Invasion History

First Non-native North American Tidal Record: 1901
First Non-native West Coast Tidal Record: 1901
First Non-native East/Gulf Coast Tidal Record:

General Invasion History:

Calyptspadix cerulea (Rope Grass Hydroid) was first described from Fort Wool, off Norfolk, Virginia (Clarke 1882). A very similar hydroid, was described from San Francisco as Bimeria franciscana,( Torrey 1902.). The chief difference between the two species was the apparent number of eggs in the gonophores, single in, G. franciscana, versus multiple eggs in C. cerulea. Calder (2019) found that the varying number of cells represented developing planulae, rather than a species difference in numbers of eggs, and that the 'G. franciscana' and 'G. cerulea', at least on the East Coast, are conspecific. Genetic comparisons need to be made among Eastern Pacific, European, and other disjunct populations of 'G. franciscana, but we will provisionally treat them as introduced populations of the Western Atlantic Calyptospadix cerulea (Calder 2019).

North American Invasion History:

Invasion History on the West Coast:

Calyptospadix cerulea was collected and described from Oakland, California, in San Francisco Bay, in 1901, as Bimeria franciscana by Torrey (Torrey 1902; USNM 43484, U.S. National Museum of Natural History 2007). It was subsequently found in "various scattered locations' in each of the three sections of San Francisco Bay (Fraser 1937). It has been collected from Suisun Bay and the Carquinez Straits, the Napa River, San Pablo Bay, and the Central and South Bays (Cohen and Chapman 2004; Cohen et al. 2005; U.S. National Museum of Natural History 2007), from brackish (mesohaline, 5-19 PSU) to marine (polyhaline-euhaline conditions, 18-35 PSU). The occurrence of C. cerulea outside San Francisco Bay is uncertain. Two specimens in the California Academy of Science collections are reported from Monterey Bay (1921, CAS 1132 and CAS 1194, California Academy of Science 2008), but no further reports of this hydroid have been made from here. In 1941, Fraser collected 'G. franciscana from the San Diego and San Nicolas Island (Fraser 1948). Ruiz et al. (unpublished data) found this species in San Diego Bay in 2000, but another survey in the same area and season (Cohen et al. 2002), sampling some of the same locations, did not find this species. Records are also known from Newport Bay (2011, Ruiz et al., unpublished date) and Morro Bay (2006, Needles and Wendt 2012). A record from Homer, Alaska (Hines et al. 2000) needs to be verified.

Invasion History on the East Coast:

Calyptospadix cerulea was described from Chesaeake Bay,(Clarke 1882), but rarely reported.  On the East Coast, 'Garveia. franciscana' was first reported in Chesapeake Bay, in the Potomac estuary by Frey (1946) as 'Bimeria tunicata', and was reported as abundant by 1967 (Cory 1967). In Delaware Bay, G. franciscana was reported by 1968 and had an apparent range expansion (or fluctuation) between 1968 and 1971 (Maurer and Watling 1972). Calder (1971) considered 'Garveia cerulea' tp be native to Chesapeake Bay, and associated with higher salinities, while 'Garveia franiciscana' was considered to be a probable invader, of unknown origin  based on its occurrences on the West Coast and Europe (Vervoort 1964; Dale Calder, personal communications).  .It was treated as a biological invader with significant impacts on fisheries and power plants (Cory 1967; Calder 1971; Andrew 1973; Thompson 1993).).  Garveia franciscana' was found in South Carolina in 1974 and was reported to be abundant in 'numerous areas across the coastal region' (Calder 1976, Calder and Hester 1978). This hydroid was abundant and widespread in the St. Johns River estuary, around Jacksonville, Florida in 2003 (Ruiz et al., unpublished data)  .Calyptospadix cerulea  has been reported from Providence River, Rhode Island, at the head of Narragansett Bay (in 2000, e( MIT Sea Grant 2003), Great Bay, New Hampshire, and Port de Grave, Newfoundland (Ruiz et al., unpublished data).  Calder (2019; Dale Calder, personal communications 1996-2005) had long suspected and finally  concluded that Garvea francisana and 'G. cerulea' were conspecific, and that Clarke's original name (Calyptospadix cerulea) had priority., this converting the hydroid from a exotic invasive species tp a probematic native.

