Cryptosula pallasiana

Overview

Scientific Name: Cryptosula pallasiana

Phylum: Bryozoa

Class: Gymnolaemata

Order: Cheilostomatida

Family: Cryptosulidae

Genus: Cryptosula

Species:

pallasiana (synonymous with Cribrilina pallasiana, Eschara pallasiana, and Lepralia pallasiana) (Gordon 2015) The distribution of C. p. is unknown because of the earlier confusion with C. okadai (now synonymy of C. zavjalovensis (see Grischenko et al. 2007)). (Mawatari 1992) [Describe here as A. iricolor]

Native Distribution

Origin Realm:

Temperate Northern Atlantic, Temperate Northern Pacific, Temperate Australasia, Tropical Atlantic

Native Region:

Origin Location:

CONFLICT: Northeast Atlantic, Northwest Atlantic, Mediterranean Sea, West Tropical Atlantic, Northeast Pacific, Northwest Pacific, Southern Australia and New Zealand Temperate Northern Atlantic From western Norway southward via British shores to the Mediterranean and Black Sea (NEMESIS 2016). STATED Native to Northeast Atlantic, Mediterranean and Ponto-Caspian (NEMESIS 2016; Lee II and Reusser 2012) STATUS STATED [Italy] Northern Venice Lagoon; Lagoon of Grado; Sacca of Barbamarco; Valli of Comacchio; Lake Fusaro; Lake Miseno; Lake Sabaudia; Stagno of Tortoli; Stagno of Santa Gilla (Ambrogi and d'Hondt 1981; Ambrogi 1985; Annoscia 1985; Occhipinti-Ambrogi 2000 and Morgana & Naviglio 1995, cited in Marchini et al 2007) STATUS STATED [US] Long Island Sound, New York; Florida; North Carolina; Indian River Lagoon; New England; Massachusetts (Altman and Whitlatch 2007; Morrisey and Miller 2009; Stachowicz et al 1999; Osman and Whitlatch 1998; Maturo 1959, cited in Marchini et al 2007; Winston 2009; Rogick and Croasdale 1949; Grosberg 1981; Harbo 2011; Gordon 1973; Rico and Gappa 2006; Carlton 2007) STATUS STATED [UK] Swansea; Poole Harbour; Isles of Man; Robin Hood's Bay; Worm's Head; Port Mooar (Amui-Vedel et al 2007; Dyrynda 1994 and Dyrynda et al 2000, cited in Marchini et al 2007; Watts and Thorpe 2006; Eggleston 1972) STATUS NOT STATED [Portugal] Ria de Aveiro (Marchini et al 2007) STATUS NOT STATED [Norway] (Gordon 1967) STATUS NOT STATED [Egypt] Alexandria; Suez Canal; Port Said Harbour; El-Ghardaqa; Abu Qir Harbour; Eastern Harbours; El-Dekheila Harbour (Abdel-Salam and Ramadan 2008) STATUS NOT STATED [Canada] Nova Scotia; New Brunswick (Harbo 2011; Rico and Gappa 2006; Marchini et al 2007; Rico and Gappa 2006; Gordon 1973; Gordon 1974; Maturo 1958; Winston 1977 and Winston 1982, cited in NEMESIS 2016; Fisheries and Oceans Canada 2011) STATUS STATED Tropical Atlantic [Mexico] East Coast (Morrisey and Miller 2009) STATUS STATED Temperate Northern Pacific [Japan] Maizure; Northern Japan (Hayami 1975; Long and Rucker 1969) STATUS STATED [USA] Narrow Strait, Kodiak, Alaska (Dick & Ross 1985) STATUS NOT STATED [Korea] First studied in 1990, sample from Nodo (Seo 1992) [Korea] Geojedo Island; Heuksando Island; Balsan (Seo and Min 2009) STATUS NOT STATED Temperate Australasia [New Zealand] Leigh (Kaselowsky et al 2005) STATUS NOT STATED Uncertain realm [China] (Tian et al 2010; Tian et al 2011; Tian et al 2014) STATUS NOT STATED

Geographic Range:

Cosmopolitan distribution (Harbo 2011; Gordon 1973; Gordon 1974; Marchini et al 2007; Rico and Gappa 2006; Winston 2009) Widespread around the world (Abdel-Salam and Ramadan 2008; Marchini et al 2007) [Western Pacific] Korea to Japan; New Zealand (Hayami 1975; Long and Rucker 1969; Kaselowsky et al 2005; Seo and Min 2009; Seo 1992) [Eastern Pacific] Alaska; Strait of Georgia, BC to California; Chile (Harbo 2011; Gordon 1973; Gordon 1974; Marchini et al 2007; Rico and Gappa 2006; Winston 2009; Carlton 2007; NEMESIS 2016; Anderson et al 2000) [Western Atlantic] New Brunswick to Florida; Mexico (Gordon 2015; Maturo 1958; Winston 1977 and Winston 1982, cited in NEMESIS 2016) [Eastern Atlantic] Norway to Portugal; Morocco; Egypt (Gordon 1967; Abdel-Salam and Ramadan 2008; Osburn 1952a) [Mediterranean; Indian Ocean] Black Sea; Red Sea (Ryland 1965, Hayward and Ryland 1979 and Hayward and Ryland 1999, cited in NEMESIS 2016; Carlton 2007)

