MEMOIRS OF THE QUEENSLAND MUSEUM a Con) VOLUME 32 PART 2 BRISBANE 1 OCTOBER, 1992 From the President’s Desk The XII International Congress of Arachnology was held at the Queensland Museum, Brisbane, from 12th to 18th July, 1992 under the joint auspices of the Trustees of the Queensland Museum and the Centre International de Documentation Arachnologique. More than 122 registrants from 32 countries and 37 accompanying persons participated. The 114 presented papers covered nearly all fields of arachnological research, reflecting the progress in our knowledge of arachnids, This progress may encounter some scepticism from particularly the older, loyal participants since every president in each presidential report underlines the ‘progress made during the last three years’. A congress program does not reflect the actual research level, but only the fields of interests of participants whose attendance at congress depends on economic or other factors. Real progress can be emphasised in the fields of biology, ecology, bio-indication and phylogeny. However, we might be regressing in taxonomy because the number of taxonomists remains stable while the numbers of identifications requested by ecological and other studies (raising new taxonomical problems) are increasing. Some arachnid groups will perhaps be ‘orphaned’ in the near future as specialists retire from active study. Arachnids are fascinating animals of great biological importance. Is there any reason not to tell this more frequently to ‘the public’? Ecologists should allow a reasonable fee for identification of their material when submitting grant applications. At the meeting of CIDA members held during the Congress, Dr R. Jocqué was elected the new CIDA president. The congress excursion was held mid-week at O’Reilly’s, Lamington Plateau, where participants could discover the rich bird and arthropod fauna of the forest, stroll through the botanical garden or have an exciting canopy walk. Brisbane was a marvellous occasion to meet friends and colleagues, but also (at least for me) a wonderful time to discover a fascinating continent and its plants and animals, and | am sure that we will come again. Thanks to all of the organising staff and the people we had the chance to meet for their friendliness and helpfulness. ‘Le Roi est mort, vive le Roi’. Hopefully we will meet in Geneva in 1995, Volker Mahnert President, Centre International de Documentation Arachnologique 1989-1992 MEMOIRS OF THE QUEENSLAND MUSEUM, VOLUME 32 CONTENTS PART | (Issued 29 June, 1992) Bruce, N.L. Anopsilana barnardi, a new species of estuarine cirolanid crustacean isopod from tropical easter Austfalia, 0.00000... cc ccc eee eutcvuuecucvs oe ee a ee Se ae ER l CHURCHILL, T,B. AND RAVEN, R.J. Systematics of the intertidal trapdoor spider genus Idioctis (Mygalomorphae: Barychelidae) in the Western Pacific with a new genus from the Northeast ............... reneetsetooacegt eG COLEMAN, R.A. Shipwreck and historical survey: Chilcott Islet, Coral Sea... .......-. 2-2. 0-e- 000-5. seetesutst St COUPER, P. J. AND INGRAM, G.J. A new species of skink of Lerista from Queensland and a re-appraisal of L, allanae Langa n)s ak oye rs yore tbs Sef FEHRE EeaD doe 1 ketal y eyed eats datele poled ala valeces: 55 DAVID, B., CHANT, D. AND FLOOD, J, Jalijbang 2 and the distribution of pecked faces in Australia............,.00.--..----. pevyetas,. GT DAVEE, P.J.F Revision of Sarmatium Dana (Cmstacea: Brachyura: Sesarminae) with descriptions of three new species ......0..0..0.0.000, PORE Se poeta pes tela sabi deunleste-o gee apie apccsy 79 Hoorer, J.N.A. AND BERGQUIST, P.R. Cymbastela, anew genus of lamellate coral reef sponges .,.,, SEER LCS SE Ape cty eda eta tele ele tela: > 99 HUGHES, N.C. AND DROSER, M.L. Trace fossils from the Phe Formation (Lower Cambrian), Zanskar Valley, northwestern India... ..... 139 Jones, D.S. Scalpellid barnacles (Cirripedia: Thoracica) from the northeastern and central eastern Australian continental shelf and slope ................ aie et dee ee: pa epet etn kig.s 145 KENNETT, R.M., GEORGES, A., THOMAS, K. AND GEORGES, T.C. Distribution of the long-necked freshwater turtle Chelodina novaeguineae and new information on its Ecology... . 6.6.6... kee cece eee ec euepeuteaeetacevetuceaeeeees 179 Lamekwy, K.J, Re-examination of the venation of Osmylopsychops spillerae Tillyard from the Triassic of Queensland 183 LAMPRELL, K, Notes on Spondylus Linnaeus with descriptions of two new Specics from Western Australia (Mollusca; Bivalvia: Spondylidae) ..;...,... Te eee koa tee nee a beled able went. 189 ROZEFELDS, A.C. The subtribe Hicksbeachiinae (Proteaceae) in the Australian Feryatyes yotlicalt eset v5 195 ROZEFELDs, A.C., CHRISTOPHEL, D.C. AND ALLEY, N.F. Tertiary occurrence of the fern Lygodium (Schizacaceae) in Australia and New Zealand, ..,..... vee, 203 RYLAND, JS. AND HAYWARD, PJ, Bryozoa from Heron Island, Great Barrier Reef... 2... 0.0.0.0 occ ce ccc ccuecee. be FPL IGS 223 SHEA, G.M. AND HUTCHINSON, MLN. A new species of lizard (Tiligua) from the Miocene of Riversleigh, Queensland........... veveeee. 303 WEsSTER, G.D, AND JELL, P.A. Permian echinoderms from Western Australia .........5.6600000-cc0cee-0e. ake bib bb ott ele a wv Sd NOTES CourEr, P.J.. Anomalopus pluto Ingram, a poorly known skink from Cape York Peninsula .,........... belere'aia. 54 Courer, P.J., WHITTIER, J., MASON, R.T, AND INGRAM, G.J. A nesting record for Coeranoscincus reticulatus (Giinther), ©... ect cece cece gua 60 DAVIE, P.J.F. The rare shrimp, Phyllognathia ceratophthalma (Balss), discovered in Moreton Bay .......2...,..,.98 RICHARDS, S.J. The tadpole of the Australopapuan frog Rana daemeli,.............., . PM adg pchiate teats Te Pye setts, 138 RICHARDS, S.J. AND JAMES, C. Poot-flagging displays of some Australian frogs... .- 2.0.0.0 ees eee ce ee cececcecce. nb ds sath « 302 CONTENTS KOTT, P. The Australian Ascidiacea part 3, Aplousobranchia (2) KOTT, P. The Australian Ascidiacea, Supplement 2 ...... 0.0.0.0... ccc cece cece cee ceeaseuueeeues 621-655 Pee eee eevee ere eve eseeteeeses see 375-620 VAN DYCK, S. AND DURBIDGE, E. A nesting community of False Water Rats Xeromys myoides on the Myora Sedgelands, North Stradbroke Island... 0.0.0... eee cc eee eet eee ee tne enact ener e eens 374 PART 2 (Issued 1 October, 1992) KOTT, P. The Australian Ascidiacea part 3, Aplousobranchia (2) ........-6 20 cece eee eee teens 375-620 KOTT, P. The Australian Ascidiacea, Supplement 2 .. 22.0.0... cect ete ee eee etter eee 621-655 THE AUSTRALIAN ASCIDIACEA PART 3, APLOUSOBRANCHIA (2) PATRICIA KOTT Kott, P. 1992 1001: The Australian Ascidiacea Part 3, Aplousobranchia (2). Menroirs of the Queensland Museum 32(2); 375-620, Brisbane. ISSN 0079-8835, In this continuing systematic revision of the taxonomy of the class Ascidiacea (subphylum Tunicata) taxa in the formerly polyphyletic family Polyclinidae Milne Edwards are treated. Included are all species recorded from Australian walters wilh a posterior abdomen containing the gonads, and usually the heart, New famulies are erected to accommodate the genera formerly grouped in the polyphyletic Euherdmaniinae, viz. Placentelidae (Placentela), Protopolyclinidae (Proropalyclinum, Monniotus, Condominium n.gen.), Ritterellidac (Ri- terella, Dumus, Pharyngodicyton), Eaherdmaniidae (Euherdmania) and Pseudadistomidae (Pseudodistoma, Anadistoma n.gen., Citorclinum), The family Polyclinidae is redefined and restricted to the genera formerly comprising the nominal subfamily Polyclininae, viz. Polyclinum, Aplidiapsis, Sidneioides, Synoicum, Merchellium and Aplidium. Larvae, observed for the majority of species, have characters significant at species, genus and family levels, and constitute valuable evidence of phylogenetic affinities. Embryos are incubated to tailed larvae internally in all taxa, usually being fertilised and beginning their development in the distal part of the oviduct, sometimes in a brood pouch constricted off from the zooid and sometimes at the base of the oviduct, Colony development is significant in this group of aplousobranch taxa in which replication is prolific. Zooids are arranged in various colonial, including cloacal, systems. In taxa with extensive cloacal spaces (Aplidium), and in those with a soft test (e.g. Polyclinwm), intra- colonial hydrostatic pressure appears to contribute to colony turgidity. Sand and other inclusions in the test add strength and contribute to diversity in colony size and shape, Parallel reduction in the size of zooids often results in convergent evolution, especially in size (number of rows of stigmata and number per row) of the branchial sac and size of larvae. Some taxa (Pseudodistoma, Polyclinum) have zooids and larvae wilh conservalive moarphol- ogy, and differences in colonial systems and colony shape and texture often determine species-level taxa. Zooid and larval morphology are more diverse in Ritterella, Synoicun Aplidiopsis and Aplidium, with differences in the atrial aperture, stigmata, stomach wall an arrangement of larval ampullae and vesicles, as well as fealures of the colony and its systems, providing characters for species determination. Other characters, such as the nature of the larval adhesive apparatus, presence of branchial papillae, length of the oesophagus, nature of the stomach wall, and shape of the posterior abdomen and arrangement of male follicles in il, are significant indicators of relationships at higher (genus or family) taxon levels. Sixty-two of the total of 110 species treated in this work are new, Many may be indigenous, but these families are not always well sampled, not often being found intertidally. Collection by SCUBA diving enables a diversity of habitats to be sampled and the range of forms treated in this work is largely the result of efforts of these divers, Other difficulties in studying this grouip, which have resulted in synonymy and misidentification, are colour variation and its post mortem change and loss, loss of colony and zooid turgor obscuring their shape and arrangement, and contractibility of zooids Sabana their morphology, The most diverse genera in these waters are in the Polyclinidae, in Which zooids are organised around well developed cloacal cavities, Aplidium is the most diverse gemns, and is well represented in both tropical and temperate waters by species with large and conspicuous colonies. OF the families with independently opening zooids without cloacal systems, Pseudodistomidae and Ritterellidae are well represented, especially in certain temperate locations. Euherdmaniidae and Protopolyclinidae are less diverse although each contains some species with wide temperate geographic ranges common in certain locations, (C0 Indo-West Pacific, Ascidiacea, Aplousobranchia, Placentelidae, Protopelyclinidae, Rinterellidae, Euherdmaniidae, Pseudodistomidae, Polyclinidae, Patricia Kotr, Queensland Museum, PQ Box 3300, South Brisbane, Queensland 4101, Australia. | April, 1992. 376 MEMOIRS OF THE QUEENSLAND MUSEUM e f Frontispiece: larvae of Aplousobranchia — a, Pseudodistoma australe (see Fig. 17g); b, Euherdmania digitata (see Fig. 13£); ¢, Polyclinum fungosum n.sp. (see Fig. 25d); d, Aplidium inflorescens n.sp. (see Fig. 82c); e, Aplidium altarium (see Fig. 62e); f, Aplidium filiforme n.sp. (see Fig. 75e). THE AUSTRALIAN ASCIDIACEA 3 CONTENTS FrrOgGHOths + oats p-ye'evalet ots v wane oe Dafoe 378 P. terranuim n.Spy. os... srolecgiebeer da0l Acknowledgements .........0000000002 005 378 P. tsutsuii Tokioka, 1954 _...-.--.--.--.- 463 Annotated Glossary... 0.600.604 pee eee 380 mt Pele a Pizon, 1908 ,... 0+ +++++4- A PUgiopstsso.40% 56-20 SRA SR TSO SR BOSS POLYCLINIDAE sensu Milne Edwards, 1842 390 A. confluata WSP. occ. ee enter een eee 468 A, mammillata NSP... eee vaeeheare 469 PLACENTELIDAE n.fam,.... ......--.-391 A, sabulosa wsp.. 0. cc ce eee 470 Sidneioiden oc cc cee eek een enne 471 PROTOPOLYCLINIDAE n.fam........... 392 S. tamaramae Kesteven, 1909 ...........- 472 Moanniotg§ = 32605 c4.¢0 305 5b Shwe tte 393 SYROICUI 4 ste scea nia neanianeanon pends 473 M. australis (Kott, 1957) 2... 0.2.0.2... .394 S. ANQUStUM NSP. 2.20 ev eee fet hands 477 M, radiatus n8p. oe. cece ee eee 396 S. arenaceum (Michaelsen, 1924) ...,..... 478 Condominium n.gen... 0.0... eee ee B96 S. atopogaster Kott, 1963... 6c eee eee 478 C. areolatum (KotL, 1963) .....-4..0062+-397 a son ee ans Millar, 1963 .-....-..+..-- mi buccinum MSP. oo. ee ee ee te Kira RITTERELLIDAE n.fam., ........... 00: 3909 S. castellatum MSP... cece eee eee eens 483 Priratetid cca oa ocd dh onle rica e 400 S. chrysanthemum 0sp. -. 02-02 e eee ees 485 R. asymmetrica Millar, 1966 ............. 402 Se CHUM DBP. 4. sacaseseesersaracz ces - 486 R. compacta 8p. .....1.1-.s2s002c 00s 403 S, CONCAVIIUM DSP. occ ee ee eee 487 RGOIMMB TES. °. psc ete etait th bein 405 7 SHEE DED on: omta ar kav im ehenia nts R. dispar Kott, 1957 00.0... cece eee 406 CORA og Sa SER ae eae lcs 250 R. multistigmata n.sp... 2.6.6. ee ee 408 BORE ICA, Phat aresscsasesitei sss S, intercedens (Sluiter, 1909)... 04.04. 49] R. papillata nsp. . 2.42.4. 228 cesses see 409 SI 493 R. pedunculata (Herdman, 1899) ......... 410 Ongistriatem NSP. «sos 61> sve ers rey rs Purine 412 S. macroglossum (Hartmeyer, 1919)... 2... 494 D. areniferus Brewin, 1952 0.00.00. 45, 413 § papillferum (Michacisen, 1930)... 499 EUHERDMANIIDAE Ritter, 1904 emend...413 Primi Mlerman, TBI) «ovo soos pe Evherdmania .. 65.6 cc ee eee tees 4\5 S suarenum ne t : ; 7 : wu etiniw i Ke : : 50t E, dentatosiphonis (Millar, 1975)... ‘416 § fropicum (Sluiter, 1909) |... 303 e a estibeie iiss 1963 2s eee cence eee pity Marchellium. 0.060 ences 504 : ra el ahaa whieh aha A a = M. albidum nsp,... 2... 05. 0- + 6-0 0-- »-+»- 505 PSEUDODISTOMIDAE n. fam........... 421 M. PANNOSUM MSP. 2-0 eevee eee sees 506 Ryeltealivedens 472 ADH EEAIN 54 ocecs led elk obt-m othcetectleth ofu mle h oho 507 SEEM ENERSY bt Sst mea Gtoben Heo Ett Fe a6 A, GCFOPOTUM MSP. . 25 oe con ee ee ee 513 LL a 42s A. altarium (Sluiter, 1909)... ....0..-2.-. 519 P. aur sig Posing a ae A. amorphaium Kott, 1963 ......0 60000000. 522 P. australe Kott, 1957 oo... eevee eee 428 A, australienseé Kott, 1963 .........6.0005 523 P.candens Wsp. . +. : Sipe eats as eedslone pi A. baeculum nsp......-..--.- a.¥i, 44° 526 - Sidi sc bee pees ie SEs Bed Bef es ae A, brevilarvacium Kott, 1963............. 527 ak Sa 2 ‘temredtenrlerier sey A. caelestis Monnivt, 1987... . . ores 528 pil Sapte > ppv yeyrss eriediaese tt at A. CLIVOSUM TSP. eee ee eee 530 : M piety Aerts Oe 9 ERIS Mee heres A. congregatum MSP. ..- 6.26 2 ee eee es 533 P. pulvinum TSP. eet oejola F srw da eerie vee ee 439 A. coniferum Kott, 1963... 2.22, .2.0-.5-5 535 Es grt te TBE 0a 82 ke ies eres i. tan A, crateriferum (Sluiter, 1909)... 6.664 .. 536 CAHENUAIMIM MSP, oe eee ee eee ee A. depressum Sluiter, 1909... .......-...4 538 A. directum Kott, 1972.0... 0.22. 224224555 539 POLYCLINIDAE emend. ..........-..-. 44] A. distaplium M.Spuo. ei. cc eee 540 Polyclinum ooo. ccc ees 443 A, élatum Kott, 1972. ...0..0...,...-..-. 541 P, fungosum Herdman, 1886 .-.........- 447 A, filiforme USP... 20. ee ee en 542 P. glabrum Sluiter, 195... 6... ee 4Ag A. flUorascum DSP. sasaes server cateuten 544 P, incrusrarum Michaelsen, 1930... ... ..-450 A, gastrolinealum O.8p.... 624624 546 P, marsupiale Koll, 1963... 20.06) 002 ee ees 452 A, gelasiniim O.8Pp.. 06.66 eee eee 547 Po nudum WSpooce cece oom ns mien 454 A, EMINATUM NSP. . =) 0 ee 548 P. OTBUUM DSP. - 6 ees 455 A. QriS€UM MSP... 6 cee en esa baa baa tan 551 P. saturnium Savigny, 1816... ........3-- 455 As INCUBGIMIN TSP eee ee P. solum MOMMOV, 666. ene 458 A, inflorescens DSP. 22 0 eo oy ee 553 P. fentatum n.sp. oo eee eee 459 A, jacksani Kott, 1963... 2.00.6. ce ee 554 377 373 A. lenticubwm tage. oo. e ee eee ee ., 555 A. lodix n.sp. trea mnt De Sar wes repeenev doe A. lunacratum nom.noV..... 2... ee S538 A, macrolobatum tsp. ,.- 2-6 eee eee ee SEL A, magnilarvurt RSP. oo eee 563 A. minisculum 18p... 0.22.2. oe ee ee 564 A. multilineaturn msp.. oe. eee eee 566 A, multiplicatum Shuiter, 1909.2... , 2... . 367 A. epacum Kott, 1963 -....-..-..-.-.--.- 570 A. OPMGLUMTTLSP, 2.40 e ee ee teen DTD A, paralinedtora W3p. occ. c cee cece eee S74 A, parastigmaticoum asp. ......--..-.--.-375 A, parvunt Kott, 963 ic ci eee ee -.+-377 AY PesrosHrn WAP... ee cea eet eee ene 37 A. protectans (Herdman, I899) . 2,222... 579 A, ritteri (Sluiter, 895) 0.0.2.0. 20.22.22... 581 A. PODUSHIIT TSP oe ce eee ene $82 A. rosarium N&pe. oe eee ee eee DBS A. rabricollum Kott, 1963,.0......22.-..- 384 A, solidum (Herdman, 1899)... 0.22.2. 586 A, tabascunt asp... oe ee ee pee SRD A. triggsense Kott, 1963 0.2... 00 A, uleute Monniot and Monniot, 1987. ...,. 592 INTRODUCTION Ascidians of the suborders Phicbobranchia (one of the 2 suborders of the order Enterogona) and Stolidobranchia (the only suborder of the order Pleurogona) were reported an in part 1 of the present work (Kott 1985); and 7 families of Aplousobranchia (the other suborder of the En- terogona) were dealt with in part 2 (Kott 1990), The following account (part 3) continues the re- view of the Aplousobranchia, dealing with genera formerly contained in the family Polyclinidae Milne Edwards, 1842. These genera are here as- signed to 6 different families, 3 of them new (Table 1). Didemnidae (Aplousobranchia). ihe only family not yet treated, will comprise part 4 of this mono- graphic treatment of the Ascidiacea in Australia. The collections on which the work is based are those set out in parts 1 and 2 (Kott 1985, 1990). Museums housing the material examined are re- ferred to by the following abbreviations: AM, Australian Museum, Sydney, New South Wiles: BM, British Museum (Natural History), London, UK; QM, Queensland Museum, Brisbane, Queensland; SAM, South Australian Museum, Adelaide, South Australia; TM, Tasmanian Mu- seum, Hobart, Tasmania; MV, Museum of Victo- tia, Melbourne, Victoria; NTM, Northern Territory Museum, Darwin, Northern Territory: WAM, Western Australian Museum. Perth, Westem Australia; USNM, LS National Museum of Natural History, Smithsonian Institution. Washington DC, USA; ZMA, Zoological Mu- MEMOIRS OF THE QUEENSLAND MUSEUM seum of Amsterdam, Amsterdam, Netherlands; ZMC. Zoological Museum, University of Copen- hagen, Copenhagen, Denmark. AMPT refers to the Australian Marine Photographic Index (Neville Coleman). All specimens referred to by a museum regis- tration number have been examined in the course of the present study. A registration number in italics refers to a sampled portion of the preceding colony. Throughout this work, the characters of species are summarised to provide a general overview of the diversity of each genus. Dichotomous keys are presented as aids to the identification of taxa at all levels. They have been prepared, as far as possible, fo reflect phylogeny, This type of key is generally used for biological material in prefer- ence to tabular keys such as those in Menmiot (1987), The latter provide too rigid a framework for the analysis of diverse biological material, communicate only a limited amount of informa- tion, lack the flexibility to distinguish between convergent and phylogenetic relationships, are open to subjective interpretation, and are unnee- essarily unwieldly and ambiguous to use. Procedures for cailection and examination of species are set outin Kott (1990). Examination of specimens is by stereascapic and compound light microscopy. All larvae are described from stained and mounted specimens taken from adult colo- nies as indicated. ACKNOWLEDGEMENTS jum grateful to those of my collegues, as sct out in parts | and 2, who, in various material ways, have contributed to the preparation and presenta- tion of these volumes; and to those wha have made Jess tangible, but equally significant contn- butions by their support and encouragement. The black and white figures were prepared from camera lucida and other sketches of the speci- mens by Stephen Cook (Figs lal, 2-6, 7bi, 8a, 9a-f, Lla-f, 13a-e, 14, 17b—3, 18a-c, 19, 21a, 24, 27, 28a, 35a,b-e); Lucille Crevola Gillespie (Figs. 7a, 17a, 22a); and John Kennedy (all other figures including the larvae of all taxa), Peta Woodgate typed the manuscript. I am grateful to them all. Sue Boyd with Christine Mateira in the Museum of Victona; Penny Berents in the Aus- tralian Museum. Loisette Marsh with Jean Thip- thorp in the Western Australian Museum, and Wolfgang Zeidler with Karen Gowlett Holmes in the South Australian Museum have been co-op- THE AUSTRALIAN ASCIDIACEA 3 379 | | § | posterior abslomen RITTERELLIDAE n, fam. al B& not constricted S| 2 5 g 38 rior abdomen g z B| 3 8 |Posten PROTOPOLYCLINIDAE n. fam. 3 3 | — Q |constricted mn |e 3 gq |2 