Exochella longirostris

Overview

Scientific Name: Exochella longirostris

Phylum: Bryozoa

Class: Gymnolaemata

Order: Cheilostomatida

Family: Exochellidae

Genus: Exochella

Species:

longirostris* [Describe here as A. iricolor]

Native Distribution

Origin Realm:

Temperate Northern Pacific, Temperate Australasia, Temperate Northern Atlantic, Tropical Atlantic, Temperate South America, Southern Ocean

Native Region:

Origin Location:

Temperate Northern Pacific [Japan] Found fossil records from the Daishaka Formation (southwestern part of Aomori City (Hayami 1975) STATUS STATED [Japan] Shirahama, Tonda, Wakayama Prefecture. (Mawatari 1952) STATUS NOT STATED [Japan] Seto Inland Sea. (Inaba 1988) STATUS NOT STATED Temperate South America [Argentina] Strait of Magellan; Santa Cruz ; Tierra del Fuego (Centurion and Gappa 2011; Gappa 1989; Gappa and Lichtschein 1988; Gappa and Sueiro 2007; Jullien 1888, cited in Hayward 1991) STATUS STATED [Chile] Ile Hoste, baie Orange, Canal du Beagle, Ile Gable, Tierra del Fuego, Punta Arenas in Magellan Stratits. (Jullien, 1888, Calvet 1904, Canu 1908, cited in Rogick 1956) STATUS NOT STATED [Chile] Magellan region. (Moyano 1999) STATUS NOT STATED [Argentina] Along the Patagonian coast from Tierra del Fuego to Peninsula Valdés. (Bastida et al. 1992) STATUS NOT STATED [Argentina] From Post-Pampeen de Punta Borja, Puerto Militar, Bhia Blanca. (Canu 1908, cited in Rogick 1956) STATUS NOT STATED [Brazil] Sta. Catharina, Parana, South of Victoria, Espirito Santo. (Marcus 1937, 1941, 1949, cited in Rogick 1956) STATUS NOT STATED [Brazil] Espirito Santo; Santa Catarina; Sao Paulo; Parania (Almeida et al 2015; Kelmo et al 2004; Marcus 1937, cited in Vieira et al 2008) STATUS NOT STATED [Chile] (Hayward 1995, cited in Winston and Woollacott 2009; Giguerolas et al 2014) STATUS NOT STATED Temperate Northern Atlantic [US] Georgia; South Carolina (Wenner et al 1984; Cheetham and Sandberg 1964; Lagaaij 1963; Moretzohn et al 2011; Winston and Maturo 2009) STATUS STATED Tropical Atlantic [Caribbean] Barbados (Winston and Woollacott 2009) STATUS NOT STATED [Brazil] Bay of Santos (Marcus 1937, 1941, 1949, cited in Rogick 1956) STATUS NOT STATED [Brazil] Bahia State (Almeida et al 2015; Kelmo et al 2004; Marcus 1937, cited in Vieira et al 2008) STATUS NOT STATED Temperate Australasia [New Zealand] Auckland (Gordon 1967) STATUS NOT STATED Southern Ocean [Antarctica] Deception Island; Patagonian assemblages, Falkland Islands, West Falkland (Barnes and De Grave 2001; Barnes and Lehane 2001; Barnes et al 2008; Hayward 1980; Hennig 1911, cited in Thomson 1977) STATUS NOT STATED [South Shetland Islands, Antarctica] Deception Island. (Barnes et al. 2008) STATUS NOT STATED Uncertain realm Gulf of Mexico (Wenner et al 1984; Cheetham and Sandberg 1964; Lagaaij 1963; Moretzohn et al 2011; Winston and Maturo 2009) STATUS STATED Gulf of Mexico and Africa. (Winston & Maturo 2009) STATUS NOT STATED [Africa] South Atlantic Ocean (Winston and Maturo 2009) STATUS NOT STATED [Mexico] Gulf of Mexico (Cheetham and Sandberg 1964; Lagaaij 1963; Moretzohn et al 2011; Winston and Maturo 2009) STATUS NOT STATED