Invasion History on the Gulf Coast:

In the Gulf of Mexico, Calyptospadix .cerulea was first reportedc as 'Bimeria tunicata was from 1943 on the 'Louisiana coast' (Fraser 1944), including Bayou Chene Fleuri and Sabine Pass (Deevey 1950,  as B. franciscana), and it was abundant by 1950 in Lake Pontchartrain (Deevey 1950; Crowell and Darnell 1955). It now occurs from Tampa Bay, Florida (Ruiz et al. unpublished data) to Galveston and Corpus Christi, Texas (Defenbaugh 1973; Ruiz et al., unpublished data). A notable occurrence was in the Crystal Springs cave system, north of Tampa Bay on the Gulf Coast of Florida (Garman et al. 1999). Deevey had noted its cosmopolitan distribution, together with that of other hydroids, suggesting the possibility of introduction.  However, C. cerulea is now regarded as native to the Northwest Atlantic (Calder 2019).

Invasion History in Hawaii:

Invasion History Elsewhere in the World:

In Europe, Calyptospadix cerulea was first found in 1920 in the Netherlands, in the Zuiderzee, before it was converted to a freshwater lake. (Vervoort 1964). Subsequently, it was found in the Netherlands at Edam, Amsterdam, Heelvoetsluis, in Belgium in the lower Scheldt (1962, Vervoort 1964) and in Germany in the Elbe River estuary (1946) Initially, it was confused with Cordylophora caspia, but was later recognized as a new species, Perigonimus megas (Kinne 1955, cited by Vervoort 1964) and the Kiel Canal (1950, Vervoort 1964). In the Mediterranean Sea, this hydroid was discovered in the Lagoon of Venice, Italy on the Adriatic Sea in 1978 (Morri 1982), where it is abundant (Mizzan 1999), and in Alexandira, Egypt in. In the Black Sea, C. cerulea was collected in Lake Varna, Bulgaria, in 1933, and is now widespread and abundant (Gomiou et al. 2002). By 1960, it was appearing in powerplants aling the Sea of Azov (Simkina 1963; Simkina 1975), and in 1962 it was collected in the Caspian Sea, which it reached by canal shipping (Aladin et al. 2002; Grigorovich et al. 2003).

There are several outlying records of Calyptospadix cerulea which may include some introductions by shipping, but some probably require re-examination of specimens, Occurrences in Brazil could either represent extensions of the native range. It was found in Paranagua Bay 2010, 25 31S; 48 30W), in 1985 and Neves et al. 2007; and 2007, Cangassu et al. 2010;) and closer to the Equator, in the small Rio Formosa estuary in Pernambuco State, Brazil (8 43 S, 35 6W 1993, Calder and Mayal 1998; Calder 2019; Teixeira and Creed 2020). Specimens of "Garviea franciscana' have been identified from Port Harcourt,, Nigeria 1957, Schuchert 2007) and Cameroon (1027, Vervoort 1964), from the east and west coasts of India, and Brisbane, Australia (Vervoort 1964), Genetic and morphological studies of these remote specimens are desirable.


Description

Calyptospadix cerulea, known as the Rope-Grass hydroid, lacks a planktonic medusa stage, undergoing sexual reproduction by means of attached gonophores. It has erect colonies, usually monosiphonic (single-stemmed), bushy, and densely branched. The colonies reach 100-300 mm in height. The branching is more or less regular and alternate, occurring at a 60 degree angle. The branches are helically arranged. The hydrocaulus (stem) arises from a mat of root-like fibers (hydrorhizae). Multiple hydrocauli may rise from the same mat. The perisarc is thick, wrinkled (especially near the base of the hydrocaulus), horn-colored, and often annulated around the base of the branches. The hydranths are partially covered by a pseudohydrotheca and end in a dome-shaped hypostome, which is surrounded by about 8-16 filiform tentacles in a single whorl. The sexes are separate, with gonophores born on the pedicels of the hydranths. The male gonophores are elongate-oval in shape and are covered with perisarc. The spermatozoa develop around a distinct spadix. The female gonophores are round or oval, also covered by perisarc, and have a large spadix. Clarke's (1882) description reported multiple eggs in each gonophore. A similar hydroid, originally described as Bimeria franciscana (Torrey 1902) from San Francisco Bay was reported to have only a single egg in each gonophore. Specimens of C. cerulea from Florida had both single and multiple clusters of cells in the gonophores, resulting from cell division (Calder 2019). Calder now considers Garveia fransciana to be conspecific with C. cerulea, subject to molecular studies of populations around the world.. ,


