Invasion History

First Non-native North American Tidal Record: 1986
First Non-native West Coast Tidal Record: 1994
First Non-native East/Gulf Coast Tidal Record: 1986

General Invasion History:

The colonial tunicate Polyandrocarpa zorritensis was described from Zorritos, Peru in 1931 (Van Name 1945) in the southeastern Pacific. The next published collections were from the southeastern Atlantic, in Brazil, at Santos and Itacurussa (near Cananeia) (Millar 1958). The native region of this tunicate could be either the Southwest Atlantic or Southeast Pacific. However, this tunicate has become widely introduced, as it is found on the East and West coasts of North America, Hawaii, Japan, and the Mediterranean. In the Mediterranean and possibly elsewhere, its growth seems to be favored by eutrophication (Brunetti and Mastrototaro 2004).

North American Invasion History:

Invasion History on the West Coast:

Polyandrocarpa zorritensis was first collected in Southern California at Oceanside Harbor in September, 1994, where it was abundant. During that same year, it was also found in Mission and San Diego Bays, but was rare (Lambert and Lambert 1998). In 1997, it was found in Alamitos Bay, Long Beach Harbor, and at King Harbor and Marina del Rey in Santa Monica (Lambert and Lambert 2003). In 2001, P. zorritensis was found on fouling plates in San Francisco Bay at two locations: Jack London Square, Oakland in the Central Bay, and in Redwood City in the South Bay (Ruiz et al. unpublished data). In 2001, it was also collected in Tomales Bay (Fairey et al. 2002). Polyandrocarpa zorritensis was found in Ensenada, Mexico, in 2000 (Lambert and Lambert 2003), and Bahia San Quintin, Baja California in 2005 (Rodriguez and Ibarra-Obando 2008).

Invasion History on the East Coast:

The date of arrival of Polyandrocarpa zorritensis on the East Coast is not known, but it was well-established in the Indian River Lagoon, near Fort Pierce, Florida by 1994 (Vazquez and Young 1996; Vazquez and Young 1998, Gretchen Lambert, personal communication). In 2005, it was found at Cape Marina, in Cape Canaveral, Florida (da Rocha, personal communication, Ruiz et al., unpublished data). In 2014, P. zorritensis was found in three marinas in Wrightsville Beach, North Carolina (Villalobos et al. 2017).

Invasion History on the Gulf Coast:

Polyandrocarpa zorritensis was reported from Clearwater Harbor, Florida, south of Tampa Bay in 2002 (Grechen Lambert, personal communication 2002), and from Port Isabel, Texas in 2004 (Lambert et al. 2005).

Invasion History in Hawaii:

In 1997, Polyandrocarpa zorritensis was found at several locations in Oahu, Hawaii, in Pearl Harbor (in 1997, Coles et al.1999a), Ke'ehi Lagoon, Honolulu (in 1997, Coles et al.1999b), and in Kaneohe Bay (in 1997, Coles et al. 2002). 

Invasion History Elsewhere in the World:

In 1991, the tunicate was found in southern Japan, at Kitakyushu, Kyushu, on the East China Sea, and at Kochi, on Shikoku (Nishikawa et al. 1993, cited by Iwasaki (2006). Polyandrocarpa zorritensis has also invaded the western Mediterranean Sea. In 1975, it was detected in the harbor of La Spezia, Italy, on the Tyrrhenian Sea (Brunetti 1978). In 2001, it was found in the Gulf of Taranto, near the tip of the Italian Peninsula (2001, Brunetti and Mastrototaro 2004). In Spanish waters, P. zorritensis has been found in the delta of the Ebro River (Turon and Perrera 1998). 


Description

Polyandrocarpa zorritensis is a colonial tunicate, whose colonies consist of a crowded mass or cluster of club-shaped zooids. The zooids are connected at the base of the colony by a tangled mass of root-like stolons. Individual zooids have an oval cross-section (meaning they are slightly compressed from side to side; Van Name 1945) and are free at their anterior end. The anterior end is rounded or truncated, and the oral and atrial siphons have four lobes, with two dark bands per lobe (Brunetti and Mastrototaro, 2004). Each zooid has its own tunic, though tunics of adjacent individuals may adhere to each other. The tunics are thin, tough and can be densely encrusted with sand. Zooids are pale brown to yellowish green in color (Lambert and Lambert 1998) and are up to 20 mm long and 3.8 mm diameter (Van Name 1945).


Taxonomy

Taxonomic Tree

Kingdom:   Animalia
Phylum:   Chordata
Subphylum:   Tunicata
Class:   Ascidiacea
Order:   Stolidobranchia
Family:   Styelidae
Genus:   Polyandrocarpa
Species:   zorritensis

Synonyms

Stolonica zorritensis (Van Name, 1931)

Potentially Misidentified Species

Ecology

General:

Life History- A colonial (or compound) tunicate consists of many zooids, bearing many or all of the organs of a solitary tunicate, but modified to varying degrees for colonial life. Colonial tunicates of the genus Polyandrocarpa have zooids resembling solitary tunicates, rounded, oval, or club-like in shape. Zooids can be crowded together, but do not coalesce. Each zooid has an oral and atrial siphon. Water is pumped into the oral siphon, through finely meshed ciliated gills on the pharynx, where phytoplankton and detritus are filtered, and passed on mucus strings to the stomach and intestines. Excess waste is expelled in the outgoing atrial water (Van Name 1945; Barnes 1983). 
 
