Order APLOUSOBRANCHIA


Compiler and date details

P. Kott, Queensland Museum, Brisbane, Queensland, Australia

Introduction

The suborder Aplousobranchia (order Enterogona) is characterised by the division of the body into thorax, abdomen and sometimes posterior abdomen. Gonads are unpaired and are in the gut loop (in the abdomen), or posterior to it. Paired epicardial sacs (embryonic endodermal outgrowths of the posterior end of the pharynx) persist in adult zooids, although (with the exception of the Cionidae) their connection with the pharynx is lost and often they are fused. The endodermal tissue of the epicardial sacs is the regenerative tissue involved in repair and replication in all but the family Clavelinidae, in which the regenerative tissue is mesodermal (Berrill 1950). Most aplousobranch families are primarily colonial; although the monotypic Cionidae (genus Ciona) always is solitary and some exceptional solitary species occur also in Diazonidae, Clavelinidae and Euherdmaniidae.

The overall tendency in this suborder to form colonies through replication, interrupts the growth of individual zooids resulting in their progressive size reduction and simplification, and subsequently to the evolution of integrated colonies with zooids arranged in complex cloacal or other systems that enhance colony interactions with the environment (Kott 1989). Internal longitudinal branchial vessels (present in Ciona and Diazonidae) are usually absent although their remains (in the form of papillae) persist in some Protopolyclinidae, Ritterellidae and Polyclinidae. Eggs are fertilised externally only in Cionidae and Diazonidae. Associated with reduction in zooid size and increase in the numbers of replicates of a single genotype, gonads are relatively small in zooids of colonial taxa, the relatively few eggs produced are fertilised internally, and embryos, brooded either in the zooids or in the colonial test, are liberated as tailed larvae.

Australian Aplousobranchia have been revised by Kott (1990, 1992a,b, 2001, 2002, 2003, 2004a-c, 2005). Family level taxa are characterised by their colonial organisation; position, size and form of the gonads; site of fertilisation and brooding of embryos; size of the thorax and condition of the branchial sac including the number of stigmata and transverse branchial vessels; length of the oesophagus and shape of the stomach; and differences in larval adhesive organs and accessory adhesive apparatus.{0244}

 

General References

Berrill, N.J. 1950. The Tunicata. Ray Society Publications 133: 1-354

Kott, P. 1990. The Australian Ascidiacea Pt 2, Aplousobranchia (1). Memoirs of the Queensland Museum 29(1): 1-266

Kott, P. 1992. The Australian Ascidiacea, supplement 2. Memoirs of the Queensland Museum 32(2): 621-655

Kott, P. 1992. The Australian Ascidiacea Pt 3, Aplousobranchia (2). Memoirs of the Queensland Museum 32(2): 377-620

Kott, P. 1998. Tunicata. 51-259, 259–261 (App. I–111), 265–292 (Index) in Wells, A. & Houston, W.W.K. (eds). Zoological Catalogue of Australia. Vol. 34. Hemichordata, Tunicata, Cephalochordata. Melbourne : CSIRO Publishing, Australia 298 pp.

Kott, P. 2001. The Australian Ascidiacea Pt 4, Didemnidae. Memoirs of the Queensland Museum 47(1): 1-410

Kott, P. 2002. Ascidiacea (Tunicata) from Darwin, Northern Territory, Australia. The Beagle, Records of the Museums and Art Galleries of the Northern Territory 18: 19-55

Kott, P. 2003. New syntheses and new species in the Australian Ascidiacea. Journal of Natural History 37: 1611-1653

Kott, P. 2004. Ascidiacea (Tunicata) in Australian waters of the Timor and Arafura Seas. The Beagle, Records of the Museums and Art Galleries of the Northern Territory 20: 37-81

Kott, P. 2004. New and little known species of Didemnidae (Ascidiacea, Tunicata) from Australia (part 1). Journal of Natural History 38(6): 731-774

Kott, P. 2004. New and little known species of Didemnidae (Ascidiacea, Tunicata) from Australia (part 2). Journal of Natural History 38(19): 2455-2526

Kott, P. 2005. New and little known species of Didemnidae (Ascidiacea, Tunicata) from Australia (Part 3). Journal of Natural History 39(26): 2409-2479

 

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Family CIONIDAE


Compiler and date details

P. Kott, Queensland Museum, Brisbane, Queensland, Australia

Introduction

The family Cionidae Lahille, 1887, containing a single genus, Ciona Fleming, 1822, is thought to be the most primitive of the extant ascidian families. It is characterised by solitary habit, a horizontal gut loop posterior to the branchial sac, a persistent opening into the posterior end of the pharynx from each of the pair of endodermal epicardial sacs, a large perforated pharynx (branchial sac) with internal longitudinal vessels, and numerous rows of stigmata.

