Alitta succinea

Invasion History

First Non-native North American Tidal Record: 1896
First Non-native West Coast Tidal Record: 1896
First Non-native East/Gulf Coast Tidal Record:

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General Invasion History:

Alitta succinea was described from the North Sea, Germany in 1847. However, Sato (2013) suggests that the East Coast of North America is a probable native region. On the East Coast, south of Cape Cod, A. succinea is the sole nereid in low-salinity waters, while in Europe (and the Gulfs of Maine and St. Lawrence) Hediste diversicolor also occurs in brackish-water habitats, and often occurs in lower salinities than A. succinea. Alitta succinea expanded its range in Europe in the 20th century (Pettibone 1963; Sato 2013), especially in the end of its northern range. It was first recorded on the west coast of Sweden in 1940 and in Denmark in 1949 (Jensen 2010). In the western Atlantic, its reported range extends into the tropics (Colombia, Venezuela, and Bahia State, Brazil) and further south through Brazil, Uruguay and northern Argentina (Pettibone 1963; Londoño-Meso et al. 2002; Orensanz et al. 2002; de Oliveira et al. 2012). This species appears to have evolved on one side or the other of the North Atlantic Ocean, but has clearly colonized many other regions. In this account we will treat the many populations as a single species, but worldwide genetic studies of this species are needed.

North American Invasion History:

Invasion History on the West Coast:

Alitta succinea was first reported from the West Coast of the US in the Oakland estuary in San Francisco Bay, California (CA) in 1896 (Carlton 1979). In San Francisco Bay, its range extends into the South Bay, San Pablo Bay, and Antioch at the western end of the Delta. In the 20th century, A. succinea spread to the north and south. In 1952-1956, A. succinea was collected in the San Gabriel River and Alamitos Bay, CA and later (1972) in Los Angeles Harbor and Newport Bay, CA (Carlton 1979). Further south, this species occurred in scattered locations on the Pacific coasts of Mexico, Costa Rica, Panama, and Colombia (Low-Pfeng and Recagno 2012; Bastida-Zavala et al. 2014; US National Museum of Natural History 2014). In 1941, A. succinea was found in Tomales Bay, CA (Carlton 1979). To the north it was found in Netarts Bay, Oregon (OR) by 1976 (Stout 1976, cited by Carlton 1979); Coos Bay, OR by 1986 (Carlton 1989; Wonham and Carlton 2005); and Puget Sound, Washington in 1998 (Cohen et al. 1998; Cohen et al. 2001).

Alitta succinea was discovered in 1936 in the Salton Sea, California. It may have been introduced as bait or accidentally with stocked marine fish. This salt lake was accidentally produced by a flood flowing through a man-made canal connected to the Colorado River, and has increased in salinity through evaporation from brackish to marine, and now to hypersaline conditions (44-60 g/l ). The Pile Worm remains the dominant macrobenthic organism in the lake (Riedel and Costa-Pierce 2005).

Invasion History in Hawaii:

Alitta succinea was collected at an unspecified location on Oahu in 1941, and in Waikiki Bay, Oahu in 1945 (Coles et al. 2002; Carlton and Eldredge 2009). A later collection was in the Alai Wai Canal, Oahu, where it was 'abundant in the mud along the banks' (Bailey-Brock and Hartman 1987, cited by Carlton and Eldredge 2009).

Invasion History Elsewhere in the World:

As noted above, A. succinea occurs on the Pacific Coast of Mexico, Central America, and Colombia. Early records (1957) from the Gulf of Panama and Colombia suggest transport through the Panama Canal (Pettibone 1963; U.S. National Museum of Natural History 2014). Other records from the Pacific coast of Baja California, the Gulf of California, Sinaloa (1st record 1972), and Costa Rica (1st record 1980) are more scattered and indicative of sporadic transport along the coast by shipping or aquaculture (Villalobos-Guerrero et al., in Low-Pfeng and Recagno 2012; Bastida-Zavala et al. 2014; US National Museum of Natural History 2014).

Alitta succinea can be considered cryptogenic on both sides of the North Atlantic, although it was first described from the island of Helgoland, Germany, in the North Sea in 1847 (Sato 2013). Its distribution appears to be spotty in some regions. Dewarumez et al. (1992) noted a single specimen in Dunkerque Harbor, France and noted that this species was not found in the fauna of Roscoff or Plymouth. It appears to have expanded its range northward in the 20th century. It was first recorded in Sweden in 1940, Denmark in 1949, and Norway, in the Oslofjord, before 1990 (Abbiati 1990).

