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about this guide | about sea squirts | colour index | species index | species pages | icons | glossary awesome ascidians a guide to the sea squirts of New Zealand Version 1

Mike Page & Michelle Kelly with Blayne Herr

1 about this guide | about sea squirts | colour index | species index | species pages | icons | glossary

about this guide

Sea squirts are amongst the more common marine invertebrates that inhabit our coasts, our harbours, and the depths of our oceans.

AWESOME ASCIDIANS is a fully illustrated invertEguide to the sea squirts of New Zealand. It is designed for New Zealanders like you who live near the sea, dive and snorkel, explore our coasts, make a living from it, and for those who educate and are charged with kaitiakitanga, conservation and management of our marine realm. It is one in a series of electronic guides on New Zealand marine invertebrates that NIWA’s Coasts and Oceans centre is presently developing.

The invertEguide starts with a simple introduction to living sea squirts, followed by a colour index, species index, detailed individual species pages, and finally, icon explanations and a glossary of terms. As new species are discovered and described, new species pages will be added and an updated version of this invertEguide will be made available online.

Each sea squirt species page illustrates and describes features that enable you to differentiate the species from each other. Species are illustrated with high quality images of the in life. As far as possible, we have used characters that can be seen by eye or magnifying glass, and language that is non technical. Information is provided in descriptive text or quick reference icons that convey information without words. Icons are fully explained at the end of this document and a glossary explains unfamiliar terms.

Mike Page is New Zealand’s only professional sea squirt taxonomist; he has a working interest in taxonomy, systematics, chemical ecology and aquaculture. For any ID advice on sea squirts you find, please email Mike your photo’s ... ([email protected])

Remember to check out http://www.niwa.co.nz/coasts-and-oceans/marine-identification-guides-and-fact-sheets

for any updated versions!

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ 2 about this guide | about sea squirts | colour index | species index | species pages | icons | glossary

a typical species page layout

taxonomic name of species taxonomic authority person(s) who first described this species

species species images classification inset images show see species index variations for arrangement

life history icons highlighting whether native or introduced, solitary or colonial, and size (scale bar)

quick id icons highlighting shape, surface detail, habitat, depth range and environment common depth range around New Zealand abundance

distribution section of coastline where species is most information commonly found details on external and internal make notes of where characters and you encountered this habitat species and let us know if you find it at a new location

key taxonomic references it could also be ... some species are difficult to tell apart without careful identification, so check the species listed here to make sure that you have the correct species

3 about this guide | about sea squirts | colour index | species index | species pages | icons | glossary about sea squirts

Sea squirts (ascidians) are amongst the most common fouling animals in ports and harbours around the world. They settle and grow in great abundance on artificial substrates such as wharf piles, seawalls, ship hulls and aquaculture structures. While most native (endemic) species are found in relatively low numbers in intertidal and most subtidal environments around New Zealand, reefs at the entrance of harbours and estuaries with high tidal flow, and cave walls, often support a rich and diverse fauna.

Introduced (invasive) species are usually highly successful, invading in great abundance and often in densities that preclude other species. They have abundant, highly mobile larvae that settle and grow quickly, competing with other species for food and space. The potential consequences of this biology, for the shellfish aquaculture industry in particular, can be serious.

Sea squirts are animals that feed by filtering the water through their body via an inhalant and exhalent siphon. Some are solitary animals, and some live in groups (colonial), some are stalked, and some encrust the substrate. Individual animals are enclosed within a leathery or gelatinous test which can be translucent. Fertilisation may be internal or external with embryos brooded in colonial and some solitary species, followed by a very short- lived free-living larval stage before settlement.

Fish, flatworms, sea urchins and sea stars are the sea squirts’ primary predators, although, in Chile, Japan, Korea , Europe and parts of Aboriginal Australia, some sea squirts are eaten by humans!

4 about this guide | about sea squirts | colour index | species index | species pages | icons | glossary solitary sea squirt Individual animals with an inhalant siphon and an exhalent siphon, often with a thick leathery test that encloses the body of the . colonial sea squirt

Groups of small animals (zooids) are embedded in a gelatinous test as a colony. Zooids can be arranged in circular or linear systems, sharing common exhalent canals and apertures. Other types can have zooids opening independently or on stalks connected to a common basal test.

5 about this guide | about sea squirts | colour index | species index | species pages | icons | glossary colour index

32 30 31 37 eumyota intestinalis Ciona savigny Molgula manhattensis

38 29 16 12 Molgula mortenseni aspersa sigillinoides Didemnum spp complex

17 11 13 25 Eudistoma elongatum Didemnum vexillum Diplosoma listerianum Pseudodistoma cereum

40 27 46 21 Pyura pachydermatina Pseudodistoma opacum Styela canopus Aplidium scabellum

19 9 24 28 Aplidium phortax Clavelina lepadiformis Synoicum occidentalis Pycnoclavella kottae

26 20 43 44 Pseudodistoma Aplidium powelli Asterocarpa humilis Cnemidocarpa bicornuta novaezelandiae

42 26 48 18 Pyura spp complex Pseudodistoma Styela plicata Aplidium benhami novaezelandiae

6 about this guide | about sea squirts | colour index | species index | species pages | icons | glossary colour index

34 36 39 47 Botryllus schlosseri Botryllus tuberatus Microcosmus squamiger Styela clava

20 23 45 33 Aplidium powelli Synoicum kuranui Cnemidocarpa nisiotus Botrylloides leachii

15 35 14 22 Hypsistozoa fasmeriana Botryllus stewartensis Lissoclinum notti Polyclinum novaezelandiae

41 21 33 10 Pyura doppelgangera Aplidium scabellum Botrylloides leachii Didemnum jucundum

7 about this guide | about sea squirts | colour index | species index | species pages | icons | glossary species index

Family Clavelinidae Clavelina lepadiformis ...... 9 Family Didemnidae Didemnum jucundum ...... 10 Didemnum vexillum ...... 11 Didemnum species complex ...... 12 Diplosoma listerianum ...... 13 Lissoclinum notti ...... 14 Family Holozoidae Hypsistozoa fasmeriana ...... 15 Sycozoa sigillinoides ...... 16 Family Polycitoridae Eudistoma elongatum ...... 17 Family Polyclinidae Aplidium benhami ...... 18 Aplidium phortax ...... 19 Aplidium powelli ...... 20 Aplidium scabellum ...... 21

Order Polyclinum novaezelandiae ...... 22 Synoicum kuranui ...... 23 Synoicum occidentalis ...... 24 Family Pseudodistomidae Pseudodistoma cereum ...... 25 Pseudodistoma novaezelandiae ...... 26 Pseudodistoma opacum ...... 27 Family Pycnoclavellidae Pycnoclavella kottae ...... 28

Family ...... 29 Family ...... 30

Class Ciona savigny ...... 31 Phylum Chordata Phylum

Subphylum Tunicata Subphylum Family Corella eumyota ...... 32 Order

Family Botryllidae Botrylloides leachii ...... 33 Family Molgulidae Botryllus schlosseri ...... 34 Botryllus stewartensis ...... 35 Botryllus tuberatus ...... 36 Molgula manhattensis ...... 37 Molgula mortenseni ...... 38 Family Pyuridae Microcosmus squamiger ...... 39 Pyura pachydermatina ...... 40 Pyura doppelgangera ...... 41 Pyura species complex ...... 42 Family Styelidae Asterocarpa humilis ...... 43 Order Stolidobranchia Cnemidocarpa bicornuta ...... 44 Cnemidocarpa nisiotus ...... 45 Styela canopus ...... 46 Stylela clava ...... 47 Styela plicata ...... 48

8 Class Ascidiaceaclassification Order Aplousobranchia Family Clavelinidae Clavelina lepadiformis Pycnoclavella kottaePycnoclavella It couldalsobe………. to Nelson atpresent restricted moorings, Harbour. floating of underside the Fouls filament. either whiteoryellow alight bulbwithaglowing givingtheappearance of visible through thetransparent test. Theendostyleand tentacles arepigmented areclearly opening separately totheoutside. Inhalantgillsac and stomach are joined by athincommonbasal test and protrude as individual heads elongateColonies have multiple zooids a distinctive medusoid shape in which The marine fauna of the British Isles amd Northwest Europe theBritish IslesamdNorthwest . Clarendon Press, Oxford: 627. The marine fauna of Millar,. 1. Scandinavia R.H.(1966)Tunicata. AscidiaceaMarine invertebrates of Hayward P.J., Ryland, J.S. (1947)Introduction toprotozoans andarthropods.