2] 8 |ctoaca 2 § 3 3 E g cloacal systems present POLYCLINIDAE emend. =] A 2 2k — 3 Z 3 6 ‘Oo “a g | a < E | & & |heart in abdomen PLACENTELIDAE a. fam. ry & 5 5 3% =i E 4 2 2 stomach at posteri id 8 § PA E jomach at posterior en 2 g 5 2 of long abdomen BURERDMSNUDAE Set L. < = 3 = | stigmata in 3 rows PSEUDODISTOMIDAE n. fam. | = Bz], & Zz ra 2 3 o s z 2 Es Z| gonads in abdomen POLYCITORIDAE g 3 3 3 2 s 2 o 3 |replicates by oesophageal budding DIDEMNIDAE = po “ 2 < 3 & replicates lobes of apertures smooth HOLOZOIDAE 3 2 generated in a a 3 vegetative =z & s stolens lobes of apertures denticulate STOMOZOIDAE > B 5 a 2 abdomen more than twice length of thorax; 3 = branchial embryos fertilised at base of oviduct; larval adhesive organs inverted tubes PYCNOCLAVELLIDAL « | 2 | E | apertures —- = 3 = | not lobed abdomen not more than twice length of thorax; bLAVERINIDAE a S 2 embryos fertilised at top of oviduct; larval adhesive organs with axial cone : r 2/3 3 2 internal longitudinal branchial vessels or forked vestiges DIAZONIDAE 3 = | solitary, and gut loop horizontal CIONIDAE a { 6 stigmata straight RHODOSOMATINAE gS gut on right side of pharynx - - CORELLIDAE & stigmata coiled 3 [g : & 2 e internal longitudinal branchial vessels represented by papillae AGNESIIDAE 5 3 Z z a - e S 3 Ly 2 2 solitary ASCIDIIDAE ea =. = ~ ° 3 be 3 gonads not embedded in the test a » | = % |pharynx fat PEROPHORIDAE = g 2 {gonads on colonial = = 3 left side only ——oeY = o g gonads embedded in the test PLURELLIDAE 5 & qa simple STYELINAE 3 33 branchial cloacal systems absent POLYZOINAE] STYELIDAE 4 S 3 tacles cloacal systems present BOTRYLLINAE Ss 2 & Qa3 o Z os F 4 3 rectangular stigmata; no renal vesicle PYURIDAE 3 =o 3 & a 6 g 3 Oo oF 3 y 3 Pharynx whole body length perforated MOLGULIDAE g 3 Fl by spiral or irregular stigmata = 4 > 3 ait ae S vo - = 4 F g ie short anterior band, perforated HEX ACROBYLIDAE n iad y circular stigmata a 4 TABLE 1. Ascidiacea — Graphic Key to families erative and prompt in response to my many re- quests for large specimen loans. During 1991 large collections from all around Australia also became available to me from the Australian Institute of Marine Science (AIMS), Townsville, through Dr Peter Murphy. This im- portant collection was made by Rob McCauley and his colleagues (AIMS Bioactivity Group), by SCUBA, as part of their Shallow Water Marine Organism Collection Program for the National Cancer Institute (1992-1996). This collection in- cludes photographs of the living colonies in situ. The voucher specimens are now registered in the collection of the Queensland Museum. Grants from ABRS (1980-82) and MS'T’ (83/1320) supported the author's field pro- gramme, The work was also supported by the ARC (419031140). ANNOTATED GLOSSARY A discussion of the morphology of aplousobranch as~ cidians and their larvae in the Australian Ascidiacea Part 2 (Kott 1990) refers particularly, although not exclusively, to families reviewed therein. Morpho- logical features, and their variations and functional implications that have not been discussed previously, ure reviewed in the glossary that follows, It should be used in conjunction with the glossaries in ips | and 2 (Kott 1985, 1990) co appreciate the full range of functional aduplations and other morphological variations in ascidians, (with the exception of the Didemnidae). adhesive organs: Each of the 4 types of adhesive organs present in the families discussed below, o¢- curs. also in other possibly related aplonsobranch families. | Deeply invaginated tubular adhesive organs, os in Pyenoclavellidac, are in Euherdmaniidae. They are arranged in a triangle when 3, or in the median line when only 2. They suggest a relationship with Pycnoclavellidae, 2.The 3 median pdhesive organs in Pseudodistamidac have a hollow, completely eversible axial protrusion in a deep epidermal cup and are either sessile or stalked, The axial protrusion is flat-topped or coni- cal, and apparently lacks the columnar cells found in the axial cones of Molozoidae (Distaplia; see Cloney 1977). When everted it is thin-walled and balloon- like, and may have closer affinities with the pdhesive organs of Pycnoclavellidae than any others. 3,The 3 median adhesive orgins in Protopolyclinidac are stalked, and have an axial cone composed of long columnar cells in adeep epidermal cup, They resem- ble the adhesive organs of Holozoidae (sce Cloncy 1977) and Didemnidae (see Turon 1991). 4. The 3 stalked adhesive organs in the antenor mid-line in Ritterellidue and Polyclinidae have a shallow, wide axial cone in a light epidermal cup. These resemble the adhesive organs of Polycitoridae, al- though they are smaller. Their relationship to the adhesive organs of Clavelinidae, and Holozoidae awaits investigation. Epidermal ampullae may also have an accessory adhesive function. (See also larvae, ampullae, ves- icles), ampullae, epidermal —, larval —: Outgrowths of the anteror larval epidermis, possibly accessory adhe- sive organs In Aplousobranehia they are absent tram Pyenoclavella, Eicherdmania ada lew other species level taxa in other genent in which size reduction appears 10 have resulicd th simphfication, Ampullac project from the base of triradial adhe- sive organs (Clavelimdas, Holozoiduc), Ampullac arise From lateral ridges along cach side of median MEMOIRS OF THE QUEENSLAND MUSEUM adhesive organs and alse from the median line alier- nabrg with them (Polyeitoridae, Prolopolyclinidae nofm,, Ritterellidae n.fam., Pseudodistomidac nfam_, Polyclinidae). Ln the new families Protopoly- clinidae (Monniones) and Riterellidae (Dumus), and in Polyclinidac (Polyelinum, Mallaig and some Aplidiim spp.) the terminal ends of the ampullae havea cap of modified, possibly secretory cells. (Sec larvae, vesicles). ampullary vesicles: see vesicles. anal opening, anus: The rectum continues anteriorly, embedded in the body wall along the dorsal border of the pharynx. Its opening into the atrial cavity is invariably bilabiate in the present aplousobrancl families. Its position relative to the atrial apertures (which arc normally antero-dorsal) appears deter- mined by selective pressures affecting the position of gonoducal openings. Faecal pellets are found in the rectum but never in the atrial cavity. Presumably, when necessary, the anus is brought into a position near the base of the atrial siphon by muscular action, $o that the faecal pellets ean be ejected directly to the exterior, Although the oviduet is sometimes diverted into a brood pouch, oris turned ventrally, the opening of the vas deferens is usually associated closely with the anus, the advantage possibly being that sperm also can be expelled directly to the exterior with the excurrent water. In Placentelidac n.fam., in which numerous em- bryos ate incubated in what probably is an elaborate brood pouch in the distal part of the oviduct (where it lies in the atrial cavity), the anus is ina primitive position in the anterior part of the atrial cavity, op- posite the second or third row of stigmata, This is in a position comparable with most Diazonidae, in which the gonodueal openings, at the base of the atrial aperture. are associated with the release of gametes direcily to the exterior for external fertilisa- tion, The anus and gonoducal openings are at mid-tho- racic level in most other genera discussed below. [n all uhese families fertilisation and incubation is either in the atrial cavity or in the distal part of the oviduct, Only in Euherdmaniidac and the (probably) related Pycnoclavellidae, both with characteristically long abdomina, are anal and gonoducal openings at the base of the atrial cavity; and in both fertilisation is at the base of the oviduct — al the posterior end of the Jong ebdomen, apertures: In most. aplousobranch ascidians, lrrespec- tive of the form of their systems, branchial apertures are 6-lobed. Exceptions in which the branchial aper- tures are 8-lobed are in some Rilterel/a spp. (viz, R, pedunewlata), Aplidium spp, (e.g, A, amorphatuen, A, pratectans) and Moreftellium, Apparcat subdivi- sion of branchial lobes in some species Aplidiwr (e.g. A, macrolabatum sp, ond A, Mneebatun n.spo results from contraction of a band of museles that extends up the centre to the tip of each Jobe. Separately opening 6-lobed atrial, as well as bran- chial, apertures interrupt the surface test of colonies THE AUSTRALIAN ASCIDIACEA 3 381 in the new families Placentclidae, Protopolyclinidac, Ritterellidae, Euherdmaniidae, and Pscudodistomi- dae. None of these contain genera with true cloacal systems although other colonial systems, or mudi- mentary cloacal systems, sometimes occur. Some narrow, stalked Ritlerellidae n.fam. (R, asynineil- rica, R. pedunculata) have the xooids in colonial systems with atrial apertures in a row along one side of the flattened, terminal edge of each stalk, and branchial apertures in a row on the opposite side. In other species (R. papillata n.sp., R, cornuta n.sp., R. compacta n.sp.) the atrial apertures are in the centre of the terminal end of lobes of the colony surrounded by the branchial apertures to form rudimentary cloa- cal systems in which excurrent water from all zooids is ejected in a single stream from the centre of the circular system. Similar rudimentary cloacal systems occur in Polycitoridae (see Kod 1989, 1990). Atleast in 8, cornuta n.sp,, ta situ photographs show the flat terminal end of the colony lobe directed away fram the oncoming current; and the excurrent steam of water from the centre of each end is entrained by the current as it passes the colony. The arrangement of the apertures of the separate zooids of Monniotuy spp. is similar to Ritrerella asytemetrica and R, pedunculata, wilh atrial apertures on the convex side of the spoon-shaped head, and branchial openings on the concave side. A large flap of testalong their upper margin directs the branchial apertures downwards (Kott 1989), Also in Protopalyclinidae n.fam,, colo- rdal systems of Condominium n.gen. have zooids arranged in single lines along flat fan-shaped lamel- Tae. Atrial apertures open in a lineon the convex sxie of the lamella, near and paralicl to the top of the colony, on the opposite side to the groove in which the branchial apertures are concealed, The mow of atrial apertares are also concealed in a grooye when zooids contract. Very likely the linear arrangement of zooids and their apertures in the hard, sandy, offen rigid testis a result of the pressures io direct and separate the respective incurrent and exeurrent ciliary sireams in these species with separately opening zooids. The compression of the atrial aperture into a transverse opening or slit and the division of its rim into an anterior and a posterior lip usually with 6 lobes on each (e.g. Monniotus, Condominium gen. nov. and some sandy Ritrerella spp.) further controls and di- tecis the incurrent and excurrent streams of water. The relatively large, usually gelatinous, separate zooids in Buherdmaniidac often have large external lobes around the apertures (c.g. E. dentatosiphontis, E. translucida n.sp.) that resemble those of Siomo- zoidae (see Kott 1990). although in the latter family the zooids are not separate, being embedded jn com- mon test, The function of these enlarged lobes around the apertures, which involve the test as well as the body wall of the zooids, is not established, It is possible that they help to direet the ciliary current, and, to some extent may act as sense organs through the pigment spots at their base which, like similor pigment spots in Crana, Diazonidae and some Phie- bobranchia (see Kott 1985, 1990), may be light sensitive, Generally, the Iobes around the apertures have musculature associated with the longitudinal muscle bands of the thorax, and the sphincter muscle that surrounds each aperture behind the lobes. Zoaids in stalked colonies of Pseudodisiomidec are arranged in the cojony like those of some species of Clavelinidac (Nephtheiz), and Holozoidac (Sigilline), all with atnal apertures uppermost wnd branchial apertures directed toward the stalk so that cach stalked head forms a single colonial (but noe cloacal) system as discussed by Kott (1989, 1990), In Polyclinidae, the zooids are arranged tn cloacal systems, cither around cloacal cavities, or along each side of canals that converge to common cloacal apertures, The atrial apertures open into tbe sides or base of these cloacal cavities or canals, The openings eilher are relatively large, exposing part of the bran- chial sac to the cloacal cavity (as in Sycozou or Distaplia); or there 1s a short, narrow protuberant siphon wilh a distinct sphincter muscle and a small terminal aperture. Each opening has a flattened ptrial lip (or tongue) projecting from ils upper rim or fram the body wall anterior to [he opening, This projection 19 Strap- of leafike, triangular or subdivided into 2 or 3, and its Gip is straight edged, pointed, bi- ut wi-dentate or serrated, Sometimes the circolar mus~ ele around the opening continues through the base of the lip, and muscle frbres from the sphincter exteod out inte the lip (e.g. Aplidiwm atewe, A. alfarinm andl the majority of Synoicurr spp.), Olher species, usu- ally with peotuberant siphons, have the sphincter muscle (and the aperture) independent of the athal lip (Aplidium crateriferwn and related species, and Polyclinwn spp.) The size of the atrial lip varies according to is role in effecting the closure of cloacal apertures uni contraction of the cloacal canals, An example can be seen in Polyclinm vaseulosum in which the anteti- orly ofiented atrial aperture is in the base of the cloacal canals, while the large atrial hp strewches along the top of the canal, and in some cases reaches and is inserted inte the (est around the cloacal aper- ture. ascending limb of gut loop: see gut, gut loup. ulrial aperture, — lip, — tongue: sce apertures: atrial cavity; In taxa in which 1 opens directly 10 the exieriut through a 6-lobed atrial aperture, the wtnal cavity, ils size varying with the size of the thorax, is arclatively contained chamber in which (often large) numbers of embryos complete their develapmeat to tailed larvac. Only in some Pseudedistoma is a sper cial incubatory pouch constricted off from the atrial cavity. In Polyclinidae the atrial cavity is a similarly com tained chamber in those species in which the atrial aperture is a small anteriorly directed circular open: ing separate from the atrial tongue (viz. many Paly- clinum, Sydneioides, and Aplidium species, anc some Synvicurn species). In the species in which the 382 MEMOIRS OF THE QUEENSLAND MUSEUM atnal fonguc is pan of the vim of the aperture and the circular muscles of the aperture diverge ont into the atrial lip, the aperture itself is less restricted, and when relaxed is a large opening exposing the atrial cavity directly to the cloacal cavity, Both types of atrial aperture are present in Aplidinm and Synoicum. No apparent correlation exists between the size and ferm of the atrial aperture and cavity and the size and number of embryos being incubated in the chamber. Only in Polyclinunt, in which small circular atrial apertures prevail, are the embryos and larvae consis- tently small (trunk appreciably less than 1 mm long). In this genus embryos also are relatively numerous {usually more than 5 being incubated at a time), Although large atrial cavities result from the rela- tively narrow branchial sacs in the wide thoraves, the embryos seldom fill the atrial cavity, and in many species they are removed entirely from it. being incubated in a broad pouch separate from the atrial cavity. Under these circumstances, the atrial cavity is not occluded by embryos. and possibly remains Functional during the period of their development. The atnal cavities of smail Aplidiurt zooids with short, narrow branchial sacs, are relatively small and are subject to further constriction by longitudinal muscles which extend the lengih of the zoords. Other variations in the configuration of the atrial cavity ocour as a result of the connectives which conduct transverse muscle fibres from the transverse: branchial vessels across the atrial oavisy to the paric- tal body wail. In the large zooids of Syricicumn cas- rellatem n.sp, a tow of these connectives appears lo divide the atrial cavity into dorsal and ventral cham- bers (see S. castellatum n.sp. below). It is possible that these chambers are present also in smaller zo- oads. but not so readily observed, In the Laxa reviewed below, the ventral pouches in the atrial cavity observed in Sycozea and Distaplia {see Kolt 1990) are not present, These are formed where the cavity extends yentrally over the perfo- rated area of pharyngeal wall but does not penetrate over the triangular areas of imperforate pharynx between the ventral ends of the rows of stigmata. (Sez also incubation of embryos) branchial papillae: possible vestiges of the branchial papillae which support the internal longitudinal branchial vessels in Phlebobrianchia, and same aplousabranch families (Cionidac, Diazonidae), oc- cur in Prolopolyclinidae fam. (Pretopolyelinum, Monniotus}, Ritterellidae afam. (Ritterella com- pacta nsp., 8 comma nsp,,&, multistigmata nsp., RX. papilkna sp.) and Polyclinidae (Polyetinumny. These brancteal papillae ane Mal, rounded to short tongue-like projections from the transverse vessels. Nothing is known of their function and their status as vestiges Of phiebobranch branchial papillae 18 speculative, branchial sac In the present group of families the branchial sac becomes progressively smaller, with fewer stigmata and fewer rows of stigmata, in paral- lel with the reloctions in zooid size, Similar trends can be observed in Polycitoridac (Pofyelior bo Eudis- toma) and Holozoidae (Sigillina to Sycozea and Dis- taplia). Placentelidae n.fam. have a large broad pharynx with many rows of numerous stigmata. In Protopolyciinidae n,fam, the number of rows of stig- mata as well as the length of each row are as great or greater (up to 20 rows of up fo 50 stigmata in Can- dommium n.gen.) than in Placentelidae nfam. In Ritteredlidae n.fam, the zooids are-reduced in size and although stigmata are in up fo 16 rows, often there are only 5, and in Dumus only 4, The number of stigmata in each row are similarly reduced from a maximum of 35 to 12. The branchial sacs (together with the solitary za- oids) of Exherdmania, which does not replicate as freely as other taxa, remain relatively large wath numerous rows of up to 100 stigmata per row, Psendodistomidae o.fam. have 3 long rows of stig- mata, ina wide pharynx. The greatest size reduction in zooids can be ob- served in the polyclinid genus Apliditum in which prolific replication occurs, and zooids are arranged in cloacal systems. In this genus the stigmata occa- sionally are in only 5 rows. Although the length of the sugmiala offen is reduced, the number of rows is not, and most species have from [2 to 20 rows of stigmata. Synoiewn species have similar reductions in size of the branchial sac. The most conspicuous difference between these genera and other polyclind genera is they reduchyn in the size of the perliyrated pharynx, Polyelinium and Aplidiopsis have rely- tively large branchial sacs. Internal longitudinal branchial vessels are present only in Cionidac and Diazonidae; and the only ves- sels in the branchial sac of all other aplousobranch taxa are transverse. The primary transverse vessels are he unperforated expanses of the pharyngeal wall between the rows of stigmata. On their mesial side, the walls of these vessels are ciliated endoderm projecting into the lumen of the pharynx, On the parietal side the epithelium is ectodermal and not ciliated. Transverse muscle fibres are in the trans- verse vessels and these join the muscles of the paric- sal body wall through connectives that cross the atrial cavity (see atrial cavity; Syroicum casrellatwm n.sp., below) Parastigmatic vessels, extending across the stig- tala halfway between the primary transverse ves- sels, Sometimes subdivide the stigmata along their length créaling additional rows, More often they occur in species with Jong stigmata and persist as inconspicuous strands which hold the delicate inter- stigmatal bars in place, and prevent distortion of the stigmata. Parastigmatic vessels occur in Protopoly- clinidae n.fam (Monaiotus spp, and Condorminiuan n.geo.) and Rittercilidacn. fam. (c.g. Riverella asyn- metrica, R, multistigmata n.sp.), Despite the laree size of Euherdmania zooids, the stigmata are rela- tively short and disposed in numerous rows. Paras- ligmnatic vessels are not known in this genus nor are they known in the short branchial sacs of Pseudodis- THE AUSTRALIAN ASCIDIACEA 3 tomidae n.fam. In