Celleporella hyalina

Overview

Scientific Name: Celleporella hyalina

Phylum: Bryozoa

Class: Gymnolaemata

Order: Cheilostomatida

Family: Hippothoidae

Genus: Celleporella (Synonymous with Schizoporella or Hippothoa (Ryland 1962b); C. hyalina likely involves a worldwide complex of similar, perhaps cryptic species. (Grischenko et al. 2007))

Species:

hyalina [Describe here as A. iricolor]

Native Distribution

Origin Realm:

Arctic, Temperate Northern Pacific, Central Indo-Pacific, Tropical Eastern Pacific, Temperate South America, Tropical Atlantic, Temperate Northern Atlantic, Temperate Australasia

Native Region:

Origin Location:

CONFLICT: Southern Australia and New Zealand Arctic Bering Sea; Chukchi Sea (Osburn 1952, cited in Dick et al. 2005; Denisenko and Kuklinski 2008; Grischenko 2002; McCartney 1997; Waeschenback et al 2012; Trott 2004; Carlton 2007; Crooks et al 2011; Maturo 1958; Rogick and Croasdale 1949; Oregon State University 1971) STATUS STATED It extends from the Arctic down European coasts as far as the Bay of Biscay and down the Pacific coast of America to California, parts of Mexico affected by cold water upwelling and Chile. (Hayward & Ryland 1999) STATUS NOT STATED Chukchi Sea; Bering Sea; East Siberian Seas (Ostrovsky 1998; Ivin and Zvyagintsev 2001; Gontar 1981; Denisenko and Kuklinski 2008; Grischenko 2002; Denisenko 2011) STATUS NOT STATED Temperate Northern Pacific Admiralty Inlet, Puget Sound, Washington, USA (De Palma 1966), Strait of Georgia, Canada (Powell 1967, etc., cited in Winston 1977) STATUS NOT STATED It certainly extends from the Arctic down European coasts as far as the Bay of Biscay and down the Pacific coast of America to California, parts of Mexico affected by cold water upwelling and Chile. (Hayward & Ryland 1999) STATUS NOT STATED Kurile Islands (Mawatari 1956) STATUS NOT STATED [Japan] Akkeshi Bay, Hokkaido. (Grischenko et al. 2007) STATUS NOT STATED [Japan] Akkeshi; Kamtschatka; Paramushir; Samani; Kushiro; Abashiri; Muroran; Mori; Shirikishinai; Hakodate; Otaru; Sea of Japan; Kurile Islands; Far Eastern Seas (Namikawa et al. 1992; Kaselowsky et al 2005; Grischenko et al 2007; Mawatari and Mawatari 1981; Grischenko and Zvyagintsev 2012; Kashin et al 2003; Gontar 1981) STATUS NOT STATED [United States] Alaska; Washington; California (Osburn 1952, cited in Dick et al. 2005; Denisenko and Kuklinski 2008; Grischenko 2002; McCartney 1997; Waeschenback et al 2012; Trott 2004; Carlton 2007; Crooks et al 2011; Maturo 1958; Rogick and Croasdale 1949; Oregon State University 1971) STATUS STATED [Mexico] Gulf of California (Hermansen et al. 2001; Soule 1961) STATUS NOT STATED [Russia] Sredhiy Island; Tchupinskaja Inlet; Possiet Bay; Siberia; Kurile Islands; Far Eastern Seas (Ostrovsky 1998; Ivin and Zvyagintsev 2001; Gontar 1981; Denisenko and Kuklinski 2008; Grischenko 2002; Denisenko 2011) STATUS NOT STATED [Hippothoa hyalina (Synonymized taxon)] Abashiri, Akkeshi, Muroran, Mori, Shirishikinai, Hakodate, Otaru (Hokkaido) and northern to middle Honshu. (Mawatari & Mawatari 1981) STATUS NOT STATED [Hippothoa hyalina (Synonymized taxon)] Seto Inland Sea (Inaba 1988) STATUS NOT STATED Central Indo-Pacific [Philippines] (Mawatari 1956) STATUS NOT STATED [Malaysia] (Mawatari 1956) STATUS NOT STATED Tropical Eastern Pacific Galapagos Island reported by Osburn (1952) under the name Hppothoa hyaline (Synonymized taxon) is another locality. STATUS NOT STATED [Galapagos] Galapagos Islands (Carlton 2007) STATUS NOT STATED It certainly extends from the Arctic down European coasts as far as the Bay of Biscay and down the Pacific coast of America to California, parts of Mexico affected by cold water upwelling and Chile. (Hayward & Ryland 1999) STATUS NOT STATED [Mexico] Pacific Coast (Hermansen et al. 2001; Soule 1961) STATUS NOT STATED Temperate South America Ria Deseado, Argentina (Amor & Pallares 1965, cited in Winston 1977) STATUS NOT STATED It certainly extends from the Arctic down European coasts as far as the Bay of Biscay and down the Pacific coast of America to California, parts of Mexico affected by cold water upwelling and Chile. (Hayward & Ryland 1999) STATUS NOT STATED [Argentina] Santa Cruz (Gappa 1989) STATUS NOT STATED [Chile] (Bishop et al. 2000; Waeschenback et al 2012)) STATUS NOT STATED Tropical Atlantic [Brazil] Bahia (Kelmo et al 2004) STATUS NOT STATED Temperate Northern Atlantic Tamar Estuary and Plymouth, England (Milne 1940, etc., cited in Winston 1977), Baltic Sea, Kattegat Area including inner area (Brog 1930, etc. and Androsova 1962, ect., cited in Winston 1977), Western Norway (Nair 1961, 1962, cited in Winston 1977) STATUS NOT STATED [United Kingdom] Isles of Man; Isles of Scilly; Menai Strait; Wales; Scotland; British Isles (Bishop et al. 2000, Eggleston 1972; Goldson et al. 2001; Gómez et al. 2007; Hermansen et al. 2001; Hoare et al. 1999; Hughes 1989; Hughes 1992; Manríquez et al 2001; Maughan et al 2000; Ryland 1959; Ryland 1960; Ryland 1962a; Waeschenback et al 2012; Goss-Custard et al 1979; Hayward 1971; Gómez et al. 2007; O'Connor et al. 1980) STATUS NOT STATED [Ireland] Cork; Galway; Wexford (Keegan et al 1987) STATUS NOT STATED [Spain] San Felipe (Gómez et al. 2007) STATUS NOT STATED [Belgium] Belgian Coast (De Blauwe 2009, cited in Gordon 2015) STATUS NOT STATED [The Netherlands] Dutch Coast (De Blauwe 2009, cited in Gordon 2015) STATUS NOT STATED [Croatia] Adriatic Basin; Danube Basin (Koletic et al 2015) STATUS NOT STATED [United States] Maine; North Carolina; Massachusetts (Osburn 1952, cited in Dick et al. 2005; Denisenko and Kuklinski 2008; Grischenko 2002; McCartney 1997; Waeschenback et al 2012; Trott 2004; Carlton 2007; Crooks et al 2011; Maturo 1958; Rogick and Croasdale 1949; Oregon State University 1971) STATUS STATED [Canada] Gulf of St. Lawrence (Miller 2012, cited in Gordon 2015) STATUS NOT STATED [Norway] Svalbard (Kuklinski et al. 2005; Kuklinski et al 2006; Waeschenback et al 2012; Wlodarska-Kowalczuk et al 2009; Porter et al. 2015) STATUS NOT STATED [Sweden] (Waeschenback et al 2012) STATUS NOT STATED [Iceland] Reykjavik (Gómez et al. 2007) STATUS NOT STATED Temperate Australasia [New Zealand] Auckland (Gordon 1967) STATUS NOT STATED Uncertain realm Cosmopolitan (Hastings 1929; Maturo 1958; Soule 1961) STATUS NOT STATED Circumglobally distributed in colder seas (McCartney 1997) STATUS NOT STATED [China] (Mawatari 1956) STATUS NOT STATED [United States] Alaska (Osburn 1952, cited in Dick et al. 2005; Denisenko and Kuklinski 2008; Grischenko 2002; McCartney 1997; Waeschenback et al 2012; Trott 2004; Carlton 2007; Crooks et al 2011; Maturo 1958; Rogick and Croasdale 1949; Oregon State University 1971) STATUS STATED

Geographic Range:

