Invasion History

First Non-native North American Tidal Record: 1978
First Non-native West Coast Tidal Record: 1978
First Non-native East/Gulf Coast Tidal Record: 1996

General Invasion History:

Caprella scaura was first described from Mauritius in 1836, and subsequently found in tropical and subtropical areas around the world. Historically, it ranged as far north as Vladivostok, Russia, in the Northwest Pacific, and south to Chile, South Africa, and southern Australia (McCain 1968). Its native region is uncertain. In other parts of the world, it has recently extended its range into warm-temperate regions, including California (Marelli 1981), South Carolina, the Gulf Coast of Florida (Foster et al. 2004), the Atlantic coast of Spain, and the Canary Islands (Guerra-Garcia et al. 2011). In the Mediterranean, there was an early report from the Thau Lagoon, France, an oyster-rearing area (Razarihelisoa 1958), but it has appeared in other regions of the Sea more recently (Krapp et al. 2006; Martínez and Adarraga 2008; Zenetos et al. 2008; Bakir and Katagan 2011). The morphological variability of this caprellid indicates that the name C. scaura is a species complex, but many of the introduced populations are morphologically similar and appear to correspond to Mayer's (1890, cited by McCain 1968) form ‘typica’. Caprella scaura is capable of long-distance dispersal on floating seaweeds and other objects. However, its worldwide distribution has probably resulted from transport by hull fouling, ballast water, and by oyster transplants and other forms of aquaculture (Foster et al. 2004; Krapp et al. 2006; Martínez and Adarraga 2008; Guerra-Garcia et al. 2011).

North American Invasion History:

Invasion History on the West Coast:

Caprella scaura was first collected on the West Coast in 1978 by Marelli (1981) from the cooling systems of power plants in San Francisco Bay and Elkhorn Slough, California. It was subsequently collected in Southern California, from the Channel Islands, Oxnard, and San Diego Bay (Fairey et al. 2002).

Invasion History on the East Coast:

Caprella scaura was collected during benthic surveys on the East Coast of Florida, in Biscayne Bay in 1996, and the St. Johns River estuary in 2001 (NOAA National Benthic Inventory 2012). In 2002, it was found in the fouling community of a floating dock in Charleston Harbor, South Carolina (Foster et al. 2004).

Invasion History on the Gulf Coast:

In 1998-2002, Caprella scaura was found near Panama City, Florida in St. Andrews Bay on jetties and channel markers (Foster et al. 2004).

Invasion History in Hawaii:

Caprella scaura was first found in Pearl Harbor, Oahu in 1929 (Coles et al. 1999a; Calton and Eldredge 2009). Subsequently, it was found in other Oahu harbors (Coles et al. 1999b), and in 2003, in Malea Harbor, Maui (Coles et al. 2004).

Invasion History Elsewhere in the World:

As noted above, the type locality of C. scaura is Mauritius. Mayer (1890, cited by Krapp et al. 2006) obtained type specimens for the subspecies typica and cornuta in Rio de Janeiro, while the type locality for his diceros and Utinomi's (1947, cited by Krapp et al. 2006) hamata form was Japan. These forms all lack a ventral spine between gnathopods 2, and may be conspecific or closely related (Krapp et al. 2006). In any event, C. scaura s. l. was distributed in the South Atlantic, Pacific and Indian Oceans by the late 19th century. By 1968, C. scaura was known from the Caribbean, the Southwest Atlantic (Brazil), the Southeast Pacific (Chile), the Southwest Pacific (Sydney, Australia), the Northwest Pacific (Vladivostok to southern Japan), and the Southeast Atlantic (South Africa) (McCain 1968). More recent records include New Caledonia (Laubitz 1991), St. Paul's Island (Southern Indian Ocean, Laubitz 1995), and Hong Kong (Guerra-Garcia and Takeuchi 2003). We consider most of this range cryptogenic, given the potential for natural dispersal, and early, undocumented transport of this caprellid by ships.

Recent expansions of C. scaura in several parts of the world appear to be introductions. In Australia, the caprellid was found in Port Philip Bay, Victoria in 1990 (Museum Victoria 2012), in Tasmania in 1980 (Guerra-Garcia and Takeuchi 2004), and in Western Australia in 1983 (Guerra-Garcia 2004). In Europe, there was an early report of C. scaura in the Mediterranean, in the Thau Lagoon, a major oyster-rearing area (Razarihelisoa 1958). However, its range in the Mediterranean and Eastern Atlantic has expanded greatly in the last two decades. Caprella scaura was seen in the Venice Lagoon in 1994 (Krapp et al. 2006), and subsequently appeared in the Ionian Sea, Greece (in 2002, Krapp et al. 2006), the Aegean Sea, Turkey (in 2008, Bakir and Katagan 2011), and Western Mediterranean locations in Spain and Italy (Martínez and Adarraga 2008). In the Eastern Atlantic, this caprellid appeared off Cadiz, Spain and in great abundance in aquaculture operations in the Canary Islands, in 2009 (Guerra-Garcia et al. 2011).