(Müller, 1776) North AtlanticOceanandSeas North 1 cm 9 Class Ascidiaceaclassification Order Aplousobranchia Family Didemnidae Didemnum jucundum Lissoclinum notti Lissoclinum It couldalsobe………. Fiordland regionandStewart Island. Native Australia. Restricted towestern andsouthern atpresenttosouthern Typically bivalves, encrusts coral trees. solitaryascidians and dead black and through exhalent apertures. thecream-coloured interiorcan be seenbelowblack; thetranslucent surface brownare visible. chocolate Surface layer to thetestiscoloureddark of covered incloselyspaced inhalant apertures, occasional exhalent apertures filmy. Test soft, Texture life. in inflated highly appear and thin are Colonies Kott, P. (2001)TheAustralian ascidiacea Part (3),Didemnidae. 4:Aplousobranchia Memoirs of theQueensland Museum , 47(1): 1–407. Memoirs of

Kott, 2001 Australia 3 cm 10 Class Ascidiaceaclassification Order Aplousobranchia Family Didemnidae Didemnum vexillum Encrusting spongesEncrusting Didemnum speciescomplex It couldalsobe………. linesandseacages.structures, marinefarm Can belocallyabundant,fouling floating boathulls, theundersidesof the test;makingitmoregelatinous thanother canals can be seenbelow thesurface. Spiculesare sparse throughout most of exhalent openings are obvious at the end of lobes and a fine open network of Common touch. the to gelatinous yet firm and coloured cream yellowto pale colony algae, encrust hydrozoans, The colonies are tube worms and mussels. extremely longdripping tendrils, sometimesmeterslong. Outgrowths the of or frond-like outgrowths the main colony. can often arise off These can form thisspeciesformextensiveof Colonies sheetsonverticalsurfaces.Cylindrical thisgenus anopaqueappearance.species of spicules can give called spicules. many High abundance of shaped structures most species of The test of Kott, P. (2001)TheAustralian ascidiacea Part (3),Didemnidae. 4:Aplousobranchia Journal of the marine Biological Association of the United Kingdom, 82: 625–628. of themarine Biological Association Journal of Kott, P. (2002)A complex didemnid ascidianfrom Whangamata, New Zealand. Memoirs of theQueenslandMuseum,47(1):1–407. Memoirs of Didemnum is crowded with minute calcite star-

Kott, 2001 Didemnum species. Northern North Pacific North Northern 1 cm 11 Class Ascidiaceaclassification Order Aplousobranchia Family Didemnidae` Didemnum encrusting spongesencrusting notti Lissoclinum Didemnum vexillum It couldalsobe………. lines, pilesaround seacages New andwharf Zealand. Common fouling marinefarm floatingstructures, boathulls, undersidesof thetest thatconnectthezooidstoacommonwaterof circulationsystem. the surrounding test.Thereareusuallycanalsorcavitiesbelow thesurface are usuallysmall(<2.0mm)longandcanbevery difficulttoremove from centre, orthroughout, givingthecolony avery crisp, friableconsistency. Thezooids spicules. Thespiculescanbefound justinthesurfacelayer testswithagelatinous of organisms Test tothinencrustations. calcareous isopaqueduetoaheavycoatingof Colonies canvary greatlyinshapefrom lobateformsovergrowing other fouling appearance. spiculescangivespicules. manythisgenus Highabundanceof anopaque speciesof most species of lambitum (Sluiter, 1900).They oftenvary by onlythesmallestdetails. Thetestof include may complex the in similarity. physical Species their to due field the in distinguished A ‘species complex’ closely related species that cannot be easily is a group of Didemnum incanum (Herdman, 1899), species complex Didemnum is crowded with minute calcite star-shaped called structures Kott, P. (2001)TheAustralian ascidiacea Part (3),Didemnidae. 4:Aplousobranchia Memoirs of theQueensland Museum , 47(1): 1–410. Memoirs of D. (Nott, 1892) and maculatum D. D. 2 cm 12 Class Ascidiaceaclassification Order Aplousobranchia Family Didemnidae Diplosoma listerianum Botrylliodes leachii Botrylliodes It couldalsobe………. and algae shellfish, including surfaces submerged barnacles. of variety a Encrusts substratum asaslimy film. removed from the test by hand, and this species is easilyremoved from the Testapertures. istransparent with small(<2mm)zooids. Zooidsareeasily seen aswhiteorgrey spotsdenselycrowded around large commonexhalent Colony extensive forms individualzooidscan be thingelatinous sheetsinwhich Brewin, B.I. thePortobello (1946) Ascidiansinthevicinityof Marine BiologicalStation,OtagoHarbour. Transactions and Proceedings of the Royal Society of New Zealand,76(2):87–131. theRoyal Society of Transactions andProceedings of Kott, P. (2001)TheAustralian ascidiacea Part (3),Didemnidae. 4:Aplousobranchia Millar, New Zealand: Ascidiacea. R.H.(1982) Themarine faunaof New Zealand Oceanographic Institute Memoir, 85: 1–117. Memoirs of theQueenslandMuseum,47(1):1–410. Memoirs of

(Milne-Edwards, 1841) Cosmopolitan main image: FloorAnthoni 3 cm 13 Class Ascidiaceaclassification Order Aplousobranchia Family Didemnidae Lissoclinum notti Didemnum vexillum Didemnum speciescomplex It couldalsobe………. Common onshallow pilesandaquaculturestructures. subtidalreefs, wharf Test Colourinlifeisopaque cream, brown is papery and easily torn. or violet. thecolony,surface andatthebaseof andhave distinctive burr-shaped ends. organisms.lobes formedonencrusted Spiculesarefound in2layers; atthe exhalent evenly apertures distributedthroughout the colony orontheapex of fragile. Zooidsarenotinmarked systems,but therearerelatively large,common and very easilytorn thin,encrusting, <2mmthick, Colonies characteristically Brewin B.I. New Zealand.Part (1958)Ascidiansof theHauraki Gulf. 12.Ascidiansof Part 3. Transactions and Proceedings of the Royal Society of New Zealand , 85(3): 455–458. theRoyal Societyof Transactions andProceedings of

Brewin, 1958 Western SouthPacific 2 cm 14 Class Ascidiaceaclassification Order Aplousobranchia Family Holozoidae Hypsistozoa fasmeriana Aplidium benhami It couldalsobe………. moderate exposure.areas of in open in depth m structures 20 to down found artificial be can flow. Colonies tidal on high with harbours and reefs coastal shallow in common Most 20 –30cmindiameter. pink toviolet. Colourinlifeisusuallyfuchsia linear andscatteredover thebody. groupscan oftenoccurinpatchy Colonies are in parallel systemsaround numerous large exhalent apertures, systems Zooids shaped. Stalks areoftennotvisible. Softand gelatinous tothetouch. individually to the substrate. Bodyattached isoften button or mushroom- body, ovoid larger much a with topped stalk fleshy short a of Colonyconsists Brewin, B.I. thePortobello (1946)Ascidians in thevicinityof MarineBiologicalStation, OtagoHarbour. Transactions and Proceedings of the Royal Society of New Zealand,76(2):87–131. theRoyal Societyof Transactions andProceedings of Millar, New Zealand: Ascidiacea. R.H.(1982) Themarine faunaof New Zealand Oceanographic Institute Memoir, 85: 1–117.