The family Cionidae, once regarded as a member of the Phlebobranchia due to its large flat branchial sac and internal longitudinal vessels, was proposed as a member of the Aplousobranchia by Kott (1969). Cionids lack the renal vesicles (derived from the epicardial sac) that crowd around the gut in Phlebobranchia and Stolidobranchia (Berrill 1950). Further, the epicardial sacs remain entire as in other Aplousobranchia, and their role in the regeneration of lost or damaged tissue is homologous with their role in the replicative process of colonial Aplousobranchia. Cionids also resemble other aplousobranchs in having the gut behind the thorax (rather than alongside), the body being divided into thorax and abdomen.

This family resembles the aplousobranch family Diazonidae most closely, having a similar soft, elastic, translucent test, large branchial sac with internal longitudinal vessels, an oviparous (externally fertilised) habit, and small larvae with few larval or adult organs other than the simple triradially arranged adhesive organs. Cionidae are separated from the Diazonidae by a horizontal (rather than vertical) gut loop. Some Diazonidae form colonies, constituting a further distinction.

Cionidae are represented in Australia by a single species, Ciona intestinalis (Linnaeus, 1767). Records of C. intestinalis are from ports and harbours in all oceans, including all Australia ports, and many estuaries (Kott 1990). However, records from Australian locations later than 1983, are only from St Vincent Gulf (South Australia), Port Phillip Bay (Victoria) and Port Kembla (New South Wales).

Species of the genus Ciona have been reviewed by Hoshino & Nishikawa (1985). Detailed accounts of morphology are given by Roule (1884) and Millar (1953).

 

General References

Berrill, N.J. 1950. The Tunicata. Ray Society Publications 133: 1-354

Fleming, J. 1822. Philosophy of Zoology; or a general view of the structure, functions, and classifications of animals. Edinburgh : Hurst Vol. 2 618 pp.

Hoshino, Z. & Nishikawa, T. 1985. Taxonomic studies of Ciona intestinalis (L.) and its allies. Publications of the Seto Marine Biological Laboratory 30(1–3): 61-79

Kott, P. 1969. Antarctic Ascidiacea. A monographic account of the known species based on specimens collected under U.S. Government auspices 1947 to 1963. Antarctic Research Series 13: i-xv 1-239

Kott, P. 1990. The Australian Ascidiacea Pt 2, Aplousobranchia (1). Memoirs of the Queensland Museum 29(1): 1-266

Lahille, F. 1886. Sur la classification des Tuniciers. Comptes Rendus (Hebdomadaires) des Séances de l'Academie des Sciences. Série D. Sciences Naturelles 102: 1573-1575

Linnaeus, C. 1767. Systema naturae, per regna tria naturae, secundum classes, ordines, genera, species, cum caracteribus, differentiis, synonymis, locis. Holmiae [= Stockholm] : L. Salvii Tomus I. Regnum Animale(pars 2) Editio duodecima, reformata [3 vols., 1766–68], pp. 533–1327.

Millar, R.H. 1953. Ciona. L.M.B.C. Memoirs on Typical British Marine Plants and Animals 35: 1-123

Roule, L. 1884. Recherches sur les ascidies simples des côtes de Provence 1. Phallusiadées. Annales du Musée d'Histoire Naturelle de Marseille, Zoologie 2(1): 1-270

 

History of changes

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Genus Ciona Fleming, 1822

 

Distribution

States

New South Wales, Queensland, South Australia, Tasmania, Victoria, Western Australia


Extra Distribution Information

Worldwide, rare in polar waters.


