Pseudopolydora paucibranchiata

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: 2019

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General Invasion History:

Pseudopolydora paucibranchiata was described from Onomichi, Japan, on the Seto Inland Sea (Okuda 1937, cited by Radashevsky 1993). It occurs in mudflats and shallow subtidal waters from Hong Kong (Wu and Lu 2007) to southern Sakhalin Island and the southern Kuril Islands, Russia (Radashevsky 1993). Populations in the Northeast Pacific (Baja California to Puget Sound), Southwest Pacific (Australia, New Zealand), and Northeast Atlantic (Norway, Portugal, Atlantic Spain, and Greece-Turkey) are generally believed to be introduced (Read 1975; Blake and Kudenov 1978; Carlton 1979; Ramberg and Schram 1982; Cohen and Carlton 1995; Simboura et al. 2008; Cacabelos et al. 2008; Rodrigues et al. 2011; Cinar et al. 2012). Likely vectors include ballast water, possibly hull fouling, and transplants of Pacific Oysters Crassostrea gigas from Japan. Most of these oysters were the Miyagi strain, from northeastern Honshu, but the morphology of P. paucibranchiata from this region has have not been studied (James Carlton, personal communication 2014). Molecular studies of P. paucibranchiata throughout its range will be useful for understanding its taxonomy, biogeography, and invasion history.

North American Invasion History:

Invasion History on the West Coast:

Pseudopolydora paucibranchiata was first reported from North American waters in Los Angeles-Long Beach Harbor, California (CA) in 1951 and was identified as 'Polydora (?) n. sp., by Olga Hartman, for the Los Angeles Regional Water Pollution Control Board (Blake and Woodwick 1975; Carlton 1979). It was also collected in 1951 in Newport Bay, and then in San Diego Bay in 1952 (Allen Hancock Foundation, cited by Carlton 1979). In the 1970s, P. paucibranchiata was found in Tomales Bay and Elkhorn Slough (1st record 1972), San Francisco Bay (1st record 1973), and Morro Bay and Santa Barbara Harbor (by 1975) (Blake and Woodwick 1975; Light 1977; Carlton 1979; California Academy of Sciences 2015). In San Francisco Bay, it is found in the Central, South, and San Pablo Bays, and ranges inland to the western end of Carquinez Strait (Cohen and Carlton 1995). It was found in Netarts Bay, Oregon (OR) (Stout 1976, cited by Carlton 1979), and later was collected in Humboldt Bay, CA (1st record 2000, Boyd et al. 2002); Coos Bay, OR (1st record 2010, Chapman et al. 2011); and Willapa Bay, Washington (1st record 1996, Cohen et al. 2002; Ferraro and Cole 2007). It has been reported as occurring in Puget Sound (Blake and Ruff 2007; Washington Department of Ecology 2014), but we have no information on specific locations.

In southern California, it has colonized most of the smaller bays and harbors from Santa Barbara to San Diego (Talley et al. 2000; Cohen et al. 2002; Fairey et al. 2002; Ranasinghe et al. 2005). In many of the intertidal mudflats and shallow, muddy bottoms where it was collected, it was an abundant or dominant polychaete (Blake and Woodwick 1975; Levin 1984; Ranasinghe et al. 2005). This polychaete has long-lived planktonic larvae, so ballast water is a likely vector for early occurrences in San Diego Bay and Los Angeles Harbor, where oyster plantings were limited. From Morro Bay northward, transplants of Pacific Oysters are a more possible vector. This polychaete is mostly known from tidal mudflats, but Barnard reported it from harbor pilings in Los Angeles-Long Beach Harbor (Barnard 1958). It seems that hull fouling of small boats, is a likely vector in the smaller harbors of southern California.