(Michaelsen, 1924) (Michaelsen, 5 cm 15 Class Ascidiaceaclassification Order Aplousobranchia Family Holozoidae Sycozoa sigillinoides Chatham Islands, andoccursasfarsouthAntarctica. This species has a wide geographic range within New Zealand, including the colony. Colourinlife translucent creamtowhitewithdots. white dots, the connecting to asinglecommonexhalent atthetopof aperture be seenthroughas thetestaslineardoublerows visible inhalantapertures of can Zooids gelatinous. soft, Texture stalks. flexible on heads ovoid Elongate Kott, P. (1990) The Australian Ascidiacea,Part 2,Aplousobranhia (1). Memoirs of theQueensland Museum, 29(1): 1–226. Memoirs of

Lesson, 1830 main andinsetimages: CrispinMiddleton Circum polar 2 cm 16 Class Ascidiaceaclassification Order Aplousobranchia Family Polycitoridae Eudistoma elongatum Eudistoma elongatum Restricted at present to the far north of the North Island. theNorth Restricted of atpresenttothefarnorth racks, mangrove roots, rocky shorelineand on shellsembedded in mud. Species occurlocallyinhighabundance in shelteredbays, growing onoyster 10 mm)creambuds, re-growing thefollowing springtolarger colonies. orange withdeveloping embryos. Coloniesregress and over-winter as small (c. opening separately to the outside. When reproductive, the zooids become overall. Zooids appear as light brown with two tiny specks, each apertures wart-like short with processes.side Test firm gelatinousand smooth is touch, to pendulous colonies tapering to a smooth stalk,sometimes Long cylindrical Kott, P. (1990)TheAustralian Ascidiacea, Part 2,Aplousobranhia (1). Memoirs of theQueensland Museum , 29(1): 1–226. Memoirs of

(Herdman, 1886) Australia main image: SeanHandley 6 cm 17 Class Ascidiaceaclassification Order Aplousobranchia Family Polyclinidae Aplidium benhami Hypsistozoa fasmeriana It couldalsobe………. Island, andFiordland. around CookStrait, Kaikoura, Portobello Peninsula, ChathamIslands, Stewart andshallowabundant on intertidal subtidalreefsdown to10mandfound fronds piles. and holdfasts in thesubtidal. Can be found Locally on wharf rocks, intertidal overgrowsLives on the undersides of bryozoans, seaweed reddish orange tobrilliantcrimson. flow.tidal high life, in of deep Colour areas in piles wharf on observed been ring that is visibleintheanimalslife. Larger, morelobate specimens have in star- zooids inhalantsiphonshaveshaped to circular systems. a distinctive Rimsof white containing heads cauliflower-shaped with stalked Colonies Brewin, B.I. thePortobello (1946)Ascidians in thevicinityof MarineBiologicalStation, OtagoHarbour. Transactions and Proceedings of the Royal Society of New Zealand,76(2):87–131. theRoyal Societyof Transactions andProceedings of Millar R.H. (1982) The marine fauna of New Zealand: Ascidiacea. Millar R.H. (1982)Themarine faunaof New Zealand Oceanographic Institute Memoir, 85: 1–117.

(Brewin, 1946) 6 cm 18 Class Ascidiaceaclassification Order Aplousobranchia Family Polyclinidae Aplidium phortax Aplidium phortax Aplidium powelli It couldalsobe………. bays, andharbours. Very inports, pilesandaquaculture structures commonspeciesfouling wharf to pink. lobesonthecolony.of Colourinlifevaries from translucent cream, to lighttan, Common exhalent are indistinct, but are often situated on theapex apertures formmeandering Zooids double-rowed, systems, along obvious attimesbranching subsurface canals. colonies. gelatinous firm, fleshy, spherical, Large Brewin, B.I. thePortobello (1946)Ascidians in thevicinityof MarineBiologicalStation, OtagoHarbour. Transactions and Proceedings of the Royal Society of New Zealand,76(2):87–131. theRoyal Societyof Transactions andProceedings of Millar R.H. (1982) The marine fauna of New Zealand: Ascidiacea. Millar R.H. (1982)Themarine faunaof New Zealand Oceanographic Institute Memoir, 85: 1–117.

(Michaelson, 1924) (Michaelson, 3 cm 19 Class Ascidiaceaclassification Order Aplousobranchia Family Polyclinidae Aplidium powelli Aplidium phortax It couldalsobe………. Island. theNorth coastline of Most commononsheltereddeepreefsdown eastern to30 m depth onthenorth and translucent. throughusually lightpink,butcanalsobeyellow theoutertest.Colourinlife visible zooidinhalant apertures of pattern and the distinctive branched Easily tofirm distinguished the touch. by the many common exhalent apertures yet Gelatinous high. cm 10 to up colonies large,fleshy,multi-lobed in Grows Brewin B.I. New Zealand. Part (1958)Ascidiansof theHauraki Gulf. 12.Ascidians of Part 3. Transactions and Proceedings of the Royal Society of New Zealand,85(3):45–458. theRoyal Societyof Transactions andProceedings of Millar R.H. (1982) The marine fauna of New Zealand:Ascidiacea. Millar R.H. (1982)Themarine faunaof

(Brewin, 1958) New Zealand Oceanographic Institute Memoir, 85: 1–117. 8 cm 20 Class Ascidiaceaclassification Order Aplousobranchia Family Polyclinidae Aplidium scabellum Aplidium scabellum Botryllus stewartensis Botryllus It couldalsobe………. IslandandChathamIslands. North recorded from eastern north itscryptic habitat in thesand. It has been but is oftenoverlooked because of reefs, rocky between seafloor sandy the on common relatively is species This colony heads. cases worm that protrude betweenare oftenassociated with polychaete the systems orcommon exhalent are visible inthecolony apertures inlife. Colonies theheadslessso, revealingof dullorange zooids beneaththesurface. No approximately 60 mm high. The test is heavilyinvested with sand; the tops Colonies are composed of closely packed flat-topped heads in a large mound Michaelsen W.Michaelsen von Inseln. (1924) Ascidiae Krikobranchiae NueseelanddenChatham- unddenAuckland- Videnskabelige Meddelelser fra Dansk naturhistorisk Forening fraDansknaturhistorisk Meddelelser Videnskabelige iKobenhavn, 77:263–434. Millar R.H. (1982) The marine fauna of New Zealand: Ascidiacea. Millar R.H. (1982)Themarine faunaof New Zealand Oceanographic Institute Memoir, 85: 1–117.

(Michaelsen, 1924) (Michaelsen, 3 cm 21 Class Ascidiaceaclassification Order Aplousobranchia Family Polyclinidae Polyclinum novaezelandiae Aplidium scabellum It couldalsobe………. substrata. artificial Found insheltered,sandy subtidal environments growing onnatural and an iridescentblue. gelatinous.is gray Colourinlife lilac-gray; thecommonsiphonsis theinsideof The surface is lightlycoated with sand and the inside soft and apertures. common exhalent siphons interspersed withsmallbutobvious inhalant Distinctive large cushion-shapedcolonieswithnumerous raised turret-shaped Brewin, B.I. New Zealand.Part (1958)Ascidians of the Stewart 11.Ascidiansof Islandregion. Transactions of the Royal Society of New Zealand,85(3): 439–453. theRoyal Society of Transactions of

Brewin, 1958 3 cm 22 Class Ascidiaceaclassification Order Aplousobranchia Family Polyclinidae Synoicum kuranui Synoicum kuranui Hypsistozoa fasmeriana It couldalsobe………. tidal flow, down to40m. Locally abundant on low sandy deepreefsandinshelteredharbourswithhigh colonies may grow intolongsausages upto1.2mlong. circular systems around common exhalent apertures. In areas of high tidal flow howeversand. Uptoeightzooidsarearranged attimesoverlainwith in up to a dozen colonies. Coloniesare smooth and gelatinous,in clustersof Mushroom tobutton-shaped colonies withoutanobvious stalk,commonlyoccurs Brewin, B.I. New Zealand.Part (1950)Ascidians of GreatBarrierIsland. 5.Ascidiansfrom theEastCoastof Transactions of the Royal Society of New Zealand,78(2–3): 354–362. theRoyal Society of Transactions of

Brewin, 1950 1 cm 23 Class Ascidiaceaclassification Order Aplousobranchia Family Polyclinidae Synoicum occidentalis subtidal habitats. Alsoknown from trawls ontheshelf-breakto300m. Infrequently found over-growing coralline paint on rock walls orshallow yellowpatchy orange, translucent. under theopaque test. Commonexhalent notvisible. Colourinlife apertures no base.Gelatinous, the at sediment incorporated fused in thetest.Zooids incircularsystemsattimesvisible cushions or several cushion flat Smooth Millar, New Zealand:Ascidiacea. R.H.(1982)The marinefaunaof New Zealand Oceanographic InstituteMemoir , 85:1-117.