Taxonomy

Taxonomic Tree

Kingdom:   Animalia
Phylum:   Cnidaria
Class:   Hydrozoa
Subclass:   Hydroidolina
Order:   Anthoathecatae
Suborder:   Filifera
Family:   Bougainvilliidae
Genus:   Calyptospadix
Species:   cerulea

Synonyms

Bimeria franciscana (Torrey, 1902)
Bimeria monodi (Billard, 1927)
Bimeria tunicata (Fraser, 1943)
Bougainvillia megas (Kinne, 1956)
Perigonimus megas (Kinne, 1956)
Garveia franciscana (Vervoort, 1964)
Calyptospadix cerulea (Clarke, 1882)

Potentially Misidentified Species


None

Bimeria vestita
Bimeria vestita is somewhat similar, but has much smaller colonies. 5-25 mm tall, and yellowish in color (Schuchert 2007).

Cordylophora caspia
This hydroid differs considerably from G. franciscana in arrangement of tentacles and other features, and resembles G. franciscana mostly in its size, bushy appearance, and occurrence in low-salinity brackish waters.

Garveia annulata
Confusion with this native Pacific species is possible. However, the stems are strongly annulated, and the hydroid is bright orange or red in color (Mills et al., in Carlton 2007).

Ecology

General:

Calyptospadix cerulea is a sessile hydrozoan which lacks a planktonic medusa stage. Colonies grow on a solid substrate, with polyps arising from a creeping stolon. The polyps form bushy structures, with many hydranths, whose tentacles capture zooplankton. The polyps produce gonophores, which produce either eggs or sperm. Colonies are diecious (single-sexed). Female gonophores usually produce a single egg. After fertilization the egg develops into a ciliated non-feeding planula larva which is released into the water column (Crowell and Darnell 1955; Bouillon et al. 2004; Schuchert 2007).

Planulae of C. cerulea settle and grow on a wide range of substrates, including shells, rock, wood, and vegetation. They can also be found on man-made substrates including pilings, buoys, fouling plates, and inside industrial water systems (Woods Hole Oceanographic Institution 1952; Calder 1971; de Rincon and Morris 2003). Calyptospadix cerulea grows in a wide range of estuarine environments, varying in salinity, temperature, currents, and oxygen. Colonies grow slowly, but survive at 1 PSUt, and grow well at 3.5 - 35 PSU (Crowell and Dayrnell 1955). However, during heavy freshwater flows in the Caloosahatchie River estuary, C. cerulea died out, and was replaced by Cordylophora caspia (Calder 2019). This hydroid survives in estuarine areas such as the Chesapeake and Delaware Bays, and estuaries in northern Europe, wher s occasionally approach 0C (Vervoort ydr1964; Watling and Maurer 1972). It does this by remaining dormant in the winter (Crowell and Darnell 1955; Calder 1992). It can tolerate temperatures as high as 37.5?C in thermal effluents (Nauman and Cory 1969). In the Sea of Azov, sexual reproduction occurred at 19.5 - 23 C and 7.5-9 ppt (Simkina 1965).

Food:

Zooplankton and small epibenthos

Consumers:

Nudibranchs

Trophic Status:

Carnivore

Carn

Habitats

General HabitatMarinas & DocksNone
General HabitatVessel HullNone
General HabitatMangrovesNone
General HabitatRockyNone
General HabitatGrass BedNone
General HabitatUnstructured BottomNone
General HabitatCanalsNone
General HabitatCoarse Woody DebrisNone
General HabitatOyster ReefNone
Salinity RangeOligohaline0.5-5 PSU
Salinity RangeMesohaline5-18 PSU
Salinity RangePolyhaline18-30 PSU
Salinity RangeEuhaline30-40 PSU
Tidal RangeSubtidalNone
Tidal RangeLow IntertidalNone
Vertical HabitatEpibenthicNone