Invasion History Elsewhere in the World:

Globally, the distribution of this species complex may be underestimated due to misidentification. In Europe, Pseudopolydora paucibranchiata was first collected in the inner Oslofjord, Norway in 1962 (Ramberg and Schram 1982; Olsgard 1999). Holte (1998) reported it from the Holandsfjord (66°N), a 'pristine' fjord. Radashevsky (2012) lists it as a species occurring in the waters around Great Britain, and the United Kingdom's National Biodiversity Network lists 19 records on the west and east coasts of Scotland (NBN Gateway 2015). The fjord and offshore records seem inconsistent with this worm's preference for shallow, comparatively warm waters (Radashevsky 1993; Blake and Woodwick 1975). Records from estuaries in northeast Spain (Ria de Vigo in 1999, Cacabelos et al. 2008); Portugal (Ria de Aveiro, Rodrigues et al. 2011), and the Mediterranean, are less surprising. In the Mediterranean, P. paucibranchiata has been found on the Greek (in 2005, Simboura et al. 2008) and Turkish (in 2009, Cinar et al. 2012) sides of the Aegean Sea; the Sea of Marmara (in 2009, Cinar et al. 2012); and in the Venice Lagoon (since 2002, Erica Keppel, personal communication; Maggiore and Keppel 2007).

The earliest record of Pseudopolydora paucibranchiata from the Southwest Pacific was by Read (1975) from Wellington Harbour, South Island, New Zealand in 1971. Inglis et al. (2006e) reported this worm from Whangarei Harbour, on the North Island. In Australia, P. paucibranchiata was first reported from Botany Bay, New South Wales in 1973 (Blake and Kudenov 1978). It was subsequently found in Port Phillip and Westernport Bays, Victoria (1st record 1975, Blake and Kudenov 1978); South Australia (1st record 1979, Hutchings and Turvey 1984), and Queensland (Saenger 1988, cited by Walker 2009; Atlas of Living Australia 2015).

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Description

Adults of Pseudopolydora paucibranchiata have 45-70 segments. The prostomium has a rounded anterior edge and extends back to chaetiger 3 as a caruncle. The prostomium bears four black eyes in a trapezoidal arrangement and an occipital tentacle is present. Some individuals have one ot wo lateral eyes. The palps are transparent and extend backwards for 20-40 chaetigers, about half of the body length. Chaetiger 1 has small notopodial lobes near the midline, but a bundle of notochaetae is absent, while the neuropodial lobes, with a small bundle of capillary setae, lie adjacent to the palps. Low parapodial lobes are present on chaetigers 2-4, but the lobes are rudimentary on chaetiger 5. This segment is otherwise unmodified, having the same size as adjacent segments. However, the notochaetae and neurochaetae of this segment are modified. An outer row of 12-14 curved major spines, with enlarged and bent tips is arranged in a J-shaped line, which is characteristic of the genus. They are accompanied by an equal number of simple spines, with unusual constrictions near their tips, forming an inner row. From chaetiger 7 rearward, to aboutchaetiger 28, , the segments bears up to 22 dorsal branchiae. From chaetiger 8 rearward, the neuropodial chaetae are hooded hooks, with up to 20 per bundle. The pygidium bears a small, ventral, cup-like disk, with a dorsal gap. The maximum reported length is 18 mm, and 0.5 mm wide (Russia, Radashevsky 1993; Radashevsky et al. 2020), but West Coast specimens are usually about 4-6 mm (Blake and Ruff 2007). Younger worms may retain larval melanophores on their dorsal surface, but these are absent in larger worms. Living worms have branching yellow chromatophores on the palps, but these disappear in preserved specimens. Larval development has been described for populations in the Sea of Japan (Myohara 1980) and California (Blake and Woodwick 1975). Description based on: Read 1975, Blake and Woodwick 1975, Blake and Kudenov 1978, Light 1978, Radashevsky 1993, Wilson 1999, Radashevsky and Hsieh 2000, Sato-Okoshi 2000, and Blake and Ruff 2007; Radashevsky et al. 2020).

The genus Pseudopolydora appears to be a complex of 'pseudocryptic' speciies with relatively small morphological differences. Occurences of 'P. paucibranchiata' from Norwegian fjords, deep waters, and sites remote from known vectors, appear to refer to cryptic native species (Radashevsky et al. 2021b).

