Invasion History

First Non-native North American Tidal Record: 1859
First Non-native West Coast Tidal Record: 1859
First Non-native East/Gulf Coast Tidal Record:

General Invasion History:

Ectopleura crocea (also known as Pinauay crocea or Tubularia crocea) was first described from 'Boston Bay', Massachusetts by Louis Agassiz in 1862 (as Parypha crocea). On the Atlantic coast of North America, its presumed native region, E. crocea has been collected from the Miramichi Estuary, New Brunswick, south to Lake Worth, Florida and Port Aransas, Texas (Fraser 1944; Deevey 1950; Defenbaugh 1973; Ruiz et al. unpublished data). It has also been collected on the Caribbean shore of South America at Santa Marta, Colombia (Wedler 1975), where we consider it cryptogenic. Ectopleura crocea has a wide global distribution and was apparently introduced to non-native locations starting in or before the 19th century (Deevey 1950; Carlton 1979; Watson 1999).

North American Invasion History:

Invasion History on the West Coast:

Even before its description in Massachusetts, this hydroid was collected from San Francisco Bay, California (as Parypha microcephala, Agassiz 1859, cited by Carlton 1979). In San Francisco Bay, it occurs in the Central Bay, the South Bay, and San Pablo Bay (Carlton 1979; Cohen and Carlton 1995; Cohen et al. 2005; Ruiz et al. unpublished data).To the south, E. crocea was collected from the major ports of San Diego (in 1876, Clark 1876, cited by Carlton 1979) and Los Angeles (in 1902, Torrey 1902, cited by Carlton 1979), before it was recorded from the smaller coastal bays: Mission Bay (in 1896, USNM 43547, U.S. Museum of Natural History 2007), Newport Bay (in 1938, Carlton 1979, USNM 43543, U.S. Museum of Natural History 2007), Kings Harbor, Santa Monica Bay (Fraser 1948, cited by Carlton 1979), and Elkhorn Slough (MacGinitie 1935, cited by Carlton 1979, Wasson et al. 2001).

To the north, E. crocea was reported from Port Simpson, British Columbia (54.6 N) in 1911. Early collections were also made from the San Juan Islands (Washington), north to the Queen Charlotte Islands, and an unspecified record from 'the Gulf of Alaska' (Fraser 1937; Carlton 1979). Mills (in Cohen et al. 1998) considered the reports from San Juan Islands to be unverified, and perhaps the result of taxonomic confusion. However, this hydroid was found on fouling plates in Ketchikan, Alaska, in 2003 (Ruiz et al. 2006). We do not know if this hydroid is established in Alaska or British Columbia. Ectopleura crocea was collected in Coos Bay, Oregon starting in 1948 (Fraser 1948, cited by Carlton 1979; Carlton 1989; Ruiz et al., unpublished data), and on fouling plates in Humboldt Bay, California in 2003 (Ruiz et al., unpublished data).

Invasion History Elsewhere in the World:

Ectopleura crocea has been reported from the tropical Pacific at Jicaron Island, Panama (Fraser 1938), from fouling plates in Salinas, Ecuador (2018, Calder et al. 2021), and the southeastern Pacific at Valparaiso, Chile (1905, Deevey 1950). In the southwestern Pacific, it was first collected in Victoria, Australia, in Port Phillip Bay (as Tubularia ralphii, Bale 1884, cited by Watson 1999). It has also been collected in Sydney Harbor and Port Kembla, in New South Wales, at Fremantle, in Western Australia (Watson 1999), at Auckland, New Zealand (Cranfield et al. 1998). In the northeast Atlantic, E. crocea appears to be an introduction. It was first collected in the Azores in 1989 (Cardigos et al. 2006) and is also known from Madeira (Wirtz 2007). It was noted as a rare occurrence on ship hulls at Plymouth, England in 1895 and 1907 (Plymouth Marine Fauna, http://www.mba.ac.uk/pmf/) and at Ipswich, England in 1959 (Rees 1963). It is not listed as a Mediterranean invader by Galil (2009), but available references suggest that it is mostly known from harbors (e.g., Villefranche-sur-mer, France, 1895, Schuchert 2010; Bay of Naples, Italy, 1892, Bouillon et al. 2004; Israel 1946, Vervoort 1993). In the southwest Atlantic, it occurs from Uruguay to Bahia Blanca, Argentina (Genzano et al. 2005). It was first reported from this region in 1971 and is considered cryptogenic there (Orensanz et al. 2003). In South Africa, it was found in Durban and Cape Town in 1947 (Ewer 1953, cited by Millard 1975, as Tubularia warreni; Schuchert 2010; (1947, Millard 1952, cited by Mead et al. 2011). In the Northwest Pacific, Tubularia sagaminea and T. mesembryanthemum, reported from Japan (Stechow 1907; Yamada 1959; Hirohito 1988) and China (Hargitt 1927, Yamada 1959. all cited by Imazu et al. 2014), are all considered synonyms of E. crocea.


