Invasive Species Compendium

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Datasheet

Hemigrapsus sanguineus
(Asian shore crab)

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Datasheet

Hemigrapsus sanguineus (Asian shore crab)

Summary

  • Last modified
  • 08 November 2018
  • Datasheet Type(s)
  • Invasive Species
  • Preferred Scientific Name
  • Hemigrapsus sanguineus
  • Preferred Common Name
  • Asian shore crab
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Arthropoda
  •       Subphylum: Crustacea
  •         Class: Malacostraca
  • Summary of Invasiveness
  • H. sanguineus is a relatively small intertidal shore crab native to cobble/boulder coastlines in the western Pacific Ocean, from Hong Kong Island to Sakhalin Island (China, Japan, Korea, Russia, 22°...

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Pictures

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PictureTitleCaptionCopyright
Hemigrapsus sanguineus (Asian shore crab); adult.
TitleAdult
CaptionHemigrapsus sanguineus (Asian shore crab); adult.
Copyright©Identity unknown-2017/via wikipedia - CC BY-SA 3.0
Hemigrapsus sanguineus (Asian shore crab); adult.
AdultHemigrapsus sanguineus (Asian shore crab); adult.©Identity unknown-2017/via wikipedia - CC BY-SA 3.0
Hemigrapsus sanguineus (Asian shore crab); adult male. Dorsal view.
TitleAdult male
CaptionHemigrapsus sanguineus (Asian shore crab); adult male. Dorsal view.
Copyright©Andrew M. Lohrer
Hemigrapsus sanguineus (Asian shore crab); adult male. Dorsal view.
Adult maleHemigrapsus sanguineus (Asian shore crab); adult male. Dorsal view. ©Andrew M. Lohrer
Hemigrapsus sanguineus (Asian shore crab); adult male. Anterior view from above (with USA 5c coin for scale = 21mm).
TitleAdult male
CaptionHemigrapsus sanguineus (Asian shore crab); adult male. Anterior view from above (with USA 5c coin for scale = 21mm).
Copyright©Andrew M. Lohrer
Hemigrapsus sanguineus (Asian shore crab); adult male. Anterior view from above (with USA 5c coin for scale = 21mm).
Adult maleHemigrapsus sanguineus (Asian shore crab); adult male. Anterior view from above (with USA 5c coin for scale = 21mm).©Andrew M. Lohrer

Identity

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Preferred Scientific Name

  • Hemigrapsus sanguineus (De Haan, 1853)

Preferred Common Name

  • Asian shore crab

International Common Names

  • English: Japanese shore crab

Local Common Names

  • Japan: isogani
  • Netherlands: blaasjeskrab
  • USA: Western Indo-Pacific crab

Summary of Invasiveness

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H. sanguineus is a relatively small intertidal shore crab native to cobble/boulder coastlines in the western Pacific Ocean, from Hong Kong Island to Sakhalin Island (China, Japan, Korea, Russia, 22°N to 49° N) (Sakai, 1976). In September 1988, it was found in Townsends Inlet, New Jersey, USA (Williams and McDermott, 1990). Since then it has spread rapidly northward, into eastern Connecticut and Rhode Island in the late 1990s, and all the way to Maine in the early 2000s. There has also been some southerly spread into North Carolina. There have been no reports of this species south of Cape Hatteras (it is absent in South Carolina, Georgia and Florida). H. sanguineus is also found to be invasive in Europe (Dauvin et al., 2009). It is considered invasive because it is able to achieve extremely high densities, with apparent negative impacts on small recruits and juveniles of several native species (barnacles, littorine snails, brachyuran crabs, mytilid bivalves) (Lohrer and Whitlatch, 2002a,b).

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Arthropoda
  •             Subphylum: Crustacea
  •                 Class: Malacostraca
  •                     Subclass: Eumalacostraca
  •                         Order: Decapoda
  •                             Suborder: Pleocyemata
  •                                 Family: Varunidae
  •                                     Genus: Hemigrapsus
  •                                         Species: Hemigrapsus sanguineus

Description

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The carapace of H. sanguineus ranges from green through to purple, orange-brown or red. It has shaded bands on its legs and red spots on its claws. A relatively small, square-shelled crab with a carapace width (CW) of 35-40 mm. Distinguishing features include the presence of three spines on each side of the carapace and in males there is a fleshy, bulb-like structure on the claws (USGS, 2002).

