Pygospio Elegans Class: Polychaeta Order: Canalpalpita, Spionida a Spionid Polychaete Worm Family: Spionidae

Phylum: Annelida Pygospio elegans Class: Polychaeta Order: Canalpalpita, Spionida A spionid polychaete worm Family: Spionidae

Taxonomy: Several subjective synonyms each segment has paired branchiae, a (synonyms based on individual opinion) are ciliary band (single or double) and two indicated for Pygospio elegans, but rarely bands of tufted cilia (Gibson and used (Spio inversa, S. rathbuni, S. seticornis Harvey 2000). and Pygospio mimutus). Posterior: Pygidium with four pigmented conical cirri (sp. elegans) Description (Fig. 3), each possessing inner tufts of Sexual Dimorphism: Males may cilia (Gibson and Harvey 2000). have small pair of elongated dorsal Parapodia: Biramous, with cirriform lobes cirri (sometimes called "auxiliary gills") beginning on the second setiger, and on setiger two (Pygospio, Blake 1975) diminishing posteriorly. Neuropodia without which are often lost in collection. interramal pouches. Size: Individuals range in size from 10–15 Setae (chaetae): Notopodia with capillary mm in length, having 50–60 segments (Light setae only. Neuropodia with simple capillary 1978). This specimen (from Coos Bay) is 5 setae on setigers 1–8 and spoon-like hooded mm in length, 0.3 mm in width and has 36–37 hooks beginning on setigers eight and nine segments (Fig. 1). (sp. elegans) (Fig. 4). Color: Light, almost white with black Eyes/Eyespots: Four eyes present. Anterior markings on anterior segments (Fig. 2), but pair widely separated and lighter in color than not on proboscis (Light 1978). distal pair (Fig. 2). Some individuals have 2– General Morphology: Body slightly flattened 8 eyespots, irregularly arranged along dorsolaterally. Fifth setiger normal and not segments (Light 1978). strongly modified as is common among Anterior Appendages: Two thick grooved spionids. palps antero-laterally are used in feeding. Body: The body can be divided into four They are relatively short and thick as major regions (from anterior to posterior) compared to the palps of other spionids. including the anterior or head, the thorax and Branchiae: Numerous branchial pairs, first abdomen and, finally, the tail or posterior appearing on setigers 11–13. No branchiae (Gibson and Harvey 2000). on posterior-most segments. Anterior: Anterior with two massive Burrow/Tube: Tube morphology is species- short dorsolaterally grooved palps specific. It is long, papery, clear, covered with (Figs. 1, 5), which are often lost during fine sand grains and approximately 1 mm in collection. Prostomium is without diameter (Bolam 2004) (Fig. 5). horns, is blunt anteriorly, with lateral Pharynx: Bears proboscis with a partially swellings and is slightly bi-lobed in eversible conical sac. some specimens. Anterior is not Genitalia: conical but taper slightly (sp. elegans) Nephridia: (Fig. 2). Paired nuchal organs present (Gibson and Harvey 2000, Fig. 1). Possible Misidentifications Trunk: Thorax with 10–12 Spionidae can be distinguished by a abranchiate setigers where each pair of long prehensile grooved palps which setiger has a dorsal ciliary band and a arise from the posterior peristomium (Blake lateral tuft of cilia. The abdomen is composed of 25–35 setigers where