Polyclinidae they are known in some species of Aplidinm with relatively large bran- chial sacs (e.g. Aplidium amorphatum, A. parastig- maficun 1.3p.). branchial tentacles: although these are not specifi- cally counted in the descriptions that follow, a ring of short, relatively stumpy tentacles of various sizes is always present at the base of the branchial siphon, protecting the incurrent aperture. Intrageneric vari- ations in the tentacles do not appear significant tax- onomically. branchial vessels: see branchial sac. brooding, brood pouch: see incubation. budding: see replication. cloacal cavities, — systems: As in Polycitoridae (see Kott 1990) some other taxa with independently opening atrial apertures and embedded zooids have their zooids arranged in incipient or rudimentary cloacal systems. Thus, in Ritterellidae nfam., R. papillara nsp, and RK, cormura n.sp. have zooids arranged in circles with their atrtal apertures in a depression in the centre of the circle and their bran- chial apertures around the periphery; and in &, com- pacta n.sp. the excurrent apertures open into a circular chamber fined with sand which appears to be an invagination in the surface of the colony. In Polyclinidae a high degree of organisation mito cloacal systems is achieved. These systems range from simple circles of zocids around circular cham- bers with acentral cloacal aperture to rows of zoaids arranged along each side of branching canals which converge to a cloacal aperture. Sometimes partitions of zooid-Free tesi occur between adjacent systems, or between parts of the same system (i.e, between cun- verging canals); but in other species the systems are crowded together. In preserved or contracted colo- nies, zovid-free areas of test, being solid and without cloacal cavities or chambers containing zooids, are prominent and usually protrude from the surface. This is an artefact, For in living. extended specimens, the cavities and zooids are expanded and the surface over them is not depressed, In Polyclinidae there are relatively few taxa (com- pared with Holozoidae) in which the colony is a single system (e.g. Aplidium australiense, Paly- clinum orbitum nsp.). More offen colonies, even those: with large, regular, citcular systems are rela- tively irregular, and contain a number of cloacal systems, each with a conspicuous cloacal aperture. The systems have the capacity to subdivide forming relatively large, bulky colonies. Despite the capacity of Polyclinidae 10 form large colonies of one or more systems, the cloacal cavities themselves are rela- tively narrow canals and small, rounded chambers rather than the extensive 3-dimensional cavities of Didemnidae. The strength and rigidity of the test, especially when this is firm and gelatinous, or contains embed- ded sand and/or calcareous particles contributes to the maintenance of colony shape including surface features such as ridges and grooves that affect the 383 organisms’ anteractions with the environment (such as direction of current flow: Kott 1989), The pressure of excurrent water in the cammon cloacal cavity (when one is present) also contribules to mainte- nance of the shape and the configuration of the surface, of the colony. Thus, bath factors coninbut- ing to the strength and ngidity of the test (such as embedded sand), and also to the turgidity of a large, 3-dimensional cloacal cavity (such as efftcient repli- cation increasing the number of zooids contributing to the internal pressure within the cloacal cavihes), constitute adaptive advantages favoring the evola- tion of large, 3-dimensional colonies. In some species of Polyclinum, the surface test has a thin, brittle layer of sand but the internal test is soft, In this case their round colony shape may be main- tained by the water pressure in the large pharynges of the crowded zooids rather than the baittleness of the surface layer of test, (See also apertures. colo- nial systems, colonies). colonial systems; In other taxa with independently opening atrial apertures, the zooats are notembedded in common test; of are arranged in colonial systems other than cloacal ones; or, rarely, do not have any obyjous systematic arrangement in the colony, Mon- niotus and Ewhendmania wooids eviher are solitary; or separate from one another, asiached to common test or stolons only at their posicrior end, ane, like separate Diazonidae. Clavelinidae, and Pycnoctivel- lidae, function as solstery individuals and do wot form systems of any kind. As with clogeal systems, other sorts of systems have zooids 40 arranged to ensure that the functioning of the whole colony exceeds the efficiency of the sum af ils component soos. Although zoojds of Rirerelfa compacta asp, R, conmuda asp. and R pepillata o.sp. are in circular Tudimentary cloacal syslems with their alrial aper- tures ba the centre, &, asyrimerrled, R, peduncwlare, R. mauftistignate rsp. und Condominium gen nov. have their zvoids arranged around fan-shaped stalked colonies, with the branchial and atnal aper- hires on oppasite sides of the colony margin. Incur- rent and excurrent streams, respectively, reinforce one another, and are Kept apart; and the long narrow siaiks enable the colony to change its anentation wilh the prevailing currents, Stalked Psevdodistema with all atrial apertures directed to the top of the cotony and the branchial apertures directed downwards wywards the point of attachment sre equally flexible with regard to changes in current flow. Other benefits from this uniform arrangement of apertures include protection of the open incurrent apertures from falling sexfj- ments (Kott 1989; see also apertures, cloacal sys- tems). volonies: The size and form of colonies reflects their phylogeny, loughness of test and amount and type of embedded material. More primitive families such as Diazonidae, Clavelinidae and Pyenoclavellidac, de not usually fonm particularly large self-supporti colonies, However, generally other aplousobranc: 384 families have firm and gelatinous test, resembling the test of phlebobranch families in consistency; or it is strengthened externally or throughout with sand or other hard (including calcareous) particles. When strengthened, large often complex self-supporting colonies are formed. Occasionally species such as Pseudodistoma fragile and P. gracilum with particu- larly soft almost fluid test also have relatively large colonies, possibly occupying sheltered habitats. The complexity of colonies of the Ascidiacea is most evident in colonies in which the embedded sand makes the test hard, brittle and allows it to form massive 3-dimensional branching colonies (e.g. A. solidum); or in intricate colonial systems (e.g. A. crateriferum, Condominum areolatum n.gen., n.sp.). However, although complexity in shape appears to require sand or other inclusions in the test, the size of colonies per se is not related to sand inclusions (e.g. gelatinous colonies of Polycitor and Aplidium become as large as some of the sand-strengthened colonies in these genera). As well as affecting the size and form of the colonies, the consistency of the test, which is directly affected by the type and quantity of material embed- ded in it affects the configuration of the surface test, which, in turn, can modify the currents flowing over the colony and influence its interactions with the environment. In many Ritterella spp. and in Monnio- tus a layer of brittle sand-embedded surface test protects the incurrent and excurrent apertures and separates the currents flowing through them. In Poly- clinidae the operation of the colony is enhanced by its ability to sort sand and to determine precisely its distribution. For instance, in Aplidium tabascum n.sp. a single line of fine sand particles outlines the margins of the firm zooid free areas that protect the branchial apertures; and in A, caelestis and A. macrolobatum n.sp. fine sand particles are embed- ded in the thin, flexible test over cloacal cavities, while coarser grains are in the rigid test between the systems. Colony size and shape does not appear affected by the development of cloacal systems, except that colo- nies consisting of a single system are a finite size and shape (e.g. Aplidium altarium n.sp.) and generally are smaller and more regular than those with multiple systems. Environmental pressures such as substrate and current flow probably have most influence on the size and shape of the colonies found in any habitat. Thus stalked species appear to occupy habitats where surge alternates the direction of water flow past the zooids; and large spherical, oval, or irregular colo- nies occupy sea floor habitats at depths where strong currents are not likely to dislodge them from sub- strates. Extensive areas of sandy substrate favour large mixed populations of Monniotus radiatus n.sp., Du- mus areniferus, Ritterella pedunculata, R. asymmet- rica, and R, papillata n.sp., which all have similar fine, sandy, finger-like colonies embedded in sand MEMOIRS OF THE QUEENSLAND MUSEUM — the individuals being more or less held together and stabilised by the sand that accumulates around them (see Kott 1989); and, at the same time, they stabilise the sand. As well as these narrow stalked colonies with hair-like test projections and basal stolons that form sandy aggregates and mats in and over the substrate (e.g. Pycnoclavella spp.: Kott 1990, Ritterella spp., Monniotus spp.), there are other types of colonies that stabilise sandy substrates and protect them from erosion. These are colonies with basal test forming root-like projections that hold them firmly in the substrate (e.g. Aplidium gemi- natum n.sp.), and colonies embedded in the substrate and forming 3-dimensional reticula and branching lamellae (e.g. Aplidium acroporum n.sp., A. petro- sum n.sp., A. solidum, and Polyclinum terranum n.sp.). (See also cloacal cavities, colonial systems). descending limb (of gut loop): see gut. dorsal lamina, — languets: No continuous fold occurs along the mid-dorsal line of the pharynx in Aplouso- branchia as in some Stolidobranchia and Phlebo- branchia. Instead pointed languets arise from the transverse vessels to the left of the mid-line, and curve over to the right. These channel the mucous food cord (gathered from the sheets moving dorsally over each side of the pharynx) as it moves posteriorly to the oesophagus. duodenum: see gut. epidermal ampullae: see ampullae. epidermal vesicles: see vesicles. excurrent aperture: see apertures. fertilisation: About half the extant Ascidiacea are soli- tary species which, with few exceptions (see below), are externally fertilised. The other half are colonial, and these, with the exception of the Diazonidae, are fertilised internally. The colonial species are pre- dominantly Aplousobranchia, although taxa of Phle- bobranchia (Perophoridae) and Stolidobranchia (Botryllinae and Polyzoinae) are included. Solitary species are predominantly Stolidobranchia and Phle- bobranchia, only 2 genera of Aplousobranchia (Ciona and Rhopalaea) being included. With the exception of Polycarpa and some Mol- gulidae and Agnesiidae, solitary Ascidiacea have long and relatively few male and female gonads, and their ducts open together near the anus, at the base of the atrial siphon. Released gametes thus are ejected, and presumably dispersed with the excurrent ciliary stream, Gametes also are released in the excurrent water as the animal rythmically squirts to irrigate the atrial and branchial cavities. In both cases waste products and gametes are projected well away from the animal where they are not likely to contaminate the incurrent stream (see Hoshino ef al. 1986, Kott 1989b), If both male and female gametes of the same mature individual are released together there would be opportunities for self-fertilisation. However there is growing evidence that individuals are generally self-sterile (Santis and Pinto 1991, Fuke 1983, Kawamura ef al. 1987, Cloney 1990, Litscher and THE AUSTRALIAN ASCIDIACEA 2 385 Honegger 1991), Further, specics are not necessarily functional hermaphrodites, and may be either pro- tandrous or protogynous. Other mechanisms may exist to prevent self-fertilisation, one being the pos- sibility that ejected sperm are dispersed further away than ova. The stimulus for release of gametes is not under- stood, although their are obvious advantages for one to exist. Under certain conditions light is known to trigger gamete release. Carlisle (1951) proposed that Ciena intestinalis released gametes and stopped se- creling mucus from the endostyle in response to the presence of a gamete of the same species in the incurrent fecding stream. Although it is an apparent advantage for mucus secretion to stop when fertili- sation is internal, it is not for externally fertilised species. In Polycarpa, short gonads and their duets are scattered over the body wall, and the openings of the ducts are removed from the atrial aperture, Presum- ably this results in longer retention of gametes of both sexes inthe atrial cavily than would be the case if they were released at the base of the atrial siphon, In certain species (P. infenaia, P. linctor, P. tinc- torella: see Kott 1985) even longer retention results from the ventral orientation of the short gonoducts, and indeed these species are internally fertilised and embryos are brooded in the atrial cavity. Certain molgulid species are analgous io Polycarpa in re- spect to the arrangement of their male and female gonoducts, and also.are internally fertilised, brood- ing, species. They all have the oviducal apertures turmed ventrally, and the openings of the-vas deferens distant trom the alrial aperture, Molgula calvaie and M. manhattensis (see Kott 1985) have short, multiple openings of the sperm duci along the length of the ovary; and M, ellistoni, M, sabulosa and Eugyraspp. {see Kott 1985) have a short yas deferens, at the proximal end of the ovary, turned ventrally. Al- though the reproductive sategy of Adagnesia spp. is not known, both gonoducts are turned ventrally (see Kott 1985) and probably embryos are brooded internally, Only one species of the Pyuridac, Pyura littoralis (see Kott 1985), has short female ducts while retaining a long vas deferens, This may be an adaptation for internal fertilisation, although ete bryos have not yet been found in the atrial cavity, [tis not known if these internally fertilised solitary species are self-sterile or not. tf diey are, the reasons for the adaptalions found in the majority uf these species, which apparently ensure the retention of sperm as well as ova in the atrial cavity, are difficult to understand, Forther, if they are selfsterile, it is possible that mucus secretion is interrapted ta allow. sperm for cross fertilisation to reach the atrial cavity through the pharynx. In Ecreimascidia turbinata, wt least, 1Lappears not lo be drawn in through the airial aperture with relaxation of body muscles following contraction and squirting (Ryland pers. cami.), Whereas only pelitively few solitary species have evolved devices for eternal fertilisation, this serat- egy is universal in colonia) species, and once it has evolved il appears never to be Jost. The family Dia- zonidac, which contains some genen retaining a solilary habil, and the related and supposedly primi- tive: solitary Cionidac: are the only aplousebranch tax externally fertilised. In colonial internally fertilised species (as in sali- tary ones) the position of each gonoduct relative bo the others and to the atrial aperture prohably is ee- lated la patterns of gamete release and lertlisalion. Although the oviduct is often displaced when it acts ax a brood pouch, the vas deferens and -anus of colonial species open together, usually part of the way up the thorax, and not necessarily close (> the atrial aperture, The atrial cavity, although it is often distended with embryas, never contains faeces out- side the rectum, and presumably faecal pellets are released by the contraction of the body wall muscles, which irrigates the atrial cavity and brings the anas close to the alrial aperture, Since the opening of the vas deferens is associated with the anus it is possible that gametes are also released inthis way. In bolozoid ener with cloacal systerns, anc in the Didemnidae, arge atrial apertures often expose anal and gonodu- cal openings direetly to the cloacal cavity. This alsa happens in some Polyclinida: (Aplidiun, Symoican anda few Polyclinum). Thus, the opening of the mabe duct of colonial, internally tertihised species (as in solitary externally fertilised ones) is usually posi honed where sperm released from it are most likely to be expelled to the exterior, either directly or through the cloacal cavity, rather than being retained in the atrial cavity for self-fertilixation. It as not know whether or not embedded colonial zoants have a pattern of rythimie squirting, like solitary and independent colonial ones, to increase means by which sperm might be released, Further, colonial, like solitary, species have been shown to be self-sterile (e.g. Diplosoma listerianuns: Ryland and Bishop 1990; Balryllus schlesseri. Sab- badin 1979), In addition to the growing evidence af evtological and chemical mechanisms for self-steril- ity, the existence of functionally gonochoric colonies jn the genus Sycozow (see Kott 1990), and of sequen- ually hermaphroditic colonics in many other axy, support the view that colonial, as well as solitary, ascidians are not generally self-feriilised, Sabbadin (1979) reports a protogynous condition for Beryl Ins. Purther evidence of protandry or protogyny ex- isis when embryos but not mature male gonads are present and when there are mature male gonads hut no embryos sygeesting that the male gonads nature either before or after the female ones (see below). If, as the evidence suggests, intemally Jertilised colonial species are not self-fertilised, sufficient con- centrations of sperm must reach the atrial cavity from outside the zooid for sore to enter the oviduct, and move to the fertilisation chamber (the brood pouch, or the proximal cad or top of the oviduct, or the atria} caviry) atthe time that oncylrs meech matierity. Thus it is probable that advantages associated with the synchronous release of gametes are as preat in inter- nally fertilised colonial species as they are ily exter nally fertilised solitary ones. especsally as colonial zpoids are small, with relatively small ovaries, few ova and often ephemeral gonads. Again, little is known of mechanisms trizgenng such a synchronous release. Further it is not known how sperm enters the atrial cavity, Ryland (pers. comm.) found that exter- nal material was not drawn into the atrial cavity of Keteinasctdia terbinate following Hs partial evacu- ation by squirting, [tis nat kown whelher or not embedded zooids do squin, however even if they do it seems unlikely thal bated in the atrial cavity, Only in Pol yes, which hos zooids with lange roomy atrtal cavities, do the 388 Adhesive organs 5 ri r No vesicles No vesicles Ampullae Pycnoclavaliidae Sigillina mjébergi Euherdmanildae No ampullae Thin-walled everling tubes MEMOIRS OF THE QUEENSLAND MUSEUM With vesicles (except In some simplitied Polyclinidae) o Vesicles separate from z » paired postero-ventral, J dorsal, and sometimes along lateral, and sometimes Pp ermal strands ra ape’ ‘ s 88 anterior epidermal strands dorsal: epidermal:t median lines Vesicles separate Vesicles separate Irom directly fram epidermis palred ventral and Pseudodistomidae everting cylinders Thin-walled or spheres Sigillina Hypodistoma Eudistoma Polycitoralla Clavelinidae wider than deep Axial cones or platforms large: Holozoidas (except: Sigillina Hypadistama) Polycitor Cystodytes moderate to large: deeper than wide Axial cones = a i] 2 s zy = o a oD 2 = £ a a — .2) w a o g a a oO 2 = a E G £ = Protopolyclinidae Axial cones small: wider than deep Ritterallidae (not aequalisiphonis group) Polyelinum Synoicurm spp. |— Synoicum spp. Aplidium luneeratum Aplidium multilineatum Aplidium omatum Aplidium protectans Aplidium robustum Aplidium tabascum Aitterella spp. (? agqualisiphonis group) Synoicurn spp. Aplidium spp. TABLE 2. Summary of epidermal elements at anterior end of larval trunk in aplousobranch families. particularly small embryos (larval trunk seldom more than 0.6 mm Jong) appear not to occlude the pharynx, even when relatively large numbers are present at a time (up to 12 embryos: P. fungosum). Nevertheless, it is in this genus that a brood pouch sometimes is present (P. marsupiale, P. tsutsuti, P. terranum n.sp.) constricted off from the body wall in a unique position, about halfway along the dorsum of the thorax at the level of the anus. It is most probable that fertilisation as well as incubation