Invasion History

First Non-native North American Tidal Record: 2013
First Non-native West Coast Tidal Record:
First Non-native East/Gulf Coast Tidal Record: 2013

General Invasion History:

Peppermint shrimps of the genus Lysmata are part of a complex of ~48 species, widely distributed in tropical and subtropical marine waters.  Lysmata vittata was described from Hong Kong by William Stimpson in 1860, and has been reported from Pacific Russia (Korn 2012; Marin et al. 2012a; Marin et al. 2012b) to the Red Sea and Persian Gulf, to eastern South Africa, as well as Singapore (Anker and DeGrave 2016); Australia (Barton et al. 2020) and Amboina, Indonesia, and New Guinea (Kubo 1951; Bruce 1976; Holthuis 1978;  cited by WoRMS Editorial Board (2022); US National Museum of Natural History 2022). However, a recent integrative taxonomic analysis found that Lysmata vittata is a complex of multiple species with the native range of Lysmata vittata sensu stricto likely spanning the temperate/subtropical Indo-West Pacific from China to southern Russia (Aguilar et al. 2022). Lysmata vittata sensu lato is considered introduced in many parts of the world, including Brazil (Alves et al. 2008; Soledade et al. 2013; Guéron et al. 2022); Caribbean Panama (Pachelle et al. 2020); New Zealand (Ahyong 2010); the Mediterranean Sea (Abdelsalam 2008); and the eastern United States (Aguilar et al. 2022). However, a careful morphological and genetic analysis indicated many records of 'Lysmata vittata' refer to other species, either L. rauli (a previous junior synonym of L. vittata) or L. cf. rauli (an undescribed sister species of L. rauli) (Aguilar et al. 2022; Guéron et al. 2022). Specifically, specimens from the Chesapeake region, Hong Kong, Taiwan and New Zealand, representing L. vittata sensu stricto were highly similar and distinct from other specimens from Hong Kong, Thailand, Panama, and Brazil (L. rauli sensu stricto) and Australia (L. cf. rauli) (Aguilar et al. 2022; Guéron et al. 2022).

In Chesapeake Bay and adjacent Atlantic waters, Lysmata vittata has been collected from the mouth to south of Tangier Island, the lower reaches of the James and York estuaries, and also in Burtons Bay, a nearby Atlantic embayment. It was collected at temperatures of 4.5 to 26.5 ºC and salinities of 18.5 to 34.5 PSU. Habitats include shallow oyster flats (~1 m) to deeper muddy channels (19.5 m). Lysmata vittata co-occurred with L. wurdemanni at many locations (Aguilar et al. 2022). Specimens of L. vittata have also been collected from Little Egg Harbor, New Jersey, Hudson River, New York, and Long Island Sound, near Norwalk, Connecticut (Robert Aguilar, personal communication; James Carlton, personal communication 2022). Lysmata vittata has also be found in New Zealand near port of Auckland (Ahyong 2010).

The likeliest introduction vector for Lysmata vittata sensu stricto in the United States and New Zealand (as well as L. rauli in Central/South America) is ballast water, although several species marketed as Lysmata vittata are sold in the aquarium trade.


North American Invasion History:

Invasion History on the East Coast:

In 2013 to 2018, 139 specimens identified as the Indo-Pacific Lysmata vittata were collected in Chesapeake Bay and in adjacent Atlantic waters, including the mouth of the Bay to south of Tangier Island, the lower reaches of the James and York estuaries, and Burtons Bay, a nearby Atlantic embayment. It was collected at temperatures of 4.5 to 26.5 ºC and salinities of 18.5 to 34.5 PSU. Lysmata vittata co-occurred with L. wurdemanni, a peppermint shrimp native to the East and Gulf Coasts from Long Island Sound to Texas (Williams 1984; Rhyne and Lin 2006), at many locations (Aguilar et al. 2022).  Specimens of L. vittata have also been collected from Little Egg Harbor, New Jersey, and Long Island Sound, near Norwalk, Connecticut, and the Hudson River, New York (Robert Aguilar, personal communication; James Carlton, personal communication 2022).


Invasion History Elsewhere in the World:

In 2006, specimens of Lysmata vittata were collected from the east and west coasts of the northern peninsula of the North Island of New Zealand (Ahyong 2010).  The shrimps were collected in marinas in harbor locations (Ahyong 2010).  Records from Egypt (Abdelsalam 2008) and Brazil (Soledade et al. 2013; Alves et al. 2018; Pachellet et al. 2020) are believed to refer to closely related species (Aguilar et al. 2022).


