Invasion History

First Non-native North American Tidal Record: 1935
First Non-native West Coast Tidal Record: 1935
First Non-native East/Gulf Coast Tidal Record: 1973

General Invasion History:

The spionid polychaete Boccardiella ligerica was described from the Loire Estuary, France and is believed to be native to brackish waters of Europe. Its range extends from the North Sea coast of Germany, to the Netherlands, France, and Portugal (Blake and Woodwick 1971; Wolff 1973; Cunha and Moreira 1995). It is believed to be introduced to the inner Baltic (Finland; Eliason and Haahtela 1969). In the Western Atlantic, this polychaete has been reported from New York to Texas, Puerto Rico, and Argentina (Blake 1983; Wern 1985; Kravitz et al. 1987; Angradi et al. 1987; Chesapeake Bay Program 1998; Orensanz et al. 2002; New York State Department of Transportation 2012). In the Northeast Pacific, it was first reported from Newport Bay, California in 1935, and was found in San Francisco Bay in 1953, where it is now very abundant (Cohen and Carlton 1995). Genetic comparisons of European, Western Atlantic, and Eastern Pacific species have not been made. This polychaete is frequently associated with the widespread invader Ficopomatus enigmaticus (Light 1977; Blake 1983). It is likely that this species has been widely distributed by ship's ballast water. Boccardiella ligerica is also abundant on the hulls of heavily fouled ships (Llansó et al. 2011).

North American Invasion History:

Invasion History on the West Coast:

A single specimen of Boccardiella ligerica was collected by Olga Hartman in Newport Bay, California (CA) in 1935 (Kudenov 1983). However, there are no further records of this polychaete from marine-estuarine waters of southern California (Light 1977). In San Francisco Bay, B. ligerica was collected in the San Pablo Channel in 1954, and later found in Suisun Bay, Honker Bay, and the Delta-Mendota (1971-193, Light 1977; Carlton 1979). By the 1970s-1980s, this polychaete was found to be a dominant form in low-salinity regions of the San Francisco estuary, including the lower Sacramento River during dry years (Siegfreid 1980; Peterson and Vayssieres 2010). Boccardiella ligerica was collected in the Bolinas Lagoon, outside San Francisco Bay in 1968 (CAS-IZ 126919, California Academy of Science 2014), but we have no information on its establishment. Established populations of B. ligerica were found in the interior of California, in the Alamo River, which flows into the Salton Sea, an athalassic salt lake (1979, Kudenov 1983). This desert river is nominally fresh, but probably has a high salt content.

Invasion History on the East Coast:

Most records of Boccardiella ligerica on the East Coast come from species lists in 'gray literature' reports, with little information on species location or habitats. It is possible that this worm has been overlooked due to its small size, and confusion with B. hamata and other spionids. Boccardiella ligerica has been reported from the St. Johns estuary, Florida (Mattson 2011); the Neuse estuary, North Carolina (Vittor Associates 1999); the Chesapeake Bay (Chesapeake Bay Program 1998); Delaware Bay (Anderson et al. 2003); and the Mullica River and Hackensack estuaries, New Jersey (Angradi et al. 2001; Meadowlands Environmental Research Institute 2004) and in the Hudson estuary at the Tappan Zee Bridge (2006 - 2009, New York State Department of Transportation 2012).  In Chesapeake Bay, B. ligerica was reported in 1990 benthic sampling data (Chesapeake Bay Program 1998; Gillett and Schaffner 2009), but was not seen during earlier collections. This worm has been found in several oligo-mesohaline sites in Virginia and Maryland: James River (Swann's Point); Pamunkey River; Rappahannock River; Patuxent River; Nanticoke River; and the Potomac River (Route 301 Bridge) (Chesapeake Bay Program 2001).

Invasion History on the Gulf Coast:

The earliest reported collection of Boccardiella ligerica from the Gulf of Mexico was in the Aransas National Wildlife Refuge, on San Antonio Bay, Texas in 1973. Only a single specimen was collected there, but an established population was found in a brackish (10-14 PSU) marsh at Sea Rim State Park, near Sabine Lake (Wern 1985). In 1984, another population was found to be established in the Crystal River estuary, on the Gulf Coast of Florida (Kravitz 1987).

