Membranipora serrilamella


Scientific Name: Membranipora serrilamella

Phylum: Bryozoa

Class: Gymnolaemata

Order: Cheilostomatida

Family: Membraniporidae

Genus: Membranipora


serrilamella *unaccepted status on WoRMS, accepted as Membranipora villosa(Bock 2015) [Describe here as A. iricolor]

Native Distribution

Origin Realm:

Temperate Northern Pacific, Temperate Northern Atlantic, Central Indo-Pacific

Native Region:

Origin Location:

Temperate Northern Pacific Virago Sound (Hincks 1882, cited in Osburn 1950) and numerous localities in British Columbia (O'Donoghue 1923, 1926, cited in Osburn) .STATUS NOT STATED From Puget Sound to California. (Robertson 1908, cited in Osburn 1950) STATUS NOT STATED [Canada] Strait of Georgia, BC; Virago Sound, BC (Marliave et al 2011; Oregon State University 1971) STATUS NOT STATED [US] Santa Barbara, California; Monterey, California; Ketchikan, Alaska; Morro Bay, California; Puget Sound, Washington (Arkema 2009; Bram et al 2005; Busch 1981; Dick et al 2005; Koenigs et al 2015; Needles and Wendt 2013; Oregon State University 1971; Reed et al 2015; Schoener and Schoener 1981; Wood and Hageman 1982; Yorke et al 2013) STATUS STATED [Japan] southern Hokkaido; southward to Wakayama Prefecture on the Pacific side; Ishikawa Prefecture on the Sea of Japan; Honshu; Paramushir; Akkeshi; Kushiro; Uchiura Bay; Mori; Muroran; Shirikishinai (Mawatari and Mawatari 1981; Mawatari 1974, cited in Dick et al 2005; Mawatari 1956; Mawatari 1973; Mawatari and Ito 1972) STATUS NOT STATED [Korea] Anhung; Wangdolcho, Uljin; Taedo Island, Tongyeong; Yujado Island, Tongyeong; Yellow Sea (Rho and Seo 1990; Seo 1996; Seo and Min 2009) STATUS NOT STATED [Kurile Island] Paramushir Island. (Mawatari 1956) STATUS NOT STATED [Korea] Wangdolcho, Uljin; Taedo Island and Yujiado Island,Tongyeong. (Seo & Min 2009) STATUS NOT STATED [Japan] Akkeshi, Kushiro (Mawatari 1956), Uchiura Bay (Hunka Bay), Hokkaido. (Mawatari 1973) STATUS NOT STATED [Japan] Kominato, Boso Peninsula, Chiba Prefecture; Shirahama, Wakayama Prefecture; Noto Peniusula. (Mawatari 1974) STATUS NOT STATED [Japan] [[as M. serrata (synonymized taxon)] Shimoda, Izu Peninsula and several localities from Onagawa Bay to Kagoshima Bay. (Okada 1934) STATUS NOT STATED [Japan] Seto Inland Sea. (Inaba 2988) STATUS NOT STATED [Membranipora villosa (synonymised species)] [Canada] Departure Bay, BC (O'Donoghue 1926) STATUS NOT STATED [Japan] [as Conopeum serrata (synonymized taxon)] Mutsu Bay. (Okada 1929) STATUS NOT STATED [Japan] [as Conopeum serrata (synonymized taxon)] Onagawa, Miyagi Prefecture. (Okada & Mawatari 1937) STATUS NOT STATED Central Indo-Pacific [China] Hong Kong. (Huang & Li 1990) STATUS NOT STATED Temperate Northern Atlantic [Croatia] Adriatic and Danube basin (Koletic et al 2015) STATUS NOT STATED Uncertain realm [Brazil] Several localities. (Marcus 1937, cited in Osburn 1950) STATUS NOT STATED

Geographic Range:

Cosmopolitan genus (Arkema 2009) [Western Pacific] Japan and Korea (Rho and Seo 1990; Mawatari and Mawatari 1981; Seo and Min 2009) [Eastern Pacific] Alaska to southern California (Harbo 2011; Carlton 2007; Oregon State University 1971; Rho and Seo 1990) [Western Atlantic] US coast (Mawatari 1956) [Eastern Atlantic] Croatia (Koletic et al 2015) [Japan] 31°N-42°N both at Pacific side and Japan Sea side. (Inaba 1988)