Colonial tunicates reproduce both asexually, by budding, and sexually, from fertilized eggs developing into larvae. Colonies vary in size and can range from small clusters of zooids to huge spreading masses. The zooids are hermaphroditic, with eggs and sperm being produced by a single individual. Eggs may be self-fertilized or fertilized by sperm from nearby animals, but many species have a partial block to self-fertilization. Depending on the life-history of the species, eggs may be externally or internally fertilized. Fertilized eggs hatch into a tadpole larva with a muscular tail, notochord, eyespots, and a set of adhesive papillae. The lecithotrophic (non-feeding, yolk-dependent) larva swims briefly before settlement. Swimming periods are usually less than a day, and some larvae can settle immediately after release, but the larval period can be longer at lower temperatures. Once settled, the tail is absorbed, the gill basket expands, and the tunicate begins to feed by filtering (Van Name 1945; Barnes 1983). 

Food:

Phytoplankton, detritus

Trophic Status:

Suspension Feeder

SusFed

Habitats

General HabitatMarinas & DocksNone
General HabitatVessel HullNone
General HabitatMangrovesNone
Salinity RangePolyhaline18-30 PSU
Salinity RangeEuhaline30-40 PSU
Tidal RangeSubtidalNone
Vertical HabitatEpibenthicNone

Life History


Tolerances and Life History Parameters

Minimum Temperature (ºC)12Field, Mediterranean Sea (Brunetti, pers. comm., cited by Lambert and Lambert 1998)
Maximum Temperature (ºC)30Field, Mediterranean Sea (Brunetti, pers. comm., cited by Lambert and Lambert 1998)
Minimum Salinity (‰)22.7Field, Mediterranean Sea (Brunetti, pers. comm., cited by Lambert and Lambert 1998)
Maximum Salinity (‰)38Field, Mediterranean Sea (Brunetti, pers. comm., cited by Lambert and Lambert 1998)
Minimum Reproductive Salinity26Salinities below 26 PSU halted the swimming of larvae (Vazquez and Young 1998)
Maximum Duration0.2Larvae, Laboratory observations (Vazquez and Young 1998)
Broad Temperature RangeNoneWarm temperate-Tropical
Broad Salinity RangeNonePolyhaline-Euhaline

General Impacts

Economic Impacts-

Polyandrocarpa zorritensis was reported to overgrow cultured oysters (Crassostrea gigas) in the Ebro Delta, Spain (Perrera et al. 1990, cited by da Rocha et al. 2009).

Ecological impacts-

Polyandrocarpa zorritensis formed 100% cover in several locations in San Diego and Mission Bays in 1994–2000, suggesting it is capable of outcompeting native and other introduced fouling species (Lambert and Lambert 2003).


Regional Impacts

NEP-VIPt. Conception to Southern Baja CaliforniaEcological ImpactCompetition
Polyandrocarpa zorritensis formed 100% cover in several locations in San Diego and Mission Bays in 1994-2000, suggestive of competition (Lambert and Lambert 2003).
P020San Diego BayEcological ImpactCompetition
Polyandrocarpa zorritensis formed 100% cover in several locations in San Diego and Mission Bays in 1994-2000, suggestive of competition (Lambert and Lambert 2003).
P030Mission BayEcological ImpactCompetition
Polyandrocarpa zorritensis formed 100% cover in several locations in San Diego and Mission Bays in 1994-2000, suggestive of competition (Lambert and Lambert 2003).
CACaliforniaEcological ImpactCompetition

Polyandrocarpa zorritensis formed 100% cover in several locations in San Diego and Mission Bays in 1994-2000, suggestive of competition (Lambert and Lambert 2003)., 

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
P064 _CDA_P064 (Ventura) 2020 Def Estab
P065 _CDA_P065 (Santa Barbara Channel) 2020 Def Estab
P040 Newport Bay 2017 Def Estab
P062 _CDA_P062 (Calleguas) 2013 Def Estab
P027 _CDA_P027 (Aliso-San Onofre) 2011 Def Estab
P110 Tomales Bay 2001 Def Estab
P090 San Francisco Bay 2001 Def Estab
NEP-V Northern California to Mid Channel Islands 2000 Def Estab
P060 Santa Monica Bay 1997 Def Estab
P050 San Pedro Bay 1997 Def Estab
NEP-VI Pt. Conception to Southern Baja California 1994 Def Estab
P023 _CDA_P023 (San Louis Rey-Escondido) 1994 Def Estab
P030 Mission Bay 1994 Def Estab
P020 San Diego Bay 1994 Def Estab