Note that conversion of the original AFD map of states, drainage basins and coastal and oceanic zones to IBRA and IMCRA regions may have produced errors. The new maps will be reviewed and corrected as updates occur. The maps may not indicate the entire distribution. See further details below.

IMCRA

Central Western Shelf Province (29), Southwest Shelf Transition (30), Southwest Shelf Province (31), Spencer Gulf Shelf Province (33), Western Bass Strait Shelf Transition (34), Bass Strait Shelf Province (35), Tasmanian Shelf Province (36), Southeast Shelf Transition (37), Central Eastern Shelf Province (38), Central Eastern Shelf Transition (39), Northeast Shelf Province (40), Northeast Shelf Transition (41)

Original AFD Distribution Data

Australian Region

Nearctic Region

Oriental Region

Palaearctic Region

Distribution References

History of changes

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Species Ciona intestinalis (Linnaeus, 1767)

 

Taxonomic Decision for Synonymy

 

Distribution

States

New South Wales, Queensland, South Australia, Tasmania, Victoria, Western Australia


Extra Distribution Information

West coast N America from south Alaska to south California, Europe and Mediterranean Sea, Greenland to southern Massachusetts, Arctic.


IMCRA

Central Western Shelf Province (29), Southwest Shelf Transition (30), Southwest Shelf Province (31), Spencer Gulf Shelf Province (33), Western Bass Strait Shelf Transition (34), Bass Strait Shelf Province (35), Tasmanian Shelf Province (36), Southeast Shelf Transition (37), Central Eastern Shelf Province (38), Central Eastern Shelf Transition (39), Northeast Shelf Province (40), Northeast Shelf Transition (41)

Ecological Descriptors

Benthic, marine.

Extra Ecological Information

On ships' hulls and harbour installations.

 

General References

Kott, P. 1990. The Australian Ascidiacea Pt 2, Aplousobranchia (1). Memoirs of the Queensland Museum 29(1): 1-266

 

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Family CLAVELINIDAE


Compiler and date details

P. Kott, Queensland Museum, Brisbane, Queensland, Australia

Introduction

Species of the family Clavelinidae, Forbes & Hanley 1848, usually are colonial with partially or completely embedded zooids, length about 10 to 100 mm, and with four to 40 rows of stigmata. A few solitary species have been described in which a connection to another zooid has not been detected. In all species, the branchial and atrial apertures have smooth rims, internal longitudinal branchial vessels are not present in the pharynx and the gut loop is of various lengths-sometimes short, with the stomach halfway along the abdomen but sometimes three or four times the length of the thorax with the stomach near the distal end of the loop. Large gonads-a sac-like to tubular ovary and a mass of small pear-shaped male follicles-are enclosed by the gut loop. Strong thoracic longitudinal muscles extend from around the apertures or from each side of the endostyle, and converge to form a band along each side of the abdomen. Zooids are very contractile. The test is soft and transparent and usually does not have sand included. Buds are formed in the isolated terminal ampullae of a posterior abdominal vascular stolon. In this family, the regenerative tissue is mesodermal. Endodermal tissue from either the gut or the epicardial sacs is not involved in replication, although epicardial sacs, as in other aplousobranch families, are present. Eggs are fertilised in the atrial cavity or in the top of the oviduct at the posterior end of the thorax, where embryos are brooded in large numbers (up to 100). They are liberated as tailed larvae. The larval trunk is from 0.5 mm to 1.5 mm long, with wide, shallow adhesive organs, arranged triradially on a robust frontal plate.

Only two genera, Clavelina Savigny, 1816, and the monotypic Nephtheis Gould, 1856, are now recognised in this family (Kott 1990). The genus Clavelina is well represented in Australian tropical and temperate waters, but is not known from the Antarctic. Nephtheis occurs in tropical waters, which are part of its western Pacific range. The Australian fauna, which includes temperature species, appears to have tropical affinities.

 

General References

Forbes, E. & Hanley, S.C.T. 1952. A History of British Mollusca and their Shells. 1848–1952. London : John van Voorst Vol. 1,2 & 4 (appendix) pp. 1-54, 369-376, 244-246.

Gould, A.A. 1856. Mollusca and shells. In, United States Exploring Expedition during the Years 1838–1842 under the Command of Charles Wilkes Atlas: 1–16, pls 1–56. London : Wiley & Putnam.

