Invasion History

First Non-native North American Tidal Record: 1938
First Non-native West Coast Tidal Record: 1938
First Non-native East/Gulf Coast Tidal Record:

General Invasion History:

Melita nitida has a wide native range in the Northwest Atlantic, from the southern Gulf of St. Lawrence to the Gulf of Mexico (Bousfield 1973) and the Caribbean coast of Colombia (Ortiz et al. 2007). This amphipod was first collected on the West Coast in San Francisco Bay, California in 1938 (Carlton 1979; Chapman 1988) and now occurs in estuaries from Howe Sound, British Columbia to the San Gabriel River, near Los Angeles (Carlton 1979; Graening et al. 2012; Murray et al. 2014). In 2000, it was first collected in European waters, in the Scheldt estuary, in the Netherlands (Faasse and van Moorsel 2003) and later (2010) in the Kiel Canal, Germany (Reichert and Beermann 2011).

There is some uncertainty about the identity of the amphipods usually identified as the Northwest Atlantic M. nitida. The Northwest Pacific M. setiflagellata Yamato 1987b is very similar, and has sometimes been considered synonymous, raising the possibility that Asian populations might have been introduced from the Atlantic (Doi et al. 2011) or that West Coast and European populations might have been introduced from Asia (Chapman 2007; Reichert and Beermann 2011; Graening et al. 2012). Currently, most authors use the name M. nitida for the introduced populations on the West Coast and Europe.

North American Invasion History:

Invasion History on the West Coast:

Melita nitida was first collected in the Northeast Pacific at Point Richmond and Lake Merritt (Oakland) in San Francisco Bay, California (CA) in 1938-41 (Carlton 1979). It is now widespread and abundant through the Bay, occurring as far upstream as Collinsville on the Sacramento River during droughts (Cohen and Carlton 1995; Peterson and Vayssieres 2010). It is abundant among tubes of the introduced polychaete Ficopomatus enigmaticus in San Francisco Bay (Carlton 1979; Chapman 1988). Melita nitida is commonly associated with Eastern Oysters (Crassostrea virginica) in its native range, and could have been transported with oyster transplants, but it also occurs in ship fouling communities (Woods Hole Oceanographic Institute 1952) and is a likely candidate for ballast water transport (Cohen and Carlton 1995).

Melita nitida has spread southward on the West Coast in a scattered fashion, to Elkhorn Slough, CA where it was first collected in 1975 (Carlton 1979; Chapman 1988; Wasson et al. 2001), the San Gabriel River estuary, and non-tidal freshwater portions of the Malibu Creek drainage, Los Angeles County in 2002 (M. Born and M. Abramson, Cadien 2007, cited by Graening et al. 2012). However, it has been introduced to many estuaries to the north, including: Coos Bay, Oregon (OR) in 1954 (Carlton 1979); Howe Sound, British Columbia in 1976 (Carlton 1979); Yaquina Bay, OR in 1986 (Chapman 1988); Alsea Bay, OR in 1987 (Chapman 1988); Willapa Bay, Washington (WA) in 1994 (Cohen et al. 2001); Puget Sound, WA in 1998 (Cohen et al. 2001); Humboldt Bay, CA in 2000 (Boyd et al. 2002); and the Columbia River, OR/WA in 2004 (2 specimens, Chapman in Sytsma et al. 2004). The times of first collection, listed above, do not suggest progressive dispersal by coastal currents, but may reflect frequency of collections in different estuaries. Beyond natural dispersal, this amphipod may have spread to West Coast estuaries by human mediated vectors including coastal shipping, bait packed in seaweed, or oyster transplants (Chapman 1988).

Invasion History Elsewhere in the World:

Melita nitida was first collected in the Northeast Atlantic in 1998, in the western Scheldt estuary, Netherlands, where it was found under boulders, often around oysters, near the low-tide mark in polyhaline waters (Faase and van Moorsel 2003). In 2010, 27 specimens were collected in the Kiel Canal, Germany, connecting the North and Baltic Seas, near the Baltic Sea entrance in Kiel (Reichert and Beermann 2011). Future establishment and spread of this amphipod in the North Sea and Baltic seems likely. In 2013-2016, M. nitida was collected at 3 sites on the French coast of the Bay of Biscay: the Gironde estuary, Arcachon Bay (an important oyster-culture site), and Hossegar Lake, near the Spanish border. Shipping is a possible vector in the Gironde estuary and recreational boating is popular on the whole coast (Gouillieux et al. 2016).


