not process food during larval stag- High intertidal core (1.1 m MLLW) produced es are used for feeding in benthic juveniles 221 animals, mid intertidal (1 m MLLW) and adults. Juvenile palps are long and reach produced 4885 animals and low intertidal setiger eight. Larval cilia are lost, anterior (0.9 m MLLW), 4113 animals (Posey 1985). eyes are arranged as in adults, prostomium is bifurcate and larval setae are replaced by Life-History Information adult setae. Larval pigment is retained, but Reproduction: The reproduction and devel- gradually lost in developing juveniles. The opment of P. kempi has been described and digestive tract is complete and the pygidium varies with geographic location (Blake and begins as a cup-shaped structure but eventu- Woodwick 1975; Strikrishnadhas and Rama- ally develops dorsal projections (Blake and moorthi 1977; Myohara 1979; Radashevskii Woodwick 1975, Fig. 19–22). 1985; Blake and Ruff 2007). Females brood Longevity: eggs within capsules that are attached in Growth Rate: long strings where the number of eggs per Food: Primarily a deposit feeder, but can shift capsule (8–38 to 100–150) and number of to suspension feeding when water currents capsules per string is highly variable (9–18 increase, by forming palps into helical shapes. to 14–24). Eggs are 99–116 µm (Blake and Predators: Fish and shorebirds. Arnofsky 1999). California populations have Behavior: When lugworm Abarenicola sp. larvae that develop by ingesting nurse eggs, disturbs surface with castings, are released from their capsules at the 15- Pseudopolydora can move its tube location setiger stage and carry out a short pelagic (Wilson 1981). Furthermore, the presence or period before settling into the benthos. Pop- smell of Abarenicola pacifica has been shown ulations from India and the Sea of Japan, on to reduce settlement of P. kempi juveniles the other hand, lack nurse cells, are re- (Woodin 1985). leased at the 3-setiger stage and undergo a long planktotrophic larval stage before set- Bibliography tlement (Blake and Ruff 2007). 1. BLAKE, J. A. 1975. Phylum Annelida: P. kempi Larva: The larvae of are recog- Class Polychaeta, p. 151-243. In: Light's nizable by many adult characteristics includ- manual: intertidal invertebrates of the cen- ing a slight modification of the fifth setiger, tral California coast. S. F. Light, R. I. prominent parapodia, setae both simple and Smith, and J. T. Carlton (eds.). University pennoned and arranged in U-shaped rows, of California Press, Berkeley. and hooded hooks with a secondary tooth 2. —. 1996. Family Spionidae, p. 81-224. In: close to main fang (Crumrine 2001). Necto- Taxonomic atlas of the benthic fauna of chaete larvae are thick and fusiform with the Santa Maria Basin and Western Santa metamorphosis occurring at 13–20 setiger Barbara Channel. Vol. 6. J. A. Blake, B. stages or 1100–1800 µm in diameter (Blake Hilbig, and P. H. Scott (eds.). Santa Bar- and Arnofsky 1999). Like developmental bara Museum of Natural History, Santa mode, larval pigment can vary between pop- Barbara, CA.

3. BLAKE, J. A., and P. L. ARNOFSKY. The polychaetous annelids of Japan. Part 1999. Reproduction and larval develop- II. Allan Hancock Foundation Publications ment of the spioniform polychaeta with Occasional Papers. 26:239-452. application to systematics and phyloge- 12. LIGHT, W. J. 1969. Polydora narica, new ny. Hydrobiologia. 402:57-106. species, and Pseudopolydora kempi cali- 4. BLAKE, J. A., and E. R. RUFF. 2007. fornica, new subspecies, two new spionids Polychaeta, p. 309-410. In: Light and (Annelida: Polychaeta) from central Cali- Smith manual: intertidal invertebrates fornia. California Academy of Sciences. from central California to Oregon. J. Carl- 13. —. 1978. Spionidae: Polychaeta, An- ton (ed.). University of California Press, nelida. Boxwood Press, Pacific Grove, CA. Berkeley, CA. 14. MYOHARA, M. 1979. Reproduction and 5. BLAKE, J. A., and K. H. WOODWICK. development of Pseudopolydora kempi 1975. Reproduction and larval develop- japonica (Polychaeta: Spionidae), with ment of Pseudopolydora paucibranchiata special reference to the polar lobe for- (Okuda) and Pseudopolydora kempi mation. Journal of the Faculty of Science (Southern) (Polychaeta: Spionidae). Bio- Hokkaido University, Series VI. Zoology. logical Bulletin. 149:109-127. 21:355-364. 6. CARLTON, J. T. 1975. Introduced inter- 15. POSEY, M. H. 1985. The effects upon the tidal invertebrates, p. 17-25. In: Light's macrofaunal community of a dominant manual; intertidal invertebrates of the burrowing deposit feeder, Callianassa cali- central California coast. S. F. Light, R. I. forniensis, and the role of predation in de- Smith, and J. T. Carlton (eds.). University termining its intertidal distribution. Ph.D. of California Press, Berkeley. University of Oregon. 7. COHEN, A. N., and J. T. CARLTON. 16. RADASHEVSKY, V. I. 1985. The Larval 1995. Nonindigenous aquatic species in development of the polychaete Pseudo- a United States estuary: a case study of polydora kempi japonica in Peter the Great the biological invasions of the San Fran- Bay of the Sea of Japan. Biologiya Morya- cisco Bay and Delta. National Sea Grant Marine Biology: 39-46. Program, Connecticut. 17. SRIKRISHNADHAS, B., and K. RAMA- 8. CRUMRINE, L. 2001. Polychaeta, p. 39- MOORTHI. 1977. Development of Pseu- 77. In: Identification guide to larval ma- dopolydora kempi (Southern, 1921) in the rine invertebrates of the Pacific North- laboratory, p. 671-677. In: Proceedings of west. A. Shanks (ed.). Oregon State Uni- the symposium on warm water zooplank- versity Press, Corvallis, OR. ton. Special Publication UNESCO/NIO. 9. FAUCHALD, K. 1977. The polychaete 18. WILSON, W. H. 1981. Sediment-mediated worms: definitions and keys to the or- interactions in a densely populated infau- ders, families, and genera. Natural Histo- nal assemblage; the effects of the poly- ry Museum of Los Angeles County Sci- chaete Abarenicola pacifica. Journal of ence Series. 28:1-190. Marine Research. 39:735-748. 10. HARTMAN, O. 1969. Atlas of the seden- 19. WOODIN, S. A. 1986. Settlement of infau- tariate polychaetous annelids from Cali- na: larval choice. Bulletin of Marine Sci- fornia. Allan Hancock Foundation, Uni- ence. 39:401-407. versity of Southern California, Los Ange- Updated 2014 les, CA. T.C. Hiebert 11. IMAJIMA, M., and O. HARTMAN. 1964.