Invasion History

First Non-native North American Tidal Record: 1951
First Non-native West Coast Tidal Record: 1951
First Non-native East/Gulf Coast Tidal Record:

General Invasion History:

Pseudopolydora cf. kempi was first described from a canal near Kolkata (formerly Calcutta), India (Southern 1921). Populations have been found in estuaries in Thailand (Angsupanich et al. 2005); Hong Kong (Lu and Wu 2007); Taiwan (Radashevsky and Kemp 2000); Japan (Imajima and Hartman 1964; Sato-Okoshi 2000); and Peter the Great Bay, Russia (Omelyanenko and Kulikova 2011). It is also listed as occurring in the Red Sea and Persian (Arabian) Gulf (Wehe and Fiege 2002). Populations on the West Coast of North America occur from British Columbia to southern California, and are presumed to be introduced (Carlton 1979; Cohen and Carlton 1995; Light 1977; Cohen et al. 2001). However, California populations (P. kempi californica, Light 1969) show morphological and developmental differences from populations in India and the Sea of Japan (P. kempi japonica Imajima and Hartman1964; Blake and Ruff 2007). Each of these forms are considered separate subspecies. The absence of Pseudopolydora cf. kempi in early polychaete surveys strongly supports introduced status for West Coast populations (Carlton 1979; Cohen and Carlton 1995). A likely source of these populations is northeastern Japan, from where the prevailing Miyagi strain of Pacific Oysters (Crassostrea gigas) was imported (Carlton 1979, James Carlton, personal communication 2015). Morphology and development of P. cf. kempi from this particular region has not been studied. Polydora cf. kempi have been reported from scattered estuarine locations around the world, including Venezuela (Chollet and Bone 2007), Atlantic Spain (Lopez-Jamar et al. 1986), South Africa (Day 1955, cited by Simon 2015), Australia (Blake and Woodwick 1978) and New Zealand (Inglis et al. 2006e). However, most of these records do not provide morphological details. More detailed morphological, developmental, and molecular data will be needed to clarify the identity and invasion history of this species complex.

North American Invasion History:

Invasion History on the West Coast:

The earliest record of Pseudopolydora cf. kempi is from 1951 at Rathtrevor Beach, Parksville, British Columbia, on the east coast of Vancouver Island, misidentified as Neopygospio laminifera (Berkeley and Berkeley 1954, cited by Carlton 1979). Early records of this polychaete are spotty, suggesting several scattered introductions with Pacific Oysters (Crassostrea gigas), e.g. San Juan Islands, Washington (WA) in 1968 (Banse 1972); Puget Sound, WA in 1979 (Gallagher et al. 1983); Yaquina Bay, Oregon (OR) (Walker 1974, cited by Carlton 1979; Hancock et al. 1977, cited by Wonham and Carlton 2005); Coos Bay, OR; Bolinas Lagoon, California (CA) in 1967 (Light 1969); Bodega Bay and Tomales Bay, CA in 1971-1972 (Blake and Woodwick 1975); and Morro Bay, CA in 1960 (Blake 1960, cited by Carlton 1979; 1960, Reish and Barnard 1967). Pseudopolydora cf. kempi was first found in San Francisco Bay in 1972-1973, and was already widespread in the Central and South Bays, San Pablo Bay, and upstream to Grizzly Bay (Chapman and Dorman 1975; Light 1977; Peterson and Vayssieres 2010; California Academy of Sciences 2015), where ballast water and oyster plantings are both possible vectors. We have a few records of this polychaete from fouling plates (Ruiz et al. unpublished data), so transport in ship fouling is also possible. The only southern California record of which we are aware is from Anaheim Bay, CA in 1970-1972 (Reish et al. 1975), where there are no documented oyster transplants or ballast water discharge. Later discoveries of this polychaete include Willapa Bay, WA (2000, Cohen et al. 2001), Columbia River estuary OR-WA (1980, Jones 1983), Tillamook Bay, OR (Golden et al. 1998, cited by Cohen 2004), and Humboldt Bay, CA (Barnhart et al.1992, cited by Boyd et al. 2002), which either have oyster culture operations, ballast water discharge, or both.