Description

Melita nitida has a slender and laterally compressed body with small and oval eyes. Coxal plates 1-4 are large and rounded below. Antenna 1 is long and robust, ~2/3 body length, with a large flagellum of many segments, whorls of stiff setae, and a small accessory flagellum of three segments. On Antenna 2, peduncle segment 5 and the flagellum are densely covered with whorls of stiff setae.

In the male, Gnathopod 1 has a short dactyl, arising on the anterior margin, with the tip curled inward, closing on the inner face of the propodus (segment 6). Gnathopod 2 is larger than Gnathopod 1 in both sexes, but the male has segments 4, 5, and 6 more enlarged, with a smooth, convex palm, and with the dactyl tip closing on the inner side of the palmar angle. The female Gnathopod 2 is similar, but smaller. The basal segments of pereiopods 5-7 are greatly expanded, especially on the posterior-distal corner. The female's Coxa 5 is ventrally extended. The pleon and urosome segments lack dorsal teeth or mucrons (spines or points in the posterior corners). However, Urosome segment 2 has two small clusters of dorsolateral spines. Uropods 1 and 2 are long, extending rearward, beyond the peduncle of Uropod 3. The telson is deeply bilobed, with each lobe having apical and subapical spines. Males reach 12 mm and females reach 9 mm. The color is grayish, with darker bands and a red spot on the head. Description based on: Holmes 1905, Mills 1964, Bousfield 1973, and Chapman 2007.

Parts of the species' range are uncertain, with records which may refer to very similar species. Sheridan (1979, cited by Chapman 1988) described three very similar species (M. elongata, M. intermedia, and M. longisetosa) from the Gulf of Mexico, but Chapman (1988) considered their status uncertain, until comparisons could be made with the lectotypes of M. nitida. Records of M. nitida from the Pacific coast of Mexico (Shoemaker 1935, cited by Chapman 1988; Hendrickx, in Low-Pfeng and Recagno 2012) may refer to an undescribed species of Melita. Melita nitida is nearly indistinguishable from the NW Pacific M. setiflagellata (Yamato 1987b). Some or all West Coast populations could be the latter (Chapman, in Carlton 2007; Graening et al. 2012). Molecular studies of the genus are desirable, but until these are performed we will treat the US West Coast populations as M. nititda.


Taxonomy

Taxonomic Tree

Kingdom:   Animalia
Phylum:   Arthropoda
Subphylum:   Crustacea
Class:   Malacostraca
Subclass:   Eumalacostraca
Superorder:   Peracarida
Order:   Amphipoda
Suborder:   Gammaridea
Family:   Melitidae
Genus:   Melita
Species:   nitida

Synonyms

Potentially Misidentified Species

Melita oregonensis
Melita oregonensis Barnard 1954 is a West Coast native of rocky shores (Chapman 2007).

Melita rylovae
Melita rylovae Bulycheva 1955 is a Northwestern Pacific species, introduced to the California coast (Chapman 20007).

Melita setiflagellata
Melita setiflagellata Yamato 1987 is a very similar species described from Japan. Some authors have treated this amphipod as conspecific with M. nitida and an introduction in Japan, but most regard it as native in Japan (Doi et al. 2011). Given the similarity, some populations of 'M. nitida' on the West Coast could actually be M. setiflagellata (Chapman 2007; Graening et al. 2012). Molecular studies of this amphipod are desirable.

Melita sulca
Melita sulca Barnard 1954 is a West Coast native of harbors, cobble bottoms, and kelp beds (Chapman 2007).

Ecology

General:

Melita nitida is a free-living gammarid amphipod found in shallow, muddy estuaries, salt marshes, mudflats, oyster beds, and fouling communities (Bousfield 1973; Borowsky 1980). In gammarid amphipods, sexes are separate, the young are brooded, and development is direct (Bousfield 1973). Females in Jamaica Bay, New York, had broods of 5-51 embryos, with a mean of 30. Embryos took 10 days to develop at 17C, and 5 days at 21C, and then were brooded as juveniles for less than 2 days before release (Borowsky 1980). Colonization of new patches of habitat is done by adults (Mungia et al. 2007).

