Sacculina

Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Protostomia
Cladus: Ecdysozoa
Cladus: Panarthropoda
Phylum: Arthropoda
Cladus: Pancrustacea
Superclassis: Multicrustacea
Classis: Thecostraca
Subclassis: Cirripedia
Superordo: Rhizocephala

Familia: Sacculinidae
Genera: Sacculina
Species (130): S. abyssicola – S. actaeae – S. aculeata – S. ales – S. americana – S. amplituba – S. anceps – S. angulata – S. anomala – S. atlantica – S. bicuspidata – S. bipunctata – S. boschmai – S. bourdoni – S. brevispina – S. bucculenta – S. bursapastoris – S. caelata – S. calappae – S. calva – S. captiva – S. carcini – S. carpiliae – S. cartieri – S. cavolinii – S. comosa – S. confragosa – S. cordata – S. crucifera – S. curvata – S. cuspidata – S. dayi – S. dentata – S. docleae – S. duracina – S. echinulata – S. elongata – S. eriphiae – S. exarcuata – S. fabacea – S. flacca – S. flexuosa – S. gerbei – S. ghanensis – S. gibba – S. glabra – S. globularis – S. gonoplaxae – S. gordonae – S. gracilis – S. granulosa – S. guineensis – S. hartnolli – S. herbstianodosa – S. hirsuta – S. hirta – S. hispida – S. holthuisi – S. hystrix – S. ignorata – S. inconstans – S. infirma – S. inflata – S. insueta – S. irrorata – S. jamaicensis – S. lata – S. leopoldi – S. leptothrix – S. lobata – S. loricata – S. margaritifera – S. micracantha – S. microthrix – S. muricata – S. nectocarcini – S. nigra – S. nodosa – S. ornatula – S. ostracotheris – S. papposa – S. pertenuis – S. phacelothrix – S. pilosa – S. pinnotherae – S. pisiformis – S. pistillata – S. pomum – S. pugettiae – S. pulchella – S. punctata – S. pustulata – S. quadrialata – S. rathbunae – S. reinhardi – S. reniformis – S. robusta – S. rotundata – S. rugosa – S. scabra – S. schmitti – S. scutigera – S. semistriata – S. senta – S. serenei – S. setosa – S. spectabilis – S. spinosa – S. striata – S. sulcata – Van Kampen & Boschma, 1925 S. surinamensis – Boschma, 1966 S. teres – Boschma, 1933 S. teretiuscula – Boschma, 1931 S. ternatensis – Boschma, 1950 S. upogebiae – Shiino, 1943 S. vankampeni – Boschma, 1931 S. verrucosa – Van Kampen & Boschma, 1925 S. vieta – Boschma, 1933 S. weberi – Boschma, 1931 S. zariquieyi
Boschma, 1947
Name

Sacculina Thompson, 1836ː 452
Type species:ː Sacculina carcini Thompson, 1836ː 452, by monotypy
ZooBank: F830BA20-6480-449A-90FE-DA0EC1A67224

Synonyms

Grapsisaccus – Giard & Bonnier, 1890ː 291, nomen nudum
Type species:ː Sacculina benedeni Kossmann, 1872ː , by monotypy
ZooBank: 0443627D-E7DF-4843-9AE5-885D3A8D91B2
Pachybdella – Diesing, 1850ː 435
Type species:ː Pachybdella rathkei Diesing, 1850ː 435
ZooBank: 926B7A28-EAAA-4E60-837F-5D4E07DC5D7F
Portunascus – Giard & Bonnier, 1890ː 291, nomen nudum
Type species:ː Portunascus corrugatus Giard & Bonnier, 1890ː 291, nomen nudum
ZooBank: FF58A54A-5BA6-4933-B45A-2E3C9AF1CF9E
Sacculinidia – Hesse, 1867ː 378
Type species:ː Sacculinidia gibbsii Hesse, 1867 by monotypy
ZooBank: 9121ED6C-3FA8-4759-A376-ABDF3FA6BECF
Sacculinida – Hesse, 1867ː 380
Type species:ː Sacculinida herbstianodosa Hesse, 1867 by monotypy
ZooBank: 9121ED6C-3FA8-4759-A376-ABDF3FA6BECF

References

Thompson, J.V. 1836. Natural history and metamorphosis of an anomalous crustaceous parasite of Carcinus maenas, the Sacculina carcini. ''The Entomological Magazine'' 3ː 452–456. BHL Reference page. [See p. 452, original description]