Life History


Tolerances and Life History Parameters

Minimum Temperature (ºC)0Based on geographical range
Maximum Temperature (ºC)37.5Field- collected in thermal effluent (Nauman and Cory 1969)
Minimum Salinity (‰)1In Lake Pontchartrain, sexual reproductive structures were seen only in the higher salinity parts of the estuary, (to 12 ppt) "It is quite likely that gonophore production is seasonal and depends on narrower ranges of environmental factors than those permissible for hydroid growth." In the Sea of Azov, asexual reproduction decreased below 8 ppt and was nearly absent at 2 ppt (Simkina 1963). Sexual reproduction occurred at 5-25 ppt in experiments, but survival of planulae at 9-15 ppt (Simkina 1965).
Maximum Salinity (‰)35Maximum Salinity, Survival - No survival occurred at 40 ppt, for animals from Sea of Azov (Simkina 1965). Colonies from the Surry Nuclear Power Plant, James River, VA, 0-15 ppt, deteriorated quickly at 30-40 ppt (Thompson 1993). Hydroids from Lake Pontchartrain LA showed "good growth" at 35 ppt (Crowell and Darnell 1955).
Minimum Reproductive Temperature19.5Development of gonophores (Simkina 1965, Sea of Azov)
Maximum Reproductive Temperature34Field data, Lake Pontchartrain LA (Crowell and Darnell 1955).
Minimum Reproductive Salinity5For sexual reproduction, Sea of Azov, Russia (Simkina 1965)
Maximum Height (mm)300(Calder 1971; Andrews 1973)
Broad Temperature RangeNoneCold temperate-Tropical
Broad Salinity RangeNoneOligohaline-Polyhaline

General Impacts

Garveia franciscana is a widespread fouling organism, which can be very abundant on man-made structures and can compete with native fouling organisms under suitable conditions. However, it can also have positive impacts and its bushy colonies may provide habitat for small fishes and invertebrates.

Economic impacts

Garveia franciscana can have negative impacts by fouling power plants and other industrial water systems. These effects are best documented in Chesapeake Bay, but they have also been reported from Venezuela (de Rincon and Morris 2003) and the Ukraine (Simkina 1963).

Industry- Fouling of power plants and other industrial water systems by G. franciscana has been reported from Chesapeake Bay (Cory 1967; Virginia Power 1992); Lake Maracaibo, Venezuela (de Rincon and Morris 2003); and the Sea of Azov, Ukraine (Simkina 1963; Simkina 1965). Fouling can block flow through water systems, cause breakdowns of traveling screens which remove debris, and speed up corrosion of metal structures (Virginia Power 1992; de Rincon and Morris 2003). Costs include shutdowns for cleaning and the cost of biocides to reduce fouling. At the Surry Nuclear power plant in Virginia, fouling by G. franciscana prompted an expensive redesign of the cooling system (Virginia Power 1992). The use of biocides to prevent fouling, such as chlorine and other chemicals, raises environmental concerns about their toxicity to other organisms (McLean 1972; Virginia Power 1992).

Fisheries- Fouling by G. franciscana has been a major problem on fishing gear in Chesapeake Bay, including crab pots and oyster trays (Andrews 1973). On the other hand, G. franciscana probably benefits commercial and sport fisheries by providing habitat for juvenile and bait fishes, shrimps, crabs, and other motile organisms in the Chesapeake Bay (Thompson 1993) and Lake Pontchartrain, Louisiana (Crowell and Darnell 1955).

Ecological Impacts

Although G. franciscana (Rope Grass Hydroid) is an abundant and sometimes dominant part of the fouling community in many estuaries, its ecological impacts are largely unknown.

Competition - Garveia franciscana and Victorella pavida (cryptogenic on the East and Gulf Coasts) overgrew most other organisms on fouling panels at Calvert Cliffs, Maryland, in summer (Abbe 1987). Garveia franciscana overlaps spatially with Cordylophora caspia, although C. caspia ranges into lower salinities (Calder 1971; Cory 1967; Thompson 1993). It also co-occurs with V. pavida (cryptogenic). Victorella pavida and G. franciscana settle at the same time at Calvert Cliffs, but G. franciscana persists longer in summer (Abbe 1987).

Habitat Change - Growths of G. franciscana provide cover for numerous amphipods, mud crabs, and other organisms in Patuxent River, Maryland (Cory 1967); James River, Virginia (Thompson 1993); and Lake Ponchartrain, Louisiana (Crowell and Darnell 1955).

Food - Garveia franciscana is fed on by nudibranchs, particularly Tenellia spp. (Abbe 1987; Cory 1967; Thompson 1993).