General Diversity:


Non-native Distribution

Invasion History:

No records of invasion (Global Invasive Species Database 2016)

Non-native Region:

Not applicable

Invasion Propens:

Not applicable

Status Date Non-native:

Not applicable

Vectors and Spread

Initial Vector:


Second Vector:


Vector Details:


Spread Rate:


Date First Observed in Japan:

Specimens collected from Paramushir since 1931 (Mawatari 1956)

Date First Observed on West coast North America:



Impact in Japan:

Kelp fouling, damage to production of Laminaria japonica, economically important edible algae (Ito and Mawatari 1972;Torii and Masuda 1972, cited in Ivin and Zvyagintsev 2001)

Global Impact:



Native Temperature Regime:

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

Native Temperature Range:

[US] 13.1 to over 18˚C (Arkema 2009) [US] 12.6-16.2°C surface temperature averages at tidal stations (Busch 1981) Muroran in Uchiura (Hunka) Bay: max 18.0ºC in summer and min 2.0ºC in winter. (Clark et al. 2003) Hong Kong: max 28.5ºC in summer and min 18.1ºC in winter. (Clark et al. 2003) Cold water, Cool temperate, Mild temperate, Warm temperate, Subtropical, Tropical (M. Otani, pers. comm.) [Membranipora villosa (synonymised species)] [US] Prefer slightly lower temperature at 2-9˚C (Soule and Soule 2012)

Non-native Temperature Regime:

Not applicable

Non-native Temperature Range:

Not applicable

Native Salinity Regime:

Mesohaline, Polyhaline, Euhaline

Native Salinity Range:

[US] 33.4 - 33.8 %o (Busch 1981) Muroran in Uchiura (Hunka Bay): max 32.0psu in dry period and min 23.0psu in wet period. (Clark et al. 2003) Hong Kong: max 34.0psu in dry period and min 10.0psu in wet period. (Clark et al. 2003) RELATED: [Cribrilina, Membranipora, Hippoporina, Schizoporella, Smittina and related genera] Supported by polyhaline water. Although Membranipora is a euryhaline genus, it is found in mesohaline water, namely 3-16%o (Hayami 1975) [Membranipora] Although Membranipora is a euryhaline genus, it is found in mesohaline water, namely 3-16psu (Borg 1931, cited in Hayami 1975). In water of more than 40psu, Mempranipora has been reported living in the Suez Canal. (Cheetham 1963, cited in Hayami 1975)

Non-native Salinity Regime:

Not applicable

Temperature Regime Survival:

Cool temperate, See details

Temperature Range Survival:

Growth rates experiences greatest increases in colony size where average temperatures were the coolest and current speeds 10-12cm/s (Arkema 2009) RELATED: [Membranipora spp.] -1.401 - 24.233ºC (OBIS 2016)

Temperature Regime Reproduction:

Mild temperate

Temperature Range Reproduction:

According to Mawatari & Ito (1972), it seems that reproduction occurs during March and August in Uchiura (Hunka) Bay of the mild temperature regime.

Salinity Regime Survival:

See details

Salinity Range Survival:

RELATED: [Membranipora spp.] 7.644 - 36.231 PPS (OBIS 2016)

Salintiy Regime Reproduction:

Polyhaline, Euhaline

Salinity Range Reproduction:


Depth Regime:

Lower intertidal, Shallow subtidal, Deep subtidal

Depth Range:

Shallow waters (Harbo 2011) Shallow water to 40m, perhaps to bottom of photic zone (Carlton 2007) Commonly occurs at 4-6m below surface (Arkema 2009) On algae and plant, directly accessible low in the intertidal (Dick et al 2005) 0-3m depth (Koenigs et al 2015) On floating fronds and down to 10 fms (about 18m) (Oregon State University 1971) 15m in depth (Seo and Min 2009) Seto Inland Sea: lower intertidal to 20-30m. (Inaba 1988) Dredged on a few occasions down to 10 fms. (Osburn 1950) It is assumed the depth is several meters because colony was collected from test panels attached to the experimental raft. (Huang & Li 1990)

