Invasion History

First Non-native North American Tidal Record: 1986
First Non-native West Coast Tidal Record: 1986
First Non-native East/Gulf Coast Tidal Record:

General Invasion History:

The estuarine amphipod Corophium heteroceratum was described from the Tanggu region, on the Bohai Sea, China (Chapman 1988; Huang 2001). It also occurs in the Yellow Sea (Wang et al. 2008). It was first collected in San Francisco Bay, California between 1985 and 1989. It occurs as far upstream as brackish Grizzly Bay, but is most abundant in subtidal muddy and silty sediments in saltier parts of the central, south, and San Pablo Bays (Cohen and Carlton 1995; Lee et al. 2003; Chapman 2007). It was also collected in Los Angeles-Long Beach Harbor, as early as 1990 (Cohen and Carlton 1995; Ranasinghe et al. 2005).

North American Invasion History:

Invasion History on the West Coast:

Corophium heteroceratum was first recognized in San Francisco Bay by 1989, and was probably collected as early as 1986 (John Chapman, personal communication, cited by Cohen and Carlton 1995). In the San Francisco estuary, it occurs in lower intertidal to subtidal muddy and silty sediments, as far up the estuary as Suisun and Grizzly Bays (mean salinity 5.9 PSU). It is more abundant in the more saline parts of San Pablo Bay, and the central and South Bays, at a mean salinity of 27.5 PSU (Cohen and Carlton 1995; Lee et al. 2003; Peterson and Vayssieres 2010). It was also collected in Tomales Bay (Fairey et al. 2002) and in Los-Angeles-Long Beach Harbors (1st record 1990, Cohen and Carlton 1995; Ranasinghe et al. 2005). Ballast water is the most likely vector for transport, since infaunal corophiid amphipods burrow in soft mud, but occasionally swim in the water column (Chapman 1988; Cohen and Carlton 1995).


Corophium heteroceratum has a slender, very depressed body, and a massive Antenna 2, which is longer than Antenna 1. The coxal plates are small and separated. The urosome segments are not fused. Pereionites 2-7 bear sharp downward projections. Both sexes have a pointed rostrum, extending beyond the eye-lobes. The eye is well-developed, but partially obscured by pigmented cuticle.

In C. heteroceratum, the first 3 segments of Antenna 1 lack conspicuous setae (photo, California Academy of Sciences 2002). In males, the first segment has one (sometimes two) medial teeth on the inner surface. Antenna 2 is proportionately larger in males than females, extending well beyond antenna 1. In males, segment 2 bears a large tooth-like excretory spout, while segment 4 has a median denticle and a variably pointed distal tooth. In females, Antenna 2 is about equal to Antenna 1. The distal end of segment 4 in females lacks a distal tooth, but has a medial spine.

The gnathopods are not especially prominent in this genus. Pereiopod 2 is longer than Pereiopod 21 and its dactyl lacks teeth. Pereiopod 7 is nearly twice as long as Pereiopod 6, and has a long curved, smooth, dactyl. The segments of the urosome are separate. Uropod 1 is the longest and 3 is the smallest. Individuals can reach a length of 9 mm (Chapman 2007). The centers of the pleonites are mottled brown and white, but the borders, and the appendages are white (photo, in California Academy of Sciences 2002). Description based on Chapman 2007.


Taxonomic Tree

Kingdom:   Animalia
Phylum:   Arthropoda
Subphylum:   Crustacea
Class:   Malacostraca
Subclass:   Eumalacostraca
Superorder:   Peracarida
Order:   Amphipoda
Suborder:   Gammaridea
Family:   Corophiidae
Genus:   Corophium
Species:   heteroceratum


Corophium heteroceratum (Yu, 1938)
Sinocorophium heteroceratum (Bousfield and Hoover, 1997)

Potentially Misidentified Species

Corophium alienensis



Corophium heteroceratum is a tube-building gammarid amphipod found in muddy substrates of estuaries and coastal waters (Chapman 1988; Cohen and Carlton 1995; Graening et al. 2012). Gammarid amphipods have separate sexes, brooded embryos, and direct development (Bousfield 1973). We have no specific information on the life history of C. heteroceratum. Nocturnal migration is common in the genus (Chapman 1988).

Corophium heteroceratum is native to the coast of northern China, with a wide temperature range (Ren and Liu 2014). It does not range as far into brackish water as C. alienense, but does penetrate into lower Suisun Bay. It is more abundant at salinities above 15 PSU, in the basins of San Francisco Bay proper (Cohen and Carlton 1995; Lee et al. 2003; Peterson and Vaysierres 2010). Corophium heteroceratum builds U-shaped tubes in shallow subtidal muddy substrates (Lee et al. 2003; Graening et al. 2012). Corophium heteroceratum is probably capable of both suspension-feeding and surface-deposit-feeding. Fishes are the primary predators of benthic amphipods in the San Francisco estuary (Feyrer et al. 2003).