Invasion History Elsewhere in the World:

In the Western Atlantic, Boccardiella ligerica was collected from the Rio Espiritu Santo estuary in Puerto Rico in 1978 (Blake 1983; US National Museum of Natural History 2014). Further south, this polychaete was collected in Uruguay, in the La Plata estuary in 1938 (Blake 1983; Orensanz et al. 2002). It is now abundant in lagoons on the Atlantic coast south of the La Plata estuary (Gappa et al. 2001; Orensanz et al. 2002). An identification of B. ligerica from South Africa (Day 1955, cited by Blake and Woodwick 1971) is considered questionable.


Boccardiella ligerica is a small infaunal polychaete, characteristic of brackish water. The animal lives in a tube constructed of mucus and sediments. The prostomium is slightly incised at the anterior margin, and bears four eyes arranged in a trapezoid. The anterior end is narrowed, and widens posteriorly, as a somewhat bulbous caruncle, extending to the 2nd or 3rd chaetiger. There is no occipital tentacle. A pair of long palps, which can extend back to the 10th or 20th chaetiger, extends from the peristomium. Chaetiger 1 is reduced, and has small notopodial lobes lacking chaetae, and only a small cluster of chaetae on the neuropodium. Chaetigers 2, 3, 4, 6, and succeeding chaetigers, have spreading bundles of chaetae in the notopodia and neuropodia. Posterior chaetigers have stouter chaetae, and a specialized recurved hook in the notopodia, while in the neuropodia, beginning in chaetiger 7, the chaetae are largely replaced by hooded hooks. Chaetiger 5 is greatly modified, twice as large as anterior or posterior chaetigers, with a pair of rows of thick, short spines, and a pair of small bundles of chaetae. Branchiae (gills) occur on segments 2, 3, 7 and the succeeding chaetigers, but they are lacking on chaetigers 4, 5, and 6 for about one-third of the length of the worm. The parapodia in posterior segments are laterally elevated, making a dorsal channel down the middle of the body. The recurved hooks on the notopodia project into this channel. The pygidium is a flattened plate with two prominent dorsal cirri. Boccardiella ligerica reaches 18 mm (Rullier 1960; Blake and Woodwick 1971; Light 1977; Light 1978; Blake 1983; Wern 1985; Blake and Ruff 2007).


Taxonomic Tree

Kingdom:   Animalia
Phylum:   Annelida
Class:   Polychaeta
Subclass:   Palpata
Order:   Canalipalpata
Suborder:   Spionida
Family:   Spionidae
Genus:   Boccardiella
Species:   ligerica


Boccardia ligerica (Ferroniere, 1898)
Boccardia ligerica? (Day, 1955)
Boccardia redeki (Horst, 1920)
Boccardia uncata (Monro, 1938)
Polydora redeki (Horst, 1920)

Potentially Misidentified Species

Boccardia proboscidea
Boccardia proboscidea Hartman 1940 is a Northeast Pacific native which inhabits rockpools, mudflats, and crevices (Blake and Ruff 2007). It is a shell-borer which has been introduced to Australia, Hawaii, South Africa, and possibly Maine (Blake and Kudenov 1978; Simon 2010).

Boccardiella hamata
Boccardiella hamata is a shell-boring spionid polychaete, described from Virginia (Webster 1879), and later reported from Rhode Island to Texas on the East and Gulf Coasts, British Columbia to Mexico on the West Coast (Dean and Blake 1966), and Pacific coasts of Hokkaido, Japan, and Russia (Radashevsky 1993).



Boccardiella ligerica is a deposit-feeding, infaunal polychaete, characteristic of brackish water. The sexes are separate. In June, in Canal de Caen, France, males were swollen with sperm, while the body cavities of the females were filled with eggs. Eggs are laid in the female's tube and hatch as ciliated, swimming morula larvae. By the 3-chaetiger stage, larvae have long chaetae, and enter the plankton. Larvae as long as 19 chaetigers were found in the plankton, but although the larvae are clearly planktotrophic, the duration of development is not known (Rullier 1960).

Boccardiella ligerica is characteristic of brackish habitats. In Texas, the worms occurred on soft mud and hard clay (Wern 1985), while Kravitz (1987) found them on fine and coarse sand bottoms in Florida. Rullier (1960) found them in empty shells of Zebra Mussels (Dreissena polymorpha) and the empty tubes of Ficopomatus enigmaticus in a canal at Caen, France. Boccardiella ligerica has also been found in more structured habitats including sunken timbers (Eliasen and Haatella 1969) and heavily fouled ships in Suisun Bay, California (Llanso et al. 2011). This worm has been reported from freshwater (possibly slightly saline) in a desert river in Imperial County, California (Kudenov 1983) and salinities as low at 0.5-0.6 PSU (Wolff 1973; Kravitz 1987). It has also been collected at 30 PSU, but appears to be rare above 18 PSU (Wolff 1973; Wern 1985; Kravitz 1987; Peterson and Vayssieres 2010).