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude
697134 Maloney et al. 2007 2005 2005-06-28 Navy - Seventh St. Def 32.6699 -117.1212
697226 Maloney et al. 2007 2005 2005-04-27 Kelco Dock Def 32.6927 -117.1470
697517 Introduced Species Study 2006 2006-08-25 Northern Corner of Harbor Def 33.9830 -118.4465
697743 Introduced Species Study 2006 2006-08-24 Yellow Condos Def 33.7283 -118.0602
697865 Introduced Species Study 2006 2006-09-12 Ski Islands Marina Def 32.7939 -117.2232
698114 Introduced Species Study 2006 2006-09-12 Hilton Resort Dock Def 32.7788 -117.2127
698115 Introduced Species Study 2011 2011-05-04 Hilton Resort Dock Def 32.7788 -117.2127
698139 ISS 2000-2002 Survey Data 2001 2001-07-11 Marina Del Rey Epifaunal 04 Def 33.9760 -118.4460
698358 Cohen et al. 2002 (So Cal Exotics RAS); Lambert and Lambert 2003 2000 2000-08-28 Oceanside Harbor Def 33.2094 -117.3947
698537 Maloney et al. 2007 2005 2005-04-27 Glorietta Bay Def 32.6796 -117.1740
698550 Cohen et al. 2002 (So Cal Exotics RAS); Lambert and Lambert 2003 2000 2000-08-28 Seaforth Landing Def 32.7644 -117.2381
698663 Introduced Species Study 2011 2011-05-03 Marine Terminal (Paco) Def 32.6584 -117.1191
698777 Maloney et al. 2007 2005 2005-06-28 Navy - 28th St. Def 32.6836 -117.1311
698843 Introduced Species Study 2006 2006-09-13 Guest Dock Def 33.2091 -117.3947
698876 ISS 2000-2002 Survey Data 2001 2001-08-14 Oceanside Epifaunal 03 Def 33.2090 -117.3956
698957 Introduced Species Study 2006 2006-09-12 Dana Inn Marina Def 32.7671 -117.2362
698962 Introduced Species Study 2011 2011-05-04 Dana Inn Marina Def 32.7671 -117.2362
699013 ISS 2000-2002 Survey Data 2000 2000-09-11 Bodega Bay Epifaunal 09 Def 38.3317 -123.0571
699093 Cohen et al. 2002 (So Cal Exotics RAS); Lambert and Lambert 2003 2000 2000-08-26 Chula Vista Boat Ramp (J Street) Def 32.6211 -117.1031
699603 Maloney et al. 2007 2005 2005-04-26 Bulk Carrier Terminal Def 32.6969 -117.1526
699604 ISS 2000-2002 Survey Data 2001 2001-10-09 Bulk Carrier Terminal Def 32.6969 -117.1526
699684 Cohen et al. 2002 (So Cal Exotics RAS); Lambert and Lambert 2003 2000 2000-08-29 King Harbor Def 33.8464 -118.3969
699697 Introduced Species Study 2006 2006-08-25 Vessel Assist Dock Def 33.9783 -118.4569
699718 Introduced Species Study 2011 2011-05-04 Seaforth Def 32.7621 -117.2365
699719 Introduced Species Study 2006 2006-09-12 Seaforth Def 32.7621 -117.2365
699757 Lambert and Lambert 1998; Lambert and Lambert 2003 1994 1994-08-27 Oceanside Harbor, by Harbor Patrol floats Def 33.2081 -117.3946
699797 Maloney et al. 2007 2005 2005-04-27 Coronado Wharf Def 32.6992 -117.1684
699798 ISS 2000-2002 Survey Data 2001 2001-10-09 Coronado Wharf Def 32.6992 -117.1684
699960 ISS 2000-2002 Survey Data 2001 2001-08-14 Oceanside Epifaunal 06 Def 33.2052 -117.3908
700074 Introduced Species Study 2011 2011-04-08 Wharf 4 Def 34.1500 -119.2100
700100 Introduced Species Study 2011 2011-05-05 Middle Harbor Yacht Slip Def 33.2106 -117.3960
700104 Introduced Species Study 2006 2006-09-13 Middle Harbor Yacht Slip Def 33.2106 -117.3960
700200 Introduced Species Study 2006 2006-08-25 Marina del Rey Harbor Entrance Def 33.9702 -118.4496
700201 Introduced Species Study 2011 2011-04-22 Marina del Rey Harbor Entrance Def 33.9702 -118.4496
700376 Introduced Species Study 2006 2006-09-15 Dana Point Harbor Mouth Slip Def 33.4594 -117.6941
700647 Introduced Species Study 2006 2006-09-12 Santa Barbara Cove Def 32.7774 -117.2484
701376 Cohen et al. 2002 (So Cal Exotics RAS); Lambert and Lambert 2003 2000 2000-08-30 Impound Marina Def 33.7639 -118.2444
701389 Introduced Species Study 2006 2006-09-13 Oceanside Commercial Fishing Dock Def 33.2057 -117.3897
701390 Introduced Species Study 2011 2011-05-05 Oceanside Commercial Fishing Dock Def 33.2057 -117.3897
701847 Introduced Species Study 2006 2006-08-21 APL Terminal Def 33.7348 -118.2479
701872 Introduced Species Study 2011 2011-05-05 Ocean Institute Dock Def 33.4622 -117.7063
701879 Introduced Species Study 2006 2006-09-15 Ocean Institute Dock Def 33.4622 -117.7063
701958 Introduced Species Study 2006 2006-08-25 Overnight Guest Dock Def 33.9761 -118.4461
701994 ISS 2000-2002 Survey Data 2001 2001-10-10 America's Cup Harbor Def 32.7239 -117.2240
701995 Introduced Species Study 2011 2011-05-03 America's Cup Harbor Def 32.7239 -117.2240
702200 Maloney et al. 2007 2005 2005-04-27 Chula Vista Marina Def 32.6225 -117.1023
702201 ISS 2000-2002 Survey Data 2001 2001-10-09 Chula Vista Marina Def 32.6225 -117.1023
702871 Cohen et al. 2002 (So Cal Exotics RAS); Lambert and Lambert 2003 2000 2000-08-26 Fiddler's Cove Def 32.6519 -117.1494