of embryos, takes place in it; or in the atrial cavity. Only very few Aplidium species (A. uteute, A. altar- ium) have a brood pouch, at the postero-dorsal corner of the atrial cavity, in which embryos are incubated. In this genus, fertilisation generally appears to be in the top of the oviduct or i the atrial cavity. incurrent aperture: see apertures. larvae: The larvae of the taxa formerly grouped in the single family Polyclinidae, as originally defined, demonstrate its poly phyletic nature. The larvae of the new families proposed below can be characterised by their size, adhesive organs, ectodermal vesicles and ectodermal ampullae (see Table 2). The Euherd- maniidae have adhesive organs that are invaginated tubes that evaginate on settlement, as in the family Pycnoclavellidae; the relatively large larvae of Pseudodistomidae n.fam., as do Polycitoridae, have large adhesive organs and ectodermal ampullae be- THE AUSTRALIAN ASCIDIACEA 3 389 tween and lateral to the median adhesive organs; the generally smaller larvae of Polyclinidae emend., Pro- topolyclinidae n.fam., and Ritterellidae n.fam. usu- ally have ectodermal vesicles which separate off from the larval epidermis into the test between and alongside the median adhesive organs. There are also differences between genera in the origin of the ec- todermal vesicles (see vesicles below). All the larvae of the families discussed below have a discrete mass of yolk cells in the middle of the trunk. Larval ocellus and otolith are present in all genera. Although Euherdmania has similar adhesive organs to Pycnoclavella, the larvae do not appear to have a tendency to loss of the ocellus as do those of Pyenoclavella. With the exception only of Euherdmania in which the invaginated tubes are triradially arranged, the 3 adhesive organs are in the anierior median line. Only Aplidium triggsense (with 12), and A. multipapilla- tum Millar, 1975 (with 8), have a different number from the dorsal mid-line. Ventrally they divide into branches each side of the endostyle. The large atrial lips of Polyclinum and species of other genera in Polyclinidae are also muscular. Branches from the ventral thoracic longitudinal mus- cles extend out into these tongues, and circular mus- cles from around the atrial apertures also are diverted out along them. These large atrial lips insert into the test to protect the atrial openings into the cloacal cavities, and in some cases, insert into the rim of common cloacal apertures and exert some control over these excurrent openings. Anteriorly, the longitudinal muscle bands extend onto the siphons, where there is exchange of fibres with circular (including sphincter) muscles, longitu- dinal fibres are then organised into bands, one ex- tending into the each lobe around the rim of the aperture. When these muscles are concentrated into the tip of each lobe they depress it and divide the tip of the lobe into 2 (e.g. Ritterella dispar, Aplidium macrolobatum n.sp.). of adhesive organs, all in the median line. muscles: With the exception of sphincter muscles around the apertures, the body musculature in the parastigmatic vessels: see branchial sac. posterior abdomen: see gonads. replication: In these well developed aplousobranch parietal body wall is usually longitudinal, although fibres are exchanged with adjacent bundles. Trans- verse fibres are present, forming a network with the longitudinal ones, only in Ritterella multistigmata n.sp. and Anadistoma n.gen. These transverse fibres may originate as branches from the longitudinal muscle bands, Also in Ritterella, oblique muscles similar to those in the Clavelinidae cross the thorax from the endostyle to the postero-dorsal corner of the thorax. Longitudinal and oblique muscles extend in a band along each side of the abdomen and posterior abdomen, as they do in most of the genera. Only some Aplidiopsis species, Polyclinum and Sid- neioides are exceptions with short longitudinal mus- cles confined to the anterior part of the thorax. In some Pseudodistomidae n.fam. (P. oriens n.sp., P. candens n,sp) at least some fibres of the longitu- dinal muscles extend along a vascular stolon poste- rior to the posterior abdomen and heart. The presence of these muscles on the long, distinct vascular stolon is reminiscent of Sigillina (Holozoidae, see Kott 1990). The presence of both a gonad-containing posterior abdomen, and a vascular stolon with mus- cles (apparently homologous with a similar organ in Sigillina) in certain Pseudodistoma species suggests that the vascular stolon of Holozoidae is a separate structure rather than being homologous with the posterior abdomen of Polyclinidae, Ritterellidae n.fam., Pseudodistomidae n.fam. and Protopolyclin- idae n.fam. In all taxa, transverse fibres are present in the transverse vessels. These are fibres that derive from longitudinal muscle bands, These muscles are seen clearly in the large zooids of Synoicum castellatum n.sp., branching off longitudinal muscles and enter- ing the transverse vessels through connectives from the parietal body wall about one-third of their length families replication takes place by horizontal divi- sion of the posterior abdomen and/or the abdomen, and usually is prolific. The process has not been specifically studied in Australian species. However the process is similar in the range of polyclinid species studied by Berrill (see 1935 for review), Nakauchi (1966-1987), and Nakauchi and Kawa- mura (1974a-1986). The process is the same in Ritterella (see Nakauchi 1977). Generally reserve food cells (trophocytes) pack the abdomen and posterior abdomen, Sometimes the thorax and abdominal organs are resorbed but in other species they remain functional during the budding process (see Ritterella pulchra: Nakauchi, 1977), The abdomen and/or posterior abdomen are then divided, by constric- tion of the epidermis, into numerous buds. A segment of epicardial tube is contained in each, and is the regenerative tissue for the new vegetative zooid which develops from each bud. In Polyclinum vegetative zooids with relatively long strap-like vegetative posterior abdomina (as opposed to the short sac-like posterior abdomen of sexual zooids) are found crowded in the growing margins of colonies (see P. vasculosum). Although moderately long, sometimes branched, vascular appendices are present in Ritterellidae, and shorter ones are present in most of the other families, these seem not to be involved in the process of replication as they are in Holozoidae. Pseudodistomidae n.fam, have a long vascular stolon with longitudinal body muscles extending along it as in Sigillina (Holozoidae). This supports the view that this family is related to Sigillina (Holo- zoidae), The method of replication in the Pseudodis- tomidae n.fam. is not known, nor is it known that the vascular appendix has a role in the process. Although Brewin (1958) found ‘bud-like masses’ lying be- neath the zooids of P. ceream, and juvenile replicate zooids are Found in the stalk of developing lobes of P. onens 1.sp., itis not known whether these developed from the vascular stolon ina process that resémbies that of Holozoidae (see Kott 1990); or if they developed from the posterior abdomen and/or abdomen of the parental zooids by horizontal strobilation, as in Poly- clinidae, A continuation of the epicardial tube into the vascular stolon of Pseudodistoma has not been found. stomach: see gul. test: see cloacal systems, colontal systems, colonies. trophocytes: see replication. yascular appendix. — process, — stolan: see repli- cation. vesicles, ampullary —, epidermal —, larval —- Hol low spheres. about 0.01 to 0.05mi Gameter with an epithelial wall, usually occur in the test of the larval trunk in Protopolyciinidac, Ritterellidac and Poly- chinidae. The fine strands attaching them to the larval epidermis sometimes sever as larvae mature. Like the enlarged epithelial cells on the tips of some ampullae the epidermal cells occasionally are larger on the outside surface of the vesicle than on its mesial wall. In Monniotus (Protopolyclinidac), Polyclinwr and Synoicum, the vesicles exclusively branch off fine stands that detach from the epithelial wall on each side of the mid-line and trail through the test. The ventral strands anse from about one half to two- thirds of the way from the anterior end of the trunk, one extending 10 the posterior end of the trunk near the posterior-horns of the haemococlic cavity where the: vesicles forma tight cluster, and another shorter ope trails anteriorly. Similar puirs of cpithelial strands from the anterior end of the dorsal surface trarl posteriorly On each side of the dorsal miid-line to the vicinity of the branchial apertare, and in Mori- niotes acoresponding pair of strands extend antero- ventrally on cach side of the adhesive organs. ‘The vesicles are especially numerous posterc-ventrally. Dorsal and ventral clusters of vesicles arising from similar test strands are present in some Jarvae of Aplidiam (A. lunacrafien nom. noy., A. miulfilinea- tun a.sp., A. ermatum asp. A. profectans n.sp., A- robustum nusp. and A. tabascam n.sp.), but the pos- tend-ventral test strand is much shorter and a posteror ventral cluster of vesicles near the base of the til does nol pocur. Larvae of the majonty of Aplidiarnn species have vesicles directly attached by their shor sometimes branched hair-like filament to lhe larval epidermis rather than to an epithelial strand trailing through the test. These vesicles are arranged in P. de- pressum Herdman, 1899) appears not properly as- signed. It has a long abdomen distinguishing it from genera of Polyclinidae. It is said to have incubatory pouches packed with embryos (Herdman, 1899). It could be a species of Pseudodistoma, no other known genus having a brood pouch as well as both a long abdomen and a posterior abdomen, although brood 445 pouches of Pseudodistoma cannot be said to be ‘packed with embryos’. The number of rows of stig- mata is not known. Polyclinum constellatum Savigny, 1816 from Mauri- tius, has flat-topped colonies attached by a small part of the basal surface. They are without sand, and have branching double rows of zooids radiating from cir- cular common cloacal apertures. Polyclinum festum: Kott and Goodbody, 1982, from Hong Kong, with large potato-shaped colonies, blue-green when liv- ing, and crowded stellate systems of zooids, may be a synonym of P. constellatum. The loaf-like P. con- stellatum; Michaelsen, 1923 from Hong Kong and other specimens from Hong Kong, Mozambique and St. Helena, all with radiating rows of zooids are probably also conspecific with P. constellatum, as Michaelsen (1923) thought them to be. The branchial sacs are relatively wide with 18 to 22 (16 to 18: Kott and Goodbody 1982) stigmata. Small papillae are present on the transverse vessels. Further probably conspecific populations are from the Atlantic (Flor- ida, the West Indies and possibly south to Rio de Janiero), and the West Indian Ocean to Ceylon (Michaelsen 1923). Polyclinum constellatum: Tokioka, 1967 has indistinct systems. Although the zooids, like those of P. con- stellatum, have 14 to 18 rows of stigmata with 15 to 20 stigmata per row (more in specimens from the Palau Is), and distinct branchial papillae, the sandy coating on all Tokioka’s examined specimens sug- gests they are not P. constellatum. Polyclinum crater Sluiter, 1909 has simple circular systems, each with the branchial apertures surround- ing a central cloacal aperture in a surface depression. The body wall has strong muscles. About 9 rows of 6 to 8 stigmata are present. The outer surface of the test is smooth and slippery. Polyclinum festum Hartmeyer, 1905 (see also Millar 1975) from Mauritius has flat-topped colonies with circular systems. Sand is present externally (around the base and outer border, and sometimes along radii across the upper surface) as in Aplidiopsis amoyensis and related species (see below). Polyclinum hospitale Sluiter, 1895, from Thursday I. is smooth on the upper surface, with debris and sand on the under surface. Sluiter did not see either cloacal openings or systems, but both are likely to be present — the atrial opening having the usual pointed lip associated with a cloacal system. The body wall is muscular — an unusual condition for Polyclinum. The gut loop (including the oesophagus) is shorter than usual for this genus, and is not twisted. There are no branchial papillae. It is probable that the colony is an Aplidium species. Polyclinum gelidus (Monniot, 1987) from New Cale- donia, with a brownish-yellow colony resembles P. vasculosum, having no branchial papillae. Monniot (1987) assigned the species to the genus Aplidiopsis, MEMOIRS OF THE QUEENSLAND MUSEUM 446 ‘ainpiode jo wu jo ued ‘y ‘ainyode jo wu Woy ayeredas ‘¢ 5 ‘qeotdon ‘n ‘ayeraduray ‘ay ‘orylowg 1saAQ-Opuy ‘qAA] OYloeg Wasa, ‘qA\ ‘ea00 ayloeg ‘gd ‘snouasiput *y i 389] snouneyad ‘uy sso auou OL FI-ZI di ie ens sseg nd wmnsopnrsva'_ “ cl “ 61‘€I-Z1 s peyouriq «TP PeZr] 0} |] vols Nd wnsqna “q pepunor « MOI ZTqQnOp oF qsa] snouneyad ‘jos c0 mMoyyeys SL‘9I-F1 a Jeno poyeu InogIeHy syop ay ‘ds'u wnpnu “¢ aqoyuaysds | SPO “ P‘07-81 “ “ “« WeNs sseq ay ‘dsuwnyqio“g ‘ds‘u asreds AJaaneyal pues — o ZET x 7 « -Plyensny wamnosg ay unypnual “gf “ L0 period S191 u “ “ Ps wingys0D ay umnuD del “d “ s0 “« 0791-41 “ Jey “ Byensny AN Wd TNSINSS PIUCUISE], yonod poo1q c0 pedrys-an3u0} LU€I-91 é A a ‘eleasny ynos avy apoidnsaipu “J spo “ 0721-8 “« “« “« ‘] vorayy dA UNTRATDS od, 90 « PI'sl “ “ “« Byegsny ynog vy Mnqoisn419ul “ ss'0 pequtod ‘Teurs pl€l-ZI ” « ASN O1teng sseg ary winsosunf ‘gq SMOI a]qnop ] UOID} 0} Aou “WOU uetuopgqe JoLa\sod “suoy — auou rLSI-+1 Ss 0} JepNoI19 Apurs ‘Avg A@arapy Nd ‘MUNJOS “gf (urur) tyduey prenisny UvIpeul AMOI/‘OU XeUL punor yuny evyided {SMO ‘OU dy asTMyDO;OHUR osuel z 1 Relive) TRAE] yetyourlg Reus rethy surmsXg — sorjing asuey orydesZ0as01g satoeds ulfeNsny WoI peplosar wnu1jo{jog Jo satoads Jo sioyowseyo Jo Areumuuns *s ATAVL THE AUSTRALIAN ASCIDIACEA 3 however its abdomen is characteristic of the genus Polyclinum. ? Polyclinum nigrum Herdman, 1906, from Sri Lanka, has a smooth shiny surface, and circular depressions in the surface, which may have contained zooids, although these are not present. There is no evidence of the genus of this. colony, The type specimen of this species from Port Jackson (AM U33, Polyclinum nigrum Herdman, 1899) and another (MV F59436) from the same locality have been re-examined and found to be Sigillina (see S. nigra: Kott 1990). The zooids are contracted, possi- bly causing Herdman to misinterpret the length of the oesophagus, Palyclinum sundaicum (Sluiter, 1909), from the west- ern Pacific (Fiji, Indonesia and Hong Kong: see Kott 1981, Kott and Goodbody 1982) are small stalked or sessile sandy colonies cach containing one or more circular systems. They are distinguished from P. fungesum and P. selum nom. noy. by their large branchial papillae. Polyclinum tralatica Shuiter, 1913 from Aru L. is an irregularly Jobed, cylindrical, gelatinous mass along a bryozoan or gorgonian stalk, Its outer surface grey with a pale violet hue. The body wall is moderately muscular. The species resembles P. vasculosum in the absence of branchial papillae and in its LO rows of about 10 to 12 stigmata Polyclinum fungosum Herdman, 1886 (Fig, 25) Polyclinum fungosum Herdman, 1886, p,190. Polyclinum marsuptale: Kott, 1976, p.60. Synoicum hypurgon: Kott, 1976, p.62. DISTRIBUTION New RECORD: Victoria (Bass Strait, QM G12748), PREVIOUSLY RECORDED: Victoria (Westernport — MV F59365 F59431-2 Kott 1976). New South Wales (Port Jackson — Herdman 1886). DESCRIPTION EXTERNAL APPEARANCE: The small (about 5mm diameter) newly recorded specimen from Bass Strait is mutilated, possibly stalked, mush- room-shaped, with a firm, translucent test, an external layer of sand, and some sand internally. The zooids are crowded, obscuring the form of the systems. Larger (to 4em diameter) colonies (MV F59431) are soft, mushroom-shaped, fixed by a.small part of base or with a small stalk. In 447 preservative they are collapsed, with a wrinkled, furrowed surface. Sand encrusts the sides and under surface, is in patches on the upper surface, and is sparse internally, INTERNAL STRUCTURE: Zooids are about 3 to 4mm long, with a narrow thorax, small horizontal c____ <4 Fars ——— Imaganee000000 ——— donno go = FIG, 25: Polyclinum fungosum — a, colony (MV F59431): b, zooid with embryos i in atrial cavity (MV F59431); ¢, dorsal part of portion of branchial sac showing, kink in transverse vessels (MV F59431); d, larva (QM G12748). Scales: a, lem, b, 0.5mm; e, 0.2mm; d, 0.1mm. 448 gut loop, and sac-like posterior abdomen, The branchial lobes are triangular and pointed, and the atrial lip is Jong and narrow. Six to 10 fine longi- tudinal muscles are present on the anterior half of the thorax. Fine circular mus¢les are present in the transverse yessels between the rows of stigmata. The branchial sac is narrow, with 12 or 13: rows of 12 to 14 short oval stigmata. A short expansion of the transverse vessels on each side of the dorsal languets in some of the zooids could have resulted from contraction of the transverse muscle fibres in the vessels. The branchial papillae on the mar- ein of the transverse vessels are small, shallow, triangular protrusions about one to every one and ahalf stigmata. The gut forms the usual smal). horizontal loop with a smooth-walled stomach, a Jong duodenal area and an oval posterior stomach. The sac-like postenor abdomen has a short neck and is pointed posteriorly. Male follicles are clustered behind and to the side of a relatively large ovary, which has up to 5 relatively large eggs and some small ones, The Bass Strait specimen contains about 12 embryos in the right pertbranchial cavity. Larvae have a row of median ampullae alternating with the adhesive organs and 4 lateral ampullae along each side. A ventral pair of rows of large epider- inal vesicles expands into 4 postero-yentral clus- tet on each side beneath the tail. Corresponding TOWS are present on each side of the mid-dersal line. The larval trunk is 0.4 ta 0.5mm long. and the broad tail barely reaches its anterior end, Each of the ampullae has a terminal cap of columnar cells, although these ane more conspicuous on the long, narrow median ampullae than on the rounded lateral ones. REMARKS The type specimen of this species (Herdman 1886) is a circular, dome-shaped cushion, I.6cm high, 3.3cm in maximum dimension. It has a hard sandy external coating, Zooids are crowded in the colony, and systems were not determined. Long, nartow stigmata are in 10 to 12 rows of about 12. Herdman did not observe any branchial papillae. The gut loop is small, with a smooth stomach. and although Herdman did not observe it to be hori- zontal, if has the short oesophagus and short mid-intestinal region that is characteristic of Patyclinum, and tt also has a polyclinid, stalked, sac-like postenor abdomen with the oyary sur- rounded by male follicles. The species apparently is assigned to the correct genus rather than be- longing to Syneicurm as Millar (1982) suggested. MEMOIRS OF THE QUEENSLAND MUSEUM Some of the zooids of the holotype contained larvae. Although the exact number present is not recorded it appears that there were more than one or 2 and that they were in the atrial cavity, as a brood pouch is not mentioned. The crowded zooids, the narrow thorax, the number of rows of stigmata, and the form of the colony of the holotype and the newly assigned and recorded specimens are all identical. The only point of difference is the presence of smal! incon- spicuous branchial papillae in the Victorian specimen. Herdman may have overlooked these as they are difficult to detect. Tropical sandy species such as P. Isutsuit and P. saturnium have broader branchial sacs with more sligmala per row, Polyclinum inerustatum n.sp. js distinguished from the present species by its slightly more stigmata per row (14) and more rows (15), and by its large tongue-like branchial papillae. The sympatric P. marsupiale also has larger branchial papillae, more rows of stigmata and more stigmata per row, and thinner more irregular colonies than the mushroom-like ones of the present species. It also has a brood pouch. Polyclinum glabrum Sluiter, 1895 (Fig. 26) Polvelinun glabewm Stuiter, 1895, p. 168; 1913, p, 71. DISTRIBUTION New Recorps: Western Australia (Montebello Is, QM GH5507), Queensland (Capricam Group, QM GHS038—4! GH5563; Townsville Harbour, QM GH2094; Lizard 1. OM GH319 GH5024 GHS036-7 GH5042), PREVIOUSLY RECORDED: Indonesia (Sluiter 1895. 