Description

Caprellid amphipods have a greatly modified body form, when compared to more familiar gammarid amphipods. The body is elongated (giving rise to the name 'skeleton shrimp'), though the abdomen is compressed. The head is partly fused with the first thoracic segment (called Pereonite 1 in amphipods). The head bears a pair of long antennae 1, somewhat shorter antennae. The 1st antennae (A1) have a 3-segmented peduncle, tipped by a flagellum with multiple segments. The 2nd antennae (A2) may be fringed with long setae, and have 3-4 segments in the peduncle, and a shorter flagellum, usually of 2 segments. A mandibular palp of several segments is present in some genera, arising between the antennae, but this is absent in Caprella. There is a small pair of gnathopods, small grasping claws with a movable finger (Gnathopod 1), on Pereonite 1. Pereonite 2 bears a much larger pair of gnathopods (Gnathopod 2), which may have conspicuous spines or setae. Pereonites 3 and 4 usually have round or club-shaped gills, while in most species, including Caprella, pereiopods are absent. Pereiopods 5, 6, and 7 are roughly equal and hook-like, for climbing and attachment, with 6 segments. Females develop oostegites, plates which form a brood pouch. Males are usually larger than females of the same species. Females and immature males can be hard to identify to species level. (Description from: Barnes 1983; Watling and Carlton, in Carlton 2007).  

Caprella scaura are distinguished by a sharply pointed, forward-pointing dorsal spine on the head. Mature male gnathopod 2 propodus may be extended distally, with a mid-palm proximal spine, distal poison tooth and triangular projection; dactylus may be sculpted However, the size of specimens can vary considerably by region and season. There are a number of morphological differences between male and females. In females, flagellum of the antennule has up to four segments, usually fewer than the male. The pereonites of females are greatly shortened compared with those of the males. Females vary in the number of body spines, with as many as one or two knobs on each pereonite. The propodus of gnathopod 2 of females is less elongated than males, and has one proximal spine, and a small distal tooth. Mature females are distinguished by the developing oostegites and brood pouch. See descriptions by: McCain 1968; Guerra-Garcia and Takeuchi 2003; Martínez and Adarraga 2008; Guerra-Garcia et al. 2011.

This 'species' shows a high degree of variation in several anatomical features. Mayer (1890, cited by McCain 1968) named six varieties (now subspecies): typica, cornuta, diceros, scauroides, spinirostris, and californica. An additional one, hamata was added by Utinomi (1947, cited by McCain 1968; Krapp et al. 2006). One of the distinguishing features of the subspecies is the absence of a ventral spine between the insertions of Gnathopods 2 in typica, cornuta, and diceros, and its presence in the other named forms, including californica, scauroides, and spinirostris. Caprella californica, from the Northeast Pacific, is now recognized as a full species (McCain 1968; Watling and Carlton, in Carlton 2007). The other ventral-spined forms, scauroides and spinirostris, may be conspecific with C. californica (Laubitz 1970, cited by Guerra-Garcia and Takeuchi 2004). The currently recognized subspecies of Caprella scaura appear to constitute a widespread species complex. Molecular studies will be needed to clarify their systematics and origin. However, specimens caught in US Atlantic and Pacific waters resemble C. s. typica (McCain 1968; Marelli 1981; Foster et al. 2004).


Taxonomy

Taxonomic Tree

Kingdom:   Animalia
Phylum:   Arthropoda
Subphylum:   Crustacea
Class:   Malacostraca
Subclass:   Eumalacostraca
Superorder:   Peracarida
Order:   Amphipoda
Suborder:   Caprellidea
Infraorder:   Caprellida
Superfamily:   Caprelloidea
Family:   Caprellidae
Genus:   Caprella
Species:   scaura

Synonyms

Caprella attenuata (Dana, 1853)
Caprella attenuata forma subtenuis (Dana, 1853)
Caprella cornuta (Dana, 1853)
Caprella cornuta forma obtusirostris (Dana, 1853)
Caprella nodosa (Templeton, 1836)
Caprella scaura forma cornuta (Mayer, 1890)
Caprella scaura forma diceros (Mayer, 1890)
Caprella scaura forma hamata (Mayer, 1890)
Caprella scaura forma typica (Mayer, 1890)