Distribution

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Native to cobble/boulder coastlines in the western Pacific Ocean, from Hong Kong Island to Sakhalin Island (China, Japan, Korea, Russia, 22°N to 49° N) (Sakai, 1976), and introduced as an exotic in the USA as a result of ship and boat hull fouling. In Europe, H. sanguineus ranges from the western part of the English Channel to Germany (Dauvin et al., 2009). It was first found in Sweden on the Gothenburg archipelago in 2012 (Berggren, 2013).

 

Distribution Table

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The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Sea Areas

Atlantic, NorthwestWidespreadIntroduced1988 Invasive Lohrer and Whitlatch, 2002a; Lohrer and Whitlatch, 2002b; Lohrer et al., 2000bExtremely high densities in the late 1990s/ early 2000s in eastern Connecticut and Rhode Island, USA. Long Island Sound.
Pacific, NorthwestWidespreadNative Not invasive Sakai, 1976Wide latitudinal range, relatively common and conspicuous shore crab

Asia

ChinaPresentNative Not invasive Sakai, 1976
-Hong KongPresentNative Not invasive Sakai, 1976; Depledge, 1984
JapanPresentPresent based on regional distribution.
-HokkaidoPresentNative Not invasive Sakai, 1976; Takahashi et al., 1985
-HonshuPresentNative Not invasive Sakai, 1976; Fukui and Wada, 1983; Fukui, 1988
-KyushuPresentNative Not invasive Sakai, 1976; Takada and Kikuchi, 1991
-ShikokuPresentNative Not invasive Sakai, 1976
Korea, Republic ofPresentNative Not invasive Hwang et al., 1993

North America

-ConnecticutPresentIntroducedmid-1990s Invasive Lohrer and Whitlatch, 2002a; Lohrer and Whitlatch, 2002b; Lohrer et al., 2000b
-DelawarePresentIntroducedlate 1980s/early 1990s Invasive Epifanio et al., 1998
-MainePresentIntroduced2001 Invasive Lohrer, 2001
-MassachusettsPresentIntroducedlate 1990s Invasive Ledesma and O'Connor, 2001
-New HampshirePresentIntroducedlate 1990s Invasive Tyrrell and Harris, 2000
-New JerseyPresentIntroducedlate 1980s/early 1990s Invasive Williams and McDermott, 1990First recorded in Townsends Inlet, New Jersey in 1988
-Rhode IslandPresentIntroducedmid/late 1990s Invasive Lohrer and Whitlatch, 2002a; Lohrer and Whitlatch, 2002b; Lohrer et al., 2000b

Europe

BelgiumPresentIntroducedDauvin et al., 2009
CroatiaPresentIntroducedSchubert, 2003
FrancePresentIntroducedBreton et al., 2002
GermanyPresentIntroducedDauvin et al., 2009
NetherlandsPresentIntroducedBreton et al., 2002
Russian FederationPresentPresent based on regional distribution.
-Russian Far EastPresentNative Not invasive Sakai, 1976
SwedenPresent, few occurrencesIntroduced2012Berggren, 2013First occurance 21/07/2012, 57.691735°N 11.635355°E. Five more individuals have been sighted since then along the Gothenburg archipelago.

History of Introduction and Spread

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The introduction of H. sanguineus to the USA was apparently accidental, possibly associated with ocean-going vessel traffic. Planktonic life-stages (zoeae and megalopae) may have arrived in ballast water in 1985 or earlier. Small H. sanguineus instars could have gone undetected at first, especially if initial densities were below a reasonable detection limit. In contrast, the transport of adult stages could have occurred as late as 1988 [from Lohrer, 2001].

Introductions

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Introduced toIntroduced fromYearReasonIntroduced byEstablished in wild throughReferencesNotes
Natural reproductionContinuous restocking
USA Asia 1985-1988 Yes Williams and McDermott (1990) Ship/boat hull fouling. Several individuals, including gravid females discovered in 1988

Habitat

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In New England, H. sanguineus is generally found in rocky intertidal habitats (Lohrer and Whitlach, 2002a). Lohrer (2001) reports that: “Susan Park at University of Delaware finds the crabs on several types of human-constructed structures in this region (rip-rap, jetties, bulkheads, wooden piers, groins, etc.). Apparently, the crabs survive in enclosed harbours as well as on exposed coast. In addition, both intertidal and subtidal oyster reefs support populations of the invasive crabs. To summarize, H. sanguineus colonizes a variety of hard substrates in the southeastern USA despite the absence of intertidal cobble habitats".