Invasion History Elsewhere in the World:

Pseudopolydora cf. kempi has been reported from many other locations around the world. In Australia, this polychaete was reported from Brisbane, in southern Queensland, and from Merrimbula and Careel Bay, New South Wales (Blake and Woodwick 1975; Hutchings and Rainer 1979). Day (1955) reported it from Mozambique, although details of their morphological descriptions differ somewhat. In a series of surveys of New Zealand harbors, P. cf. kempi was found at one location, in Whangarei Harbour, on the North Island (Inglis et al. 2006e). Other reports are from the Caribbean coast of Venezuela (Chollet and Bone 2007) and from La Coruna Harbor, Spain (Lopez-Jamar et al. 1986). We have treated these records as introductions on our bioregion maps, provisionally, but detailed molecular and morphological studies will be needed to determine the identity of these spionids.


Description

Pseudopolydora cf. kempi has been subdivided into several subspecies, which show differences in adult morphology and larval development. The status of these subspecies is unresolved, due to scanty descriptions and the absence of type specimens. With future work, this taxon may be split into several cryptic species (Radashevsky and Hsieh 2000; Sato-Okoshi 2000). Here, we will treat it as a single taxon, Pseudopolydora cf. kempi.

Adults of Pseudopolydora cf. kempi have up to 38 (California) or 45-53 (Japan, Taiwan, and Australia) chaetigers. The prostomium is incised or bilobed and narrows to a caruncle extending back to chaetigers 3-4. There are usually 4 (but sometimes 6) eyes on the prostomium, and usually an occipital tentacle (but missing in Light (1969) specimen). The palps are transparent, and extend backwards for 10-20 segments. Chaetiger 1 has short notopodial lobes and neuropodial lamellae, but lacks notochaetae. This segment does have neurochaetae. Chaetiger 2 and the succeeding segments have rounded notopodal lobes, with 2-3 transverse rows of thin capillary notochaetae, with shorter, thicker chaetae in the first row. The neuropodia of chaetigers 2-7 have dense bundles of chaetae. Chaetiger 5 is only slightly modified, with a J-shaped row of spines between the notopodial and neuropodial chaetae (a genus character). Branchiae start on the 7th chaetiger, and continue for about half the body length. Beginning at chaetiger 8, the neuropodia have varying numbers of hooded hooks (7-18, California, Light 1969; up to 22, Taiwan, Radashevsky and Hsieh 2000; 25-30, Australia, Blake and Woodwick 1978) double-toothed hooded hooks, without accompanying capillary chaetae. The branchiae decrease in size toward the posterior. The pygidium is a round, flaring disc, with a notch on the ventral side, flanked by two finger-like processes on each side. The reported maximum length ranges from 6.5 (California, Light 1969) to 22 mm (Japan, Sato-Okoshi 2000). The worms are white to tan in color, with black pigments in the space between the chaetae of the first 5-6 chaetigers, and sometimes with a pair of dorsal spots on the chaetigers. Description based on: Southern 1921, Imajima and Hartman 1964, Light 1969, Light 1978, Blake and Kudenov 1978, Hutchings and Rainer 1979, Radashevsky and Hsieh 2000, and Sato-Okoshi 2000.

Populations of Pseudopolydora 'kempi' from different localities show small differences in morphology. Populations in Japan and California have been designated as separate subspecies) P. kempi japonica (Imajima and Hartman 1964) and P. kempi californica (Light 1969), respectively, but authors diverge on the significance of these differences (Sato-Okoshi 2000; Radashevsky and Hsieh 2000). Pseudopolydora cf. kempi from India, the Sea of Japan, and California differ in the number and size of nurse eggs providing food for developing embryos, and in the length of the planktonic larval stage (Blake and Woodwick 1975; Myohara 1979; Rdashevsky 1985; Blake and Ruff 2007). The morphology and larval development of P. cf. kempi from the Pacific coast of Japan, the source of most oyster imports and much shipping, and a likely source of North American populations, has not been studied (James Carlton, personal communication).