Phytoplankton, detritus

Trophic Status:

Suspension Feeder



General HabitatUnstructured BottomNone
Salinity RangeMesohaline5-18 PSU
Salinity RangePolyhaline18-30 PSU
Salinity RangeEuhaline30-40 PSU
Tidal RangeSubtidalNone
Vertical HabitatEpibenthicNone
Vertical HabitatEndobenthicNone

Tolerances and Life History Parameters

Minimum Salinity (‰)4.9Field data. Mean salinity, Fresh-Brackish-Estuarine Transtion Zone, Suisun Bay (Lee et al. 2003)
Maximum Salinity (‰)35Average marine salinity
Maximum Length (mm)9Chapman 2007)
Broad Temperature RangeNoneCold temperate-Warm temperate
Broad Salinity RangeNoneMesohaline-Euhaline

General Impacts

Corophium heteroceratum is an abundant benthic amphipod in the more saline parts of San Francisco Bay (Lee et al. 2003; Peterson and Vayssierres 2010). Its feeding habits are not well known, but many species of Corophium amphipods are capable of both feeding on surface deposits and suspended phytoplankton. It is potentially an important grazer on phytoplankton and as a food for fishes.

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
NWP-4a None 0 Native Estab
NEP-V Northern California to Mid Channel Islands 1986 Def Estab
NEP-VI Pt. Conception to Southern Baja California 1990 Def Estab
P050 San Pedro Bay 1990 Def Estab
P090 San Francisco Bay 1986 Def Estab
P093 _CDA_P093 (San Pablo Bay) 1986 Def Estab
P110 Tomales Bay 2001 Def Estab

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude


Bousfield, E. L.; Hoover, P. M. (1997) The amphipod superfamily Corophioidea on the Pacific coast of North America. Part V. Family Corophiidae: Corophiinae, new subfamily. Systematics and distributional ecology., Amphipacifica 2(3): 67-139

California Academy of Sciences 2002 <i>Corophium heteroceratum</i> Yu, 1938. <missing URL>

Chapman, John W. (1988) Invasions of the Northeast Pacific by Asian and Atlantic Gammaridean amphipod crustaceans, including a new species of Corophium, Journal of Crustacean Biology 8(3): 364-382

Chapman, John W. (2007) The Light and Smith Manual: Intertidal invertebrates from Central California to Oregon (4th edition), University of California Press, Berkeley CA. Pp. 545-611

Cohen, Andrew N. and 10 authors (2005) <missing title>, San Francisco Estuary Institute, Oakland CA. Pp. <missing location>

Cohen, Andrew N.; Carlton, James T. (1995) Nonindigenous aquatic species in a United States estuary: a case study of the biological invasions of the San Francisco Bay and Delta, U.S. Fish and Wildlife Service and National Sea Grant College Program (Connecticut Sea Grant), Washington DC, Silver Spring MD.. Pp. <missing location>

Fairey, Russell; Dunn, Roslyn; Sigala, Marco; Oliver, John (2002) Introduced aquatic species in California's coastal waters: Final Report, California Department of Fish and Game, Sacramento. Pp. <missing location>

Graening, G. O.; Rogers, D. Christopher; Holsinger, John R.; Barr, Cheryl; Bottorff, Richard (2012) Checklist of inland aquatic Amphipoda (Crustacea: Malacostraca) of California, Zootaxa 3544: 1-27

Holman, Luke E. de Bruyn, Mark Creer, Simon Carvalho, Gary Robidart, Julie; Rius, Marc (2019) Detection of introduced and resident marine species using environmental DNA metabarcoding of sediment and water, Scientific Reports 9: Published online

Huang, Zongguo (Ed.), Junda Lin (Translator) (2001) Marine Species and Their Distributions in China's Seas, Krieger, Malabar, FL. Pp. <missing location>

Jimenez, H.; Ruiz, G. M. (2016) Contribution of non-native species to soft-sediment marine community structure of San Francisco Bay, California, Biological Invasions Publshed online: <missing location>

Lee, Henry II; Thompson, Bruce; Lowe, Sarah (2003) Estuarine and scalar patterns of invasion in the soft-bottom benthic communities of the San Francisco estuary., Biological Invasions 5: 85-102

Peterson, Heather A.; Vayssieres, Marc (2010) Benthic assemblage variability in the upper San Francisco estuary: A 27-year retrospective, San Francisco Estuary and Watershed Science <missing volume>: published online

Ranasinghe, J. Ananda and 6 authors. (2005) The prevalence of non-indigenous species in southern California embayments and their effects on benthic macroinvertebrate communities, Biological Invasions 7: 679-686

Wang, Xiao-Chen and 5 authors (2008) Macrobenthic ecology of the intertidal zones of Chajiandao, Dakouhedao and Wangzidao of Yellow River estuary in Autumn, Chinese Journal of Zoology 21: 77-82