Glycera Robusta Phylum: Annelida Class: Polychaeta Order: Phyllodocida the Large Proboscis Worm Family: Glyceridae

Glycera robusta Phylum: Annelida Class: Polychaeta Order: Phyllodocida The large proboscis worm Family: Glyceridae

Taxonomy: There are relatively few glycerid other glycerids, Glycera robusta contains both genera and Glycera contains the largest coelomic cell hemoglobin and myoglobin number of species. Several authors have within the body wall musculature (Terwilliger attempted to divide this genus into subgenera and Garlick 1978). or genera. Groups have been divided based Anterior: Glycerids are on proboscideal organs (but no formal genera characterized by their conical, designated) by Hartman (1950) and recent annulated and elongate prostomia that reviews have synonymized species taper to a fine point anteriorly (Blake (Böggemann 2002) or split the genus into and Ruff 2007). The prostomium is many sibling species (e.g. O’Connor 1987). small and bears 10 biannulate rings, Glycera robusta, however, is unique in its the first being approximately one third universal stability as a valid taxon with a of the total length (Fig. 2). The reliable description (Scamit 2002; Blake and prostomium is longer than it is wide Ruff 2007). (Hilbig 1997) (Fig. 2) and the basal prostomial ring is fused with the Description peristomium (Blake and Ruff 2007). Size: The largest of the Glyceridae, up to Trunk: Segments posterior to 800 mm in length and 22 mm in width peristomium are considerably wider (Berkeley and Berkeley 1942; Hilbig 1997) than anterior most segments (Hilbig and can have up to 300 segments (Hartman 1997) (Fig. 2). The body bears 1968). The illustrated specimen, from South numerous, tightly packed segments. Slough of Coos Bay, was 500 mm in length Posterior: Anal end is narrow and and 20 mm in width. tapers to a fine point, adorned with a Color: Dark red, but can be yellow-brown. pair of small cirri (Fig. 1) (Berkeley and Their color gives rise to the common name of Berkeley 1942). the family Glyceridae – the blood worms Parapodia: Inconspicuous, biramous (Fig. 4) (Blake and Ruff 2007). and one-tenth of total body width in mid-body General Morphology: Long, stout and stiff regions (Hilbig 1997). Pre- and post-acicular worms with numerous densely packed lobes are equally bifid, the former resembling segments and a conical and annulated the ventral cirrus (Fig. 4) (Blake 1975) while anterior that tapers to a point (Glyceridae, the post-acicular lobes are short (Hartman Blake and Ruff 2007). Glycera robusta is 1968). The dorsal parapodial margin with dorsoventrally flattened in cross-section, is blister-like, fleshy branchia (see Branchiae widest in anterior regions, and gradually below) (Fig. 4). tapers to a point posteriorly (Hilbig 1997) (Fig. Setae (chaetae): Notosetae simple, slender 1). and finely serrated capillaries and neurosetae Body: Members of the family Glyceridae lack are compound spinigers that are slightly wider a separate circulatory system and their than notosetae (Hilbig 1997) (Fig. 5). coelomic fluid contains hemoglobin Eyes/Eyespots: No eyespots are visible, (Terwilliger et al. 1976; Morris et al. 1980) that although small eyespots may be present on is visible through the thin body wall. Unlike the terminal prostomial ring (Glyceridae, Blake and Ruff 2007).

A publication of the University of Oregon Libraries and the Oregon Institute of Marine Biology Individual species: http://hdl.handle.net/1794/12662 and full 3rd edition: http://hdl.handle.net/1794/18839 Email corrections to [email protected] Anterior Appendages: The anteriormost also have two jaws, not four, and a row of prostomial ring bears two pairs of small and denticles (called chevrons, Böggemann et bifurcate terminal cirri, but no other anterior al. 2012). The genus Glycera is appendages are present (Blake and Ruff characterized by its pointed and annulated 2007) (Fig. 2). prostomium, four small anterior cirri, Branchiae: Blister-like branchiae begin on peristomium fused to basal prostomial rings, setiger 23 (Fig. 1, 4). The branchiae protrude a cylindrical proboscis with proboscideal from dorsal parapodial walls from setiger 23– organs and four fang-like jaws terminally 34 and (in large specimens) branchiae are (Hilbig 1997). Members of this genus also present along the ventral parapodial walls have biramous parapodia with two pre-setal beyond setiger 35 (Hilbig 1997). The and 1–2 post-setal lobes, short dorsal cirri presence of branchiae ventrally and dorsally and elongate ventral cirri. They usually was once thought to be characteristics of have branchiae, simple notosetae and different species, but was found to be a compound neurosetae. There are currently character that varies with individual size five local Glycera species (Blake and Ruff (Berkeley and Berkeley 1942; Hilbig 1997). 2007). Burrow/Tube: G. americana, with four-lobed Pharynx: Bears large and powerful parapodia and branched, retractile proboscis (up to 26 mm long) (Hartman branchiae (Hartman and Reish 1950), is 1968). When fully everted, four terminal black found intertidally to 120 m. G. nana chitinous jaws are visible, each jaw composed (=capitata), another large species (but only of a hook-like fang and a V-shaped support up to 100 mm), with two pre-setal (called an aileron). The ailerons in G. robusta parapodial lobes, and one post-setal lobe, consist of a thick outer ramus and a very thin but no branchiae. G. macrobranchia inner ramus (Hilbig 1997). The proboscis (=convoluta) has a single non-retractile epithelium is densely covered with pear- branchia and 14 to 16 annulations in the shaped papillae, called proboscideal organs prostomium (unlike 10 in G. robusta). G. (Berkeley and Berkeley 1942; Blake and Ruff dibranchiata has two finger-like branchiae, 2007) (Fig. 3). These proboscideal organs one above and one below the setal lobe, are oval to flask shaped, bearing 6–8 ridges this species is commonly harvested for bait (Hilbig 1997) (Fig. 3). in Canada and along the eastern coast of Genitalia: the United States. G. tenuis has but one Nephridia: pre-setal parapodial lobe on its posterior setiger, is only 80 mm in length, when Possible Misidentifications mature, and 13–16 proboscideal organs Distinctive characters of the Glyceridae (Blake and Ruff 2007). include a pointed and annulated prostomium with two pairs of anterior Ecological Information appendages and a long, powerful proboscis Range: Type locality is California (Hartman with four hook-shaped jaws and accessory 1968). Known range includes the western ailerons (Böggemann et al. 2012). The (Japan) and eastern Pacific (Washington to other proboscis worm family, the southern California), however, G. robusta is Goniadidae, is morphologically similar to the not currently in Puget Sound Keys (e.g. Glyceridae and identification requires Kozloff 1974). examination of the parapodia and Local Distribution: Coos Bay distribution proboscis. The Goniadidae have bodies includes many sites in the bay, such as South divided into three parts by different types of Slough and Fossil Point, and outside the bay, parapodia and their bodies are usually more in the small sandy beaches at Cape Arago cylindrical and slender than is seen in the (Hartman and Reish 1950). Glyceridae (Hilbig 1997; Blake and Ruff Habitat: Glycera robusta preferred 2007). The everted proboscis of goniadids substrates include beds of black mud