See Dauvin et al. (2009) for further information on habitat preferences in Europe.

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Multiple
Littoral
Coastal areas Principal habitat
Intertidal zone Principal habitat
Salt marshes Secondary/tolerated habitat
Brackish
Estuaries Secondary/tolerated habitat
Marine
 
Benthic zone Secondary/tolerated habitat

Biology and Ecology

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Reproductive Biology

Fully grown H. sanguineus females can likely produce >50,000 eggs at a time (Fukui, 1988; McDermott 1998b). The eggs of H. sanguineus are of intermediate size (0.015 mm3), relative to other co-occurring grapsid crabs in Japan (Fukui, 1988). The eggs hatch into larvae, which proceed to molt through five zoeal stages to become megalopae (Hwang et al., 1993). Megalopae can be found in the water column or on the benthos (Hwang et al., 1993; Lohrer, 2000). After settlement and metamorphosis, the first crab stage is usually between 1.6 and 2.0 mm carapace width (Epifanio et al., 1998; Lohrer, 2000). The time from hatching to first instar undoubtedly varies with latitude, but takes 25+ days under optimal conditions (warm water of high salinity) (Hwang et al., 1993; Epifanio et al., 1998) [from Lohrer, 2001].

Physiology and Phenology

In temperate areas, females first begin to produce eggs in the spring (Takahashi et al., 1985; Fukui, 1988; Lohrer and Whitlatch, 1997; Saigusa and Kawagoye, 1997; Ledesma and O’Connor, 2001). Gravid females release their eggs when water temperatures are warm and larval settlement usually occurs in late summer and fall (Fukui, 1988; Lohrer, 2000). Settled crabs grow rapidly at first, probably molting four or five times in their first month on the benthos (Epifanio et al., 1998). Although winter temperatures likely slow growth considerably, the growth rate of these crabs diminishes after maturity regardless of temperature effects (Fukui, 1988) [from Lohrer, 2001].

Nutrition

There were no explicit investigations of the diet of native H. sanguineus populations until the late 1990s (Lohrer et al., 2000b). Most Japanese researchers assumed that the crab was largely herbivorous. Depledge (1984) observed H. sanguineus grazing algae from rock surfaces, but never performed gut content analyses. In central Japan, H. sanguineus often picks at Caulacanthus okamurai, a red algal turf attached to large cobbles in intertidal areas. However, several small invertebrate species live inside the matrix formed by stout filaments of the alga. It is not known whether the crab incidentally consumes the invertebrates with the alga, or whether it targets animal food items specifically. In any case, as much as 40% of the diet of H. sanguineus is composed of small invertebrates (including mussels, snails, amphipods, and polychaetes).

In North America, several investigators have reported the regular consumption of animal food items by H. sanguineus (Lohrer and Whitlatch, 1997; McDermott, 1999; Lohrer et al., 2000b; Brousseau et al., 2001; Ledesma and O’Connor, 2001). Males of this species have larger claws than females and can consume larger prey items (McDermott, 1999; Brousseau et al., 2001). However, both males and females apparently focus on relatively small prey items (i.e. species with small maximum sizes and/or recruits and juveniles of larger species) (Lohrer et al., 2000b). Although taxonomic differences obviously exist between northeastern Asia and northeastern North America, many of the same types of food items are encountered by H. sanguineus in both regions (e.g. ephemeral green algae, mytilid bivalves, littorinid snails, gammarid amphipods, nereid worms). Several species of intertidal barnacle exist in central Japan, but none were observed inside native H. sangineus guts (Lohrer et al., 2000b). However, in North America, barnacles are regularly found in the guts of wild-caught crabs (Lohrer and Whitlatch, 1997; McDermott 1999; Lohrer et al., 2000b; Ledesma and O’Connor 2001). The barnacles and molluscs that H. sanguineus consumes in eastern North America may have softer shells than their Asian counterparts, and may be more susceptible to predation by invasive shore crabs (Lohrer et al., 2000b) [from Lohrer, 2001].