Regional Impacts

NA-ET3Cape Cod to Cape HatterasEconomic ImpactIndustry
Calyptospadix cerulea is a major fouling organism in the Chesapeake Bay region. The organism with the highest quantified cost to power plants in the Chesapeake Bay is C. cerulea, which is a major fouler of power plants along mesohaline waters of Chesapeake Bay (Cory 1967; Virginia Power 1992) and elsewhere in the world (Simkina 1963). At two power plants in Chesapeake Bay, Chalk Point (Patuxent River) and Morgantown (Potomac River), the cost of biocides to control fouling dominated by C. cerulea was 95,000 to $180,000 per year in 1995-97 (Krueger 1997, personal communication). At the Surry Nuclear Power Plant on the lower James River, the weight of the hydroids caused breakdowns of the traveling screens which remove debris. Aggregates of hydroids blocked water flow in the main condensers, in the circulating water systems used for cooling during routine operation, and in the service water systems, which would be used in shutdown after an accident. 'Almost daily' cleaning was required during warm weather to keep water flowing through the service system. In 1992, the projected cost of 'doing nothing' about the biofouling problem, except routine cleaning and repairs was $37.5 million projected over the plant's remaining license period (to the year 2013), or $3.4 million per year. The operators of the plant instead undertook an extensive reconstruction of the cooling and screen system, which was intended to reduce fouling problems, with a projected cost of $23.6, or $2.1 million per year (Virginia Power 1992). One major consequence of the invasion of exotic fouling organisms is the increased use of biocides and antifouling coatings in power plants and industrial water systems. These chemicals are toxic substances used to kill microbial and invertebrate organisms on surfaces exposed to natural waters. Among coatings used in power plants are copper oxide paints, organotin compounds, as well as nontoxic coatings that inhibit attachment of organisms (Virginia Power 1992). At the Surry Nuclear Power Plant, several biocides in addition to chlorine were tested against G. franciscana, including ammonium hydroxide, hydrogen peroxide, sodium bromide-hypochlorite mixture, a surfactant mixture ('ClamTrol'), and chemically induced anoxia. However, at doses allowed by their U.S. EPA permit, these were ineffective (Virginia Power 1992).
NA-ET3Cape Cod to Cape HatterasEconomic ImpactFisheries
Fisheries- Costs: Fouling by Calyptospadix cerulea has been a major problem on fishing gear, including crab pots and oyster tayCalyptospadix ceruleaCalyptospadix cerulea probably benefits commercial and sport fisheries by providing habitat for juvenile and bait fishes, shrimps, crabs, and other motile organisms (Thompson 1993).
NA-ET3Cape Cod to Cape HatterasEcological ImpactCompetition
Competition- Although Calyptospadix cerulea (Rope Grass Hydroid) is an abundant and sometimes dominant part of the fouling community in parts of Chesapeake Bay, its ecological impacts are largely unknown. Calyptospadix cerulea and Victorella pavida overgrew most other organisms on fouling panels at Calvert Cliffs, Maryland, in summer (Abbe 1987).
NA-ET3Cape Cod to Cape HatterasEcological ImpactHabitat Change
Growths of Calyptospadix cerulea provide cover for numerous amphipods, mud crabs, and other organisms in the Patuxent River (Cory 1967) and James River (Thompson 1993).
NA-ET3Cape Cod to Cape HatterasEcological ImpactFood/Prey
Food/Prey- Calyptospadix cerulea is fed on by nudibranchs, particularly Tenellia sp. (Cory 1967; Abbe 1987; Thompson 1993).
CAR-INorthern Yucatan, Gulf of Mexico, Florida Straits, to Middle Eastern FloridaEcological ImpactHabitat Change
May provide shelter for invertebrates and small fishes (Crowell and Darnell 1955).
MED-XNoneEconomic ImpactIndustry
Power plant fouling (Simkina 1963)
CAR-IIINoneEconomic ImpactIndustry
Fouling of industrial plants using Lake Maracaibo water and submerged structures (de Rincon and Morris 2003).
M130Chesapeake BayEconomic ImpactIndustry
Calytpspadix cerulea is a major fouling organism in the Chesapeake Bay region. Industry- The organism with the highest quantified cost to power plants in the Chesapeake Bay is C. cerulea, which is a major fouler of power plants along mesohaline waters of Chesapeake Bay (Cory 1967; Virginia Power 1992) and elsewhere in the world (Simkina 1963). At two power plants in Chesapeake Bay, Chalk Point (Patuxent River) and Morgantown (Potomac River), the cost of biocides to control fouling dominated by C. cerulea was $95,000 to $180,000 per year in 1995-97 (Krueger 1997, personal communication). At the Surry Nuclear Power Plant on the lower James River, the weight of the hydroids caused breakdowns of the traveling screens which remove debris. Aggregates of hydroids blocked water flow in the main condensers, in the circulating water systems used for cooling during routine operation, and in the service water systems, which would be used in shutdown after an accident. 'Almost daily' cleaning was required during warm weather to keep water flowing through the service system. In 1992, the projected cost of 'doing nothing' about the biofouling problem, except routine cleaning and repairs was $37.5 million projected over the plant's remaining license period (to the year 2013), or $3.4 million per year. The operators of the plant instead undertook an extensive reconstruction of the cooling and screen, which was intended to reduce fouling problems, with a projected cost of $23.6, or $2.1 million per year (Virginia Power 1992). One major consequence of the invasion of exotic fouling organisms is the increased use of biocides and antifouling coatings in power plants and industrial water systems. These chemicals are toxic substances used to kill microbial and invertebrate organisms on surfaces exposed to natural waters. Among coatings used in power plants are copper oxide paints, organotin compounds, as well as nontoxic coatings that inhibit attachment of organisms (Virginia Power 1992). At the Surry Nuclear Power Plant, several biocides in addition to chlorine were tested against G. franciscana, including ammonium hydroxide, hydrogen peroxide, sodium bromide-hypochlorite mixture, a surfactant mixture ('ClamTrol'), and chemically induced anoxia. However, at doses allowed by their U.S. EPA permit, these were ineffective (Virginia Power 1992).
M130Chesapeake BayEconomic ImpactFisheries
Fisheries- Costs: Fouling by Calyptospadix cerulea has been a major problem on fishing gear, including crab pots and oyster trays (Andrews 1973). Benefits: Calyptspadix cerulea probably benefits commercial and sport fisheries by providing habitat for juvenile and bait fishes, shrimps, crabs, and other motile organisms (Thompson 1993).
M130Chesapeake BayEcological ImpactCompetition
Competition- Although Calyptspadix cerulea (Rope Grass Hydroid) is an abundant and sometimes dominant part of the fouling community in parts of Chesapeake Bay, its ecological impacts are largely unknown.n. Calyptspadix cerulea and Victorella pavida overgrew most other organisms on fouling panels at Calvert Cliffs, Maryland, in summer (Abbe 1987).
M130Chesapeake BayEcological ImpactHabitat Change
Growths of Calyptospadix cerulea provide cover for numerous amphipods, mud crabs, and other organisms in the Patuxent River (Cory 1967) and James River (Thompson 1993).
M130Chesapeake BayEcological ImpactFood/Prey
Food/Prey- Calyptospadix cerulea is fed on by nudibranchs, particularly Tenellia sp. (Abbe 1987;Cory 1967; Thompson 1993).

