Invasion History

First Non-native North American Tidal Record: 1950
First Non-native West Coast Tidal Record: 1950
First Non-native East/Gulf Coast Tidal Record:

General Invasion History:

Bugulina stolonifera was described from Milford Haven, Wales in 1960, but it was confined to docks and a boat warmed by thermal effluents (Ryland 1960). Ryland considered the species to be a possible introduction, although this was complicated by a pre-1875 sample labelled as being from the 'British coast'. Ryland noted samples of this species from Marseilles and Naples (Ryland 1960). Bugulina stolonifera appears to be a recent introduction in many parts of the world, mostly in harbors.  It is native to the Northwest Atlantic Ocean (McCann et al. 2019).  It was reported from North Carolina, under the name Bugula californica, a Pacific species which it closely resembles (Maturo 1958; Ryland 1991). In the northwest Atlantic, it ranges from New Hampshire to the Gulf of Mexico, Bermuda, and Jamaica (Winston 1982; Ryland 1991; Creary 2001; Ruiz et al., unpublished data).

North American Invasion History:

Invasion History on the West Coast:

Soule (1980, cited by Cohen and Carlton 1995) found that bryozoans from California ports, previously identified as Bugulina californica, were actually B. stolonifera. Bugulina californica is a distinct, but very similar species confined to the North Pacific, and apparently to open coasts. Identifications of B. californica from California harbors are presumed to refer to B. stolonifera (Cohen and Carlton 1995). Bugulina stolonifera (as B. californica) was collected by Osburn in Newport Bay in the 1940s (Osburn 1950), and was reported by Reish (1972, cited by Cohen and Carlton 1995) in Newport Bay, Huntington Harbor, Alamitos Bay and Los-Angeles-Long Beach harbors. This bryozoan was present in San Francisco Bay by 1978 (Soule 1980, cited by Cohen and Carlton 1995). Currently, B. stolonifera has been found throughout California from San Diego Bay (Ruiz et al. unpublished data) to Humboldt Bay (Ruiz et al. unpublished data), including Morro Bay (Needles 2007), Monterey Bay-Elkhorn Slough (Wasson et al. 2001; de Rivera et al. 2005), Tomales Bay, and Bodega Harbor (de Rivera et al. 2005). Bugulina stolonifera was collected in Puget Sound (Cohen et al. 1998), but has not been collected in Coos Bay, Oregon, or Willapa Bay, Washington (Wonham and Carlton 2005).

Invasion History in Hawaii:

Bugulina stolonifera was first collected in Kaneohe Bay, Oahu in 1935 and in 1940 in Pearl Harbor (Carlton and Eldredge 2009, as B. californica). It was also collected on a barge towed from San Diego to Oahu (Godwin 2003).

Invasion History Elsewhere in the World:

Bugulina stolonifera is now distributed worldwide. It was found at the Pacific side of the Panama Canal (Powell 1971), and in New Zealand in 1961 (Gordon and Mawatari 1992). In Australia, museum specimens collected in the 1880s, from Port Phillip Bay, Victoria, have been identified as B stolonifera. This species is now found on most of the coastline of the southern half of Australia (Keough and Ross 1999). In the Northwest Pacific, this species was first collected (as B. californica) in Tokyo Bay in 1960, and in 1997 as B. stolonifera, in Ise Bay at Nagoya, Japan (Scholz et al. 2003). It ranges from Vladivostok (Zvyagintsev et al. 2003), to Hong Kong and Hainan, China (Morton 1987; Seo 1993; Huang 2001). In the Indian Ocean, B stolonifera has been reported from the Bay of Bengal (Visakhaptnam, India; Rao and Ganapati 1976) and from the Arabian Sea (Pakistan; Javed and Tirmizi 1993). In 2016, B. stolonifera was found in the Galapagos on Santa Cruz and Baltra Islands (Carlton et al. 2019; McCann et al. 2019). 
 