A publication of the University of Oregon Libraries and the Oregon Institute of Marine Biology Individual species: http://hdl.handle.net/1794/12681 and full 3rd edition: http://hdl.handle.net/1794/18839 Email corrections to [email protected] 1996). Two other polychaete families have Ecological Information long palps: the Magelonidae, with adhesive Range: Known range includes the northern palps (not long and flowing) and with Atlantic, northern Pacific and western Canada flattened spade-like prostomiums and the to California (Light 1978). Chaetopteridae, which have palps, but their Local Distribution: Coos Bay collection bodies are very obviously divided into three sites include South Slough. Individuals also quite different regions, which is not the case collected in the Columbia River estuary. in spionids. The similar family Cirratulidae, Habitat: Lives in papery sandy tubes in mud may also have a large pair of palps, but they or sand flats and is considered an indicator have tentacular filaments, which are lacking species for slight organic pollution (domestic in the spionids. Spionids also have hooded sewage, Germany (Anger 1977)). Their hooks in posterior segments, may or may ability to eat suspended as well as deposited not have prostomial appendages or matter increases survivability in variable branchiae, the prostomium is well environments (Taghon et al. 1980) (see developed and fused with peristomium, the Food). pharynx is without jaws and the setae are Salinity: Found at salinities ranging from 30 mostly simple (Blake 1975). Often certain to 28 in Coos Bay, but also tolerant of segments are highly modified and have salinities as low as 2 (Bolam 2004; Blake special setae, for example prostomial horns 2006). are present in some genera. Spionid Temperature: parapodia are biramous, with acicula (see Tidal Level: Only occurring at high tide level Leitoscoloplos pugettensis) and sometimes in Coos Bay. have stout saber setae. There are 19 local Associates: Associated species include the spionid genera (Blake and Ruff 2007). sabellid polychaete, Chone ecuadata (Blake The genus Pygospio is and Ruff 2007). characterized by branchiae beginning Abundance: South Slough, June abundance posterior to setiger 10 and species lack the was measured in cores (15 cm diameter x 13 dramatically modified fifth setiger of some cm depth) at three tidal heights. High spionids. Pygospio californica is the most intertidal core (1 m MLLW) produced 5–7 closely related species to P. elegans. It is animals, mid intertidal (1 m MLLW) produced green in life and is twice the size of P. 6–16 animals and low intertidal (0.9 m elegans (Hartman 1969). Pygospio MLLW), 13–20 animals. This species is most californica's prostomium has a tapered abundant in April (Posey 1985). conical tip and is not bi-lobed. Furthermore, P. californica has a narrow caruncle that Life-History Information reaches to the first setiger, where P. Reproduction: Both sexual and asexual (by elegans has no caruncle. Brown spots exist fragmentation or architomy) reproduction on the proboscis in P. californica and a have been reported for P. elegans (Blake and reddish brown ventral stripe runs down the Arnofsky 1999). Asexual reproduction has first 10–20 segments (Light 1978). The been shown to increase in frequency with paired branchiae in P. californica begin on warmer water temperatures (Armitage 1979; setiger 19, not on 11–13 as in P. elegans. Rasmussen 1953). Spermatophores are P. californica is found on intertidal sand flats mushroom-shaped and have a long tail (Blake and the only record in Oregon is from and Arnofsky 1999). Eggs are released Umpqua estuary, it is currently only reported through nephridial pores into egg capsules from central California (Blake and Ruff within the adult tube (Soderstrom, (Hartman 2007). 1941)). Asexual reproduction occurs in Other small tube dwelling spionids March–October while sexual reproduction include the genus Polydora, whose tubes takes place in winter months (November– are mucoid (Hartman 1969), and December) in False Bay, Washington Pseudopolydora kempi, which also has a (Fernald et al. 1987). However, sexual mucoid tube, a J-shaped row of falcigers on reproduction has been reported in Coos Bay setiger five and a cup-shaped pygidium. in April. Larvae are adelphophagic, ingesting

Hiebert, T.C. 2014. Pygospio elegans. 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. the unfertilized nurse eggs with which they Bibliography share a cocoon. Larva: Pygospio elegans larval development 1. ANGER, K. 1977. Benthic proceeds in a variety of ways (as in invertebrates as indicators of organic Pseudopolydora kempi) and has been pollution in the Western Baltic Sea. confirmed to be variations across a single Internationale Revue der Gesamten species (= poecilogony, Blake and Arnofksy Hydrobiologie und Hydrographie. 1999; Morgan et al. 1999; Blake 2006; 62:245-254. Kesäniemi et al. 2012). Larvae can ingest 2. ARMITAGE, D. L. 1979. The Ecology nurse eggs and hatch from their capsule at and reproductive cycle of Pygospio the 3-setiger stage and live as planktonic elegans Claparède (Polychaeta: larvae before settling at 17–20 setiger stage. Spionidae) from Tomales Bay, On the other hand, larvae can remain in the California: A Thesis. University of the egg capsule until they are large and hatch out Pacific. as a non-pelagic larva (Blake 2006, Fig. 3. BLAKE, J. A. 1975. Phylum Annelida: 13.17). These varying developmental modes Class Polychaeta, p. 151-243. In: may correspond to season, where Light's manual: intertidal invertebrates planktotrophic larvae are more common in of the central California coast. S. F. spring (Rasmussen 1973; Fernald et al. Light, R. I. Smith, and J. T. Carlton 1987). Larval characters include, lack of (eds.). University of California Press, modification on setiger five, prostomium not Berkeley. distally pointed, with frontal horns and 4. —. 1996. Family Spionidae, p. 81-224. branchiae limited to middle and posterior In: Taxonomic atlas of the benthic most setigers (except setiger two in males) fauna of the Santa Maria Basin and (Crumrine 2001). Western Santa Barbara Channel. Vol. Juvenile: 6. J. A. Blake, B. Hilbig, and P. H. Longevity: High mortality when disturbed by Scott (eds.). Santa Barbara Museum castings of polychaete Abarenicola (Wilson of Natural History, Santa Barbara, CA. 1981). 5. BLAKE, J. A. 2006. Spionida, p. 565- Growth Rate: Pygospio elegans can fully 638. In: Reproductive biology and regenerate their anterior including phylogeny of annelida. Enfield, prostomium and palps in 9–12 days post Science Publishers. Vol. 4. Science ablation (Lindsay et al. 2007). Publishers, Enfield. Food: A deposit, suspension and filter 6. BLAKE, J. A., and P. L. ARNOFSKY. feeder, searching mud surface and water for 1999. Reproduction and larval food. Individuals can switch from deposit to development of the spioniform suspension feeding when water flow velocity polychaeta with application to increases (Taghon et al. 1980). A most systematics and phylogeny. versatile feeder, P. elegans can filter feed by Hydrobiologia. 402:57-106. building a mucus net within or above its tube 7. BLAKE, J. A., and E. R. RUFF. 2007. (Fauchald and Jumars 1979; Bolam 2004). Polychaeta, p. 309-410. In: Light and Predators: Fish and shorebirds are major Smith manual: intertidal invertebrates predators because this species lives at high from central California to Oregon. J. tide level. Carlton (ed.). University of California Behavior: Moves tube when disturbed by Press, Berkeley, CA. activity of the large lugworm Abarenicola sp. 8. BOLAM, S. G. 2004. Population (Wilson 1981). Dense patches or structure and reproductive biology of aggregations of P. elegans are ephemeral, Pygospio elegans (Polychaeta: but have a significant effect on the immediate Spionidae) on an intertidal sandflat, infaunal community (Bolam and Fernandes Firth of Forth, Scotland. Invertebrate 2003). Biology. 123:260-268. 9. BOLAM, S. G., and T. F. FERNANDES. 2003. Dense