The peppermint shrimp Lysmata vittata (East Asian Lined Shrimp; Red-Striped Shrimp) is one of ~48 named species within Lysmata, which includes obligate cleaner shrimps with distinctive coloration, and free-living omnivorous shrimps. Lysmata vittata is a small shrimpreaching total lengths of 18–30 mm. The body is semi-transparent, and marked by fine red longitudinal stripes varying in pigment density  Marin et al. 2021a; Aguilar et al. 2022). The rostrum is thin and reaches the second article of the dorsal peduncle. The rostrum has 6–8 dorsal teeth and 2–6 ventral teeth (Aguilar et al. 2022). There is a pterygostomial tooth on the buccal edge of the carapace. The dorsal antennule is uniramous, lacking an accessory branch. The antennule outer edge bears a short stylocerite (spiny process) which reaches the mid-length of the peduncle.  The antennal scale is longer than the peduncle and is three times as long as it is wide (Marin et al. 2012a; Agular et al. 2022). The second pereiopod has 18–23 articles (Aguilar et al. 2022).

Specimens reported from Australia, Brazil, Panama, and Mediterranean Egypt refer to other species once grouped in the Lysmata vittata species complex (Aguilar et al. 2022; Guéron et al. 2022). These species vary in some of the details listed here, including number of pereiopod articles, accessory branch status, or coloration. Some of these correspond to Lysmata rauli (Laubenheimer & Rhyne 2010) described from Brazil), but assumed native to the eastern Pacific , as well as L. cf. rauli also found in Hong Kong and Egypt (Aguilar et al. 2022).




Taxonomic Tree

Kingdom:   Animalia
Phylum:   Arthropoda
Subphylum:   Crustacea
Class:   Malacostraca
Subclass:   Eumalacostraca
Superorder:   Eucarida
Order:   Decapoda
Suborder:   Pleocyemata
Infraorder:   Caridea
Superfamily:   Alpheoidea
Family:   Lysmatidae
Genus:   Lysmata
Species:   vittata


Hypolysmata vittata (Stimpson , 1860)
Nauticaris unirecedens (Spence Bate , 1888)
Hippolysmata vittata var. subtilis (Thallwitz, 1891)

Potentially Misidentified Species

Lysmata cf. rauli

Lysmata rauli has been considered a junior synonym of Lysmata vittata, but Aguilar et al. (2022) and Guéron et al. (2022) clearly show they are distinct and resurrect L. rauli to full species status. Both authors also provide strong evidence that Lysamta rauli is a complex of sister species. Lymata vittata sensu stricto can be distinguished from the Lysmata rauli complex by the lack of both an accessory branch on the dorsal antennule and dark transverse bars on the abdomen.

Lysmata rathbunae

Native from Florida to Yucatan, with differing spination and setal counts, and oblique and transverse stripes.

Lysmata wurdemanni

Native from Long Island to Yucatan (WoRMS Editorial Board (2022)). Lysmata wurdemanni is distiguished from L. vittata by many features of spination, setation and segmentation. The carapace has large transverse and oblique red bands (Rhyne and Lin 2000).



The peppermint shrimp Lysmata vittata (East Asian Lined Shrimp; Red-Striped Shrimp) is a marine shrimp, native from temperate to subtropical marine waters. In the Chesapeake Bay, it occurred at temperatures of 4.5 to 26.5 ºC and salinities of 18.5 to 34.5 PSU (Aguilar et al. 2022).  In the Pos'yet Bay, Russia and in the Sea of Japan, this shrimp inhabits a cool-temperate location (Marin et al. 2012a).  This shrimp is reported from a variety of habitats, including shallow oyster flats, and deeper muddy channels (Aguilar et al. 2022); boulders on a sandy bottom (Marin et al. 2012a), the 'under-stone' habitat at the edge of coral reefs (Bruce 1976), and algae and stones in tidepools (Bruce 1976).  Aquarium websites describe this species as an omnivorous, occasionally showing cleaning behavior by removing parasites from fishes, and removing pest species such as the anemone Exaiptasia pallida.