General Diversity:

Probably comprises a global species complex, possibly involving a combination of regional endemic species, upon which ship-fouling introductions have been added (Mead et al 2011, cited in NEMESIS 2016)

Non-native Distribution

Invasion History:

Yes, see inv_propens

Non-native Region:

Northeast Pacific, Northwest Pacific, Tropical Eastern Pacific, Eastern Indo-Pacific, Central Indo-Pacific, Southern Australia and New Zealand, Northeast Atlantic, Mediterranean, Northwest Atlantic, East Tropical Atlantic, West Tropical Atlantic, Southwest Atlantic, Magellanic, Southern Africa

Invasion Propens:

CONFLICT: Northeast Atlantic, Northwest Atlantic, Mediterranean Sea, West Tropical Atlantic, Northeast Pacific, Northwest Pacific, Southern Australia and New Zealand Temperate Northern Pacific Various southern California bays. (Cohen & Carlton 1995) *Introduced Mexico, Vancouver Island, Bodega Harbor, and Coos Bay. (Cohen & Carlton 1995) *Introduced Straits and Sounds, Humbolt Bay. (Wonham & Carlton 2005) *Introduced Northwest Pacific (NEMESIS 2016) *Cryptogenic [Canada] Vancouver Island; Strait of Georgia (Gordon 2015; NEMESIS 2016; Anderson et al 2000; Crawford and Irvine 2011) *Introduced [USA] Narrow Strait, Alaska; San Francisco Bay; Long Beach, California (Dick and Ross 1985; NEMESIS 2016; Blum et al 2007; Crooks et al 2011; Gordon 2015) *Introduced Widely introduced to the Northeast Pacific (Vancouver Island to San Diego) (NEMESIS 2016) *Introduced North Pacific Ocean, Japanese Sea, Coastal Waters of Southeast Alaska and British Columbia, Elkhorn Slough (Gordon 2015) *Alien Cryptogenic species in BC, Canada (Victoria) (Crawford and Irvine 2011) *Cryptogenic Cryptogenic in the Northwest Pacific (Russia-Japan-Korea-China (Zvyagintsev 1985, Huang 2001, Seo 1998, cited in NEMESIS 2016) *Cryptogenic The distribution in Northwest Pacific is also considered as cryptogenic. (NEMESIS 2016) *Cryptogenic East Sea, South Sea and Yellow Sea in Korea. (Seo & Min 2009) *Note: Though C. p. is considered as cryptogenic in the northwest Pacific, first record of the species in Korea in 1999 may mean that this species is introduced species in Korea. Tropical Atlantic Cryptogenic in the East Tropical Atlantic (Russia-Japan-Korea-China (Zvyagintsev 1985, Huang 2001, Seo 1998, cited in NEMESIS 2016) *Cryptogenic Sierra Leone (NEMESIS 2016) *Cryptogenic Temperate South America Puerto Belgrano and QuequÃ©n, Argentina. (Orensanz et al. 2002, Gappa 2000 cited in NEMESIS 2016) *Introduced Rio de Janeiro, Brazil. (Marcus 1942, cited in NEMESIS 2016) *Introduced [Argentina] Quequen Port; Patagonian Harbour (NEMESIS 2016; Brankevich et al 1988; Rico and Gappa 2006) *Introduced [Brazil] South of Rio de Janeiro (NEMESIS 2016) *Introduced Widely introduced to the Southwest Atlantic (Brazil-Argentina) (Australia-New Zealand) (NEMESIS 2016) *Introduced Argentina, Rio de la Plata, Uruguay (Gordon 2015) *Alien Temperate Southern Africa Table Bay Harbor, South Africa. (Mead et al. 2011, cited in NEMESIS 2016) *Introduced [South Africa] (Lee II and Reusser 2012; Gordon 2015) *Alien Temperate Australasia Every ports from Marsden Point to Bluff including Whangateau Harbour, Aotea Lagoon, Akaroa, New Zealand. (Cranfield et al. 1998) *Adventive [New Zealand] Auckland, Takapuna side of the Waitemata Harbour (Gordon 1967; Morrisey and Miller 2009; Floerl et al 2009; NEMESIS 2016) *Introduced [Australia] Records of occurrences from regions: AUS-IV, AUS-IX, AUS-V, AUS-VII, AUS-VIII, AUS-X, and AUS-XII (NIMPIS 2016) *Non-native [Australia] Port Adelaide, South Australia; Gulf of St. Vincent; Port Phillip; Westernport Bays; Sydney Harbor;; Western Australia (Brock 1985; NEMESIS 2016; Gordon & Mawatari 1992, cited in Marchini et al 2007; Vail and Wass 1981; Keough 1998; Lee II and Reusser 2012; NEMESIS 2016) *Introduced New Zealand (Gordon 2015) *Alien Port phillip Bay, Sydney Harbor, and Port Adelaide in Australia. (Keough & Ross 1999) *Likely introduced Tropical Eastern Pacific Oaxaca, Mexico. (Osburn 1952b) *Note: Osburn (1952b) notes that earlier writers did not mention c.p. on the Pacific coast except from Homer, Alaska, but is a fairly common species in southern California. Cryptogenic? Eastern Indo-Pacific FAILED introduction to Hawaii (NEMESIS 2016) *FAILED INTRODUCTION Central Indo-Pacific Port Hackland, Queensland, Australia (Brock 1985; NEMESIS 2016; Gordon & Mawatari 1992, cited in Marchini et al 2007; Vail and Wass 1981; Keough 1998; Lee II and Reusser 2012; NEMESIS 2016) *Introduced Coral Sea (Gordon 2015) *Alien Temperate Northern Atlantic US coast of Atlantic Ocean (Gordon 2015) *Alien [Northeast Atlantic] Morocco to Sierra Leone (NEMESIS 2016) *Cryptogenic Cryptogenic in the Mediterranean (Morrisey and Miller 2009) *Cryptogenic Cryptogenic in the Northwest Atlantic (Lee II and Reusser 2012) *Cryptogenic Cryptogenic: First collected in Northwest Atlantic in 1855 in Great Egg Harbor, New Jersey (Leidy 1855, cited in Fofofoff et al 2003) *Cryptogenic East coast of the USA (NEMESIS 2016) *Cryptogenic Cryptogenic in the Northwest Atlantic (Russia-Japan-Korea-China (Zvyagintsev 1985, Huang 2001, Seo 1998, cited in NEMESIS 2016) *Cryptogenic West Atlantic: Nova Scotia to North Carolina (Hayward & Ryland 1999), though that in the west Atlantic is considered as cryptogenic (NEMESIS 2016). *Cryptogenic Uncertain realm Australia, Brazil, Canada, Mexico, South Africa, South Atlantic Ocean (Gordon 2015) *Alien Gulf of Mexico (NEMESIS 2016) *Cryptogenic Southwest Pacific (Australia-New Zealand) (NEMESIS 2016) *Introduced

