Smittoidea prolifica

Overview

Scientific Name: Smittoidea prolifica

Phylum: Bryozoa

Class: Gymnolaemata

Order: Cheilostomatida

Family: Smittinidae

Genus: Smittoidea

Species:

prolifica [Describe here as A. iricolor]

Native Distribution

Origin Realm:

Temperate Northern Pacific

Native Region:

Origin Location:

Temperate Northern Pacific [Japan] Fossil records from northern Japan (Hayami 1975) STATUS STATED [US] San Francisco; San Diego Bay; Baja California (Crooks et al 2011; Sorensen et al 2013; Markert et al 2015) STATUS STATED San Francisco Bay. (Altman 2011) STATED Southern California: Newport Harbor, Hankock Stations, Santa Catalina Island, Point Fermin, Santa Cruz Island, Albatoross Station 2945. (Osburn 1952) STATUS NOT STATED Lower California: San Ignacio Lagoon. (Osburn 1952) STATUS NOT STATED Baja California. (Soule & Soule 1964, cited in Markert et al. 2015) STATUS NOT STATED [Western Pacific] Note: the records from the Western Pacific are doubtful and might refer to other species. (De Blauwe & Faasse 2004) STATUS NOT STATED [Japan] [Smittina reticulata (Synonymized taxon)] Mutsu Bay; Fukura, Awaji Island; off Manazuru, Hatsushima Island and Manazuru-zaki in Sagami Bay. (Okada & Mawatari 1936) STATUS NOT STATED [Korea] Sokotra rock in Yellow Sea. (Rho & Seo 1986) STATUS NOT STATED [Korea] All coast of southern Korea. (Seo & Min 2009) STATUS NOT STATED [Korea] Wolseong; Maldo Island; Gaerinyeo Island, Gageodo Island; Hangaechang in Munseom Island, Jeju-do; Cheju-do (Seo and Min 2009; Rho and Seo 1986) STATUS STATED [Mexico] Gulf of California (Soule 1961; Soule and Soule 1964; van den Brink and Wijsman 2010) STATUS NOT STATED CONFLICT: [Antarctic] Rogick (1956) described species as an Antarctic species but accuracy is questioned (Hara 2001) STATUS NOT STATED

Geographic Range:

[Western Pacific] Japan and all coasts of southern Korea (Seo and Min 2009; Soule and Soule 1977; Markert et al 2015; Haydar 2010; De Blauwe and Faasse 2004; van den Brink and Wijsman 2010; Rho and Seo 1986; Seo 1998) [Eastern Pacific] San Francisco Bay, Channel Islands to Baja California; Western Mexico (Carlton 2007; Markert et al 2015; Rho and Seo 1986; Seo 1998; Soule 1961; Soule and Soule 1964) [Eastern Atlantic] Germany; The Netherlands; North Sea (De Bauwe and Faasse 2004; De Blauwe et al 2014; Gollasch et al 2009; Haydar 2010; Markert et al 2015) [The Pacific Coast of North America] Hancock Station 1449-42, Newport Harbor, southern California: 34°35'47"N, 117°52'55"W. (Osburn 1952) [Korea] Sokotra rock in Yellow Sea: 32°08N, 125°10'E. (Rho & Seo 1986) CONFLICT: Rogick (1956) described species as an Antarctic species but accuracy is questioned (Hara 2001)

General Diversity:

Type locality is the Gulf of California (De Blauwe and Faasse 2004, cited in van den Brink and Wijsman 2010)

Non-native Distribution

Invasion History:

Yes (De Blauwe & Faasse 2004, Markert et al. 2015)

Non-native Region:

Northeast Atlantic

Invasion Propens:

Temperate Northern Atlantic [The Netherlands] North Sea (Porter et al 2015; De Blauwe and Faasse 2004; De Blauwe and Faasse 2014; Gollasch et al 2009; Haydar 2010; Haydar and Wolff 2011; Vanagt et al 2013) *Introduced Dutch Delta in the southwest of the Netherland. (De Blauwe & Faasse 2004) *Introduced Harbor region of Rotterdam (Faasse et al. 2013), wind park off Ijmuiden and the WaddenSea in the Netherland. (Vanagt et al. 2013 and Gittenberger et al. 2015, respectively, cited in Markert et al. 2015) *Non-native [Germany] North Sea coast (Markert et al 2015) *Non-native Jade Weser Port and Wihelmshaven in Germany. (Markert et al. 2015) *Non-native

Status Date Non-native:

[The Netherlands] First record of species dates from August 8, 1998; possibly in 1995 or earlier (De Blauwe and Faasse 2004; Haydar 2010; Markert et al 2015) *see notes [Germany] First records in German coastal waters from November 2014 (Markert et al 2015) [The Netherland] Schelphoek, October 1995 (First record in Europe); the harbor region of Rotterdam, 2008 (Faasse et al. 2013); the Prinses Amaliawindpark off IJmuiden, 2011. (Vanagt et al. 2013, cited in Faasse et al. 2013). [The Netherland] Neeltje Jans, August 1998; Goesse Meer, September 1998 and April 2001; Anna Frisopolder, November 1999; Goesse Sas, August 2000; Zierikzee, August 2000; Katse Hoek, June 2001; Westkapelse Kreek, October 2001. (De Blauwe & Faasse 2004) [Germany] Jade Weser Port and Wilhelmshaven, November 2014. (Markert et al. 2015)

Vectors and Spread

Initial Vector:

Aquaculture and Fisheries

Second Vector:

Hull fouling (Commercial, military)

Vector Details:

Most likely route of introduction is suggested to be via shellfish imports (Porter et al 2015; De Blauwe and Faasse 2004) It is less likely that species reached The Netherlands on ship's hull or in ballast water; all collecting localities except one are situated around Oosterschelde, shellfish culture center in The Netherlands (De Blauwe and Faasse 2004) It is likely to settle on ships and in the future (secondary) introductions via ship's hulls may occur as well (De Blauwe and Faasse 2004; Markert et al 2015) S. p. is most likely introduced to Northeast Atlantic waters along with imports of the Pacific oyster for aquaculture purposes. (De Blauwe & Faasse 2004) Secondary introduction via ship hulls seems to be more likely, as S. p. was found only at the commercial harbor Jade Weser Port and at the naval port of Wilhelmshaven. (Makert et al. 2015) Unlikely that transport was via recreational crafts (Markert et al 2015) Aquaculture or stock (Gollasch et al 2009) Oysters as vector into Europe (Haydar 2010; Haydar and Wolff 2011) RELATED: Main vector of transport (into Norwegian waters) is highly likely to be leisure craft, as evidenced by the frequent observation of non-native and invasive species in recreational marinas. Some of the doorknocker species may enter Norwegian waters via the aquaculture route (Porter et al 2015)

Spread Rate:

Limited ability to disperse naturally and that there has been little facilitated transport within its current range in the Netherlands (van den Brink and Wijsman 2010)

Date First Observed in Japan:

Fossil records of species from late Pliocene to early Pleistocene (Hayami 1975)

Date First Observed on West coast North America:

[US] First described from waters off southern California by Osburn in 1952 (Soule and Soule 1964)

Impacts

Impact in Japan:

NF

Global Impact:

The ecological impact of invasive S. p is assumed to be negligible (De Blauwe and Faasse 2004, Gittenberger et al. 2015, cited in Markert et al. 2015), but appropriate studies have not been performed in Europe to date (Merkert et al. 2015).

Tolerences

Native Temperature Regime:

Mild temperate, Warm temperate, Subtropical, Tropical

Native Temperature Range:

Warm temperate to tropical range (Soule 1961) Collected at site with 26˚C (Soule and Soule 1964) Mild temperate, Warm temperate (M. Otani, pers. comm.)