Boccardiella ligerica has been reported to feed on phytoplankton and microzooplankton (Hempel 1957, cited by Wolff 1973). It also probably feeds on particles on the sediment surface, as do some related species of Polydora. Particles are trapped on the mucous covered, ciliated palps, and transported to the mouth (Fauchauld and Jumars 1979). Given its abundance in some estuarine habitats (Wolf 1973; Cohen and Carlton 1995; Peterson and Vayssieres 2010), this species may play an important role in estuarine food webs, as a consumer of phytoplankton and detritus, and as food for small fishes and predatory invertebrates in low salinity regions.


Detritus, phytoplankton, benthic microalgae

Trophic Status:

Deposit Feeder



General HabitatUnstructured BottomNone
General HabitatSalt-brackish marshNone
General HabitatCanalsNone
General HabitatCoarse Woody DebrisNone
General HabitatVessel HullNone
Salinity RangeLimnetic0-0.5 PSU
Salinity RangeOligohaline0.5-5 PSU
Salinity RangeMesohaline5-18 PSU
Salinity RangePolyhaline18-30 PSU
Tidal RangeSubtidalNone
Tidal RangeLow IntertidalNone
Vertical HabitatEndobenthicNone
Vertical HabitatEpibenthicNone

Tolerances and Life History Parameters

Minimum Temperature (ºC)0Assumed winter temperature, Finland (Eliason and Haahtela 1969).
Maximum Temperature (ºC)31Wern 1985
Minimum Salinity (‰)0.6Field, Kravitz 1987.
Maximum Salinity (‰)30Kravitz 1983
Maximum Length (mm)18Blake and Woodwick 1971
Broad Temperature RangeNoneCold temperate-Warm temperate
Broad Salinity RangeNoneLimnetic-Euhaline

General Impacts

No impacts have been reported for Boccardiella ligerica. It has developed very high population densities in low-salinity regions where native spionids are rare, such as low-salinity parts of the San Francisco estuary (Peterson and Vayssieres 2010).

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
NEA-IV None 0 Native Estab
NEA-II None 0 Native Estab
NEA-V None 0 Native Estab
B-X None 1960 Def Estab
NA-ET3 Cape Cod to Cape Hatteras 1990 Def Estab
CAR-IV None 1978 Def Estab
CAR-I Northern Yucatan, Gulf of Mexico, Florida Straits, to Middle Eastern Florida 1973 Def Estab
SA-II None 1938 Def Estab
NEP-VI Pt. Conception to Southern Baja California 1935 Def Unk
NEP-V Northern California to Mid Channel Islands 1954 Def Estab
B-IX None 1972 Def Estab
M130 Chesapeake Bay 1990 Def Estab
M080 New Jersey Inland Bays 1998 Def Estab
P040 Newport Bay 1935 Def Failed
P090 San Francisco Bay 1954 Def Estab
CAR-VII Cape Hatteras to Mid-East Florida 1999 Def Estab
G290 San Antonio Bay 1973 Def Estab
G260 Galveston Bay 1978 Def Estab
G250 Sabine Lake 1985 Def Estab
S050 Cape Fear River 1999 Def Estab
G074 _CDA_G074 (Crystal-Pithlachascotee) 1984 Def Estab
M060 Hudson River/Raritan Bay 2004 Def Estab
P093 _CDA_P093 (San Pablo Bay) 1954 Def Estab
P095 _CDA_P095 (Tomales-Drakes Bay) 1968 Def Unk
S180 St. Johns River 1998 Def Estab
M090 Delaware Bay 2003 Def Estab
B-III None 0 Crypto Estab
NEA-III None 1970 Crypto Estab
SA-I None 1997 Def Estab
CAR-III None 0 Def Estab
WA-IV None 1955 Crypto Estab

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude


Anderson, Mark G.; Smith, Joseph A. M.; Wilson, Bartholomew D. (2011) Delaware River basin priority conservation areas and recommended conservation strategies, The Nature Conservancy, Partnership for the Delaware Estuary and Natural Lands Trust, <missing place>. Pp. not numbered

Angradi, T.R.; Hagan, S. M.; Able, K. W. (2001) Vegetation type and the intertidal macroinvertebrate fauna of the brackish marsh: Phragmites vs. Spartina., Wetlands 21(1): 75-92

