Botrylloides niger

Invasion History

First Galapagos Record: 2015

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General Invasion History:

Botrylloides niger was described from Bermuda by Herdman in 1886 (Van Name 1945) and has a wide distribution in the tropical and subtropical Atlantic, Pacific, and Indian Oceans. Some of the uncertainty in its origin and range comes from taxonomic confusion with similar species. There seems to be considerable uncertainty about the identity and range of Botrylloides niger. Monniot and Monniot (1997) refer to this species as Botryllus niger (Herdman 1886) but consider ‘B. nigrum’ (Herdman 1906) from Sri Lanka to refer to a separate species. Some records of ‘B. nigrum’ may refer to Botryllus gregalis and Botryllus magnicoecus. This species was reported from South Africa by Sluiter (1898, cited by Van Name 1945) but is not included in Monniot et al.'s (2001) tunicate monograph, and is not included on our map. Records from Australia, as Botrylloides nigrum magnicoecum, are now identified as Botrylloides anceps (Kott 2005). Early records from the Mediterranean (1954, Israel 1958, Tunisia) may have been misidentifications of B. leachi (Izquierdo-Muñoz et al. 2009) Consequently, our map has a slightly less widespread range than that given by da Rocha and Kremer (2005). 

In the Western Atlantic, Botrylloides niger ranges from North Carolina and Bermuda to southern Brazil (Van Name 1945, da Rocha and Kremer 2005). Records are fewer in the Eastern Atlantic, but B. niger is known from Morocco (Monniot 1967; Millar 1968); Madeira (Gestoso et al. 2017), the Cape Verde Islands (Monniot and Monniot 1994), and Angola (2001, Pestana et al. 2017). A genetic study by Sheets et al. (2016) found that Botrylloides niger has a lower level of genetic divergence across its species range.  Genetic diversity was greatest in the tropical Western Atlantic, supporting its possible role as a native region. However, B. niger was established in many Indo-Pacific localities, including the Red Sea and Gulf of Aden, the Persian Gulf, Singapore, Hong Kong, the Philippines, Palau and Tahiti (Van Name 1945; U.S. National Museum of Natural History 2014; Goodbody 1982; Monniot et al. 1985; Sheets et al. 2016; Lambert 2002). 

Invasion History in the Galapagos:

Botrylloides niger was collected in 2015 and 2016 on mangrove roots in Tortuga Bay, on docks at Puerto Ayora, Franklin’s Bay, and on Baltra Island, from settlement plates and cages (Lambert et al. 2019). 

Invasion history elsewhere in the world:

Invasion history for elsewhere in the world has not been summarized for this species at this time.

Description

Botrylloides niger is a colonial tunicate that forms extensive branching systems, often forming dense, crowded colonies, whose limits can be difficult to determine. The zooids are elongate and curved, and about 1.5 to 1.85 mm. They are placed at right angles to the surface. The mantle has many longitudinal muscle bands. The branchial apertures are rounded, while atrial aperture varies from rounded to a transverse slit, which may be produced into a short languet. Adults have 8 larger tentacles, with 8 or more small, or rudimentary tentacles in the intervals. The two lateral tentacles of the larger order are usually largest. There are 3 internal longitudinal vessels on each side, separated by 2 to 4 stigmata, and from the endostyle and dorsal lamina by 5–6 stigmata. The stomach is roughly funnel-shaped with 9–-10 glandular folds. The testis is smaller, with 6–10 lobes. Many colonies have male organs only, but some have a large egg or embryo on one both sides of the body. Egg diameter is ~0.26 mm (Van Name 1945). Colonies in Jamaican mangrove areas are flattened and dark-orange red in color (Goodbody and Webber 2003). In the South Pacific and South Atlantic, the zooids are orange, against a darker background. Specimens in Bahrain has yellow zooids against a black background (Monniot and Monniot 1997). 

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Taxonomy

Ecology

General:

Life History- A colonial (or compound) tunicate consists of many zooids, bearing most or all the organs of a solitary tunicate but modified to varying degrees for colonial life. Colonial tunicates of the genera Botrylloides have small zooids, usually not organized in systems, and fully embedded in a mass of tunic material. Each zooid has an oral siphon and an atrial canal, opening to a shared cloacal chamber. Water is pumped into the oral siphon, through finely meshed ciliated gills on the pharynx, where phytoplankton and detritus is 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. Buds can form from the body wall of the zooid. 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. Eggs are internally fertilized, and embryos are incubated in a brood pouch. Once they are mature, 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). 
 
Botrylloides niger is found in tropical and subtropical environments at average seasonal temperatures of 16–30 degrees C (Sheets et al. 2016). In experiments, Botrylloides niger showed 80% mortality and signs of stress (reduced heart rate, distended cloaca) at 24 PSU, but no mortality at 28 PSU. This tunicate is known from a wide range of natural and artificial surfaces, including rocks, seagrasses, coral reefs, oyster beds, marinas, floats, and vessel hulls. It often grows on other organisms (Van Name 1945; Woods Hole Oceanographic Institution and United States Navy Department 1952; Rodrigues and da Rocha 1993; Goodbody and Webber 2003; Skinner et al. 2003). Structures than from natural surfaces, (Simkanin et al. 2012). Dock floats are especially favored habitats, probably because their motion provides rapid water exchange, and a fresh supply of food-laden water (Glasby 2001). Other colonized man-made structures include pilings, piers, aquaculture structures, and boat hulls (Carman et al. 2010; Davidson et al. 2010; Simkanin et al. 2012). Natural habitats include rocky reefs, bivalve colonies, seaweeds, and eelgrass (White and Orr 2011; Simkanin et al. 2012; Wong and Vercaemer 2012; Carman et al. 2016). Predation may limit or slow the spread of?B. niger. 

Food:

Phytoplankton

Trophic Status:

Suspension Feeder

SusFed

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Habitats

General Habitat	Grass Bed	None
General Habitat	Oyster Reef	None
General Habitat	Mangroves	None
General Habitat	Coral reef	None
General Habitat	Marinas & Docks	None
General Habitat	Vessel Hull	None
General Habitat	Rocky	None
Salinity Range	Polyhaline	18-30 PSU
Salinity Range	Euhaline	30-40 PSU
Tidal Range	Subtidal	None
Vertical Habitat	Epibenthic	None

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Life History

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Tolerances and Life History Parameters

Minimum Temperature (ºC)	16.4	Field, Mediterranean Sea (Sheets et al. 2016)
Maximum Temperature (ºC)	30.3	Field, Mazatlan, Nayarit, Mexico (Sheets et al. 2016)
Minimum Salinity (‰)	28	Experimental, Boca de Toro, Panama (Dijkstra and Simkanin 2016)
Broad Temperature Range	None	Subtropical-Tropical
Broad Salinity Range	None	Polyhaline-Euhaline

General Impacts

Ecological Impacts- Botrylloides niger occurs on other organisms, including bivalves, but its thin colonies are not known to harm bivalves (Rodrigues et al. da Rocha 1993; da Rocha et al. 2009).  It is a common organism in hull fouling and harbor structures (Woods Hole Oceanographic Institution, United States Navy Dept. Bureau of Ships 1952; Rodrigues, and da Rocha 1993).  However, serious economic impacts have not been reported. 

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