Non-native Temperature Regime:

Mild temperate, Warm temperate, Subtropical, Tropical

Non-native Temperature Range:

8.6˚C - 10.4 ˚C (Markert et al 2015) Warm temperate to tropical range (Soule 1961) Temperature in November at Jade Weser Port was 10.4ºC and that at Wilhelmshaven was 8.6ºC. (Markert et al. 2015) Rotterdam: max 20.0ºC in summer and min 2.0ºC in winter. (Clark et al. 2003) Mild temperate (M. Otani, pers. comm.)

Native Salinity Regime:

Mesohaline, Polyhaline, Euhaline, Hypersaline

Native Salinity Range:

S. p. is found not only in euhaline and polyhaline regions, but also in mesohaline. (Winston 1977) Seasonally hypersaline (Sorensen et al 2013)

Non-native Salinity Regime:

Freshwater, Oligohaline, Mesohaline, Polyhaline, Euhaline

Temperature Regime Survival:

NF

Temperature Range Survival:

NF

Temperature Regime Reproduction:

NF

Temperature Range Reproduction:

NF

Salinity Regime Survival:

NF

Salinity Range Survival:

NF

Salintiy Regime Reproduction:

Polyhaline, Euhaline

Salinity Range Reproduction:

NF

Depth Regime:

Lower intertidal, Shallow subtidal, Deep subtidal

Depth Range:

Intertidal to >100m (Carlton 2007; Soule and Soule 1964) Along shore; also found from along shore down to 45 fms (about 81m) (Soule 1961; van den Brink and Wijsman 2010; De Blauwe and Faase 2004; Markert et al 2015; Seo and Min 2009) Collected in widely differing habitats, ranging from sublittoral boulders and shellgrounds to shallow intertidal pools to brackish water bodies without an open connection to the sea; low substrate specificity, as it was collected from wood, the underside of boulders, living bivalves and empty shells and from algae (De Blauwe and Faasse 2004) [The Pacific Coast of North America] Down to 45 fms. (Osburn 1952) [The Netherland] Shallow intertidal pools to sublittoral. (De Bauwe & Faasse 2004) [Germany] 5m to 12m. (Markert et al. 2015) [Korea] Gaerinyeo Island, Gageodo Island: 18-23m. (Seo & Min 2009) [Korea] Hangaechang in Munseom Island, Jeju-do: 28m. (Seo & Min 2009)

Non-native Salinity Range:

Native Abundance:

Common

Reproduction

Fertilization Mode:

See details

Reproduction Mode:

Hermaphrodite/monoecious

Spawning Type:

NA

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:

Almost every zooid is reproductively active and hermaphroditic, except the youngest two or three rows. Each zooid contains a brood chamber which remains in contact with the neighboring zooid. Each brood chamber has a pore; can reproduce both asexually and sexually (van den Brink and Wijsman 2010) Fertile eggs develop into pelagic larvae which then settle onto appropriate substrate and develop into a new colony (van den Brink and Wijsman 2010) RELATED: [Order: Cheilostomata] Free spawning species produce the characteristic triangular cyphonautes larva. These larvae are long-lived and planktotrophic. The larval body is enclosed in a membranous shell; the size can be up to little over 1 mm. Cyphonautes larvae are not keyed out - if possible at all. (van Couwelaar 2003) [Gymnolaemates] Internal fertilization, whether intracoelomic or intraovarian, is obligatory (Temkin 1994 and 1996, cited in Ostrovsky 2013) [Gymnolaemates] Differ from most organisms in that sperm-egg fusion does not stimulate egg activation. Egg activation may not occur until "spawned" outside of maternal zooid (Temkin 1991) [Bryozoans] While sperm is spawned through pores in lophophore tentacles, eggs are usually harbored inside the body wall, and are internally fertilized by sperm, coming in on lophophore feeding currents (Brusca and Brusca 2003, cited in Rouse 2011; Kozloff 1990, cited in Rouse 2011) [Bryozoans] Colonial hermaphrodites, with testes (spermatogenic tissue) and ovaries developing either within the same zooid (zooidal hermaphroditism) or in different zooids within the same colony (zooidal gonochorism) (Ostrovsky 2013) Members of the phylum Bryozoa are hermaphroditic. Both fertilization and egg brooding may either be internal or external (Ruppert et al. 2004) [Bryozoa] All bryozoan colonies are hermaphroditic. Autozooids may be dioecious; or monoecious, and protandrous or protogynous. (Hayward & Ryland 1999) [Bryozoa] Reproduces asexually by budding. (Mawatari 1976)