Barry A. Vittor & Associates, Inc. (1999) <missing title>, U.S. Department of Commerce, National Oceanic and Atmospheric Administration, Mobile AL. Pp. <missing location>

Blake, James A. (1983) Polychaetes of the family Spionidae from South America, Antarctica, and adjacent seas and islands, Biology of the Antarctic Seas 39(3): 205-288

Blake, James A.; Kudenov, Jerry D. (1978) Spionidae (Polychaeta) from southeastern Australia and adjacent areas with a revision of the genera, Memoirs of the National Museum in Victoria 39: 171-280

Blake, James A.; Ruff, R. Eugene (2007) The Light and Smith Manual: Intertidal invertebrates from Central California to Oregon (4th edition), University of California, Berkeley CA. Pp. 309-410

Blake, James A.; Woodwick, Keith H. (1971) A review of the genus Boccardia Carazzi (Polychaeta: Spionidae) with descriptions of two new species, Bulletin of the Southern California Academy of Sciences 70(1): 31-42

California Academy of Sciences 2005-2015 Invertebrate Zoology Collection Database. <missing URL>

Carlton, James T. (1979) History, biogeography, and ecology of the introduced marine and estuarine invertebrates of the Pacific Coast of North America., Ph.D. dissertation, University of California, Davis. Pp. 1-904

Carlton, James T. 1998 <i>Boccardiella ligerica</i>. <missing URL>

Chainho, P. and 5 authors (2006) Seasonal and spatial patterns of distribution of subtidal benthic invertebrate communities in the Mondego River, Portugal – a poikilohaline estuary, Hydrobiologia 555: 59-74

Chesapeake Bay Program 1998-2001 Benthic Database.. <missing URL>

Cohen, Andrew N.; Carlton, James T. (1995) Nonindigenous aquatic species in a United States estuary: a case study of the biological invasions of the San Francisco Bay and Delta, U.S. Fish and Wildlife Service and National Sea Grant College Program (Connecticut Sea Grant), Washington DC, Silver Spring MD.. Pp. <missing location>

Cunha, M. R.; Moreira, M. H. (1995) Macrobenthos of Potamogeton and Myriophyllum beds in the upper reaches of Canal de Mira (Ria de Aveiro, NW Portugal): community structure and environmental factors, Netherlands Journal of Aquatic Biology 29(3-4): 377-390

Dauer, Daniel; Lane, Michael; Llanso, Robert J. (2002) <missing title>, Chesapeake Bay Program, Annapolis MD. Pp. 1-65

Day, J. H. (1955) The Polychaeta of South Africa. Part e: Sedentary species of Cape shores and estuaries, Journal of the Linnean Society of London- Zoology 42: 407-452

Dean, David; Blake, James A. (1966) Life-history of Boccardia hamata (Webster) on the East and West coasts of North America, Biological Bulletin 130(3): 316-330

Eliason, A.; Haahtela, I. (1969) Polydora (Boccardia) redeki Horst from Finland., Annales Zoologici Fennici 6: 215-218

Fauchald, Kristian; Jumars, Peter A. (1979) The diet of worms : A study of polychaete feeding guilds, Oceanography and Marine Biology, an Annual Review 17: 193-284

Gappa, J. Lopez ; Tablado, A.; Fonalleras, M. C.; Adami, M. L. (2001) Temporal and spatial patterns of annelid populations in intertidal sediments of the Quequen Grande estuary (Argentina), Hydrobiologia 455: 61-69

Gillett, David J.; Schaffner, Linda C. (2009) Benthos of the York River, Journal of Coastal Research Special Issue(57): 80-98

Gollasch, Stephan; Rosenthal, Harald (2005) Bridging divides: Maritime canals as invasion corridors, Springer, Dordrecht, Netherlands. Pp. 5-90

Jensen, Kathe R. 2015 NOBANIS – Invasive Alien Species Fact Sheet – Boccardiella ligerica – From: Identification key to marine invasive species in Nordic waters:NOBANIS,11/15/2014. <missing URL>

Kravitz, Michael J. (1987) First record of Boccardiella ligerica (Ferronniere) (Polychaeta: Spionidae) from the east coast of North America, Northeast Gulf Science 9(1): 39-42

Kudenov, Jerry D. (1983) First record of Boccardia ligerica (Polychaeta: Spionidae) from Imperial County, California., Bulletin of the Southern California Academy of Sciences 82(3): 144-148

Leppakoski, Erkki; Olenin, Sergei (2000) Non-native species and rates of spread: lessons from the brackish Baltic Sea., Biological Invasions 2: 151-163