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
NA-ET3 Cape Cod to Cape Hatteras 1882 Native Estab
CAR-VII Cape Hatteras to Mid-East Florida 0 Native Estab
CAR-I Northern Yucatan, Gulf of Mexico, Florida Straits, to Middle Eastern Florida 1943 Native Estab
NEP-V Northern California to Mid Channel Islands 1901 Def Estab
CIO-I None 1932 Crypto Estab
CIO-III None 1932 Crypto Estab
CIO-V None 1932 Crypto Estab
NEA-II None 1920 Def Estab
MED-VII None 1978 Def Estab
AUS-XII None 1959 Crypto Estab
MED-X None 1960 Def Estab
CASP Caspian Sea 1962 Prb Estab
B-III None 1950 Def Estab
SA-III None 1993 Crypto Estab
MED-IX None 1933 Def Estab
CAR-III None 1924 Native Estab
SEP-H None 1935 Def Estab
WA-III None 1927 Def Estab
NEP-VI Pt. Conception to Southern Baja California 1941 Def Estab
NA-ET2 Bay of Fundy to Cape Cod 2001 Native Estab
G070 Tampa Bay 2002 Native Estab
N110 Saco Bay 1968 Native Estab
G260 Galveston Bay 1948 Native Estab
M130 Chesapeake Bay 1882 Native Estab
S180 St. Johns River 2001 Native Estab
G130 Pensacola Bay 2002 Native Estab
M020 Narragansett Bay 2000 Native Unk
N130 Great Bay 2001 Native Estab
P090 San Francisco Bay 1901 Def Estab
P020 San Diego Bay 1941 Def Estab
G268 _CDA_G268 (Austin-Oyster) 1948 Native Estab
G250 Sabine Lake 1948 Native Estab
G310 Corpus Christi Bay 2000 Native Estab
G170 West Mississippi Sound 1955 Native Estab
G074 _CDA_G074 (Crystal-Pithlachascotee) 1999 Native Estab
S056 _CDA_S056 (Northeast Cape Fear) 1974 Native Estab
S060 Winyah Bay 1974 Native Estab
S070 North/South Santee Rivers 1974 Native Estab
S080 Charleston Harbor 1974 Native Estab
S090 Stono/North Edisto Rivers 1974 Native Estab
S076 _CDA_S076 (South Carolina Coastal) 1974 Native Estab
S100 St. Helena Sound 1974 Native Estab
S110 Broad River 1974 Native Estab
S120 Savannah River 1974 Native Estab
M100 Delaware Inland Bays 1972 Native Estab
P093 _CDA_P093 (San Pablo Bay) 1901 Def Estab
SA-II None 1985 Crypto Estab
S190 Indian River 2005 Native Estab
WA-II None 1957 Def Estab
P061 _CDA_P061 (Los Angeles) 1941 Def Estab
CIO-II None 0 Crypto Estab
MED-V None 1980 Def Estab
P080 Monterey Bay 1921 Def Unk
P070 Morro Bay 2006 Def Estab
P040 Newport Bay 2011 Def Estab
G330 Lower Laguna Madre 2008 Native Estab
PAN_PAC Panama Pacific Coast 1935 Def Estab
PAN_CAR Panama Caribbean Coast 1924 Native Estab
NA-S3 None 0 Native Estab
NA-S2 None 0 Native Estab