Latitude/Altitude Ranges

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Latitude North (°N)Latitude South (°S)Altitude Lower (m)Altitude Upper (m)
49 22

Air Temperature

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Parameter Lower limit Upper limit
Mean maximum temperature of hottest month (ºC) 0 30
Mean minimum temperature of coldest month (ºC) -10 0

Water Tolerances

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ParameterMinimum ValueMaximum ValueTypical ValueStatusLife StageNotes
Salinity (part per thousand) 33 35 Optimum 25-35 tolerated. Larval tolerances differ to those of adults
Turbidity (JTU turbidity) Optimum Does not live in really muddy places
Water temperature (ºC temperature) Optimum 5-30 tolerated

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Sacculina Parasite Adult to species Yamaguchi and Aratake, 1997

Notes on Natural Enemies

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In Asia, a rhizocephalan parasite (Sacculina polygenea) attacks H. sanguineus and prevalence can be as high as 80% in some locations (Yamaguchi and Aratake, 1997).

Impact Summary

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CategoryImpact
Environment (generally) Negative

Economic Impact

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The apparent replacement of Carcinus maenas [European green crab] by H. sanguineus has ecological and economic implications. Although both crabs are non-indigenous, green crabs are pests that impact shellfish resources throughout New England. Green crabs consume more animal matter than H. sanguineus (Elner, 1981; Ropes, 1989; Lohrer and Whitlatch, 1997; Lohrer et al., 2000b) and present a significant selective pressure on a variety of snails, clams, and mussels (Vermeij, 1982a,b; Leonard et al., 1999; Trussell and Smith, 2000; Smith and Jennings, 2000; Whitlow and Dochtermann, 2001). On a per capita basis, green crabs consume more juvenile blue mussels (Mytilus edulis) than H. sanguineus (Lohrer and Whitlatch, 2002b). However, since the densities of H. sanguineus are often 60-80 times greater than those of C. maenas, the collective effects of H. sanguineus are now more important to juvenile blue mussels in rocky intertidal habitats. The invasion by H. sanguineus probably represents a net negative influence on blue mussel populations in the rocky intertidal zone, despite the fact that C. maenas (a more voracious predator) is relatively rare in this habitat type now [from Lohrer, 2001].

Environmental Impact

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Impact on Biodiversity

Lohrer and Whitlach (2002a) examined the relationship between the European green crab (Carcinus maenas) and H. sanguineus in New England over a 4-year period. During the study, a sharp decline in C. maenas was observed that coincided with an increase in H. sanguineus populations. Lohrer and Whitlach (2002b) reported that in less than 10 years after its appearance in Long Island Sound, H. sanguineus had become the numerically dominant species over C. maenas.

Risk and Impact Factors

Top of page Invasiveness
  • Proved invasive outside its native range
  • Has a broad native range
  • Abundant in its native range
  • Highly adaptable to different environments
  • Capable of securing and ingesting a wide range of food
  • Highly mobile locally
  • Long lived
  • Fast growing
  • Has high reproductive potential
  • Gregarious
Impact outcomes
  • Altered trophic level
  • Ecosystem change/ habitat alteration
  • Modification of natural benthic communities
  • Monoculture formation
  • Reduced native biodiversity
  • Threat to/ loss of native species
Impact mechanisms
  • Competition - monopolizing resources
  • Herbivory/grazing/browsing
  • Interaction with other invasive species
  • Predation
Likelihood of entry/control
  • Highly likely to be transported internationally accidentally
  • Difficult/costly to control

Similarities to Other Species/Conditions

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Hemigrapsus is a common genus present in many coastal areas all over the world. Crabs most similar to H. sanguineus include Hemigrapsus penicillatus (which some have said might be a new species called Hemigrapsus takanoi, see Asakura et al. (2005) for further details). The differences between H. sanguineus and H. penicillatus are described by Breton et al. (2002).

References

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Asakura A; Watanabe S, 2005. Hemigrapsus takanoi, a new species, a sibling species of the common Japanese intertidal crab H. penicillatus (Decapoda: Brachyura: Grapsidae). Journal of Crustacean Biology, 25(2):279-292.

Berggren; Matz S, 2013. New discoveries of crustaceans on the Swedish west coast. (Nya fynd av kräftdjur vid den svenska västkusten.) Fauna & Flora, 108(3):42-44.

Breton G; Faasse M; Noel P; Vincent T, 2002. A new alien crab in Europe, Hemigrapusus sanguineus (Brachyurea: Grapsidae). Journal of Crustacean Biology, 22(1):184-189.

