Boccardia Proboscidea Class: Polychaeta, Sedentaria, Canalipalpata

Phylum: Annelida

Boccardia proboscidea Class: Polychaeta, Sedentaria, Canalipalpata

Order: Spionida, Spioniformia A burrowing spionid worm Family: Spionidae

Taxonomy: Boccardia proboscidea’s senior Trunk: subjective synonym, Polydora californica Posterior: Pygidium is a round, flaring (Treadwell, 1914) and an un-typified name, disc with four unequal lobes where dorsal Spio californica (Fewkes, 1889) were both lobes are smaller (Fig. 4) (Hartman 1969). suppressed in 2012 by the International Parapodia: Biramous after first setiger. Commission on Zoological Nomenclature Podia on the first setiger are not lobed, small (ICZN, case 3520). The widely cited and and inconspicuous. The second setiger's used name, Boccardia proboscidea parapodial lobes become twice as large as (Hartman, 1940) was conserved (ICZN the first's, and continue to worm posterior. 2012). Setae (chaetae): All setae are simple and include bunches of short, capillary spines to se- Description tiger six (except for modified setiger five) Size: Specimens up to 30–35 mm in length (Figs. 5a, b). A transverse row of and 1.5 mm in width, where length extends approximately eight neuropodial uncini with age (Hartman 1940). The illustrated (hooded hooks) with bifid (two-pronged) tips specimen has approximately 130 segments begins on setiger seven and continues to (Fig. 1). posterior end (Fig. 5e), with bunches of Color: Yellow-orange with red branchiae capillary setae below them (until setiger 11). and dusky areas around prostomium and Notosetae of setiger five are heavy, dark and parapodia (Hartman 1969). Sato-Okoshi arranged vertically in two rows of five with and Okoshi (1997) report black pigment fol- pairs of long, falcate spines (Fig. 5c) and lowing the prostomial ridge and palpal shorter brush-topped clubs (Fig. 5d) while the grooves. neurosetae are capillary (Hartman 1969). All General Morphology: Spionid polychaete notosetae are capillary except for those of with thick anterior palps, conspicuous seg- setiger five. mentation. Eyes/Eyespots: Up to six (4–6) eyespots be- Body: Long, depressed, somewhat flattened tween palpal bases (Fig. 3). body that tapers posteriorly (Hartman 1940). Anterior Appendages: Long, simple, longitu- First setiger small capillary setae in bunches dinally grooved tentacle-like palps (family Spi- (Fig. 5a). Setiger five is modified with two onidae) (Fig. 1). kinds of dark, strong setae in notopodia. Branchiae: The gill-like structures in this spe- Setiger five is almost twice the length of cies are long, single vascular processes that setiger four (Figs. 2, 3) (Hartman 1940). are present on setigers 2–4, and from setiger Anterior: Prostomium long, rounded, seven to (almost) the worm posterior end without medial groove and snout-like (Figs. 2, 3). (species proboscidea, Hartman 1940). Burrow/Tube: Individuals build vertical, U- Caruncle (sensory organ) present and shaped burrows in rocky shale or amongst extends to setiger three (Fig. 3) (Hartman and within bivalve shells (Bailey-Brock 2000). 1969).

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. Boccardia proboscidea. 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.

