Invasion History

First Non-native North American Tidal Record: 1865
First Non-native West Coast Tidal Record: 1920
First Non-native East/Gulf Coast Tidal Record: 1865

General Invasion History:

Cordylophora caspia was first described from the Caspian Sea by Pallas in 1771 and is believed to be native to the Black Sea-Caspian Sea region (Briggs 1931; Naumov 1969; Hutchinson 1993). Shipping has spread C. caspia through much of the world and this hydroid is now known from temperate and tropical coastal regions of every continent (except Antarctica), and from many fresh waters as well (Arndt 1984; Hutchinson 1993; Naumov 1969; Slobodkin and Bossert 1991).

Genetic studies (Folino-Rorem et al. 2009) indicate that multiple (at least four) genetic lineages of Cordylophora spp., possibly representing cryptic species, have been introduced in Europe, North America and South America. Most of these lineages have been widely distributed, and multiple lineages can occur at the same site. One lineage (2B) was confined to the Pacific Coast of North America, although it co-occurred with a more-widely distributed lineage (1B). Lineages differed ecologically- 1A was found only in freshwater, 2A and 2B only in brackish waters, while 1B was found at both fresh and brackish sites. This study was wide-ranging, but did not include the Ponto-Caspian basin, the presumed region of origin of the Cordylophora species complex (Folino-Rorem et al. 2009). Until the genetic diversity of the complex is further clarified we will continue to treat it as a single species of Ponto-Caspian origin.

North American Invasion History:

Invasion History on the West Coast:

Sometime between the 1920s and the 1950s, Cordylophora caspia was discovered in Pacific coast estuaries including San Francisco Bay, the Columbia River, and Puget Sound (Hand and Gwillam 1951; Carlton 1979; Cohen and Carlton 1995). The earliest record appears to be from freshwater – Lake Union, Seattle, Washington (Hand and Gwilliam 1951; Carlton 1979). In 1930, it was found at Antioch on the San Joaquin River (Hand and Gwilliam 1951), and has subsequently been found at many locations in the Delta and in other fresh and brackish tributaries of San Francisco Bay (Hand and Gwillam 1951; Carlton 1979; Cohen and Carlton 1995; Cohen et al. 2005). This hydroid has been found in many fresh and brackish Pacific tributaries, including Elkhorn Slough, California (in 1998, Wasson et al. 2001); Humboldt Bay, California (in 1968, Mace and Mackie 1970; Carlton 1979); Coos Bay, Oregon (in 1959, Mace and Mackie 1970; Carlton 1979); Alsea Bay, Oregon (in 1975, Carlton 1979); the Columbia River estuary (in 1965, Haertel and Osterberg 1967; Carlton 1979); Willapa Bay, Washington (Cohen et al. 2001); brackish Puget Sound tributaries (Cohen et al. 1998; Cohen et al. 2001); and Albert Head Lagoon in Victoria, British Columbia (in 1967, Mace and Mackie 1970). Folino-Rorem et al. (2009) found two genetic lineages of C. caspia on the West Coast, one (1B) widespread in North America and Europe, the other (2B) confined to the Pacific Coast. Both lineages were found at Pittsburg, California, in the inner San Francisco estuary (Folino-Rorem et al. 2009).

Invasion History on the East Coast:

The first published record of Cordylophora caspia in North America was that of Leidy (1870), in which he described its occurrence in the Schuykill River, a Delaware River tributary in Philadelphia, but he remarked on finding it at Newport, Rhode Island 'some years earlier'. It apparently spread up and down the coast, reaching brackish ponds on Martha's Vineyard in 1872 (Verrill and Smith 1873), and Chesapeake Bay by 1877 (Clarke 1878; Bibbins 1892). The spread of this hydroid on the East coast seems to have been quite spotty. The first record for the Miramichi estuary, New Brunswick, was 1912 (Fraser 1944); it was found in 1926 in Back Bay, Virginia (an arm of Currituck Sound) (USNM 42191, U.S. National Museum of Natural History 2007); in 1928 in Pamlico Sound, North Carolina (Pearse 1936); and in 1932 in Charles River, Massachusetts (Blake 1932). However, first records for some well-travelled and populated estuaries were much later: 1972 for the Hudson River, New York (Mills et al. 1997); 1974 for the Cooper and Ashepoo Rivers, South Carolina (Calder and Hester 1978); and 1995 for the Connecticut River (Smith et al. 2002). It is possible that this hydroid was overlooked in some estuaries for decades before its published discovery.