1913). DESCRIPTION EXTERNAL APPEARANCE; The colonies from shallow reefal habitats are irregular cushions, or robust, spreading sheets, up to 1em thick and 6cm in maximum dimension, with soft, transparent test entirely free of sand, and orange zooids, Collector's notes describe them as ‘soft orange’ and ‘chinese orange’ (Ridgeway 1886). In pre- servative these colonies are transparent and yel- lowish to a translucent, soft grey or beige colour. Common cloacal apertures are relatively crowded, especially around the borders of the colony. Some protrude from the surface on eoni- cal prominences, but others are sessile, irregular and extensive openiigs, Atrial lips of zooids are THE AUSTRALIAN ASCIDIACEA, 3 in the test around the rim of the cloacal openings. tn some preserved zooids, the tips of the atrial lips contain dark pigment. Cloacal systems are crowded, and zooids also are crowded along each side of the canals that radiate from the cloacal apertures. u \ Leis ey rene sv gl \ a \ \ - : \ ah 8 \, \ x 4 ‘ < FIG. 26: Polyclinum glabrum — a, colony (QM GH319); b, zooid relaxed with embryos in atrial cavity, anterior rows of stigmata obscured (QM GH319); ¢, portion of branchial sac showing dorsal Janguets and branchial papillae on the transverse ves- sels (QM GH319), d, larva (QM GH5565). Scales: a, 2cm; b, 0.5mm: ce, 0.2mm: d, 0.4mm. 449 The colony from the Montebello Is is a large (maximum dimension 4cm) flattened slab which, with a light coating of sand on one side, appears to have been lying on its side when collected. Originally it probably stood upright, attached to pebbles by 1ts narrow base. Systems open on both sides of the colony, which has an irregularly subdivided surface, INTERNAL STRUCTURE: The zooids are about 4mm Jong, with a large thorax, a small horizontal gut Joop, and a spherical to tapering (QM GH2094) posterior abdomen. The atrial lip is long and moderately broad, and contains fine longitudinal muscles, It extends along the roof of the cloacal canal toward the cloacal aperture and ends in a single point or a fringe of up to 6 minute papillae. A small median papilla projects from the body wall just behind the atrial aperture. Fine longitudinal muscles are present over the anterior part of the thorax. There are 12 or 13 rows of up to 19 oval stigmata with the maximum number in rows 2 to.6. Rounded branchial papillae are on the transverse vessels, about one and a half per stigma, The gut loop is small! and horizontal. The posterior abdomen is attached to it by a relatively long neck. The posterior abdomen is either sac- like or (sometimes) it tapers to a short vascular stolon that projects from its posterior end behind the heart. Three embryos are in the posterior end of the atrial cayily of specimens from the Capricorn Group collected in December, Lizard I. speci- mens collected in July, and 6 in those from Mon- tebello Is collected in August. Larvae have a long (1.2mm) trunk with median and lateral ampullae anteriorly, and dorsal and ventral clusters of ec- todermal vesicles along each side of the mid-line. REMARKS _ Sluiter (1895) described the type specimen as a large naked cushion, less than !em thick, with a smooth surface, without conspicuous systems, and with conspicuous branchial papillae. Al- though he believed this colony to be juvenile, both male and female gonads are present. The cloacal systems in the present specimens are so crawded that they are hard to distinguish, and in View of the similarities in the branchial sac, and the smooth-surfaced colonies without sand, they seem to be conspecific with the Indonesian spe- cies. The irregular shape and soft consistency of the colony distinguishes it from the more regular and robust P. constellatum. The branchial papillae, the more numerous stigmata per row, and the 450 transparent test of the living specimens distin- guish it from P. vasculosum. The larval trunk is unusually long for this genus. Sluiter’s specimen from Ambon (1895) was grey in preservative with a reddish-violet glim- mer, but this does not occur in other known colonies. Polyclinum incrustatum Michaelsen, 1930 (Fig. 27. Plate 6a,b) Polyclinum neptunium: Michaelsen, 1930, p.542 f. incrustatum. Kott, 1963, p.83 (part, South Austra- lian specimen). DISTRIBUTION New RECORDS: South Australia (Golden I., SAM E2145 QM GH4185; Hopkins I., QM GH4192; Mar- garet Brock Reef, SAM E2531; Perforated I., QM GH4141; The Gap, QM GH4163; Cathedral Rock, QM GH4145 GH5209; Nuyts Arch., SAM E2528). Victo- ria (Portland Harbour, MV 59434 F59437), PREVIOUSLY RECORDED: Western Australia (Bunbury, Albany — Michaelsen 1930). South Australia (Reevesby I. — Kott 1963). DESCRIPTION EXTERNAL APPEARANCE: The colonies are thick sheets, sometimes with swellings, high con- voluted ridges and irregularities on the surface. The common cloacal apertures are sessile or on small raised prominences about Smm apart. The zooids open all around the ridges and irregulari- ties, as well as on the flat part of the colony. Sand is present in a single layer in the surface test. Sometimes it is interrupted by the prominences raising the cloacal apertures above the surface, but in other specimens these are also covered with sand, Embedded sand is only sparse in the internal test between the zooids, but the grains are mod- erately crowded between the zooid systems. Cloacal cavities are small, circular, and are con- fined to the area beneath the cloacal aperture. The zooids are arranged around these cavities. The atrial lip of each zooid in the system extends across the roof of the cavity and is inserted into the test around the rim of the cloacal aperture. The branchial apertures open in a small surface de- pression surrounding each cloacal prominence. INTERNAL STRUCTURE: Zooids are about 8mm long, with the narrow thorax about three-quarters of the total length. A small horizontally oriented gut loop has an almost spherical to long, cylindri- cal posterior abdomen connected to it by a short MEMOIRS OF THE QUEENSLAND MUSEUM narrow neck. There is a narrow vascular stolon from the posterior end of the posterior abdomen. The atrial lip is long and narrow. It has denticu- lations on its free tip. The usual muscular atrial siphon projects forward to open into the base of the cloacal cavity around its outer margin. The anterior part of the zooid, including the branchial lobes and atrial lip, contains black pig- ment. Fine longitudinal muscles (about 8) extend from the branchial siphon to about halfway down the thorax. Fifteen rows of 13 or 14 longish-oval stigmata have transverse vessels with flat tongue- shaped papillae along the margin, about one to every stigma. These are delicate and hard to see without staining. The oesophagus is short, and curved ventrally to the small, smooth-walled, horizontal stomach. The distal end of the gut loop is bent to the left. The narrow neck connecting the posterior abdo- men is attached in the pole of the gut loop in the concavity of this distal bend of the loop. It appears to have been drawn up onto this side of the gut loop, as the vas deferens moves to the outer curve of loop and follows the curve of the rectum where it lies adjacent to the duodenum, stomach and oesophagus. The rectum terminates in a 2-lipped anus halfway up the branchial sac. Up to 20 male follicles surround the ovary in the sexual posterior abdomen. The long (3mm) vegetative posterior abdomen is in colonies col- lected in April from Perforated I. (QM GH4141) and in February from Franklin I. (SAM E2528). Up to 7 embryos are clumped together in the atrial cavity of zooids from Portland Harbour (MV F59434) but the collection date is not re- corded. Larvae have a trunk nearly 0.6mm long, with median and lateral ampullae, and postero- ventral and antero-dorsal vesicles. REMARKS Zooids of this species are relatively large and that, together with their distinct circular systems helps to distinguish them from other species. Michaelsen (1930) believed specimens from Western Australia with embedded sand through- out to be a form of P. neptunium Hartmeyer, 1912 for which the type locality is South Africa. Hart- meyer’s species has a more or less spherical stalked head, and, with the exception of one speci- men (which has sand in the lower half of the colony), is without sand either on the surface or embedded (see also Millar 1962). The abdomen is clavate when mature, and long and cylindrical when immature, the zooids are in circular to dou- ble-row systems, and have conspicuous branchial THE AUSTRALIAN ASCIDIACEA 3 451 COOSLU) SWUNHANd | FIG. 27: Polyclinum incrustatum — a, general view of colony (QM GH4185); b, close-up view of surface of colony with common cloacal apertures surrounded by branchial apertures (MV F59434); c, zooid in colony showing atrial lip in test above cloacal cavity, and branchial aperture amongst sand adhering to outer surface (QM GH4185); d, abdomen and vegetative posterior abdomen (QM GH4141); e, portion of branchial sac showing branchial papillae (QM GH4141). Scales: a, 2cm; b, 2mm; c-e, 0.5mm. 452 papillac. Michacisen (1920) and Kott (1963) as- signed the western and southern Australian mate- rial to this species largely on the basis of the long vegetative abdomen, which now is known to oc- cur in other species of this genus. Michaelsen identified 2 forms, Polyclinum nepiuniuer £ typicum being the form in which sand is absent altogether or present only in the basal layer; and P. neptuniim f, incrustatum with sand throughout. The present sessile specimens from South Australia, with moderate amounts of embedded sand between the systems, with an almost continuous Jayeron the surface, and with strictly circular systems and raised cloacal aper- tures, appear to be conspecific wath P. preptereiuyn: Michaelsen, 1930 f, incrustatum recorded from Bunbury and Albany, but not with the stalked South African species. The stalked specimen from Western Australia (Kott 1963), does have characters in common with the South African species, but is distinguished by the flattened up- per surface. It resembles Polyclinwa circulatan Sluiter, 1909, and may be a synonym. There is insufficient data available for the Shark Bay specimens of P. neptunium f. typicum [see Michaelsen 1930) with double mws systems and sand in the lower half of the colony to assign it to a species. Polyclinum marsuptale Kott, 1963 (Fig. 28a—. Plate 6e,d) Polyclinum marsupiale Kott, 1963, p.83 (part, not specimen from Heron [.? Apliadum spp.); 1976, p.62 (< Polyclinum fungosun), DISTRIBUTION Type Locauity: Western Australia (Fremantle — Michaclsen 1930), FUTHER Recorps: None. DESCRIPTION (after Michaelsen 1930) EXTERNAL APPEARANCE: The colonies are is- regular, flat topped cushions or sheets, with broad ridges and furrows on the surface, up to Sem in maximum dimension and 1.8em high. The test is THE AUSTRALIAN ASCIDIACEA 3 soft. but with a tough, thin outer cuticle, Zooids (from 4 to 11) are arranged in circular to oval systems. The cloacal apertures are sessile, and depressed into a groove. Sparse sand is present on the margin, but the upper surface of the colony is naked. INTERNAL STRUCTURE: The zooids are about icm long, of which about &mm is posterior abdo- men, The thorax 1s longer than the abdomen. The branchial lobes are inconspicuous. The atrial si- phon has the tongue extending from the upperrim of the aperture. Two loose bands of muscle extend along each side of the zooid to its posterior end. The stomach is smooth-walled, A brood pouch is constricted off from the postero-dorsal corner of the thorax. It always contains 2 embryos. REMARKS Despite the difference in colonies and the ab- sence of the faecal pellets that occur tn the type specimen from New Zealand, Michaelsen (1930) assigned the Western Australian specimen to S. hypurgon on the basis of the similarities in the zooids, However there are differences between the New Zealand and Fremantle colonies, and they are not conspecific. A new species is erected to accommodate the Fremantle specimens re- ferred to S. hypurgon f. galei: Michaelsen, 1930. The species is distinguished from others which have the atrial lip associated directly with the rim of the aperture by the smooth stomach wall, ab- sence of a dorsal papilla, and the flat-topped irregular colony with surface ridges and furrows. Kott (1963), interpreting the dark oval bodies embedded in the test as faecal pellets, assigned colonies — one each from Heron I. and Rottnest 1. — to the present species. Re-examination of these specimens (AM Y1382 U3991) has shown them to be Aplidium lobatum. Embryos are incubated in a brood pouch (as they do in the present species) in S. sacculunt n. sp.. which has lobed colonies, a dorsal papilla and a mulberry-like stomach wall distinguishing it. The broad ridges and furrows on the surface are reminiscent of those in Aplidium crateriferum and A. lobatem n. sp. However the latter species has a distinctive zooids which more closely re- semble those of $. papilliferum (see below) than of the présentl species. Synoicum intercedens (Sluiter, 1909) (Fig. 50) Morchellium inlercedens Stuiter, 1909, p08. 39] Synoicum intercedens: Monniot and Monniot, L987, p.al. 2Synoicum suesanum Michaelsen, 1919, 9.27, DISTRIBUTION NEW RECORDS: Queensland (Capricorn Group, OM GH5084-5101 GH510S—7 GH5180 GH5182 GHS193 GH5253 GH5256 GH5282 GH5499 GH5566 GH557 1-2 GH5591-2 GH5608 GH5672)- PREVIOUSLY RECORDED: Indonesia {Sluiter 1909). Moorea, Tahiti (Monniot and Monniot 1987). Although at present there are no records from Australian locations, other than those front the southem end of the Great Barrier Reef, it is prob- able that the species will be found generally around the northern part of the continent and in the western Pacific. DESCRIPTION EXTERNAL APPEARANCE: The colonies form cushions up to 3cm Jong with the upper surface divided into low, rounded swellings to domes from 3 to 10mm in diameter, projecting up to $mm from the solid sandy basal common test and outer margin of the colony. Each of these protu- berances has a Jarge central sessile common cloa- cal aperture. One specimen has a few randomly distnbuted test swellings each containing a single system of juvenile zooids on the surface of a mass of firm gelatinous basal test (QM GH5090) and probably is a regenerating colony, Sometimes only a shallow crease is in the surface test deline- ating the circular area that is homologous with a rounded projection in other specimens. Sand sometimes lies in the crease between the surface swellings, and is present in the basal test, Up to 10 zooids are in each system. The test is particularly opaque and firm in con- sistency in larger colonies, bul is transparent and soft but firm in small ones. The living colonies are a chestnut or claret colour, or ‘claret brown’, of ‘chocolate brown’, or ‘aster purple’ (Ridgeway 1886), with ‘madder brown’ celis throughout the test. Some small colonies. when living, are de- scribed as ‘red chernes projecting from sary base and margin’, Zooids are ‘burnt carmine’ or ‘aster purple’ (Ridgeway 1886), or brownish- black. Basally colonies are yellowish, probably owing to the embedded sand showjng through the brown test. In preservative the colonics are light yellowish, cloudy and translucent and the zooids are reddish-brown, INTERNAL STRUCTURE: Zooids are up to lem jong. Six rounded lobes surround the branchial aperture. The atrial siphon is relatively large, its 402 MEMOIRS OF THE QUEENSLAND MUSEUM upper border is produced into a long lip with 3 small lobes on its tip. Sometimes the atrial lip curves around to form a tube with its Jumen continuous with that of the siphon itself. A small papilla projects from the dorso-median line near the base of the posterior border of the atrial si- phon- Eight fine longitudinal and oblique muscles are on the thorax and these extend along each side of the abdomen and posterior abdomen in a nar- row band. There are 14 to 17 rows of 12 to 14 stigmata. The oesophagus is curved, entering the relatively short dorsal border of the stomach, The stomach wall is covered with rounded promi- nences. (mulberry-like). The duodenal area is wide, the mid-intestine inflated and an oval pos- terior stomach is in the bend of the gut loop. The anal opening is about one-third of the distance up the thorax. Gonads are in the middle third of the posterior abdomen, the small ovary behind the most ante- nor of the testis follicles, which extend back in a single series, or are bunched together when mus- cles contact. Two embryos (but not tailed larvae) are in the posterior part of the atrial cavity of specimens collected in November (QM GH5087). Speci- mens collected in April and May have 3 embryos including a tailed larvae lined up in the atnal cavity (QM GH5088 GH5091). The larval trunk is about 0.50mm long. It has median and lateral ampullae alternating with the adhesive organs. Epidermal vesicles branch off strands that extend along each side of the mid-dorsal line as in Paly- clinum, but ventral vesicles appear to be median. The lateral ampullae sometimes appear to haye lost their connection with the epidermis. REMARKS The species is distinguished from others by its surface swellings each containing a circular sys- tem, and its mulberry-like stomach, The surface swellings are very variable and they are not al- ways conspicuous, Thus the fact that Sluiter (1909) did not observe them may not be signifi- cant. Other characters of the newly recorded ma- terial conform with the type specimen, and the Australian and Indonesian specimens appear ta be conspecific. Syroicum intercedens: Monniot and Monniot, 1987, has fewer rows of stigmata than either the type or the present Australian specimens, and the number of stigmata per row FIG. 50: Synoicum intercedens — a, colony (QM GH5087); b, zooid (QM GH5087); ¢, larva (QM GHS5107). Scales: a, 3mm; b, 1mm; c, 0.1mm. THE AUSTRALIAN ASCIDIACEA 3 has not been reported. Otherwise these French Polynesian specimens appear to conform with the type material. Synoicum suesanum Michaelsen, 19}9 has similar colonies with circular systems, and 13 to 15 rows of stigmata with 10 to 12 per row. Michaelsen (1919) distinguished the species by the depression separating the systems. How- ever that depression sometimes occurs in S. inter- cedens, There is no known character separating S. suesanum from the present one. The new Australian temperate species Synoicum sacculum n. sp. has a mulberry-like stomach, but its systems are not separated fram one another by grooves or depressions, and a single embryo is incubated in a stalked brood pouch. The tropical Synoicum suarenum n. sp. which is sympatric with S, intercedens and is the same dark colour has, like 5. sacculum n.sp., many circular systems in each of its Wedge- shaped lobes, and lacks grooves or subdivisions which, in the present species, tend to isolate the systems from one another, Synoicium suarenum nsp. is further distinguished by its smooth- walled stomach and larger larval trunk. Synoicum macroglossum (Hartmeyer, 1919) and §. tropicum (Sluiter, 1909) are also distin- guished From the present species by their smooth- walled stomachs. Synoicum longistriatum n.sp. (Fig. 51) DISTRIBUTION Tyrek Locatity: Western Australia (off Cockburn Sound, 32°8 155°08'E CSIRO Cruise 144/DM4/63, 110m, coll. L. Thomas 28.8.63, holotype QM GH5583. FURTHER RECORDS: None. DESCRIPTION EXTERNAL APPEARANCE: The colony is an up- night, almost cylindrical lobe, slightly expanded terminally, It is 7cm high and about 3cm in di- ameter. Basally there are sandy root-like projec- tions of the test. The lower third of the lobe, comprising the lower part of the stalk, is Irans- versely wrinkled, and covered with a layer of sand, The middle third has a layer of finer sand and is not wrinkled. The naked upper third has openings of the crowded zooids. These, seen through the transparent test, appear to be arranged in crowded longitudinal rows converging to cloa- cal apertures on the terminal free end, The crowded zooids are perpendicular at the surface, but posteriorly they curve down into the stalk of the colony, $93 INTERNAL STRUCTURE: In this colony a large quantity of sand fills the pharynx of most of the zooids. obscuring its structure, The branchial ap- ertures haye 6 lobes, and atrial apertures are large openings with the upper rim extended out to form a tongue. About 12 fine longitudinal muscles are in the thoracic body wall, These extend along the length of the abdomen and posterior abdomen. Stigmata are in 16.rows of about 16. The stomach is large and voluminous, without longitudinal folds, but sometimes collapsing into irregular or horizontal ones. Double rows of male follicles are in part of the long posterior abdomen of some of the zooids, although the ovary was not detected. FIG, 51: Synoicum longistriatum n.sp. (holotype QM GH5583) — colony. Scale: lem. 494 REMARKS The