703057 Introduced Species Study 2011 2011-05-03 San Diego Bay Cruise Ship Terminal Def 32.7168 -117.1759
703058 Maloney et al. 2007 2005 2005-04-26 San Diego Bay Cruise Ship Terminal Def 32.7168 -117.1759
703101 Maloney et al. 2007 2005 2005-04-26 Switzer Creek Def 32.7043 -117.1615
703349 ISS 2000-2002 Survey Data 2001 2001-10-09 Coronado Cays Def 32.6274 -117.1329
703501 Introduced Species Study 2006 2006-09-13 North Corner of Harbor Def 33.2122 -117.3954
703515 Cohen et al. 2002 (So Cal Exotics RAS); Lambert and Lambert 2003 2000 2000-08-26 Shelter Island Def 32.7100 -117.2342
703624 Cohen et al. 2002 (So Cal Exotics RAS); Lambert and Lambert 2003 2000 2000-08-31 Long Beach Yacht Club Def 33.7534 -118.1138
703661 Maloney et al. 2007 2005 2005-04-26 SIO Nimitz Marine Facility Def 32.7078 -117.2368
703759 Introduced Species Study 2006 2006-09-14 Balboa North Channel Def 33.6097 -117.8957
703842 Maloney et al. 2007 2005 2005-04-26 Shelter Island Marina Def 32.7180 -117.2255
704340 Introduced Species Study 2011 2011-04-21 Super Mexico Pier Def 33.7708 -118.2113
704644 Maloney et al. 2007 2005 2005-06-28 Navy - Carrier Base Def 32.7062 -117.1886
757293 Lambert and Lambert 2003 1997 King Harbor Def 33.8464 -118.3969
757294 Lambert and Lambert 2003 1998 King Harbor Def 33.8464 -118.3969
757295 Lambert and Lambert 2003 1998 Impound Marina, Long Beach Def 33.7639 -118.2444
757296 Lambert and Lambert 2003 1997 Long Beach Yacht Club Def 33.7534 -118.1138
757297 Lambert and Lambert 1998; Lambert and Lambert 2003 1995 1995-11-25 Oceanside Harbor, by Harbor Patrol floats Def 33.2081 -117.3946
757298 Lambert and Lambert 1998; Lambert and Lambert 2003 1996 1996-10-12 Oceanside Harbor, by Harbor Patrol floats Def 33.2081 -117.3946
757299 Lambert and Lambert 1998; Lambert and Lambert 2003 1997 1997-04-06 Oceanside Harbor, by Harbor Patrol floats Def 33.2081 -117.3946
757300 Lambert and Lambert 2003 1997 Oceanside Harbor, by Harbor Patrol floats Def 33.2081 -117.3946
757301 Lambert and Lambert 1998; Lambert and Lambert 2003 1994 1994-10-09 Bahia Point, Mission Bay Def 32.7761 -117.2468
757302 Lambert and Lambert 1998; Lambert and Lambert 2003 1995 1995-11-25 Bahia Point, Mission Bay Def 32.7761 -117.2468
757303 Lambert and Lambert 1998; Lambert and Lambert 2003 1996 1996-11-03 Bahia Point, Mission Bay Def 32.7761 -117.2468
757304 Lambert and Lambert 2003 1997 Bahia Point, Mission Bay Def 32.7761 -117.2468
757305 Lambert and Lambert 2003 1998 Bahia Point, Mission Bay Def 32.7761 -117.2468
757306 Lambert and Lambert 2003 1997 South Shores Boat Ramp, Mission Bay Def 32.7644 -117.2168
757307 Lambert and Lambert 2003 1998 South Shores Boat Ramp, Mission Bay Def 32.7644 -117.2168
757308 Lambert and Lambert 1998; Lambert and Lambert 2003 1994 1994-10-09 Dana Landing, Mission Bay Def 32.7674 -117.2357
757309 Lambert and Lambert 2003 1997 Dana Landing, Mission Bay Def 32.7674 -117.2357
757310 Lambert and Lambert 1998; Lambert and Lambert 2003 1995 1995-03-26 Seaforth Landing Def 32.7644 -117.2381
757311 Lambert and Lambert 1998; Lambert and Lambert 2003 1996 1996-11-03 Seaforth Landing Def 32.7644 -117.2381
757312 Lambert and Lambert 1998; Lambert and Lambert 2003 1997 1997-04-06 Seaforth Landing Def 32.7644 -117.2381
757313 Lambert and Lambert 2003 1997 Seaforth Landing Def 32.7644 -117.2381
757314 Lambert and Lambert 2003 1998 Seaforth Landing Def 32.7644 -117.2381
757315 Lambert and Lambert 1998; Lambert and Lambert 2003 1994 1994-10-09 Shelter Island, San Diego Bay Def 32.7100 -117.2342
757316 Lambert and Lambert 1998; Lambert and Lambert 2003 1995 1995-03-26 Shelter Island, San Diego Bay Def 32.7100 -117.2342
757317 Lambert and Lambert 1998; Lambert and Lambert 2003 1995 1995-09-30 Shelter Island, San Diego Bay Def 32.7100 -117.2342
757318 Lambert and Lambert 1998; Lambert and Lambert 2003 1996 1996-11-03 Shelter Island, San Diego Bay Def 32.7100 -117.2342
757319 Lambert and Lambert 1998; Lambert and Lambert 2003 1997 1997-04-06 Shelter Island, San Diego Bay Def 32.7100 -117.2342
757320 Lambert and Lambert 2003 1997 Shelter Island, San Diego Bay Def 32.7100 -117.2342
757321 Lambert and Lambert 2003 1998 Shelter Island, San Diego Bay Def 32.7100 -117.2342
757322 Lambert and Lambert 1998; Lambert and Lambert 2003 1994 1994-10-09 Harbor Island, San Diego Bay Def 32.7253 -117.2064
757323 Lambert and Lambert 1998; Lambert and Lambert 2003 1995 1995-03-26 Harbor Island, San Diego Bay Def 32.7253 -117.2064
757324 Lambert and Lambert 1998; Lambert and Lambert 2003 1996 1996-11-03 Harbor Island, San Diego Bay Def 32.7253 -117.2064
