Jassa marmorata

Invasion History

First Non-native North American Tidal Record: 1949
First Non-native West Coast Tidal Record: 1949
First Non-native East/Gulf Coast Tidal Record:

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General Invasion History:

Jassa marmorata was described from Narragansett Bay, Rhode Island in 1903 (Holmes 1905). It has been widely confused with J. falcata, which is known, with certainty, only from the Northeast Atlantic (Conlan 1990). Reports of J. falcata around the world, have been found to refer to Jassa marmorata (Conlan 1990). Here, we assume a North Atlantic origin for J. marmorata, possibly, but not certainly, from the Northwest Atlantic (Cohen and Carlton 1995; Chapman 2000). In the Northwest Atlantic, J. marmorata ranges from the Gulf of St. Lawrence to the Gulf of Mexico (Bousfield 1973; Conlan 1990). In the Northeast Atlantic, it ranges from Sweden to Senegal, throughout the Mediterranean and Black Sea (Conlan 1990; Sezgün et al. 2001; Sorbe et al. 2002). Jassa marmorata is now known from both sides of the temperate South Atlantic, North Pacific, and South Pacific (Conlan 1990; Hong 1993; Poore and Storey 1999). Both Jassa marmorata and J. falcata are tube-dwelling and attach to algae, rocks, pilings, buoys, ship hulls, and ballast water (Woods Hole Oceanographic Institution 1952; Bousfield 1973; Lincoln 1979; Conlan 1990; Cohen and Carlton 1995).

Determining the range of J. marmorata is problematic, due to the presence of similar species [e.g. J. falcata and in the Northeast Atlantic; J. staudei and J. slatteryi in the Northeast Pacific, and many other species elsewhere (Conlan 1990)]. A recent molecular study found that many morphologically identified 'J. marmorata from the Northeast Pacific were actually the native species J. slatteryi and J. staudei', while, all the specimens from North Carolina belonged to an undescribed new species (Pilgrim and Darling 2010). 

North American Invasion History:

Invasion History on the West Coast:

The date of arrival of Jassa marmorata on the West Coast is not known- in addition to historical confusion with J. falcata, at least 8 native or cryptogenic species of Jassa are present (Chapman 2007). One early record of 'J. falcata' from Coos Bay, Oregon, collected in 1949, probably refers to J. marmorata (Barnard 1954, cited by Carlton 1979; Conlan 1990). By the 1950s, it was very abundant in Los Angeles-Long Beach Harbor, where it was a major organism fouling pilings (Barnard 1958). The first San Francisco Bay record is from Oakland in 1977 (Carlton 1979). It now occurs in the Central, South, and San Pablo portions of the Bay (Cohen et al. 2005; Carr et al. 2011). Its current range along the coast extends from Bahia de Los Angeles, Baja California, Mexico (28.9 N) to Point Slocum, Alaska (57.5 N) (Conlan 1990). Within this distribution it has been reported from most of the major saltwater shipping ports (San Diego, Los Angeles-Long Beach, Port Hueneme; San Francisco, Humboldt Bay, Coos Bay, Yaquina Bay, Puget Sound, Victoria) (Barnard 1958; Carlton 1979; Conlan 1990; Cohen et al. 1998; Fairey et al. 2002) and from many smaller bays and harbors, receiving only recreational and fishing traffic (Tijuana Estuary, Alamitos Bay, Morro Bay, Elkhorn Slough, Halfmoon Bay, Tomales Bay, Bodega Harbor, Crescent City Harbor, Willapa Bay, Barkley Sound) (Carlton 1989; Conlan 1990; Cohen et al. 2001; Fairey et al. 2002; Jeff Crooks, personal communication 2005). The relatively late reports of this genus suggest that it was introduced to the West Coast by shipping in the 20th century, rather than by oysters in the 19th century (Carlton 1979; Cohen and Carlton 1995).