Bugulina stolonifera is introduced in the South and East Atlantic. It was collected from Santos, Brazil (Marcos 1937), and in Argentina, from Mar del Plata and Cabo Blanco (Gappa 2000; Orensanz et al. 2002). On the west coast of Africa, it was found in Ghana (Cook 1968). The date of this bryozoan's arrival in European waters is not known (Ryland 1960). In Europe it was described from Milford Haven, Swansea, Wales, and said to occur in 'southern ports' in the British Isles (Ryland 1960). It is also known from the Netherlands (Wolff 2005), Belgium (Kerckhoff et al. 2007), Brittany (France), and Portugal (Marchini et al. 2007). In the Mediterranean, it was reported from the "Western Mediterranean" (Gautier 1958, cited by Winston 1977), and more specifically from the Etang de Berre, France (Mars 1966, cited by Winston 1977), from the Naples region and the Lagoon of Venice, Italy (Sacchi 1961a, b; Carrada and Sacchi 1966, cited by Winston 1977), and Turkish coast of the Aegean Sea (Kocak and Kucuksezgin 2000). Bugulina stolonifera also occurs in the Azores (Cardigos et al. 2006).

Description

Colonies of Bugulina stolonifera form compact tufts 30–40 mm in height, and gray-buff in color. Typically, there are 3–5 stolons arising from the site of the ancestrula, from which new colonies arise. Tips of the branches show slight spiral growth. The bases of branches consist of two parallel series of zooids (bifurcation type 4; of Ryland 1960, Hayward and Ryland 1998). The zoecia (zooids) are long and slender, with the frontal membrane occupying 1/2 to 3/4 of the length. The zooids are 500–740 µm in length and 165–190 µm in width. The outer distal margin forms a large spine, with a small slender spine below. Another spine is located on the inner distal angle. The aviculariae are attached to the outer distal margin, a little below the spines. The length of the avicularia is about equal to the width of the zooids, and the beaks are curved downward. The polyps have 13–14 tentacles. The ooecia are globular, and the embryos are yellow (Hayward and Ryland 1998). The ancestrula is symmetrical, with a short frontal membrane, 2–3 spines on each distal angle, and a median proximal spine (Description from: Ryland 1960; Gordon and Mawatari 1992; Hayward and Ryland 1998; De Blauwe 2009; Winston and Hayward 2012). 

The taxonomy of the Bugulidae has recently been revised. Twenty-four species, including B. flabellata, B. fulva, and B. simplex are now in the new genus Bugulina (Fehlauer-Ales et al. 2015). 

Taxonomy

Taxonomic Tree

Kingdom:   Animalia
Phylum:   Bryozoa
Class:   Gymnolaemata
Order:   Cheilostomata
Suborder:   Anasca
Family:   Bugulidae
Genus:   Bugulina
Species:   stolonifera

Synonyms

Bugula avicularia (McDougall, 1943)
Bugula californica (Maturo, 1958)
Bugula stolonifera (Ryland, 1960)

Potentially Misidentified Species

Bugulina californica
Bugulina californica is a distinct but very similar species confined to the North Pacific, and apparently to the open coast. Identifications of 'B. californica' from the Northwest Atlantic, and from Pacific harbors are presumed to refer to B. stolonifera (Cohen and Carlton 1995).

Ecology

General:

Life History- Bugulina stolonifera is a bush-like, lightly calcified bryozoan, composed of many individual zooids. The zooids feed by extending the ciliated tentacles of the lophophore as a funnel, creating a current, and driving food particles into their mouths. The food is guided along the tentacles and through the pharynx by the cilia. Larger food particles can be moved or captured by flicking or contracting the tentacles. The zooids are hermaphroditic, and produce large yolky eggs, which hatch into lecithotrophic larvae, which are planktonic for short periods (less than 1–2 days). Larvae settle on a substrate and metamorphose into the first zooid of a colony, an ancestrula (Barnes 1983).

Ecology- Bugulina stolonifera attaches to wood, pilings, dock floats, ships' hulls and other hard substrates. Most of the available records are from artificial substrates (Maturo 1958; Ryland 1960; Gordon and Mawatari 1992; Hayward and Ryland 1998; Ryland et al. 2011).