Non-native Salinity Range:

Native Abundance:

Common, Abundant


Fertilization Mode:

See details

Reproduction Mode:


Spawning Type:


Development Mode:

Planktotrophic planktonic larva (feeding)

Asexual Reproduction:

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

Reproduction Details:

In hermaphrodite zooids of Membranipora serrilamella, ovaries are always formed later than testes, with male and female reproductive phases either somewhat overlapping or separated by a time gap (Hagerman 1983, cited in Ostrovsky 2013) In mature colonies, there is a peripheral zone of young zooids without gonads, a subperipheral zone of zooids in which testes are developing (situated among sterile ones) and a more inner zone (or several belt zones) consisting of simultaneously hermaphrodite (and sterile) zooids; finally the colony centre is represented by the oldest zooids with degenerated polypides without gonads (Ostrovsky 2013) Broadcaster (Mawatari 1975; Mawatari and Mawatari 1975; Hagerman 1981, cited in Ostrovsky 2013; Ostrovsky and Porter 2011) During growth phase, the oocyte increases in volume ca. 6000-8000-fold. Towards the end of the vitellogenic phase, oocytes lose contact with the subovarian space and move into the centro-apical ovulatory zone where they are partially exposed to the zooidal cavity and their oolemma forms numerous microvilli (Ostrovsky 2013) Mature primary oocytes are shaped as a biconcave elongated disc 85.8-101μm (Ostrovsky 2013) Fertilization in malacostegans occurs during or shortly after ovulation. Sperm enter the coelorm of fertile zooids via the intertentacular organ, through which eggs are also spawned (Temkin 1994 and 1996, cited in Ostrovsky 2013) Ovaries are always formed later than testes, with male and female reproductive phases either somewhat overlapping or separated by a time gap (Hageman 1983, cited in Ostrovsky 2013) Fertilization in malacostegans occurs during or shortly after ovulation; Sperm enter the coelom of fertile zooids via the intertentacular organ, through which eggs are also spawned (Temkin 1994, 1996 cited in Ostrovsky 2013) Hermaphroditism is the usual mode of reproduction in M. s. (Mawatari 1975) RELATED: [Membranipora] Studies on Membranipora membranacea shows that it has planktotrophic developmental pattern (Temkin 1991) [Membraniporidea] Shed numerous small eggs directlyto the sea (Hayward & Ryland 1998) and fetilize in the sea. (Mawatari 1976) These develop into shelled, planktorophic larvae, termed cyphonautes, which feed and grow during several weeks or months spent in the plankton. (Hayward & Ryland 1998) [Membraniporidea] Fertilization of M. s. may take place externally. (Mawatari 1975) [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] Non-brooding bryozoans feed during the larval stage, while the larvae of brooding bryozoans do not, since these larvae tend to settle soon after release (Hill 2001) [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 1998) [Bryozoa] Reproduces asexually by budding. (Mawatari 1976) CONFLICT For many Gymnolaemate bryozoa, eggs are not activated when fused with sperm at internal site. Instead, egg must be broadcasted before egg activation takes place. In this way, Membranipora spp. is described as being broadcasters despite fertilization being internal process (Temkin 1991; Ostrovsky 2013)

Adult Mobility:


Adult Mobility Details:

Sessile invertebrate relying on currents to feed (Arkema 2009) RELATED: [Bryozoa] The abundance and taxonomic diversity of benthic bryozoan faunas are directly related to substratum. (Hayward & Ryland 1998) [Bryozoa] Bryozoan colonies are sessile (Hayami 1975) [Bryozoa] Bryozoans are a phylum of sessile, colonial suspension feeders found throughout the world in both marine and freshwater environments. (Tilbrook 2012)

Maturity Size:

Thin crusts to 1mm high; more than 7.5cm diameter (Harbo 2011) Ranging in size from small, circular, young colonies to large expanses of coalesced colonies on Laminarian algae (Dick et al 2005) Largest specimen (colony) was about 25cm in diameter (Mawatari 1973) Zooids: 0.53-1.20mm long by 0.23-0.41mm wide (Dick et al 2005)

Maturity Age:


Reproduction Lifespan:




Broods per Year:


Reproduction Cues:

Discharge of ova of M. s. is induced by light. (Mawatari 1975) 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 1998) [Bryozoa] In various degrees of intensity according to the species temperature also stimulates sexual reproduction. (Winston 1977)

Reproduction Time:

Many settled larvae found especially in May and August at Mori, Japan (Mawatari 1973) Cyponautes larvae were obtained at least until August since March in 1971. (Mawatari & Ito 1972) No particular season during which Cyphonautes are discharged for they have been found and collected from early settled colonies from beginning of May to middle of September (O'Donoghue 1926) [Membranipora villosa (synonymised species)] Appeared more plentiful in morning (10-noon than earlier or in the afternoon) (O'Donoghue 1926)


The number of ovulated oocytes formed by a single fertile zooid may be, according to different sources, from 20-25 to 40 or more (Ostrovsky 2013) When ova are closely arranged under the frontal membrane, more than 40 ova are simutaneously observed in one and the same zooid. (Mawatari 1975)

Egg Size:

Oocytes with a diameter of 85.8-101 µm and 100 µm (Ostrovsky 2013) Diameter of unfertilized ovum is about 100μm. (Mawatari 1975) RELATED: [Gymnolaemata] About 200µm (Woollacott and Zimmer 1977)

Egg Duration:

About one day after the discharge of ovum, a hatched larva begins to swim. (Mawatari 1975)

Early Life Growth Rate:

Intertentacular organ develops at the onset of oogenesis, whole process lasts about 2 days (Ostrovsky and Porter 2011) Oocyte increases in volume 6000-8000 fold (Ostrovsky 2013) RELATED: [Gymnolaemata] Two phases of larvae metamorphosis: first stage about 20mins; second stage 1-6 days (Woollacott and Zimmer 1977) [Membranipora villosa (synonymised species)] Loss of Cyphonautes shell occurs between 40-60hrs after fixation; second stage of development where 4 or 5 incipient partitions appear takes 44-50 hours after fixation; next 48hrs is mostly growth of size of the zoarium (O'Donoghue 1926)

Adult Growth Rate:

Growth rates experiences greatest increases in colony size where average temperatures were the coolest and current speeds 10-12cm/s (Arkema 2009)

Population Growth Rate:

Colonies with a diameter of 1.2-6.6mm increment their diameter 0.2-2.4mm in three days in August. (Mawatari 1975)

Population Variablity:

Shortly before the metamorphosis is completed, formation of the first daughter zooid subsequently occurs. And then peripheral buds begin to grow successively one after another from the anteriormost pair to posterior ones. If these peripheral buds would rapidly grow, a colony soon becomes disk-shaped. (Mawatari 1973)



Kelp forest, Macroalgal beds, Rocky intertidal, Rocky subtidal, Fouling

Habitat Type:

Epibenthic, Epiphytic, Epizoic


Rock, Biogenic, Artificial substrate


Exposed, Semi-exposed, Protected

Habitat Expansion:


Habitat Details:

On kelps and algae (Harbo 2011) Common on algae; blades of Macrocystis and Laminaria (Carlton 2007; Busch 1981; Koenigs et al 2015; Mawatari 1973; Reed et al 2015; Yorke et al 2013; Yoshioka 1982) Occurs commonly in kelp forests, grows on kelp blades (Arkema 2009) Offshore oil platform (Bram et al 2005) Laminarian algae, including plants (Dick et al 2005) Floating fronds of kelp (Mawatari 1956; Oregon State University 1971) Rocky shores, on Laminarian blades (Millen 1987) Encrusting on broader shells (Rho and Seo 1990) Attached to fouling panels, made of white formica undersurface (Schoener and Schoener 1981) Excessively abundant on the floating fronds of kelp. (Osburn 1950) M. s. was found on the frond of a brown alga, Undaria pinnatifida and Laminaria japonica at Uchiura (Hunka) Bay, Hokkaido. (Mawatari 1973) A large colony of M. serrata (Synonymized taxon) attached to a seaweed at Shimoda, Izu Peninsula. (Okada 1934) Colony was collected from test panels attached to the experimental raft. (Huang & Li 1990) M. s. probably posesses strong preference for the laminarians, and the colonies were exclusively found on the fronds of Laminariaceae, namely, Undaria pinnatifida, Costaria costata, Alaria crassiforia, Laminaria japonica and Laminaria spp. (Mawatari 1975) [Conopeum serrata (Synonymized taxon)] Attached to Ulva. (Okada 1929)