Taxonomy

Taxonomic Tree

Kingdom:   Animalia
Phylum:   Annelida
Class:   Polychaeta
Subclass:   Palpata
Order:   Canalipalpata
Suborder:   Spionida
Family:   Spionidae
Genus:   Pseudopolydora
Species:   cf. kempi

Synonyms

Neopygospio laminifera (Berkeley & Berkeley, 1954)
Pseudopolydora kempi californica (Light, 1969)
Pseudopolydora kempi japonica (Imajima & Hartman, 1964)
Pseudopolydora kempi kempi (Southern, 1921)

Potentially Misidentified Species

Pseudopolydora bassarginensis
Pseudopolydora bassarginensis (Zachs 1933) was described from the Sea of Japan. Is distribution is not completely known, but it has been reported from Willapa Bay, Washington and Coos Bay, Oregon (Cohen et al. 2001; Chapman et al. 2011).

Pseudopolydora paucibranchiata
Pseudopolydora paucibranciata is another species complex, which shows morphological and developmental differences from P. cf. kempii and also differences in habitat, being restricted to more saline regions of estuaries, though overlapping with P. kempi in some areas (Blake and Woodwick 1975).

Ecology

General:

Pseudopolydora cf. kempi is a complex of at least two, and probably more cryptic species, with very different developmental patterns across its geographical range. The different forms are similar in occurring in estuarine habitats, constructing mud and mucus tubes, having separate sexes, and laying eggs in capsules, deposited in strings, inside the tubes. The capsules contain both developing embryos, and nurse eggs, which become fragmented as the embryos develop (Blake and Woodwick 1975; Srikrishnada and Ramamoorthi 1977; Myohara 1979). California populations lay strings of 15-20 eggs, and Myohara's diagram also shows ~15 capsules. California (Tomales and Morro Bay) populations differ greatly in developmental patterns from Pseudopolydora cf. kempi in the Sea of Japan or India. The Japanese and Indian animals hatch from the egg-capsule at the 3-chaetiger stage and spend 2-4 weeks in the plankton, settling at ~18 setigers (Myohara 1979; Srikrishnada and Ramamoorthi 1977; Radshevsky 1985). Developing California animals consume most of the neighboring eggs in the capsule, and hatch at 10-12 setigers, spending a few days in the plankton, and settling at 13-15 chaetigers. Development in California P. kempi is largely lecithotrophic (Blake and Woodwick 1975). It is possible that other West Coast populations may show development patterns described from Asian populations.

Pseudopolydora cf. kempi has been reported from brackish estuaries and coastal waters in cold-temperate to tropical waters (Berkeley and Berkeley 1951; Srikrishnada and Ramamoorthi 1977; Light 1978). In India, P. cf. kempi larvae have been collected at salinities of 1.6-34.8 PSU (Srikrishnada and Ramamoorthi 1977), while in San Francisco Bay, P. cf. kempi occurs abundantly upstream into low salinity portions of Grizzly Bay (Peterson and Vayssieres 2010). Pseudopolydora cf. kempi occurs on intertidal mudflats and soft sand or mud substrates (Blake and Woodwick 1975; Gallagher and Wells 1983). It is a deposit-feeder, using its palps to pick up particles from the sediment surface (Gallagher and Wells 1983; Hentschel 1998). It is not clear whether P. cf. kempi can switch to suspension-feeding, as some other spionids do. Pseudopolydora cf. kempi is probably an important prey for fishes and benthic invertebrates. In a Japanese estuary, sublethal predation by flounders (Platichthys bicoloratus) was common, with predators biting off parts of worms, which subsequently regenerated (Tomiyama et al. 2007).

Food:

Benthic microalgae, detritus, phytoplankton

Trophic Status:

Deposit Feeder

DepFed

Habitats

General HabitatUnstructured BottomNone
General HabitatSalt-brackish marshNone
General HabitatCanalsNone
Salinity RangeMesohaline5-18 PSU
Salinity RangePolyhaline18-30 PSU
Salinity RangeEuhaline30-40 PSU
Tidal RangeSubtidalNone
Tidal RangeLow IntertidalNone
Vertical HabitatEndobenthicNone
Vertical HabitatEpibenthicNone