Food:

Phytoplankton

Trophic Status:

Suspension Feeder

SusFed

Habitats

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

Life History

Tolerances and Life History Parameters

Maximum Temperature (ºC)30.2Field, US East & West Coast marinas (Lord et al. 2015)
Minimum Salinity (‰)26Field data, coastal lagoons, Italy (Occhipinti Ambrogi 1983)
Maximum Salinity (‰)40Field salinity (Shark Bay, Western Australia) (Wyatt et al. 2005)
Minimum Duration0Larval period
Maximum Duration1Wendt 2000
Maximum Height (mm)40New Zealand, Gorodon and Mawatari 1992
Broad Temperature RangeNoneCold temperate-Tropical
Broad Salinity RangeNonePolyhaline-Euhaline

General Impacts

Bugulina stolonifera is a frequent component of fouling communities on port structures and ship hulls (Ryland 1960). In marinas in the Aegean Sea, Turkey, it was the only species growing in the most stressful conditions (low oxygen, high organic input (Kocak 2007). 

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
NA-ET2 Bay of Fundy to Cape Cod 0 Crypogenic Established
CAR-VII Cape Hatteras to Mid-East Florida 0 Crypogenic Established
CAR-I Northern Yucatan, Gulf of Mexico, Florida Straits, to Middle Eastern Florida 0 Crypogenic Established
SEP-H None 1930 Non-native Established
MED-II None 1958 Non-native Established
MED-III None 1966 Non-native Established
NWP-3a None 1950 Non-native Established
NWP-3b None 1960 Non-native Established
NEP-V Northern California to Mid Channel Islands 1978 Non-native Established
NEP-VI Pt. Conception to Southern Baja California 1950 Non-native Established
NEP-III Alaskan panhandle to N. of Puget Sound 1998 Non-native Established
MED-VII None 1937 Non-native Established
NEA-V None 1979 Non-native Established
NEA-II None 1960 Non-native Established
IP-1 None 1993 Non-native Established
CIO-II None 1967 Non-native Established
SP-XXI None 1940 Non-native Established
NA-ET3 Cape Cod to Cape Hatteras 0 Crypogenic Established
AUS-VIII None 1885 Non-native Established
NZ-IV None 1961 Non-native Established
AUS-X None 2007 Non-native Established
AUS-VII None 1977 Non-native Established
AUS-IV None 2008 Non-native Established
NZ-VI None 1992 Non-native Established
NWP-2 None 1979 Non-native Established
SA-I None 1980 Non-native Established
SA-II None 1937 Non-native Established
MED-VI None 2000 Non-native Established
WA-II None 1968 Non-native Established
NA-ET4 Bermuda 0 Crypogenic Established
AUS-V None 2002 Non-native Established
NWP-4a None 1973 Non-native Established
AUS-III None 2002 Non-native Established
P050 San Pedro Bay 1972 Non-native Established
P040 Newport Bay 1950 Non-native Established
P062 _CDA_P062 (Calleguas) 2003 Non-native Established
P064 _CDA_P064 (Ventura) 2004 Non-native Established
P080 Monterey Bay 1998 Non-native Established
P090 San Francisco Bay 1978 Non-native Established
P112 _CDA_P112 (Bodega Bay) 2004 Non-native Established
P290 Puget Sound 1998 Non-native Established
P110 Tomales Bay 2004 Non-native Established
N130 Great Bay 0 Crypogenic Established
M020 Narragansett Bay 0 Crypogenic Established
P020 San Diego Bay 2000 Non-native Established
NEP-IV Puget Sound to Northern California 2003 Non-native Established
P130 Humboldt Bay 2001 Non-native Established
CAR-II None 0 Crypogenic Established
NEA-VI None 1998 Non-native Established
P070 Morro Bay 2005 Non-native Established
CAR-IV None 0 Crypogenic Established
AUS-II None 1999 Non-native Established
CIO-I None 2000 Non-native Established
RS-3 None 2003 Non-native Established
NEA-III None 0 Non-native Established
NA-S2 None 0 Crypogenic Established
WA-I None 2010 Non-native Established
CAR-III None 0 Crypogenic Established
SA-III None 2015 Non-native Established
PAN_PAC Panama Pacific Coast 1930 Non-native Established
PAN_CAR Panama Caribbean Coast 0 Crypogenic Established
SEP-Z None 2016 Non-native Established
NEA-IV None 2009 Non-native Unknown
P060 Santa Monica Bay 2015 Non-native Established
WA-IV None 2016 Non-native Established