Chan, B.K-K., Dreyer, N., Gale, A.S., Glenner, H., Ewers-Saucedo, C., Perez-Losada, M., Kolbasov, G.A., Crandall, K.A. & Høeg, J.T. 2021. The evolutionary diversity of barnacles, with an updated classification of fossil and living forms. Zoological Journal of the Linnean Society 193(3): 789–846. DOI: 10.1093/zoolinnean/zlaa160 Open access Reference page. [See p. 38, discussion]
Diesing, K.M. 1850. Systema helminthum. Vol. 1. pp. [i]–xiii [+ii], 1–680. Vindobonae [Vienna]: sumptibus Academiae Caesareae Scientiarum. Online Google Books Open access Reference page.
Giard, A. & Bonnier, J. 1890. Prodrome d'une monographie des épicarides du Golfe de Naples. Bulletin Scientifique de la France et de la Belgique 22ː 367–391. BHL Reference page. [See p. 291, as Grapsisaccus and Portunascus]
Gurney, R.H., Rybakov, A.V., Høeg, J.T. & Kuris, A.M. 2006. Sacculina nectocarcini, a new species of rhizocephalan (Cirripedia: Rhizocephala) parasitising the red rock crab Nectocarcinus integrifrons (Decapoda: Brachyura: Portunidae). Zootaxa 1332: 37–50. Preview (PDF). Reference page.
Hesse, C-E. 1867. Observations sur des crustacés rares ou nouveux des cotes de France (quatorzième article). Annales des Sciences Naturelles, Zoologie et paléontologie sér. 5 8ː 377–381. BHL Reference page.
Høeg, J.T., Noever, C., Rees, D.J., Crandall, K. & Glenner, H. 2020. A new molecular phylogeny-based taxonomy of parasitic barnacles (Crustacea: Cirripedia: Rhizocephala). Zoological Journal of the Linnean Society 190: 632–653. DOI: 10.1093/zoolinnean/zlz140 Open access pdf Reference page. [See p. 643, discussion]

Sacculina is a genus of barnacles that is a parasitic castrator of crabs. They belong to a group called Rhizocephala. The adults bear no resemblance to the barnacles that cover ships and piers; they are recognised as barnacles because their larval forms are like other members of the barnacle class Cirripedia. The prevalence of this crustacean parasite in its crab host can be as high as 50%.[2][3]
Habitat

Sacculina live in a marine environment. During their larval stage they are pelagic, but as they form into adults they live as ectoparasites on crabs. Their primary host is the green crab, which is native to the Eastern Atlantic Ocean. Though these crabs have spread to other bodies of waters, it is not believed that Sacculina barnacles have traveled with them to these new localities.[4]
Anatomy

The body of the adult parasite can be divided into two parts: one part is called the "externa" where the bulbous reproductive organ of the parasite sticks out of the abdomen of the host. The other part is called the "interna" which is inside the host's body. This part is composed of root-like tendrils that wrap themselves around the host's organs, which gives its group name of Rhizocephala, meaning "root-head". Through microCT scans, these roots have been discovered to wrap around certain organs of the body, with most around the hepatopancreas of crustaceans. This area is primarily for absorbing nutrients, which would explain why most concentrate in that region. In a similar species called Briarosaccus roots were seen extending to the brain and central nervous system, which is a hypothesis to help explain how parasites like these can manipulate their hosts' behavior.
Life cycle

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The female Sacculina larva finds a crab and walks on it until she finds a joint. She then molts into a form called a kentrogon, which then injects her soft body into the crab while her shell falls off. The Sacculina grows in the crab, emerging as a sac, known as an externa, on the underside of the crab's rear thorax, where the crab's eggs would be incubated. Parasitic Sacculina destroy a crab's genitalia, rendering the crab permanently infertile.

After this invasion of the Sacculina, the crab is unable to perform the normal function of molting. This results in a loss of nutrition for the crab, and impairs its overall growth. The natural ability of regrowing a severed claw that is commonly used for defense purposes is therefore lost after the infestation of Sacculina.

The male Sacculina 'larva' looks for a female Sacculina on the underside of a crab. He then implants his cells into a pocket in the female's body called the "testis", where the male cells then produce spermatozoa to fertilize eggs.