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude
2539 Frey 1946 1946 1946-01-01 Lower Cedar Point Native 38.7650 -76.0503
2540 Cory 1967 1963 1963-01-01 Chalk Pt. Native 38.3189 -76.4053
2541 Calder 1971 1965 1965-01-01 Norfolk Native 36.8689 -76.2739
2542 Calder 1971 1965 1965-01-01 Surry Power Station (Hog Island) Native 36.9414 -76.4439
2543 Abbe 1987 1970 1970-01-01 Calvert Cliffs Native 38.5250 -76.4103
2544 Ruiz et al. unpublished data 1997 1997-01-01 SERC dock, Edgewater Native 38.8647 -76.5150
2545 Cory 1967 1963 1963-01-01 Solomons Native 38.3183 -76.4544
2546 Watling and Maurer 1972 1968 1968-01-01 Woodland Beach Native 39.0500 -75.1500
2547 Watling and Maurer 1972 1968 1968-01-01 Bayside Native 38.7903 -75.1625
2548 Watling and Maurer 1972 1968 1968-01-01 Rehoboth Native 38.6000 -75.1008
2549 Watling and Maurer 1972 1968 1968-01-01 Murderkill River Native 39.0583 -75.3969
2550 Watling and Maurer 1972 1968 1968-01-01 St. Jones River Def 39.0658 -75.4011
2551 Pfitzenmeyer 1976; Lippson et al. 1979 1976 1976-01-01 Morgantown Native 37.9389 -76.2528
2552 Calder 1976; Calder and Hester 1978 1974 1974-01-01 Georgetown Native 33.2750 -79.2569
2553 Calder 1976; Calder and Hester 1978 1974 1974-01-01 Santee River Def 33.1667 -79.3000
2554 Calder 1976; Calder and Hester 1978 1974 1974-01-01 'oligo-meso' zone, Charleston Native 32.8850 -79.8767
2555 Calder 1976; Calder and Hester 1978 1974 1974-01-01 Cooper River Native 32.8967 -79.9600
2556 Calder 1976; Calder and Hester 1978 1974 1974-01-01 Edisto River Native 32.5617 -80.3950
2557 Calder 1976; Calder and Hester 1978 1974 1974-01-01 Edisto River Native 32.5667 -80.3950
2558 Garman 1999 1997 1997-01-01 Crystal Springs Beach marine caves Native 28.0911 -82.7800
2559 Crowell and Darnell 1955 1955 1955-01-01 Slidell Native 30.2750 -89.7811
2560 Deevey 1950 1950 1950-01-01 Freeport Native 28.9539 -95.3594
2561 Deevey 1950 1950 1950-01-01 Houston Ship Channel Native 29.5694 -94.9364
2562 Deevey 1950 1950 1950-01-01 Sabine Pass Native 29.7333 -93.8942
2563 Defenbaugh 1973 1955 1955-01-01 Missisippi Sound Native 30.3503 -89.1528
2564 Torrey 1902 1902 1902-01-01 Oakland Def 37.7933 -122.2894
2622 Ruiz et al. unpublished 2002 (DOD) 2002 2002-01-01 'Regatta Pointe' Native 27.5107 -82.5755
2635 Ruiz et al. unpublished 2002 (DOD) 2002 2002-01-01 'BayBoro Yacht Club' Native 27.7593 -82.6360
2636 Ruiz et al. unpublished 2002 (DOD) 2002 2002-01-01 'Mariner's Club' Native 27.7272 -82.4748
2638 Ruiz et al. unpublished 2002 (DOD) 2002 2002-01-01 Interbay Moorings Native 27.8585 -82.3810
2640 Ruiz et al. unpublished 2002 (DOD) 2002 2002-01-01 Outbound Texas City Dike Native 29.3762 -94.8473