Cosmopolitan distribution in temperate zone epialgal communities; circumglobally distributed in colder seas (Hughes 1989; McCartney 1997; Hayward 1929; Maturo 1958; Waeschenback et al 2012; Soule 1961) [Western Pacific] Far Eastern Sea; Japan; China; Philippine; Malaysia; New Zealand (Namikawa et al. 1992; Kaselowsky et al 2005; Grischenko et al 2007; Mawatari and Mawatari 1981; Grischenko and Zvyagintsev 2012; Kashin et al 2003; Gontar 1981; Gordon 1967; Hermansen et al 2001; Mawatari 1956) [Eastern Pacific] Chukchi Sea; Alaska to Chile, South America (Bishop et al 2000; Waeschenback et al 2012; Dick et al 2005; Oregon State University 1971) [Western Atlantic] Maine to North Carolina (Waeschenback et al 2012; Dick et al 2005; Denisenko and Kuklinski 2008; Grischenko 2002; Carlton 2007; Oregon State University 1971) [Eastern Atlantic] Iceland; Sweden to Spain; Mediterranean Sea (Gómez et al 2007; Waeschenback et al 2012) [Japan] Aininkappu Cape in Akkeshi Bay: 42°59.6'N, 144°51.3'E. (Grischenko et al. 2007) [Japan] Aikappu Cape, tip in Akkeshi Bay: 43°00.4'N, 144°50.1'E. (Grischenko et al. 2007) [Japan] Daikokujima Island, north side in Akkeshi Bay: 42°57.3'N, 144°52.5'E. (Grischenko et al. 2007) [Japan] Daikokujima Island, west side in Akkeshi Bay: 42°57.0'N, 144°52.1'E. (Grischenko et al. 2007) [Japan] Mabiro Cape in Akkeshi Bay: 42°58.6'N, 144°53.2'E. (Grischenko et al. 2007) [Japan] From 34°N to 60°N both at Pacific and Japan Sea side. (Inaba 1988)

General Diversity:

Species complex that may have at least 11 major genetically divergent lineages worldwide (Dick et al. 2005; Waeschenback et al 2012; Porter et al 2015) Strong population structure, with significant isolation by distance. NE Atlantic C. hyalina is structured into two main parapatric lineages that appear to have had independent Pleistocene histories (Gómez et al 2007) A northern Atlantic Species; C. hyalina sensu lato in the eastern Pacific is a complex of species (Carlton 2007) Two main parapatric lineages for NE Atlantic Celleporella hyalina. Range expansions have resulted in two contact zones in Spain and W. Ireland. Lineage 1 is found from Ireland to Spain...recent population expansion into the Irish Sea, S. Ireland, S. England and Spain. Lineage 2 is found from Iceland to Spain. (Gómez et al. 2007)

Non-native Distribution

Invasion History:

Yes (Hewitt et al. 2004)

Non-native Region:

Southern Australia and New Zealand

Invasion Propens:

CONFLICT: Southern Australia and New Zealand Temperate Australasia Port Phillip Bay, Australia. (Hewitt et al. 2004) *Introduced RELATED: Temperate Northern Atlantic [Celleporella species] Described as nonindigenous recent invaders in the Venice lagoon (Italy) (Occhipinti Ambrogi & d'Hondt 1996, Hewitt et al. 1999, Boyd et al 2002, cited in Gómez et al. 2007) Temperate Australasia [Celleporella species] Described as nonindigenous recent invaders in Port Philip Bay (Australia) (Occhipinti Ambrogi & d'Hondt 1996, Hewitt et al. 1999, Boyd et al 2002, cited in Gómez et al. 2007) Temperate Northern Pacific [Celleporella species] Described as nonindigenous recent invaders in Humboldt Bay (California, USA) (Occhipinti Ambrogi & d'Hondt 1996, Hewitt et al. 1999, Boyd et al 2002, cited in Gómez et al. 2007)

Status Date Non-native:

First record in Port Phillip Bay, Australia is 1889. (Hewitt et al. 2004)

Vectors and Spread

Initial Vector:

Hull fouling (not specified), Natural dispersal

Second Vector:

NF

Vector Details:

Rafting of colonies on detached seaweed or other drifting substrata (Goldson et al 2001) Data does not suggest human-mediated dispersal has impacted significantly on C. hyalina populations (Gómez et al 2007) Hull fouling (Hewitt et al. 2004)

Spread Rate:

Considered to have low dispersal abilities in NE Atlantic (Gómez et al 2007) From Pleistocene, range expansions from independent lineages have expanded to two contact zones in Spain and W. Ireland. Lineage 1 is found from Ireland to Spain and has low haplotype diversity, with closely related haplotypes, suggesting a recent population expansion into the Irish Sea, Ireland, S England and Spain. Lineage 2 is found from Iceland to Spain and has high haplotype diversity. (Gómez et al 2007)

Date First Observed in Japan:

Considered to date back to Pleistocene (Gómez et al 2007) Kamtshatka samples collected in June 1912 (Mawatari 1956)

Date First Observed on West coast North America:

Considered to date back to Pleistocene (Gómez et al 2007) [Alaska, US] Dick et al conducted their surveys in 2003 (Dick et al 2005)

Impacts

Impact in Japan:

[Japan] Marine fouling, affects corrosion in different materials and hydrodynamic loads in constructions; also attribute positive aspects as active filters of polluted water (Kashin et al 2003)

Global Impact:

NF

Tolerences

Native Temperature Regime:

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

Native Temperature Range:

Inhabits cold-temperate to polar oceans circumglobally (Gómez et al. 2007; McCartney 1997) Colonies were maintained at 15°C, within 5°C of mean summer sea-surface temperature maxima at all native localities ( Gómez et al. 2007) Collected at water temperatures from -1.516 to 11.845°C (OBIS 2015) Historically been treated as a widely distributed species. Osburn (1952) noted it as a "truly cosmopolitan species, occurring around the world and from the Arctic...to the tropics" (cited in Dick et al. 2005) [Santa Cruz, Argentina] Water temperatures are typically subantarctic and influenced by the Malvinas Current, varying from about 4.5°C in winter to 13.5°C in summer (Gappa 1989) [Kongsfjorden, Norway] Surface water temperature of below -0.5°C (Kuklinski et al 2006) [Japan] Akkeshi Bay: The lowest recorded water temperature was -1.4ºC in February 2003, the highest 21.1ºC in August 2004. (Grischenko et al. 2007) Cold water, Cool temperate, Mild temperate, Warm temperate, Subtropical, Tropical (M. Otani, pers. comm.)

Non-native Temperature Regime:

Cool temperate, See details

Non-native Temperature Range:

Water temperature in Port Phillip Bay is ranging from 21ºC in summer to 11ºC in winter. (Thresher 1999) Cool temperate (M. Otani, pers. comm.)

Native Salinity Regime:

Polyhaline, Euhaline

Native Salinity Range:

Collected at salinity levels of 31.235-35.203 PSU (OBIS 2015) [Santa Cruz, Argentina] Salinity varies around 33%o (Gappa 1989) [Kongsfjorden, Norway] Salinity above 34.4 psu (Svendsen et al 2002, cited in Kuklinski et al 2006) [Japan] Akkeshi Bay: relatively constant, about 30psu; it ranged from 26psu in June 2003 to 31psu in August 2004. (Grischenko et al. 2007) It is also said that C. h. is tolerant of conditions ranging from near-oceanic to estuarine. (Grischenko et al. 2007) RELATED Cribrilina, Membranipora, Hippoporina, Schizoporella, Smittina and related genera are supported by polyhaline water. Although Membranipora is an euryhaline genus, it is found in mesohaline water, namely 3-16%o (Hayami 1975)

Non-native Salinity Regime:

Polyhaline, Euhaline

Temperature Regime Survival:

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

Temperature Range Survival:

Collected at water temperatures from -1.516 to 11.845°C (OBIS 2015) Evidence of inverse relationship between temperature and size in larval paraenchymal cells and colony modules (autozooids) but no inverse temperature-size relationship observed for lengths of tentacles or their epidermal cells, or the volumes of whole colonies at reproductive maturity. Tested temperatures were 10°C and 18°C. Its suggested that variation in colony volume closely reflected the variation in number, not sizes, of autozooids (Atkinson et al. 2006) Growth rate and growth efficiency were significant reduced at 22°C when compared to 15°C and 19°C, with growth frequently ceasing completely at 22°C during the 8-15 day interval. The growth rate measured at 19°C was overall significantly higher than that measured at 15°C, with exception of pH 7.3 treatment. Growth efficiency was not significantly different between 15°C and 19°C (Pistevos et al 2011) Historically been treated as a widely distributed species. Osburn (1952) noted it as a "truly cosmopolitan species, occurring around the world and from the Arctic...to the tropics" (cited in Dick et al. 2005)