Adult Mobility:

Sessile

Adult Mobility Details:

Encrusting species (Markert et al 2015) RELATED: [Bryozoa] The abundance and taxonomic diversity of benthic bryozoan faunas are directly related to substratum. (Hayward & Ryland 1999) [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:

Colony: 1-2 cm round (De Blauwe and Faasse 2004) Autozooids: 0.50-0.70 x 0.20-0.26mm (De Blauwe and Faasse 2004; van den Brink and Wijsman 2010) Zooid: 448 x 236µm (Markert et al 2015)

Maturity Age:

Almost every zooid is reproductively active except the youngest two or three rows (van den Brink and Wijsman 2010) Extremely early start of reproduction, indicated by number of ovicell forming at a colony size of two square millimetres (Markert et al 2015)

Reproduction Lifespan:

NF

Longevity:

Indication that species is a short lived species (Markert et al 2015)

Broods per Year:

NF

Reproduction Cues:

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

Reproduction Time:

Colonies usually form during summer by asexual reproduction through budding (van den Brink and Wijsman 2010) Extremely early start of reproduction, indicated by number of ovicell forming at a colony size of two square millimetres (Markert et al 2015)

Fecundity:

Grows and reproduce rapidly (De Blauwe and Faasse 2004) High fecundity; most zooids carry ovicells, even in early astogenic stages. Some cases of one autozooid as maternal to two ovicells (Markert et al 2015)

Egg Size:

RELATED: Ovicell: 0.20-0.25 wide (De Blauwe and Faasse 2004) [Gymnolaemata] About 200µm (Woollacott and Zimmer 1977)

Egg Duration:

NF

Early Life Growth Rate:

Grows and reproduce rapidly (De Blauwe and Faasse 2004) RELATED: [Gymnolaemata] Two phases of larvae metamorphosis: first stage about 20mins; second stage 1-6 days (Woollacott and Zimmer 1977)

Adult Growth Rate:

Grows and reproduce rapidly (De Blauwe and Faasse 2004)

Population Growth Rate:

Grows and reproduce rapidly (De Blauwe and Faasse 2004)

Population Variablity:

NF

Habitat

Ecosystem:

Sediment subtidal, Rocky intertidal, Rocky subtidal, Oyster reef, Macroalgal beds, Flotsam, Fouling

Habitat Type:

Epibenthic, Epiphytic; Epizoic; Under rock

Substrate:

Sand, Mixed sediments, Cobble, Rock, Biogenic, Artificial Substrate

Exposure:

Semi-exposed, Protected

Habitat Expansion:

NF

Habitat Details:

Fouling panels (Crooks et al 2011) Collected in widely differing habitats, ranging from sublittoral boulders and shellgrounds to shallow intertidal pools to brackish water bodies without an open connection to the sea; low substrate specificity, as it was collected from wood, the underside of boulders, living bivalves and empty shells and from algae (De Blauwe and Faasse 2004) Encrusting stones, shells and stems. Common species on piles and floats and along the shore and down to 45 fms (~81m) (De Blauwe and Faasse 2004) Found on boulders (De Blauwe et al 2014) Encrusting Crassostrea gigas (oyster) and Crepidula fornicata (American slipper limpet)(Markert et al 2015) Capacity to grow on plastics; colonise suitable hard substrates in protected areas (Markert et al 2015) Encrusts other bryozoans (Rho and Seo 1986) Encrusting blades of marine algae and shells of mollusks (Soule 1961) Site with bottom substrate: coarse sand and shell (Soule and Soule 1964) Wide range of habitats including sublittoral boulder and shellgrounds, shallow intertidal pools and brackish water bodies; substrates including wood, underside of boulders, wharf piles, buoys and other floats, living bivalves, empty shells and algae (van den Brink and Wijsman 2010) Found on piles and floats. (Osburn 1952) Known from the Pacific coast and the Gulf of California encrusting stones, shells and stems. (Osburn 1952) Collected from wood, the underside of boulders, living bivalves and empty shells and from algae (Sargassum muticum). (De Blauwe & Faasse 2004) Colony encrusting the other bryozoans. (Rho & Seo 1986) Semi-exposed, Flotsam (M. Otani, pers. comm.)