Invasion History Elsewhere in the World:

In the Southwest Atlantic, Jassa marmorata has been collected from Cananeia, Brazil (25.0°S) to Chubut Province, Argentina 42.5°S (Conlan 1990; Alonso de Pina 2005). Conlan's paper gives no dates, but it was collected at least as early as 1968 at Santa Clara del Mar, Buenos Aires Province, Argentina (Alonso de Pina 2005). In South Africa, J. marmorata may have been seen (as J. falcata) as early as 1908-1914, but was definitely identified from specimens collected in Table Bay, Cape Town, in 1948 (Conlan 1990; Mead et al. 2011b). On the Indian Ocean side, it was reported from Durban in 1951 (Conlan 1990; Robinson et al. 2005).

In the Southwest Pacific, Jassa marmorata was collected in 1885 in Lyttleton, New Zealand (as J. falcata) (Chilton 1921; Conlan 1990). In Australia, it is known from Port Jackson (Sydney, New South Wales) (Conlan 1990), Port Phillip, and Westernport Bays, Victoria (in 1997, Poore and Storey 1999), and from Blackman's Bay, in Hobart, Tasmania (Conlan 1990). It is present on the coast of Chile from Valparaiso to the Straits of Magellan (Conlan 1990; Gonzlaez 1991). In the Northwest Pacific, J. marmorata is known from South Korea (1st record 2004, Lim et al. 2008); the Yellow Sea of China; Vladivostok, Russia; and the Pacific Coast of Japan (Conlan 1990; Hong 1993). The earliest date of record for the Northwest Pacific is not known, but it was reported from Russian waters of the Sea of Japan by the 1930s (Gurjanova 1938, cited by Doi et al. 2011).

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Description

Jassa marmorata exhibits dramatic sexual dimorphism, especially manifest in the large 2nd gnathopod of the male. Males vary in size (overall and the size of feeding appendages), particularly the 2nd gnathopod, maturing either as 'major' forms with very large gnathopods and large 'thumbs', or 'minor' forms which are smaller overall with much smaller gnathopods. The major forms are thought to be more aggressive in mating, while the latter are thought to function as 'sneaks' (Conlan 1990; Conlan et al. 2021).

Both the major and minor forms have weak rostrums, with medium-large round eyes located near the ends of the prominent anterior lobes. The coxa plates are medium-sized, and mostly separated from plates 2, 3, and 4, which are deeper than 1 and 5. Coxal plate 2 is short and angular. The anterior lobe of plate 5 is more than 3X deeper than the plate's posterior lobe. Both antennae are fairly robust, with Antenna 2 being longer than Antenna 1. Segments 2 and 3 of the peduncle of antenna 1 are roughly equal. The peduncle and the flagellum of Antennae 1 and 2 have numerous long setae on the posterior margin. Segment 5 of Antenna 2 bears simple setae towards the proximal end, but plume-like setae distally. The flagellum of Antenna 2 is very short.

The Gnathopod 1 is similar between the sexes, and 'major' and 'minor' males. The propodus (Segment 6) is oval, narrowing distally, with a convex palm. The dactyl is curved with a finely serrated inner margin. The propodus (segment 2) of gnathopods 2 of a 'major' male is greatly inflated into two lobes, with the main, upper body resembling the palm of a hand, and a lower lobe resembling a thumb. 'Minor' males are smaller and have a smaller propodus, varying in shape, but with a much less developed thumb (Holmes 1905; Conlan 1989). Gnathopod 2 in females is much larger than Gnathopod 1, and the palm of the propodus has a convex inner margin, with proximal and distal protuberances, lined with clusters of setae. Females and juvenile males lack the thumb, which only develops on the male’s last molt.