In the Great Lakes, C. caspia was first collected in Lake Erie in 1956, and was abundant by the 1960s (Davis 1957; Hubschman 1971; Hubschman and Kishler 1972). It has been collected from Lake Ontario (Rochester, New York) (Folino-Rorem et al. 2009) to Duluth, Michigan at the western end of Lake Superior (Grigorevich et al. 2003), and in the Finger Lakes, in the Great Lakes Basin. Genetic analysis indicates that C. caspia has been spread largely through sexually produced larvae which have settled on boats, ships, and barges (Darling and Folino-Rorem 2009).

Invasion History on the Gulf Coast:

Cordylophora caspia was collected in Shreveport, Louisiana, in the Red River in 1918, about 600 km from the Gulf; and by 1944, was collected in Lake Pontchartrain (Fraser 1944). It has subsequently been found in tributaries along the northern shore of the Gulf from the Suwanee River, Florida (Mason et al. 1994) to the Sabine River, Texas (McClung et al. 1978). In the Escambia River, Florida (in 1952-53), C. caspia was 'an abundant population occurring on saw grass stalks and roots' (Wurtz and Roback 1955). It was found at 23 locations in the New Orleans area and Vermillion Parish, Louisiana, including tidal fresh and brackish waters (Poirrier and Denoux 1973).

This hydroid has spread widely through the fresh waters of the Mississippi Basin. In 1909, it was collected in the Illinois River (Havana, Illinois) (Smith 1910), and spread in a scattered fashion to Kentucky (in 1922, Garman, cited by Hubschman and Kishler 1972), Louisiana (in 1918, Poirrier and Denoux 1973), Oklahoma (in~1968, Ransom 1981), and Kansas (in 1980, Ransom 1981).

Invasion History in Hawaii:

Cordylophora caspia was collected in 1967, in Kaneohe Bay, Oahu (Powers 1971, cited by Coles et al. 2002), and in 1974-1975 in an anchialine pond (connected to the sea through porous rock) on Maui (Cooke 1977, cited by Carlton and Eldredge 2009).

Invasion History Elsewhere in the World:

In Europe, Cordylophora caspia was probably first introduced in the late 17th century through canals linking the waterways of the Baltic and Black Seas (Olenin 2002). It spread rapidly, and was widespread in inland waters and estuaries in northern Europe, from Finland to the British Isles by the late 19th century (Allman 1872; Arndt 1984; Jensen and Knudsen 2005; Wolff 2005). More recently, C. caspia spread north to Bergen and Stavanger, Norway (by 1985, Hopkins 2002) and south to the Guadalquivir River, Spain (by 2001, Escot et al. 2003, cited by Garcia-Berthou et al. 2007), the Po River delta, Italy (Morri and Bianchi 1983), and other Mediterranean lagoons. It is widespread in the major freshwater rivers of Europe, including the Rhine (Vervoort 1964; Roos 1979), the Elbe (Nehring 2006), Weser, Oder, and Danube basins (Bij de Vaate et al. 2002).

In Central America, C. caspia was first collected on the Caribbean side of the Panama Canal in the Gatun Locks in 1925 (Hildebrand 1939; Fraser 1944; USNM 43378, U.S. National Museum of Natural History 2007). Subsequently, it was collected on the Pacific side in the Pedro Miguel Locks in 1975 (Arndt 1984, Cohen 2006; USNHM 89237, U.S. National Museum of Natural History 2007). Specimens have also been collected from Gamboa, Panama, on Gatun Lake, and they belonged to the widespread freshwater genotype 1A (Folino-Rorem et al. 2009). In South America, it is known from the coast and freshwaters of Brazil (Arndt 1984; Grohmann 2008; Farrapeira et al. 2011), Uruguay, Argentina (Grohmann 2008), and from Chilean fjords (in 2006, Galea 2007, Folino-Rorem et al. 2009).