Geographic Range:

[Western Pacific] Japan; New Zealand (Hayami 1975; Gordon 1967) [Eastern Pacific] Chile (Giguerola et al 2014; Winston and Woollacott 2009) [Western Atlantic] South Carolina, US to Argentina; Falkland Islands (Wenner et al 1984; Cheetham and Sandberg 1964; Centurion and Gappa 2011; Hayward 1991; Moretzsohn et al 2011; Giguerola et al 2014; Winston and Woollacott 2009; Moyano 1999) [Eastern Atlantic] Africa (Winston and Maturo 2009) [Antarctica] (Barnes and De Grave 2001; Barnes and Lehane 2001; Barnes et al 2008; Hayward 1980; Hennig 1911, cited in Thomson 1977) Southern hemisphere: ranging from 20°33'S to about 55°40'S. (Rogick 1956) Marguerite Bay, Antarctica: approximately 68°30'S, 68°30'W. (Rogick 1956) Around Japan: 30°N-43°N both at Pacific side and Japan Sea side. (Inaba 1988)

General Diversity:

Antarctica (species complex?) (Moretzsohn et al 2011; Winston and Maturo 2009)

Non-native Distribution

Invasion History:

See details

Non-native Region:

Northwest Atlantic

Invasion Propens:

Temperate Northern Atlantic [US] Mississippi River on the Louisiana Gulf Coast (Cheetham and Sandberg 1964) *Discussed as surprising to see as its a South Atlantic species (did not describe as invasive)

Status Date Non-native:

[US] First time seen in US (specifically Louisiana Gulf Coast) (Cheetham and Sandberg 1964)

Vectors and Spread

Initial Vector:

NF

Second Vector:

NF

Vector Details:

NF

Spread Rate:

Typically from South America. Known range extended northward to within 25 degrees of the equator (Marcus 1949, cited in Cheetham and Sandberg) and observed for the first time in the northern hemisphere (Cheetham and Sandberg 1964)

Date First Observed in Japan:

Fossil records of species in Japan dating since early Pliocene (Hayami 1975)

Date First Observed on West coast North America:

NF

Impacts

Impact in Japan:

NF

Global Impact:

NF

Tolerences

Native Temperature Regime:

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

Native Temperature Range:

-1.439 to 24.233°C (OBIS 2015) [Argentina] Puerto Deseado, Santa Cruz had water temperatures about 4.5°C in winter to 13.5°C in summer (Gappa 1989) Along the Patagonian coast from Tierra del Fuego to Peninsula Valdés: 4.6ºC-13ºC. (Bastida et al. 1992) Marguerite Bay: 30.2ºF (-1.0ºC) at the bottm of 85-100 fathoms deep. (Rogick 1956) Tropical (M. Otani, pers. comm.)

Non-native Temperature Regime:

NF

Non-native Temperature Range:

NF

Native Salinity Regime:

Euhaline

Native Salinity Range:

33.165-36.149 PPS (OBIS 2015) [Argentina] Puerto Deseado, Santa Cruz has salinity around 33%o (Gappa 1989)

Non-native Salinity Regime:

NF

Temperature Regime Survival:

See details

Temperature Range Survival:

-1.439 to 24.233°C (OBIS 2015)

Temperature Regime Reproduction:

NF

Temperature Range Reproduction:

NF

Salinity Regime Survival:

Euhaline

Salinity Range Survival:

33.165-36.149 PPS (OBIS 2015)

Salintiy Regime Reproduction:

Polyhaline, Euhaline

Salinity Range Reproduction:

NF

Depth Regime:

Lower intertidal, Shallow subtidal, Deep subtidal

Depth Range:

0-4m depth (Moretzsohn et al 2011; Cheetham and Sandberg 1964; Lagaaij 1963) 0-200m (Figuerola et al 2014) [Mexico] Collected from 70fms (128.1m) (Lagaaij 1963, cited in Hayami 1975) [Caribbean] Barbados: Sampled from 80fms (146.4m) (Winston and Woollacott 2009) Epibenthic encrusting; 4m (Winston and Maturo 2009) Bay of Santos, Brazil: 20m. (Marcus 1937, cited in Rogick 1956) South of Victoria, Espirito Santo: 35m. (Marcu 1949, cited in Rogick 1956) Along the Patagonian coast from Tierra del Fuego to Peninsula Valdés: 55-95m. (Bastida et al. 1992) Tierra del Fuego: 19m. (Jullien 1888, cited in Rogick 1956) Marguerite Bay, Antarctica: 85-100 fathoms. (Rogick 1956) Seto Inland Sea, Japan: lower intertidal zone to 20-30m. (Inaba 1988)

Non-native Salinity Range:

Native Abundance:

Common, See details

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:

Ovicells present (Ostrovskii 2004) Macrolecithal oocytes (Ostrovsky 2013) RELATED: [Gymnolaemates] Internal fertilization, whether intracoelomic or intraovarian, is obligatory (Temkin 1994 and 1996, cited in Ostrovsky 2013) [Gymnolaemates] Differ from most organisms in that sperm-egg fusion does not stimulate egg activation. Egg activation may not occur until "spawned" outside of maternal zooid (Temkin 1991) [Bryozoans] 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 1999) [Bryozoa] Reproduces asexually by budding. (Mawatari 1976)

Adult Mobility:

Sessile

Adult Mobility Details:

Epibenthic encrusting (Morezsohn et al 2011) RELATED: [Bryozoa] The abundance and taxonomic diversity of benthic bryozoan faunas are directly related to substratum. (Hayward & Ryland 1999) [Bryozoa] Bryozoans are a phylum of sessile, colonial suspension feeders found throughout the world in both marine and freshwater environments. (Tilbrook 2012)

Maturity Size:

Colony area approximately 0.5cm2 (Centurion and Gappa 2011) Colony thickness approximately 0.32mm (Gappa 1989) Zooecia: 0.52-0.60mm (Hayami 1975) Autozooid: 0.49x0.37mm (Hayward 1991)

Maturity Age:

NF

Reproduction Lifespan:

NF

Longevity:

NF

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:

NF

Fecundity:

NF

Egg Size:

RELATED: [Exochella] Growing oocyte: 19.0x15.0μm; Early vitellogenic: 70.0x55.0μm; Mature oocyte: 145.0x105.0μm (Ostrovsky 2013) [Gymnolaemata] About 200µm (Woollacott and Zimmer 1977)

Egg Duration:

NF

Early Life Growth Rate:

Oocyte growth was minimum 8-fold (Ostrovsky 2013) RELATED: [Gymnolaemata] Two phases of larvae metamorphosis: first stage about 20mins; second stage 1-6 days (Woollacott and Zimmer 1977)

Adult Growth Rate:

NF

Population Growth Rate:

NF

Population Variablity:

NF

Habitat

Ecosystem:

Sediment subtidal, Rocky intertidal, Rocky subtidal, Macroalgal beds, Fouling, Other

Habitat Type:

Epibenthic, Epiphytic, Epizoic

Substrate:

Mud, Gravel, Cobble, Rock, Biogenic, Artificial substrate

Exposure:

Semi-exposed, Protected

Habitat Expansion:

NF

Habitat Details:

Encrusting on pebbles weighing 60-80g (Centurion and Gappa 2011) Found in mudlumps in Louisiana (Cheetham and Sandberg 1964) Collected during experiments on acrylic panels (Gappa 1989) [Japan] Admixed in shell bed (Hayami 1975) [New Zealand] Found on shells and/ or barnacle plates (Gordon 1967) Epibenthic encrusting. (Winston & Maturo 2009) Found on rocks, gravels, and algae. (Inaba 1988) Semi-exposed, Protected (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] Competition for space between bryozoan species , U. alvareziana dominated over this species may be related to colony size (Gappa 1989) DISTURBANCE [Disturbance] El Nino events may be responsible for species disappearing in Northern Bahia, Brazil (Kelmo et al 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 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:

EPIBIONTS [Epibionts] Sponge spicules are matted over one colony, calcareous worm tubes and occasional bryozoan zoids (of other species) are present on other colonies. However, most of the colony surface is free of extraneous growths. (Rogick 1956)

References and Notes

References:

Almeida, A. C. S., Alves, O., Peso-Aguiar, M., Dominguez, J., & Souza, F. (2015). Gymnolaemata bryozoans of Bahia State, Brazil. Marine Biodiversity Records, 8, e120. Doi: 10.1017/s1755267215000743 Barnes, D. K. A., & De Grave, S. (2001). Ecological biogeography of southern polar encrusting faunas. Journal of Biogeography, 28(3), 359-365 Barnes, D. K. A., & Lehane, C. (2001). Competition, mortality and diversity in South Atlantic coastal boulder communities. Polar Biology, 24(3), 200-2008 Barnes, D. K. A., Linse, K., Enderlein, P., Smale, D., Fraser, K. P. P., & Brown, M. (2008). Marine richness and gradients at Deception Island, Antarctica. Antarctic Science, 20(3), 271-279. Doi: 10.1017/s0954102008001090. http://journals.cambridge.org/action/displayFulltext?type=1&fid=1875136&jid=ANS&volumeId=20&issueId=03&aid=1875128 Bastida, R., Roux, A., & Martinez, D. E. (1992). Benthic communities of the argentine continental-shelf. Oceanologica Acta, 15(6), 687-698. http://archimer.ifremer.fr/doc/00101/21234/18851.pdf Centurión, R., & Gappa, J. L. (2011). Bryozoan assemblages on hard substrata: species abundance distribution and competition for space. Hydrobiologia, 658(1), 329-341 Cheetham, A. H., & Sandberg, P. A. (1964). Quaternary Bryozoa from Louisiana Mudlumps. Journal of Paleontology, 38(6), 1013-1046 Figuerola, B., Gordon, D. P., Polonio, V., Cristobo, J., & Avila, C. (2014) Cheilostome bryozoan diversity from the southwest Atlantic region: Is Antarctica really isolated?. Journal of Sea Research, 85, 1-17 Gappa, J. J. L. (1989). Overgrowth competition in an assemblage of encrusting bryozoans settled on artificial substrata. Marine Ecology Progress Series, 51, 121-130 Gappa, J. J. L., & Lichtschein, V. (1988). Geographic distribution of bryozoans in the Argentine Sea (South-Western Atlantic). Oceanologica Acta, 11(1), 89-99 Gappa, J. J. L., & Sueiro, M. C. (2007). The subtidal macrobenthic assemblages of Bahía San Sebastián (Tierra del Fuego, Argentina). Polar Biology, 30(6), 679-687. Doi: 10.1007/s00300-006-0225-3 Global Invasive Species Database. http://www.iucngisd.org/gisd/ Access Date: 11-Mar-2016. Gordon, D. P. (1967). A Report on the Ectoproct Polyzoa of some Auckland Shores. Tane, 13, 43-76 Hayami, T. (1975). Neogene Bryozoa from northern Japan. Tohoku Univ., Sci. Rep., 2nd ser. (Geol.), 45(2), 83-126 Hayward, P. J. (1980). Cheilostomata (Bryozoa) from the South Atlantic. Journal of Natural History, 14(5), 701-721. Doi: 10.1080/00222938000770591 Hayward, P. J. (1991). Systematic studies on some Antarctic and sub-Antarctic Ascophora (Bryozoa: Cheilostomata). Zoological Journal of the Linnean Society, 101(4), 299-335 Hill, K. (2001) Smithsonian Marine Station at Fort Pierce. Retrieved from http://www.sms.si.edu/irlspec/Electr_bellul.htm 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. 