colony is distinctive and unique in this genus, superficially resembling that of Distaplia australiensis (see Kott 1990). However, the zo- oids are characteristic of the Polyclinidae. The Aplidium species with longitudinal double-row systems converging to the top of the head, viz. A. australiense, A. geminatum n.sp., A, inflorescens n.sp. and A. paralineatum n.sp. are distinguished by their numerous stomach folds. The shape of the colony and its sandy surface resembles S. atopogaster, but the form of the systems differ. It is unfortunate that this unique specimen is not well preserved, Nevertheless as it is a unique colony form and unlikely to be confused with other species in this genus, it is considered appro- priate for it to be designated holotype. It is prob- able that damage was caused to the specimen by the way it was collected (beam trawl), rather than its treatment after collection. MEMOIRS OF THE QUEENSLAND MUSEUM Synoicum macroglossum (Hartmeyer, 1919) (Fig. 52. Plate 9a) Macroclinum macroglossum Hartmeyer, 1919, p.126, DISTRIBUTION New Recorbs: Queensland (Heron 1., QM GHS5055; Mackay, QM GH5421). PREVIOUSLY RECORDED: Western Australia (Cape Jaubert — Hartmeyer 1919). DESCRIPTION EXTERNAL APPEARANCE: One colony (QM GHS5055) is a large flat-topped cushion 6cm in diameter, 2cm high, with rounded margins and a vertical border. Sand is present on the base and around the border, but not on the upper surface. Sparse sand is present internally in the lower half of the colony, but it becomes even more. sparse toward the centre, as if it was gradually moving in from the sandy outer border. The colony from FIG, 52: Synoicum macroglossum — a, colony (QM GH5421); b, whole zooid severed between thorax and posterior abdomen (QM GHS5055); ¢, d, thorax and atrial aperture, with atrial lip shown in different positions (QM GH5421 GH5055); e, portion of right branchial wall (QM GH5055). Scales: a, 2cm; b, c, 1mm; d, e, 0.5mm. THE AUSTRALIAN ASCIDIACEA 3 Mackay is an upright cone, 6cm high, with sand around the base and sparsely embedded through the test. The test is translucent ard gelatinous but firm and rigid in both specimens. Circular to long, Jow swellings, with shallow depressions between them are on the upper surface, The small and inconspicuous branchial openings can be seen around the peniphery of the swellings, and in the depressions between them, Large cloacal aper- tures occur in the depressions, especially where 2 of more converge. These apertures either are ses- sile, or they protrude on surface swellings created by thickened test over the circular cloacal cavities that are formed at the junctions of the cloacal cinals. Internally the posterior ends of the zooids criss- cross one another. The test is pale yellowish- brown to greenish-yellow in preservative, and the preservative itself is stained yellow-orange. One living specimen (QM GH5055) was, ‘mummy brown’ internally and ‘seal brown’ (Ridgeway 1886) on the surface. The other (from Mackay) is said to have been black when living. INTERNAL STRUCTURE: Zooids are long (to 1.5cm) and narrow. The thorax is about one-third of the length of the zooid, and the abdomen and about half of the gonad bearing part of the poste- rior abdomen together comprise another third. Eight longitudinal muscles extend the length of the thorax, and are reduced to 2 or 3 each side of the abdomen and posterior abdomen. ‘The bran- chial aperture is on a trumpet-shaped siphon that appears to be fixed in the finm surface layer of test. A sphincter is around the base of the siphon. The jong, muscular atrial lip has a dentate tip, and is difficult to remove from the test where it lies over the top of the cloacal cavity, It anses from the body wall opposite the first row of stigmata, antenor to the muscular atrial aperture which lies opposite the third row of stigmata. A sphincter is also present around the atna) aperture. When contracted, the atrial lip curves over the aperiure. A papilla projects from the body wall just behind the atrial siphon opposite the fifth row of stig- mata. Nineteen rows of 12 to 14 fairly long stig- mata are in the branchial sac, although the number reduces posteriorly. A wide strip of unperforated lest is each side of the endostyle, There is a minute papilla on each transverse vessel at about the level of the third ventral stigma on each side of the body. A row of robust dorsal languets are to the left of the dorsal mid-line, The gut loop is vertical and narrow — the oesophagus curves ventrally to enter the bilater- 495 ally symmetrical, smooth-walled stomach. The duodenum is large, the mid-intestine inflated and a Jarge oval posterior stomach is in the bend of the gut loop, The bilabiate anus opens at the level of the fourth row of stigmata from the posterior end of the thorax. A very short gonad-free part of the postenor abdomen is behind the gut loop. A small avary is antenior to the oval testts follicles which are in 2 longitudinal series, altemmating with one another, Sometimes (but not always) an extent of the pos- terior end of the posterior abdomen is free of gonads. A V-shaped heart is in the posterior ex- tremity of the posterior abdomen. The newly re- corded specimens collected in November, have well-developed ovanes, but no embryos were found. REMARKS The species is distinguished by its protruding atrial siphon, separate atrial lip, narraw zooid and robust cushroan-shaped colony. The type speci- men differs from the present one in the presence of a layer of sand on the upper surfaces as well as around the margin; and in the presence of up to 14 rows of stigmata rather than the 18 of the present easter Australian colonies. Hartmeyer (1919) did not observe the small branchial papilla foward the ventral end of each transverse vessel, but these are inconspicuous and obscured unless the thorax is completely relaxed, The dorsal papilla and separate atrial lip are like S. papilliferum (Michaelsen, 1930) and the zo- oids also have similar numbers of stigmata, How-~ ever, although Michaelsen believed the dorsal papilla was unique to his species (see below), it occurs in both Polyclinum and Syicicum, Al- though the conspicuous cloacal apertures and long curving double row systems of the present tropical species may be different from S. papillif- erum (in which Michaelsen did not find any cloa- cal apertures and could not determine the formn of the systems}, at present the only character defi- nitely known to separate the species is the pres- ence of sand adhering to the extemal test of the temperate S. papilliferum, Species of Aplidium with similar zooids (e.g. A. lobatwn, A. cratif- ertum) lack a median dorsal papilla, and have different systems from those of the present spe- cies. Synoicum chrysanthemum has similar sys- tems, but the colonies aré different. 496 MEMOIRS OF THE QUEENSLAND MUSEUM Synoicum obscurum n.sp. (Fig. 53) DISTRIBUTION Tyre LocaLity: Victoria (Bass Strait, reef off Ninety- mile Beach, 90km SW of Lakes Entrance, coll. J. Watson R3E/36 1977, holotype QM G12726, paratype QM G12727). FURTHER RECORD: NSW (Arrawarra, QM GH5589), DESCRIPTION EXTERNAL. APPEARANCE: The colonies are thick mats with sand crowded throughout making them hard and brittle. The mats are made up of sandy cone-shaped, upright lobes which adhere closely to one another. The terminal free ends of these lobes project from the upper surface of the colony as rounded swellings, each of which ac- commodates a circular system of up to 12 zooids. Zooids are tightly enclosed in the sand and hard to remove from it although the thin channels that accommodate the zooids are rigid. The zooids surround a shallow common cloacal cavity (in the terminal end of each lobe) which opens through a common cloacal aperture in the centre. Bran- chial apertures are inside, and protected by, the rounded margin of the terminal end of each lobe, The cloacal cavities beneath the upper surface of colony confer a loose appearance to the surface layer of brittle sand-filled test. INTERNAL STRUCTURE: The zooids are rela- tively muscular, with about 20 longitudinal bands on the thorax. These muscles extend posteriorly to form a band along each side of the mid-ventral line. Zooids are relatively long, even when con- tracted the thorax, abdomen and gonad-bearing anterior part of the posterior abdomen are about 6mm long, while the long posterior part of the posterior abdomen, without gonads in it, extends into the sandy base of the colony, The 6 branchial lobes are rounded. An atrial siphon protrudes from the dorsal surface, and a flat atrial tongue with a tridentate tip is produced out from the upper rim of the opening. The lower rim of the atrial opening is divided into 3 small lobes, A pointed median dorsal papilla is at the base of the atrial siphon. About 20 rows of up to 20 stigmata appear to be present, although the pharynx is too contracted for accurate counts. Irregular internal glandular areolations of the stomach wall are present, especially on the mesial side of the stomach (against the intestine), A few FIG. 53: Synoicum obscurum n.sp. (holotype QM G12726) —a, colony; b, zooid;¢, larva, tail removed, Scales: a, 5mm; b, 1mm; ¢, 0.1mm. THE AUSTRALIAN ASCIDIACEA 3 of these shallow glandular areas sometimes form irregular longitudinal ridges which resemble the longitudinal folds of Aplidiwn. The duodenal, mud-intestinal and posterior stomach regions are characteristic for the genus. In the type material, a large ovum is present behind the gut loop, anterior ta a short double longitudinal series (about 8 per series) of testis follicles. The posterior abdomen continues poste- rior to the testis follicles, which protrude dorsally from it as 4 result of the contraction of the ventral muscles. Four or 5 embryos are crowded in the alnal cavity. The larval trunk is 0.6mm long. The anterior end is obscured by numerous narrow ectodermal ampullae. There are 3 median adhesive organs, and an ocellus and otolith are in the cerebral vesicle. REMARKS A similar colony form is found in many species of Aplidium (e.g. A. bacculum n.sp., A. filiforme nsp.). However, despite the presence of some longitudinal glandular ridges in the stomach wall, the present species is distinguished from the ge- nus Aplidium by the shape of its zooids with a relatively primitive lobed atrial opening and the median dorsal papilla. Syroictn durvin has colonies consisting of sandy lobes, but the test above each system is naked, and each lobe has 2 or 3 systems. Synoicum papilliferum (Michaclsen, 1930) Muacroclinum papilliferum Michaelsen, 1930, p.530. Synoiewm papilliferum: Millar, 1966, p.360, Not Kott, 1963, p.87 (< Polyelinum tenuatumn n. sp.). 1972, p.l6; 1972b, p.l77 (< &. saveulin asp.) DISTRIBUTION New REcorDS: None. PREVIOUSLY RECORDED: Western Australia (Bunbury — Michaelsen 1930). Victoria (Nepean Peninsula — Millar 1966). DESCRIPTION (after Michaelsen 1930) EXTERNAL APPEARANCE: Colonies ate cushion shaped or irregular sheets, up to 3em diameter and 2.4cm high. Sometimes they are fixed by a nar- rowed base. The preserved colonies are transpar- ent and yellowish, with an external layer of sand. Some sand is present intemally but it is not as crowded as itis externally. The outer layer of test is tough, Cloacal cavities are present at the level 497 of the atrial apertures, but cloacal apertures were nol seen, INTERNAL STRUCTURE: The zooids are verti- cally arranged in the test, with their posterior abdomina criss-crossing, The body wall is deli- cate with about [{) longitudinal muscle bands. The branchial siphon is conspicuous with a well developed sphincter at tts base, The short atrial siphon, surrounded by a sphincter muscle, pro- jects laterally fram the body wall a short distance behind the atrial lip, which is narrow, but not Jong, and its tip is produced into 2 or 3 small papillae. A conspicuous median papilla projects from the body wall just behind the atrial siphon. Sixteen of 17 rows of 9 or 10 stigmata are in the long, narrow thorax. The oesophagus curves to enter the smooth stomach which js about halfway down the abdomen, A short gonad-free area is behind the abdomen, in front of the single row of male follicles that occupies the rest of the narrow posterior abdomen. When the posterior abdomen contracts, the male follicles are drawn up into a double row, Two short vascular stolons project from the end of the zoord. REMARKS Although Michaelsen thought the papilla was unique, it does occur in species of Polyclimonr as well as in other Synoicum species, Nevertheless Kott (1963; 1972a.b) followed Michaelsen and assigned specimens of S. sacculum n. sp. and Polyclinue tenuatum to this species because, like §. intercedens and S. sucrenum n.sp.. they had a dorsal papilla, However, all these species are readily distinguished from the present one by having the atrial lip extending from the anterior tim of the aperture. Synoicum macroglossum has both a separate atnal lip and a dorsal papilla, but is distinguished from the present species by its conspicuous cloacal systems with raised cloacal apertures and the absence of an extemal layer of sand. Although they differ in having a dorsal papilla, zooids of the present species. with their long, narrow thorax, long muscular branchial siphon, and atrial siphon separate from the atrial lip, resemble those of certain Aplidium spp. (e.g. A. cratiferum, A. lobatum, see below), The stomach folds of some of these Aplidium spp. are few and indistinct, but they all have distinct cloacal sys- tems that distinguish them from the present spe- cies. Millar (L966) records that his specimens of S. papilliferum had all the features of the type speci- mens, except the dorsal papilla. However, as there 498 is no other known. temperate species of Synoicum with similar characters, it is possible that the papilla was obscured in the material he examined, Synoicum prunum (Herdman, 1899) (Fig. 54. Plate 9b) Psammaplidium prunum Herdman, 1899, p.84. Synoicum prunum: Kott, 1963, p87, DISTRIBUTION New ReEcorps; New South Wales (Arrawarra, QM GH5394). Queensland (Bribie 1., QM G2102; off Cape Tribulation, QM GH799). PREVIOUSLY RECORDED: New South Wales (Port Jackson — AM G2101 Herdman 1899), DESCRIPTION EXTERNAL APPEARANCE: Like the holotype, the colony from Cape Tribulation is upright, oval, converging to a narrow sandy base produced into sandy roots. It is about 7em high and 4cm in diameter, The specimen from Bribie I, is massive (14cm long and 9cm maximum diameter), lumpy, oval and football-shaped overall, but with deep creases dividing it into lobes, It has no obvious point of fixation. The internal test always is firm and gelatinous with long tough fibres in it. [tis almost opaque in preservative, while the outside layer is tough and hard with occasional patches of sand embedded. The long thread-like zooids are found withdrawn from the surface and criss-cross one another through the internal test. Their arrangement is obscure and the form of the systems was not determined. INTERNAL STRUCTURE: Both newly recorded colonies are in a vegetative stage, many develop- ing buds being present. Adult zooids are thin and thread-like. The very long, thin posterior abdo- men is up to 2cm, being many times the length of the rest of the zooid (only about 2mm long when contracted), The contracted thoraces (QM GH799) are slightly longer than the abdomen, and when extended (QM G2102) they are at least twice the length of the abdomen. About 12 fine longitudinal muscles are on the thorax and these extend the length of the zooid in a band on each side. A single fleshy atrial lip continuous with the anterior rim of the atnal aperture has a straight tip with a single, small, pointed, median tooth, The branchial Jobes are only shallow, A small dorsal papilla arising from the body wall behind the atrial aperture can be seen in the contracted zo- oids, but was not detected in the extended ones. MEMOIRS OF THE QUEENSLAND MUSEUM Fourteen dorsal Janguets were counted in the specimen from Cape Tribulation, indicating that there are 15 rows of stigmata, the same number as in the Bribie I. zooids. The thoraces. are char- acteristically narrow and up to 8 siigmata per row were counted. The abdomen is small, The narrow and characteristically long oesophagus curves to enter the smooth-walled, smal] stomach which is halfway down the abdomen. The relatively long duodenum, inflated mid-intestine and posterior FIG, 54; Synoicum prunum — a, colony outline (QM GH799); b, thorax and abdomen (QM GH799); ec, larva (holotype AM G2J01), Seales: a, 2cm: b. 0.2mm; ¢, 0. [mm. THE AUSTRALIAN ASCIDIACEA 3 stomach are distal to the stomach in the descend- ing limb of the gut loop. Gonads are not devel- oped in the newly recorded specimens. They are present in the long posterior abdomen of the type specimen, which also has 2 embryos in the atrial cavity. The larval trunk is 0,55 to 0.65min long, It has median and lateral ectodermal ampullae altemat- ing with the 3 median adhesive organs, and patches of ectodermal vesicles antero-dorsally and postero-yentrally. REMARKS The large, almost naked colonies of this species are unique in the Australian fauna. are some dark, greenish-black colonies of Poly- clinum Vasculosum recorded from Moreton Bay that reach the same size as the present species and superficially can be confused with it, Synoicum citrum has naked but sessile colonies, smaller than those of S. prunum, with larger zooids, fewer rows of stigmata and more stigmata per row, Synoicum sacculum n. sp. (Fig. 55. Plate 9o-c) Swieicum papilliferum: Kot 1972a, p16, 1972b, p.177; 1976, p.62. DISTRIBUTION Type Locaity: South Australia (Waldegrave !. in caves, 2m strong waves, coll. 8. Shepherd 25.11.73, holotype SAM E2556, paratypes QM GH5044), Vic- tora (Flinders, jetty piles, Sm, coll. N. Coleman 18.6.77 AMPI 201, paratype QM G10163). FURTHER RECORDS: South Australia (St. Vincent Gulf, West I, — Kott 1972a; Thorny Passage, SAM E2491), Tasmania (W. Granville Harbour, SAM E2552). The Tasmanian specimen was taken from a depth of 425m. DESCRIPTION EXTERNAL APPEARANCE: The colony consists of crowded sessile, flat-topped to rounded lobes about lem in diameter rising from a sandy basal membrane or test mass. Sometimes the lower half of the lobe is constricted to form a thick stalk with some sand embedded. Sand is sparse internally in the upper part of each lobe. Sand is sometimes present on the upper surface, although the test above the zooid systems is usually left naked. The test is very sofi internally. Zooids form circular systems of 2 19 3mm diameter with up to 10 zooids around a central cloacal aperture. The cloacal apertures are abowl 2em apart, and are 494 evenly spaced on the top of cach lobe, In one small (1.7cm high) flat-topped colony (SAM E2491) the branchial openings are around the caiter margins of surface depressions, and sessile cloacal apertures are in the centre. The test is raised in firm rounded ridges between the depres- sions. In life the lobes are bright red, with bright red zooids. In preservative the test is yellowish and translucemt, and the preservative is stained a yel- lowish-red. INTERNAL STRUCTURE: The zooids are 2 to 3mm long, with a short posterior abdomen, The thorax, abdomen and posterior abdomen are cach about one third of the total length. There is a fine, short posterior vascular stolon. About 6 fine lon- gitudinal and oblique muscles are on the thorax and these continue along the length of the zeoid, Strong muscles extend fram the branchial siphon along the atrial siphon (which often is long) onta the atrial lip which continues out from the anterior rim of the opening. The straight tip of the atrial hip has 3 narrow finger-like papillae evenly spaced along it. A papilla is in the mid-dorsal line behind the atrial aperture, sometimes carried out along the siphon when that is produced dorsally. About 8 short stigmata are in 9 or 10 rows. The oesophagus fs long and curved, and the stomach, about halfway down the abdomen has mulberry- like swellings in its wall. The duodenal area is long, the mid-intestine inflated, and an oval stom- ach is in the bend of the gut loop, The arnal opening is opposite the third last row of stigmata. Colonies with large mature male follicles in the short posienor abdomen have about 20 bunched around the ovary (SAM E2491, QM GH5044), These colonies also have one lo 3 embryos, up to the tailed larval stage, present in a brood pouch constricted off from the postcro-dorsal corner of the thorax, The larval trunk is almost spherical, 0.4mm in diameter, with otolith and ocellus, postero-ven- tral vesicles, and media and lateral ampullae anteriorly. The tail is wound about three quarters of the way around it. REMARKS The form and