When a female Sacculina is implanted in a male crab, it interferes with the crab's hormonal balance. This sterilizes it and changes the bodily layout of the crab to resemble that of a female crab by widening and flattening its abdomen, among other things. The female Sacculina then forces the crab's body to release hormones, causing it to act like a female crab, even to the point of performing female mating dances. If the parasite is removed from the host, female crabs will normally regenerate new ovarian tissue, while males usually develop complete or partial ovaries instead of testes.[5]

Although all energy otherwise expended on reproduction is directed to the Sacculina, the crab develops a nurturing behavior typical of a normal female crab. The natural hatching process of a crab consists of the female finding a high rock and grooming its brood pouch on its abdomen and releasing the fertilized eggs in the water through a bobbing motion. The female crab stirs the water with her claw to aid the flow of the water. When the hatching larvae of Sacculina are ready to emerge from the brood pouch of female Sacculina, the crab performs a similar process. The crab shoots them out in pulses, creating a large cloud of Sacculina larvae. The crab uses the familiar technique of stirring the water to aid in flow.[6]
Life span

Sacculina are primarily host dependent so their life span matches that of their hosts. Crabs usually have a life span anywhere from 1 to 2 years.[7]
Biological control agents

Sacculina has been suggested to be used as a type of biological control agent to help reduce the populations of the invasive green crab. This is controversial because Sacculina can also use native crab species as their host and there would be no way to control or stop them from attacking native species.[4]
Species

More than 100 species of Sacculina are currently recognised:[8]

Sacculina abyssicola Guérin-Ganivet 1911
Sacculina actaeae Guérin-Ganivet 1911
Sacculina aculeata Boschma, 1928
Sacculina ales Kossmann, 1872
Sacculina americana Reinhard, 1955
Sacculina amplituba Phillips, 1978
Sacculina anceps Boschma, 1931
Sacculina angulata van Kampen & Boschma 1925
Sacculina anomala Boschma, 1933
Sacculina atlantica Boschma, 1927
Sacculina beauforti Boschma, 1949
Sacculina bicuspidata Boschma, 1931
Sacculina bipunctata Kossmann, 1872
Sacculina boschmai Reinhard, 1955
Sacculina bourdoni Boschma, 1960
Sacculina brevispina van Kampen & Boschma 1925
Sacculina bucculenta Boschma, 1933
Sacculina bursapastoris Kossmann, 1872
Sacculina caelata Boschma, 1931
Sacculina calappae van Kampen & Boschma 1925
Sacculina calva Boschma, 1933
Sacculina captiva Kossmann, 1872
Sacculina carcini Thompson, 1836
Sacculina carpiliae Guérin-Ganivet 1911
Sacculina cartieri Kossmann, 1872
Sacculina cavolinii Kossmann, 1872
Sacculina comosa Boschma, 1931
Sacculina compressa Boschma, 1931
Sacculina confragosa Boschma, 1933
Sacculina cordata Shiino, 1943
Sacculina crucifera Kossmann, 1872
Sacculina curvata Boschma, 1933
Sacculina cuspidata Boschma, 1949
Sacculina dayi Boschma, 1958
Sacculina dentata Kossmann, 1872
Sacculina docleae Huang & Lützen, 1998
Sacculina duracina Boschma, 1933
Sacculina echinulata van Kampen & Boschma 1925
Sacculina elongata Boschma, 1933
Sacculina eriphiae Smith, 1906
Sacculina exarcuata Kossmann, 1872
Sacculina fabacea Shiino, 1943
Sacculina flacca Boschma, 1931
Sacculina flexuosa Kossmann, 1872
Sacculina gerbei Guérin-Ganivet 1911
Sacculina ghanensis Boschma, 1971
Sacculina gibba Boschma, 1933
Sacculina glabra van Kampen & Boschma 1925
Sacculina globularis Boschma, 1970
Sacculina gonoplaxae Guérin-Ganivet 1911
Sacculina gordonae Boschma, 1933
Sacculina gracilis Boschma, 1931
Sacculina granifera Boschma, 1973
Sacculina granulosa Boschma, 1931
Sacculina guineensis Boschma, 1971
Sacculina hartnolli Boschma, 1965
Sacculina herbstianodosa (Hesse, 1867)
Sacculina hirsuta Boschma, 1925
Sacculina hirta Boschma, 1933
Sacculina hispida Boschma, 1928
Sacculina holthuisi Boschma, 1956
Sacculina hystrix van Kampen & Boschma 1925
Sacculina ignorata Boschma, 1947
Sacculina imberbis Shiino, 1943
Sacculina inconstans Boschma, 1952
Sacculina infirma Boschma, 1953
Sacculina inflata Leuckart, 1859
Sacculina insueta Boschma, 1966
Sacculina irrorata Boschma, 1934
Sacculina jamaicensis Boschma, 1966
Sacculina lata Boschma, 1933
Sacculina leopoldi Boschma, 1931
Sacculina leptothrix Boschma, 1933
Sacculina lobata Boschma, 1965
Sacculina loricata Boschma 1955
Sacculina margaritifera Kossmann, 1872
Sacculina micracantha Boschma, 1931
Sacculina microthrix Boschma, 1931
Sacculina muricata Boschma, 1931
Sacculina nectocarcini Gurney et al. 2006
Sacculina nigra Shiino, 1943
Sacculina nodosa Boschma, 1931
Sacculina ornatula Boschma, 1951
Sacculina ostracotheris Boschma, 1967
Sacculina papposa van Kampen & Boschma 1925
Sacculina pertenuis Boschma, 1933
Sacculina phacelothrix Boschma, 1931
Sacculina pilosa Kossmann, 1872
Sacculina pilosella van Kampen & Boschma 1925
Sacculina pinnotherae Shiino, 1943
Sacculina pisiformis Kossmann, 1872
Sacculina pistillata Boschma, 1952
Sacculina pomum Kossmann, 1872
Sacculina pugettiae Shiino, 1943
Sacculina pulchella Boschma, 1933
Sacculina punctata Boschma, 1934
Sacculina pustulata Boschma, 1925
Sacculina quadrialata Boyko & van der Meij, 2018
Sacculina rathbunae Boschma, 1933
Sacculina reinhardi Boschma 1955
Sacculina reniformis Boschma, 1933
Sacculina robusta Boschma, 1948
Sacculina rotundata Miers, 1880
Sacculina rugosa van Kampen & Boschma 1925
Sacculina scabra Boschma, 1931
Sacculina schmitti Boschma, 1933
Sacculina scutigera Huang & Lützen, 1998
Sacculina semistriata van Kampen & Boschma 1925
Sacculina senta Boschma, 1933
Sacculina serenei Boschma, 1954
Sacculina setosa van Kampen & Boschma 1925
Sacculina spectabilis Boschma, 1948
Sacculina spinosa van Kampen & Boschma 1925
Sacculina striata Boschma, 1931
Sacculina sulcata van Kampen & Boschma 1925
Sacculina surinamensis Boschma, 1966
Sacculina teres Boschma, 1933
Sacculina teretiuscula Boschma, 1931
Sacculina ternatensis Boschma, 1950
Sacculina upogebiae Shiino, 1943
Sacculina vankampeni Boschma, 1931
Sacculina verrucosa van Kampen & Boschma 1925
Sacculina vieta Boschma, 1933
Sacculina weberi Boschma, 1931
Sacculina zariquieyi Boschma, 1947