Brousseau DJ; Filipowicz A; Baglivo; JA, 2001. Laboratory investigations of the effects of predator and size on prey selection by the Asian crab, Hemigrapsus sanguineus. Journal of Experimental Marine Biology and Ecology, 262:199-210.

Brousseau DJ; Korchari PG; Pflug C, 2000. Food preference studies of the Asiatic shore crab (Hemigrapsus sanguineus) from western Long Island Sound. In: Marine Bioinvasions: Proceedings of the First National Conference, Cambridge, MA, 24-27 January 1999 [ed. by Pederson J] Cambridge, MA, USA: MIT Sea Grant College Program, 200-207.

Dauvin JC; Rius AT; Ruellet T, 2009. Recent expansion of two invasive crabs species Hemigrapsus sanguineus (de Haan, 1835) and H. takanoi Asakura and Watanabe 2005 along the Opal Coast, France. Aquatic Invasions, 4(3):451-465. http://www.aquaticinvasions.ru/2009/AI_2009_4_3_Dauvin_etal.pdf

Depledge MH, 1984. Cardiac activity in the intertidal crab Hemigrapsus sanguineus (de Haan). Asian Marine Biology, 1:115-123.

Elner RW, 1981. Diet of green crab Carcinus maenas (L.) from Port Hebert, southwestern Nova Scotia. Journal of Shellfish Research, 1(1):89-94.

Epifanio CE; Dittel AI; Park S; Schwalm S; Fouts A, 1998. Early life history of Hemigrapsus sanguineus, a non-indigenous crab in the Middle Atlantic Bight. Marine Ecology Progress Series, 170:231-238.

Fukui Y, 1988. Comparative studies on the life history of the grapsid crabs (Crustacea, Brachyura) inhabiting intertidal cobble and boulder shores. Publication of the Seto Marine Biological Laboratory, 33((4/6)):121-162.

Fukui Y; Wada K, 1983. Intertidal Anomura and Brachyura and their distributions on the south coast of Tanabe Bay. The Nanki Biological Society, 25(2):159-167.

Hwang SG; Lee C; Kim CH, 1993. Complete larval development of Hemigrapsus sanguineus (Decapoda, Brachyura, Grapsidae) reared in laboratory. Korean Journal of Systematic Zoology, 9(2):69-86.

Ledesma ME; O'Connor NJ, 2001. Habitat and diet of the non-native crab Hemigrapsus sanguineus in southeastern New England. Northeastern Naturalist, 8(1):63-78.

Leonard GH; Bertness MD; Yund PO, 1999. Crab predation, waterborne cues, and inducible defenses in the blue mussel, Mytilus edulis. Ecology, 80:1-14.

Lohrer AM, 2000. Mechanisms and consequences of an exotic crab species invasion. PhD thesis. Storrs, Connecticut, USA: Department of Ecology and Evolutionary Biology, University of Connecticut.

Lohrer AM, 2001. The invasion by Hemigrapsus sanguineus in eastern North America: a review. Aquatic Invaders, 12(3):1-11.

Lohrer AM; Fukui Y; Wada K; Whitlatch RB, 2000. Structural complexity and vertical zonation of intertidal crabs, with focus on the habitat requirements of the invasive Asian shore crab, Hemigrapsus sanguineus (de Haan). Journal of Experimental Marine Biology and Ecology, 244:203-217.

Lohrer AM; Wada K; Fukui Y; Whitlatch RB, 2000. Home and Away: comparisons of resource utilization by a marine species in native and invaded habitats. Biological Invasions, 2(1):41-57.

Lohrer AM; Whitlatch RB, 1997. Ecological studies on the recently introduced Japanese shore crab (Hemigrapsus sanguineus), in eastern Long Island Sound. In: Proceedings of the Second Northeast Conference on Nonindigenous Aquatic Nuisance Species, Burlington, VT, 18-19 April 1997 [ed. by Balcom N] Groton, Connecticut, USA: Connecticut Sea Grant College Program, 49-60.

Lohrer AM; Whitlatch RB, 2002. Interactions among aliens: apparent replacement of one exotic species by another. Ecology, 83(3):719-732.

Lohrer AM; Whitlatch RB, 2002. Relative impacts of two exotic brachyuran species on blue mussel populations in Long Island Sound. Marine Ecology Progress Series, 227:135-144.