Pharynx: California and has no notosetae on setiger Genitalia: one, only neurosetae (Blake and Ruff 2007). Nephridia: Its bristle-topped setae (on setiger five) have a small accessory tooth at the distal Possible Misidentifications end and the posterior notopodia have Spionid polychaetes are distin- acicular setae (Blake 1975). This species guished by their long palps. Two other pol- bores in coralline algae, hermit crab shells ychaete families have long palps: the and the jingle shell Pododesmus sp. Magelonidae, with adhesive palps (not Boccardia columbiana resembles B. long and flowing) and with flattened spade- proboscidea most closely. Its chief like prostomiums and the Chaetopteridae difference is that the fascicles of fine setae which have palps, but their bodies are very on setiger one are long and fanned forward, obviously divided into three quite different where they are short on B. proboscidea. regions. This species is reddish-brown, and bores The genus Boccardia contains 22 into wood pilings and coarse algae. species (Simon et al. 2010) and members Boccardia proboscidea was the only of this genus and the genus Boccardiella member of this genus found in Oregon by have branchiae on setigers anterior to se- Hartman and Reish (Hartman and Reish tiger five. These two local genera differ in 1950). the kinds of modified setae on the fifth se- Boccardia polybranchia and B. tricus- tiger, Boccardiella species have one pa have two kinds of setae on setiger five. (simple and falcate) while Boccardia have Boccardia polybranchia is green to reddish- two (one simple and falcate and the sec- yellow in color and has a notched prostomi- ond expanded and club-like) (Blake and um. Its first setiger lacks notosetae, it has Ruff 2007). only 60–80 segments and a pygidium that is Boccardiella hamata (=Boccardia a thick ring. Boccardia polybranchia is a uncata) has recurved spines, rather than cosmopolitan species that lives in estuarine straight bifid uncini, on its posterior para- mud and is reported in western Canada podia and the pygidium has two lappets (Hartman 1969) but not between central (Hartman 1969). It is common in oyster California and Oregon (Blake and Ruff beds and builds tubes in mudflats or bores 2007). Boccardia tricuspa has two kinds of holes into hermit crab and bivalve shells setae on setiger five, which are falcate and and is reported from central California to tridentate (not bruso-topped). Its branchiae Oregon (Blake and Ruff 2007) and in Van- are anterior to setiger five and are small and couver Island, Canada (Sato-Okoshi and inconspicuous. It bores in molluscs and is Okoshi 1997). Boccardiella truncata is usually a more southern species than B. green in color, has a saucer-like pygidium proboscidea. and a truncate anterior end. It is not usually estuarine and is, instead, more common Ecological Information intertidally in sandstone cliffs in northern Range: Type locality is Caspar, California. California (Blake and Ruff 2007). NE Pacific range, western Canada south to Two species of Boccardia have both southern California (Hartman 1969). falcate and brush-topped setae on setiger Local Distribution: Coos Bay distribution in- five as seen in B. proboscidea. Boccardia cludes the outer rocky coast and offshore berkeleyorum is reported from central (Hartman and Reish 1950).

Habitat: Burrows are found amongst Mytilus adelphophagy (consumption of one embryo spp. (mussel) colonies and individuals by another) on smaller larvae. These inhabit a variety of niches (Hartman 1940). developmental strategies were summarized Salinity: Collected at 30 and exhibits a by Oyarzun et al. (2011) and consists of great toleration for salinity variation females which produce capsules with 1) (Hartman 1940). planktotrophic larvae, 2) planktotrophic larvae Temperature: Boccardia proboscidea resi- and nurse eggs and 3) planktotrophic larvae, dence in tidepools is evidence of a wide nurse eggs and adelphophages larvae. The temperature tolerance (Hartman 1940). variation in capsule composition depends on Tidal Level: High rocky intertidal pools and female behavior and latitude, where in high intertidal crevices (Blake 1975). individuals at higher latitudes produce more Associates: Mytilus spp. and its accompa- adelphophages larvae which emerge at nying organisms in rocky crevices. An addi- advanced stages and settle quickly (Oyarzun tional associate is the small, red harpacti- et al. 2011). coid copepod, Tigriopus sp. (Hartman 1940). Larva: Larval stages found in plankton in the Abundance: The only Boccardia species summer (Hartman and Reish 1950) and larval found in Oregon by Hartman and Reish, development is described (Blake and (1950) where it is the most common Kudenov 1981; Gibson and Smith 2004). member of a common family (Hartman Larvae develop through trochophore and 1940). metatrochophore stages and have a single dorsal chromatophore in young stages and Life-History Information two beginning at the 5-setiger stage (Blake Reproduction: Eggs, in five or more cap- and Kudenov 1981). Larvae also possess sules of 50 eggs each, are deposited in a three pairs of black eyes. Once larvae reach tube and aerated during development by the nectochaete stage, they can be identified adult's rhythmic movement (Hartman 1940). by adult characters (e.g. modified setae on Egg capsules are present from August to setiger five, branchiae beginning on setiger) October in Barkley Sound, Canada (Sato- (Crumrine 2001). Okoshi and Okoshi 1997). Embryonic de- Juvenile: Metamorphosis is gradual and oc- velopment is rapid and easily occurs in the curs between the 13–15 setiger stages where lab where capsules in the same tube often larval structures are lost and juvenile features exhibit different development modes. For are gained. Juvenile features include setal this reason, the larval development of B. morphology like that of the adults where the proboscidea is a model for poecilogony length and number increase with growth. (differing developmental strategies within the Branchiae develop first on setigers 7–9 and same species, Gibson et al. 1999) among appear later in anterior and posterior setigers. spionids (Blake and Ruff 2007). Woodwick The juvenile pygidium is like that of the adult, (1977) reported two egg sizes, 100 and 150 comprising four lobes (Gibson and Smith µm, which emerge as planktotrophic 2004). (emerging at 3-setiger stage) and Longevity: lecithotrophic (emerging at up to 15-setiger Growth Rate: stage) larvae, respectively. Later, Blake and Food: Spionids feed by sweeping their tenta- Kudenov (1981) observed nurse eggs within cles across the surface of substrate where capsules which are ingested by and support particles are collected and wiped on the un- larvae within the capsule, coupled with derside of prostomium (Dales 1967). Boccar-