Temperature Regime Reproduction:

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

Temperature Range Reproduction:

Half-lives of spermatozoa in dilute suspension reported as 1.2 h at 12°C, 16°C and 18°C (Manríquez et al. 2001; Bishop and Pemberton 2006) Temperature was not found to significantly influence the rate of decline in fertility of suspended allosperm (Manríquez et al. 2001) Reproductive investment increased overall with increasing temperature and decreasing pH. At pH 7.3, the index increased significantly between 22°C and 15°C and 19°C. (Pistevos et al 2011) Over 8 day interval, study did not show masculinisation in response to increased temperature. Gender allocation moved significantly towards females between 19°C and 22°C. Index did shift towards males between the ambient pH level of 8.1 and both 7.6 and 7.3. Results may be affected by the slow of growth at 22°C where few zooids of any kind were produced (Pistevos et al. 2011)

Salinity Regime Survival:

Euhaline

Salinity Range Survival:

Collected at salinity levels of 31.235-35.203 PSU (OBIS 2015)

Salintiy Regime Reproduction:

Polyhaline, Euhaline

Salinity Range Reproduction:

NF

Depth Regime:

Lower intertidal, Shallow subtidal, Deep subtidal

Depth Range:

[Isles of Man, UK] Samples were collected from shore down to 100 m depth (Eggleston 1972) [UK] Isles of Scilly: 9-52m (Hayward 1971) [Svalbard, Norway] Species dominated in shallow (0-40m) areas (18%) (Kuklinski et al 2005) [Galapagos] 10-15 fms (18-27m) and shoreline (Hastings 1929) Inhabits the low intertidal-shallow subtidal zone of cold-temperate to polar oceans circumglobally (Gómez et al. 2007) Intertidal species living at maximum water depths of about 50m (Ryland and Hayward 1977 and Hayward and Ryland 1979, cited in Hermansen et al 2001) Sampling depth for species: 0 - 538m (OBIS 2015) From intertidal to > 130m (Carlton 2007; Oregon State University 1971) Akkeshi Bay: lower intertidal zone. (Grischenko et al. 2007) The Isle of Man: intertidal to sublittoral: to 55m. (Eggleston 1969, cited in Hayward & Ryland 1999)

Non-native Salinity Range:

Native Abundance:

Common, Abundant

Reproduction

Fertilization Mode:

Internal

Reproduction Mode:

Hermaphrodite/monoecious

Spawning Type:

NA

Development Mode:

Lecithotrophic planktonic larva (non-feeding)

Asexual Reproduction:

Budding/fragmentation (Splitting into unequal parts. Buds may form on the body of the “parent”)

Reproduction Details:

Species is hermaphroditic and spermcast (Bishop and Pemberton 2006; Hoare et al. 1999; Pemberton et al 2003) Populations can substantially or completely prevent selfing by incompatibility mechanisms; Little evidence of self-fertilization in colonies studied, typically although not exclusively an obligate outbreeder (Hoare et al. 1999; Hughes et al 2002, cited in Bishop and Pemberton 2006; Hoare and Hughes 2001; Hughes et al. 2002b) Long-distance fertilizations could occur, implying utilization of dilute spermatozoa (Yund and McCartney 1994, cited in Bishop and Pemberton) [Chile] Female zooids are only produced when genetically different colonies are cultured together (Cancino et al 1991, cited in Bishop et al. 2000) although male zooids are present in isolated colonies (Bishop et al 2000) [UK] Cultured in reproductive isolation; Female zooids may be produced, but their oocytes remain small and are not ovulated. Single brief exposure to a compatible mate or allosperm results in outcrossed progeny (Manríquez 1999, cited in Bishop et al 2000) Maximum production of zygotes could occur down to sperm concentrations of 50 sperm per ml (Pemberton et al. 2003, cited in Bishop and Pemberton 2006) Spermatozoa can be captured by autozooids and stored in the absence of female zooids; although site of storage is unknown; Stored spermatozoa can be subsequently used to fertilize eggs some weeks later following colony maturation; Sperm shed externally while eggs are retained and fertilized internally (Bishop and Pemberton 2006; McCartney 1997; Hughes et al 2002, cited in Bishop and Pemberton 2006) Sperm may precociously penetrate immature ovarian oocytes, with male pronucleus remaining separate in the cytoplasm until syngamy occurs near the time of passage of the egg from the coelom of the maternal zooid to the site of brooding (Ostrovsky 1998, cited in Bishop and Pemberton 2006) Passage of inseminated eggs into ovicells to commence development may occur up to several weeks after uptake of allosperm by adult colonies (Manríquez 1999, cited in Bishop and Pemberton 2006) Separate male, female and feeding zooidal polymorphs; During mating, sperm enter the coelom of a zooid via the dorsal coelomopore and precociously penetrate ovarian primary oocytes in the early stages of growth (Temkin 1996 and Ostrovsky 1998, cited in Bishop et al 2000; Bishop et al. 2000; Ostrovsky et al 2009) Colonies had lower budding rates when grown close to neighbouring colonies than when isolated, but the ratio of reproductive zooids to autozooids was similar (Cancino and Hughes 1987) Male zooids were always produced first (Cancino and Hughes 1987, Hughes et al. 2002) Evidence of self-fertilisation, albeit with a drastic reduction in larval viability, has been reported for Welsh colonies kept in reproductive isolation; Self-fertilisation may be available as an emergency option but naturally, species routinely outcross and limits self-fertilisation (Hunter and Hughes 1993b; Goldson et al. 2001; Hughes et al 2009) Autozooids capture sperm during filter-feeding, and eggs, presumably after fertilization, pass from the female zooids into special chambers called ovicells, where the embryos undergo placental brooding until released as free-swimming larvae (Gómez et al. 2007) Each colony is founded by a sexually produced larva which metamorphoses to form the primary zooid or ancestrula. In mature colonies, male and female gonozooids are budded from an underlying layer of sterile feeding zooids.(Hughes 1992; Hughes 1987) Unlike many bryozoans, colonies of C. hyalina in nature do not propagate by fission, and genetically distinct colonies are therefore best regarded as modular 'individuals' rather than clones, analogous to a population of unitary aclonal animals (Hughes 1989; Hughes 1992; Hughes et al. 2004) Evidence that when under stress, species increases the ratio of male to female modules in response to diverse environmental stressors; possibly to enhance siring success under competition (Hughes et al. 2003; Hughes et al. 2009; McCartney 1997) Produce sexual zooids through process known as frontal budding, form layer on top of the basal autozooid layer; Because sexual zooids only occur in a single layer, the number of sexual zooids is maximally limited by the area of the underlying layer of feeding zooids (Ryland and Gordon 1977; Cancino and Hughes 1988, cited in McCartney 1997; Hughes 1987) After entering the ovicell, the embryo received an additional input of nutrients from the female zooid - placental system (Hughes 1987b) The eggs in C. hyalina are microlecithal, produced sequentially and receive extra embryonic nutrition during embryogenesis in a specialized brood chamber; Lecithotrophic species (Ostrovsky 1998; Temkin 1991) Completion of the ovicell took approximately four days from the stage at which the zooid was first recognizable as a female (Hughes 1987) Brooding in ovicell with presence of embryophore (Ostrovsky 2013) RELATED: [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) [Celleporella hyalina] Fertilisation occurs internally during late vitellogenesis. Colonies bear embryos for a few weeks only and died soon after all the larvae had been released. (Hayward & Ryland 1999)

Adult Mobility:

Sessile

Adult Mobility Details:

Sedentary species, sessile clonalial invertebrate which encrusts macroalgae (Cancino and Hughes 1987; Hoare et al. 1999) Sessile, dispersal is assumed to occur through the nonfeeding, yolk-rich larvae in a short planktonic phase. Nature of C. hyalina's common substratum (macroalgae) makes it likely that rafting on detached floating fronds could be a significant means of dispersal, enabling colonization of new and distant habitats (Gómez et al. 2007) RELATED: Rafting and fouling may be more important for dispersal than larval motility although it is unknown how far populations can travel on storm-severed algal fronds or other flotsam and jetsam (Hoare et al. 1999; Watts et al 1998) [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:

[US] Largest colony observed 1.5 cm across in Ketchikan, Alaska (Dick et al. 2005) Zooids: 0.48-0.63mm long (average=0.542mm) by 0.23-0.28 mm wide (average=0.248mm) (Dick et al. 2005) Orifice: 0.13-0.16mm long (average=0.140mm) by 0.11-0.14mm wide (average = 0.124mm) (Dick et al. 2005) Females zooid bodies are 80-100μm long (Hughes 1987b)

Maturity Age:

Male sexual maturity is marked by appearance of male zooids at about four weeks after settlement. Female zooids first appear some seven to ten days later (McCartney 1977) Potential partners placed together and maintained for 2 weeks. Each colony was therefore able to act as both a male and female in each mating ( Gómez et al. 2007) A period of 3 weeks for all allosperm recipients bore mature embryos in a laboratory setting; Colonies repeatedly exposed to allosperm began to produce embryos in week 3 and to reach a sustainable level at week 5 (Hughes et al 2002)

Reproduction Lifespan:

C. hyalina can be maintained on artificial substrata for >18 months with repeated seasonal cycles of reproduction (Cancino 1983, cited in Cancino and Hughes 1987) Allosperm can be captured and stored for several weeks and used to fertilise a series of eggs (Manríquez 1999, cited in Manríquez et al 2001) Observed half-life of about 1 hr for water-borne sperm of Celleporella hyalina. Allosperm older than 12hr in all but one case failed to produce embryos. The 1 case produced 1 embryo (Manríquez et al. 2001) Male and female zooids are budded and function simultaneously throughout remainder of a colony's life (McCartney 1977) There are three separate generations during the year, with peak embryo production during February-March, May-August, and October-November. (Eggleston 1972, cited in Hayward & Ryland 1999)

Longevity:

[UK] maximum life span of 4 months (Cancino 1986, cited in Cancino and Hughes 1987) Normally life span constrained to a time scale of months by deterioration of the substratum or competitive overgrowth (Cancino 1986, cited in Hughes et al. 2003; Seed and O'Connor 1981) [Laboratory conditions] C. hyalina can be maintained on artificial substrata for >18 months (Cancino 1983, cited in Cancino and Hughes 1987) [Laboratory] First autozooids of colonies recruited in October 1980 fed for 3-4 weeks, then degenerated and regenerated within 1-2 weeks. During 3 months of observation, the zooids underwent 3 cycles of degeneration and regeneration. The maximum number of regeneration cycles that autozooids are able to survive remains unknown but the centers of colonies became inactive after 3-4 months, corresponding to the longevity of female zooids (Cancino and Hughes 1987) Male zooids had a life span of 2-3months whereas female zooids were active for 3-4 months (Cancino and Hughes 1987) The life cycle is completed quickly and colonies live for less than a year. (Hayward & Ryland 1999)

Broods per Year:

Female zooids were active for 2-4 months and produced two to three larvae each; Only one egg per cycle is brooded per female zooid (Cancino and Hughes 1987; McCartney 1997) Three period of peak reproduction each year. These are probably separate generations as each colony bears embryos for only a few weeks and dies soon after all the larvae are released (Eggleston 1972; Ostrovsky 1998) In the observed colony, female zooids became inactive and presumably senescent, after a maximum of four successive brooding cycles (Hughes 1987b)

Reproduction Cues:

Vitellogenic egg growth leading to ovulation is triggered by receipt of compatible allosperm; Can acquire and store water-borne allosperm and use it to fertilize ova for a period of 3-6 weeks after reaching female sexual maturity (Bishop et al 2000, cited in Bishop and Pemberton 2006; Hughes et al. 2002) Unrelated or half-sib spermatozoa trigger the budding of additional female zooids (Hughes et al 2002, cited in Bishop and Pemberton 2006) Egg activation and nuclear fusion are delayed until the grown egg is ovulated and passes from the zooidal coelorn (Temkin 1996, cited in Bishop et al 2000) Physical contact between different colonies is not necessary for female maturation to be initiated, exposure to water conditioned by another clone is sufficient (Bishop et al. 2000) Independent adult ramets of the same clone do not trigger each other's egg growth; factor triggering oocyte growth is present in cross-water; factor can be removed by 0.45μm filtration and contents of the seminal vesicle of a compatible clone pipetted in a minute total volume can trigger oocyte growth (Bishop et al. 2000) Although release and uptake of non-gametic courier cells (or cell components) cannot be totally discounted, the direct involvement of allosperm in triggering vitellogenic oocyte growth appears by far the stronger probability (Bishop et al. 2000) Colonies began sexual reproduction at a size of about 57 autozooids, but at different ages depending on growth conditions; relationship of total reproductive zooids and number of autozooids became linear beyond a colony size of 500-700 autozooids (Cancino and Hughes 1987) Larval release may be controlled by light conditions, where larvae released early in the morning to leave sufficient time for them to encounter suitable substratum and settle during daylight. This would enable the larvae, then in their photonegative phase, to detect shaded microhabitats (Cancino et al. 1991) Larval release occurred almost entirely during daylight, beginning shortly after sunrise and peaking approximately 1.5hrs later, by which 30-40% of total release had taken place. Tides has a secondary effect by controlling the amount of light reaching colonies (Cancino et al. 1991) 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) [Celleporella hyalina] Peak larval release occurs shortly after sunrise, allowing larvae maximum opportunity to find a suitable settlement site within their 12 hour free-swimming period. (Cancino et al. 1994, cited in Hayward & Ryland 1999)

Reproduction Time:

Larvae released after 91 days of colonies growing (Atkinson et al. 2006) The proportion of sexual zooids that were reproductively active was greatest in June and least in winter (December - February) (Cancino and Hughes 1987) C. hyalina can be maintained on artificial substrata for >18 months with repeated seasonal cycles of reproduction (Cancino 1983, cited in Cancino and Hughes 1987) Colonies showed a peak of sexual activity during June 1981; reproduced sexually at higher rates from April to September than during the rest of the year (Cancino and Hughes 1987) Each brooding period about 3-4 weeks in duration, during which the embryo is retained within the ovicell (Cancino and Hughes 1987) Three periods of peak reproduction (February and March, May to August and October and November) each year. These are probably separate generations as each colony bears embryos for only a few weeks and dies soon after all the larvae are released (Eggleston 1972) Maximum reproductive activity occurs in late summer and early autumn (Cancino 1986, cited in Goldson et al. 2001) Reproduction, larval settlement and metamorphosis extend over the whole summer and at least the beginning of autumn in the White Sea, but seasonal observations are necessary to characterize the annual cycle (Ostrovsky 1998; Ryland 1959) There are three separate generations during the year, with peak embryo production during February-March, May-August, and October-November. (Eggleston 1972, cited in Hayward & Ryland 1999) Embryo presented in the specimens of 66.6% collected in Akkeshi Bay during the periods 2-7 June and 3-6 July . (Grischenko et al. 2007)

Fecundity:

When colonies had accumulated 1000-2000 autozooids, the number of reproductive zooids (males + females) per autozooid became asymptotic in the ratio of 1.0:2.0:3.6 in conditions of restricted, semi-restricted, and unrestricted water flow (Cancino and Hughes 1987) The proportion of sexual zooids that were reproductively active was greater among colonies experiencing greater water flow (Cancino and Hughes 1987) Fertility of released sperm decays exponentially, with a half-life of about 1.2hrs (Manríquez et al. 2001; Goldson et al. 2001) Only one egg per cycle is brooded per female zooid (McCartney 1997) Maximum of five larvae per female in colonies raised under laboratory conditions (Cancino 1983, cited in Hughes 1987b) Each female zooid is capable of brooding a maximum of four successive embryos. (Hayward & Ryland 1999)

Egg Size:

Eggs remain in female body for 2-3 days before transference to newly-completed ovicell and is about 80μm in diameter at this stage. By this time, egg is probably fertilized. Embryo spends 12-14 days in the ovicell and the larva is approximately 200μm in diameter at maturity (Hughes 1987b) Eggs undergo a 15-fold increase in volume during the approximately 16-day brooding cycle in the ovicell (Hughes 1987b; McCartney 1994, cited in McCartney 1997) Ovulated egg is approximately volume 2.68x10^5 μm^3 but is greater size within the ovicell (Hughes 1987b) RELATED: Ovicell: 0.18-0.23mm long (average = 0.195mm) by 0.18-0.26mm wide (average=0.214mm) (Dick et al. 2005) [Gymnolaemata] About 200µm (Woollacott and Zimmer 1977)

Egg Duration:

Eggs appeared in the body of females at the moment the ovicells were empty or when an old larva was about to leave the ovicell (Cancino and Hughes 1987) Approximately 16 day brooding cycle in the ovicell (McCartney 1977) Embryo spends 12-14 days in the ovicell and the larva is approximately 200μm in diameter at maturity (Hughes 1987b) Passage of inseminated eggs into ovicells to commence development may occur up to several weeks after uptake of allosperm by adult colonies (Manríquez 1999, cited in Bishop and Pemberton 2006) Colonies bear embryos for a few weeks only. (Hayward & Ryland 1999)

Early Life Growth Rate:

[Laboratory] Colonies initially grew at similar rates for all treatments (water flow rates), with a mean instantaneous rate of 0.125 per zooid per day from March to May 1980; Colonies in greater water flow eventually grew larger, produced more sexual zooids and had a greater reproductive zooids to autozooids ratio than those experiencing less water flow (Cancino and Hughes 1987) Colonies established in October 1980 and placed in restricted and unrestricted water flow grew exponentially for the first 50 days, but growth rate declined during the next 2 months (mid December - mid February) before regaining a higher level in the spring; winter mean instantaneous growth rate was greater in restricted than in unrestricted flow (Cancino and Hughes 1987) Colonies began producing males within a month, at a mean colony size of 57 autozooids (Cancino and Hughes 1987) Sexual maturation occurs early in colonial development (Cancino and Hughes 1987) Embryo spends 12-14 days in the ovicell and the larva is approximately 200μm in diameter at maturity. About a 13-15 fold increase in volume during the brooding period (Hughes 1987b; Ostrovsky 1998) RELATED: [Gymnolaemata] Two phases of larvae metamorphosis: first stage about 20mins; second stage 1-6 days (Woollacott and Zimmer 1977)

Adult Growth Rate:

Reproductive zooids/autozoooids became asymptotic after colonies reached 1000-2000 autozooids, ratio 1.0:2.0:3.6 for colonies grown in restricted, semi-restricted and unrestricted water flow (flow rates 1.0:1.8:3.0) (Cancino and Hughes 1987) Female zooids appeared as colonies grew larger, sex ratio rising as colony size increased to about 1000 autozooids (Cancino and Hughes 1987) Growth rate of 0.13 per day, regardless of flow rates (Cancino and Hughes 1987) [Menai Strait, UK] Growth rate was 0.07-0.10 per day on microscope slides, in relation to colony size; on macroalgae, the average specific growth rate was 0.06 per day, in relation to colony size (Hermansen et al. 2001) Feeding rate, and thus growth- seems to decrease with increasing colony size, which may be due to changes in shape parameter (Bishop and Bahr 1973, cited in Hermansen et al. 2001) Larvae takes 3-4 weeks to develop in natural conditions in North Wales (Cancino and Hughes 1988, cited in Ostrovsky 2013), although developmental time can be shorter, just 12-14 days (Hughes 1987, cited in Ostrovsky 2013)

Population Growth Rate:

Colonies grew faster from April to September than during the rest of the year (Cancino and Hughes 1987) [Menai Strait, UK] Growth rate was 0.07-0.10 per day on microscope slides, in relation to colony size; on macroalgae, the average specific growth rate was 0.06 per day, in relation to colony size (Hermansen et al. 2001)

Population Variablity:

Significant variation in larval behaviour patterns has been reported for geographically distant populations (Orellana et al 1996, cited in Hunter et al 1999) Exhibited genetic differentiation over distances as small as 10m; results indicate that a relationship exists between genetic and geographic distance, leading to localised population structure (Goldson et al 2001) NE Atlantic C. hyalina is structured into two main parapatric lineages that appear to have had independent Pleistocene histories (Gómez et al 2007)

Habitat

Ecosystem:

Rocky intertidal, Rocky subtidal; Mussel reef, Macroalgal beds, Kelp forests, Fouling

Habitat Type:

Epibenthic, Epiphytic, Epizoic

Substrate:

Cobble, Rock, Biogenic, Artificial substrate

Exposure:

Exposed, Semi-exposed, Protected

Habitat Expansion:

NF

Habitat Details:

[UK] Normally lives on the temporary frondal tissue of the kelp Laminaria saccharina L. where it is an enforced ephemeral; occurs as an epiphyte of laminarian and fucoid algae; encrusting Fucus serratus L. plants (Cancino 1986, cited in Cancino and Hughes 1987; Eggleston 1972; Hughes 1989; O'Connor et al 1980) [US] Alaska: Collected colonies growing intertidally on rocks, as well as subtidal on mussel shells (Dick et al. 2005) [US] Beaufort: Reported to settle on almost any substrate; only found in the area on larger algae, particularly the browns such as Sargassum and Zonaria (Maturo 1958) [US] Collected on brown algae: Ascophyllum nodosum, F. vesiculosus spiralis, Laminaria agardhii, and Sargassum filipendula (Rogick and Croasdale 1949) [Norway] Svalbard and Kongsfjorden: Dominated algal substrates, comprising 50% of colonies; also on large rocks...colonies were not detected from substrates of soft bottom or small rocks in the study (Kuklinski and Barnes 2005, cited in Kuklinski et al. 2005; Kuklinski et al 2006) [New Zealand] Found on stones and boulders, shells and/or barnacle plates, algae and glass (Gordon 1967) Found on more exposed shores found in small amounts on a number of algae, especially in Laminaria holdfasts, but in certain sheltered localities it forms heavy incrustations on the fronds of L. saccharina. (Ryland 1959; Ryland 1962a; Hayward 1971) On rocky shores, found in small amounts on a number of algae, most zoaria are found on the rhizoids of Laminaria digitata and L. hyperborea (Ryland 1959) Where L. saccharina grows in abundance, then the polyzoan forms extensive crustations over the thallus; larvae tends to select concavities than convexities on the frond (Ryland 1959) Larvae settle on the fronds of Fucus serratus L. and Laminaria saccharina; macroalgae (Goldson et al. 2001; Hayward and Ryland 1979, cited in Hermansen et al. 2001) Sessile modular colonies live epiphytically, typically on macroalgal fronds, often in large numbers. Most often found in relatively sheltered areas, such as coastal bays, inlets, straits, rías, and fjords (Gómez et al. 2007) Habitat in Millepora coriacea (Linneaus 1767) Described as an early successional species, found more abundant at the more disturbed sites in the study; also described as abundant on boulders and kelp in high flow rapid sites (Maughan et al 2000) Artificial substrata often used to rear colonies in laboratory experiments and other studies (Manríquez et al 2001; Maughan et al 2000; Hunter and Hughes 1993a) Substrata observed on: stones and boulders, shells, hydroids, polyzoans, algae (Ryland 1962a) Encrusts rock, shell, algae, hermit crabs (Carlton 2007) Common on red algae fronds (Oregon State University 1971) Essentially on species of algae in shallow water, often in shelter reaching great abundance on Laminaria, especially L. saccharina, on which the colonies clump in the dimples on the frond; occasionally on stones and shells. (Hayward & Ryland 1999) C. h. is found mainly on rocks and also found on substrata are shells, algae, concrete, plastic debris, and other bryozoans such as Alcyonidium alcilobatum, Flustrellidra akkesiensis, and Flustrellidra corniculata (Grischenko et al. 2007)

Trophic Level:

Suspension feeder

Trophic Details:

Is a suspension feeder on phytoplankton (Hughes 1992) 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:

Within a colony, there is a complete separation between feeding functions and reproduction functions (Hughes and Hughes 1986) Feeds using a tentaculate organ called a lophophore (Pistevos et al 2011) [Laboratory] Survived and grew on a range of algal diets. Chlorophyte algae proved least suitable, cryptophyte Rhodomonas baltica found to be outstanding as a foodstuff (Hunter and Hughes 1993a) 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:

ECOLOGY: Colonies grown in unrestricted (faster) water flow eventually became larger than those grown in restricted water flow; water flow can affect the size of colonies and potentially, reproduction (Cancino and Hughes 1987; Hughes and Hughes 1986) COMPETITION: [Menai Straits, UK] Avoided Halidrys siliquosa in the study (Ryland 1959, cited in Ryland 1962a) [Northern Ireland, UK] Celleporella hyalina tended to co-occur with Electra pilosa but avoided Flustrellida hispida and Alcyonidium hirsutum (O'Connor et al 1980) [Chile] A reduction in initial colony size (may be caused by extended period of larval swimming past 4hrs) has been found to affect eventual reproductive output and increase the likelihood of competitive overgrowth (Ryland 1981 and Stocker 1991, cited in Goldson et al 2001; Cancino and Hughes 1987) Out of 51 times observed, 38 had instances where Anomia ephippium undercutted C. hyalina. Anomia ephippium had high undercut rates throughout the study (Maughan et al 2000) Dense algal growth at 1-2ft below water surface may restrict larvae settlement whereas lower below surface (3-4ft), settlement was more equal on different surfaces of panels; species may be selecting shaded surfaces away from light (Ryland 1960) Fertilization competition as sessile invertebrates. Success rates highly correlate with number of male zooids produced by colonies (Yund and McCartney 1994) PREDATION [Predation] Dironapicta feeds on Celleporella hyalina (Carlton 2007) RELATED: PREDATION [Predation] Destruction of zooids by predators (Harvell's paper was for another species but cited by Hughes et al. 2003 when discussing potential environmental stress to bryozoans and C. hyalina) (Harvell 1984, cited in Hughes et al. 2003) [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:

NF

References and Notes

References:

Atkinson, D., Morley, S. A., & Hughes, R. N. (2006). From cells to colonies: at what levels of body organization does the ‘temperature-size rule’ apply?. Evolution & Development, 8(2), 202-214. Doi: 10.1111/j.1525-142X.2006.00090.x Bishop, J. D. D., Manríquez, P. H., & Hughes, R. N. (2000). Water-borne sperm trigger vitellogenic egg growth in two sessile marine invertebrates. Proceedings of the Royal Society B: Biological Sciences, 264(1449), 1165-1169. Doi: 10.1098/rspb.2000.1124 Bishop, J. D. D. & Pemberton, A. J. (2006). The Third Way: Spermcast Mating in Sessile Marine Invertebrates. Integrative and Comparative Biology, 46(4), 398-406. Doi: 10.1093/icb/icj037 Cancino, J. M. (1983). Demography of Animal Modular Colonies (Modular Colonies) (Doctoral dissertation). Retrieved from ProQuest Dissertation and Theses (DX92348). Cancino, J. M. & Hughes, R. N. (1987). The effect of water flow on growth and reproduction of Celleporella hyalina (L.). (Bryozoa: Cheilostomata). Journal of Experimental Marine Biology and Ecology, 112(2), 109-130. Doi: 10.1016/0022-0981(87)90112-2 Cancino, J. M., Hughes, R. N., & Ramirez, C. (1991). Environmental Cues and the Phasing of Larval Release in the Bryozoan Celleporella hyaline (L.). Proceedings: Biological Sciences, 246(1315), 39-45. Carlton, J. T. (Ed.). (2007). The Light and Smith Manual: intertidal invertebrates from central California to Oregon. Los Angeles, CA: Univ of California Press. César-Aldariz, J., Fernández-Pulpeiro, E., & Reverter-Gil, O. (1999). A new species of the genus Celleporella (Bryozoa: Cheilostomatida) from the European Atlantic coast. Journal of the Marine Biological Association of the United Kingdom, 79(1), 51-55 Crooks, J. A., Chang, A. L., & Ruiz, G. M. (2011). Aquatic pollution increases the relative success of invasive species. Biol Invasions, 13(1), 165-176. Doi: 10.1007/s10530-010-9799-3 Denisenko, N. V. (2011). Bryozoans of the East Siberian Sea: history of research and current knowledge of diversity. In P. N. W. Jackon & M. E. S. Jones (Eds.), Annals of Bryozoology 3: aspects of the history of research on bryozoans (pp. 1-15). Dublin, UK: International Bryozoology Association. Denisenko, N. V. & Kuklinski, P. (2008). Historical development of research and current state of bryozoans diversity in the Chukchi Sea. In P. N. W. Jackson (Ed.), Annals of Bryozoology 2: Aspects of the History of Research on Bryozoans (pp. 1-15). Dublin, UK: International Bryozoology Association. Dick, M. H., Grischenko, A. V., & Mawatari, S. F. (2005). Intertidal Bryozoa (Cheilostomata) of Ketchikan, Alaska. Journal of Natural History, 39(43), 3687-3784. Doi: 10.1080/00222930500415195 Dick, M. H., & Ross, J. R. P. (1985). Intertidal Cheilostome Bryozoans in Rock-Pile Habitat at Narrow Strait, Kodiak, Alaska, In C. Nielsen & G. P. Larwood (Eds.), Bryozoa: Ordovician to Recent : Papers Presented at the 6th International Conference on Bryozoa, Vienna 1983 (p.87-93). Fredensborg, DN: Olsen & Olsen. Eggleston, D. (1972). Patterns of reproduction in the marine Ectoprocta of the Isle of Man. Journal of Natural History, 6(1), 31-38. Doi: 10.1080/00222937200770041 Goldson, A. J., Hughes, R. N., & Gliddon, C. J. (2001). Population genetic consequences of larval dispersal mode and hydrography: a case study with bryozoans. Marine Biology, 138, 1037-1042. Doi: 10.1007/s002270000511 Gómez, A., Hughes, R. N., Wright, P. J., Carvalho, G., & Lunt, D. H. (2007). Mitochondrial DNA phylogeography and mating compatibility reveal marked genetic structuring and speciation in the NE Atlantic bryozoans Celleporella hyalina. Molecular Ecology, 16, 2173-2188. Doi: 10.1111/j.1365-294x.2007.03308.x Gordon, D. (2015). Celleporella hyalina (Linnaeus, 1767). Retrieved from http://marinespecies.org/aphia.php?p=taxdetails&id=111397. Gordon, D. P. (1967). A report on the ectoproct Polyzoa of some Auckland shores. TANE, 13, 43-76. Goss-Custard, S., Jones, J., Kitching, J.A. & Norton, T. A. (1979). Tide pools of Carrigathorna and Barloge Creek. Phil. Trans. R. Soc. B., 287(1016), 1-44. Doi: 10.1098/rstb.1979.0051 Grischenko, A. V. (2002). History of investigations and current state of knowledge of bryozoans species diversity in the Bring Sea. In P. N. W. Jackson & M. E. S. Jones (Eds.). Annals of Bryozoology: Aspects of the History of Research on Bryozoans (pp. 97-116). Dublin, UK: International Bryozoology Association. Grischenko, A. V., Dick, M. H., & Mawatari, S. F. (2007). Diversity and taxonomy of intertidal Bryozoa (Cheilostomata) at Akkeshi Bay, Hokkaido, Japan. Journal of Natural History, 41(17-20), 1047-1161. Doi: 10.1080/00222930701391773. Grischenko, A. V. & Zvyagintsev, A. Y. (2012). On the State of Inventory of the Bryozoan Fauna in Peter the Great Bay of the Sea of Japan in Light of Detection of the Cheilostome Bryozoans Callopora sarae and Microporella trigonellata. Russian Journal of Biological Invasions, 3(3), 172-179. Doi: 10.1134/S2075111712030034 Hastings, A. B. (1929). Cheilostomatous Polyzoa from the Vicinity of the Panama Canal collected by Dr. C. Crossland on the Cruise of the S. Y. ‘St. George’. Journal of Zoology, 99(4), 697-740. Doi: 10.1111/j.1096-3642.1929.tb01453.x Hayward, P. J. (1971). The marine fauna and flora of the Isles of Scilly Bryozoa and Entoprocta, Journal of Natural History, 5(5), 481-489. Doi: 10.1080/00222937100770341 Hayward PF & Ryland JS (1999) Cheilostomatous Bryozoa part 2. Hippothooidea - Celleporoidea. Synopses of the British Fauna (New Series). Barnes RSK & Crothers JH (eds.) No. 14 (Second Edition). The Linnean Society of London and The Estuarine and Coastal Sciences Association by Field Studies Council: 416pp. Hermansen, P., Larsen, P. S., & Riisgård, H. U. (2001). Colony growth rate of encrusting marine bryozoans (Electra pilosa and Celleporella hyalina). Journal of Experimental Marine Biology and Ecology, 263(1), 1-23. Doi: 10.1016/S0022-0981(01)00243-X Hewitt CL et al. (2004) Introduced and cryptogenic species in Port Philip Bay, Victoria, Australia. Marine Biology 144: 183-202. Hill, K. (2001) Smithsonian Marine Station at Fort Pierce, Retrieved from http://www.sms.si.edu/irlspec/Electr_bellul.htm. Hoare, K. & Hughes, R. N. (2001). Inbreeding and hermaphroditism in the sessile, brooding bryozoans Celleporella hyalina. Marine Biology, 139(1), 147-162. Doi: 10.1007/s002270100566 Hoare, K., Hughes, R. N., & Goldson, A. J. (1999). Molecular genetic evidence for the prevalence of outcrossing in the hermaphroditic brooding bryozoans Celleporella hyalina. Marine Ecology Progress Series, 188, 73-79. Doi: 10.3354/meps188073 Hughes, D. J. (1987a). Experimental study of growth and reproduction in marine bryozoans (Doctoral dissertation). Retrieved from ProQuest Dissertation and Theses (U018284). Hughes, D. J. (1987b). Gametogenesis and embryonic brooding in the cheilostome bryozoans Celleporella hyalina. J. Zool., London, 212(4), 691-711. Doi: 10.1111/j.1469-7998.1987.tb05965.x Hughes, D. J. (1989). Variation in Reproductive Strategy Among Clones of the Bryozoan Celleporella hyalina. Ecological Monographs, 59(4), 387-403. Doi: 10.2307/1943073 Hughes, D. J. (1992). Genotype-Environment Interactions and Relative Clonal Fitness in a Marine Bryozoan. Journal of Animal Ecology, 61(2), 291-306. Doi: 10.2307/5322 Hughes, D. J. & Hughes, R. N. (1986). Life history variation in Celleporella hyalina (Bryozoa). Proceedings of the Royal Society of London, Series B 228(1251), 127-132. Doi: 10.1098/rspb.1986.0046 Hughes, R. N., Manríquez, P. H., & Bishop, J. D. D. (2002). Female investment is retarded pending reception of allosperm in a hermaphroditic colonial invertebrate. Proceedings of the National Academy of Sciences of the United States of America, 99(23), 14884-14886. Doi: 10.1073/pnas.162339699 Hughes, R. N., Manríquez, P. H., Bishop, J. D. D., & Burrows, M. T. (2003). Stress promotes maleness in hermaphroditic modular animals. Proceedings of the National Academy of Sciences of the United States of America, 100(18), 10326-10330. Doi: 10.1073/pnas.1334011100 Hughes, R. N., Manríquez, P. H., Morley, S., Craig, S. F., & Bishop, J. D. D. (2004). Kin or self-recognition? Colonial fusibility of the bryozoans Celleporella hyalina. Evolution & Development, 6(6), 431-437. Doi: 10.1111/j.1525-142X.2004.04051.x Hughes. R. N., Gómez, A., Wright, P. J., Moyano, H. I., Cancino, J. M., Carvalho, G. R., & Lunt, D. H. (2008). Molecular phylogeny supports division of the ‘cosmopolitan’ taxon Celleporella (Bryozoa; Cheilostomata) into four major clades. Molecular Phylogenetics and Evolution, 46, 369-374. Doi: 10.1016/j.ympev.2007.08.014 Hughes, R. N., Wright, P. J., Carvalho, G. R., & Hutchinson, W. F. (2009). Patterns of self compatibility, inbreeding depression, outcrossing, and sex allocation in a marine byrozoan suggest the predominating influence of sperm competition. Biological Journal of the Linnean Society, 98, 519-531. Doi: 10.1111/j.1095-8312.2009.01312.x Hughes, R. N., Wright, P. J., Manriquez, P. H. (2002). Predominance of obligate outbreeding in the simultaneous hermaphrodite Celleporella hyalina sensu lato. Bryozoan Studies, 159-162 Hunter, E. (1991). Variation of growth and reproduction in a marine bryozoans (Doctoral dissertation). Retrieved from ProQuest Dissertations and Theses database (U065680). Hunter, E. & Hughes, R. N. (1993a). Effects of diet on life-history parameters of the marine bryozoans, Celleporella hyalina (L.). Journal of Experimental Marine Biology and Ecology, 167(2), 169-177. Doi: 10.1016/0022-0981(93)90029-N Hunter, E. & Hughes, R. N. (1993b). Self-fertilisation in Celleporella hyalina. Marine Biology, 115(3), 495-500. Doi: 10.1007/BF00349848 Hunter, E., & Hughes, R. N. (1994). The Influence of temperature, food ration and genotype on zooid size in Celleporella hyalina (L.). In P. J. Hayward (Ed.). Biology and Palaeobiology of Bryozoans, Proceedings of 9th International Bryozoology Conference (pp. 83-86). Swansea, UK: University of Wales. Hunter, E., Shimizu, K., & Fusetani, N. (1999). Role of protein in larval swimming and metamorphosis of Bugula neritina (Bryozoa: Cheilostomatida). Marine Biology, 133, 701-707 Inaba A (1988) Fauna and Flora of the Seto Inland Sea. Second edition II. Mukaishima Marine Biological Station of Hiroshima University: 1-475. (in Japanese) Ivin, V. V., & Zvyagintsev, A. Y. (2001). The fouling of the structures for algal mariculture. The