Trophic Level:

Suspension feeder

Trophic Details:

Food of M. s. is assumed to be particulate organic matter and phytoplankton. (Macdonald et al. 2010) 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 1998) [Cheilostomata] Main food is diatom, protozoans and etc. and unappropriate sized particles are ejected (Mawatari 1976)

Forage Mode:


Forage Details:

Feeding success highest at 10-12cm/s current flow speeds when compared to 0-5cm/s and >20cm/s speeds (Arkema 2009) Feeds in a zone characterized by vertical gradients in water flow and particle concentration, often referred to as the boundary layer (Denny 1988, cited in Arkema 2009) Food of M. s. is assumed to be particulate organic matter and phytoplankton. (Macdonald et al. 2010) 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 1998) [Cheilostomata] Main food is diatom, protozoans and etc. and unappropriate sized particles are ejected (Mawatari 1976)

Natural Control:

PREDATION [Predation] Juvenile molluscan predation (Carlton 2007) [Predation] Nudibranchs (Dick et al 2005; McDonald and Nybakken 1997; Millen 1987) [Membranipora villosa (synonymised species)] [Predation] Attack by Corambe sp. (O'Donoghue 1926) COMPETITION [Competition] Competition for space (Seed and O'Connor 1981) PARASITES [Membranipora villosa (synonymised species)] [Parasites] If acute, death of one polyp and in consequence, the discontinuance of this line of growth; may be caused by tiny Ciliates or Nemathelminthes (O'Donoghue 1926) 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