Lohrer AM; Whitlatch RB; Wada K; Fukui Y, 2000. Using niche theory to understand invasion success: a case study of the Asian shore crab, Hemigrapsus sanguineus. In: Marine Bioinvasions: Proceedings of the First National Conference, Cambridge, MA, 24-27 January 1999 [ed. by Pederson J] Cambridge, MA, USA: MIT Sea Grant College Program, 57-60.

McDermott JJ, 1998. The western Pacific brachyuran Hemigrapsus sanguineus (Grapsidae) in its new habitat along the Altlantic coast of the Untied States: reproduction. Journal of Crustacean Biology, 55(2):289-298.

McDermott JJ, 1998. The western Pacific brachyuran Hemigrapsus sanguineus (Grapsidae), in its new habitat along the Altlantic coast of the Untied States: geographic distribution and ecology. ICES Journal of Marine Science, 55(2):289-298.

McDermott JJ, 1999. The western Pacific brachyuran Hemigrapsus sanguineus (Grapsidae) in its new habitat along the Atlantic coast of the United States: feeding, cheliped morphology and growth. In: Crustaceans and the biodiversity crisis. Proceedings of the Fourth International Crustacean Congress, Amsterdam, the Netherlands, 20-24 July 1998 [ed. by Schram F, von Vaupel Klein J] Brill, Leiden, The Netherlands, 425-444.

Ropes JW, 1989. The food habits of five crab species at Pettaquamscutt River, Rhode Island. US Fish and Wildllife Service Fisheries Bulletin, 87:197-204.

Saigusa M; Kawagoye O, 1997. Circatidal rhythm of an intertidal crab, Hemigrapsus sanguineus: synchrony with unequal tide height and involvement of a light response mechanism. Marine Biology, 129:87-96.

Sakai T, 1976. Crabs of Japan and the Adjacent Seas. Tokyo, Japan: Kodansha Ltd.

Schubert CD, 2003. Scientia Marina., 195-200.

Smith LD; Jennings JA, 2000. Induced defensive responses by the bivalve Mytilus edulis to predators with different attack modes. Marine Biology, 136:461-469.

Takada Y; Kikuchi T, 1991. Seasonal and vertical variation of the boulder shore fauna in Amakusa. Publication from the Amakusa Marine Biological Laboratory, 11(1):1-17.

Takahashi K; Miyamoto T; Mizutori Y; Ito M, 1985. Ecological studies on rocky shore crabs in Oshoro Bay. Scientific Reports of the Hokkaido Fisheries Experimental Station, 27:71-89.

Trussell GC; Smith LD, 2000. Induced defenses in response to an invading crab predator: an explanation of historical and geographic phenotypic change. Proceedings of the National Academy of Sciences, 97:2123-2127.

Tyrrell MC; Harris LG, 2000. Potential impact of the introduced Asian shore crab, Hemigrapsus sanguineus, in Northern New England: diet, feeding preferences, and overlap with the green crab, Carcinus maenas. In: Marine Bioinvasions: Proceedings of the First National Conference, Cambridge, MA, 24-27 January 1999 [ed. by Pederson J] Cambridge, MA, USA: MIT Sea Grant College Program, 208-220.

USGS, 2002. Non-indigenous species information bulletin: Hemigrapsus sanguineus. http://fl.biology.usgs.gov/hemigrapsus.pdf

Vermeij GJ, 1982. Environmental change and the evolutionary history of the periwinkle (Littorina littorea) in North America. Evolution, 36:561-580.

Vermeij GJ, 1982. Phenotypic evolution in a poorly dispersing snail after the arrival of a predator. Nature, 299:349-350.

Whitlow L; Dochtermann J, 2001. Phenotypic plasticity of native soft-shell clams in response to chemical and physical stimuli from invasive green crab predation. In: International Conference on Marine Bioinvasions, New Orleans, LA, 9-11 April 2001.

Williams AB; McDermott JJ, 1990. An eastern United States record for the Western Indo-Pacific crab, Hemigrapsus sanguineus (Crustacea: Decapoda: Grapsidae). Proceedings of the Biological Society of Washington, 103(1):108-109.

Yamaguchi T; Aratake H, 1997. Morphological modifications caused by Sacculina polygenea in Hemigrapsus sanguineus (De Haan)(Brachyura: Grapsidae). Crustacean Research, 26:125-145.

Contributors

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23/06/08 Original text by:

Andrew Lohrer, NIWA, Gate 10 Silverdale Road, Hamilton, 3216, New Zealand

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