Potentially Misidentified Species

Caprella californica
NE Pacific native, formerly considered a subspecies of C. scaura (Cabezas et al. 2014)

Caprella penantis
Cosmopolitan, a species complex

Caprella scauroides
NW Pacific native, Taiwan

Caprella simia
NW Pacific native, introduced to California

Ecology

General:

Life History – Males and females are morphologically distinct. The males are larger, more robust, and armed with larger gnathopods, probably an adaptation to competition for females and for guarding themselves during molting, which precedes mating. The young are brooded by the female in an egg-pouch formed by large plates (oostegites) on the 3rd and 4th pereionites (Turcotte and Sainte Marie 2009). Brooding takes about four days at 24⁰C (Lim and Alexander 1986). Development is direct, with the newborn juveniles having the general form of the adults. The 1st instar of Caprella scaura are quite small (~1.2 mm), lacking the distinctive head spine, and having less developed gnathopods, however perieopods with long dactyls, specialized for clinging to the mother are observed even in small individuals (Lim and Alexander 1986). The young are dependent on maternal care and cling to females for about a week after birth. Brooding females and females with newborns are more likely to fight with males than are non-reproducing females (Lim and Alexander 1986; Aoki 1999).

Ecology – Caprellids can feed in a variety of ways, including filtering small particles from the water, browsing on small filamentous algae, scraping tissue from large algae, scavenging, and predation (Turcotte and Sainte Marie 2009). An examination of the gut contents (specimens from Mauritius, Australia, Greece, and Chile) found that 99.8% of their diet was detritus (Guerra-Garciana and de Figeroa 2009). A later study found a dietary shift during development, with juvrniles doing more predatory feeding, and then shifting to detritus feeding as adults (Ros et al. 2014). Caprella scaura is known for using a wide range of habitats, including seaweeds, seagrasses, sponges, hydroids, bryozoans (Zoobotryon verticillatum, Bugula neritina), and manmade structures such as buoys, floating docks, and aquaculture systems (McCain 1968; Thiel et al. 2003; Foster et al. 2004; Guerra-Garcia et al. 2011; Ros et al. 2013; Ros et al. 2016; Molina et al. 2017). In short term experiments, C. scaura tolerated salinities as low as 7.5 PSU (Cockman and Albone 1987, cited by Ros et al. 2015), but on the Mediterranean and Atlantic coasts of Spain,C. scaura predominated over C. equilibria at salinities of 36-38 PSU (Ros et al. 2015).

Food:

Phytoplankton, detritus

Trophic Status:

Suspension Feeder

SusFed

Habitats

General HabitatGrass BedNone
General HabitatCoarse Woody DebrisNone
General HabitatOyster ReefNone
General HabitatMarinas & DocksNone
General HabitatRockyNone
General HabitatUnstructured BottomNone
Salinity RangePolyhaline18-30 PSU
Salinity RangeEuhaline30-40 PSU
Tidal RangeSubtidalNone
Vertical HabitatEpibenthicNone


Tolerances and Life History Parameters

Minimum Temperature (ºC)7Field, Mar Piccolo, Italy, Mediterranean Sea (Prato et al. 2013)
Maximum Temperature (ºC)30Field, Mar Piccolo, Italy, Mediterranean Sea (Prato et al. 2013)
Minimum Salinity (‰)7.5Experimental, 24 hr LC 50, Swan River, Western Australia, Cockman and Albone 1987, cited by Ros et al. (2015).
Maximum Salinity (‰)38Field, Mar Piccolo, Italy, Mediterranean Sea (Prato et al. 2013)
Minimum Length (mm)5.6For mature breeding females,Mar Piccolo, Italy, Mediterranean Sea, Prato et al. 2013)
Maximum Length (mm)23Mar Piccolo, Italy, Mediterranean Sea (Prato et al. 2013)
Broad Temperature RangeNoneWarm temperate-Tropical
Broad Salinity RangeNonePolyhaline-Euhaline

General Impacts

Caprella scaura has reached very high densities in some introduced locations, including Cadiz Harbor (Guerra-Garcia et al. 2011) and Roses Bay, Spain (Martinez and Adaragga 2008); the Venice Lagoon (Krapp et al. 2006); and in aquaculture operations in the Canary Islands (Guerra-Garcia et al. 2011). In Cadiz Harbor, C. scaura apparently displaced native caprellids and other amphipods (Guerra-Garcia et al. 2011). While this caprellid has been associated with fish culture, negative impacts on aquaculture have not been reported.