Status Date Non-native:

First non-native North American Marine/Estuarine record: 1943 (NEMESIS 2016) First Non-native West coast marine/estuarine record: 1943, Hunter's Point Shipyard in San Francisco Bay, US (Carlton 1979 and Cohen and Carlton 1995, cited in NEMESIS 2016) [Alaska, US] Collected in 1951 (James Carlton, personal communication, cited in NEMESIS 2016) [California, US] Present many years before 1943 (James Carlton, personal communication 2013, cited in NEMESIS 2016) [United States] San Francisco Bay: 1944-1947. (Cohen & Carlton 1995) [United States] Newport Bay: 1944. (Scheer 1945, cited in NEMESIS 2016) [United States] Alamitos Bay: 1962. (Reish 1963, cited in NEMESIS 2016) [United States] Los Angeles Harbor: 1965. (Bnta 1970, cited in NEMESIS 2016 [United States] Morro Bay: 2001. (Fairey et al. 2002, cited NEMESIS 2016) [United States] Monterey Bay: before 1952. (Osburn 1952b) [United States] Elkhorn Slough: 1988. (Wasson et al. 2001, cited in NEMESIS 2016) [United States] Tomales Bay and Fort Bragg Harbor: 2000. (Fairey et al. 2002, cited in NEMESIS 2016) [United States] Bodega Harbor: before 1975 (Standing 1975) and 2000 (Fairey et al. 2000, including former referece, both were cited in NEMESIS 2016). [United States] Humbolt Bay: 2000. (Boyd et al. 2002, cited NEMESIS 2016) [United States] Coos Bay: 1998. (Wonham & Carlton 2005) [United States] Padilla Bay: 2003. (de Rivera et al. 2005, cited in NEMESIS 2016) [Canada] Pendrell Sound, Vancouver Island. (Powell 1970, cited in NEMESIS 2016) [Argentina] Puerto Belgrano and QuequÃ©n: 1978. (Orensanz et al. 2002, Gappa 2000, cited in NEMESIS 2016) [Brazil] Reported in 1942 (NEMESIS 2016) [Australia] Hobsons Bay and Port Phillip Heads, Port Phillip Bay: late 1800'S. First collected in Australian waters in Port Phillip Bay in 1890 (Keough & Ross 1999) [New Zealand] Port of Wanganui, Napier, and Dunedin: at least 1890s. New Zealand at 'almost every port' in 1895 (Gordon & Mawatari 1992) [South Africa] Collected in Table Bay Harbor, South Africa in 1947 (Mead et al. 2011, cited in NEMESIS 2016) [Korea] First studied in 1990, sample from Nodo (Seo 1992) [China] Collected in 2009 (Tian et al 2010; Tian et al 2011; Tian et al 2014) CONFLICT: Cryptogenic: First collected in Northwest Atlantic in 1855 in Great Egg Harbor, New Jersey (Leidy 1855, cited in Fofofoff et al 2003)

Vectors and Spread

Initial Vector:

Hull fouling (not specified), Ballast water, Aquaculture and Fisheries

Second Vector:

Vector Details:

Hull fouling; Aquaculture & Fisheries: Atlantic Oyster (Lee II and Reusser 2012) Common in ship fouling (Ryland 1964, cited in NEMESIS 2016) Western coast of Canada: Aquaculture (oysters) and ship fouling (Anderson et al 2000) [Australia] Accidental with deliberate translocations of fish or shellfish (Gordon 2015) Fisheries and Aquaculture; Vessels (NIMPIS 2016) Competitive fouling organism likely introduced via ballast water or hull fouling (Morrisey and Miller 2009) Probably all of New Zealand's non-indigenous bryozoans have arrived via fouling of ships' hulls or (more recently) sea chests (internal hull recesses housing ballast and cooling water intakes (Coutts & Dodgshun 2007; Floerl et al 2009) [United States] San Francisco Bay: hull fouling and in shipments of Atlantic oyster. (Cohen & Carlton 1995) [United States] Northern Pacific region: hull fouling and commercial oyster industry. (Wonham & Carlton 2005) [New Zealand] Hull fouling. (Cranfield et al. 1998) [Australia] Port Phillip Bay: hull fouling. (Hewitt et al. 2004)

Spread Rate:

Newer records does not reflect recent expansion of the species, but rather simply a lack of prior reporting (James Carlton, personal communication, 2013, cited in NEMESIS 2016) Expanded its southern limit of distribution in the southwest Atlantic by 7Â° of latitude (Lopez Gappa 2000, cited in Rico and Gappa 2006) Newly found for first time in Yellow Sea, materials collected in 1999 (Seo and Min 2009)

Date First Observed in Japan:

Unknown but have been observed as early as 1966 (Long and Rucker 1969) Fossil records from late Pliocene (Hayami 1975) The distribution of C. p. is unknown because of the earlier confusion with C. okadai (now synonymy of C. zavjalovensis (see Grischenko et al. 2007)). (Mawatari 1992)

Date First Observed on West coast North America:

Kaneohe Bay, Hawaii: 1966 (Lee II and Reusser 2012) First recorded at Hunter's Point Shipyard in San Francisco Bay in 1943, subsequently at Newport Bay in 1944. Species undoubtly present many years prior (NEMESIS 2016) Sitka, Alaska in 1951 (Jame Carlton, personal communication, cited in NEMESIS 2016)

Impacts

Impact in Japan:

Global Impact:

[Australia] Water abstraction or nuisance fouling; outcompetes native species for resources and/or space (Gordon 2015) C. p. is a common species in ship and dock fouling communities (Ryland 1965, Zvyagintsev 1985, cited in NEMESIS 2016); however, there are no economic and ecological impacts reported for this bryozoan. (NEMESIS 2016) C. p. is not reported in the literature as a strong competitor, so its impact is unlikely to be substantial. (Keough & Ross 1999)

Tolerences

Native Temperature Regime:

Cold water, Cool temperate, Mild temperate, warm temperate, Subtropical, Tropical

Native Temperature Range:

Considered a boreal-tropical species (Dick and Ross 1985) Broad temperature range: Cold temperate to warm temperate (NEMESIS 2016; Winston 2009) Sampled at temperature: 6.894-16.565°C (OBIS 2015) Subtropical; 13.9 to 20.5˚C (Kaselowsky et al 2005) Cold water, Cool temperate, Mild temperate, Warm temperate (M. Otani, pers. comm.)

Non-native Temperature Regime:

Cold water, Cool temperate, Mild temperate, Warm temperate, Subtropical, Tropical

Non-native Temperature Range:

Considered a boreal-tropical species (Dick and Ross 1985) Collected at sites with surface temperatures 12.5-25.7°C and 19.5-34.2°C (Brock 1985) Temperate subtidal site at Port Phillip Bay (Keough 1998) Maximum temperate: 24.7°C; cold temperate -warm temperate (Lord et al 2015, cited in NEMESIS 2016; NEMESIS 2016) Temperature sampled: 6.894-16.565°C (OBIS 2015) Vancouver, BC: max 13.0ºC in summer and min -1.0ºC in winter. (Clark et al. 2003) San Francisco: max 20.0ºC in summer and min 11.0ºC in winter. (Clark et al. 2003) San Diego: max 22.0ºC in summer and min 13.0ºC in winter. (Clark et al. 2003) Water temperature in Port Phillip Bay is ranging from 21ºC in summer to 11ºC in winter. (Thresher 1999) Cold water, Cool temperate, Mild temperate, Warm temperate (M. Otani, pers. comm.)

Native Salinity Regime:

Polyhaline, Euhaline

Native Salinity Range:

Salinity: Brackish, Polyhaline (alpha) and Marine (beta) (Lee II and Reusser 2012) Tolerant of brackish water; Polyhaline-Euhaline (NEMESIS 2016) Salinity range from 19-38%o (NEMESIS 2016) Salinity ranged from 18.330-39.053 PPS (Obis 2015) Found at stations with salinity between 36-39 (Marchini et al 2007)

Non-native Salinity Regime:

Oligohaline, Mesohaline, Polyhaline, Euhaline

Temperature Regime Survival:

Cool temperate, Mild temperate, Warm temperate, Subtropical, Tropical

Temperature Range Survival:

Maximum temperature: 24.7°C (Lord et al 2015, cited in NEMESIS 2016) Broad temperature range: Cold temperate-warm temperate (NEMESIS 2016) [Laboratory] Colonies grown at 14°C had 2x growth rate for zooids than at 18°C (Amui-Vedel et al 2007) [Laboratory] Zooids grown at 14°C were on average 23.79μm longer and 18.55 μm wider than those grown at 18°C; number of tentacles significantly higher at 14°C than 18°C (Amui-Vedel et al 2007) Sampled at temperature: 6.894-16.565°C (OBIS 2015) Cool temperate, Mild temperate, Warm temperate, Subtropical, Tropical (M. Otani, pers. comm.)