Trophic Level:

Suspension feeder

Trophic Details:

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:

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:

COMPETITION [Competition] Other encrusting bryozoan species always were more numerous; never seen to overgrow other invertebrates (De Blauwe and Faasse 2004) 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 1999) [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 1999) [Predation] [Bryozoa] Other than opisthobranchs as a predator, amphipods, isopods, mites and pycnogonids have all been recorded preying on bryozoan colonies. (Hayward & Ryland 1999) 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:

Altman S (2011) Diversity, invasiblity, and resouce use in marine fouling commuities of San Francisco Bay. Doctoral thesis in University of Maryland: 1-142. 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. Crooks, J. A., Chang, A. L., & Ruiz, G. M. (2011). Aquatic pollution increases the relative success of invasive species. Biological Invasions, 13(1), 165-176. Doi: 10.1007/s10530-010-9799-3. De Blauwe, H., & Faasse, M. (2004). Smittoidea prolifica Osburn, 1952 (Bryozoa, Cheilostomatida), a Pacific bryozoan introduced to The Netherlands (Northeast Atlantic). Biologie, 74, 33-39. De Blauwe, H., Kind, B., Kuhlenkamp, R., Cuperus, J., Weide, B. v. d., & Kerckhof, F. (2014). Recent observations of the introduced Fenestrulina delicia Winston, Hayward & Craig, 2000 (Bryozoa) in Western Europe. Studi Trentini di Scienze Naturali, 94, 45-51 Faasse M, van Moorsel G, Tempelman D (2013) Moss animals of the Dutch part of the North Sea and coastal waters of the Netherlands (Bryozoa). Nederlandse Faunistische Mededelingen 41: 1-14. http://www.repository.naturalis.nl/document/621036 Gollasch, S., Haydar, D., Minchin, D., Wolff, W. J., & Reise, K. (2009). Introduced aquatic species of the North Sea coasts and adjacent brackish waters. In G. Rilov, & J. E. Crooks (Eds.). Biological Invasions in Marine Ecosystems (pp. 507-528). Heidelberg: Springer Berlin. Hara, U. (2001). Bryozoans from the Eocene of Seymour Island, Antarctic Peninsula. In: A. Gaździcki (Ed.). Palaeontological Results of the Polish Antarctic Expeditions. Part III. Palaeontologia Polonica 60 (pp.33-156). Haydar, D. (2010). What is natural? The scale and consequences of marine bioinvasions in the North Atlantic Ocean. Retrieved from ResearachGate website: https://www.researchgate.net/profile/Deniz_Haydar/publication/44711479_What_is_natural__the_scale_and_consequences_of_marine_bioinvasions_in_the_North_Atlantic_Ocean/links/09e4150d0a68129aa3000000.pdf#page=42 Haydar, D., & Wolff, W. J. (2011). Predicting invasion patterns in coastal ecosystems: relationship between vector strength and vector tempo. Marine Ecology Progress Series, 431, 1-10. Doi: 10.3354/meps09170 Hayami, T. (1975). Neogene Bryozoa from northern Japan. Tohoku Univ., Sci. Rep., 2nd ser. (Geol.), 45(2), 83-126. http://ci.nii.ac.jp/els/110004646784.pdf?id=ART0007368357&type=pdf&lang=jp&host=cinii&order_no=&ppv_type=0&lang_sw=&no=1458033798&cp 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. Hill, K. (2001) Smithsonian Marine Station at Fort Pierce. Retrieved from http://www.sms.si.edu/irlspec/Electr_bellul.htm Markert, A., Matsuyama, K., Rohde, S., Schupp, P., & Wehrmann, A. (2015). First record of the non-native Pacific bryozoan Smittoidea prolifica Osburn, 1952 at the German North Sea coast. Marine Biodiversity, 1-7. Doi: 10.1007/s12526-015-0415-8 Mawatari S (1976) Bryozoa (Ectoprocta). In: Animal systematics. Uchida T (ed.) Nakayama-shoten Co. Ltd., Tokyo: 35-229. (in Japanese) Okada Y & Mawatari S (1936) Bryozoa fauna collected by the "Misago" during the zoological survey around Izu Peninsula (II). Science Report of the Tokyo Bunrika Daigaku Section B Zoology and Bottany 3: 53-73. 