Arkema, K. K. (2009). Flow-mediated feeding in the field: consequences for the performance and abundance of a sessile marine invertebrate. Marine Ecology Progress Series, 388, 207-220. Doi: 10.3354/meps08140 Bock, P. (2015). Membranipora serrilamella Osburn, 1950. In: P. Bock, & D. Gordon (Eds.). World List of Bryozoa. Retrieved from Bram, J. B., Page, H. M., & Dugan J. E. (2005). Spatial and temporal variability in early successional patterns of an invertebrate assemblage at an offshore oil platform. Journal of Experimental Marine Biology and Ecology, 317(2), 223-237. Doi: 10.1016/j.jembe.2004.12.003 Busch, S. J. (1981). Ecology and distribution of the benthic community on the Monterey breakwater, Monterey, California (Doctoral Dissertation). Retrieved from Naval Postgraduate School Dudley Knox Library database. (T199253). Carlton, J. T. (Ed.). (2007). The Light and Smith Manual: intertidal invertebrates from central California to Oregon. Los Angeles, CA: Univ of California Press. Clarke C, Hillard R, Junqueira AOR, Neto ACL, Polglaze J, Raaymakers S (2003) Ballast water risk assessment, Port of Sepetiba, Fedral Republic of Brazil. GloBallast Monograph Series 14: 1-63 + 7 Appendices. 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 Feder, H. M., Turner, C. H., & Limbaugh, C. (1974). Observations on Fishes Associated with Kelp Beds in Southern California. Retrieved from State of California Department of Fish and Game website: Global Invasive Species Database. Access Date: 16-Mar-2016. Gontar, V. I. (1981). On the Cheilostomata (Bryozoa) of the Kurile Islands. In G. P. Larwood, & C. Nielsen (Eds.), Recent and Fossil Bryozoa: Papers Presented at the 5th International Conference on Bryozoa, Durham, 1980 (pp.101-103). Fredensborg: Olsen&Olsen. Harbo, R. M. (2011). Whelks to Whales: Coastal Marine life of the Pacific Northwest. Madeira Park, BC: Harbour Publishing Co. Ltd. Hayami, T. (1975). Neogene Bryozoa from northern Japan. Tohoku Univ., Sci. Rep., 2nd ser. (Geol.), 45(2), 83-126. Hayward PF & Ryland JS (1998) Cheilostomatous Bryozoa part I. Aeteoidea - Cribrilinoidea. Synopses of the British Fauna (New Series). Barnes RSK & Crothers JH (eds.) No. 10 (Second Edition). The Linnean Society of London and The Estuarine and Coastal Sciences Association by Field Studies Council: 366pp. Hill, K. (2001) Smithsonian Marine Station at Fort Pierce. Retrieved from Huang Z & Li C (1990) The bryozoan foulers of Hong Kong and neighbouring waters. Proceedings of the Second International Marine Biological Workshop: The Marine flora and Fauna of Hong Kong and Southern China, Hong Kong, 1986. Morton B (ed.). Hong Kong University Press: 737-781. 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. Koenig, C., Miller, R. J., & Page, H. M. (2015). Top predators rely on carbon derived from giant kelp Macrocystis pyrifera. Marine Ecology Progress Series, 537, 1-8. Doi: 10.3354/meps11467 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 Macdonald TA, Burd BJ, Macdonald VI, van Roodselaar A (2010) Taxonomic and Feeding Guild Classification for the Marine Benthic Macroinvertebrates of the Strait of Georgia, British Columbia. Canadian Technical Report of Fisheries and Aquatic Sciences 2874: 1-63pp. Marliave, J. B., Gibbs, C. J., Gibbs, D. M., Lamb, A. O., & Young, S. J. F. (2011). Biodiversity stability of shallow marine benthos in Strait of Georgia, British Columbia, Canada through climate regimes, overfishing and ocean acidification. In O. Grillo, & G. Venora (Eds.), Biodiversity Loss in a Changing Planet (pp. 49-74). Rijeka, Croatia: InTech, Mawatari, S. (1956). Cheilostomatous bryozoa from the Kurile Islands and the neighbouring districts. Pacific Science, 10(2), 113-135 Mawatari, S. F. (1973). The post-larval development of Membranipora serrilamella Osburn (Bryozoa, Cheilostomata). Proc. Jap. Soc. Syst. Zool., 9, 45-54 Mawatari S (1974) Studies on Japanese anascan Bryozoa. 3. Division Malacostega (1). Bulletin of the National Science Museum 17: 17-52. Mawatari SF (1975) The life history of Membranipora serrilamella Osburn (Bryozoa, Cheilostomata). Bulletin of the Liberal Arts and Science Course. Nihon University School of Medicine 3: 19-57. Mawatari S (1976) Bryozoa (Ectoprocta). In: Animal systematics. Uchida T (ed.) Nakayama-shoten Co. Ltd., Tokyo: 35-229. (in Japanese) Mawatari, S. F., & Itô, T. (1972). The Morphology of Cyphonautes Larva of Membranipora serrilamella Osburn from Hokkaido. Journal of the Faculty of Science Hokkaido University Series VI. Zoology, 18(3), 400-405. 