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Ascidiacea World Database

By Noa Shenkar, Arjan Gittenberger, Gretchen Lambert, Marc Rius, Rosana

Moreira da Rocha, Billie J Swalla, Xavier Turon
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Background

Ascidians (Phylum: Chordata, Class: Ascidiacea), or sea squirts, are the largest
and most diverse class of the sub-phylum Tunicata (also known as Urochordata).
They comprise approximately 3000 described species found in all marine
habitats from shallow water to the deep sea. There are no freshwater species,
and most cannot tolerate salinities below about 20. Recent phylogenomic
studies suggest that they are actually the sister group to the vertebrates (Bourlat
et al., 2006), although this conflicts with rRNA and mitochondrial data (Swalla
and Smith, 2008). Adult ascidians bear little resemblance to typical chordates,
though their short-lived non-feeding tadpole larvae clearly exhibit the four
fundamental characteristics of the phylum: a dorsal tubular nerve cord,
notochord, rudimentary pharyngeal gill slits and a post-anal tail. Another
important character is the presence of the endostyle in the pharynx that will
evolve as the thyroid gland in vertebrates. Following settlement, the
lecithotrophic larvae undergo metamorphosis during which they lose all of these
characteristics except for the endostyle and the gill slit rudiments in the pharynx
(Millar, 1971), which become functional and multiply to form the branchial sac.
The name "tunicate" (sub-phylum Tunicata) comes from the polysaccharidecontaining tunic that envelops the animal and forms a somewhat flexible
skeleton (Swalla and Cameron, 2010). Various proteins and blood cells occur in
the tunic, and spicules in a few species (Monniot et al., 1991; Hirose 2009).
Ascidians filter their food from the water-column via an oral siphon that brings
water into the branchial sac where food items such as microalgae are filtered
onto a mucus net; water, feces and gametes are expelled through an excurrent
siphon. Particles suspended in the current are trapped along the wall of the
branchial sac in a mucus net produced by the endostyle. The net pores range
from 0.1 to 1 m, allowing ascidians to filter even very small particulate matter,
primarily in the range of 0.5 to 10 m diameter (Bak et al., 1998; Bone et al.,

2003). Several solitary species are cultured for food in Japan, Korea, France and
Chile (Lambert, 2005) or extracted straight from rocky shores for human
consumption or bait (Branch et al. 2010).

During the past two decades enormous progress has been achieved in the fields
of development, evolution, immunology, natural products and ecology of
ascidians. Their small genome, small cell number and (usually) short life-cycle
make them an attractive model system for developmental biologists (Dehal et
al., 2001; Nishida and Sawada 2001). Investigating the phylogenetic position of
the subphylum Tunicata in relation to the other subphyla in the phylum Chordata
is crucial to the understanding of possible mechanisms of chordate evolution
(Swalla et al., 2000; Zeng and Swalla, 2005). The study of self/non-self
recognition in ascidians provides important information regarding the
evolutionary origin of the vertebrate immune system (Khalturin and Bosch,
2007). In addition, ascidians provide a fertile ground for studies in the field of
natural products (Wang et al., 2007 for review), and play an important role in
marine bioinvasions across the globe (Locke and Carman, 2009).

Systematics

Kowalevsky discovered the chordate nature of the ascidian tadpole larva in 1866;
the Ascidiacea were then reclassified as chordates rather than as molluscs (Raff
and Love, 2004). Following Lahille (1886), the class Ascidiacea is now divided
into three orders based on the structure of the adult branchial sac:
Aplousobranchia (colonial), Phlebobranchia and Stolidobranchia. This is the
current designation used by most ascidian taxonomists. All ascidians are
hermaphrodites, having both male and female gonads, though many are not selffertile. There are both solitary and colonial species; numerous excellent
anatomical illustrations can be found in Monniot et al. (1991).

Ascidian systematics is the domain of specialists, but a keen observer can

differentiate the orders and most families, and recognize the well-described
common species. Although you cannot judge an ascidian by its cover,
underwater photographs can provide much information and assist in
distinguishing one species from another (Monniot et al., 1991) especially if the
fauna of the given region has already been studied. The invaluable monographs
published by Van Name (1945), C. and F. Monniot (Monniot et al., 1991; Monniot
and Monniot, 1996, 2001 and many others) and Kott (1985, 1990, 1992, 2001
and others) greatly assist in identifying the ascidian fauna worldwide to species
level. Tabular keys to the families and genera of the world can be found in
Monniot et al. (1991). A comprehensive listing of nearly all the publications on

ascidians since 1995 can be found by clicking on the various newsletters listed at
http://depts.washington.edu/ascidian