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. (2013). Evolution of Sexual Reproduction in Marine Invertebrates – Example of gymnolaemate bryozoans. Dordrectht: Springer Netherlands. Doi: 10.1007/978-94-007-7146-8 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(3), 157-169. Rho, B. J., & Seo, J. E. (1986). A systematic study on the marine bryozoans in Cheju-do. Korean Journal of Zoology, 29(1), 31-60. http://www.koreascience.or.kr/search/articlepdf_ocean.jsp?url=http://ocean.kisti.re.kr/downfile/volume/zsk/HGDMB7/1986/v29n1/HGDMB7_1986_v29n1_31.pdf&admNo=HGDMB7_1986_v29n1_31 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., and Barnes, R.D. (2004). Invertebrate Zoology: A functional evolutionary approach. Ann Arbor, MN: Thomson Brooks/Cole. Seo, J. E. (1998). Taxonomy of the marine bryozoans from Namhaedo Island and its adjacent waters, Korea. The Korean Journal of Systematic Zoology, 14(4), 414-424. Seo, J. E., & Min, B. S. (2009). A faunistic study on cheilostomatous bryozoans from the shoreline of South Korea, with two new species. The Korean Journal of Systematic Zoology, 25(1), 19-40. Sorsen, K., Swope, B., & Kirtay, V. (2013). Marine Ecologic Index Survey of San Diego Bay (Technical Report 2038). Retrieved from Southern California Coastal Water Research Project Westminster website: http://oai.dtic.mil/oai/oai?verb=getRecord&metadataPrefix=html&identifier=ADA603815 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 Soule, D. F., & Soule, J. D. (1964). The Ectoprocta (Bryozoa) of Scammon’s Lagoon, Baja California, Mexico. American Museum Novitates, 2199, 1-56. Soule, J. D., & Soule, D. F. (1977). Fouling and bioadhesion: life strategies of bryozoans. In R. M. Woollacott, & R. L. Zimmer (Eds.). Biology of Bryozoans (pp. 437-457). Elsevier Temkin, M. H. (1991). Fertilization in the Gymnolaemate Bryozoa (Doctoral dissertation). Retrieved from ProQuest Dissertations and Theses database. (DP23819). 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 Van Couwelaar, M. (2003). Zooplankton and Micronekton of the North Sea. Retrieved from http://species-identification.org/species.php?species_group=zmns&menuentry=groepen&id=102&tab=refs van den Brink, A. M., & Wijsman, J. W. M. (2010). High risk exotic species with respect to shellfish transports from the Oosterschelde to the Wadden Sea (No. C025/10). Retrieved from Wageningen website: http://library.wur.nl/WebQuery/wurpubs/fulltext/143113 Vanagt, T., Van de Moortel, L.,& Faasse, M. (2013). Development of hard substrate fauna in the Princess Amalia Wind. Retrieved from PressPage website: http://content.presspage.com/uploads/566/2011036ecoastreporthardsubstratepawpfinal.pdf 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 Woollacott, R. M., & Zimmer, R. L. (Eds.). (1977). Biology of Bryozoans. New York, NY: Academic Press

Literature:

Moderate level of information; data from comparable regions or older data (more than 10 years) from the area of interest

Notes:

*De Blauwe and Faasse (2004) indicates that the species may have been misidentified as S. reticulata in previous publications for the Netherlands; may have existed in the Netherlands prior to 1998