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 McDonald, G. R., & Nybakken, J. W. (1997). List of the Worldwide Food Habits of Nudibranchs. Veliger, 40(2), 4-426 Millen, S. V. (1987). The nudibranch genus Adalaria, with a description of a new species from the northeastern Pacific. Canadian Journal of Zoology, 65(11), 2696-2702. Doi: 10.1139/z87-409 Needles, L. A., & Wendt, D. E. (2013). Big changes to a small bay: introduced species and long-term compositional shifts to the fouling community of Morro Bay (CA). Biol Invasions, 15, 1231-1251. Doi: 10.1007/s10530-012-0362-2 Nielsen, C., & Worsaae, K. (2010). Structure and occurrence of cyphonautes larvae (Bryozoa, Ectoprocta). Journal of Morphology, 271(9), 1094-1109. Doi: 10.1002/jmor.10856. OBIS. Ocean Biogeographic Information System. Access date: 15-09-2016 *Note: genus level data Okada Y (1929) Report of the biological survey of Mutsu Bay. 12. Cheilostomatous Bryozoa of Mutsu Bay. Science Report of the Tohoku Imperial University, Ser. 4, 4: 11-35. Okada Y (1934) Bryozoa fauna in the vicinity of the Shimoda Marine Biological Station. Science Reports of the Tokyo Bunrika Daigaku 26: 1-20. Okada Y & Mawatari S (1937) On the collection of Bryozoa along the coast of Onagawa Bay and its vicinity, the northern part of Honshu, Japan. Science Reports of the Tohoku Imperial University, Ser. 4 (Biology) 3: 433-445. Osburn RC (1950) Bryozoa of the Pacific coast of America. Part 1. Cheilostomata - Anasca. The University of Southern California Publication. Allan Hancock Pacific Expedition 14: 1-269. 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., & Porter, J. S. (2011). Pattern of occurrence of supraneural coelomopores and intertentacular organs in Gymnolaemata (Bryozoa) and its evolutionary implications. Zoomorphology, 130(1), 1-15. Doi: 10.1007/s00435-011-0122-3 Reed, D. C., Carlson, C. A., Halewood, E. R., Nelson, J. C., Harrer, S. L., Rassweiler, A., & Miller, R. J. (2015). Patterns and controls of reef-scale production of dissolved organic carbon by giant kelp Macrocystis pyrifera. Limnology and Oceanography, 60(6), 1996-2008. Doi: 10.1002/Ino.10154 Rho, B. J., & Seo, J. E. (1990). A systematic study on the marine bryozoans in Korea 7. Suborder Anasca. The Korean Society of Systematic Zoology, 6(1), 145-159 Rouse, S. (2011). Aetea anguina. Bryozoa of the British Isles. Retrieved from Ruppert, E.E., Fox, R.S., and Barnes, R.D. (2004). Invertebrate Zoology: A functional evolutionary approach. Ann Arbor, MN: Thomson Brooks/Cole. Schoener, A., & Schoener, T. W. (1981). The Dynamics of the Species-Area Relation in Marine Fouling Systems: 1. Biological Correlates of Changes in the Species-Area Slope. The American Naturalist, 118(3), 339-360 Seed, R., & O’Connor, R. J. (1981). Community organization in marine algal epifaunas. Annual Review of Ecology and Systematics, 12, 49-74 Seo, J. E. (1996). Two new species of Membraniporoidea (Bryozoa: Cheilostomata) from Korea. Animal Systematics, Evolution and Diversity, 12(1), 45-51 Seo, J. E., & Min, B. S. (2009). A faunistic study on cheilostomatous bryozoans from the shoreline of South Korea, with two new species. Korean J. Syst. Zool., 25(1), 19-40. Temkin, M. H. (1991). Fertilization in the Gymnolaemate Bryozoa (Doctoral dissertation). Retrieved from ProQuest Dissertations and Theses database. (DP23819). Temkin, M. H. (1996). Comparative fertilization biology of gymnolaemate bryozoans. Marine Biology, 127(2), 329-339 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. Winston JE (1977). Distribution and ecology of estuarine ectoprocts: A critical review. Chesapeake Science, 18: 34‐57. doi:10.2307/1350363. Wood, R. L., & Hageman, G. S. (1982). The fine structure of cellular junctions in a marine bryozoans: gap junctions. Journal of Ultrastructure Research, 79(2), 174-188. Doi: 10.1016/S0022-5320(82)90029-6 Woollacott, R. M., & Zimmer, R. L. (Eds.). (1977). Biology of Bryozoans. New York, NY: Academic Press Yorke, C. E., Miller, R. J., Page, H. M., & Reed, D. C. (2013). Importance of kelp detritus as a component of suspended particulate organic matter in giant kelp Macrocystis pyrifera forests. Marine Ecology Progress Series, 493, 113-125. Doi: 10.3354/meps10502 Yoshioka, P. M. (1982). Predator-induced polymorphism in the bryozoan Membranipora membranacea (L.). J. Exp. Mar. Biol. Ecol., 61(3), 233-242. Doi: 10.1016/0022-0981(82)90071-5.


Substantial scientific information; non-peer-reviewed information; data specific to the region; supported by recent data (within the last 10 years) or research


Yoshioka (1982) concluded that M. villosa and M. serrilamella are synonyms of M. membranacea. However, Soule et al. (1995) retained the name M. serrilamella , arguing that it is morphologically distinct from Atlantic M. membranacea; furthermore, they did not include M. villosa in their synonymy of M. serrilamella and provided separate illustrations of the two (Dick et al 2005)