Hiebert, T.C. 2015. Glycera robusta. 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. (Ricketts and Calvin 1971), gravelly sand 3. BLAKE, J. A., and R. E. RUFF. 2007. (Hartman 1968), and sand and cobble Polychaeta, p. 309-410. In: The Light sediments (Blake and Ruff 2007). and Smith manual: intertidal Salinity: invertebrates fromc California to Temperature: Oregon. J. T. Carlton (ed.). University Tidal Level: Intertidal and shelf depths of California Press, Berkeley, CA. (Hartman 1968; Blake and Ruff 2007). 4. BOEGGEMANN, M. 2002. Revision of Associates: the Glyceridae GRUBE 1850 Abundance: (Annelida: Polychaeta). Abhandlungen der Senckenbergischen Life-History Information Naturforschenden Gesellschaft:1-249. Reproduction: The reproduction and 5. BOEGGEMANN, M., C. BIENHOLD, development of G. robusta is not known. and S. M. GAUDRON. 2012. A new Most glycerids become epitokous in spring species of Glyceridae (Annelida: and summer months (Morris et al. 1980) and 'Polychaeta') recovered from organic females release lens-shaped oocytes substrate experiments at cold seeps in (Fernald et al. 1987). the eastern Mediterranean Sea. Larva: Development proceeds via an Marine Biodiversity. 42:47-54. eyeless trochophore larva. These 6. CRUMRINE, L. 2001. Polychaeta, p. planktotrophic larvae feed on diatoms and 39-77. In: Identification guide to larval detritus, eventually developing to epibenthic marine invertebrates of the Pacific stages and become predatory once their jaws Northwest. A. Shanks (ed.). Oregon are fully formed (Fernald et al. 1987). Many State University Press, Corvallis, OR. of the locally known Glycera species produce 7. FERNALD, R. L., C. O. HERMANS, T. nectochaete larvae that are characterized by C. LACALLI, W. H. WILSON, JR, and a long, pointed and annulated prostomium as S. A. WOODIN. 1987. Phylum is seen in the adults (Fig. 14, Crumrine 2001). Annelida, Class Polychaeta, p. 138- Juvenile: 195. In: Reproduction and Longevity: development of marine invertebrates Growth Rate: of the northern Pacific coast. M. F. Food: Glycerids are mainly carnivorous Strathmann (ed.). University of (Crumrine 2001). Washington Press, Seattle, WA. Predators: 8. HADERLIE, E. C. 1980. Polychaeta: Behavior: Individuals use their proboscis to the marine annelid worms, p. 448-489. burrow quickly. In: Intertidal invertebrates of California. R. H. Morris, D. P. Abbott, and E. C. Haderlie (eds.). Stanford University Bibliography Press, Stanford, CA. 9. HARTMAN, O. 1968. Atlas of the 1. BERKELEY, E., and C. BERKELEY. Errantiate Polychaetous Annelids from 1942. North Pacific Polychaeta, chiefly California. Allan Hancock Foundation, from the west coast of Vancouver University of Southern California, Los Island, Alaska, and Bering Sea. Angeles. Canadian Journal of Research. 10. HARTMAN, O., and D. J. REISH. 20:183-208. 1950a. Goniadidae, Glceridae and 2. BLAKE, J. A. 1975. Phylum Annelida: Nephtyidae. Allan Hancock Pacific Class Polychaeta, p. 151-243. In: Expedition. 15:1-182. Light's manual: intertidal invertebrates 11. —. 1950b. The marine annelids of of the central California coast. S. F. Oregon. Oregon State College, Light, R. I. Smith, and J. T. Carlton Corvallis, Oregon. (eds.). University of California Press, 12. HILBIG, B. 1997. Family Glyceridae, Berkeley. p. 187-205. In: Taxonomic atlas of the