Invertebrates, algae, anemones


Fishes, crabs, anemones


Other shrimps, Lysmata wurdenanni

Trophic Status:



Life History

The peppermint shrimp Lysmata vittata (East Asian Lined Shrimp; Red-Striped Shrimp) is a protandrous simultaneous hermaphrodite, maturing first as a male, and then reproducing as a functional euhermaphrodite, with an ovotestes producing eggs but still mating as a male (Chen et al. 2019). Juveniles become functional males with a carapace length of 3.7 to 7.5 mm; and at larger sizes, 5.8 to 8.5 mm, become euhermaphrodites with both viable eggs and sperm (Chen et al. 2019). After fertilization, the eggs are extruded and carried below the abdomen. The eggs hatch into a planktonic Zoea 1, and molt and grow through 8 molts to a Zoea IX and then to a megalopa. Each zoeal stage took 3-5 days at 28 ºC, for an overall larval development of 24–40 days (Marin et al. 2012a, Yang and Kim 2010). The megalopa molts into a juvenile before becoming a functional male (Marin et al. 2012; Chen et al. 2019).

Note that Chen et al.’s ‘Lysmata vittata ‘, from Xiamen, China could be Lysmata rauli, or another species. Reproductive studies on verified L. vittata are desirable.


Tolerances and Life History Parameters

Minimum Depth (m)0.5Chesapeake Bay, Aguilar et al. 2022
Maximum Depth (m)20Chesapeake Bay, Aguilar et al. 2022
Minimum Salinity (‰)18.5Aguilar et al. 2022
Maximum Salinity (‰)34.5Chesapeake Bay, Aguilar et al. 2022
Minimum Reproductive Temperature25Temperature for larval rearing. Korea, Yang et al. Kim 2010
Maximum Reproductive Temperature28

Temperature for gonad development studies, Taiwan, Chen et al. 2010

Minimum Duration24Lab rearing, 8 zoeal stages, Korea, Yang and Kim 2010
Maximum Duration40Lab rearing, 8 zoeal stages, Korea, Yang and Kim 2010
Minimum Length (mm)18Male, protandrous hermpahrodite (Marin et al. 2012)
Maximum Length (mm)30Mature Euhermaphrodite, Marin et al. 2012
Broad Temperature Range1Cold temperate-Tropical
Broad Salinity Range1Polyhaline-Euhaline

General Impacts

Lysmata vittata has few reported ecological or economic impacts. Shrimp marketed as “L. vittata” are valued in the aquarium trade for their colorful appearance and as a predator on pest anemones (e.g., Exaiptasia pallida) and flatworms (Vaughan et al. 2018; Barton et al. 2020.). However, many of these shrimps may reflect Lysmata rauli or another species and not L. vittata sensu stricto (Aguilar et al. 2022).

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
NWP-2 None 1860 Native Estab
NWP-3a None 0 Native Estab
NWP-4a None 0 Native Estab
NWP-4b None 0 Native Estab
M130 Chesapeake Bay 2013 Def Estab
NA-ET3 Cape Cod to Cape Hatteras 2013 Def Estab
M128 _CDA_M128 (Eastern Lower Delmarva) 2013 Def Estab
NZ-IV None 2006 Def Estab
M080 New Jersey Inland Bays 2022 Def Unk
M040 Long Island Sound 2022 Def Unk
M060 Hudson River/Raritan Bay 2021 Def Unk

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude


Ahyong, Shane T. (2010) New species and new records of Caridea (Hippolytidae: Pasiphaeidae) from New Zealand, Zootaxa 2372(2372): 341-357

Abdelsalam, Khaled Mahmood (2018) First record of the exotic lysmatid shrimp Lysmata vittata (Stimpson, 1860) (Decapoda: Caridea: Lysmatidae) from the Egyptian Mediterranean coast, Mediterranean Marine Science 19(1): 124-131

Aguilar, Robert; Prakash, Sanjeevi; Ogburn, Matthew B.; Pagenkopp Lohan, Katrina M.; MacDonald, Kenneth S. III; Driskell, Amy C.; Ahyong, Shane T.; (2022) Unresolved taxonomy confounds invasive species identification: the Lysmata vittata Stimpson, 1860 (Decapoda: Caridea: Lysmatidae) species complex and recent introduction of Lysmata vittata sensu stricto in the western Atlantic, Journal of Crustacean Biology 42(1): 1-18

Alves, Douglas Ferandes Rodrigues; Marcondes Lima, Daniel Jose, Hiorse, Gustavo Luis; Martinez, Pablo Ariel; Dolabella, Sivio Santana; Barros-Alves, (2018) Morphological and molecular analyses confirm the occurrence of two sympatric Lysmata shrimp (Crustacea, Decapoda) in the southwestern Atlantic, Zootaxa 4526(1): 41-55

Barton, Jonathan A.; Humphrey, Craig; Bourne, David G.; Hutson, Kate S. (2020) Biological controls to manage Acropora-eating flatworms in coral aquaculture, Aquaculture Environment Interactions 12: 61-66