Cordylophora caspia has been introduced to freshwater lakes and brackish estuaries in New Zealand (1st record 1885, Cranfield et al. 1998), Australia (1st Record 1922, Briggs 1931, Hewitt, personal communication), Iraq (Shatt-al-Arab estuary) and Shanghai, China (Arndt 1984).


Description

Cordylophora caspia grows in erect, branching colonies, growing from a single stem, arising from stolons attached to the substrate. Sometimes one colony will grow on top of another. The shoots branch regularly or irregularly. Frequently, there is one main stem and shorter side branches. Annulations (rings) are present near the base of the stems and branches. A colony may have 40 or more hydranths. The hydranths are spindle-shaped when relaxed, ovoid when contracted, about 1-2 mm high, with a conical or bullet-shaped hypostome. There are scattered (usually 14-16, sometimes up to 27) tentacles. The gonophores are oval, arising from the stem or branches, and contain 7-16 eggs. Hydranths are white or pale pink and the stems are yellowish-brown. Stems are typically 30-45 mm in size, but may reach 150 mm or more (Calder 1971; Schuchert 2004; Calder 2010).


Taxonomy

Taxonomic Tree

Kingdom:   Animalia
Phylum:   Cnidaria
Class:   Hydrozoa
Subclass:   Hydroidolina
Order:   Anthoathecatae
Suborder:   Filifera
Family:   Cordylophoridae
Genus:   Cordylophora
Species:   caspia

Synonyms

Bimeria baltica (Stechow, 1923)
Cordylophora americana (Leidy, 1870)
Cordylophora lacustris (Stechow, 1927)
Cordylophora whiteleggei (von Lendenfeld, 1886)
Tubularia caspia (Pallas, 1771)
Tubularia cornea (Agardh, 1816)
Cordylophora albicola (Kirchenpauer, in Busk, 1861)
Cordylophora lacustris var. otagoensis (Fyfe, 1929)

Potentially Misidentified Species

Garveia franciscana
Vervoort (1964) lists several cases in which these species have been confused. However, he notes morphological differences and attributes the confusion to the fact that these are the only two large hydroids species that occur abundantly in temperate brackish waters.

Ecology

General:

Cordylophora caspia is a sessile hydrozoan which lacks a planktonic medusa stage. Colonies grow on a solid substrates, 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 produce multiple eggs, typically 7-16, which are brooded and fertilized by sperm from the water column. The eggs develop into ciliated non-feeding planula larvae (Schuchert 2004). These larvae probably spend less than a day in the water before settlement (Sommer 1992).

Planulae of C. caspia settle and grow on a wide range of substrates, including shells, rock, wood, and vegetation. This hydroid has been found on a number of plants, including submerged plants (Ceratophyllum demersum- Coontail; Nitella sp.; Potamogeton sp.- Pondweeds; Elodea sp.- Waterweed; Vallisneria americana- Wild Celery), stalks of floating plants (Nymphaea odorata- White Water Lily), and roots and stems of emergents (Alternanthera philoxeroides- Alligatorweed; Phragmites australis - Common Reed) (Clarke 1878; Poirrier and Denoux 1973; Calder 1978; Roos 1979). It has also been reported from shells of living freshwater (native Unionidae, introduced Dreissena spp.- Zebra and Quagga Mussels) and brackish-water mussels (Mytilopsis leucophaeta- Dark False Mussel) (Calder 1978; Curry et al. 1981; Walton 1996). It has also been found on man-made substrates including old automobiles and nylon ropes (Roos 1979), buoys and ships (Woods Hole Oceanographic Institution 1952), and doubtless many others.