757325 Lambert and Lambert 1998; Lambert and Lambert 2003 1997 1997-04-06 Harbor Island, San Diego Bay Def 32.7253 -117.2064
757326 Lambert and Lambert 2003 1997 Harbor Island, San Diego Bay Def 32.7253 -117.2064
757327 Lambert and Lambert 2003 1998 Harbor Island, San Diego Bay Def 32.7253 -117.2064
757328 Lambert and Lambert 1998; Lambert and Lambert 2003 1994 1994-10-09 Fiddler's Cove, San Diego Bay Def 32.6519 -117.2342
757329 Lambert and Lambert 1998; Lambert and Lambert 2003 1995 1995-03-26 Fiddler's Cove, San Diego Bay Def 32.6519 -117.2342
757330 Lambert and Lambert 1998; Lambert and Lambert 2003 1995 1995-09-30 Fiddler's Cove, San Diego Bay Def 32.6519 -117.2342
757331 Lambert and Lambert 1998; Lambert and Lambert 2003 1996 1996-11-03 Fiddler's Cove, San Diego Bay Def 32.6519 -117.2342
757332 Lambert and Lambert 1998; Lambert and Lambert 2003 1997 1997-04-06 Fiddler's Cove, San Diego Bay Def 32.6519 -117.2342
757333 Lambert and Lambert 2003 1997 Fiddler's Cove, San Diego Bay Def 32.6519 -117.2342
757334 Lambert and Lambert 2003 1998 Fiddler's Cove, San Diego Bay Def 32.6519 -117.2342
757335 Lambert and Lambert 1998; Lambert and Lambert 2003 1994 1994-10-09 24th Street Boat Launch, San Diego Bay Def 32.6495 -117.1108
757336 Lambert and Lambert 1998; Lambert and Lambert 2003 1995 1995-09-30 24th Street Boat Launch, San Diego Bay Def 32.6495 -117.1108
757337 Lambert and Lambert 2003 1997 24th Street Boat Launch, San Diego Bay Def 32.6495 -117.1108
757338 Lambert and Lambert 2003 1998 24th Street Boat Launch, San Diego Bay Def 32.6495 -117.1108
757339 Lambert and Lambert 1998; Lambert and Lambert 2003 1994 1994-10-09 Chula Vista Boat Ramp (J Street) Def 32.6211 -117.1031
757340 Lambert and Lambert 1998; Lambert and Lambert 2003 1995 1995-09-30 Chula Vista Boat Ramp (J Street) Def 32.6211 -117.1031
757341 Lambert and Lambert 1998; Lambert and Lambert 2003 1997 1997-04-06 Chula Vista Boat Ramp (J Street) Def 32.6211 -117.1031
757342 Lambert and Lambert 2003 1997 Chula Vista Boat Ramp (J Street) Def 32.6211 -117.1031
757343 Lambert and Lambert 2003 1998 Chula Vista Boat Ramp (J Street) Def 32.6211 -117.1031
757344 de Rivera et al. 2005 2003 Dana Marina, Mission Bay Def 32.7675 -117.2365
757345 de Rivera et al. 2005 2003 Seaforth Marina, Mission Bay Def 32.7630 -117.2373
757346 de Rivera et al. 2005 2003 Chula Vista Marina, San Diego Bay Def 32.6244 -117.1038
767428 Ruiz et al., 2015 2013 2013-07-19 SeaWorld Marina, Mission Bay, CA, California, USA Def 32.7676 -117.2314
767483 Ruiz et al., 2015 2013 2013-08-04 Bahia Resort Marina, Mission Bay, CA, California, USA Def 32.7731 -117.2478
767498 Ruiz et al., 2015 2013 2013-07-31 Campland on the Bay, Mission Bay, CA, California, USA Def 32.7936 -117.2234
767514 Ruiz et al., 2015 2013 2013-08-01 Hyatt Resort Marina, Mission Bay, CA, California, USA Def 32.7634 -117.2397
767529 Ruiz et al., 2015 2013 2013-08-03 Mission Bay Sport Center, Mission Bay, CA, California, USA Def 32.7857 -117.2495
767542 Ruiz et al., 2015 2013 2013-07-30 Hilton Resort Docks, Mission Bay, CA, California, USA Def 32.7791 -117.2128
767569 Ruiz et al., 2015 2013 2013-08-05 Paradise Point Resort, Mission Bay, CA, California, USA Def 32.7730 -117.2406
767684 Ruiz et al., 2015 2013 2013-07-17 Naval Station San Diego, San Diego Bay, CA, California, USA Def 32.6867 -117.1333
767698 Ruiz et al., 2015 2013 2013-07-24 NAB ACU-1 Docks, San Diego Bay, CA, California, USA Def 32.6786 -117.1615
767710 Ruiz et al., 2015 2013 2013-07-25 Navy Ammo Dock, Pier Bravo, San Diego Bay, CA, California, USA Def 32.6939 -117.2276
767721 Ruiz et al., 2015 2013 2013-07-21 Cabrillo Isle Marina, San Diego Bay, CA, California, USA Def 32.7272 -117.1995
767734 Ruiz et al., 2015 2013 2013-07-22 Coronado Cays Marina, San Diego Bay, CA, California, USA Def 32.6257 -117.1309
767751 Ruiz et al., 2015 2013 2013-07-18 NAB Fiddlers Cove, San Diego Bay, CA, California, USA Def 32.6524 -117.1486
767769 Ruiz et al., 2015 2013 2013-07-26 Pier 32 Marina, San Diego Bay, CA, California, USA Def 32.6516 -117.1077
767776 Ruiz et al., 2015 2013 2013-07-20 Chula Vista Marina, San Diego Bay, CA, California, USA Def 32.6252 -117.1036
767791 Ruiz et al., 2015 2013 2013-07-28 Marriott Marquis and Marina, San Diego Bay, CA, California, USA Def 32.7059 -117.1655
769288 Ruiz et al., 2021b 2017 2017-07-28 Port of LB Pier F, San Pedro Bay, California, USA Def 33.7473 -118.2131
769312 Ruiz et al., 2021b 2018 2018-08-06 Shoreline Public Marina, San Pedro Bay, California, USA Def 33.7586 -118.1856
769355 Ruiz et al., 2021b 2018 2018-08-06 Shoreline Public Marina, San Pedro Bay, California, USA Def 33.7586 -118.1856