More definite introductions of A. succinea occurred in the Caspian Basin. In 1940, nereid polychaetes from the Sea of Azov were introduced to the Caspian Sea. These were initially identified as A. succinea (Spassky 1945) and later reported to be all Hediste diversicolor (eg. Aladin et al. 2002; Grigorovich et al. 2003). Recent collections and historical records indicate that both species were introduced and established (Ghasemi et al. 2013). Alitta succinea was also introduced to the Aral Sea, and was abundant by 1974 (Proskurina 1980, cited by Sato 2013).

In the Northwest Pacific, A. succinea was first recorded in Tokyo Bay in 1964. It is now abundant in Japanese and Korean estuaries which are subject to pollution and eutrophication (Sato 2013). It has been reported from Vladivostok, Russia in the Sea of Japan (Khlebovich 1996, cited by Sato 2013); Masan Bay, South Korea; Tianjin, China, on the Bohai Sea (Huang 2001; Sato 2013); and Hong Kong (Wang and Huang 1993).

In Australia, A. succinea was first collected in Port Phillip Bay in 1978 (Wilson 1999). Records from other parts of the continent are based on records of Nectoneanthes oxypoda (Sato 2013). It occurs sporadically in estuaries near Melbourne (Wilson 1999). This polychaete has also been introduced to the southeast coast of South Africa. Day (1967, cited by Mead et al. 2011b) found it in Durban Bay, and several bays near Port Elizabeth.

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Description

Alitta succinea has an elongate, cylindrical body, divided into up to 160 segments. The prostomium is pear-shaped, with four eyes, two frontal antennae, and a pair of stout conical palps. The prostomium is flanked by four pairs of tentacular cirri. The posterior pair of tentacular cirri is longest, and can reach back to chaetigers 4-15. Ventrally and anteriorly, the muscular extendable proboscis consists of two rings, terminating in a pair of amber-colored jaws, with 4-9 teeth each. It is possible to see it only when the organism is relaxed. The proboscis is marked by patches of amber-colored denticles (paragnaths) arranged in species-specific patterns (see Pettibone (1963), Sato (2013) and Villalobos-Guerrero and Carrera-Parra (2015), for descriptions of these patterns in A. succinea). These patterns and details of chaetal structure (acicula, homogomph, heterogomph, spiniger, falciger), are needed for identification of species (Pettibone 1963; Blake and Ruff in Carlton 2007; Sato 2013; Villalobos-Guerrero and Carrera-Parra 2015), but will not be dealt with here.

The parapodia vary greatly in form from anterior to posterior, with the posterior appendages being longer and more elongated in shape. The two anterior-most parapodiae are not fully biramous. The subsequent anterior parapodia are divided into two branches, which are in turn divided into smaller lobes, called ligules. The dorsal lobe is called the notopodium. It has a dorsal cirrus, which does not extend beyond the ligules. The dorsal ligule is large and triangular, the lowest (ventral) one is smaller, and the middle one, called the prechaetal lobe, is smallest, about 1-2 to 2/3 the size of the ventral ligule, and bears a bundle of thin chaetae. The neuropodium has a dorsal ligule, a broader median post chaetal lobe, with a bundle of thicker chaetae, a ventral ligule, and a ventral cirrus. The parapodia start to change in the middle of the worm and are longer and different in structure in the posterior region. The dorsal ligule is long and strap-shaped, with the dorsal cirrus near the tip, the middle ligule reduced or absent, and the lower one short and conical. The neuropodium is more like that of the anterior region, except that the postchaetal ligule is missing (Pettibone 1963; Sato 2013).

Specimens of A. succinea range up to 170 mm. The worm is brownish anteriorly, with the prostomium and bases of the parapodia darkly pigmented. The rest of the body can be greenish, greenish-yellow, or pale reddish, and sometimes with white or dark dots. A red dorsal blood vessel runs down the midline of the body. This species is found in a wide range of habitats, including mud and sand bottoms, oyster beds, fouling communities, etc., and often occurs in brackish waters (Pettibone 1963).

Alitta succinea undergoes a dramatic morphological change (epitoky) when breeding, with both sexes changing into a 'heteronereis' form. The segments become compressed in the antero-posterior direction, so that the worm's body is shorter (14-55 mm in males; 30-75 mm in females). The eyes are enlarged, especially in males. The body becomes divided into three sections, an anterior section of 13-18 segments with unchanged chaetae, a middle section of 29-56 segment, with flattened parapodia and paddle-like chaetae, and a posterior 'tail' with un-modified chaetae. The body of the male becomes bright red, with a white tail, while the female is paler, white to yellow-green. During this mating period, the adults swarm and swim at the surface (Pettibone 1963).