Kott, P. 1990. The Australian Ascidiacea Pt 2, Aplousobranchia (1). Memoirs of the Queensland Museum 29(1): 1-266

Savigny, J.C. 1816. Recherches anatomiques sur les ascidies composées et sur les ascidies simples—Système de la classe des Ascidies pp. 1–239. In Mémoires sur les Animaux sans Vertèbres, Pt 2. Paris : G. Dufour.

 

History of changes

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Genus Clavelina Savigny, 1816

 

Taxonomic Decision for Synonymy

 

Distribution

States

New South Wales, Northern Territory, Queensland, South Australia, Tasmania, Victoria, Western Australia


Extra Distribution Information

Worldwide except Antarctic waters.


Note that conversion of the original AFD map of states, drainage basins and coastal and oceanic zones to IBRA and IMCRA regions may have produced errors. The new maps will be reviewed and corrected as updates occur. The maps may not indicate the entire distribution. See further details below.

IMCRA

Timor Transition (1), Lord Howe Province (14), Northern Shelf Province (25), Northwest Shelf Transition (26), Northwest Shelf Province (27), Central Western Shelf Transition (28), Central Western Shelf Province (29), Southwest Shelf Transition (30), Southwest Shelf Province (31), Great Australian Bight Shelf Transition (32), Spencer Gulf Shelf Province (33), Western Bass Strait Shelf Transition (34), Bass Strait Shelf Province (35), Tasmanian Shelf Province (36), Southeast Shelf Transition (37), Central Eastern Shelf Province (38), Central Eastern Shelf Transition (39), Northwest Province (4), Northeast Shelf Province (40), Northeast Shelf Transition (41), Central Western Transition (5)

Original AFD Distribution Data

Australian Region

Oriental Region

Palaearctic Region

Distribution References

History of changes

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Species Clavelina amplexa Kott, 2002

 

Distribution

States

Northern Territory


Note that conversion of the original AFD map of states, drainage basins and coastal and oceanic zones to IBRA and IMCRA regions may have produced errors. The new maps will be reviewed and corrected as updates occur. The maps may not indicate the entire distribution. See further details below.

IMCRA

Timor Transition (1), Northern Shelf Province (25), Northwest Shelf Transition (26)

Original AFD Distribution Data

Australian Region

Ecological Descriptors

Benthic, marine.

 

History of changes

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Species Clavelina arafurensis Tokioka, 1952

 

Distribution

States

Northern Territory, Western Australia


Note that conversion of the original AFD map of states, drainage basins and coastal and oceanic zones to IBRA and IMCRA regions may have produced errors. The new maps will be reviewed and corrected as updates occur. The maps may not indicate the entire distribution. See further details below.

IMCRA

Timor Transition (1), Northern Shelf Province (25), Northwest Shelf Transition (26), Northwest Shelf Province (27)

Original AFD Distribution Data

Australian Region

Oriental Region

Ecological Descriptors

Benthic, marine.

Extra Ecological Information

Under ledges.

 

General References

Kott, P. 1990. The Australian Ascidiacea Pt 2, Aplousobranchia (1). Memoirs of the Queensland Museum 29(1): 1-266 [38]

 

History of changes

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Species Clavelina australis (Herdman, 1899)

 

Distribution

States

New South Wales, Queensland, Victoria


Note that conversion of the original AFD map of states, drainage basins and coastal and oceanic zones to IBRA and IMCRA regions may have produced errors. The new maps will be reviewed and corrected as updates occur. The maps may not indicate the entire distribution. See further details below.

IMCRA

Western Bass Strait Shelf Transition (34), Bass Strait Shelf Province (35), Southeast Shelf Transition (37), Central Eastern Shelf Province (38), Central Eastern Shelf Transition (39)

Original AFD Distribution Data

Australian Region

Ecological Descriptors

Benthic, marine.

 

General References

Kott, P. 1990. The Australian Ascidiacea Pt 2, Aplousobranchia (1). Memoirs of the Queensland Museum 29(1): 1-266 [39]

 

History of changes

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Species Clavelina baudinensis Kott, 1957

 

Distribution

States

Western Australia


Note that conversion of the original AFD map of states, drainage basins and coastal and oceanic zones to IBRA and IMCRA regions may have produced errors. The new maps will be reviewed and corrected as updates occur. The maps may not indicate the entire distribution. See further details below.