Light, William J. (1977) Spionidae (Annelida: Polychaeta) from San Francisco Bay, California: a revised list with nomenclatural changes, new records and comments on related species from the Northeastern Pacific Ocean., Proceedings of the Biological Society of Washington 90(1): 66-88

Light, William J. (1978) <missing title>, Boswood Press, Pacific Grove CA. Pp. <missing location>

Llansó, Roberto J.; Sillett, Kristine; Scott, Lisa (2011) <missing title>, Versar, Inc., Columbia MD. Pp. <missing location>

Mattson, Robert A. (2011) <missing title>, St. Johns River Water Management District, Palatka FL. Pp. unpaged

Meadowlands Environmental Research Institute (2004) <missing title>, ENSR Inc., <missing place>. Pp. unpages

New York State Department of Transportation (2012) <missing title>, New York State Department of Transportation, Albany NY. Pp. F1-F87

New York State Dept. of Environmental Conservation 2000 New York State Amphibian and Reptile Atlas Project. <missing URL>

Olenin, Sergej, Leppakoski, Erkki (1999) Non-native animals in the Baltic Sea: alteration of benthic habitats in coastal inlets and lagoons., Hydrobiologia 393: 233-243

Orensanz, Jose Maria and 14 other authors (2002) No longer the pristine confines of the world ocean: a survey of exotic marine species in the southwestern Atlantic, Biological Invasions 4(1-2): 115-143

Paavolo, Marjo; Laine, Ari O.; Helavuori, Markius; Kraufvelin, Patrik (2008) Profiling four brackish water harbours: zoobenthic composition and invasion status., Boreal Environment Research 13: 159-175

Peterson, Heather A.; Vayssieres, Marc (2010) Benthic assemblage variability in the upper San Francisco estuary: A 27-year retrospective, San Francisco Estuary and Watershed Science <missing volume>: published online

Radashevsky, Vasily I. (1993) Revision of the genus Polydora and related genera from the North West Pacific (Polychaeta: Spionidae), Publications of the Seto Marine Biological Laboratory 36(1/2): 1-60

Radashevsky, Vasily I. (2012b) Spionidae (Annelida) from shallow waters around the British Islands: an identification guide for the NMBAQC Scheme with an overview of spionid morphology and biology, Zootaxa 3152: 1-35

Rullier, Francois (1960) Morphologie et developpement du Spionidae (annelide polychete) Polydora (Boccardia) redeki Horst., Cahiers de Biologie Marine 1: 231-244

Siegfried, Clifford A.; Kopache, Mark E.; Knight, Allen W. (1980) The benthos of a portion of the Sacramento River (San Francisco Bay estuary) during a dry year, Estuaries 3(4): 296-307

Simon, C. A.; Worsfold, T. M.; Lange, L.; Sterley, J (2010) The genus 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(3): 585–598

Simon, Carol A. 2015 A checklist of the Spionidae (Annelida: Polychaeta) of Southern Africa, including South Africa, Namibia, Mozambique and Madagascar.. <missing URL>

U.S. National Museum of Natural History 2002-2021 Invertebrate Zoology Collections Database.

Valanko, Sebastian; Norkko, Alf; Norkko, Joanna (2010) Strategies of post-larval dispersal in non-tidal soft sediment communities, Journal of Experimental Marine Biology and Ecology 384: 51-60

Webster, H. E. (1879) Annelida chaetopoda of the Virginia coast, Transactions of the Albany Institution 9: 202-269

Wern, Judy O. (1985) First record of the spionid polychaete Boccardiella ligerica (Ferronniere, 1898) from the Gulf of Mexico, Contributions in Marine Science 28: 125-128

Wijnhoven, Sander; Hummel, Herman (2011) Patterns in macrozoobenthic assemblages indicate the state of the environment: insights from the Rhine-Meuse estuary, Marine Ecology Progress Series 436: 29-50

Wolff, W. J. (1973) The estuary as a habitat: An analysis of data on the soft-bottom macrofauna of the estuarine area of the rivers Rhine, Meuse, and Scheldt, Zoologische Verhandelingen 126: 4-242

Wong, Eunice; Kupriyanova, Elena K.; Hutchings, Pat; Capa, María; Radashevsky, Vasily; ten Hove, Harry A. (2014) A graphically illustrated glossary of polychaete terminology: invasive species of Sabellidae, Serpulidae and Spionidae, Memoirs of Museum of Victoria 71: 327-342