Bugulina stolonifera

Invasion History

First Galapagos Record: 2016

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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from New Hampshire to the Gulf of Mexico, Bermuda, and Jamaica (Winston 1982; Ryland 1991; Creary 2001; Ruiz et al., unpublished data).

Invasion History in the Galapagos:

Bugulina stolonifera was found in the Galapagos Islands, in 2016 in Franklin’s Bay, Santa Cruz Island, and on Baltra Island (McCann et al 2019). 

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). 

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Taxonomy

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

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Habitats

General Habitat	Marinas & Docks	None
General Habitat	Coarse Woody Debris	None
General Habitat	Oyster Reef	None
General Habitat	Rocky	None
General Habitat	Vessel Hull	None
Salinity Range	Polyhaline	18-30 PSU
Salinity Range	Euhaline	30-40 PSU
Tidal Range	Subtidal	None
Tidal Range	Low Intertidal	None
Vertical Habitat	Epibenthic	None

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Life History

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Tolerances and Life History Parameters

Maximum Temperature (ºC)	30.2	Field, US East & West Coast marinas (Lord et al. 2015)
Minimum Salinity (‰)	26	Field data, coastal lagoons, Italy (Occhipinti Ambrogi 1983)
Maximum Salinity (‰)	40	Field salinity (Shark Bay, Western Australia) (Wyatt et al. 2005)
Minimum Duration	0	Larval period
Maximum Duration	1	Wendt 2000
Maximum Height (mm)	40	New Zealand, Gorodon and Mawatari 1992
Broad Temperature Range	None	Cold temperate-Tropical
Broad Salinity Range	None	Polyhaline-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). 

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