References

Thompson, J. V. (1836). "Natural history and metamorphosis of an anomalous crustaceous parasite of Carcinus maenas, the Sacculina carcini". The Entomological Magazine. 3: 452–456.
Ross, Piper (2007). Extraordinary animals : an encyclopedia of curious and unusual animals. Westport, Conn.: Greenwood Press. ISBN 978-0313339226. OCLC 191846476.
Leung, Tommy (2016-10-06). "Peltogaster sp". Parasite of the Day. Retrieved 2017-11-30.
"The Parasitic Sacculina That Bends Its Host to Its Own Will". Today I Found Out. 2013-10-07. Retrieved 2017-11-30.
General Parasitology
Zimmer, Carl (2001). Parasite rex : inside the bizarre world of nature's most dangerous creatures (1st Touchstone ed.). New York: Simon & Schuster. ISBN 074320011X. OCLC 47903774.
"Sacculina carcini". Animal Diversity Web. Retrieved 2017-11-30.

"World Register of Marine Species, genus Sacculina". Retrieved 2021-09-01.

Sources
Boschma, H. (1955). "The described species of the family Sacculinidae". Zoologische Verhandelingen. 27 (1): 1–76.

(cites many earlier papers by Boschma and others including other sources for the above list)

Guérin-Ganivet, J. (1911). "Contribution a l'étude systématique et biologique des Rhizocéphales". Travaux scientifiques du Laboratoire de Zoologie et de Physiologie Maritimes de Concarneau. 3 (7): 1–97.
Gurney, R. H.; Rybakov, A. V.; Høeg, J. T.; Kuris, A. M. (2006). "Sacculina nectocarcini, a new species of rhizocephalan, a new species of rhizocephalan(Cirripedia: Rhizocephala) parasitising the red rock crab Nectocarcinus integrifrons (Decapoda: Brachyura: Portunidae)(Decapoda: Brachyura: Portunidae)". Zootaxa. 1332: 37–50. doi:10.11646/zootaxa.1332.1.2.
van Kampen, Pieter Nicolaas; Boschma, Hilbrandt (1925). Die rhizocephalen der Siboga-expedition (in German). Leiden: Brill. OCLC 6845128.