769953 Ruiz et al., 2021b 2018 2018-08-08 Port of LB Matson, San Pedro Bay, California, USA Def 33.7729 -118.2178
769969 Ruiz et al., 2021b 2018 2018-08-08 Port of LB Matson, San Pedro Bay, California, USA Def 33.7729 -118.2178
769977 Ruiz et al., 2021b 2018 2018-08-08 Port of LB Matson, San Pedro Bay, California, USA Def 33.7729 -118.2178
770534 Ruiz et al., 2021b 2017 2017-08-04 Hill's Boat Service, Newport Bay, California, USA Def 33.6026 -117.8984
770548 Ruiz et al., 2021b 2017 2017-08-04 Hill's Boat Service, Newport Bay, California, USA Def 33.6026 -117.8984
770570 Ruiz et al., 2021b 2017 2017-08-04 Hill's Boat Service, Newport Bay, California, USA Def 33.6026 -117.8984
770580 Ruiz et al., 2021b 2017 2017-08-04 Hill's Boat Service, Newport Bay, California, USA Def 33.6026 -117.8984
770588 Ruiz et al., 2021b 2017 2017-08-04 Marina Park, Newport Bay, California, USA Def 33.6085 -117.9222
770611 Ruiz et al., 2021b 2017 2017-08-04 Marina Park, Newport Bay, California, USA Def 33.6085 -117.9222
770623 Ruiz et al., 2021b 2017 2017-08-04 Marina Park, Newport Bay, California, USA Def 33.6085 -117.9222
770727 Ruiz et al., 2021b 2017 2017-08-03 Rhine Wharf Public Pier, Newport Bay, California, USA Def 33.6148 -117.9266
774454 Ruiz et al., 2022 2015 2015-07-29 Waves-Tahiti A, Marina del Rey, California, USA Def 33.9729 -118.4511
774455 Ruiz et al., 2022 2015 2015-07-29 Waves-Tahiti B, Marina del Rey, California, USA Def 33.9745 -118.4502
774456 Ruiz et al., 2022 2015 2015-07-27 Esprit Marina, Marina del Rey, California, USA Def 33.9757 -118.4497
774457 Ruiz et al., 2022 2015 2015-07-27 Neptune Marina, Marina del Rey, California, USA Def 33.9758 -118.4565
774458 Ruiz et al., 2022 2015 2015-07-28 Catalina Yacht Club, Marina del Rey, California, USA Def 33.9818 -118.4440
774459 Ruiz et al., 2022 2015 2015-07-28 Catalina Yacht Club, Marina del Rey, California, USA Def 33.9818 -118.4440
774460 Ruiz et al., 2022 2015 2015-07-28 Catalina Yacht Club, Marina del Rey, California, USA Def 33.9818 -118.4440
774461 Ruiz et al., 2022 2015 2015-07-28 Catalina Yacht Club, Marina del Rey, California, USA Def 33.9818 -118.4440
774462 Ruiz et al., 2022 2015 2015-07-31 Del Rey Marina-Basin D, Marina del Rey, California, USA Def 33.9807 -118.4505
774463 Ruiz et al., 2022 2015 2015-07-31 Del Rey Marina-Basin D, Marina del Rey, California, USA Def 33.9807 -118.4505
774464 Ruiz et al., 2022 2015 2015-07-31 Del Rey Marina-Basin D, Marina del Rey, California, USA Def 33.9807 -118.4505
774465 Ruiz et al., 2022 2015 2015-07-31 Del Rey Marina-Basin D, Marina del Rey, California, USA Def 33.9807 -118.4505
774466 Ruiz et al., 2022 2015 2015-07-31 Del Rey Marina-Basin D, Marina del Rey, California, USA Def 33.9807 -118.4505
774467 Ruiz et al., 2022 2015 2015-07-31 Del Rey Marina-Basin E, Marina del Rey, California, USA Def 33.9825 -118.4513
774468 Ruiz et al., 2022 2015 2015-07-31 Del Rey Marina-Basin E, Marina del Rey, California, USA Def 33.9825 -118.4513
774469 Ruiz et al., 2022 2015 2015-07-31 Del Rey Marina-Basin E, Marina del Rey, California, USA Def 33.9825 -118.4513
774470 Ruiz et al., 2022 2015 2015-07-31 Del Rey Marina-Basin E, Marina del Rey, California, USA Def 33.9825 -118.4513
774471 Ruiz et al., 2022 2015 2015-07-30 Dolphin Marina, Marina del Rey, California, USA Def 33.9782 -118.4529
774472 Ruiz et al., 2022 2015 2015-07-30 Dolphin Marina, Marina del Rey, California, USA Def 33.9782 -118.4529
774473 Ruiz et al., 2022 2015 2015-08-01 Holiday Marina, Marina del Rey, California, USA Def 33.9797 -118.4549
774474 Ruiz et al., 2022 2015 2015-08-01 Holiday Marina, Marina del Rey, California, USA Def 33.9797 -118.4549
774475 Ruiz et al., 2022 2015 2015-08-01 Holiday Marina, Marina del Rey, California, USA Def 33.9797 -118.4549
774476 Ruiz et al., 2022 2015 2015-08-01 Holiday Marina, Marina del Rey, California, USA Def 33.9797 -118.4549
774477 Ruiz et al., 2022 2015 2015-08-01 Holiday Marina, Marina del Rey, California, USA Def 33.9797 -118.4549
774478 Ruiz et al., 2022 2015 2015-07-28 Santa Monica Yacht Club, Marina del Rey, California, USA Def 33.9781 -118.4455
774479 Ruiz et al., 2022 2015 2015-07-28 Santa Monica Yacht Club, Marina del Rey, California, USA Def 33.9781 -118.4455
774480 Ruiz et al., 2022 2015 2015-07-28 Santa Monica Yacht Club, Marina del Rey, California, USA Def 33.9781 -118.4455
774481 Ruiz et al., 2022 2015 2015-07-28 Santa Monica Yacht Club, Marina del Rey, California, USA Def 33.9781 -118.4455
774482 Ruiz et al., 2022 2015 2015-07-28 Santa Monica Yacht Club, Marina del Rey, California, USA Def 33.9781 -118.4455