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) 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 Lagaaij, R. (1963). New additions to the bryozoans fauna of the Gulf of Mexico. Publications of the Institute of Marine Science, University of Texas, 9, 162-236 Mawatari S (1976) Bryozoa (Ectoprocta). In: Animal systematics. Uchida T (ed.) Nakayama-shoten Co. Ltd., Tokyo: 35-229. (in Japanese) Moretzsohn, F., Brenner, J., Michaud, P., Tunnell, J. W., & Shirley, T. (2011). Biodiversity of the Gulf of Mexico Database (BioGoMx). Retrieved from http://gulfbase.org/biogomx/biospecies.php?species=Spp-68-0186 Moyano G., H. I. (1999). Magellan Bryozoa: A review of the diversity and of the Subantarctic and Antarctic zoogeographical links. Scientia Marina, 63(Supl. 1), 219-226. http://scimar.icm.csic.es/scimar/pdf/63/sm63s1219.pdf Ostrovskii, A. N. (2004). Brood chambers (ovicells) of cheilostome bryozoans (bryozoa: Gymnolaemata): structure, research history, and modern problematic. Russian Journal of Marine Biology, 30(Suppl.1), S43-S55 OBIS. (2015). Ocean Biogeographic Information System. Retrieved from http://iobis.org/mapper 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 Rogick M (1956) Studies on marine Bryozoa. VIII. Exochella longirostris Jullien 1888. Biological Bulletin 111: 123-128. http://www.biolbull.org/content/111/1/123.full.pdf 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. Temkin, M. H. (1991). Fertilization in the Gymnolaemate Bryozoa (Doctoral dissertation). Retrieved from ProQuest Dissertations and Theses database. (DP23819). Thomson, M. R. A. (1977). An annotated bibliography of the paleontology of Lesser Antarctica and the Scotia Ridge. New Zealand Journal of Geology and Geophysics, 20(5), 865-904. Doi: 10.1080/00288306.1977.10420686 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 Vieira, L. M., Migotto, A. E., & Winston, J. E. (2008). Synopsis and annotated checklist of Recent marine Bryozoa from Brazil. Zootaxa, 1810, 1-39 Waters, A. W. (1905). Bryozoa from near Cape Horn. Journal of the Linnean Society of London, Zoology, 29(191), 230-251. Doi: 10.1111/j.1096-3642.1905.tb00040.x Wenner, E. L., Hinde, P., Knott, D. M., & Van Dolah, R. F. (1984). A Temporal and Spatial Study of Invertebrate Communities Associated with Hard-Bottom Habitats in the South Atlantic Bight (NOAA Technical Report NMFS 18). Retrieved from the US NOAA website: http://spo.nmfs.noaa.gov/tr18opt.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 Winston JE & Maturo Jr. FJ (2009) Bryozoans (Ectoprocta) of the Gulf of Mexico. In: Gulf of Mexico - origins, waters, and biota vol.1: Biodiversity. Felder DL & Camp DK (eds.) , Texas A & M University Press, College Station, Texas: 1147-1164. http://biogomx.org/sites/default/files/pdfs/chapters/68-Winston%20and%20Maturo%202009-Bryozoans%20of%20the%20GoMx.pdf Winston, J. E., Vieira, L. M., & Woollacott, R. M. (2014). Scientific Results of the Hassler Expedition. Bryozoa. No. 2. Brazil. Bulletin of the Museum of Comparative Zoology, 161(5), 139-239 Winston, J. E., & Woollacott, R. M. (2009). Scientific Results of the Hassler Expedition. Bryozoa. No. 1. Barbados. Bulletin of the Museum of Comparative Zoology, 159(5), 239-300. Doi: 10.3099/0027-4100-159.5.239 Woollacott, R. M., & Zimmer, R. L. (Eds.). (1977). Biology of Bryozoans. New York, NY: Academic Press

Literature:

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

Notes:

Possibly previously described as Smittia tricuspis (Hincks) although WoRMS did not list as so (Waters 1905)