Description

Ectopleura crocea, also commonly known as Pinauay or Tubularia crocea, is a hydrozoan which lacks a medusa stage. Its colonies grow from branching stolons, in tangled masses up to 100–120 mm in height, and consist of up to several hundred unbranched stems, with one hydranth per stalk. The perisarc is wrinkled with a few annulations, but there are no joints in the stalk. The hydranth is vase-shaped with a long hypostome. The tentacles are threadlike, in two whorls of 20–24 each. The proximal-whorl tentacles are larger and longer than those in the distal whorl. The female gonophores are carried on 12–16 blastostyles, hanging below the tentacles. The female gonophores produce eggs, which develop into planktonic actinula larvae resembling miniature hydranths, usually with four tentacles. Production of these larvae may vary regionally; being rare (West Coast, Fraser 1937) or frequent (Chesapeake Bay, Calder 1971). The male gonophores are oval or spherical, without apical processes. The body of the hydranth is pink (description from: Fraser 1937; Calder 1971; Watson 1999; Schuchert 2010).

The correct genus name for this hydroid is disputed. Marques and Migotto (2000) published a cladistic analysis of the genus Ectopleura which supported the monophyly of the genus, but found that genus consisted of two subclades, and put several widespread species including E. larynxE. crocea, and E. marina into a new genus, Pinauay. Schuchert (2010) considers the split to be unjustified. Imazu et al. (2014) review the taxonomy and distribution of this hydroid, using the name E. crocea. They tentatively support the synonymy of Western Atlantic E. crocea with E. ralphi, as described from Australia and based on Brazilian specimens, but suggest that worldwide morphological and genetic comparisons are needed.


Taxonomy

Taxonomic Tree

Kingdom:   Animalia
Phylum:   Cnidaria
Class:   Hydrozoa
Subclass:   Hydroidolina
Order:   Anthoathecatae
Suborder:   Filifera
Family:   Tubulariidae
Genus:   Ectopleura
Species:   crocea

Synonyms

Parypha microcephala (L. Agassiz, 1859)
Paryphya crocea (L. Agassiz, 1862)
Pinauay crocea (Marques and Migotto, 2000)
Tubularia crocea (Allman, 1872)
Tubularia mesembryanthemum (Allman, 1871)
Tubularia polycarpa (Allman, 1872)
Tubularia ralphi (Bale, 1884)
Tubularia gracilis (von Lendenfeld, 1885)
Tubularia australis (Stechow, 1924)
Tubularia warreni (Ewer, 1953)
Tubularia sagamina (Stechow, 1907)
Ectopleura media (Fraser, 1938)

Potentially Misidentified Species

Ectopleura integra
Native to the Galapgos, releases medusae (Calder et al. 2019)

Ectopleura marina
West Coast on open shores (Mills et al., in Carlton 2007)

Ecology

General:

The hydroid Ectopleura crocea lacks a free-living medusa. It does have a short-lived (~24 hours) planktonic lecithotrophic larval form, known as an actinula, resembling a small sea-anemone. It grows on a solid substrate, with polyps arising from branching, creeping stolons. The polyps grow as single stalks, each bearing a hydranth, whose tentacles capture zooplankton. The polyps produce bunches of gonophores, which produce either eggs or sperm. Colonies are diecious (single-sexed). Female gonophores produce multiple eggs, typically 2–4, which are brooded and fertilized by sperm in the water column. The egg develops in the gonophore through the planula stage into an actinula (Barnes 1983; Bouillon et al. 2004; Schuchert 2010). Larvae spend about 24 hours in the water column (as Ectopleura mesembryanthemum; Yamashita et al. 2003).

This hydroid occurs on a variety of substrates, including rocks, shells, concrete, pilings, buoys, jetties, pipes, and ships’ hulls (Fraser 1944; Woods Hole Oceanographic Institution 1952; Calder 1971; Gosner 1978). It is characteristic of harbors and polluted waters (Bouillon et al. 2004; Schuchert 2010). In South Carolina, it was found at a salinity range of 23–34 PSU (Calder 1976). The occurrences of Ectopleura crocea in Salinas, Ecuador, with mean water temperature of 24 °C, increases the known temperature tolerance of this hydroid (Calder et al. 2021).