References

Brunetti, Riccardo (1978) A colonial ascidian new to the Mediterranean., Vie et Milieu 28-29(4): 647-652

Brunetti, Riccardo; Mastrototaro, Francesco (2004) The non-indigenous stolidobranch ascidian Polyandrocarpa zorritensis in the Mediterranean: description, larval morphology and pattern of vascular budding, Zootaxa 528: 1-8

California Department of Fish and Wildlife (2014) Introduced Aquatic Species in California Bays and Harbors, 2011 Survey, California Department of Fish and Wildlife, Sacramento CA. Pp. 1-36

Carman, Mary, and 8 authors (2011) Ascidians at the Pacific and Atlantic entrances to the Panama Canal, Aquatic Invasions 6(4): 371-380

Coles S. L., DeFelice R. C., Eldredge, L. G. (1999a) Nonindigenous marine species introductions in the harbors of the south and west shores of Oahu, Hawaii., Bishop Museum Technical Report 15: 1-212

Coles, S. L.; DeFelice, R. C. : Eldredge, L. G. (2002a) Nonindigenous marine species in Kaneohe Bay, Oahu, Hawai`i, Bishop Museum Technical Report 24: 1-364

Coles, S. L.; DeFelice, R. C.; Eldredge, L. G.; Carlton, J. T. (1999b) Historical and recent introductions of non-indigenous marine species into Pearl Harbor, Oahu, Hawaiian Islands., Marine Biology 135(1): 147-158

Dalby, James E. Jr.; Young, Craig M. (1992) Role of early post-settlement mortality in setting the upper depth limit of ascidians in Florida epifaunal communities, Marine Ecology Progress Series 80: 221-228,

de Rivera, Catherine, and 27 authors (2005) Broad-scale non-indigenous species monitoring along the West Coast in National Marine Sanctuaries and National Estuarine Research Reserves report to National Fish and Wildlife Foundation, National Fish and Wildlife Foundation, Washington, D.C.. Pp. <missing location>

Dias, G. M.; Rocha, R. M.; Lotufo, T. M. C.; Kremer, L. P. (2013) Fifty years of ascidian biodiversity research in Sao Sebastiao, Brazil, Journal of the Marine Biological Association of the United Kingdom 93(1): 273-282

Fairey, Russell; Dunn, Roslyn; Sigala, Marco; Oliver, John (2002) Introduced aquatic species in California's coastal waters: Final Report, California Department of Fish and Game, Sacramento. Pp. <missing location>

Iwasaki, Keiji (2006) Assessment and Control of Biological Invasion Risks., Shoukadoh Book Sellers,and IUCN, Kyoto and Gland, Switzerland. Pp. 104-112

Izquierdo-Muñoz, Andrés; Díaz-Valdés, Marta; Ramos-Esplá, Alfonso A. (2009) Recent non-indigenous ascidians in the Mediterranean Sea., Memoirs of the Museum of Victoria 4(1): 59-64

Lambert, C. C.; Lambert, G. (1998) Non-indigenous ascidians in southern California harbors and marinas., Marine Biology 130: 675-688

Lambert, Charles C; Lambert, Gretchen (2003) Persistence and differential distribution of nonindigenous ascidians in harbors of the Southern California Bight., Marine Ecology Progress Series 259: 145-161