Millar, 1982 South West Pacific Ocean 4 cm 24 Class Ascidiaceaclassification Order Aplousobranchia Family Pseudodistomidae Pseudodistoma cereum Most commonincoastaldeepreefsdown to30m. apparent on theapex smalllumps. Colourinlifecream, yellow of orfaintpink. exterior. Zooidsopenseparately totheexterior; zooidare openingsfor each onthe appearance.but softtothetouch andcartilaginous, Fingers arestiff Small testcellsmay be numerous insomecoloniesgivingtheman opaque encrusting. more become and flatten but may they mmhigh, environments 50 energy to high in up stalks on fingers rounded fat form usually Colonies Michaelsen W.Michaelsen von Inseln. (1924)Ascidiae Krikobranchiae Nueseeland den Chatham- undden Auckland- Brewin, B.I. New Zealand.Part (1958)Ascidians of the Stewart 11.Ascidiansof Islandregion. Videnskabelige Meddelelser fra Dansk naturhistorisk Forening fra Dansk naturhistorisk Meddelelser Videnskabelige i Kobenhavn, 77: 263–434. Transactions of the Royal Society of New Zealand , 85(3):439–453. the Royal Society of Transactions of

Michaelson, 1924 Michaelson, 4 cm 25 Class Ascidiaceaclassification Order Aplousobranchia Family Pseudodistomidae Pseudodistoma novaezelandiae Aplidium benhami It couldalsobe………. Islands.the Kermadec Island. This species is also common around the North coast of eastern the north common inexposed shallow coastal reefs, generally down to10 m depth on groupsColonies occur inpatchy (20–30 cm diameter) on coralline paint.Very orange.Colour inlifebrightpeach Inhalant and individual zooidsopendirectlyonsurface.exhalent siphonsof Stalkedan ovoid colonialascidianwith tosphericalhead,testclear,no sand. Brewin, B.I. New Zealand.Part (1950)Ascidians of GreatBarrierIsland. 5.Ascidiansfrom theEastCoastof Transactions of the Royal Society of New Zealand,78(2–3): 354–362. theRoyal Society of Transactions of

(Brewin, 1950) 2 cm 26 Class Ascidiaceaclassification Order Aplousobranchia Family Pseudodistomidae Pseudodistoma opacum Pseudodistoma cereum It couldalsobe………. epiphytic redalgae growing onthesurface. boulders. Coloniescan have intertidal Typically theundersidesof encrusts openingontothesurface. Colourinlifetantodullpeach. apertures membrane that basal gives rise to small flat-topped lobes. ofaspreading Test semi-transparent with yellow zooids, up made mats irregular fleshy Thick Brewin, B.I. New Zealand.Part (1950)Ascidians of GreatBarrierIsland. 5.Ascidiansfrom theEastCoastof Transactions of the Royal Society of New Zealand,78(2–3): 354–362. theRoyal Society of Transactions of

(Brewin, 1950) main andinsetimages: BrentCopp 10 cm 27 Class Ascidiaceaclassification Order Aplousobranchia Family Pycnoclavellidae Pycnoclavella kottae Clavelina lepadiformis Clavelina It couldalsobe………. Three KingsIslands. exposure, where it can be locally abundant. Restricted Cape and to North Found occasionally on walls high overhangs and the edge in areas of of colony head.Colourinlifeis goldandslightlytranslucent onthesiphons. opening halfway down and an atrial (exhalent) the opening at the top of narrow basal mat. Stalksarelongand slender; heads have anoral (inhalant) A distinctive colonialascidian that hasnumerous pod-like heads arisingfrom a Millar, R.H.(1960) Ascidiacea.

Millar, 1980 Discovery Reports, 30:1–160. South West Pacific Ocean 10 cm 28 Class Ascidiaceaclassification Order Phlebobranchia Family Ascidiidae Ascidiella aspersa Ascidiella aspersa Corella eumyota It couldalsobe………. theSouthIsland. structures, inbothmarineandestuarineenvironments of This speciesisfound onshallow pilesand submerged subtidal rock, wharf tentacles aresmooth. and covered in smallbumps. The gill slitsare elongate, and not folded, the exhalent siphononethirdto halfway down oneside. Testtranslucent, isthin, Sack-like ovoid theanimal,andan bodyan inhalantsiphonatthetopof with Brewin, B.I. thePortobello (1946) Ascidiansinthevicinityof Marine BiologicalStation,OtagoHarbour. Transactions and Proceedings of the Royal Society of New Zealand,76(2):87–131. theRoyal Society of Transactions andProceedings of Kott, P. (1985)TheAustralian andStolidobranchia. Ascidiacea 1.Phlebobranchia Millar R.H. (1982) The marine fauna of New Zealand: Ascidiacea. Millar R.H. (1982)Themarine faunaof New Zealand Oceanographic Institute Memoir, 85: 1–117.

(Müller, 1776) Memoirs of theQueenslandMuseum,23:440. Memoirs of Cosmopolitan inset image:Gittenberger Arjan main image: DennisGordon 2 cm 29 Class Ascidiaceaclassification Order Phlebobranchia Family Cionidae Ciona intestinalis Ciona savigny It couldalsobe………. pontoons around New Zealand. pilesand Often found inhighabundance on aquaculture structures, wharf thebody sideof wall.found oneach folded, and tentacles are smooth. Sixbroad bands are longitudinal muscle and lemon yellow pigment spots on siphon rim. Gillslitsare elongate, not end, anterior flexible, the at pigment green light gelatinous, with transparent, Body elongate, tapering towards two closely spaced siphons. Test is soft, hasorangesavigny pigmentspotsonthesiphonrim. Ciona Two Cionaareknown toco-occur speciesof in New and harbours. Zealandports intestinalis haslemonyellow pigmentspotsonthesiphonrimwhileCiona Brewin B.I. New Zealand. Part (1950)Ascidiansof IV. Christchurch. Ascidiansinthevicinityof Transactions and Proceedings of the Royal Society of New Zealand , 78(2-3): 344–353. theRoyal Societyof Transactions andProceedings of

(Linnaeus, 1767) Cosmopolitan main image: Keith Hiscock 4 cm 30 Class Ascidiacea Class Phlebobranchia Family Cionidae Herdman, W.A. (1882)Report onthe Tunicata collectedduring thevoyage H.M.S. Challenger of duringthe years 1873–1876, Part 1,Ascidiae simplices. classification Ciona savigny Ciona savigny Ciona intestinalis It couldalsobe………. pontoons around New Zealand. pilesand Often found inhighabundance on aquaculture structures, wharf the body side of wall.Six broad bands are found longitudinal muscle on each on thebody wall.slits areelongate, Gill notfolded, and tentacles are smooth. flecks pigment white yellowor and rim, siphon on spots pigment orange and end, anterior flexible, the at pigment green light gelatinous, with transparent, Body elongate, tapering towards two closely spaced siphons. Test is soft, hasorangesavigny pigmentspotsonthesiphonrim. Ciona intestinalis Two Cionaareknown toco-occur speciesof in New and harbours. Zealandports Brewin B.I. New Zealand. Part (1950)Ascidiansof IV. Christchurch. Ascidiansinthevicinityof Transactions and Proceedings of the Royal Society of New Zealand,78(2-3):344–353. theRoyal Societyof Transactions andProceedings of haslemonyellow pigmentspotsonthesiphonrimwhileCiona