Temperature Regime Reproduction:

Temperature Range Reproduction:

Salinity Regime Survival:

Mesohaline, Polyhaline, Euhaline

Salinity Range Survival:

[Italy] Frequent in samples from localities with higher range of salinities (not less than 18-20%o) (Ambrogi and d'Hondt 1981) Associated with sites more directly influenced by the sea, showing high salinity (range 22.5-34%o ) and high current velocities; Somewhat tolerant of low salinity, more than Bugula species (Ambrogi 1985) Tolerant of brackish water (NEMESIS 2016) Minimum salinity 19%o (Occhipinti Ambrogi and d'Hondt 1981; NEMESIS 2016) Maximum salinity 38%o (NEMESIS 2016) Salinity ranged from 18.330-39.053 PPS (Obis 2015) Salinity as low as 10psu is sublethal for C. p. (Turpaeva 1960, cited in Gordon & Mawatari 1992)

Salintiy Regime Reproduction:

Polyhaline, Euhaline

Salinity Range Reproduction:

Depth Regime:

Mid intertidal, Lower intertidal, Shallow subtidal, Deep subtidal, See details

Depth Range:

CONFLICT: Maximum depth 35m (NIMPIS 2016), but also listed deeper than 35m Occurs in estuaries, coastal bay, nearshore and shelf regimes (Lee II and Reusser 2012) Benthic depth range: intertidal and subtidal (sub-shallow) preferred; observed in sub-deep subtidal as well and occurring at 0-60m (Lee II and Reusser 2012) Intertidal and shallow subtidal (Harbo 2011; Watts and Thorpe 2006; Dick and Ross 1985; Keough 1998; Gordon 1973; NIMPIS 2016; Rico and Gappa 2006) Intertidal to 60m depth (Carlton 2007; Kaselowsky et al 2005) Sampled depth range: 0-130m (OBIS 2015) Fairly shallow water (Ryland 1965, cited in Abdel-Salam and Ramadan 2008) Fairly common in the littoral zone (Osburn 1952a) British shores: from just below MTL and into the shallow sublittoral. (Hayward & Ryland 1999) Hueneme, California: 29 fms and Anacapa Island, California. (Osburn 1952b) Mid intertidal (M. Otani, pers. comm.)

Non-native Salinity Range:

Native Abundance:

Abundant, Common

Reproduction

Fertilization Mode:

Internal

Reproduction Mode:

Hermaphrodite/monoecious

Spawning Type:

Development Mode:

Lecithotrophic planktonic larva (non-feeding)

Asexual Reproduction:

Budding/fragmentation (Splitting into unequal parts. Buds may form on the body of the â€œparentâ€)

Reproduction Details:

Sexual and asexual reproduction; colonies consist of replicated series of zooids, each budded asexually from a predecessor. Founding zooid metamorphoses from the sexually produced larva. Hermaphroditic (Gordon 2015; Amui-Vedel et al 2007) Hermaphrodite; Asexual reproduction: Budding (Lee II and Reusser 2012) Early development: live birth w/ nutritional supply; Juvenile development dispersal: larval phase is planktonic larvae (Lee II and Reusser 2012) Ovicells absent (small avicularium is sometimes present proximal to the aperture) (Kozloff 1996; Carlton 2007; Abdel-Salam and Ramadan 2008) Brooded in body cavity (Carlton 2007) Internal brood sac (Hincks 1861, cited in Ostrovsky 2013; Eggleston 1972) Lecithotrophic larvae (Ostrovsky 2013) RELATED: [Gymnolaemates] Internal fertilization, whether intracoelomic or intraovarian, is obligatory (Temkin 1994 and 1996, cited in Ostrovsky 2013) [Gymnolaemates] Differ from most organisms in that sperm-egg fusion does not stimulate egg activation. Egg activation may not occur until "spawned" outside of maternal zooid (Temkin 1991) [Bryozoans] While sperm is spawned through pores in lophophore tentacles, eggs are usually harbored inside the body wall, and are internally fertilized by sperm, coming in on lophophore feeding currents (Brusca and Brusca 2003, cited in Rouse 2011; Kozloff 1990, cited in Rouse 2011) [Bryozoans] Colonial hermaphrodites, with testes (spermatogenic tissue) and ovaries developing either within the same zooid (zooidal hermaphroditism) or in different zooids within the same colony (zooidal gonochorism) (Ostrovsky 2013) [Bryozoans] Members of the phylum Bryozoa are hermaphroditic. Both fertilization and egg brooding may either be internal or external (Ruppert et al. 2004) [Bryozoans] The first zooid in a colony is called the ancestrula. It is from this individual that the rest of the colony will grow asexually from the budding (Hill 2001) [Bryozoa] All bryozoan colonies are hermaphroditic. Autozooids may be dioecious; or monoecious, and protandrous or protogynous. (Hayward & Ryland 1999) [Bryozoa] Reproduces asexually by budding. (Mawatari 1976)