Melita nitida is known from temperatures of 0-32C and salinities of 0-35 PSU (Bousfield 1973; Sheridan 1979; Chapman 1988). It is known from a wide range of habitats, including: intertidal mudflats and algal masses, intertidal rocks and debris, clumps of hydroids and bryozoans, floats and pilings, buoys, and crevices under oysters and other bivalve shells (Bousfield 1973; Sheridan 1979; Chapman 1988; Mungia et al. 2007). It was also found in the fouling of a retired cargo ship moored in Suisun Bay, California (Llansó et al. 2011). Melita nitida in the Indian River Lagoon, Florida, fed mostly on seagrass epiphytes, and to a lesser degree on seagrass debris (Zimmerman et al. 1979). Opportunistic predation and scavenging is common in gammarid amphipods, but has not been reported for M. nitida. Likely predators include crabs, shrimps, fishes, and shorebirds.

Food:

Epiphytic algae, seagrass debris

Consumers:

Fishes

Trophic Status:

Herbivore

Herb

Habitats

General HabitatCoarse Woody DebrisNone
General HabitatTidal Fresh MarshNone
General HabitatOyster ReefNone
General HabitatMarinas & DocksNone
General HabitatSalt-brackish marshNone
General HabitatGrass BedNone
Salinity RangeLimnetic0-0.5 PSU
Salinity RangeOligohaline0.5-5 PSU
Salinity RangeMesohaline5-18 PSU
Salinity RangePolyhaline18-30 PSU
Salinity RangeEuhaline30-40 PSU
Tidal RangeSubtidalNone
Tidal RangeLow IntertidalNone
Tidal RangeMid IntertidalNone
Vertical HabitatEpibenthicNone


Tolerances and Life History Parameters

Minimum Temperature (ºC)0Based on geographical range
Maximum Temperature (ºC)32Field data, Florida, (Sheridan 1979, cited by Faasse and van Moorsel 2003)
Minimum Salinity (‰)0Field data, San Francisco Bay (Cohen and Carlton 1995)
Maximum Salinity (‰)35Field data, New England (Bousfield 1973); California (Chapman 1988).
Minimum Length (mm)5Minimum size of females (Borowsky 1980)
Maximum Length (mm)12Adult females, males 9 mm (Bousfield 1973)
Broad Temperature RangeNoneCold temperate-Tropical
Broad Salinity RangeNoneTidal Limnetic-Euhaline

General Impacts

Melita nitida is common in many West Coast estuaries. It is probably a prey item for fishes and shorebirds; however, no specific impacts have been reported.

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
P045 _CDA_P045 (Santa Ana) 2007 Non-native Established
NEP-VI Pt. Conception to Southern Baja California 2007 Non-native Established
P130 Humboldt Bay 2000 Non-native Established
P080 Monterey Bay 1975 Non-native Established
NEP-IV Puget Sound to Northern California 1954 Non-native Established
P093 _CDA_P093 (San Pablo Bay) 1938 Non-native Established
P090 San Francisco Bay 1938 Non-native Established
NEP-V Northern California to Mid Channel Islands 1938 Non-native Established