Herdman, 1882 Zoology of the Challenger Expedition, 6(17): 1–296. Zoology of North PacificNorth main image: ChrisWoods 4 cm 31 Class Ascidiaceaclassification Order Phlebobranchia Family Corellidae Corella eumyota Molgula spp. Molgula spp. Ascidiella aspersa It couldalsobe………. piles, ropesto wharf andothersubmerged around structures New Zealand. Prefers calmprotected waters, found inshallow subtidalenvironments attached inhalantsiphons. but somehave brightpeach inlife,through Gutandgonads oftenvisible cartilaginous. thetest.Colourless spiral,not folded, gills andoral tentaclessmooth.Test transparent, smooth, smaller exhalent siphon side, individualsareoftenfound in groups. animal, Inhalantsiphonat top of Body oval toelongate, tothesubstrate onright laterally compressed,attached Millar, SouthAfrican ascidians. descriptions of R.H.(1962) Further of theway⅓ of down thebody. the side of Gillslits

Traustedt, 1882 Annals of the SouthAfrican Museum,46(7): 113–221. Annals of inset image: Natural HistoryMuseum,UK Cosmopolitan main image: JudithOakley 3 cm 32 Class Ascidiaceaclassification Order Stolidobranchia Family Botryllidae Botrylloides leachii Botrylloides Botryllus schlosseri Botryllus It couldalsobe………. ships. harbours throughout New Zealand.May have beenintroduced by sailing early piles, moorings, andisvery and and wharf Encrusts jetties commoninports thecolony.the borderof the test between the zooid systems bodies and are visible near the surface of orange andcream.Thetestistransparent, softandgelatinous. Smallgranular Colour inlifeishighlyvariable, ranging from typicallypurpletogreen Systemsconnect to numerousinhalant apertures. commonexhalent apertures. lightpigmentationaround zooidsareusuallyobvious the because of systems of overgrowing otherspecies, givingcoloniesa lobate appearance. Parallel andupto20cmdiameter,Colonies areencrusting, about3-5mmthick often Brewin B.I. thePortobello (1946)Ascidians inthevicinityof MarineBiologicalStation, OtagoHarbour. Transactions and Proceedings of the Royal Society of New Zealand,76(2):87–131. theRoyal Society of Transactions andProceedings of Kott P. (1985) TheAustralian andStolidobranchia. Ascidiacea1.Phlebobranchia Millar R.H. (1982) The marine fauna of New Zealand: Ascidiacea. Millar R.H. (1982)Themarine faunaof New Zealand Oceanographic Institute Memoir, 85: 1–117.

(Savigny, 1816) Memoirs of theQueenslandMuseum,23:440. Memoirs of North AtlanticOceanandSeas North 5 cm 33 Class Ascidiaceaclassification Order Stolidobranchia Family Molgulidae Botryllus schlosseri Botryllus Botrylloides leachii Botrylloides It couldalsobe………. been introduced by sailingships. early and is very and harbours throughout commoninports New Zealand. May have moorings, piles, mooringpontoons,Encrusts jetties, and wharf undersidesof life, coloniescan vary widely incolour, but are usually orange, green or purple. distinguished by circularzooidsystemsaround commonexhalent In apertures. , butcanbe leachii oftenco-occurring with Botrylloides 3 mmthick, Colonies Kott P. (1985) TheAustralian andStolidobranchia. Ascidiacea1.Phlebobranchia

(Pallas, 1766) Memoirs of theQueensland Museum , 23:440. Memoirs of Cosmopolitan inset image: ChrisWoods 1 cm 34 Class Ascidiaceaclassification Order Stolidobranchia Family Molgulidae Botryllus stewartensisBotryllus Botrylloides leachii Botrylloides It couldalsobe………. New Zealand. Found occasionally on reefs in sheltered coves and on walls in fiords in Southern present, sedimentisconfinedtotheoutertest. is sandyColour inlife or cream.Thetestisdelicateandsoft,when violet, zooids. Morphology and colour inlifevary with sedimentlevels and exposure. lobehasacentral commonexhalent withacircleof basal mat. Each aperture Colonies arelow sandy lobesapproximately 10 mm high,tightlypacked ona Brewin B.I. New Zealand. Part (1958)Ascidiansof theStewart 11.Ascidians of Islandregion. Kott P. (1985)The Australian and Stolidobranchia. Ascidiacea1.Phlebobranchia Transactions of the Royal Society of NZ,85(3):439–453. the Royal Societyof Transactions of Memoirs of the Queensland Museum, 23: 440. Memoirs of

Brewin, 1958 Australia 1 cm 35 Class Ascidiaceaclassification Order Stolidobranchia Family Molgulidae Botryllus tuberatus tuberatus Botryllus Botryllus schlosseri Botryllus It couldalsobe………. and Nelsonharbour. and shallowintertidal subtidalenvironments around Wellington southcoast andoysters asmussels in otherorganismsThis speciestypicallyencrusts such test. shaped appearance. Colourinlifebrilliantorange and cream in a transparent flower- a colony the giving systems, circular spaced widely in arranged are Colonies are small, delicate, very thinand transparent. Bright orange zooids Kott P. (1985) TheAustralian andStolidobranchia. Ascidiacea1.Phlebobranchia

Ritter &Forsyth, 1917 Memoirs of theQueensland Museum , 23:440. Memoirs of Circum equitorial 0.5 cm 36 Class Ascidiaceaclassification Order Stolidobranchia Family Molgulidae Molgula manhattensis Molgula manhattensis Molgula mortenseni It couldalsobe………. the seafloor. PresentlyrestrictedtotheManukau Harbour. Tolerant highsedimentand low of salinity. May occur in large aggregations on cream. longer atthebase, formingroot-like processes.translucent Colour inlife to thetest;hairsare usually hairsonthe surface of Sediment adherestoshort looks like cellophane.slits spiral, Gill gillsfolded, and oral tentaclesbranched. close together ontheuppersurface. Test semi-translucent andrelatively tough, Body small,spherical to oval, inhalantand exhalent siphonsrelatively longand Kott, P. (1985)TheAustralian and Stolidobranchia. Ascidiacea I.Phlebobranchia Memoirs of theQueensland Museum , 23:440. Memoirs of

(De Kay, 1843) Cosmopolitan inset image: MisjelDeclee main image: Keith Hiscock 4 cm 37 Class Ascidiaceaclassification Order Stolidobranchia Family Molgulidae Molgula mortenseni Molgula mortenseni Molgula manhattensis It couldalsobe………. shells andotherlarge solitaryascidians. asharbour seabeds around Newsuch Zealand.Oftenepizoic,livingamong Can befound individualsinhighsedimentenvironments inaggregations of to cream. the body wall. adherent sand, otherwise translucent white Colour inlifethat of Akidneyfolded, through and tentacles branched. can often beseenclearly gills spiral, slits Gill grains. sand fine with coated flexible, thin, test surface, Body ovoid toglobular, 1–2 cm diameter, closetogether apertures onupper Brewin, B.I. New Zealand.Part (1951)Ascidians of theHauraki Gulf. 6.Ascidiansof Part 2. Transactions and Proceedings of the Royal Society of New Zealand,79(1):104–113. theRoyal Societyof Transactions andProceedings of Kott, P. (1985) The Australian andStolidobranchia. AscidiaceaI. Phlebobranchia

Memoirs of the Queensland Museum, 23: 440. Memoirs of (Michaelsen, 1922) (Michaelsen, illustration sourcesfrom: Brewin (1951) South West Pacific Ocean 0.5 cm