Cordylophora caspia grows in a vast range of aquatic environments, varying in salinity, temperature, currents, oxygen, etc. Survival tolerances vary greatly among populations as a result of both genetics and acclimation (Arndt 1984; Kinne 1956). Experimental limits were 24⁰C for C. caspia from Germany (Kinne 1956) and 30+⁰C for animals collected near Woods Hole, Massachusetts (Fulton 1962). Optimal temperatures for asexual growth appeared to be 11-18⁰C for German populations (Kinne 1956), 18-26⁰C for MA populations (Fulton 1962), 16-25⁰C for San Francisco estuary populations (Meek et al. 2012), and 23-30⁰C for colonies from Iraq (Arndt 1984). At least one genetic lineage of C. caspia (1A) ranges from Europe (UK and Germany) to the tropics (Panama) (Folino-Rorem et al. 2009). This hydroid is known to grow abundantly in fresh and brackish waters, and can tolerate exposure to full seawater (Kinne 1958; Mace and Mackie 1970; Arndt 1984). Tolerance to seawater may vary genetically. One lineage in Folino-Rorem et al.'s (2009) study was restricted to fresh water (lineage 1A), one to brackish water (2), and one inhabited both (1B). Cordylophora caspia is also tolerant of hypoxia, and had optimal growth at 40% of saturation (Fulton 1962). In the San Francisco estuary, it was associated with middle levels of oxygen concentration, low salinity, and low transparency (Wintzer et al. 2011a). This hydroid can respond to unfavorable conditions by regressing into a dormant state, consisting of bodies of tissue (menonts) in the stolons and stems, which serve as a diapause stage (Kinne 1956; Roos 1979; Jormalainen et al. 1994). In regions with milder climates, such as San Francisco Bay (Wintzer et al. 2011a) and South Carolina (Calder 1992) the hydroid is active all year, with extended polyps. In more severe climates, such as the Netherlands and Finland, hydroids may be dormant in winter (Roos 1979; Jormalainen et al. 1994) and sometimes also during periods of high predation in mid-summer (Jormalainen et al. 1994).

Food:

Zooplankton; Small epibenthos

Consumers:

Tenellia adspersa; amphipods, fishes

Trophic Status:

Carnivore

Carn

Habitats

General HabitatVessel HullNone
General HabitatSwampNone
General HabitatSalt-brackish marshNone
General HabitatOyster ReefNone
General HabitatFresh (nontidal) MarshNone
General HabitatGrass BedNone
General HabitatCanalsNone
General HabitatRockyNone
General HabitatMarinas & DocksNone
General HabitatTidal Fresh MarshNone
General HabitatNontidal FreshwaterNone
General HabitatCoarse Woody DebrisNone
Salinity RangeLimnetic0-0.5 PSU
Salinity RangeOligohaline0.5-5 PSU
Salinity RangeMesohaline5-18 PSU
Salinity RangePolyhaline18-30 PSU
Tidal RangeSubtidalNone
Vertical HabitatEpibenthicNone


Tolerances and Life History Parameters

Minimum Temperature (ºC)0Based on geographic range
Maximum Temperature (ºC)30Experimental upper limits were 24 C for C. caspia from Germany (Kinne 1956) and 30+ C for animals collected near Woods Hole MA (Fulton 1962)
Minimum Salinity (‰)0This hydroid grows and reproduces in freshwater.
Maximum Salinity (‰)35Upper salinity ranges are based on experimental survival. The upper limit for sexual reproduction was 27 ppt ( Kinne 1958). Field salinity ranges are generally much lower (Arndt 1984; Lippson et al. 1979; Poirrier and Denoux 1973; Ruiz et al. unpublished data).
Minimum pH6.2Field data from TX (McClung and Davis 1983) and LA (Poirrier and Denoux 1973). Optimal growth in experiments occurred at 6.8-8.6, but no growth occurred at 5.1 (Fulton 1962).
Maximum pH8.6None
Minimum Duration0.5Sommer 1992
Maximum Duration1Sommer 1992
Maximum Height (mm)150Stems are typically 30 45 mm, but may reach 150 mm or more (Calder 1971; Schuchert 2004; Calder 2010)
Broad Temperature RangeNoneCold-temperate-Tropical
Broad Salinity RangeNoneFresh-Polyhaline