Bruce, A. J. (1976) A report on a small collection of shrimps from the Kenya National Marine Parks from at Malindi with notes on selected species, Zoologische Verhandelingen 145(1): 1-72

Chen, Daming; Liu, Fang; Zhu, Zhihuang; Lin, Qi; Zeng, Chaoshu; Ye, Haihui (2019) Ontogenetic development of gonads and external sexual characters of the protandric simultaneous hermaphrodite peppermint shrimp, Lysmata vittata (Caridea: Hippolytidae), PLOSOne 14(4): e0215406

Gueron, Rodrigo; Almeida, Alexandre Oliveira; Aguilar, Robert; Ogburn, Matthew B.; Prakash, Sanjeevi ; Baeza, J. Antonio (2022) Delimiting species within the Lysmata vittata (Stimpson, 1860) (Decapoda: Lysmatidae) species complex in a world full of invaders, Zootaxa 5150(2): 189–216

Holthuis, L. B. (1978) A collection of decapod crustacea from Sumbs, Lesser Sunda Islands , Zoologische Verhandelingen 162(1): 3-55

Kubo, Isuo (1951) Some Macruruous decapod crustacea found in Japanese waters, with descriptions of four new species. S, Journal of the Tokyo University of Fisheries 28(2): 259-289

Laubenheimer, Helio; Rhyne, Andrew L. (2010) Lysmata rauli, a new species of peppermint shrimp, (Decapoda: Hippolytidae) from the southwestern Atlantic*, Zootaxa 2372(1): 298–304

Marin, Ivan; Korn, Olga N.; Kornienko, Elena S. (2012a) The Caridean Shrimp Lysmata vittata (Stimpson, 1860) (Decapoda: Hippolytidae): A New Species for the Fauna of Russia, Russian Journal of Marine Biology 38(4): 359-363

Marin, Ivan; Korn, Olga N.; Kornienko, Elena S. (2012b) On the presence of the caridean shrimp Lysmata vittata (Decapoda, Hippolytidae, in the Russian waters of the Sea of Japan , Crustaceana 85(1): 123-124

Pachelle, Paulo P.; Carvalho, Lena; , Alves, Douglas F. R.; Anker, Arthur (2020) A revision of the Brazilian species of Lysmata Risso, 1816 (Decapoda: Caridea: Lysmatidae), with discussion of the morphological characters used in their identification, Zootaxa 4789(1): 55-90

Rhyne, Andrew L. ; Lin, Junda (2006) A western Atlantic Peppermint Shrimp complex: Redescription of Lysmata wurdemanni, description of four new species, and remarks on Lysmata rathbunae (Crustacea, Decapoda, Hippolytidae., Bullerin of Marine Science 70(1): 165-204

Santos, Rafael de Carvalho; Alves, Douglas Fernandes Rodrigues; Carvalho-Batista, Abner (2021) Shrimps of genus Lysmata Risso, 1816 (Caridea: Lysmatidae) from Queimada Grande Island region, southeastern Brazil, Nauplius 20(e2021032): 1-29

Soledade, Guidomar Oliveira; Baeza, Juan Antonio; Boehs, Guisla; Sabrina; Simões, Morilhas; Santos, Patricia Souza; da Costa, Rogerio Caetano; Ol (2013) A precautionary tale when describing species in a world of invaders: morphology, coloration and genetics demonstrate that Lysmata rauli is not a new species endemic to Brazil, but a junior synonym of the Indo-Pacific L. vittata, Journal of Crustacean Biology 33(1): 66-77

Stimpson, W. A. (1860) rodromus descriptionis animalium evertebratorum, quae in Expeditione ad Oceanum Pacificum Septemtrionalem, a Republica Federata missa, C. Ringgold et J. Rodgers, observavit et descriptist, Proceedings of the Academy of Natural Sciences of Philadelphia 12: 22-47

Vaughan, David B.; Grutter, Alexandra S.; Hudson, Kate S. (2018) Cleaner shrimp are a sustainable option to treat parasitic disease in farmed fish, Scientific Reports 8: 13959

Williams, Austin B. (1984) Shrimps, Lobsters, and Crabs of the Atlantic Coast of the Eastern United States, Maine to Florida, Smithsonian Institution Press, Washington, DC. Pp. <missing location>

Yang, Hoi Jeong; Kim, Chang Hyun (2010) Zoeal Stages of Lysmata vittata (Decapoda: Caridea: Hippolytidae) Reared in the Laboratory, Korean Journal of Systematic Zoology 26(3): 261-278