38 image credit: illustration sourced from Brewin (1951) Brewin from sourced illustration credit: image Class Ascidiaceaclassification Order Stolidobranchia Family Pyuridae Microcosmus squamigerMicrocosmus juvenile Cnemidocarpanisiotus Pyura speciescomplex It couldalsobe………. and exposed locations, New predominantlyaround Zealand. northern Usually occurs in large aggregates on rock, concrete and cave walls insheltered in lifeorange withmaroon onwrinkles, orange andmaroon-striped siphons. left gonad crosses the gut loop. overbranched, Colour the descending limb of and tough, attimeshard and occasionally brittle.slits simple, Gill tentacles Body elongate wart-like to oval, on short apertures siphons. Test leathery Kott, P. (1985)TheAustralian and Stolidobranchia. Ascidiacea I.Phlebobranchia Memoirs of theQueensland Museum , 23:440. Memoirs of

Michaelsen, 1927 Michaelsen, Cosmopolitan main image: Griffiths Charles 1 cm 39 Class Ascidiaceaclassification Order Stolidobranchia Family Pyuridae Pyura pachydermatina pachydermatina Pyura Styela clava It couldalsobe………. 30 m. than greater to down intertidal the from floor sea the on forests dense form open coast, and in harbours with high tidal flow. In optimal conditions they can Sea tulipsgrow inhighenergy New environments insouthern Zealandonthe in lifecream,tinged withmaroon alonglongitudinalridges andinsiphons. thebody. sideof oneach gonads in pairedblocks Colour tentacles branched, that are more pronounced in smallerindividuals. Gillslitselongate, gillsfolded, thehead smooth, thrownSurface of undulatinglongitudinalridges intothick wrinkled. Large inhalant and exhalent head. siphons are obvious at the top of head. Stalks generally smoothorhorizontallywrinkled,never longitudinally distinguished byThe ‘seatulip’iseasily itslongleatherystalkandbulbous Brewin, B.I. thePortobello (1946) Ascidiansinthevicinityof Marine BiologicalStation,OtagoHarbour. Transactions and Proceedings of the Royal Society of New Zealand,76(2):87–131. theRoyal Societyof Transactions andProceedings of Millar, New Zealand:Ascidiacea. R.H.(1982)The marinefaunaof New Zealand Oceanographic InstituteMemoir , 85:1–117.

(Herdman, 1881) main image: FloorAnthoni 6 cm 40 Class Ascidiaceaclassification Order Stolidobranchia Family Pyuridae Rius, M.,Teske, P.R. (2013) Crypticdiversity incoastalAustralasia: habitat-forming siblingspecies. amorphologicalandmitonuclear genetic analysisof Pyura doppelgangera Pyura Pyura praeputialis It couldalsobe………. to 12m.Restricted atpresenttotheFar North. platforms, sometimesoccupying 100% cover. May be found subtidally down Individuals canbevery large andoftenformdense aggregates onintertidal are brightreddishorange. the sandy,Colour inlifeis that of test, may encrusted be quite green, siphons wall visiblefrom exterior. Gillslitselongate, gillsfolded, tentacles branched. or cross-shapedcruciform siphons are visible by the bright reddish orange body tough, thick, cartilaginous, coated with sand and algal filaments. When inflated, thebody.representing siphonssetinthedepressed upper surfaceof Test Large, solitary, stumpy, ascidian with two large chalice-shaped mounds Zoological Journal of theLinneanSociety , 168:597-611. Zoological Journal of

Rius &Teske, 2013 main andinsetimages: Roger Grace Australasia 7 cm 41 Class Ascidiaceaclassification Order Stolidobranchia Family Pyuridae Pyura Pyura Microcosmus squamiger Cnemidocarpa nisiotus It couldalsobe…… piles around New Zealand. Found growing toshelldebris andfouling ontheseabedattached wharf are oftenpigmentedwithdeeppurple, andsiphonrimstripedwhiteorpeach. orange test.Siphons withdarker burnt onraised sectionsof in lifepalepeach thebody sideof wallgonad oneach may bearranged Colour inpairedblocks. warty, finely wrinkled. Gill slits elongate, gills folded, tentacles branched. A long thebody.close together or ateitherendof Test tough,leathery, deeplyfurrowed, Body elongate, oval siphons set reasonably to banana-shaped with long muscular (Sluiter,subuculata P. 1900)and Brewin, 1946. cancellata the smallestdetails. Speciesin thecomplex includeP. Brewin, rugata 1948, P. physical their to due field the in similarity.distinguished They often vary only by A ‘speciescomplex’ closelyrelatedspecies that cannot beeasily is agroup of species complex Brewin, B.I. thePortobello (1946)Ascidians in thevicinityof MarineBiologicalStation, OtagoHarbour. Transactions and Proceedings of the Royal Society of New Zealand,76(2):87–131. theRoyal Societyof Transactions andProceedings of Millar, New Zealand: Ascidiacea. R.H.(1982) Themarine faunaof New Zealand Oceanographic Institute Memoir, 85: 1–117. main andinsetimages: AnneFrijsinger &MatVestjens 4 cm 42 Class Ascidiaceaclassification Order Stolidobranchia Family Styelidae Asterocarpa humilis Asterocarpa bivalves around New Zealand. pilesandfoulingThe speciesoccursinthesubtidalunderboulders, onwharf thebodyeither sideof wall. are elongate, tentacles smooth, and gonads appear in star-shaped clusters on and algae, and is translucent and usually grey to buff-coloured. The gill slits bands. The test is smooth and flexible, at times encrusted with sponges, hydroids Body havelongitudinal globularwithmaroon eightwhiteinternal siphonswhich Brewin B.I. thePortobello (1946)Ascidians inthevicinityof MarineBiologicalStation, OtagoHarbour. Transactions and Proceedings of the Royal Society of New Zealand,76(2):87–131. theRoyal Society of Transactions andProceedings of Millar R.H. (1982) The marine fauna of New Zealand: Ascidiacea. Millar R.H. (1982)Themarine faunaof Kott P. (2). (1992) TheAustralian ascidiacea3.Aplousobranchia New Zealand Oceanographic Institute Memoir, 85: 1–117. Memoirs of theQueenslandMuseum,32(2):375–620. Memoirs of

(Heller, 1878) Tasmania /Indo-Pacific main image:Schories Dirk 0.5 cm 43 Class Ascidiaceaclassification Order Stolidobranchia Family Styelidae Cnemidocarpa bicornuta Cnemidocarpa nisiotus It couldalsobe………. Cnemidocarpa nisiotus. abundant on shallowpiles. Generally co-occurs with reefsandwharf Very common in ports, harbours, and coastal environments. May belocally hydrozoans, bryozoans andfilamentousalgae. light orangeColour inlife wrinkled. longitudinally tocream. Oftenfouled with tubular, sometimesbent backwards end.Test at their terminal leatheryand tothebody wallare smooth.Gonads attached underthegillsac are longand pigment on theorange siphonlining. Gillslitsare elongate, folded, tentacles are covered inwarty processes. Characterised by magenta four bandsof siphon nearer to the base. Siphons are separated by a distinctive saddle, and Banana-shaped with the inhalant siphon usually at the top, and the exhalent Brewin B.I. thePortobello (1946)Ascidians inthevicinityof MarineBiologicalStation, OtagoHarbour. Transactions and Proceedings of the Royal Society of New Zealand,76(2):87–131. theRoyal Societyof Transactions andProceedings of Millar, New Zealand:Ascidiacea. R.H.(1982)The marinefaunaof New Zealand Oceanographic InstituteMemoir , 85:1–117.