Adult Mobility:

Sessile

Adult Mobility Details:

Colonial on hard substratum epifauna (Gordon 2015; NEMESIS 2016) RELATED: [Bryozoa] The abundance and taxonomic diversity of benthic bryozoan faunas are directly related to substratum. (Hayward & Ryland 1999) [Bryozoa] Bryozoans are a phylum of sessile, colonial suspension feeders found throughout the world in both marine and freshwater environments. (Tilbrook 2012)

Maturity Size:

Colony: 20cm wide (Harbo 2011) Colonies are 600 to 1000Î¼m x 350-500Î¼m in size (NEMESIS 2016) Zooids measure on average 0.8mm in length and 0.44mm in width (NIMPIS 2016) Zooids moderately large; 0.65x0.35-0.45mm (Osburn 1952a) Colony size up to 3cm in diameter. Zooids are 0.43-0.65mm long and 0.26-0.45mm wide (Abdel-Salam and Ramadan 2008)

Maturity Age:

Completion of metamorphosis (formation of a functional juvenile lophophore) took approximately 4 days for Cryptosula pallasiana (Kosman and Pernet 2011)

Reproduction Lifespan:

Embryos most abundant during the summer at British coast. (Heyward & Ryland 1999) Settlement recorded My to December in western Norway, with peak July and September. (Ryland 1963, cited in Hayward & Ryland 1999) Settlement in Britain continues through October. (Hayward & Ryland 1999) Settlement occurs in spring, summer and autumn in Port Phillip Bay. (Keough & Ross 1999) Settlement is heaviest during spring and autumn months at Lyttelton, New Zealand. (Skerman 1958, cited in Gordon & Mawatari 1992)

Longevity:

Polypides survive for 2-10 (15-72 days) weeks in aquarium conditions; duration of regression takes place over 6-17 days (Gordon 2013; Gordon 1973, cited in Gordon 2013) Dead zooid within a colony may be regenerated as a whole new zooid by one or more neighbouring zooids (Jebram 1977) Considered a perennial species, long-lived species at least 2-3 years (Eggleston 1972; Kuklinski and Taylor 2006)

Broods per Year:

In an experiment, 19 larvae were collected from one colony of Cryptosula pallasiana (Kosman and Pernet 2011)

Reproduction Cues:

RELATED: [Bryozoans] Experiments often used light as a cue to collect embryos/larvae (Woollacott and Zimmer 1977) [Bryozoa] In coastal species light is an important stimulus to larval release, and many cheilostomates shed larvae during the first few hours of daylight. (Hayward & Ryland 1999) [Bryozoa] In various degrees of intensity according to the species temperature also stimulates sexual reproduction. (Winston 1977)

Reproduction Time:

[Alaska, US] October, November and December samples were found with embryos, but may be due to tide levels and greater species diversity at lower intertidal ranges (Dick and Ross 1985) [California, US] Avery Point: recruitment levels of at least 50 individuals per 100cm2 per week at some point between June and October; Similarly in summer 2002 (Altman and Whitlatch 2007) July is period of active growth for C. pallasiana (Amui-Vedel et al 2007) Larval release can occur from May to December in Europe (Ryland 1965, cited in Gordon 1973) [US] Larvae reported in August and September; also found in June and July (Barrois 1877, cited in Rogick and Croasdale 1949) Embryos most abundant during the summer at British coast. (Heyward & Ryland 1999) Settlement recorded My to December in western Norway, with peak July and September. (Ryland 1963, cited in Hayward & Ryland 1999) Settlement in Britain continues through October. (Hayward & Ryland 1999) Settlement occurs in spring, summer and autumn in Port Phillip Bay. (Keough & Ross 1999) Settlement is heaviest during spring and autumn months at Lyttelton, New Zealand. (Skerman 1958, cited in Gordon & Mawatari 1992)

Fecundity:

Egg Size:

Early and mid-stage embryos: 160.0x120.0; 180.0x150.0Î¼m (Ostrovsky 2013) RELATED: [Gymnolaemata] About 200Âµm (Woollacott and Zimmer 1977)

Egg Duration:

Early Life Growth Rate:

Specific growth rates: at 14Â°C: 0.034+/- 0.0097; at 18Â°C: 0.066+/- 0.0158 (Amui-Vedel et al 2007) [Laboratory] Zooids grown at 14Â°C were on average 23.79Î¼m longer and 18.55 Î¼m wider than those grown at 18Â°C (Amui-Vedel et al 2007) [UK] Average length of zooids in July (20Â°C) was significantly longer than in January (5Â°C); ; zooids measured could have been budded at an unknowable period prior to the collection date (Amui-Vedel et al 2007) Cryptosula pallasiana released larvae of an average volume of 0.0086 Â± 0.0001 mm^3; completion of metamorphosis (time to formation of a functional juvenile lophophore) took 4 days (Kosman and Pernet 2011) RELATED: [Gymnolaemata] Two phases of larvae metamorphosis: first stage about 20mins; second stage 1-6 days (Woollacott and Zimmer 1977)

Adult Growth Rate:

Specific growth rates: at 14Â°C: .0017-0.050, average 0.034+/- 0.0097; at 18Â°C: 0.041-0.092, average 0.066+/-0.0158 (Amui-Vedel et al 2007) [Laboratory] Zooids grown at 14Â°C were on average 23.79Î¼m longer and 18.55 Î¼m wider than those grown at 18Â°C; approximately 4 weeks to reach maximum colony size (Amui-Vedel et al 2007) [UK] Average length of zooids in July was significantly longer than in January; zooids measured could have been budded at an unknowable period prior to the collection date (Amui-Vedel et al 2007) [New Zealand] Spring settling colonies measured 12 cm2 (Skerman 1958 and 1959, cited in Menon and Nair 1972) Brooded larvae of Cryptosula pallasiana in Nova Scotia were developing in the aquarium for approximately 30 days (Gordon 1977, personal communications, cited in Ostrovsky 2013)

Population Growth Rate:

[US] Avery Point: recruitment levels of at least 50 individuals per 100cm2 per week at some point between June and October; Similarly in summer 2002 (Altman and Whitlatch 2007) [Laboratory] Colonies reached a maximum of nearly 300 completely formed feeding zooids after a period of 4 weeks at 18Â°C; only one colony reached maximum of slightly more than 100 zooids at 14Â°C (Amui-Vedel et al 2007)

Population Variablity:

Habitat

Ecosystem:

SAV, Rocky intertidal, Rocky subtidal, Mussel reef, Oyster reef, Kelp forest, Fouling, Macroalgal beds

Habitat Type:

Epibenthic, Epiphytic, Epizoic

Substrate:

Rock, Gravel, Cobble, Biogenic, Hardpan, Artificial Substrate

Exposure:

Exposed, Semi-exposed, Protected

Habitat Expansion:

Habitat Details:

VARIABILITY: Substratum preferences may not be constant throughout the entire range of the species; around British coasts it is commonest species on stones but occur rarely on algae, yet in some Norwegian fjords, it encrusts littoral fucoids and is not be found on stones (Ryland 1962b; Gordon 1967) Unconsolidated ecosystem: Sub-aquatic vegetation; Consolidated ecosystem: Rocky intertidal, subtidal rocky, oyster/mussel reef, fouling, kelp forest (Lee II and Reusser 2012) Consolidated substrate: rock, biogenic (oyster, kelp, and rooted aquatic), artificial substrate (pilings, hull/ballast, and others) (Lee II and Reusser 2012) Epibenthic, Epiphytic and epizoic (Lee II and Reusser 2012) [Canada] Species found on rocks and algae (Gordon 1973; Gordon 1974) [Alaska, US] Found in intertidal rock-pile habitat in Narrow Strait (Dick and Ross 1985) [California, US] Collected from floating docks in Long Beach (Kosman and Pernet 2011) [Mass, US] Collected on green algae species: Enteromorpha intestinalis, Ulva Lactuca var. rigida; on brown algae species: Ascophyllum mackaii, Ascophyllum modosum, F. svesiculosus spiralis and Laminaria agardhii; encrusts on rocks shells and algae; more commonly and forms larger colonies on the harder substrates than on algae (Rogick and Croasdale 1949) [Korea] Found on stones and shells (Seo 1992; Seo and Min 2009) [Australia] Found at sheltered site at Port Phillip Bay(Keough 1998) [New Zealand] Found on stones and boulders, shells and/ or barnacle plates, wharf piles and ascidian tests (Gordon 1967) [Portugal] Floating pontoons and wooden piles, ports (Marchini et al 2007) [Italy] Occurs on mussels, barnacles, aquatic macrophytes and rotten wood; dominant in marine parts of channels in the Northern Venice Lagoon (Ambrogi and d'Hondt 1981) Well known fouling species; Collected on boats, pontoons, oysters and cement aggregate tiles (Brock 1985) Attached to rocks, shells, floats and pilings, seaweeds, seagrasses and tunicates in the intertidal and shallow subtidal (Harbo 2011) On eelgrass blades, rocks, oysters, pilings, ships' hulls, and other hard substrates (NEMESIS 2016) Substrate: bedrock, concrete, oysters, reef, wood (NIMPIS 2016) Found encrusting on any surface that will provide attachment, including ships' hulls; widespread particularly in ports, harbours, and estuarine situations (Ryland 1965 and Gordon & Mawatari 1992, cited in Abdel-Salam and Ramadan 2008) Greater on sandstone surfaces, less on concrete and least on wooden surfaces. More on horizontal undersides and less on vertical panels; in harbours where it is sheltered and residential area (Glasby 2000; Glasby and Connell 2001) Found on panels on rocky reefs more than on panels on pontoons at all sites (Glasby and Connell 2001) Abundant on stones and shells, rock overhangs, and wooden pile (Ryland 1962a; Ryland 1962b) Relatively common in the sounds and estuaries (Maturo 1958) Boat's hull; piece of clothing; Encrusting on anything that will afford attachment (Seo and Min 2009; Osburn 1952a) Substratum preferences may not be constant throughout the entire range of the species; around British coasts it is commonest species on stones but occur rarely on algae, yet in some Norwegian fjords, it encrusts littoral fucoids and is not found on stones (Ryland 1962b; Gordon 1967) C. p. is found on stones, shells and other hard substrata as well as kelp holdfast and Himanthalia buttons in British coast. (Hayward & Ryland 1999) C. p. is found on rock, shell, ascidians, glass, pilings, and painted and ferro-cement hulls of pleasure craft and fishing boat. (Gordon & Mawatari 1992) C. p. occurs away from piers and other artificial structures. It is found commonly in the very low intertidal zone of boulder fields. (Keough & Ross 1999) Exposed, Semi-exposed (M. Otani, pers. comm.)