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Taxonomy

Ecology

General:

Pseudopolydora paucibranchiata is an estuarine spionid polychaete, which dwells in mud and mucus tubes in sediment, and in fouling communities on hard surfaces. The sexes are separate. Males and females mature at about 30-35 mm (Radashevsky et al. 2020). Eggs are laid in capsules, linked in strings attached to the interior of the tubes. Capsules in California and Japanese estuaries contained up to 35-50 eggs, with 7-10 capsules on a string. Larvae are released from the capsules at the 3-chaetiger stage, and feed in the plankton (Blake and Woodwick 1975; Myohara 1980). California larvae settle and metamorphose at about 13-17 chaetigers (Blake and Woodwick 1975), while Japanese larvae were reported to settle at 23-33 chaetigers, and a length of 1.5-2.8 mm, about 2 weeks after hatching at 23C (Myohara 1980). Blake and Woodwick (1975) do not give the development time, but indicate that the pattern of development was planktotrophic and longer than the short lecithotrophic larval period which they observed in P. kempi. The Japanese larvae matured 24-31 days after hatching, at 5-10 mm length and 45-50 chaetigers (Myohara 1980). Pseudopolydora paucibranchiata reaches a maximum size of 18 mm (Radashevsky 1993).

Pseudopolydora paucibranchiata is known from cold-temperate to subtropical estuarine-marine habitats from the tropics to cold-temperate waters. Its habitats include subtidal and intertidal soft substrates of sand or mud, where it oncstructs tubes up to 45 mm long, extending up to 10 mm above the surface (Radshevsky et al. 2020).. It is also known from salt marshes, and fouling communities in marinas and harbors (Barnard 1958; Blake and Woodwick 1975; Myohara 1980; Ferraro and Cole 2007). This polychaete does not seem to extend very far into brackish or hypersaline conditions, compared to some other spionids. We found reports over a range of about 15-37 PSU (Ferraro and Cole 2007; Simboura et al. 2010). It was found, with P. cornuta, in the polluted portions of Hong Kong (Lu and Wu 2007) and Izmir Bay, Turkey (Dagli et al. 2011; Cinar et al. 2012), and in harbors contaminated with copper (Neira et al. 2013). However, in the Oslofjord, Norway, abundance of P. paucibranchiata was negatively correlated with copper content (Olsgard 1999). Adults of P. paucibranchiata are capable of shifting between deposit-feeding detritus and surface-microalgae to suspension-feeding (phytoplankton and suspended particles). Their ciliated palps can either stretch over the sediment surface to pick up sediment particles, or wave in the current to pick up phytoplankton (Shimeta 1996). Pseudopolydora paucibranchiata can reach high abundances, and is probably important prey for fishes and benthic invertebrates.

Food:

Phytoplankton, benthic microalgae, detritus

Consumers:

Fish, crabs, shrimps

Trophic Status:

Deposit Suspension Feeder

DepSusFed

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Habitats

General Habitat	Unstructured Bottom	None
General Habitat	Salt-brackish marsh	None
General Habitat	Marinas & Docks	None
Salinity Range	Polyhaline	18-30 PSU
Salinity Range	Euhaline	30-40 PSU
Tidal Range	Subtidal	None
Tidal Range	Low Intertidal	None
Vertical Habitat	Endobenthic	None
Vertical Habitat	Epibenthic	None

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Tolerances and Life History Parameters

Minimum Salinity (‰)	15	Field, minimum salinity for Willapa Bay sediment (Ferraro and Cole 1996)
Maximum Salinity (‰)	37	Typical Aegean Sea Salinity
Minimum Reproductive Temperature	11	Start of spawning, Peter the Great Bay (Radashevsky et al. 2020).
Minimum Duration	10	Laboratory, Japan, 23 C (Myohara 1980)
Maximum Duration	24	Laboratory, Japan, 23 C (Myohara 1980)
Minimum Length (mm)	5	Minimum size at maturity (Blake and Woodiwick 1975; Myohara 1980)
Maximum Length (mm)	18	Russia (Radashevsky 1993; Radashevsky et al. 2020)
Broad Temperature Range	None	Cold temperate-Subtropical
Broad Salinity Range	None	Polyhaline-Euhaline

General Impacts

Ecological Impacts

Pseudopolydora paucibranchiata can be abundant in soft sediments and fouling communities, and has been found in some polluted habitats in San Diego Bay (California), Izmire Bay (Turkey) and Hong Kong (Lu and Wu 2007; Dagli et al. 2011; Cinar et al. 2012; Neira et al. 2013). It may have some ability to displace native species in stressful environments, but it is not regarded as a strong 'pollution indicator' to the same extent as Polydora cornuta.

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