The planktotrophic larval stages were described by Banse (1954) and Kinne (1954). Hansen (1999) shows an illustration of a 6-chaetiger larva.

Villalobos-Guerrero and Carrera-Parra (2015) re-described Alitta succinea, using exclusively material from the North Sea, Germany, near the type locality, whereas previous authors had combined features from multiple locations. They consider the globally reported 'A. succinea' to be a complex of species of unresolved native-invasion status. They examined worms previously A. succinea from Eastern Tropical Pacific in Mexico and Guatemala, and restored an earlier species name, A. acutifolia. Their work suggests that detailed morphological and genetic examination of 'A. succinea' populations worldwide will be needed to resolve their identity and invasion status.

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Taxonomy

Ecology

General:

Alitta succinea is found in a wide range of intertidal and subtidal, brackish to marine habitats. Reproduction involves a dramatic metamorphosis (called epitoky), influenced by temperature, salinity, and (possibly) photoperiod (Kinne 1954; Fong 1991). Sexes are separate, and males usually transform at a smaller size (14-55 mm) than females (30-75 mm). The eyes become larger, especially in males. During metamorphosis, the body structure is modified and divided into three sections. The mid-section becomes modified for swimming, with paddle-like chaetae (Pettibone 1963). These modified adults 'heteronereis' swarm, often in immense numbers at the surface. The worms are attracted to light and swim rapidly, often in a gyrating motion, with males swimming faster and often pursuing females. Eggs and sperm are released into the water. Males and females die after spawning- this species is semelparous. Swarming has been observed both in daylight hours and at night (Pettibone 1963). In experiments in San Francisco Bay, higher salinity (20 vs. 5 PSU) and temperature (~18 vs. ~15°C) resulted in increased rates of metamorphosis. Increasing salinity from 5 to 20 PSU caused a 3-fold increase in metamorphosis. No photoperiod effects were seen in these experiments (Fong 1991). On the East Coast (MA-NC), swarming has been seen at various times from March to September (Pettibone 1963). On the West Coast, in San Leandro Creek, San Francisco Bay, spawning occurred in late August-early September (Fong 1991).

Females may carry 80,000 to 100,000 eggs (Elton 1958). Fertilized eggs develop into swimming trochophores and develop segments and protruding chaetae. Larvae are planktotrophic and feed on phytoplankton. When individuals have 8-10 chaetigerous segments, they settle. This takes about 10-14 days at 20-24°C (Banse 1954; Kinne 1954; Hansen 1999). They do not tolerate salinity lower than 14.5 PSU (Kinne 1954) and this could be the explanation for the species' biogeographical distribution.

Alitta succinea is reported to occur from cold-temperate to tropical habitats, although genetic comparisons of temperate and tropical populations have not been made. In northern parts of its range, it inhabits estuaries which are ice-covered in winter. Adults tolerate salinities as low as 2.5 PSU (Freel et al. 1973) and as high as 65 PSU (Kuhl and Oglesby 1979). Populations have survived in athalassic salt lakes such as the Salton Sea, Caspian Sea, and Aral Sea, although their salt compositions differ considerably from normal sea water (Kuhl and Oglesby 1979; Ghasemi et al. 2013). Reproductive metamorphosis in adults from San Francisco Bay occurs in salinities as low as 5 PSU, although some populations from elsewhere show impaired larval development at salinities below 8-11 PSU (Fong 1991). This polychaete can occur in 'very foul mud' (Pettibone 1963) and has colonized harbors in Japan where native nereids have been eliminated by development or pollution (Sato 2013). It can even tolerate brief period of hypoxia (1-3 days) (Kristensen 1983).

Alitta succinea favors sheltered habitats, including: coves, estuaries, marshes, and mangrove areas. In sand, mud, or peat, this worm digs a broad U-shaped burrow. It also occurs under rocks, among oyster shells, barnacles, mussels, and sponges (Pettibone 1963). This worm is omnivorous, feeding on algae, detritus, diatoms, small invertebrates (including oyster larvae), and carrion (Fauchald and Jumars 1979; Barnes et al. 2010; de Oliveira et al. 2012). In its native and introduced ranges, it is an important food of crabs, fishes, and shorebirds (Pettibone 1963; Riedel and Costa-Pierce 2005; Ghasemi et al. 2013). On the East Coast of the US, it is the dominant nereid polychaete in estuarine waters, but in European waters, it dominates a salinity zone intermediate between that of the more marine Alitta virens and the more brackish-water Hediste diversicolor (Wolff 1974; Sato 2013). In San Francisco Bay, it occupies zones intermediate between the more marine Nereis vexillosa and the brackish-fresh Hediste limnicola (Cohen and Carlton 1995). In European waters, A. succinea and H. diversicolor are often mutually exclusive (Wolff 1973).