(Sluiter, 1900) 2 cm 44 Class Ascidiaceaclassification Order Stolidobranchia Family Styelidae Cnemidocarpa nisiotus Cnemidocarpa bicornuta It couldalsobe………. occurs with Can belocallyabundant on shallow piles. Generally co- reefsand wharf Very common in ports, harbours and coastal environments around New Zealand. siphons. maroon siphonliningsand four pale yellow to white longitudinalbands in the flask-shaped thebody sideof wall. brown three gonads on each Colourinlifedark tosilty, with are there and smooth, are tentacles slits, elongate with leathery, usuallyfouled withhydroids, bryozoans andalgae.are folded Gills Body wrinkled, large irregular warty processes occur around the siphons. Test Oval shaped body with two siphons approximately a body half length apart. Brewin B.I. the Portobello (1946)Ascidiansinthevicinity of Marine BiologicalStation,OtagoHarbour. Cnemidocarpa bicornuta. Transactions and Proceedings of the Royal Society of New Zealand,76(2):87–131. theRoyal Societyof Transactions andProceedings of Millar, New Zealand: Ascidiacea. R.H.(1982) Themarine faunaof New Zealand Oceanographic Institute Memoir, 85: 1–117.

(Sluiter, 1900) main image: FloorAnthoni 1.5 cm 45 Class Ascidiaceaclassification Order Stolidobranchia Family Styelidae Styela canopus Pyura speciescomplex It couldalsobe………. Nelson Harbour. pilesinlowOccurs subtidally onwharf abundance, present known distribution, with purplishtinges. outside ovary. Colourinlifecream to tan,stripeswhite, orange orburnt brown slits elongate,leathery test.Gill gillsfolded, tentaclessmooth,testisfollicles the siphonsandupperbody; thesemay beobscured by wrinklesinthetough thebody. of distinct ontheback down Finestripesrun theexternal surface of tubercles occurring around thesiphonsandlongitudinalwrinkles,becoming less the body,on the top of one slightly larger than the other. Test tough with warty Body small,erect,oblong, withnostalkand closelyspaced twosiphons short Kott, P. (1985)TheAustralian andStolidobranchia. Ascidiacea I.Phlebobranchia

(Savigny, 1816) Memoirs of theQueensland Museum , 23:440. Memoirs of main andinsetimages: Rosana Rocha Cosmopolitan 1 cm 46 Class Ascidiaceaclassification Order Stolidobranchia Family Styelidae Styela clava Pyura pachydermatina It couldalsobe………. the seabed.Canbelocallyabundant. and farms, marine and pontoons marina as such structures artificial on Settles and sediment. follicles outside ovary. Colourin life cream to tan, often covered with epiphytes down thestalk.Gillsfolded, gillslitselongate, and tentacles smooth. Testis at thetoparound thesiphons. testcreased longitudinally and Posterior of half thebody.together atthetopof Test leatheryand conical, withwarty swellings body, tosubstratum by anchored root-like processes. siphonsareclose Short stalk, generallyIndividuals usuallywithashort nolonger thanthecylindrical

Kott, P. (1985)TheAustralian andStolidobranchia. Ascidiacea I.Phlebobranchia Herdman, 1881 Memoirs of theQueensland Museum , 23:440. Memoirs of Cosmopolitan 4 cm 47 Class Ascidiaceaclassification Order Stolidobranchia Family Styelidae Styela plicata Often occursindenseclustersandisrarely fouled withotherorganisms. orange tinges. smooth, and testis follicles outside ovary. Colourinlifedullwhitewithburnt knobbed, pleatedappearance.elongate,slits Gill folded, gills tentacles subdivided by are further ridges horizontalcreases, which givingita distinctly longitudinal into divided test. Test cartilaginous thick firm, a with ovoid Body Kott, P. (1985)TheAustralian and Stolidobranchia. Ascidiacea I.Phlebobranchia

(Lesueur, 1823) Millar, New Zealand: Ascidiacea. R.H.(1982) Themarine faunaof New Zealand Oceanographic Institute Memoir, 85: 1–117. Memoirs of theQueenslandMuseum,23:440. Memoirs of Cosmopolitan inset images: John Borom 2 cm 48 about this guide | about sea squirts | colour index | species index | species pages | icons | glossary icons morphology

hemispherical with brain- ball spherical, globular brain like corrugations

rounded elongate, long tubular sausage- loaf sausage hemispherical shaped colonies

without definable shape, often with lobed surface, closely packed flat topped amorphous lobed cluster potato or tuber-shaped, lobes joined by basal mat massive

many single bodies on spreading over substratum, thick encrusting medusa long stalks arising from a more than 20 mm thick narrow basal mat

widely-spaced siphons spreading over substratum, thin encrusting solitary saddle with low saddle in less than 5 mm thick between

elongated body with a finger-like, often arising short narrow stem, siphons fingers from an encrusting or solitary stalked vase closely spaced at anterior restricted base, digitate end wandering along and oval bulbous body with 2 above substratum meandering solitary stalked siphons on a long narrow attached at intervals, stem repent low, laterally elongate, stalked with club-shaped oval shaped, with 2 stalked grouped bodies attached to a solitary mound siphons, separated by common basal mat about ½ a body length

rounded body, siphons stalked simple single stalked bodies solitary rounded often close together at the anterior end

vertically elongated body bunched vase-shaped grapes solitary oblong with 2 siphons at the ante- individuals joined basally rior end

49 about this guide | about sea squirts | colour index | species index | species pages | icons | glossary icons surface

even, hairless, silky, can be bearing small flattened smooth warty slightly undulating bumps or tubercles, verrucose

zooid apertures line hairs projecting from the subdermal canals radiating body of solitary ascidians, radial systems hairy and branching away from often holding sand grains, common cloacal apertures hirsute

zooid apertures form rings common cloacal apertures circular systems around common cloacal raised lobes raised at the terminal end of apertures lobes

prickly bundles of very long gelatinous and see-through, spiny spicules projecting from the transparent translucent test of solitary ascidians

siphons raised above the irregularly pitted and ridged rough wrinkled siphons body wall, wrinkled and surface, often tough, rugose often warty

sandy sediment incorporated star-shaped carbonate sand in test into test of colonial ascidians, spicules granules visible in and on feels granular the test

bearing irregularly parallel zooid oral apertures deeply wrinkled ribs and grooves along the parallel systems in parallel lines along body wall subdermal canals

zooids open separately test surface with ridges in a honeycomb no systems forming paired openings on honeycomb pattern low humps in the test

icons habitat very fine muddy and silty hard substrate such as mud sediments derived from rock mudstone, sandstone, basalt, terrigenous rocks, soils and compressed carbonates clays

living or growing on the shell, stone, and pebble rubble epizoic external surface of an rubble animal

anything man-made such as small coarse grains of worn artificial sand mooring blocks, mussel lines, silica, rock, and shell substratum wharf piles

50 about this guide | about sea squirts | colour index | species index | species pages | icons | glossary

icons environment

exposed shoreline zone between sand and rubble spread over high and low tides, including rock underlying hard substrate, flats, pools, overhangs, crevices, organisms attached to basement intertidal covered rock organisms exposed to wave rock susceptible to inundation and action, temperature extremes, full scouring from wave surge and illumination, and desiccation currents, and subdued illumination

composed of a variety of zone below the low tide, including sedimentary substrates including rock flats, slopes, walls, crevices, coarse gravels, shell hash and overhangs, boulder fields, sands to finer sand, mud, and subtidal seabed organisms exposed to wave silts, organisms susceptible to surge and currents, and subdued inundation and scouring from illumination wave surge and currents, and subdued illumination seabed raised into a bank of compacted rubbles and other underwater cliffs and slopes, carbonate materials including organisms exposed to wave wall bank shell, kina and sealace hash, surge and currents, and subdued organisms exposed to wave illumination surge and currents, and subdued illumination underwater caves, shelves and overhangs, organisms may indents experience wave surge, subdued illumination, or near darkness

icons life history

species first described from one animal bound by a single solitary native and only found New Zealand test waters, endemic

species first described from multiple animals bound by a outside of New Zealand waters colonial introduced single test and is found in New Zealand and other locations, invasive

since first described in NZ, this range species has been recorded extention elsewhere