colour of the colonies is distinc- tive, as are the alrial lip from the anterior rim of the opening with finger-like papillae on its tip, the mulberry-like stomach, and the short posterior abdomen with bunched male follicles. The spherical larval trunk also is unusual, The brood pouch constricted off from the bedy wall is remi- niscent! of the one described for Macroclinum 500 MEMOIRS OF THE QUEENSLAND MUSEUM THE AUSTRALIAN ASCIDIACEA 3 iryjrergon f. galei: Michaelsen, 1930 although the Jalter contains 2 embryos, its stomach is smooth, and its colonies are not lobed. A dorsal papilla similar to the one in the present species, a short posterior abdomen and a similar alrial aperture occurs in the tropical species 5, suarenum n. sp. However in the latter species the stomach is smooth, the larvae are not spherical and a brood pouch is not constricted off from the body wall, Synoicum erectum, a species that overlaps the South Australian range of the present one, has sandy colonies consisting of single lobes rather than the many lobes of the present species. The zooids can be distinguished by their longer pos- icrior abdomen, longer larval trunk and absence of a brood pouch. Synoicum suarenum i. sp. (Fig, 56, Plate 9f,2) Synoteum kuranuk Kott, 1981, piso. Syanicum ?clavatum: Millar, 1975, p.255. DISTRIBUTION Type Locauiry: Queensland (Capricorn Group, Wil- son Reef, rabble zone, low tide, coll, P. Kot Jan 1977, holotype QM GH5080; Heron 1, rubble zone, low tide, coll. P. Kott 4.11.86, paratype QM GH5070). FURTHER RECORDS: Queensland (Capricorn Group, QM GH5066-9 GH5071-9 GH5081-3 GH5176 GH5178; Hardy Reef, QM GH5377; Lizard 1, OM (H5277 GH5586); Fiji (Kott 1981). Indonesia (Millar 1975). DESCRIPTION EXTERNAL APPEARANCE: Flat-topped wedge- to top-shaped lobes, crowded together and nar- rowing to the base where they are joined by common basal test or by connecting strands Lo adjacent lobes. Sometimes the basa] stalk portion of the lobe is branched and there are 2 heads from a basal common stalk. Colonies from Hardy Reef have long stalks (longer than those from Heron 1.); narrowing toward the base. When alive and inflated the separate lobes that make up the col- ony appear confluent. The separate lobes are more apparent when il is removed from the sub- strate, The flat-topped lobes are up to 2.5em high and up to 3.0cm in diameter at the top. In life colonies have a characteristic appearance resem- bling a pig’s kidney beth in colour and form. They sot are always opaque, shiny cherry coloured, crim- son, and a dark maroon, pliim colour, or ‘pansy' ‘pomegranate-", 'Indiun-' to ‘aster-purple’ (Ridgeway 1886). In preservative colonies be- come brownish-black, but the colour eventually is lost altogether. At first the preservative is stained orange-yellow, and clouds of hrownish- black pigment cells are in the test. Circular clow- cal systems of 4 to 8 zooids are evenly spaced over the surface, The central common cloacal apertures are in slight surface depressions about 2.5mm apart, and each system is about 2.5mm in diameter. The rim of each common cloacal aper- ture has small denticulations, into which the den- ticulations on the tip of the atrial lips of each zooid fit. The cloacal chambers are shallow and circu- lar. being almost completely open to the exterior through the large cloacal aperture. The testis firm and gelatinous, Sand is absent from the surface, and from the intemal test. Up to 15 systems eccur in one lobe of the colony. INTERNAL STRUCTURE: The zooids are less than Icm in length, even when distended, and the thorax is half the total length of the zooid. The posterior abdomen is generally relatively shart for a species of this genus. One or 2 short yasctlar appendages are al the posterior end of the poste- rior abdomen. The 6 branchial iobes are small and pointed. The ginal siphon is short but wide, with its upper border produced into a short, broad tongue with 3 to § shart pointed teeth on its broad, straight outer edge. There is a median pointed papilla projecting out from the dorsal line behind the atrial aperture. About i2 fine longitudinal and oblique thoracic muscles in bands of moderate width extend along each side of the thorax and abdomen and are inserted into a projecting horn on each side of the posterior abdomen. Sixtecn tows of 10 short, oval stigmata are in the bran- cual sac, each row reducing in length dorsally and ventrally. The oesophagus is curved and the dorsal border of the stomach between the ceso- phageal and duodenal openings Is relatively short, The stomach wall is smooth externally, but internally it has some glandular areolations. The duodenal area is long, the tnid-intestine inflated and an oval posterior stormach is in the bend of the gut loop, The bilabiate anus is about one-third of the distance up the thorax, The posterior abdomen is continuous with the abdomen and the gonads are in its posterior half, there being a narrow gonad-free neck of variable length behind the gut FIG. 55: Synoicum sacculam n.sp.— a, b, colonies (paratype QM G10163, a, showing circular systems; bh, colony outline): c, zo0id with embryo in brood pouch (SAM E2491); d, thorax (paratype QM G1G163); e, larva (paratype SAM 65044). Scales: a, 2mm, b, lem; ¢, d, 0.5mm, ¢, 0.1mm, 502 MEMOIRS OF THE QUEENSLAND MUSEUM loop. The ovary is mixed in with the anterior testis Embryos (up to 3) are in the posterior end of the follicles, which are either in one or 2 irregular atrial cavity of zooids collected in January, April longitudinal series or, When muscles are con- and May, Tailed larvae have a trunk about 0.6mm tracted, bunched in the posterior end of the pos- long, Median and lateral ampullae are present. A terior abdomen, giving it a sac-like appearance. _postero-ventral cluster of epidermal vesicles is in There are about 20 testis follicles, the larval test each side of the mid-line, near the FIG. 56: Synoicum suarenum nsp. — a, colony with longer lobes (QM GH5377); b, colony with short lobes (holotype QM GHS080); c, colony from above showing systems (QM GH5067),; d, zooid (QM GH5067); e, posterior abdomen showing gonads bunched in posterior end (paratype QM GH5070); f, larva (QM GH5073). Scales: a, lcm; b, 2cm; ¢, 5mm; d, e, 0.5mm; f, 0.2mm. THE AUSTRALIAN ASCIDIACEA 3 base of the tail. These separate from an epidermal strand that trails pasteriorly through the test, from each side of the mid-line ventral to the adhesive o¥gans, A similar strand of epidermis from which epidermal vesicles separate off into the dorsal test above the endostyle is on cach side of the mid- dorsal line behind the adhesive organs. The tail winds about two-thirds of the distance around the trunk. The lateral ampullae separate from the rest of the larval ectoderm and become large and spherical. REMARKS In those colonies with male follicles clumped in the posterior end of the zooid, the anterior part of the posterior abdomen is relatively narrow, resembling the neck of a palyclinid posterior abdomen. However, the species has characters of Synoicum rather than Polyclinum, viz. the mid-in- testine is large and inflated, the gut loop straight and relatively long, and the cloacal systems small. Synoicum clavatum (Oka, 1927) from Japan resembles the present species in some characters, but it differs in having larger round-topped rather than flat-topped lobes with longer, narrow stalks. There is some confusion regarding the characters of the Japanese species. however, for Nishikawa (1990) records 20 to 24 Jongitudinal muscles and 10 to 12 stigmata, but Tokioka (19546) reported. only 12 muscles and 20 stigmata (see Tokioka 1954b, Pl. IVF). The colours of living colonies of the Japanese populations, which could provide more evidence of their relationships with one another and with the present species, have not been recorded. Synoicum clavatum: Millar, 1975 from the Philippines has flat-topped lobes containing cir- cular cloacal systems similar to those of the newly recorded Australian material; and the zonids have & to 9 longitudinal muscles similar to those from Heron I. However, although they are in 15 rows like the Australian material, Millar reports about 20 stigmata per row in his specimens, The flat- topped Philippine colonies, therefore, are only questionably conspecific with the Australian colonies, The dorsal papilla has not been recarded for Philippine or Japanese specimens. Syivicumn kuranui Brewin, 1950b from Great Barrier J. (New Zealand) are sandy capitate lobes with crimson zooids and areolated stomach walls and are readily distinguished from the present spe- cies. Synoicuim kuranui: Kott, 1963 from Heron L is nol conspecific with either the present one or with the New Zealand species (see 5S. durum, above). The sympatric §. imrercedens has sessile cushion-like colonies with sandy margins. Their upper surface with swellings corresponding to lhe systems, larger sysiems, longer atrial tongues, a mulberry-like stomach and a smaller larva. Syneicum suarenunt 1s a tropical species and it differs from the temperate §. sacculum by the shape of the colony lobes (sessile in S. sacculun), the smooth stomach wall (mulberry-like in 5, sacculum), and the small, almost rudimentary cloacal systems with the wide cloacal aperture exposing the cloacal cavity to the exterior, as in some Fudistoma spp. {angolanum group: Kort 1990), Synoicum tropicum (Sluiter, 1909) (Fig. 57) Atopogaster tropicum Sluitec, 1909, p. 107, DisSTRIBUTION New RECORD: Westetmm Australia (NW Shelf, QM GH5056). PREVIOUSLY RECORDED: Indonesia (Sluiter 190%), The species is known only from the type and from the newly recorded portion of a colony, DESCRIPTION EXTERNAL APPEARANCE: The specimen is a wedge-shaped slice of what appears to have heen a large, upnght lobe, about 12cm high. The testis firm and the surface naked. Zooids ane arranged in circular systems of $ to 20. The branchial apertures are around the margin of a circular depression about Smim diameter in the surface fest. A conspicuous common cloacal aperture protrudes on a conical prominence in the centre of each circle of zovids, The test is white and opaque, with a patch of dark pigment where each zooid opens to the surface. Common cloacal apertures are about 7mm apart, an area of test without zooid openings separating the circles of branchial apertures from one another. INTERNAL STRUCTURE: The zooids tn the pre- sent piece of a colony have not been adequately fixed and cannot be dissected or manipulated without disintegrating. They are, as in the type specimen, perpendicular to the surface, The zo- oids of the type colony {Siuiter 1909) have 12 rows of 8 to 10 stigmata. The gut forms a simple loop, the stomach wall is smooth, and there is a long posterior abdomen. REMARKS The present specimen agrees with the type in having circular depressions on the surface with branchial apertures around the margin of each depression, protruding cloacal apertures, and no sand either embedded or adhering to the surface. The circular surface depression is characteristic. Although circular systems with a large circular cloacal cavities surrounded by zooids occur in other species of Synoicumt, the well-spaced and depressed circles of the present species are un- usual, The horizontal stomach folds that characterised the genus Atopogaster (to which Sluiter assigned this species), are artefacts that occur in species with large, smooth stomachs and Afepogaster is here proposed as a junior synonym of Synoicum. Genus Morchellium Giard, 1872 Type species: Amarauctum argus Milne Ed- wards, 1842. The known species have a relatively long oe- sophagus. A vertical, barrel-shaped stomach half- way down the abdomen is smooth or has mulberry-like swellings in the stomach wall, Go- nads are in the posterior abdomen with the ovary anterior to the serially arranged testis follicles. MEMOIRS OF THE QUEENSLAND MUSEUM Rows of stigmata are numerous. The atrial aper- ture is wide and open, often forming a protuberant siphon with an enlarged uppper rim. Neither the atrial ip nor the sphincter muscle are as well developed as they are in Aplidium and Poly- clinum. The posterior abdomen is narrow and continuous with the abdomen, there being no constriction between them, The number of rows of stigmata are in the range known for other genera of the Polyclinidae. How- ever the number per row (20 to 30) generally exceeds that of all other genera of this family, being almost as numerous as in Polycitoridae and Sigillina spp. The number of stigmata per row in Protopolyclinidae, and in some iiterella spp., are of this order. In Euherdmaniidae and Placen- telidae they are more, and in Pseudodistomidae less numerous. Although they are not present in the type spe- cies, the Australian species of this genus have parastigmatic vessels in the branchial sac. A few species of Aplidium —e.g. A, caeruleum (Sluiter, 1906) — and Synoicum atepogaster also have parastigmatic vessels (see Kott 1969) and so do certain Protopolyclinidae (Monniotus spp., Con- dominium) and certain Ritterella spp. (see above) FIG. 57: Synoicum tropicum (QM GHS5056) — a, part of colony; b, circular systems with protuberant branchial apertures surrounding a larger, central common cloacal aperture, cach depressed into the surface. Scales: a, 2cm; b, 2mm. THE AUSTRALIAN ASCIDIACEA 3 as well as holozoid genera Distaplia, Sycozea and Hypsistozoa, Thus na profound phylogenetic sig- nificance can be attached to the presence of these vessels in Morchellium species, Parastigmatic vessels hold in place the long, parallel interstig- matic bars that occur in most species with particu- larly long, narrow stigmata. These generally occur when there are few rows of stigmata rela- tive to the size of the branchial sac. Probably the presence of parastigmatic vessels, preventing dis- FIG. 58: Morchellium albidum n.sp. (holotype QM GH5582) — a, part of upper surface showing zooid systems: b, zooid, with detached postenor abdomen showing ventral muscle band, Scales; a, 4mm; b, 0.5mm, S05 tortion of the stigmata, maintaining the dimen- sions of their apertures and the effective operation of the cilia lining them, is a convergent character in the taxa referred to above. The transverse thoracic muscles of the Austra- lian species M. pannosum n.sp. are unique in the Polyclinidac, although they are known in Sigillina (Holozoidae), Polycitoridae (see Kott 1990) and some Ritterellidae (sce above), The pigment spots in each of the branchial lobes (as in the Diazonidae and Ewherdmania spp.) ate also unsual. Neither of these characters occurs in M. albidum n.sp. Thus, they do not seem to be ge- nenc characters as the genus js currently defined. Nor do they occur in Synoicwm. Morchellium argus (type species) has 4 sym- metrically arranged unusual pigment spots around the apertures (Berrill 1950), but these are not homologous with the diazonid-like pigment spots of M, pannosum n,sp, The large atrial aperture, and barrel-shaped stomach without longitudinal folds found in both Australian species are similar in certain Synoicum spp. from which Morchellium is distinguished only by its 8 branchial lobes. The type species is from the English Channel and the west coast of Britain, Ireland and France. The species discussed below are the only 2 of this genus described from Australia. They differ trom one another principally in their colonies, M. pannosum nu.sp. having sandy lobes, each with a single system, and M, albidum n.sp. having nu- merous circular systems in each cushion-like col- ony. Morchellium partitionis (Monniot, 1987) from New Caledonia, the only species of thus genus previously recorded from the tropical western Pacific. is divided into sandy claviform lobes and is 2cm high, The external test is thickly encrusted with sand. Its colony form is similar to that of M, pannosum n.sp., and it has a similar number of raws of stigmata (13 to 14) and a similar number of stigmata per row (18). However its lack of parastigmatic vessels and its mulberry-like stom- ach distinguish it. Morchellium albidum n.sp. (Fig. 58) DISTRIBUTION Tyre Locauiry: South Australia (West Bay, Wedge I,, southern Spencer Gulf, Earthwatch site 2, coll. A. Butler 16.2.88 QM GH5582). FURTHER RECORDS: None. 506 DESCRIPTION EXTERNAL APPEARANCE: The colony is a large, oval bolster 4em long and 3cm wide. Zooids are arranged in crowded, circular systems of up to 12, with large sessile common cloacal apertures in the centre of each. The test is firm and transparent and the zooids, seen clearly from the surface, are translucent-white in preservative. An opaque white deposit is in the surface test between the systems, INTERNAL. STRUCTURE: Zooids are large and robust. However. the thorax and abdomen to- gether are as little as [.4mm long when con- tracted, The posterior abdomen is longer, The rim of the branchial aperture has 8 conspicuous rounded lobes, The atrial aperture protrudes from the dorsal surface, and its anterior rim is some- times produced into a strap-like tongue, with 3 pointed terminal lobes. In other specimens the atrial lip is shorter and broader, the 3 terminal lobes being more conspicuous than the tongue itself. A median dorsal papilla is present behind the aperture. The thorax has about 12 Jongitudinal muscles which forma wide band along the ventral border of the abdomen and posterior abdomen. When contracted this ventral band draws the pos- terior abdomen up alongside the abdomen, Stigmata are in about 12 rows of probably up to 12, but the thorax is too contracted to allow accu- tate counts to be made. Each raw of stigmata ts crossed by a parastigmatic vessel. The stomach is voluminous, and collapsed into irregular, more or less horizontal folds, The pas- terior abdomen is. about the same length as the rest of the zooid. It contains a double senes of male follicles and an ovary anterior to these. REMARKS The zooids of this species resemble those of Morchellium pannosum nsp., as well as Synoicum atopogaster and S$. concavituin. The naked bolster-like colonies differ from all these except the last, which, however, has firmer test, less crowded systems, and lacks parasngmatic vessels. The present species also has fewer rows of shgmata, and fewer per row than Syneicum atopogaster or Morchellium pannosum n.sp. Morchellium pannosum n.sp. (Fig. 39) DISTRIBUTION Tyre LOCALITY: Victoria (Port Phillip Heads, 13m, fast current, coll. J. Watson 10,9.77, holotype QM G12722). MEMOIRS OF THE QUEENSLAND MUSEUM The species is said to be common off Port Phillip Heads. DESCRIPTION EXTERNAL APPEARANCE; The newly recorded specimen is an irregular mass of sandy test, up to 3cm high, with the upper half of the colony di- vided into flattened to cylindrical lobes about lcm in diameter. FIG. 59. Morchellium pannesum n.sp. (holotype QM G12722) — a, part of colony; b, whole zooid; e, anterior part of zooid showing atrial lip, atrial aper- lure, branchial aperture and tentacles, and position of neural complex. Scales: a, Smm; b, Imm; ¢, 0.5mm, THE AUSTRALIAN ASCIDIACEA 3 A bare area with a large usually long. narrow opening in the centre is on the upper surface of each of the lobes, which contain single, or occa- sionally 2 cloacal systems, The cloacal cavities are relatively shallow, The test is very soft, jelly-like, Haccid and translucent. Internally, sand is present in the hase of the colony, but not elsewhere. The living zo- oids are reported to have been mauve. INTERNAL STRUCTURE; The zooids are robust. About 20 strong longitudinal muscle bands on the thorax extend along on each side of the ventral line of the abdomen and posterior abdomen in 2 wide bands, On the thorax there also appears to be a layer of transverse muscle fibres over the lengitudinal thoracic bands. In these contracted zooids (about 6mm long), the abdomen is shorter than the thorax, which is about the same length as the posterior abdomen, The latter curves ventrally as a result of the strong ventral longitudinal bands. The mm of the branchial aperture appears to have 8 very shallow lobes.in which there are faded pigment spots, although in some contracted specimens the rim Jooks puckered and gathered rather than lobed. A fleshy tridentate lip extends out from the upper nm of the atrial aperture. There are about 16 branchial tentacles of about 3 size groups, each group in a circle at slightly different levels in the base of the branchial siphon, About 15 rows each of about 20 long stigmata are folded along a parastigmatic vessel, The stomach is large, smooth-walled, and has imegular trans- verse folds that probably result from its collapse, The duodenum is moderately long, and a poste- rior stomach is in the bend of the gut loop, The voluminous rectum extends halfway up the tho- rax, A double row of male follicles are in the middle