Regional Impacts

NEA-VNoneEcological ImpactCompetition
In the harbor of Cadiz, Spain, Caprella scaura has apparently replaced native Caprella equilibra or Caprella dilatata and apparently reduced the abundance of other native amphipods (Guerra-García et al. 2011). On the Atlantic and Mediterranean coasts of Spain, C. scaura tended to replace C. equilibria at salinities at 36-38 PSU, and higher temperatures (Ros et al. 2015)
MED-INoneEcological ImpactCompetition
On the Atlantic and Mediterranean coasts of Spain, C. scaura tended to replace C. equilibria at salinities at 36-38 PSU, and higher temperatures (Ros et al. 2015).
MED-IINoneEcological ImpactCompetition
On the Atlantic and Mediterranean coasts of Spain, C. scaura tended to replace C. equilibria at salinities at 36-38 PSU, and higher temperatures (Ros et al. 2015)
WA-INoneEcological ImpactCompetition
In experiments in Madeira, Caprella scaura tended to displace the cryptogenic C, equilibia, and this effect was greater at 29 C compared to 29 C, suggesting that C. scaura's abundance will be favored by climate warming.

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
EA-V None 1836 Crypto Estab
SEP-B None 0 Crypto Estab
AUS-X None 0 Crypto Estab
SP-XXI None 1929 Def Estab
AUS-VII None 1929 Crypto Estab
NWP-2 None 0 Crypto Estab
AUS-IX None 1980 Def Estab
NWP-3b None 0 Crypto Estab
NWP-3a None 0 Crypto Estab
NWP-4a None 0 Crypto Estab
NWP-4b None 0 Crypto Estab
AUS-VIII None 1990 Def Estab
SA-II None 1890 Crypto Estab
CAR-IV None 1968 Crypto Estab
MED-VII None 1994 Def Estab
NEP-V Northern California to Mid Channel Islands 1978 Def Estab
AUS-IV None 1983 Def Estab
CAR-I Northern Yucatan, Gulf of Mexico, Florida Straits, to Middle Eastern Florida 1996 Def Estab
CAR-VII Cape Hatteras to Mid-East Florida 2001 Def Estab
SP-IV None 1989 Crypto Estab
WA-IV None 0 Crypto Estab
WA-V None 0 Crypto Estab
EA-IV None 0 Crypto Estab
SEP-H None 1968 Crypto Estab
MED-IV None 2006 Def Estab
P080 Monterey Bay 1978 Def Estab
P090 San Francisco Bay 1978 Def Estab
NEP-VI Pt. Conception to Southern Baja California 2001 Def Estab
P020 San Diego Bay 2001 Def Estab
P050 San Pedro Bay 2001 Def Estab
P060 Santa Monica Bay 2001 Def Estab
G110 St. Andrew Bay 1998 Def Estab
S200 Biscayne Bay 1996 Def Estab
S180 St. Johns River 2001 Def Estab
S080 Charleston Harbor 2002 Def Estab
SEP-C None 0 Crypto Estab
MED-II None 1958 Def Estab
P062 _CDA_P062 (Calleguas) 2001 Def Estab
NEA-V None 2009 Def Estab
WA-I None 2009 Def Estab
MED-VI None 2008 Def Estab
AUS-XII None 0 Crypto Estab
MED-III None 2004 Def Estab
MED-I None 2011 Def Estab
P070 Morro Bay 2011 Def Unk
P040 Newport Bay 2011 Def Estab
P023 _CDA_P023 (San Louis Rey-Escondido) 2011 Def Estab
NEA-VI None 2013 Def Estab
NA-ET3 Cape Cod to Cape Hatteras 2018 Def Unk
M128 _CDA_M128 (Eastern Lower Delmarva) 2018 Def Unk
MED-VIII None 2012 Def Estab
NEA-II None 2009 Def Estab

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude
767664 Ruiz et al., 2015 2013 2013-07-16 Naval Base Point Loma, San Diego Bay, CA, California, USA Def 32.6886 -117.2343
768001 Ruiz et al., 2015 2012 2012-08-23 Sausalito Marine Harbor, San Francisco Bay, CA, California, USA Def 37.8609 -122.4853
768228 Ruiz et al., 2015 2012 2012-09-13 San Leandro Marina, San Francisco Bay, CA, California, USA Def 37.6962 -122.1919

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