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude
697186 Introduced Species Study 2005 2005-10-19 Mare Island Strait - Navy Non-native 38.1015 -122.2695
697190 Introduced Species Study 2010 2010-07-29 Mare Island Strait - Navy Non-native 38.1015 -122.2695
697297 Introduced Species Study 2010 2010-06-11 Cal Maritime Academy/Vallejo Non-native 38.0661 -122.2299
697321 Introduced Species Study 2006 2006-08-10 Tomales Bay Boat Launch Non-native 38.1991 -122.9220
697499 Boyd et al. 2002 (Humboldt Bay Report) 2002 Klopp Lake Non-native 40.8553 -124.0919
697574 Introduced Species Study 2005 2005-10-07 Benicia Waterfront Non-native 38.0401 -122.1385
697575 Introduced Species Study 2010 2010-06-29 Benicia Waterfront Non-native 38.0401 -122.1385
697682 Introduced Species Study 2010 2010-07-13 Port Sonoma/Petaluma R. Non-native 38.1157 -122.5026
697825 Introduced Species Study 2005 2005-10-04 Berkeley Marina Non-native 37.8676 -122.3172
698265 Boyd et al. 2002 (Humboldt Bay Report) 2002 Hookton Slough Non-native 40.6775 -124.2218
698325 Introduced Species Study 2005 2005-10-20 Loch Lomond Marina Area Non-native 37.9720 -122.4832
698441 Introduced Species Study 2006 2006-11-01 B-Dock Non-native 36.8027 -121.7851
698451 Introduced Species Study 2005 2005-10-06 Santa Fe Channel - Front Non-native 37.9101 -122.3644
698460 Introduced Species Study 2010 2010-06-28 Santa Fe Channel - Front Non-native 37.9101 -122.3644
698516 Introduced Species Study 2010 2010-07-14 Romberg Tiburon Center Non-native 37.8906 -122.4458
698857 Cohen et al. 2005 (SF Bay Area RAS) 2004 2004-05-25 Port Sonoma, San Pablo Bay Non-native 38.1156 -122.5026
699048 Introduced Species Study 2010 2010-07-13 Petaluma River Turning Basin Non-native 38.2344 -122.6354
699214 Introduced Species Study 2005 2005-09-08 Treasure Island Non-native 37.8149 -122.3702
699379 Introduced Species Study 2005 2005-07-06 Coyote Point Non-native 37.5920 -122.3210
699525 Cohen et al. 2005 (SF Bay Area RAS) 2004 2004-05-26 Point San Pablo Yacht Harbor, San Pablo Bay Non-native 37.9624 -122.4188
699555 Introduced Species Study 2010 2010-05-31 Redwood Creek - Shipping Non-native 37.5120 -122.2109
699765 Introduced Species Study 2010 2010-06-30 Mare Island Strait - Marina Non-native 38.1051 -122.2667
699769 Introduced Species Study 2005 2005-10-19 Mare Island Strait - Marina Non-native 38.1051 -122.2667
699873 Introduced Species Study 2005 2005-09-07 Redwood Creek - Marina Non-native 37.5021 -122.2130
700009 Introduced Species Study 2005 2005-06-09 McNears Beach Non-native 37.9962 -122.4556
700065 Boyd et al. 2002 (Humboldt Bay Report) 2002 Southport Landing Non-native 40.6952 -124.2494
700187 Introduced Species Study 2006 2006-11-01 MLML Small Boats Non-native 36.8041 -121.7860
700524 Introduced Species Study 2005 2005-10-05 Sea Plane Lagoon Non-native 37.7761 -122.2998
700795 Introduced Species Study 2005 2005-09-09 Sea Plane Harbor Non-native 37.6349 -122.3848
700866 Introduced Species Study 2005 2005-10-18 Pacheco Creek Oil Pier Non-native 38.0489 -122.0903
700970 J. Chapman, pers. comm., in Carlton 1979 1977 Elkhorn Slough General Location Non-native 36.8086 -121.7856
700975 Chapman 1988 1975 Elkhorn Slough at Kirby Park Non-native 36.8398 -121.7435
701121 Chapman 1988 1938 Point Richmond Non-native 37.9230 -122.3940
701340 Introduced Species Study 2005 2005-10-06 Santa Fe Channel - Back Non-native 37.9207 -122.3684
701438 Introduced Species Study 2010 2010-06-30 Hercules Wharf Non-native 38.0231 -122.2928
701443 Introduced Species Study 2005 2005-10-19 Hercules Wharf Non-native 38.0231 -122.2928
702167 Introduced Species Study 2005 2005-10-04 Oakland Outer Harbor Non-native 37.8217 -122.3145
702904 Boyd et al. 2002 (Humboldt Bay Report) 2002 Bracut Non-native 40.8313 -124.0845
702944 Introduced Species Study 2006 2006-08-10 Golden Hinde Small Marina Non-native 38.1078 -122.8623
702979 Cohen et al. 2005 (SF Bay Area RAS) 2004 2004-05-25 Petaluma River Turning Basin, San Pablo Bay Non-native 38.2355 -122.6382
703261 Introduced Species Study 2005 2005-11-15 China Camp Non-native 38.0025 -122.4617
703399 Introduced Species Study 2005 2005-10-05 Oakland Inner Harbor - Small marinas Non-native 37.7847 -122.2669
703784 Introduced Species Study 2005 2005-06-10 Hayward Landing Non-native 37.6447 -122.1543
703903 Introduced Species Study 2010 2010-06-30 Rodeo Marina Non-native 38.0394 -122.2717