Food:

Algae, detritus, invertebrates

Consumers:

Fishes, crabs, birds

Competitors:

other nereid polychaetes

Trophic Status:

Omnivore

Omni

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Habitats

General Habitat	Coarse Woody Debris	None
General Habitat	Unstructured Bottom	None
General Habitat	Oyster Reef	None
General Habitat	Marinas & Docks	None
General Habitat	Mangroves	None
General Habitat	Vessel Hull	None
General Habitat	Grass Bed	None
Salinity Range	Oligohaline	0.5-5 PSU
Salinity Range	Mesohaline	5-18 PSU
Salinity Range	Polyhaline	18-30 PSU
Salinity Range	Euhaline	30-40 PSU
Salinity Range	Hyperhaline	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 Temperature (ºC)	0	Based on geographical range
Maximum Temperature (ºC)	34	Maximum temperatures were monthly means (August) recorded in a powerplant effluent in the Patuxent River estuary, Maryland, where N. succinea was present (Nauman and Cory 1969).
Minimum Salinity (‰)	2.5	Survival at the minimum salinity was observed in animals from Colorado Lagoon, Long Beach CA Bay at 20 ºC (Freel et al. 1973).
Maximum Salinity (‰)	65	The maximum salinity was tested with worms from the Salton Sea using “Instant Ocean” artificial seawater (Kuhl and Oglesby 1979).
Minimum Reproductive Salinity	14.5	Larval development was unsuccessful at salinities below 14.5 PSU (Kinne 1954).
Maximum Reproductive Salinity	50	Epitokal maturation and normal embryonic and larval development in Salton Sea animals was seen at salinities as high as 45 ppt (Kuhl and Oglesby 1979)
Minimum Duration	10	20-24 C, 14-16 PSU, animals from Kiel Bay, Germany (Banse 1954).
Maximum Duration	14	20-24 C, 14-16 PSU, animals from Kiel Bay, Germany (Banse 1954).
Minimum Length (mm)	14	Male heteronereids, 14-55 mm; Female heteronereids, 30-75 mm (Pettibone 1963)
Maximum Length (mm)	170	Pettibone 1963
Broad Temperature Range	None	Cold-temperate-Tropical
Broad Salinity Range	None	Mesohaline-Euhaline

General Impacts

Alitta succinea is a relatively large, burrowing omnivorous polychaete. In the areas in which it has been introduced, it coexists with native nereid worms, but the degree to which those species are affected by competition, or displaced, is unknown. In European estuaries, A. succinea appears to be restricted to intermediate salinity ranges between the more marine Alitta virens and the more brackish Hediste diversicolor, and appears to require somewhat warmer conditions (Pettibone 1963). In San Francisco Bay, it is intermediate in salinity range between Nereis vexillosa and the brackish-fresh Hediste limnicola. It is possible that it has 'compressed' the ranges of the two natives through competition (Cohen and Carlton 1995). In Japan, A. succinea has invaded habitats where native nereids (Hediste japonica; Nectoneanthes uchiwa) have been disturbed by development, eutrophication, and pollution (Sato 2013). However, the role of competition has not been studied in these nereids. Alitta succinea is a major prey item of migratory shorebirds, and may be more important than native nereids because of its large size (Recher 1966; Iwamatsu et al. 2007).

More definite impacts have been observed when A. succinea has been introduced to saline lakes lacking connections to the sea, and lacking large polychaetes. Alitta succinea is a large burrowing animal, and is both a substantial food item for fishes and birds, and also has the potential to alter food and nutrient webs in salt lakes such as the Salton Sea, Sea of Azov, and the Caspian Sea (Riedel and Costa-Pierce 2005; Ghasemi et al. 2013).

Economic Impacts 

Fisheries- Alitta succinea was introduced to the Caspian, Aral, and Salton Seas to enhance fisheries in these saline lakes, and has provided an additional prey item for native and introduced fishes (Kuhl and Oglesby 1979; Riedel and Costa-Pierce 2005; Ghasemi et al. 2013). In the Salton Sea, an accidental, artificial body of salt water in a desert region of California, it provided the basis of the food chain which supported introduced gamefishes (Anisotremus davidsoni, Baidiella incistia, and Cynoscion xanthulus) from the Gulf of California (Kuhl and Oglesby 1979). As salinity increased, the marine fishes were replaced by hybrid Tilapia (Oreochromis mossambicus X urolepis) still largely dependent on A. succinea as food (Riedel and Costa-Pierce 2005). Similarly, introduced A. succinea and Hediste diversicolor remained abundant and a food source for the remaining fishes during a catastrophic increase in salinity and desiccation of the Aral Sea (Aladin et al. 2008; Proskurina 1980, cited by Sato 2013).