Yellow Sea, 7(1), 61-69. Kaselowsky, J., Scholz, J., Mawatari, S. F., Probert, P. K., Gerdes, G., Kadagies, N., & Hillmer, G. (2005). Bryozoans and microbial communities of cool-temperate to subtropical latitudes- paleoecological implications. Facies, 50, 349-361. Doi: 10.1007/s10347-004-0034-5 Kashin, I. A., Bagaveeva, E. V., & Chaplygina. S. F. (2003). Fouling Communities of Hydrotechnical Constructions in Nakhodka Bay (Sea of Japan). Russian Journal of Marine Biology, 29(5), 267-283 Keegan, B. F., O’Connor, B. D. S., McGrath, D., Könnecker, G. & Ó Foighil, D. (1987). Littoral and benthic investigations on the south coast of Ireland II. The macrobenthic fauna off Carnsore Point. Proceedings of the Royal Irish Academy, 87B: 1-14. Kelmo, F., Attrill, M. J., Gomes, R. C. T., & Jones, M. B. (2004). El Niño induced local extinction of coral reef bryozoan species from Northern Bahia, Brazil. Biological Conservation, 118(5), 609-617 Koletić, N., Novosel, M., Rajević, N., & Franjević, D. (2015). Bryozoans are returning home: recolonization of freshwater ecosystems inferred from phylogenetic relationships. Ecology and Evolution, 5(2), 255-264. Doi: 10.1002/ece3.1352 Kuklinski, P., Gulliksen, B., Lөnne, O. J., & Weslawski, J. M. (2005). Composition of bryozoans assemblages related to depth in Svalbard fjords and sounds. Polar Biology, 28, 619-630. Doi: 10.1007/s00300-005-0726-5 Kuklinski, P., Gulliksen, B., Lөnne, O. J., & Weslawski, J. M. (2006). Substratum as a structuring influence on assemblages of Arctic bryozoans. Polar Biology, 29, 625-661. Doi: 10.1007/s00300-005-0102-5 Linnaeus, C. (1767). Systema naturae sive regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Laurentii Salvii, Holmiae. 12th ed. v. 1 (pt 2): 1286. Retrieved from http://www.biodiversitylibrary.org/item/83650#5 López Gappa, J.J. (1989). Overgrowth competition in an assemblage of encrusting bryozoans settled on artificial substrata. Marine Ecology Progress Series, 51(1), 121-130. Mandríquez, P. H. (2000). Mate choice and reproductive investment in the cheilostome bryozoans Celleporella hyalina (L. (BL) (Doctoral dissertation). Retrieved from ProQuest Dissertations and Theses database (U133674). Manríquez, P. H., Hughes, R. N., & Bishop, J. D. D. (2001). Age-dependent loss of fertility in water-borne sperm of the bryozoans Celleporella hyalina. Marine Ecology Progress Series, 224, 27-92. Doi: 10.3354/meps224087 Maturo, F. J. (1957). A study of the Bryozoa of Beaufort, North Carolina, and Vicinity. Journal of the Elisha Mitchell Scientific Society, 73, 11-68. Maughan, B. C. & Barnes, D. K. A. (2000). Seasonality of Competition in Early Development of Subtidal Encrusting Communities. Marine Ecology, 21(3-4), 205-220. Doi: 10.1046/j.1439-0485.2000.00703.x Mawatari, S. (1956). Cheilostomatous bryozoa from the Kurile Islands and the neighbouring districts. Pacific Science, 10(2), 113-135 Mawatari S (1976) Bryozoa (Ectoprocta). In: Animal systematics. Uchida T (ed.) Nakayama-shoten Co. Ltd., Tokyo: 35-229. (in Japanese) Mawatari, S. F., & Mawatari, S. (1981). A preliminary list of Cheilostomatous Bryozoans collected along the coast of Hokkaido. Proc. Jap. Soc. Syst. Zool., 21, 41-58 McCartney, M. A. (1997). Sex Allocation and Male Fitness Gain in a Colonial, Hermaphroditic Marine Invertebrate. Evolution, 51(1), 127-140. Doi: 10.2307/2410966 Navarrete Z., A., & Cancino, J. M. (2005). Morphological differentiation in the Celleporella hyalina (Linnaeus, 1767) complex (Bryozoa: Cheilostomata) along the Chilean coast. Bryozoan Studies 2004. Doi: 10.1201/9780203970799.ch20 Namikawa, H., Mawatari, S. F., & Calder, D. R. (1992). Role of the tentaculozooids of the polymorphic hydroid Stylactaria conchicola (Yamada) in interactions with some epifaunal space competitors. J. Exp. Mar. Biol. Ecol., 162(1), 65-75. Doi: 10.1016/0022-0981(92)90125-T Namikawa, H., Mawatari, S. F., & Calder, D. R. (1993). Reproduction, planula development, and substratum selection in three species of Stylactaria (Cnidaria: Hydrozoa) from Hokkaido, Japan. Journal of Natural History, 27(3), 521-533. Doi: 10.1080/00222939300770291 O’Connor, R. J., Seed, R., & Boaden, P. J. S. (1980). Resource space partitioning by the Bryozoa of a Fucus serratus L. community. Journal of Experimental Marine Biology and Ecology, 45(2), 117-137. Doi: 10.1016/0022-0981(80)90055-6 OBIS. (2015). Ocean Biogeographic Information System. Retrieved from http://iobis.org/mapper Oregon State University. (1971). Oceanography of the Nearshore Coastal Waters of the Pacific Northwest Relating to Possible Pollution. Retrieved from http://nepis.epa.gov/Exe/ZyPDF.cgi/9100GCEO.PDF?Dockey=9100GCEO.PDF Osburn RC (1952) Bryozoa of the Pacific coast. Part 2, Cheilostomata-Ascophora. The University of Southern California Publication. Allan Hancock Pacific Expedition 14: 271-611. Ostrovsky, A. N. (1998). Comparative studies of ovicell anatomy and reproductive patterns in Cribrilina annulata and Celleporella hyalina (Bryozoa: Cheilostomatida). Acta Zoologica, 79(4), 287-318 Ostrovsky, A. N. (2013). Evolution of Sexual Reproduction in Marine Invertebrates – Example of gymnolaemate bryozoans. Dordrectht: Springer Netherlands. Doi: 10.1007/978-94-007-7146-8 Ostrovsky, A. N., Gordon, D. P., & Lidgard, S. (2009). Independent evolution of matrotrophy in the major classes of Bryozoa: transitions among reproductive patterns and their ecological background. Marine Ecology Progress Series, 378, 113-124. Doi: 10.3351/meps07850 Pemberton, A. J., Hughes, R. N., Manríquez, P. H., & Bishop J. D. (2003). Efficient utilization of very dilute aquatic sperm: sperm competition may be more likely than sperm limitation when eggs are retained. Proceedings of the Royal Society of London B: Biological Sciences, 270(Suppl 2), S223-S226. Doi: 10.1098/rsbl.2003.0076 Pistevos, J.C.A., Calosi, P., Widdicombe, S., & Bishop, J. D. D. (2011). Will variation among genetic individuals influence species responses to global climate change?. Oikos, 120(5), 675-689. Doi: 10.1111/j.1600-0706.2010.19470.x Porter, J. S., Jones, M. E. S., Kuklinski, P., & Rouse, S. (2015) First records of marine invasive non-native Bryozoa in Norwegian coastal waters from Bergen to Trondheim. BioInvasions Records, 4 (in press) Reverter Gil, O. (1994). Briozoos de la Ria del Ferrol (Doctoral dissertation). Retrieved from ProQuest Dissertations and Theses database (C484009). Rogick, M. D., & Croasdale, H. (1949). Studies on Marine Bryozoa, III. Woods Hole Region Bryozoa Associated with Algae. Biological Bulletin, 96(1), 32-69 Rouse, S. (2011). Aetea anguina. Bryozoa of the British Isles. Retrieved from http://britishbryozoans.myspecies.info/content/aetea-anguina-linnaeus-1758 Ruppert, E.E., Fox, R.S., & Barnes, R.D. (2004). Invertebrate Zoology: A functional evolutionary approach. Ann Arbor, MN: Thomson Brooks/Cole. Ryland, J. S. (1959). Experiments on the Selection of Algal Substrates by Polyzoan Larvae. Journal of Experimental Biology, 36(4), 613-631. Ryland, J. S. (1960). Experiments on the influence of light on the behaviour of polyzoan larvae. Journal of Experimental Biology, 37(4), 783-800. Ryland, J. S. (1962a). The Association Between Polyzoa and Algal Substrata. Journal of Animal Ecology, 31(2), 331-338. Doi: 10.2307/2145 Ryland, J. S. (1962b). Biology and identification of intertidal Polyzoa. Field Studies, 1(4), 33-51. Seed, R. & O’Connor, R. J. (1981). Community Organization in Marine Algal Epifaunas. Annual Review of Ecology and Systematics, 12, 49-74. Soule, J. D. (1961). Results of the Puritan-American Museum of Natural History Expedition to Western Mexico. 13. Ascophoran Cheilostomata (Bryozoa) of the Gulf of California. American Museum Novitates, 2053, 1-66 Temkin, M. H. (1991). Fertilization in the Gymnolaemate Bryozoa (Doctoral dissertation). Retrieved from ProQuest Dissertations and Theses database. (DP23819). Thresher RE (1999) The physical environment of Port Philip Bay. In: Marine biological invasions of Port Philip Bay, Victoria. Hewitt CL, Campbell ML, Thresher RE, Martin RB (eds.). CRIMP Technical Report 20: 25-31. Tilbrook KJ (2012) Cheilostomata: first records of two invasive species in Australia and the northerly range extension for a third. Check List 8: 181-183. http://www.checklist.org.br/getpdf?NGD192-11 Trott, T. J. (2004). Cobscook Bay Inventory: A Historical Checklist of Marine Invertebrates Spanning 162 Years. Northeastern Naturalist, 11(2), 261-324. Waeschenbach, A., Porter, J. S., & Hughes, R. N. (2012). Molecular variability in the Celleporella hyalina (Bryozoa; Cheilostomata) species complex: evidence for cryptic speciation from complete mitochondrial genomes. Molecular Biology Reports, 39(9), 8601-8614. Doi: 10.1007/s11033-012-1714-9 Watts, P.C., Thorpe, J. P., & Taylor, P. D. (1998). Natural and anthropogenic dispersal mechanisms in the marine environment: a study using cheilostome Bryozoa. Philos Trans R Soc Lond B Biol Sci, 353(1367), 453-464. Doi: 10.1098/rstb.1998.0222 Winston JE (1977). Distribution and ecology of estuarine ectoprocts: A critical review. Chesapeake Science, 18:, 34‐57. doi:10.2307/1350363. https://fau.digital.flvc.org/islandora/object/fau%3A6214/datastream/OBJ/view/Distribution_and_ecology_of_estuarine_ectoprocts__A_critical_review.pdf Włodarska-Kowalczuk, M., Kukliński, P., Ronowicz, M., Legeżyńska, J., & Gromisz, S. (2009). Assessing species richness of macrofauna associated with macroalgae in Arctic kelp forests (Hornsund, Svalbard). Polar Biology, 32, 897-905. Doi: 10.1007/s00300-009-0590-9 Woollacott, R. M., & Zimmer, R. L. (Eds.). (1977). Biology of Bryozoans. New York, NY: Academic Press Wright, P. J. (2004). Reproductive compatibility and speciation in Celleporella hyalina sensu lato (Doctoral dissertation). Retrieved from ProQuest Dissertations and Theses database. (U200630). Yund, P. O. & McCartney, M. A. (1994). Male Reproductive Success in Sessile Invertebrates: Competition for Fertilizations. Ecology, 75(8), 2151-2167. Doi: 10.2307/1940874.

Literature:

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

Notes:

Celleporella hyalina is a cryptic species complex composed of numerous genetic, reproductively isolated lineages that occupy mostly allopatric regions around the globe, yet are morphologically very similar (Gómez et al. 2006, cited in Gómez et al. 2007) Called a pioneer species (Kulkinski et al 2006; Maughan et al 2000)