703905 Introduced Species Study 2005 2005-10-19 Rodeo Marina Non-native 38.0394 -122.2717
703970 Introduced Species Study 2010 2010-05-31 Railroad Bridge Non-native 37.4602 -121.9750
703980 Introduced Species Study 2005 2005-09-07 Railroad Bridge Non-native 37.4602 -121.9750
704520 Introduced Species Study 2010 2010-06-30 Napa Valley Marina Non-native 38.2198 -122.3119
704521 Introduced Species Study 2005 2005-10-19 Napa Valley Marina Non-native 38.2198 -122.3119
718569 California Academy of Sciences Invertebrate Zoology Collection Database 1986 1986-09-18 Grizzly Bay Non-native 38.1162 -122.0388
718570 California Academy of Sciences Invertebrate Zoology Collection Database 1976 1976-08-25 South of Coyote Hills Slough mouth Non-native 37.5582 -122.1280
718571 California Academy of Sciences Invertebrate Zoology Collection Database 2001 2001-05-04 Petaluma Marina, outer floating dock Non-native 38.2294 -122.6131
718573 California Academy of Sciences Invertebrate Zoology Collection Database 2001 2001-04-13 Vallejo Marina, Vallejo Non-native 38.1071 -122.2692
718574 California Academy of Sciences Invertebrate Zoology Collection Database 2001 2001-05-04 Petaluma Marina, inner floating dock Non-native 38.2313 -122.6141
718582 California Academy of Sciences Invertebrate Zoology Collection Database 2002 2002-03-26 Port Sonoma Non-native 38.1186 -122.4983
758602 Chapman 1988 1941 1941-06-15 Lake Merritt Non-native 37.8025 -122.2578
758603 Light 1941, cited in Carlton 1979a 1941 Lake Merritt Non-native 37.8025 -122.2578
758604 Chapman 1988 1942 1942-03-28 Lake Merritt Non-native 37.8025 -122.2578
758605 Carlton 1979a 1953 Redwood City Non-native 37.5032 -122.2146
758606 Bousfield and Carlton 1967 1960 Lake Merritt Non-native 37.8025 -122.2578
758607 Chapman and Dorman 1975 1966 1966-12-22 Lake Merritt Non-native 37.8025 -122.2578
758608 California Academy of Sciences Invertebrate Zoology Collection Database 1971 1971-11-21 Lake Merritt Non-native 37.8025 -122.2578
758609 California Academy of Sciences Invertebrate Zoology Collection Database 1976 1976-08-25 Mouth of Coyote Hills Slough Non-native 37.5628 -122.1314
758610 California Academy of Sciences Invertebrate Zoology Collection Database 1976 1976-08-26 South of Coyote Hills Slough mouth Non-native 37.5582 -122.1280
758611 Chapman 1979; unpublished manuscript cited in Carlton 1979a 1979 San Francisco Bay Non-native 37.8494 -122.3681
758612 Yale Peabody Museum of Natural History Collections Database 2015 1985 Muzzi Marsh Non-native 37.9279 -122.5070
758613 Cohen and Chapman 2005 2005 2005-11-27 Dolphin # 11 Non-native 38.0532 -122.3293
758614 Heiman and Micheli 2010; Hobbs et al. 2015 2002 Elkhorn Slough (Upper) Non-native 36.8547 -121.7600
768140 Ruiz et al., 2015 2012 2012-09-06 Loch Lomond Marina, San Francisco Bay, CA, California, USA Non-native 37.9736 -122.4802
770898 Ruiz et al., 2021a 2017 2017-09-26 Oyster Point Marina, San Francisco Bay, California, USA Non-native 37.5869 -122.3237
771080 Ruiz et al., 2021a 2017 2017-09-19 Richmond Marina Bay Yacht Harbor, San Francisco Bay, California, USA Non-native 37.9129 -122.3494
771122 Ruiz et al., 2021a 2017 2017-09-19 Richmond Marina Bay Yacht Harbor, San Francisco Bay, California, USA Non-native 37.9129 -122.3494
771342 Ruiz et al., 2021a 2017 2017-09-21 San Leandro Marina, San Francisco Bay, California, USA Non-native 37.6977 -122.1912
771350 Ruiz et al., 2021a 2017 2017-09-21 San Leandro Marina, San Francisco Bay, California, USA Non-native 37.6977 -122.1912
771379 Ruiz et al., 2021a 2017 2017-09-21 San Leandro Marina, San Francisco Bay, California, USA Non-native 37.6977 -122.1912
771649 Ruiz et al., 2021a 2018 2018-09-12 Benicia Marina, San Francisco Bay, California, USA Non-native 38.0453 -122.1539
771936 Ruiz et al., 2021a 2018 2018-09-24 Ballena Isle Marina, San Francisco Bay, California, USA Non-native 37.7654 -122.2845
772387 Ruiz et al., 2021a 2018 2018-09-28 Loch Lomond Marina, San Francisco Bay, California, USA Non-native 37.9723 -122.4818
772396 Ruiz et al., 2021a 2018 2018-09-28 Loch Lomond Marina, San Francisco Bay, California, USA Non-native 37.9723 -122.4818
772446 Ruiz et al., 2021a 2018 2018-09-28 Loch Lomond Marina, San Francisco Bay, California, USA Non-native 37.9723 -122.4818

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