General Impacts

Economic impacts 

Cordylophora caspia has been reported from the cooling water systems of power plants in Illinois (Folino 2000), Brazil (Grohmann 2009), Finland (Vuorinen et al. 1984), Luxembourg (Massard and Geimer 1990), and the Ukraine (Simkina 1963). Fouling by C. caspia has required the shutdown of generators for cleaning and the use of toxic chemicals such as chlorine to prevent fouling (Folino 2000; Grohmann 2008). Other impacts are possible. It occurs in fouling on boats, ships, and buoys (Woods Hole Oceanographic Institution 1952), but is not widely reported as a ship fouling problem.

Ecological impacts

Cordylophora caspia is now a significant biomass component of the fouling community in the fresh-mesohaline regions of many estuaries around the world. It is also the only erect compound hydroid occurring in inland freshwater lakes (Pennak 1978; Hutchinson 1993). However, information on the ecological impact of its invasion is scarce, and mostly anecdotal, although some experimental studies have been conducted in Chesapeake Bay (see Von Holle and Ruiz 1997; Von Holle unpublished data).

Competition - Cordylophora caspia is a potential competitor for space in fouling communities. In field experiments on fouling plates (Key Bridge, Patapsco River, Maryland), where laboratory-grown colonies of C. caspia were added, abundances of the bryozoan Victorella pavida (cryptogenic), the entoproct Loxosomatoides laevis (introduced), and the protozoans Metafolliculina sp., and Stentor sp. were reduced (Von Holle and Ruiz 1997; Von Holle unpublished data).

Food/Prey - Cordylophora caspia is an important food for nudibranchs, which include many specialized predators of hydroids. Cordylophora caspia is apparently eaten by the nudibranch Tenellia adspersa, cryptogenic on the East Coast of North America, but widely introduced elsewhere (Gaulin et al. 1986; Chester 1996). Despite its nematocysts, C. caspia is also eaten by some generalized predators, such as amphipods (Roos 1979). Extensive feeding by the introduced amphipod Gammarus tigrinus (native in Chesapeake Bay) on C.caspia was reported in Dutch freshwaters by Roos (1979). In the San Francisco Bay estuary, C.caspia comprised 18-23% of the diet of the introduced Shimofuri goby (Tridentiger bifasciatus) (Matern and Brown 2005).

Predation -Although colonies of Cordylophora caspia in many bodies of water represent a substantial biomass of predators on zooplankton and mobile epibenthos (Bibbins 1892; Arndt 1984; Roos 1979), their role as predators has rarely been studied quantitatively. However, C. caspia predates on settling Zebra Mussel (Dreissena polymorpha) veligers, selecting smaller veligers, even as their filaments increase overall rates of settlement (Folino-Rorem and Stoeckel 2006).

Habitat Change - Cordylophora caspia colonies are dense and bushy, and constitute a substantial structural alteration to surfaces of wood, plants, rocks, etc., which may provide some protection from predators and currents (Roos 1979). In field experiments, the addition of laboratory-grown colonies of C. caspia resulted in increased abundances of Amphibalanus improvisus, Alitta succinea, and corophiid amphipods on fouling plates (Von Holle and Ruiz 1997; Von Holle unpublished data). In the Great Lakes basin, colonies of C. caspia provide substrate for settlement of larval Zebra Mussels (Dreissena polymorpha) (Folino-Rorem et al. 2006). This hydroid has been recorded as a fouling organism on living native freshwater bivalves (Amblema plicata, Potamilus purpuratus) in Louisiana (Curry et al. 1981), Zebra Mussels in the Great Lakes region (Folino-Rorem et al. 2006), and Zebra Mussels and Dark False Mussels (Mytilopsis leucophaeta) in the Hudson River (Walton 1996). However, impacts of C. caspia on these bivalves were not reported. 