IMCRA

Central Western Shelf Province (29), Southwest Shelf Transition (30), Southwest Shelf Province (31)

Original AFD Distribution Data

Australian Region

Ecological Descriptors

Benthic, marine.

Extra Ecological Information

2–12 m.

 

General References

Kott, P. 1990. The Australian Ascidiacea Pt 2, Aplousobranchia (1). Memoirs of the Queensland Museum 29(1): 1-266 [41]

 

History of changes

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Species Clavelina cylindrica (Quoy & Gaimard, 1834)

 

Taxonomic Decision for Synonymy

 

Distribution

States

South Australia, Tasmania, Victoria, Western Australia


Note that conversion of the original AFD map of states, drainage basins and coastal and oceanic zones to IBRA and IMCRA regions may have produced errors. The new maps will be reviewed and corrected as updates occur. The maps may not indicate the entire distribution. See further details below.

IMCRA

Northwest Shelf Province (27), Central Western Shelf Transition (28), Central Western Shelf Province (29), Southwest Shelf Transition (30), Southwest Shelf Province (31), Great Australian Bight Shelf Transition (32), Spencer Gulf Shelf Province (33), Western Bass Strait Shelf Transition (34), Bass Strait Shelf Province (35), Tasmanian Shelf Province (36), Southeast Shelf Transition (37), Northwest Province (4), Central Western Transition (5)

Original AFD Distribution Data

Australian Region

Ecological Descriptors

Benthic, marine.

Extra Ecological Information

In storm debris, washed up or floating free in surface currents.

 

General References

Kott, P. 1990. The Australian Ascidiacea Pt 2, Aplousobranchia (1). Memoirs of the Queensland Museum 29(1): 1-266 [42]

 

History of changes

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Species Clavelina dagysa (Kott, 1957)

 

Distribution

States

Western Australia


Note that conversion of the original AFD map of states, drainage basins and coastal and oceanic zones to IBRA and IMCRA regions may have produced errors. The new maps will be reviewed and corrected as updates occur. The maps may not indicate the entire distribution. See further details below.

IMCRA

Central Western Shelf Transition (28), Central Western Shelf Province (29), Southwest Shelf Transition (30), Southwest Shelf Province (31), Northwest Province (4), Central Western Transition (5)

Original AFD Distribution Data

Australian Region

Ecological Descriptors

Benthic, marine.

Extra Ecological Information

To 20 m.

 

General References

Kott, P. 1990. The Australian Ascidiacea Pt 2, Aplousobranchia (1). Memoirs of the Queensland Museum 29(1): 1-266 [45]

 

History of changes

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Species Clavelina fecunda (Sluiter, 1904)

 

Distribution

States

Northern Territory, Queensland, Western Australia


Extra Distribution Information

West Pacific Ocean.


Note that conversion of the original AFD map of states, drainage basins and coastal and oceanic zones to IBRA and IMCRA regions may have produced errors. The new maps will be reviewed and corrected as updates occur. The maps may not indicate the entire distribution. See further details below.

IMCRA

Timor Transition (1), Northern Shelf Province (25), Northwest Shelf Transition (26), Central Western Shelf Province (29), Southwest Shelf Transition (30), Southwest Shelf Province (31), Northeast Shelf Province (40), Northeast Shelf Transition (41)

Original AFD Distribution Data

Australian Region

Oriental Region

Ecological Descriptors

Benthic, marine.

Extra Ecological Information

Black sand, coral, Lithothamnion [Rhodophyta] bank, shallow waters to 20 m.

 

General References

Kott, P. 1990. The Australian Ascidiacea Pt 2, Aplousobranchia (1). Memoirs of the Queensland Museum 29(1): 1-266 [47]

 

History of changes

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Note: the generation of this complete preview for APLOUSOBRANCHIA was cancelled at Clavelina fecunda (Sluiter, 1904) owing to the initation of another complete preview for Demansia torquata (Günther, 1862). Only one complete preview may be executed at a time. Previews were generated for 13 of 571 taxa.