Tolerances and Life History Parameters

Maximum Temperature (ºC)29Venezuela (Chollet and Bone 2007)
Minimum Salinity (‰)1.6Field observation, Vellar estuary, India (Srikrishnada and Ramamoorthi 1977)
Maximum Salinity (‰)37Venezuela (Chollet and Bone 2007)
Minimum Duration2Lecithotrophic form, California, nearly direct development, released from egg capsule at 10-13 chaetigers (Blake and Woodwick 1975)
Maximum Duration28Planktotrophic form, Japan (Myohara 1979, released from capsule at 3 chaetigers.
Minimum Length (mm)6.5Length of holotype, California (Light 1969). New adults are probably smaller,
Maximum Length (mm)22Japan, Sato-Okoshi 2000. San Francisco Bay animals averaged 12 mm (Light 1978).
Broad Temperature RangeNoneCold temperate-tropical
Broad Salinity RangeNoneMesohaline-Euhaline

General Impacts

Ecological impacts

Pseudopolydora cf. kempi is frequently abundant in subtidal brackish waters in Asian waters and the West Coast of North America. It is a potential prey item for fishes and other predators (Tomiyama et al. 2007). Together with other tube-building invertebrates, this worm has an ecological impact in mudflats and soft-substrate habitats by adding structure to relatively homogeneous environments, facilitating the recruitment of other invertebrates (Gallagher et al. 1983).

Regional Impacts

NEP-IIIAlaskan panhandle to N. of Puget SoundEcological ImpactHabitat Change
The tube-building polychaetes Hobsonia florida, Pseudopolydora kempi and taniads (Tanais) sp. (Crustacea, Peracarida) facilitate the recruitment of other taxa to 10 cm-2 patches (Gallagher et al. 1983).
P290Puget SoundEcological ImpactHabitat Change
The tube-building polychaetes Hobsonia florida, Pseudopolydora kempi and taniads (Tanais) sp. (Crustacea, Peracarida) facilitate the recruitment of other taxa to 10 cm-2 patches (Gallagher et al. 1983).

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
NEP-IV Puget Sound to Northern California 1974 Def Estab
NWP-4a None 0 Native Estab
NWP-3a None 0 Native Estab
NEP-VI Pt. Conception to Southern Baja California 1972 Def Estab
NEP-III Alaskan panhandle to N. of Puget Sound 1951 Def Estab
NWP-2 None 0 Native Estab
NEP-V Northern California to Mid Channel Islands 1960 Def Estab
CIO-II None 0 Native Estab
EA-IV None 0 Crypto Estab
NWP-3b None 0 Native Estab
NWP-4b None 0 Native Estab
AUS-X None 1975 Def Estab
P170 Coos Bay 1977 Def Estab
P293 _CDA_P293 (Strait of Georgia) 2011 Def Estab
EAS-I None 0 Native Estab
P210 Yaquina Bay 1974 Def Estab
P270 Willapa Bay 2000 Def Estab
P230 Netarts Bay 1976 Def Estab
P290 Puget Sound 1979 Def Estab
P292 _CDA_P292 (San Juan Islands) 1968 Def Estab
P110 Tomales Bay 1972 Def Estab
P070 Morro Bay 1960 Def Estab
P095 _CDA_P095 (Tomales-Drakes Bay) 1967 Def Estab
P260 Columbia River 1980 Def Estab
P240 Tillamook Bay 0 Def Estab
P130 Humboldt Bay 1992 Def Estab
P112 _CDA_P112 (Bodega Bay) 1971 Def Estab
P050 San Pedro Bay 1972 Def Estab
NZ-IV None 0 Def Estab
NEA-V None 0 Def Estab
CAR-III None 2001 Def Estab
AUS-XII None 1972 Def Estab
NWP-5 None 0 Native Estab
P090 San Francisco Bay 1972 Def Estab
RS-3 None 0 Native Estab
RS-2 None 0 Native Estab
RS-1 None 0 Native Estab
AG-1 None 0 Native Estab
AG-5 None 0 Native Estab
AG-4 None 0 Native Estab
AG-3 None 0 Native Estab

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude
768153 Ruiz et al., 2015 2012 2012-09-06 Loch Lomond Marina, San Francisco Bay, CA, California, USA Def 37.9736 -122.4802

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