Literature cited

Bak RPM, Joenje M, de Jong I, Lambrechts DYM, Nieuwland G (1998) Bacterial

suspension feeding by coral reef benthic organisms. Mar Ecol Prog Ser 175: 285
288
Bone Q, Carre C, Chang P (2003) Tunicate feeding filters. J Mar Biol Ass UK 83:
907-919
Khalturin K, Bosch TCG (2007) Self/nonself discrimination at the basis of chordate
evolution: limits on molecular conservation. Curr Opin Immunol 19: 4-9
Bourlat SJ, Juliusdottir T, Lowe CJ, Freeman R, Aronowicz J, Kirschner M, Lander
ES, Thorndyke M, Nakano H, Kohn AB, Heyland A, Moroz LL, Copley RR & Telford
MJ (2006) Deuterostome phylogeny reveals monophyletic chordates and the new
phylum Xenoturbellida. Nature 444: 85-88
Branch GM, Griffiths CL, Branch ML, Beckley LE (2010) Two Oceans: A guide to
the marine life of southern Africa. Struik Publishers, Cape Town, 456 pp.
Dehal P et al. (+ 86 additional authors) (2002) The draft genome of Ciona
intestinalis: Insights into chordate and vertebrate origins. Science 298: 21572167
Hirose E (2009) Ascidian tunic cells: morphology and functional diversity of free
cells outside the epidermis. Invertebr Biol 128: 8396
Kott P (1985) The Australian Ascidiacea. Part 1: Phlobranchia and
Stolidobranchia. Memoirs of the Queensland Museum 23: 1-440
Kott P (1990) The Australian Ascidiacea. Part 2: Aplousobranchia (1). Memoirs of
the Queensland Museum 29 (1): 1-226
Kott P (1992) The Australian Ascidiacea. Part 3: Aplousobranchia (2). Memoirs of
the Queensland Museum 32 (2): 375-620
Kott P (2001) The Australian Ascidiacea. Part 4: Aplousobranchia (3),
Didemnidae. Memoirs of the Queensland Museum 47 (1): 1-407
Lahille MF (1886) Sur la classification des Tuniciers. C. R. Acad. Sci. Paris 102:
15731575
Lambert G (2005) Ecology and natural history of the protochordates. Can J Zool
83: 34-50

Locke A, Carman M (2009) An overview of the 2nd International Invasive Sea

Squirt Conference: What we learned. Aquat Inv 4: 1-4
Millar RH (1971) The biology of ascidians. Adv Mar Biol 9: 1-100
Monniot C, Monniot F, Laboute P (1991) Coral reef ascidians of New Caledonia.
ORSTOM, Paris
Monniot C, Monniot F (1996) New collections of ascidians from the western
Pacific and Southeastern Asia. Micronesia 29 (2): 133-279
Monniot C, Monniot F (2001) Ascidians from the tropical western Pacific.
Zoosystema 23 (2): 201-383
Nishida H, Sawada K (2001) Macho-1 encodes a localized mRNA in ascidian eggs
that specifies muscle fate during embryogenesis. Nature 409 : 724729
Perrier E (1898) Note sur la Classification des Tuniciers. CR Acad Sci Paris 126: 1
5.
Raff RA, Love AC (2004) Kowalevsky, Comparative Evolutionary Embryology, and
the Intellectual Lineage of Evo-Devo. J Exp Zool (Mol Dev Evol) 302B: 19-34
Swalla BJ, Cameron CJ (2010) Tunicata in Phylocode. Tunicata in Phylocode: A
Phylogenetic Code of Biological Nomenclature. Version 2b P. D. Cantino and K. de
Queiroz, eds.
Swalla BJ, Cameron CB, Corley LS, Garey JR (2000) Urochordates are
monophyletic within the deuterostomes. Syst Biol 49: 52-64
Swalla BJ, Smith AB (2008) Deciphering Deuterostome Phylogeny: Molecular,
morphological and palaeontological perspectives. Phil Trans R Soc B 363: 15571568
Van Name WG (1945) The North and South American ascidians. Bull Am Mus Nat
Hist 84: 1476
Wang WF, Namikoshi M (2007) Bioactive nitrogenous metabolites from ascidians.
Heterocycles 74: 53-88
Zeng L, Swalla BJ (2005) Molecular phylogeny of the protochordates: chordate
evolution. Can J Zool 83 (1): 2433
Citation

By downloading or consulting data from this website, the visitor acknowledges

that he/she agrees to the following:

If data are extracted from this website for secondary analysis resulting in a
publication, the website should be cited as follows:

Shenkar, N.; Gittenberger, A.; Lambert, G.; Rius, M.; Moreira Da Rocha, R.; Swalla,
B.J.; Turon, X. (2015) Ascidiacea World Database. Accessed at
http://www.marinespecies.org/ascidiacea on 2015-03-02
If any data constitutes a substantial proportion of the records used in secondary
analyses (i.e. more than 25% of the data are derived from this source, or the
data are essential to arrive at the conclusion of the analysis), the
editors/managers of the database should be contacted. It may be useful to
contact us directly in case there are additional data that may strengthen the
analysis or there are features of the data that are important to consider but may
not have been apparent from the metadata.
Sumber : http://www.marinespecies.org/ascidiacea/ (Diakses 2 Maret
2015)