Regional Impacts

NEP-VNorthern California to Mid Channel IslandsEcological ImpactFood/Prey
In the San Francisco Bay estuary, C. caspia comprised 18-23% of the diet of the introduced Shimofuri goby (Tridentiger bifasciatus) (Matern and Brown 2005).
P090San Francisco BayEcological ImpactFood/Prey
In the San Francisco Bay estuary, C. caspia comprised 18-23% of the diet of the introduced Shimofuri goby (Tridentiger bifasciatus) (Matern and Brown 2005)
CACaliforniaEcological ImpactFood/Prey
In the San Francisco Bay estuary, C. caspia comprised 18-23% of the diet of the introduced Shimofuri goby (Tridentiger bifasciatus) (Matern and Brown 2005)., In the San Francisco Bay estuary, C. caspia comprised 18-23% of the diet of the introduced Shimofuri goby (Tridentiger bifasciatus) (Matern and Brown 2005)

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
P080 Monterey Bay 1998 Non-native Established
P135 _CDA_P135 (Mad-Redwood) 1975 Non-native Established
P114 _CDA_P114 (Gualala-Salmon) 1975 Non-native Established
P130 Humboldt Bay 1968 Non-native Established
NEP-IV Puget Sound to Northern California 1959 Non-native Established
P093 _CDA_P093 (San Pablo Bay) 1951 Non-native Established
NEP-V Northern California to Mid Channel Islands 1930 Non-native Established
P090 San Francisco Bay 1930 Non-native Established

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude
698428 Wasson et al. 2001 (Elkhorn Slough Survey) 1998 Elkhorn Slough Station 1 (North side of bridge over Old Salinas River channel; south of Moss Landing) Non-native 36.7908 -121.7906
698900 Hand and Gwilliam 1951; Cohen and Carlton 1995 1930 San Joaquin River at Antioch Non-native 38.0172 -121.8078
699641 Fraser 1948; R.C. Miller, pers. comm., in Hand and Gwilliam 1951; Miller 1958 1946 Lake Merced, San Francisco Bay Non-native 37.7202 -122.4962
702982 Cohen et al. 2005 (SF Bay Area RAS) 2004 2004-05-25 Petaluma River Turning Basin, San Pablo Bay Non-native 38.2355 -122.6382
704507 Cohen et al. 2005 (SF Bay Area RAS) 2004 2004-05-28 Napa Valley Marina, San Pablo Bay Non-native 38.2200 -122.3128
704526 Introduced Species Study 2010 2010-06-30 Napa Valley Marina Non-native 38.2198 -122.3119
711912 J. Standing, pers. comm. 1975, in Carlton 1979 1975 Freshwater Lagoon Non-native 41.2692 -124.0914
711913 C. Hand, pers. comm. 1968, in Mace and Mackie 1970, cited in Carlton 1979 1968 Humboldt Bay Non-native 40.7500 -124.2083
715810 Wintzer et al. 2011 2007 Boynton Slough Non-native 38.2129 -122.0532
759493 H. Kirby, Jr. (U.C. Berkeley), pers. comm., in Hand and Gwilliam 1951 1951 San Pablo Reservoir Non-native 37.9288 -122.2457
759494 H. Kirby, Jr. (U.C. Berkeley), pers. comm., in Miller 1958 1946 Richmond Wastewater Filtration Plant Non-native 37.9190 -122.3779
759495 Aldrich 1961 1955 San Joaquin River at Antioch Non-native 38.0258 -121.7547
759496 Aldrich 1961 1955 San Joaquin River at Antioch Non-native 38.0258 -121.7547
759497 Aldrich 1961 1955 San Joaquin River at Bradford Non-native 38.0944 -121.6711
759498 J. Standing and C. Mills, pers. comm. 1975, in Carlton 1979 1975 Salmon Creek Non-native 38.3523 -123.0616
759499 Wintzer et al. 2011 2007 Montezuma Slough Non-native 38.1712 -121.9602
759500 Wintzer et al. 2011 2008 Boynton Slough Non-native 38.2129 -122.0532
759501 Wintzer et al. 2011 2008 Montezuma Slough Non-native 38.1712 -121.9602
759502 Wintzer et al. 2011 2008 Suisun Slough Non-native 38.1385 -122.0813

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