Lambert, Gretchen (2019) Fouling ascidians (Chordata: Ascidiacea) of the Galápagos: Santa Cruz and Baltra Islands, Aquatic Invasions 14: 132-149

Lambert, Gretchen; Faulkes, Zen; Lambert, Charles C.; Scofield, Virginia L. (2005) Ascidians of South Padre Island, Texas, with a key to species., Texas Journal of Science 57(3): 251-262

Locke, Andrea (2009) A screening procedure for potential tunicate invaders of Atlantic Canada., Aquatic Invasions 4(1): 71-79

Lopez-Legentil, Susanna; Legentil, Miquel L.; Erwin, Patrick M.; Turon, Xavier (2015) Harbor networks as introduction gateways: contrasting distribution patterns of native and introduced ascidians, Biological Invasions 17: 1623-1638
DOI 10.1007/s10530-014-0821-z

Lord, Joshua P.; Calini, Jeremy M.; Whitlatch, Robert B. (2015) Influence of seawater temperature and shipping on the spread and establishment of marine fouling species, Marine Biology 162: 2481-2492

Low-Pfeng, Antonio; Recagno, Edward M. Peters (2012) <missing title>, Geomare, A. C., INESEMARNAT, Mexico. Pp. 236

Lutaenko, Konstantin A.; Furota,Toshio; Nakayama, Satoko; Shin, Kyoungsoon; Xu, Jing (2013) <missing title>, Northwest Pacific Action Plan- Data and Information Network Regional Activity Center, Beijing, China. Pp. <missing location>

Mastrototaro, F.; D’Onghia, G.; Tursi, A. (2008) Spatial and seasonal distribution of ascidians in a semi-enclosed basin of the Mediterranean Sea., Journal of the Marine Biological Association of the United Kingdom 88(5): 1053-1061

Millar, R. H. (1958) Some ascidians from Brazil., Annals and Magazine of Natural History 13(1): 497-514

Monniot, Françoise (2018) Ascidians collected during the Madibenthos expedition in Martinique: 2. Stolidobranchia, Styelidae, Zootaxa 2: 291-318

Nichols, Claire L.; Lambert, Gretchen; Nydam, Marie L. (2023) Continued persistence of non-native ascidians in Southern California harbors and marinas, Aquatic Invasions 18(1): 1-22
https://doi.org/10.3391/ ai.2023.18.1.101962

Otani, Michio (2006) Assessment and Control of Biological Invasion Risks., Shoukadoh Book Sellers and IUCN, Kyoto, Japan, and Gland, Switzerland. Pp. <missing location>

Pestana, Lueji Barros; Dias, Gustavo Muniz; Marquesa, Antonio Carlos (2017) A century of introductions by coastal sessile marine invertebrates in Angola, South East Atlantic Ocean, Marine Pollution Bulletin 125: 426-a432

Quintanilla, Elena; Thomas Wilke; Ramırez-Portilla, Catalina; Sarmiento, Adriana; Sanchez, Juan A. () , None <missing volume>: <missing location>

Ramalhosa Patrício; Gestoso, Ignacio Rocha Rosana M.; Lambert, Gretchen; Canning-Clode, João (2021) Ascidian biodiversity in the shallow waters of the Madeira Archipelago: Fouling studies on artificial substrates and new records, Regional Studies in Marine Science 43(1901672): Published online

Rodriguez, Laura F.; Ibarra-Obando, Silvia E. (2008) Cover and colonization of commercial oyster (Crassostrea gigas) shells by fouling organisms in San Quintin Bay, Mexico, Journal of Shellfish Research 27(2): 337-343

Ruiz, Gregory M.; Geller, Jonathan (2018) Spatial and temporal analysis of marine invasions in California, Part II: Humboldt Bay, Marina del Re, Port Hueneme, and San Francisco Bay, Smithsonian Environmental Research Center & Moss Landing Laboratories, Edgewater MD, Moss Landing CA. Pp. <missing location>

Ruiz, Gregory; Geller, Jonathan (2021) Spatial and temporal analysis of marine invasions: supplemental studies to evaluate detection through quantitative and molecular methodologies, Marine Invasive Species Program, California Department of Fish and Wildlife, Sacramento CA. Pp. 153 ppl.

Stabili, Loredana; Licciano, Margherita; Longo, Caterina; Lezzi, Marco; Giangrande, Adriana (2015) The Mediterranean non-indigenous ascidian Polyandrocarpa zorritensis: Microbiological accumulation capability and environmental implications, Marine Pollution Bulletin 101: 146-152

Streit, Olivia T; Lambert, Gretchen; Erwin, Patrick M.; Lopez-Legentil, Susanna (2021) Diversity and abundance of native and non-native ascidians in Puerto Rican harbors and marinas, Marine Pollution Bulletin 167(112262): Published online

Tracy, Brianna M.; Reyns, Nathalie B. (2014) Spatial and temporal patterns of native and invasive ascidian assemblages in a Southern California embayment, Aquatic Invasions 9: In press

Van Name, Willard G. (1945) The North and South American ascidians, Bulletin of the American Museum of Natural History 84: 1-462

Vazquez, Elsa; Young, Craig M. (1996) Responses of compound ascidian larvae to haloclines., Marine Ecology Progress Series 133: 179-190

Vazquez, Elsa; Young, Craig M. (1998) Ontogenetic changes in phototaxis during larval life of the Ascidian Polyandrocarpa zorritensis, Journal of Marine Biology and Ecology 231: 267-277