51 about this guide | about sea squirts | colour index | species index | species pages | icons | glossary glossary amorphous without definable shape, often with lobed surface, potato or tuber-shaped, massive ampullae blind terminal expansion of the epidermal vessels, often flask-shaped in the Botryllidae anterior front apertures openings of the body to the exterior for exchange of water, inhalant ‘mouth’ (branchial) aperture, exhalent (atrial) aperture artificial substratum anything man-made such as mooring blocks, mussel lines, wharf piles ball spherical, globular bank seabed raised into a bank of compacted rubbles and other carbonate materials including shell, kina and sealace hash, organisms exposed to wave surge and currents, and subdued illumination brain hemispherical with brain-like corrugations cartilaginous having the texture of cartilage, firm and tough yet flexible circular systems zooid apertures form rings around common cloacal apertures covered rock sand and rubble spread over underlying hard substrate, organisms attached to basement rock susceptible to inundation and scouring from wave surge and currents, and subdued illumination deeply wrinkled bearing irregularly parallel ribs and grooves along the body wall environment physical, chemical, ecological, behavioural, and other conditions experienced by an organism epizoic living or growing on the external surface of an animal fingers finger-like, often arising from an encrusting or restricted base, digitate firm requires some pressure to compress, firm fleshy feels like skin or edam cheese, dense, slightly stretchy, cellular material more abundant than fibrous material gelatinous jelly-like, slippery gill sac organ used for both the exchange of gasses (breathing) and collection of food gonad reproductive structure granular sand papery texture due to presence of calcareous spicules in the test grapes bunched vase-shaped individuals joined basally habitat environment and local situation an organism lives in hairy hairs projecting from the body of solitary ascidians, often holding sand grains, hirsute honeycomb test surface with ridges in a honeycomb pattern indents indentations in the substrate such as underwater caves, shelves and overhangs, organisms may experience wave surge, subdued illumination, or near darkness intertidal exposed shoreline zone between high and low tides, including rock flats, pools, overhangs, crevices, organisms exposed to wave action, temperature extremes, full illumination, and desiccation loaf rounded elongate, hemispherical lobed cluster closely packed flat-topped lobes joined by basal mat meandering wandering along and above substratum attached at intervals, repent medusa many single bodies on long stalks arising from a narrow basal mat morphology shape mud very fine muddy and silty sediments derived from terrigenous rocks, soils and clays no systems zooids open separately forming paired openings on low humps in the test opaque impenetrable by light parallel systems zooid oral apertures in parallel lines along subdermal canals posterior back radial systems zooid apertures line subdermal canals radiating and branching away from common cloacal apertures raised lobes common cloacal apertures raised at the terminal end of lobes rock hard substrate such as mudstone, sandstone, basalt, compressed carbonates rough irregularly pitted and ridged surface, often tough, rugose rubble shell, stone, and pebble rubble sand in test sandy sediment incorporated into test of colonial ascidians, feels granular sand small coarse grains of worn silica, rock, and shell sausage long tubular sausage-shaped colonies seabed composed of a variety of sedimentary substrates including coarse gravels, shell hash and sands to finer sand, mud, and silts, organisms susceptible to inundation and scouring from wave surge and currents, and subdued illumination smooth even, hairless, silky, can be slightly undulating solitary mound low, laterally elongate, oval shaped, with 2 siphons, separated by about ½ a body length solitary oblong vertically elongated body with 2 siphons at the anterior end solitary rounded rounded body, siphons often close together at the anterior end solitary saddle widely-spaced siphons with low saddle in between solitary stalked vase elongated body with a short narrow stem, siphons closely spaced at anterior end solitary stalked oval bulbous body with 2 siphons on a long narrow stem spicules star-shaped carbonate granules visible in and on the test

52 about this guide | about sea squirts | colour index | species index | species pages | icons | glossary

spiny prickly bundles of very long spicules projecting from the test of solitary ascidians stalked grouped stalked with club-shaped heads attached to a common basal mat stalked simple single stalked bodies subdermal canal a canal that connects zooids together around a common cloacal aperture (exhalent) subtidal zone below the low tide, including rock flats, slopes, walls, crevices, overhangs, boulder fields, organisms exposed to wave surge and currents, and subdued illumination surface patterning or ornamentation on the surface of the body of an animal tentacle tentacles surround the inhalant (branchial) aperture; they can be simple or branched and are important characters at the genus level test a protein coating surrounding the body, it can be tough and leathery in some solitary species, or a gelatinous matrix surrounding zooids in colonial species testis follicle sacs that contain sperm; these are usually cream-coloured and the ovary is orange, containing eggs thick encrusting spreading over substratum, more than about 20 mm thick thin encrusting spreading over substratum, less than about 5 mm thick translucent lets light through the test, but not enough to perceive distinct details through it. transparent test of both colonial and solitary ascidians can be gelatinous, apearing see-through, translucent wall underwater cliffs and slopes, organisms exposed to wave surge and currents, and subdued illumination warty bearing small flattened bumps or tubercles, verrucose wrinkled siphons siphons raised above the body wall, wrinkled and often warty zooids small individual seasquirts of the same species living communally in a common test, often forming systems to pump water, or opening individually to the exterior

53 about this guide | about sea squirts | colour index | species index | species pages | icons | glossary acknowledgements

This guide is dedicated to the late Patricia Mather (nee Kott) in acknowledgement of her lifetime contribution to the taxonomy of Southern Hemisphere ascidians. Our knowledge of the New Zealand ascidian fauna is richer for the early works of Sluiter, Michaelsen and more recently, those of Brewin and Millar. Many of the images in this guide were taken during NIWA’s Marine Biotechnology Programme collection voyages; many thanks to Vicky Webb for having the foresight to support our research in this area. This research was funded by NIWA under Coasts and Oceans Research Programme 2 Marine Biological Resources: Discovery and definition of the marine biota of New Zealand (2012/2013 SCI). image credits

Unless stated otherwise, all images in this guide are the work of Mike Page. Permission to use images from the following photographers is gratefully acknowledged: Craig Depree, NIWA Hamilton, New Zealand; Sean Handley, NIWA Nelson, New Zealand; Dennis Gordon, NIWA Wellington, New Zealand; Chris Woods, NIWA Christchurch, New Zealand; Roger Grace, Algies Bay, New Zealand; Floor Anthoni, Seafriends, Leigh, New Zealand; Arjan Gittenberger, University of Leiden, Leiden, Netherlands; Keith Hiscock, Marine Biological Association, Plymouth, United Kingdom; Misjel Decleer, Bruges, Belgium; Judith Oakley, Oakley Intertidal, Wales, United Kingdom; Natural History Museum, London, United Kingdom; Charles Griffiths, Capetown University, Capetown, South Africa; Rosana Rocha, Universidade Federal do Paraná Curitiba, Brazil; John Borom, Weeks Bay Reserve Foundation, Alabama, United States of America; Dirk Schories, Universidad Austral de Chile, Chile, South America; Anne Frijsinger and Mat Vestjens, Kapolnasnyek, Hungary.

further reading

Berrill, N.J. (1950) The Tunicata, Royal Society Publications 133: 1–354.

Kott, P. (1989) Form and function in the Ascidiacea, Bulletin of Marine Science 45 (2): 253–276.

Kott, P. (1998) Tunicata, in A. Wells & W.W.K. Houston (eds), Zoological catalogue of Australia, CSIRO Publishing, Melbourne: 51–252.

Kott, P., Bradford-Greive, J.M., Esnal, G.B., Murdoch, R.C. (2009) 23. Phylum Tunicata: Sea squirts, salps, appendicularians, in D.P. Gordon (ed), New Zealand Inventory of Biodiversity Volume 1 Kingdom Animalia: Radiata, Lophotrochozoa, and Deuterostomia, Canterbury University Press: 409–430.

Monniot, C., Monniot, F., Laboute, P. (1991) Coral reef ascidians of New Caledonia, Editions de L’ORSTOM, Paris 30: 1–248.

Produced for NIWA by TC Media Ltd www.tcmedia.co.nz July 2013 54