Invasion History

First Non-native North American Tidal Record: 2000
First Non-native West Coast Tidal Record: 2000
First Non-native East/Gulf Coast Tidal Record:

General Invasion History:

Bugulina cf. foliolata is a bryozoan, described from the vicinity of Rio de Janeiro, Brazil (Vieira et al. 2012), and had been identified earlier as B. flabellata, native to European waters. It is also morphologically close to B. fulva and B. aquilirostris, both North Atlantic species (Ryland 1960). A bryozoan identified as 'B. flabellata' was found in Southern California (Cohen et al. 2002; Ruiz et al., unpublished data). In later work, it was identified as B. fulva (de Riviera et al. 2005; Linda McCann, personal communication). More recently, Linda McCann has identified this bryozoan as Bugulina cf. foliolata, as tentatively matching Vieira et al.'s (2012) description.

North American Invasion History:

Invasion History on the West Coast:

Bugulina cf. foliolata was first reported in 2000 as B. flabellata in Long Beach Harbor by Cohen et al. (2002), and in San Diego Bay in a separate SERC collecting program (Ruiz et al. unpublished data). In 2003 it was collected in a marina in Monterey Bay, and in 2004 it was collected in Channel Islands Harbor, Oxnard, (de Rivera et al. 2005). Other collections were in 2010 in San Francisco Bay Marina (Ruiz et al. unpublished) and in 2017 in Long Beach Harbor (Ruiz and Geller 2021). This bryozoan is probably widely distributed, but overlooked on the California coast. In Brazil, B. foliolata was reported 'on stones, shells, and other bryozoans at 7–20 meters depth' (Vieira et al. 2012).

Cohen et al. (2002) reported two unidentified Bugula, sp.1, resembling Bugulina turbinaia. and Bugula, sp. 2, resembling Bugulina fulva from Puget Sound. The identity of these forms is uncertain.


Colonies of Bugulina cf. foliolata form flat fans of strap-like branches 2–8 zooids wide, arranged in fivefold clusters. The zooids are elongate (about 4x long as wide). The frontal membrane occupies almost the entire frontal area. The distal corners of the zooids are sharply pointed. The zooids at the outer edges of the branches have thicker lateral and distal walls. The outer and terminal zooids have three outer and two inner distal spines. The inner zooids have two outer and two inner distal spines. The lower spines curve towards the middle of the zooid. The avicularia are dimorphic and pedunculate, with a round body and a hooked beak. They are positioned about 1/4 of the way down the walls of the zooids. The avicularia on the outer zooids are large than those on the inner zooids. The ovicells are globular, and distinct, and are centered in the midline of the zooids. The colonies are reddish-brown, and in Vieira et al.'s photographs have branches about 15–20 mm long (Vieira et al. 2012).  

Bugulina cf. foliolata has been confused with the North Atlantic species B. fulva and B. flabellata, but Bugulina foliolata has longer and narrower zooids, and differences in position and shape of the avicularia (Vieira et al. 2012). The identification is somewhat tentative, and has not been confirmed by molecular methods (Linda McCann, personal communication).


Taxonomic Tree

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


'Bugula flabellata' (Marcus, 1938)

Potentially Misidentified Species

Bugulina flabellata

Gray 1848, described from England, Amphi-Atlantic distribution

Bugulina fulva

Described from Britain, Ryland 1960, Amphi-Atlantic



Bugulina cf. foliolata is a bushy, fouling bryozoan, with long, strap-like branches, composed of numerous 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. Bugulina cf. foliolata was reported from 'stones, shells, and other bryozoans' (Vieira et al. 2012). When first identified as Bugula flabellata and B. fulva, it was reported from fouling plates and marine pilings (Cohen et al. 2002) 


Phytoplankton, Detritus



Trophic Status:

Suspension Feeder



General HabitatRockyNone
General HabitatMarinas & DocksNone
Salinity RangePolyhaline18-30 PSU
Salinity RangeEuhaline30-40 PSU
Tidal RangeSubtidalNone
Vertical HabitatEpibenthicNone

Life History

In Bugulina cf. foliolata, as in other similar bryozoans, the zooids are hermaphroditic and produce large yolky eggs that hatch into lecithotrophic larvae, which are planktonic for short periods (less than 1 day). Larvae settle on a substrate and metamorphose into the first zooid of a colony, an ancestrula (Barnes 1983).

Tolerances and Life History Parameters

Minimum Depth (m)7Sao Paulo and Rio de Janeiro States, Brazil. Atlantic Ocean (Vieira et al. 2012)
Maximum Depth (m)20Sao Paulo and Rio de Janeiro States, Brazil. Atlantic Ocean (Vieira et al. 2012)
Maximum Length (mm)20Length of branches, estimated from photographs (Vieira et al. 2012)
Broad Temperature RangeNoneSubtropical-Warm Temperate
Broad Salinity RangeNonePolyhaline-Euhaline

General Impacts

No ecological or environmental impacts have been reported for Bugulina cf. foliolata.

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
SA-II None 0 Native Estab
P050 San Pedro Bay 2000 Def Estab
P020 San Diego Bay 2000 Def Estab
P062 _CDA_P062 (Calleguas) 2004 Def Estab
NEP-VI Pt. Conception to Southern Baja California 2000 Def Estab
NEP-V Northern California to Mid Channel Islands 2003 Def Estab
P080 Monterey Bay 2003 Def Estab
P090 San Francisco Bay 2010 Def Estab

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude


Barnes, Robert D. (1983) Invertebrate Zoology, Saunders, Philadelphia. Pp. 883

Cohen, Andrew N. and 12 authors (2002) Project report for the Southern California exotics expedition 2000: a rapid assessment survey of exotic species in sheltered coastal waters., In: (Eds.) . , Sacramento CA. Pp. 1-23

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>

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.

Ryland, J. S. (1960) The British species of Bugula (Polyzoa)., Proceedings of the Zoological Society of London 134: 65-105

Vieira, Leandro M.; Winston, Judith E.; ,Fehlauer-Ale, Karin H. (2012) Nine new species of Bugula Oken (Bryozoa: Cheilostomata) in Brazilian shallow waters, PLOS ONE 7(7): e40492