Food:

Zooplankton, small epibenthos

Consumers:

Nudibranchs

Competitors:

Trophic Status:

Carnivore

Carn

Habitats

General HabitatOyster ReefNone
General HabitatCoarse Woody DebrisNone
General HabitatMarinas & DocksNone
General HabitatRockyNone
General HabitatVessel HullNone
Salinity RangePolyhaline18-30 PSU
Salinity RangeEuhaline30-40 PSU
Tidal RangeSubtidalNone
Vertical HabitatEpibenthicNone

Life History


Tolerances and Life History Parameters

Minimum Temperature (ºC)0Based on geographical range
Maximum Temperature (ºC)30Charleston Harbor SC (Calder 1992)
Minimum Salinity (‰)23Field distribution, SC (Calder1976)
Maximum Salinity (‰)34Field distribution, SC (Calder1976)
Maximum Duration1Yamashita et al. 2003, for Ectopleura mesembryanthemum
Broad Temperature RangeNoneCold temperate-Tropical
Broad Salinity RangeNonePolyhaline-Euhaline

General Impacts

Ectopleura crocea is frequently an abundant fouling organism in its native and introduced ranges. It seems to prefer man-made structures and is tolerant of polluted waters (Schuchert 2010). It is also occurs on mussel shells and around mussel beds, and is a potential competitor with mussels and a possible predator on their larvae (Okamura 1986; Fitridge 2011).

Economic impacts

Ectopleura crocea has been reported from pilings, buoys, jetties, pipes, and ship hulls (Fraser 1944; Woods Hole Oceanographic Institution 1952; Calder 1971; Gosner 1978). It is probably an important contributor to fouling communities because of its size and frequent abundance. However, specific impacts on shipping have not been reported.

Fisheries- Ectopleura crocea fouls cultured mussels (Mytilus galloprovincialis) in Port Phillip Bay, Australia, with adverse effects on their growth and condition, possibly due to competition for food, and on the recruitment of larvae due to predation (Fitridge 2011).

Ecological Impacts

Competition-Ectopleura crocea was a dominant form on fouling plates in San Francisco Bay (Okamura 1986).

Habitat Change- The degenerating stalks of Ectopleura crocea provided a filamentous surface for metamorphosing larvae of Mytilus spp. on fouling plates in San Francisco Bay (Okamura 1986), and in Port Phillip Bay, Australia (Fitridge 2011).

Impacts in the Galapagos Islands

Impacts are unknown in the Galapagos Islands.


Regional Impacts

NEP-VNorthern California to Mid Channel IslandsEcological ImpactCompetition
Ectopleura crocea was a dominant form on fouling plates in San Francisco Bay in late summer and fall, occupying up to 60% of the plates' surface, reaching a peak in October (Okamura 1986).
NEP-VNorthern California to Mid Channel IslandsEcological ImpactHabitat Change
The degenerating stalks of Ectopleura crocea provided a filamentous surface for metamorphosing larvae of Mytilus spp. on fouling plates in San Francisco Bay in winter, facilitating the establishment of dense populations in spring (Okamura 1986).
P090San Francisco BayEcological ImpactCompetition
Ectopleura crocea was a dominant form on fouling plates in San Francisco Bay in late summer and fall, occupying up to 60% of the plates' surface, reaching a peak in October (Okamura 1986).
P090San Francisco BayEcological ImpactHabitat Change
The degenerating stalks of Ectopleura crocea provided a filamentous surface for metamorphosing larvae of Mytilus spp. on fouling plates in San Francisco Bay in winter, facilitating the establishment of dense populations in spring (Okamura 1986).
CACaliforniaEcological ImpactCompetition
Ectopleura crocea was a dominant form on fouling plates in San Francisco Bay in late summer and fall, occupying up to 60% of the plates' surface, reaching a peak in October (Okamura 1986)., Ectopleura crocea was a dominant form on fouling plates in San Francisco Bay in late summer and fall, occupying up to 60% of the plates' surface, reaching a peak in October (Okamura 1986).
CACaliforniaEcological ImpactHabitat Change
The degenerating stalks of Ectopleura crocea provided a filamentous surface for metamorphosing larvae of Mytilus spp. on fouling plates in San Francisco Bay in winter, facilitating the establishment of dense populations in spring (Okamura 1986)., The degenerating stalks of Ectopleura crocea provided a filamentous surface for metamorphosing larvae of Mytilus spp. on fouling plates in San Francisco Bay in winter, facilitating the establishment of dense populations in spring (Okamura 1986).