dia proboscidea eats small copepods (Polychaeta : Spionidae), with multiple de- (Hartman 1940) and is a voracious predator velopment modes. Marine Biology. on algal particles, Bryozoa, Hydrozoa, other 134:743-751. attached and free-swimming animals 8. GIBSON, G. D., and H. L. SMITH. 2004. (Hartman 1940). From embryos to juveniles: morphogene- Predators: sis in the spionid Boccardia proboscidea Behavior: A colonial burrower, B. probos- (Polychaeta). Invertebrate Biology. cidea can be seen with tentacles protruding 123:136-145. from burrow (Ricketts and Calvin 1971). 9. HARTMAN, O. 1940. Boccardia proboscidea, a new species of spionid worm from Bibliography California. Journal of the Washington 1. BAILEY-BROCK, J. H. 2000. A new rec- Academy of Science. 30:382-397. ord of the polychaete Boccardia probos- 10. —. 1969. Atlas of the sedentariate poly- cidea (family Spionidae), imported to Ha- chaetous annelids from California. Allan wai'i with oysters. Pacific Science. 54:27- Hancock Foundation, University of South- 30. ern California, Los Angeles, CA. 2. BLAKE, J. A. 1975. Phylum Annelida: 11. HARTMAN, O., and D. J. REISH. 1950. Class Polychaeta, p. 151-243. In: Light's The Marine annelids of Oregon. Oregon manual: intertidal invertebrates of the State College, Corvallis, Oregon. central California coast. S. F. Light, R. I. 12. INTERNATIONAL COMMISSION ON ZO- Smith, and J. T. Carlton (eds.). University OLOGICAL NOMENCLATURE. 2012. of California Press, Berkeley. Opinion 2303. (Case 3520) Boccardia pro- 3. BLAKE, J. A., and J. D. KUDENOV. boscidea Hartman, 1940 (Annelida, Spio- 1981. Larval development, larval nutrition nidae): specific name conserved. Bulletin and growth for two Boccardia species of Zoological Nomenclature. 69:232-234. (Polychaeta: Spionidae) from Victoria, 13. OYARZUN, F. X., A. R. MAHON, B. J. Australia. Marine Ecology Progress Se- SWALLA, and K. M. HALANYCH. 2011. ries. 6:175-182. Phylogeography and reproductive varia- 4. BLAKE, J. A., and R. E. RUFF. 2007. tion of the poecilogonous polychaete Boc- Polychaeta, p. 309-410. In: The Light cardia proboscidea (Annelida: Spionidae) and Smith manual: intertidal inverte- along the West Coast of North America. brates from central California to Oregon. Evolution & Development. 13:489-503. J. T. Carlton (ed.). University of California 14. RICKETTS, E. F., and J. CALVIN. 1971. Press, Berkeley, CA. Between Pacific tides. Stanford University 5. CRUMRINE, L. 2001. Polychaeta, p. 39- Press, Stanford, California. 77. In: Identification guide to karval nar- 15. SATO-OKOSHI, W., and K. OKOSHI. ine unvertebrates of the Pacific North- 1997. Survey of the genera Polydora, Boc- west. A. Shanks (ed.). Oregon State Uni- cardiella and Boccardia (Polychaeta, Spio- versity Press, Corvallis, OR. nidae) in Barkley Sound (Vancouver Is- 6. DALES, R. P. 1967. Annelids. land, Canada), with special reference to Hutchinson & Co., Ltd., London. boring activity. Bulletin of Marine Science. 7. GIBSON, G., I. G. PATERSON, H. TAY- 60:482-493. LOR, and B. WOOLRIDGE. 1999. Molec- 16. SIMON, C. A., T. M. WORSFOLD, L. ular and morphological evidence of a sin- LANGE, and J. STERLEY. 2010. The ge- gle species, Boccardia proboscidea nus Boccardia (Polychaeta: Spionidae)

associated with mollusc shells on the south coast of South Africa. Journal of the Marine Biological Association of the United Kingdom. 90:585-598. 17. WOODWICK, K. H. 1977. Lecithotrophic larval development in Boccardia proboscidea Hartman, p. 347-371. In: Essays on polychaetous annelids in memory of Dr. Olga Hartman. D. J. Reish and K. Fauchald (eds.). Allan Hancock Founda- tion, University of Southern California, Los Angeles, California. Updated 2014 T.C. Hiebert