Pereiopods 3-4 have segment 4 expanded anteriorly and distally. Segments 5 and 6 are short. Pereiopods 5-7 are much longer. Uropods l and 2 have the outer ramus, roughly equal to the inner ramus. Uropod 2 extends slightly beyond Uropod 3, and has blunt rami, proximal to a blunt terminal spine (Holmes 1905; Lincoln 1979; Conlan 1990; Chapman 2007). 'Major' males range up to 10-12 mm in overall length (mean=9), while some males, presumably minors, are as small as 4 mm. Females are 3.3 -11 mm, but with mean lengths of 8.2 (Conlan 1990; Beermann and Purz 2013, Helgoland, North Sea). The 'live' color is reddish brown, marbled with patches of white on the carapaces, antennae, and gnathopods. Above description based on Holmes 1905, Lincoln 1979, Conlan 1990, and Chapman 2007.

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Taxonomy

Ecology

General:

Jassa marmorata is a widespread tube-dwelling amphipod. Gammarid amphipods have separate sexes, brooded embryos, and direct development (Bousfield 1973). Gravid females were seen year-round in Jamaica Bay, New York, but were most abundant in May-August (Franz 1989). Females are larger and more fecund in winter and spring, with monthly mean lengths of 5.7-6.3 mm and 32-44 embryos per female in January through May, versus lengths of 4.6 to 5.4 mm and 16 to 26 embryos in June through December (Franz 1989). In German populations, brood size increased from 2-5 at ~ 5 mm to 125-175 embryos per female at 11 mm (Beerman and Purz 2013).

Amphipods of the genus Jassa not only have strong sexual dimorphism, but also have dimorphism within males, with 'major' forms having large gnathopods and large 'thumbs', and smaller 'minor' forms having much smaller gnathopods and thumbs. The major forms are thought to be more aggressive in mating, while the latter are thought to function as 'sneaks', mating furtively (Borowsky 1985; Conlan 1990; Clark 1997a; Clark and Caudill 2001). The morphological differences are determined by diet quality, rather than genetics (Kurdziel et al. 2002). Major males perch on the females' tubes, display their gnathopods, and drive away other major males, while minor males 'sneak' in, not challnging major males. Males mate multiple times, while females mate only once. Both morphs have equal mating success (Clark 1997).

Jassa marmorata is widely distributed in temperate and subtropical regions, and can survive temperatures from 0-27C (Franz 1989). It is known mostly from polyhaline-marine salinities (18-38 PSU), but has been collected at salinities as low as 12 PSU (Cohen et al. 2001). It secretes threads of 'amphipod silk', to which sediment detritus is attached, to form its tubes. When the amphipods are abundant, they can form a turf, roughly one body-length long. Individuals move inside the tube, and can 'somersault' to emerge at either end. The amphipods partially emerge at the mouth of the tube, extending their antennae. In still water, animals create water movement by beating their pleopods. The setae of the antennae capture food particles, and are also used for tube-building (Dixon and Moore 1997). Jassa marmorata prefers sites with currents sufficient to bring a good supply of food particles, but not strong enough to dislodge the animals (Borowsky 1985). One of the more unusual habitats colonized by J. marmorata are the pipes and tanks of flowing seawater systems in laboratories (Borowsky 1985; Dixon and Moore 1997).  It is more often found in habitats including rocks, oyster reefs, eelgrass beds, marinas, docks, buoys, jetties, shipwrecks and ships' hulls (Bousfield 1973; Franz 1989; Conlan 1990; Carr et al. 2011). In Jamaica Bay, New York, the population was largest in spring and was found growing on bryozoans and other fouling organisms (Franz 1989). Jassa marmorata appears to be primarily a suspension feeder on phytoplankton and detritus, but also preys on ostracods and other small crustaceans (Bousfield 1973; Dixon and Moore 1997). Potential predators include shrimps, crabs, and fishes (Franz 1989).