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
P130 Humboldt Bay 2003 Non-native Established
P070 Morro Bay 1986 Non-native Established
P060 Santa Monica Bay 1948 Non-native Established
NEP-IV Puget Sound to Northern California 1948 Non-native Established
P040 Newport Bay 1938 Non-native Established
P080 Monterey Bay 1935 Non-native Established
P050 San Pedro Bay 1902 Non-native Established
P030 Mission Bay 1896 Non-native Established
NEP-VI Pt. Conception to Southern Baja California 1876 Non-native Established
P020 San Diego Bay 1876 Non-native Established
P093 _CDA_P093 (San Pablo Bay) 1865 Non-native Established
P090 San Francisco Bay 1859 Non-native Established
NEP-V Northern California to Mid Channel Islands 1859 Non-native Established

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude
697086 Introduced Species Study 2005 2005-08-25 Ferry Terminal Pier Non-native 37.7945 -122.3917
697120 Wasson et al. 2001 (Elkhorn Slough Survey) 1998 Elkhorn Slough Station 9 (Kirby Park) Non-native 36.8398 -121.7435
697247 Cohen et al. 2005 (SF Bay Area RAS) 2004 2004-05-27 Pete's Harbor, San Francisco Bay Non-native 37.5006 -122.2242
697960 Introduced Species Study 2011 2011-04-21 Draw Bridge Non-native 33.7645 -118.2428
698246 Cohen et al. 2005 (SF Bay Area RAS) 2004 2004-05-23 Pier 39, San Francisco Bay Non-native 37.8114 -122.4098
699924 Introduced Species Study 2010 2010-05-31 Dumbarton Bridge Non-native 37.5070 -122.1168
700129 Wasson et al. 2001 (Elkhorn Slough Survey) 1998 Elkhorn Slough Station 5 (South Marsh Trail) Non-native 36.8193 -121.7378
700370 Wasson et al. 2001 (Elkhorn Slough Survey) 1998 Elkhorn Slough Station 3 (Skipper's) Non-native 36.8104 -121.7863
700457 Wasson et al. 2001 (Elkhorn Slough Survey) 1998 Elkhorn Slough Station 4 (Vierra's) Non-native 36.8090 -121.7841
700583 Introduced Species Study 2011 2011-05-16 Harbormaster Launch Ramp Non-native 36.6023 -121.8907
700604 Torrey 1902 1902 San Pedro [Los Angeles] Harbor Non-native 33.7371 -118.2765
700705 MacGinitie 1939 1938 1938-11-15 Newport Bay Non-native 33.6092 -117.9067
700953 MacGinitie 1935 1935 Salinas River, near confluence with Elkhorn Slough Non-native 36.8065 -121.7851
701171 Agassiz 1865 1859 San Francisco [Bay] Non-native 37.8494 -122.3681
702470 Clark 1876; Torrey 1902 1875 San Diego Bay Non-native 32.6717 -117.1439
703368 Fraser 1948 1948 off Redondo Beach Non-native 33.8396 -118.3932
703638 Cohen et al. 2005 (SF Bay Area RAS) 2004 2004-05-28 Rodeo Marina, San Pablo Bay Non-native 38.0391 -122.2711
712144 Ruiz et al., unpublished data (DOD) 2003 Cabrillo Isle Marina, San Diego Non-native 32.7265 -117.2009
712149 Reish 1972 1972 Los Angeles/Long Beach Harbor Complex Non-native 33.7632 -118.2526
712151 Campbell 1922 1920 Long Beach Non-native 33.7669 -118.1883
712155 Ruiz et al., unpublished data (DOD) 2000 Treasure Island Dock, San Francisco Non-native 37.8203 -122.3626
712156 Ruiz et al., unpublished data (DOD) 2000 Hunters Point, San Francisco Non-native 37.7165 -122.3632
712157 Ruiz et al., unpublished data (DOD) 2000 Dumbarton Swing (Railway) Bridge Non-native 37.4976 -122.1098
712158 Ruiz et al., unpublished data (DOD) 2000 Berkeley Fishing Pier, Berkeley Non-native 37.8600 -122.3222
712166 Ruiz et al., unpublished data (DOD) 2000 Port of Richmond Terminal 4, Richmond Non-native 37.9090 -122.3878
712167 Ruiz et al., unpublished data (DOD) 2000 Port of Oakland, Oakland Non-native 37.8102 -122.3230
712168 Ruiz et al., unpublished data (DOD) 2000 Romberg Tiburon Center, Tiburon Non-native 37.8909 -122.4463
712169 Ruiz et al., unpublished data (DOD) 2000 Port of San Francisco Pier 50 Non-native 37.7727 -122.3865