A publication of the University of Oregon Libraries and the Oregon Institute of Marine Biology Individual species: http://hdl.handle.net/1794/12681 and full 3rd edition: http://hdl.handle.net/1794/18839 Email corrections to [email protected] aggregations of Pygospio elegans 17. LIGHT, W. J. 1978. Spionidae: (Claparede): effect on macrofaunal Polychaeta, Annelida. Boxwood Press, community structure and sediments. Pacific Grove, CA. Journal of Sea Research. 49:171-185. 18. LINDSAY, S. M., J. L. JACKSON, and 10. CRUMRINE, L. 2001. Polychaeta, p. S. Q. HE. 2007. Anterior regeneration 39-77. In: Identification guide to larval in the spionid polychaetes Dipolydora marine invertebrates of the Pacific quadrilobata and Pygospio elegans. Northwest. A. Shanks (ed.). Oregon Marine Biology. 150:1161-1172. State University Press, Corvallis, OR. 19. MORGAN, T. S., A. D. ROGERS, G. 11. FAUCHALD, K., and P. A. JUMARS. L. J. PATERSON, L. E. HAWKINS, 1979. Diet of worms: a study of and M. SHEADER. 1999. Evidence for polychaete feeding guilds. poecilogony in Pygospio elegans Oceanography and Marine Biology. (Polychaeta: Spionidae). Marine 17:193-284. Ecology Progress Series. 178:121- 12. FERNALD, R. L., C. O. HERMANS, T. 132. C. LACALLI, W. H. WILSON, JR, and 20. POSEY, M. H. 1985. The effects upon S. A. WOODIN. 1987. Phylum the macrofaunal community of a Annelida, Class Polychaeta, p. 138- dominant burrowing deposit feeder, 195. In: Reproduction and Callianassa californiensis, and the role development of marine invertebrates of predation in determining its intertidal of the northern Pacific coast. M. F. distribution. Ph.D. University of Strathmann (ed.). University of Oregon. Washington Press, Seattle, WA. 21. RASMUSSEN, E. 1953. Asexual 13. GIBSON, G. D., and J. M. L. reproduction in Pygospio elegans HARVEY. 2000. Morphogenesis Claparède (Polychaeta sedentaria). during asexual reproduction in Nature. 171:1161-1162. Pygospio elegans Claparede 22. —. 1973. Systematics and ecology of (Annelida, Polychaeta). Biological the Isefjord marine fauna (Denmark) Bulletin. 199:41-49. with a survey of the eelgrass (Zostera) 14. HARTMAN, O. 1941. Polychaetous vegetation and its communities. annelids. Part III. Spionidae. Some Ophelia. 11:1-507. contributions to the biology and life 23. TAGHON, G. L., A. R. NOWELL, and history of Spionidae from California. P. A. JUMARS. 1980. Induction of Allan Hancock Foundation of Scientific suspension feeding in spionid Research. Allan Hancock Pacific polychaetes by high particulate fluxes. Expeditions. Reports. 7:299. Science (New York, N.Y.). 210:562-4. 15. —. 1969. Atlas of the sedentariate 24. WILSON, W. H. 1981. Sediment- polychaetous annelids from California. mediated interactions in a densely Allan Hancock Foundation, University populated infaunal assemblage; the of Southern California, Los Angeles, effects of the polychaete Abarenicola CA. pacifica. Journal of Marine Research. 16. KESÄNIEMI, J. E., P. D. RAWSON, S. 39:735-748. M. LINDSAY, and K. E. KNOTT. 2012. Phylogenetic analysis of cryptic speciation in the polychaete Pygospio elegans. Ecology and Evolution. 2:994-1007.