Food:

Phytoplankton; Detritus

Consumers:

Shrimps, crabs, fishes

Trophic Status:

Suspension Feeder

SusFed

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Habitats

General Habitat	Coarse Woody Debris	None
General Habitat	Oyster Reef	None
General Habitat	Marinas & Docks	None
General Habitat	Rocky	None
General Habitat	Vessel Hull	None
General Habitat	Unstructured Bottom	None
General Habitat	Grass Bed	None
Salinity Range	Mesohaline	5-18 PSU
Salinity Range	Polyhaline	18-30 PSU
Salinity Range	Euhaline	30-40 PSU
Tidal Range	Subtidal	None
Tidal Range	Low Intertidal	None
Vertical Habitat	Epibenthic	None

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Tolerances and Life History Parameters

Minimum Temperature (ºC)	-2	Based on geographical range
Maximum Temperature (ºC)	27	Jamaica Bay, New York (Franz 1989)
Minimum Salinity (‰)	12	Field Data, Willapa Bay WA (Cohen et al. 2001)
Maximum Salinity (‰)	38	Typical Mediterranean salinity
Minimum Length (mm)	3.3	Some mature females were as small as 3.3 mm, but minor males were as small as 4 mm (Conlan 1990; Beermann and Purz 2013).
Maximum Length (mm)	12	Major males, Helgoland, Germany, average length 9mm (Beermann and Purz 2013). Some females reached 11 mm, but averaged 8.2 (Beermann and Purz 2013).
Broad Temperature Range	None	Cold temperate-Subtropical
Broad Salinity Range	None	Mesohaline-Euhaline

General Impacts

Jassa marmorata is a widespread tube-dwelling amphipod, which rapidly colonizes artificial and natural habitats, and can develop dense populations, especially where swift currents bring a steady flow of phytoplankton and other food particles. This amphipod is important prey for shrimps, fishes, and crabs (Bousfield 1973; Franz 1989; Beerman and Purz 2013). There are few reported impacts for this species. 

Economic Impacts

Shipping- Jassa marmorata fouls pilings in Los Angeles-Long Beach Harbor, covering them with masses of tubes covered with sediment, but perhaps provides a benefit by discouraging boring organisms (Barnard 1958).

Ecological Impacts

Habitat Change- Dense masses of amphipod tubes may discourage settlement by boring organisms and other foulers, such as the tunicate Ciona spp. (Barnard 1958). They can also provide habitat for other organisms, such as predatory polychaetes (Barnard 1958).

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Regional Impacts

NEP-VI	Pt. Conception to Southern Baja California		Economic Impact	Shipping/Boating
Jassa marmorata fouls pilings in Los Angeles-Long Beach Harbor, covering them with masses of tubes covered with sediment, but perhaps provides a benefit by discouraging boring organisms (Barnard 1958).
NEP-VI	Pt. Conception to Southern Baja California		Ecological Impact	Habitat Change
Dense masses of amphipod tubes may discourage settlement by boring organisms and other foulers, such as the tunicate Ciona intestinalis (Barnard 1958). They also may provide habitat for other organisms, such as predatory polychaetes (Barnard 1950).
P050	San Pedro Bay		Economic Impact	Shipping/Boating
Jassa marmorata fouls pilings in Los Angeles-Long Beach Harbor, covering them with masses of tubes covered with sediment, but perhaps provides a benefit by discouraging boring organisms (Barnard 1950).
P050	San Pedro Bay		Ecological Impact	Habitat Change
Dense masses of amphipod tubes may discourage settlement by boring organisms and other foulers, such as the tunicate Ciona intestinalis (Barnard 1950). They also may provide habitat for other organisms, such as predatory polychaetes (Barnard 1950).
CA	California		Ecological Impact	Habitat Change
Dense masses of amphipod tubes may discourage settlement by boring organisms and other foulers, such as the tunicate Ciona intestinalis (Barnard 1950). They also may provide habitat for other organisms, such as predatory polychaetes (Barnard 1950).
CA	California		Economic Impact	Shipping/Boating
Jassa marmorata fouls pilings in Los Angeles-Long Beach Harbor, covering them with masses of tubes covered with sediment, but perhaps provides a benefit by discouraging boring organisms (Barnard 1950).

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