712170 Ruiz et al., unpublished data (DOD) 2000 Port of San Francisco Pier 23 Non-native 37.8045 -122.3985
712171 Ruiz et al., unpublished data (DOD) 2000 Port of San Francisco Pier 96 Non-native 37.7420 -122.3695
715014 Lambert 2005 2003 Kuiper Oyster Raft, Mad River Slough Non-native 40.8722 -124.1490
759521 National Museum of Natural History, Invertebrate Zoology Collection Database 1895 1895-07-08 San Diego [Bay] Non-native 32.6717 -117.1439
759522 National Museum of Natural History, Invertebrate Zoology Collection Database 1895 1895-07-08 Coronado (San Diego Bay) Non-native 32.7070 -117.1900
759523 California Academy of Sciences, Invertebrate Zoology Collection Database 1896 1896-07-04 San Pedro Non-native 33.7371 -118.2765
759524 Torrey 1902 1902 Oakland Harbor Non-native 37.7989 -122.3334
759525 Torrey 1904 1904 San Diego Bay Non-native 32.6717 -117.1439
759526 National Museum of Natural History, Invertebrate Zoology Collection Database 1912 1912-04-08 Albatross Station 5766 (Alameda Channel) Non-native 37.7931 -122.2896
759527 National Museum of Natural History, Invertebrate Zoology Collection Database 1912 1912-10-07 San Francisco Bay Non-native 37.8494 -122.3681
759528 MacGinitie 1935 1927 Salinas River, near confluence with Bennett Slough Non-native 36.8156 -121.7875
759529 California Academy of Sciences, Invertebrate Zoology Collection Database 1930 Elkhorn Slough Non-native 36.8086 -121.7856
759530 Whedon 1943, cited in WHOI 1952 1940 San Diego Bay Non-native 32.6717 -117.1439
759531 Graham and Gay 1945 1941 Fruitvale Avenue Bridge Non-native 37.7689 -122.2296
759532 Carpelan 1957 1951 Floodgate at Charleston Slough Non-native 37.4557 -122.1009
759533 Mariscal 1965 1965 Palo Alto Yacht Harbor Non-native 37.4584 -122.1052
759534 Campbell and Campbell 1968 1968 Newport Bay Non-native 33.6092 -117.9067
759535 Turner et al. 1969 1960 Santa Monica Artificial Reef Non-native 34.0094 -118.5297
759536 Reish 1972 1972 Marina del Rey Non-native 33.9722 -118.4522
759537 Reish 1972 1972 Newport Bay Non-native 33.6092 -117.9067
759538 Reish 1972 1972 Alamitos Bay Non-native 33.7545 -118.1231
759539 Behrens and Tuel 1977 1974 Pete's Harbor Non-native 37.5006 -122.2242
759540 Behrens and Tuel 1977 1974 Port of Redwood City Non-native 37.5124 -122.2106
759541 California Academy of Sciences, Invertebrate Zoology Collection Database 1976 1976-08-26 Elkhorn Slough at Kirby Park Non-native 36.8398 -121.7435
759542 California Academy of Sciences, Invertebrate Zoology Collection Database 2015 2003 2003-10-10 Pete's Harbor Non-native 37.5006 -122.2242
759543 Cohen and Chapman 2005 2005 2005-11-27 Buoy #2 (Central Bay) Non-native 37.8904 -122.4176
759544 Cohen and Chapman 2005 2005 2005-11-27 Buoy #2 (South Bay) Non-native 37.6958 -122.3414
759545 Cohen and Chapman 2005 2005 2005-11-27 Buoy # 8 Non-native 38.0293 -122.3718
759546 Cohen and Chapman 2005 2005 2005-11-27 Dumbarton Bridge (pylon) Non-native 37.5031 -122.1230
759547 Cohen and Chapman 2005 2005 2005-11-27 San Mateo Bridge (pylon) Non-native 37.5835 -122.2515
759548 Carlton and Hodder 1995 1987 1987-09-22 Humboldt Bay Non-native 40.7864 -124.1922
759549 Carlton and Hodder 1995 1987 1987-09-25 San Francisco Bay Non-native 37.8494 -122.3681
767666 Ruiz et al., 2015 2013 2013-07-16 Naval Base Point Loma, San Diego Bay, CA, California, USA Non-native 32.6886 -117.2343
768035 Ruiz et al., 2015 2012 2012-08-27 Port of San Francisco Pier 31, San Francisco Bay, CA, California, USA Non-native 37.8078 -122.4060
768168 Ruiz et al., 2015 2012 2012-09-05 Port of Oakland, San Francisco Bay, CA, California, USA Non-native 37.7987 -122.3228

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