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude
767360 Ruiz et al., 2015 2012 2012-08-22 Tomales-Marshall, Bodega Bay, California, USA Non-native 38.1514 -122.8888
767373 Ruiz et al., 2015 2012 2012-08-21 Tomales-Nick's Cove, Bodega Bay, California, USA Non-native 38.1980 -122.9222
767404 Ruiz et al., 2015 2012 2012-08-17 Tomales- Shell Beach, Bodega Bay, California, USA Non-native 38.1163 -122.8713
767420 Ruiz et al., 2015 2013 2013-07-19 SeaWorld Marina, Mission Bay, CA, California, USA Non-native 32.7676 -117.2314
767453 Ruiz et al., 2015 2013 2013-07-29 Mission Bay Yacht Club, Mission Bay, CA, California, USA Non-native 32.7778 -117.2485
767473 Ruiz et al., 2015 2013 2013-08-04 Bahia Resort Marina, Mission Bay, CA, California, USA Non-native 32.7731 -117.2478
767503 Ruiz et al., 2015 2013 2013-08-01 Hyatt Resort Marina, Mission Bay, CA, California, USA Non-native 32.7634 -117.2397
767520 Ruiz et al., 2015 2013 2013-08-03 Mission Bay Sport Center, Mission Bay, CA, California, USA Non-native 32.7857 -117.2495
767562 Ruiz et al., 2015 2013 2013-08-05 Paradise Point Resort, Mission Bay, CA, California, USA Non-native 32.7730 -117.2406
767599 Ruiz et al., 2015 2013 2013-09-05 Launch Ramp, Morro Bay, CA, California, USA Non-native 35.3577 -120.8508
767610 Ruiz et al., 2015 2013 2013-08-29 Moorings, Morro Bay, CA, California, USA Non-native 35.3619 -120.8548
767621 Ruiz et al., 2015 2013 2013-08-31 Morro Bay Marina, Morro Bay, CA, California, USA Non-native 35.3641 -120.8532
767639 Ruiz et al., 2015 2013 2013-09-03 State Park Marina, Morro Bay, CA, California, USA Non-native 35.3459 -120.8423
767651 Ruiz et al., 2015 2013 2013-09-04 Tidelands, Morro Bay, CA, California, USA Non-native 35.3602 -120.8521
767662 Ruiz et al., 2015 2013 2013-07-16 Naval Base Point Loma, San Diego Bay, CA, California, USA Non-native 32.6886 -117.2343
767703 Ruiz et al., 2015 2013 2013-07-25 Navy Ammo Dock, Pier Bravo, San Diego Bay, CA, California, USA Non-native 32.6939 -117.2276
767756 Ruiz et al., 2015 2013 2013-07-26 Pier 32 Marina, San Diego Bay, CA, California, USA Non-native 32.6516 -117.1077
767782 Ruiz et al., 2015 2013 2013-07-28 Marriott Marquis and Marina, San Diego Bay, CA, California, USA Non-native 32.7059 -117.1655
767796 Ruiz et al., 2015 2011 2011-09-15 Richmond Marina Bay Yacht Harbor, San Francisco Bay, CA, California, USA Non-native 37.9117 -122.3494
767809 Ruiz et al., 2015 2011 2012-09-20 San Leandro Marina, San Francisco Bay, CA, California, USA Non-native 37.6979 -122.1912
767816 Ruiz et al., 2015 2011 2011-09-20 San Francisco Marina, San Francisco Bay, CA, California, USA Non-native 37.8067 -122.4432
767829 Ruiz et al., 2015 2011 2011-09-14 Coyote Point Marina, San Francisco Bay, CA, California, USA Non-native 37.5880 -122.3160
767839 Ruiz et al., 2015 2011 2011-09-16 Loch Lomond Marina, San Francisco Bay, CA, California, USA Non-native 37.9724 -122.4796
767849 Ruiz et al., 2015 2011 2011-09-13 Oyster Point Marina, San Francisco Bay, CA, California, USA Non-native 37.6725 -122.3864
767865 Ruiz et al., 2015 2011 2011-09-13 Redwood City Marina, San Francisco Bay, CA, California, USA Non-native 37.8046 -122.3985
767875 Ruiz et al., 2015 2011 2012-09-15 Berkeley Marina, San Francisco Bay, CA, California, USA Non-native 37.8758 -122.3181
767884 Ruiz et al., 2015 2011 2012-09-19 Sausalito Marine Harbor, San Francisco Bay, CA, California, USA Non-native 37.8609 -122.4853