Trophic Level:

Suspension feeder

Trophic Details:

Suspension feeder (Lee II and Reusser 2012) Is a suspension feeder on phytoplankton (Hughes 1992) Suspension feeder, feeds on phytoplankton (NEMESIS 2016) RELATED: [Bryozoans] Suspension feeder...filter phytoplankton less than 0.045mm in size from the water column. (Hill 2001) [Bryozoa] Many phytoplankton species are cleary unsuitable as food for bryozoans. (Hayward & Ryland 1999) [Cheilostomata] Main food is diatom, protozoans and etc. and unappropriate sized particles are ejected (Mawatari 1976)

Forage Mode:

Generalist

Forage Details:

Feeds on phytoplankton. Feeds by extending ciliated tentacles of the lophophore as a funnel, creating a current, and driving food particles into their mouths (NEMESIS 2016) RELATED: [Bryozoans] Suspension feeder...filter phytoplankton less than 0.045mm in size from the water column. (Hill 2001) [Bryozoa] Many phytoplankton species are cleary unsuitable as food for bryozoans. (Hayward & Ryland 1999) [Cheilostomata] Main food is diatom, protozoans and etc. and unappropriate sized particles are ejected (Mawatari 1976)

Natural Control:

POLLUTION [Pollution] Species is frequently found near channel mouths, where water circulation is very active and clearer waters are present. Species is affected adversely by turbid and polluted environments (Ambrogi 1985; Nair 1961, cited in Ambrogi 1985) PREDATION [Predation] Grazing organisms such as sea urchins and fish (Gordon 2015) [Predation] Fed on by Okenia plana, Onchidoris muricata (Carlton 2007) [Predation] Fish, crabs, etc. (Altman and Whitlatch 2007) [Predation] Eaten by various nudibranchs, sea urchins and fish (Morrisey and Miller 2009) [Predation] Juveniles did not seem to be affected by any of the predators (Osman and Whitlatch 1998) COMPETITION [Competition] Intra- and interspecific competition (Altman and Whitlatch 2007) [Competition] Competition and overgrowth from sponges, algae, and tunicates (Gordon 2015) [Competition] Associated with Ciona intestinalis-free communities (Blum et al 2007) [Competition] Susceptible to invasions when growing alone (Stachowicz et al 1999) [Competition] In a study, Botryllus schlosseri was found to be 100% successful in overtopping and killing >50% of C. pallasiana (Grosberg 1981) DISTURBANCE [Disturbance] Storm events (Altman and Whitlatch 2007) RELATED: PREDATION [Predation] [Bryozoa] Browsers and grazers, including sea urchins, fish, crabs and some prosobranchs, are known to include bryozoans in their diet. (Hayward & Ryland 1998) [Predation] [Bryozoa] Bryozoans are also the prey of very many small, selective predators, some of which may be adapted to a very narrow spectrum of prey species. Among them opisthobranch predators of bryozoans are well known. (Hayward & Ryland 1998) [Predation] [Bryozoa] Other than opisthobranchs as a predator, amphipods, isopods, mites and pycnogonids have all been recorded preying on bryozoan colonies. (Hayward & Ryland 1998) EPIBIONTS [Epibionts] [Cheilostomata] It is frequently observed in Japan that several species of hydroids flourish on Cheilostomata cause damages to them. (Mawatari 1976)

Associated Species:

References and Notes

References:

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M., Hayward, P. J., & Porter, J. S. (2007). Zooid size and growth rate of the bryozoans Cryptosula pallasiana Moll in relation to temperature, in culture and in its natural environment. Journal of Experimental Marine Biology and Ecology, 353(1), 1-12. Doi: 10.1016/j.jembe.2007.02.020 Anderson, E. P., Austin, W. C., Byers, S., & Lipovsky, S. (2000). Literature Review on Non-indigenous Invertebrates of the Strait of Georgia. Retrieved from http://www.dfo-mpo.gc.ca/Library/336461.pdf Annoscia, E. (1985). Bryozoan Studies in Italy in the Last Decade: 1973-1982. In C. Nielsen & G. P. Larwood (Eds.), Bryozoa: Ordovician to Recent: Papers Presented at the 6th International Conference on Bryozoa, Vienna 1983 (pp. 1-7). Fredensborg, Denmark: Olsen&Olsen Blum, J. C., Chang, A. L., LiljesthrÃ¶m, M., Schenk, M. E., Steinberg, M. K., & Ruiz, G. M. (2007). The non-native solitary ascidian Ciona intestinalis (L.) depresses species richness. 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Literature:

Extensive scientific information; peer-reviewed information; data specific to the location; supported by long-term datasets (10 years or more)

Notes:

The distribution of C. p. is unknown because of the earlier confusion with C. okadai (now synonymy of C. zavjalovensis (see Grischenko et al. 2007)). (Mawatari 1992)

JTMD Species Summary