Ecological Impacts

Food/Prey- As noted above, A. succinea is a major prey item for fishes, especially in salt lakes with no native large polychaetes, such as the Caspian, Aral, and Salton Sea. As a large burrowing animal it has the potential to return buried carbon and nutrient resources to the foodweb (Kuhl and Oglesby 1979; Riedel and Costa-Pierce 2005; Swan et al. 2007; Aladin et al. 2008; Proskurina 1980, cited by Sato 2013). Alitta succinea is also an important prey item for migratory shorebirds, such as sandpipers and plovers in its native and introduced range. It may be more important than native nereids in its introduced ranges, because of its large size and wide environmental tolerances (Recher et al. 1966; Iwamatsu et al. 2007). In the Salton Sea, A. succinea is a major prey item for migrating populations of the Eared Grebe (Podiceps nigricollis). Die-offs of the worm due to anoxia and hydrogen sulfide build-up in sediments, have resulted in die-offs of birds on the Salton Sea (Anderson et al. 2007).

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Regional Impacts

CASP	Caspian Sea		Ecological Impact	Food/Prey
Alitta succinea, together with Hediste diversicolor, were added as fish forage to enhance Caspian Sea fisheries in the 1940s. By 1949, the pelagic heteronereis spawning stage of A. succinea had become a major food item of planktivorous fishes (shad, Alosa saposhnikovi and sprat, Clupeonella engrauliformis). This worm is probably also a major food source for benthic fishes, such as sturgeon, Striped Mullet (Mugil cephalus), and gobies (Neogobius sp.)(Karpinsky et al. 2005, cited by Ghasemi et al. 2013).
NWP-3b	None		Ecological Impact	Food/Prey
Alitta succinea was a major prey item for five species of migrating shorebirds on estuarine mudflats in Fujimae-Higata on Ise Bay, Honshu, Japan. The bird species were Dunlin (Calidris alpina), Red-necked Stint (C. ruficollis), Grey Plover (Pluvialis squatarola), Whimbrel (Numenius phaeopus), and Black-headed Gull (Larus ridibundus). Alitta succinea dominated the polychaete biomass at some times and locations because of its larger body size than the native Hediste diadroma (Iwamatsu et al. 2007).
P090	San Francisco Bay		Ecological Impact	Food/Prey
Alitta succinea was eaten by 11 species of shorebirds, at Palo Alto CA, on South San Francisco Bay. For four of these species [Semipalmated Plover (Charadrius semipalmatus), Long-Billed Dowitcher (Limnodromus scolopaceus), Dunlin (Calidris alpina), Marbled Godwit (Limosa fedoa)], A. succinea was more than 50% of gut contents (Recher 1966).
NEP-V	Northern California to Mid Channel Islands		Ecological Impact	Food/Prey
Alitta succinea was eaten by 11 species of shorebirds at Palo Alto CA, on South San Francisco Bay. For four of these species [Semipalmated Plover (Charadrius semipalmatus), Long-Billed Dowitcher (Limnodromus scolopaceus), Dunlin (Calidris alpina), Marbled Godwit (Limosa fedoa)] A. succinea was more than 50% of gut contents (Recher 1966).
CASP	Caspian Sea		Economic Impact	Fisheries
Alitta succinea, together with Hediste diversicolor, were added as fish forage to enhance Caspian Sea fisheries in the 1940s (Spassky 1945; Ghasemi et al. 2013)
CA	California		Ecological Impact	Food/Prey
Alitta succinea was eaten by 11 species of shorebirds at Palo Alto CA, on South San Francisco Bay. For four of these species [Semipalmated Plover (Charadrius semipalmatus), Long-Billed Dowitcher (Limnodromus scolopaceus), Dunlin (Calidris alpina), Marbled Godwit (Limosa fedoa)] A. succinea was more than 50% of gut contents (Recher 1966)., Alitta succinea was eaten by 11 species of shorebirds, at Palo Alto CA, on South San Francisco Bay. For four of these species [Semipalmated Plover (Charadrius semipalmatus), Long-Billed Dowitcher (Limnodromus scolopaceus), Dunlin (Calidris alpina), Marbled Godwit (Limosa fedoa)], A. succinea was more than 50% of gut contents (Recher 1966).

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