2641 Ruiz et al. unpublished 2002 (DOD) 2002 2002-01-01 Port of Pensacola Native 30.4016 -87.2110
2643 Ruiz et al. unpublished 2002 (DOD) 2002 2002-01-01 Quiet Waters Native 30.3361 -88.1808
2645 Ruiz et al. unpublished 2002 (DOD) 2002 2002-01-01 Santa Rosa Yacht Club Native 30.3535 -87.1553
2646 Ruiz et al. unpublished 2002 (DOD) 2002 2002-01-01 Island Moorings Marina, Corpus Christi Native 27.8070 -97.0872
2650 Ruiz et al. unpublished 2002 (DOD) 2002 2002-01-01 Teekwood Marina Native 29.2989 -94.9034
2651 US National Museum of Natural History 2011 1974 1974-01-26 Miraflores Locks, Upper Chamber Floor Def 8.9986 -79.5953
2652 US National Museum of Natural History 2011 1974 1974-01-05 Gatun Locks, Lower East Chamber, Wall Native 9.2778 -79.9250
2676 Ruiz et al. unpublished 2002 (DOD) 2000 2000-01-01 Chula Vista Marina, San Diego/ Def 32.6256 -117.1032
2678 Ruiz et al. unpublished 2002 (DOD) 2000 2000-01-01 NAB Fiddlers Cove, San Diego Def 32.6517 -117.1379
2683 Ruiz et al. unpublished 2002 (DOD) 2000 2000-01-01 Norfolk Naval Station Native 36.9413 -76.3298
2685 Ruiz et al. unpublished 2002 (DOD) 2001 2001-01-01 River City Brewing Company, Jacksonville Native 30.3191 -81.6608
2686 Ruiz et al. unpublished 2002 (DOD) 2001 2001-01-01 Jones College, Jacksonville Native 30.3876 -81.6105
2687 Ruiz et al. unpublished 2002 (DOD) 2001 2001-01-01 Jaxport, Blount Island, Jacksonville Native 30.3876 -81.5512
2688 Ruiz et al. unpublished 2002 (DOD) 2001 2001-01-01 Dames Point, Bulk Aggregate Terminal, Jacksonville Native 30.4093 -81.5825
2691 Ruiz et al. unpublished 2002 (DOD) 2001 2001-01-01 Talleyrand-Jaxport Terminal, Jacksonville/ Native 30.3570 -81.6195
2697 Ruiz et al. unpublished 2002 (DOD) 2001 2001-01-01 Browns Creek Bridge-Clapboard, Jacksonville Native 30.4177 -81.5330
2698 Ruiz et al. unpublished 2002 (DOD) 2001 2001-01-01 Mayport Naval Station, Turning Basin Native 30.3682 -81.4016
2709 Ruiz et al. unpublished 2002 (DOD) 2001 2001-01-01 USCG, Mayport Shrimp Docks Native 30.3982 -81.4148
2712 Ruiz et al. unpublished 2002 (DOD) 2001 2001-01-01 Pier 68 Marina, Jacksonville Native 30.3925 -81.6470
2714 Ruiz et al. unpublished 2002 (DOD) 2001 2001-01-01 Blount Island Berths 22 and 20, Jacksonville Native 30.3956 -81.5495
6904 Cohen et al. 2005 2004 2004-01-01 Rodeo Marina Def 38.0391 -122.2711
6905 Cohen et al. 2005 2005 2005-01-01 Napa River Def 38.2261 -122.3076
6906 Cohen et al. 2005 2004 2004-01-01 Napa Valley Marina Def 38.2200 -122.3128
6908 Fraser 1948 1941 1941-01-01 San Nicolas Island Def 33.2495 -119.5004
6909 MIT Sea Grant 2003) 2000 2000-08-14 India Point Native 41.8200 -71.3900

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