767899 Ruiz et al., 2015 2011 2011-09-21 South Beach Harbor, San Francisco Bay, CA, California, USA Non-native 37.7797 -122.3871
767912 Ruiz et al., 2015 2011 2011-09-20 Jack London Square Marina, San Francisco Bay, CA, California, USA Non-native 37.7947 -122.2822
767926 Ruiz et al., 2015 2011 2011-09-22 Ballena Isle Marina, San Francisco Bay, CA, California, USA Non-native 37.7676 -122.2869
767947 Ruiz et al., 2015 2011 2011-09-12 Paradise Cay Yacht Harbor, San Francisco Bay, CA, California, USA Non-native 37.9156 -122.4769
767960 Ruiz et al., 2015 2011 2011-09-12 Corinthian Yacht Club, San Francisco Bay, CA, California, USA Non-native 37.8103 -122.3228
767975 Ruiz et al., 2015 2012 2012-08-24 Richmond Marina Bay Yacht Harbor, San Francisco Bay, CA, California, USA Non-native 37.9134 -122.3523
767997 Ruiz et al., 2015 2012 2012-08-23 Sausalito Marine Harbor, San Francisco Bay, CA, California, USA Non-native 37.8609 -122.4853
768051 Ruiz et al., 2015 2012 2012-09-11 Ballena Isle Marina, San Francisco Bay, CA, California, USA Non-native 37.7676 -122.2869
768073 Ruiz et al., 2015 2012 2012-08-30 Oyster Point Marina, San Francisco Bay, CA, California, USA Non-native 37.6633 -122.3817
768098 Ruiz et al., 2015 2012 2012-08-29 Coyote Point Marina, San Francisco Bay, CA, California, USA Non-native 37.5877 -122.3174
768121 Ruiz et al., 2015 2012 2012-09-04 Redwood City Marina, San Francisco Bay, CA, California, USA Non-native 37.5023 -122.2130
768145 Ruiz et al., 2015 2012 2012-09-06 Loch Lomond Marina, San Francisco Bay, CA, California, USA Non-native 37.9736 -122.4802
768163 Ruiz et al., 2015 2012 2012-09-05 Port of Oakland, San Francisco Bay, CA, California, USA Non-native 37.7987 -122.3228
768186 Ruiz et al., 2015 2012 2012-09-07 Jack London Square Marina, San Francisco Bay, CA, California, USA Non-native 37.7940 -122.2787
768224 Ruiz et al., 2015 2012 2012-09-13 San Leandro Marina, San Francisco Bay, CA, California, USA Non-native 37.6962 -122.1919
768242 Ruiz et al., 2015 2012 2012-09-12 Emeryville, San Francisco Bay, CA, California, USA Non-native 37.8396 -122.3133
768264 Ruiz et al., 2015 2013 2013-08-15 Ballena Isle Marina, San Francisco Bay, CA, California, USA Non-native 37.7656 -122.2858
768288 Ruiz et al., 2015 2013 2013-08-20 Coyote Point Marina, San Francisco Bay, CA, California, USA Non-native 37.5877 -122.3163
768308 Ruiz et al., 2015 2013 2013-08-22 Jack London Square Marina, San Francisco Bay, CA, California, USA Non-native 37.7926 -122.2746
768326 Ruiz et al., 2015 2013 2013-08-23 Loch Lomond Marina, San Francisco Bay, CA, California, USA Non-native 37.9723 -122.4829
768347 Ruiz et al., 2015 2013 2013-08-13 Oyster Point Marina, San Francisco Bay, CA, California, USA Non-native 37.6639 -122.3821
768368 Ruiz et al., 2015 2013 2013-08-14 Redwood City Marina, San Francisco Bay, CA, California, USA Non-native 37.5024 -122.2134
768391 Ruiz et al., 2015 2013 2013-08-19 Richmond Marina Bay Yacht Harbor, San Francisco Bay, CA, California, USA Non-native 37.9138 -122.3522
768428 Ruiz et al., 2015 2013 2013-08-21 San Leandro Marina, San Francisco Bay, CA, California, USA Non-native 37.6980 -122.1908
768442 Ruiz et al., 2015 2013 2013-08-16 Sausalito Marine Harbor, San Francisco Bay, CA, California, USA Non-native 37.8611 -122.4851

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