of the posterior abdomen, REMARKS In addition to the number of branchial lobes, the species distinctive characters are its fleshy body wall, strong (including transverse) muscles, roomy, transversely folded stomach, parastig- malic vessels, and distinctive lobed colony with flaccid test and large naked cloacal apertures. The transversely folded stomach is similar to that of Syroicym elopegester and S. trepicumt (Sluiter, 1909). Tt was previously thought to be characteristic of the genus Atopogasier (< Synoicum). Zooids resemble those of Synoicum atepogas- ter Kott, 1963 which has a similar number of stigmata, parastigmatic vessels and longitudinal muscles thal draw the posterior abdomen up alongside the abdomen, The present species is distinguished by its 8 branchial lobes and their associated pigment spots, Iransverse thoracic muscles and single cloacal system per lobe. Genus Aplidium Savigny, 1816 Type species! Aplidiun: lobatum Savigny, 1816, Zoaids usually are small and thread-like with a relatively, narrow thorax and gonads in a poste- rior abdomen that is continuous with (rather than constricted off fram) the abdomen. There are 6 or sometimes 8 branchial lobes, The atrial tongue is relatively small and rises from the anterior mm of the opening or from the body wall anterior to it. The atrial cavity also is relatively small. Longitu- dinal muscles.extend the length of the thorax and continue onto the abdomen and posterior abdo- men. Transverse muscles are in the transverse branchial vessels. joining with the longitudinal muscles in the parietal body wall through alrial connectives. The gul loop is vertical with a verti- cal rather than curved oesophagus and a vertical barrel-shaped stomach with parallel longitudinal folds in its wall. In Aplidiumt, like Synaicue, hut unlike Polyclinum, Aplidiopsis, and other Fami- lies of Aplousobranchia with a short abdomen, the post-duodenal or proxtmal part ef the mid-in- testine — between the duodenum and posterior stomach — has more or less the same diameter as the duodenum rather than being narrower, A shon distal section of mid-intestine lics between the oval posterror stomach (which usually is in the bend of the gut-loop) and the rectum. Often there are small pockets (caeca) each side of the proxt- mal end of the rectum, and these form a rectal valve, The anus opens-ane third to halfway up the atrial cavity. Larvae are small, the trunk usually less than Imm and most often less than 0.8mm. Three small, shallow adhesive organs on long slender stalks are present in the antenor mid-line. Median and lateral ectodermal ampullae sometimes are present anteriorly. and usually there are ectoder- mal vesicles at the anterior end of the trunk. These are 2,3 or more deep in a wide band, or inasingle senes, forming an arc along each side of the median adhesive organs. Sometimes they also occur in the median line between the adhesive organs (Fig_ 60). The vesicles separale from, or sometimes remain directly attached to the ecto- dem by, fine, sometumes branched, stalks. This MEMOIRS OF THE QUEENSLAND MUSEUM 508 ‘(oneumuEsseIp ‘satsads ouo Aue Joj Jaquinu astsaid May) Jou nq ‘uontsod aaneyal sayy Sutmoys) “dds wnipydy ut Yun [eAIE] ay) JO pus JOLayue at) Jo syuowsys jewuapidq ‘99 ‘O14 Variations in antero-median elements ws 3 asuasSiy4y 'y| a lunpros 'y| S asuarmaysny | © du winsodoiy c icin oe sapatsaa ex) 5 | w pyndure rc 8 a g 8 ueS10 aaisaype = —_ a & | sjoqurds 0] day 3 | a| 3 a Ie | Br ag Es u 2 |Se ze aynayn ge wurttanyyn “Y gs rf a a] | 3 ax BS 2 a BE baal g Bg lo & ae oo & 3s Bx Roa 2a B§ w [Fe = |86 =e i26 aig? rt gf & “a 3 L A ” 3 3 Ela ne g leo 3 ( a \ i é S judg-g wyessup~ i a ee eg —*] sajJ1saa OU Aysouaysod saptsaa Ajrousaysod saysisaa. SaPIseA Jo ae ajdyynu sajaIsaA 30 Dae aydurs sapisen au avyndune jesayey eyndure jesaye] ou sjualuaya [PJa}e]-O1a}ue ut suonEE A THE AUSTRALIAN ASCIDIACEA 3 constitutes a distinction from Polyclinum in which the only vesicles are those that branch off long strands of ectoderm which trail] back through the larval lest, one on each side of the mid-dorsal and mid-ventral lines and form postero-ventral und antero-dorsal patches of vesicles. These patches of vesicles occasionally also occur in Aplidium, but are seldom the only ones present, Asin Polyclinum, the larval ocellus and otolth in Aplidium are well developed. Colonies often are large, but appear to be well integrated, with zooids organised into cloacal systems and with an external morphology of the colony that enhances its interactions with the external environment. Thus. branchial openings muy be protected in depressed parts of the surface test, at the base of ridges or swellings of the Surface test (e.g. A. caelestis, A. crateriferum). Cloacal apertures usually are elevated above the colony on conical prominences (Kott 1989), The test of Aplidiwe is gelatinous, but often js rela- tively firm, and sometimes is further strengthened by dense inclusions of embedded sand. Many species appear to sort the sand embedded tn the test, large sand particles being found in the base und margins of the colony while smaller particles are around the branchial and cloacal apertures, where the test could be expected to be more flexible, The cloacal cavities generally are shallow, with only a thin layer of surface test over them. Atrial apertures always are at the anterior cnd of the dorsal surface, and variations in their position usually are a result of contraction of body muscu- lature. Deeper cloacal cavities sometimes de- velop, and the atrial opening into the base of the cloacal cavity with the atrial lip inserted above it is accommodated by the separation of the atrial lip from the aperture. Thicker surface test is ac- commodated by a long branchial siphon fe.g. A. crateriferum), The genus Sidnywm Savigny, 1816 (type spe- cies Sidnyum turbinatum Savigny, 1816) is dis- tinguished from Aplidium only by the 3-tobed (rather than 6-lobed) branchial apertures. The zooids form circular aystems which open on the upper surface of soft, cushion-shaped or lobed colonies with transiucent test, and with or without enerusting sand. The best known species of this genus are the type and the related §. elegans (see Berrill 1950), which both are common in some European locations. In addition to the 2 related European species, the known species with usually (but not always) 8 branchial lobes are: Syioicum appendiculatum Michaelsen, 1923 from the Azores. Although assigned to Sidnywn by Monniot (1974), the species with its deeply curved oesopha- gus bilaterally symmetrical stomach with furrows (rather than folds), seems appropriately placed in the genus Synaicwn. Sidnyum pentatrema Monniot, 1972 from Bermuda and Guadaloupe (see Monmiot 1983) appears to be an Aplidivm species in which the number of bran- chial lobes varies from 6 to 9, although & is the number which occurs most often (Manmniot 1972. 1983). These lobes are not equal in size, and probably several have subdivided to produce the higher num- ber, which aceardingly cannot be said to representa teliable plesiomorphic character indicating generic status for the taxon. Sidnyam pentatrema: Monniot and Monniot, 1987, known only from 2 small sand covered lobes trom Preach Polynesia probably is convergent rather than conspecific with the Azores specics. Apilidiam mernogensis (Brewin, 1956) from Chatham f.. New Zealand has characters in common with both S. pentatrema and §. appendiculatum. Monniat (1972) concluded that & appendicalarum from the Azores is distinct from A. menrocensis, but the telia- tionships between the latter and the Tahitian S, per fatrema: Monniot and Monniot, 1987 and A. mernooensis: Monniot, 1987 from New Caledonia ave not clear (see A, filifornie n.sp.. below), Aplidium amorphatum Kott, 1963 and A, protertans (Herdman, 1899) have (see below) 8 branchial lobes, Their other characters (including the stomach folds) conform with the genus Aplidium, Sidnvum indicum Renganathan and Monniot, 1984, a massive gelatinous colany from the Indian Qoean, has thread-like zooids that have the usual charac- teristics of Aplidium meluding a Folded stomach (see AL. prorectans, below). Thus, of the 8 species that are possible candi- dates for the genus Sidnyum, 3 — S. pentatrema Monniot, 1972: §. pentatrema: Manniot and Monntol. 1987; and A. mernooensis (Brewin, 1956) — with 5 rows of stigmata and 5 stomuch folds, probably are the result of convergent evo- lution, The number of branchial lobes in at least one of these species is unstable. The species ap- pear at least as closely related to specics of Aplidium as to one another. Of the other 5 known species, the 2 closely related and sympatric Euro- pean ones do not seem to have closer relation- ships with the Australian A. amorphanen and A. protectans, and the Indian Ocean A. indicurn than with other Aplidium species, Further, in the diverse genus Aplidium, both adult and larval morphological differences be~ tween species and groups of species appear to 510 transcend (in evolutionary terms) the few differ- ences between Sidnyum and Aplidium. Accord- ingly, in this work the genus Sidnyum is considered a synomym of Aplidium. Aplidium has a close relationship with Synoicum. The difference between them disap- pears when, in some Aplidium species, the stom- ach wall is distended and the 5 folds flatten out. Further the larvae are similar, and it is probable that these two polyclinid genera are closely re- lated phylogenetically. Resolution of the taxonomy, and identification of species of the genus Aplidium has been, and continues to be, a problem. Owing to post-mor- tem changes and other artefacts of fixation and preservation, the reported appearance of living specimens is often difficult to reconcile with their changed appearance in preservative. In the pre- served specimens, systems often are obscured by sand, and colours oxidise or are lost altogether. Zooids, already small and convergent, contract and withdraw from the surface, thus obscuring both their morphology and their arrangement in opaque and often distorted colonies. This confu- sion is compounded by a degree of plasticity in the shape of colonies which grow to accommo- date a variety of substrates and conditions. Fur- ther the number of specimens ayailable for examination does not always include a range of age groups or material from a variety of popula- tions so that intraspecific variation is often not known, or has not been recognised. Difficulties also exist in finding reliable taxonomic characters in these small, simplified zooids, and in their crowded colonies. The number of rows of stigmata, stomach folds, stigmata per row, and the configuration of the atrial aperture and lip, are important characters for species identification. Also of significance are characters of the colonies such as the distribution of pigment, consistency of the test, distribution and nature of test inclusions, form of the systems, shape of the colony and the presence or absence of a stalk. Usually of only limited significance at the species level are the relative lengths of differ- ent parts of the zooid (especially the posterior abdomen), the number of longitudinal muscle bands and the number and arrangement of testis follicles, which either are affected by the flexibil- ity and contractability of the zooids, or by vari- ations in form that occur with growth and sexual maturity. In identifying species of this genus, particular care needs to be paid to the counting of stomach folds. The whole thickness of the stomach wall is MEMOIRS OF THE QUEENSLAND MUSEUM pleated, and accordingly folds are internal as well as external. Internal pleats seen through the wall and counted with the external ones may explain the high intraspecific variation in the number of folds sometimes recorded. A transverse section of the stomach enables accurate counts to be made. The number of rows of stigmata is obscured when a part or the whole of the branchial sac is con- tracted. Inaccurate counts of the number of rows and the number per row occur when, in contracted sacs, stigmata are folded, and the interstigmatic bars and the perforations are crowded together. Fully relaxed branchial sacs are desirable for ac- curate determination of their structure, and dis- section to expose the stigmata is essential, as shadows seen through the contracted body wall are not a reliable way to count the rows. Nishikawa (1990) points out that the genus Aplidium is one of the largest in the Ascidiacea. Aplidium species abound in temperate as well as tropical seas. Because of the difficulties in resolv- ing the taxonomy of species in this genus some of the approximately 200 recorded may be inva- lid. Others have been lumped together, more criti- cal examination showing that many previously considered conspecific taxa are actually distinct from one another. Most of the previous works on the Aplidium fauna of Australia are based on relatively small collections from limited areas, viz. from New South Wales (Herdman 1899), northwestern Aus- tralia (Hartmeyer 1919, Millar 1963), south-west- ern Australia (Michaelsen 1930), Port Phillip Bay (Millar 1966), eastern and north-eastern Australia (Herdman 1886, Herdman and Riddell 1913, Kott 1966, 1972c), South Australia (Kott 1972a,b, 1975 and 1976). Kott (1963) dealt with Austra- lia-wide collections. Sluiter (1909) in his report on the Siboga collection encountered only 5 spe- cies of Aplidium of which 4 occur in Australia. Only 25 species of Aplidium spp. have previously been reported from Australia. The number of specimens available for exami- nation in the present study has made it possible to more accurately determine species parameters and resolve synonymy. The identity of most of the 16 previously known indigenous Australian species and the 4 recorded also from the Western Pacific has been confirmed, however the 2 spe- cies said to be conspecific with species recorded from New Zealand, 3 from South Africa, and a pantropical species (A. lobatum), were wrongly assigned. No Aplidium species has been found to have a range from Australia to either South Africa or New Zealand, and the number of apparently THE AUSTRALIAN ASCIDIACEA 3 indigenous Australian species has been signifi- cantly increased as a result of the present review. Aplidium, with 46 known species, including a relatively high indigenous component, now Te- corded from Australia, is one of the most diverse am of the Ascidiacea in these waters (Tables 9), Key to species of Aplidium recorded from Australia 1. Stomach folds more than 8 .......cs::see ene 2 Stomach folds not more than 8 ...,.........cce00 25 2. Cloacal apertures few and terminal or central; parallel lin mG or radial cloacal canals Cloacal apertnres | numerous sand scattered over the colony surface; no long, parallel longilu- dinal or radial cloacal canals ,,......2.4..se01e 9D 3. Colonies Mat-topped sandy lobes adhering lo one another ........01..-. ... A. congregalum N.sp, Colonies not sai sry lobes a meng to one another .. sees pesetepaereer A 4. Colony with thin, leathery stalk. A. australiense Colony without thin leathery stalk ............0 5 5. Stomach folds Jess than 20 Stomach folds more than 20... 6. Stomach folds 16..............A. brevilarvacium Stomach folds more than 10............. A, parvum 7. Colony sessile ortop-shaped, without a stalk... Sess adbacbanthasaabarsnataasoadssseothossechosbathacch A. allarium Colony not sessile or top-shaped, with a stalk..8 8. Head of colony with sand externally; larval trunk with epidermal vesicles .,..0......6000. eater Cogpaydsreelapdorbaplcedas d0eBpebe A, geminatum 1.sp. Head of colony without sand externally; larval trunk without epidermal vesicles ......,...-.0:.0++- sesoctbesoavbavoad pvseseersersereseseeeAhs inflorescens N.Sp. 9. Stigmata in 5 rOWS -.......ceeesessverseieemsersseres LO Stigmata in more than 5 rows .......... 10. Atrial aperture witha lip .. A. minisculumn.sp- Atrial aperture withouta lip...,...A. depressum 1]. Zooids in long conspicuously branching dou- ble-TOW SYSLEMS «00... .:ccseseeseeseseeeeeesseeseeneeee 12 Zooids not in long ar aa tal nee double-row SySteMs -......ccerceeceeees 14 12. Sand embedded throughout....,....... A, elatum Sand notembedded throughout... 13 13. Branchial siphon surrounded by wide band of white, opaque cells; larvae with single series of epidermal vesicles each side of anterior PHVA TAHT 5 sep siees psa gepavingnon'ngey A. multiplicatum Branchial siphon not surrounded by wide band of white, opaque cells; larvae with multiple series of epidermal vesicles each side of ante- THOT MIG-HINE «00... eseseneereeneereerereens A, OPACumM 14, Stomach folds not more than 15 .W.............. 15 Stomach folds more than 15 .......).:.;csasease U7 15_ Sand embedded in the test .............0c. eee 16 Sand not embedded in the test ,.........c.c0ccseeevee sottistieerninenienmene A. fluorescum n.sp, 16. Sand embedded in surface layer of test; zooids in conspicuous circular sysiems.... A. jacksoni Sand embedded in basal layer of test; zovids not in conspicuous circular mee the ’ Antibbaatiateecominacseayacsets wh friggsense 17.. Stigmata more than 20 per row; brood pouch present ... sagsonsegesnsasesecperwespevere red by LER E Stigmata loss thew 20 per TOW; cai pouch not PIRESE RE cB reerecersteta seas aneatanesinsrasnenmpsaaeuiuenas 18 18. Stigmata in more than 12 rows wee 1D Stigmata in 12 tows or fewer 19, Stomach folds 18; stigmata not more than 8 POL TOW ve ceuiesne A. Belasinum TLSp, Stomach folds 20 or more; stigmata more than BPEL LOW 4: ssscisveisssssinsepeocsensensensensessesenatnneree OU 20. Stomach folds oblique ... ro aulsabidevoeWquevegvontehdeniepessieh A. gastrolineamumn n. sp. Stomach folds ot ObQUe wo... cesta slodelebeontabsssuoesrsvotassvainsissinssnt Aa POBUSTUIN SP. 21. Stomach folds 30, oblique ... * A, multilineatum n, sp. Stomach folds Tess ‘shan 30, not oblique ...... 22 22. Systems circular. .cseciessissssessessse rene 2 Systems not circular... A. resarium n.sp. 23. Sand on upper surface of colony ,,...;.++ bata APRIR TONE Tye retire pine store A. ineubatum nsp, Sand not on upper surface of colony ,.....,.... 24 24. Stigmata more than 10 per row ... A. /odixn.sp. Stigmata not more than LO per row ........-..06 betutermteenersies A. OFNGTUM NSP. 25. Atnal lip separate [rom rim of aperture ,..,.. 26 Atrial lip not separate from tim of aperture .35 26. Colonies undivided, upright and regular .... 27 Colonies not ondivided, upright and regular 512 MEMOIRS OF THE QUEENSLAND MUSEUM 27, Colony aay (eb cepeteetestereeeey A Magnilarvuni n.sp. Colony not + sandy cxlamally acct A. coniferum 28, Systems CHrCular 0... ccesscpeescersepeeresenerseee Systems mot chrcubar .....c..2204:e css cseneeceepseeeep ene 29. A single system percolony lobe ... A. directum More than a single system per colony labe . 30 30. Branchial siphon with bulging branchial sphincter... ites A, erateriferum Branchial siphon 9 without bulging branchial sphincter .....c..00...A, /unacratum nom. nov. 31.. Cloacal systems irregular; cloacal apertures at junction of 2 or 3 caMAls wc. scsi 32 Cloacal systems regular: cloacal apertures in centre of radially converging canals........... 33 32. Common cloacal canals surround circular to polygonal zooid-free areas .. Abba derabesemsseataegaty 1 Ay “Tenticuliam a, sp. Common cloacal canals ds not surround circu- lar to polygonal zooid-free areas ......, peas keeparseueniatiessevinsserreesavincssaivesstivecn tle caelestis 33. Stomach lolds 5.......c.:ecececeenpererseeneeen D4 Stomach folds 8 o..ccse A. griseum n.sp. 34. Stigmatain 18 rows.............A. clivosum n.sp. Stigmata in 12 rows ....cc0.. A. rubricollum 35. Stigmata in 4 or 5 roWS wie 36 Stigmata in more than 5 roWS ....ceerreee yen D9 36. Colony a3 dimensional reticulum; 8 stomach Paldsy .vrcccccc pete eocee: A. acroporum f.sp. Colony not a 3 dimensional reticulum; 5 stom- BCH FOS 0... ns scssesceencpgnesceperneondernssnenesnenas OF 37. Stigmata in 4 rows... A, distaplinm n.sp. Stigmata in 5 TOWS cs sceeene ss puceeneecre OR 38. Stalks hides one system per terminal branch serine A baceulum n,sp. Stalks 1 not ong more than one system per ter- minal lobe... ... A, filifarme n.sp. 39. Sand sparse or absent from internal test..... 40 Sand moderate or crowded in internal test ,. 42 40, Branchiallobes 8 ,,......0.00 sassadtessaitnevedh sainavve 41 Branchial lobes 6 ..,. A. parastigmaticum n.sp. 41. Parastigmatic vessels present... A amiorphatne Parastigmalic vessles not nidabaie's xe uA. protec ans 42, Colony asingJe system .A. paralineatum nsp Colony nota single system voce ae 43 43. Stigmata more than 10 per TOW prscsreseerene CONE eared SHS TTA A. macrolaburum nisp. Stigmata less than 10 per row ..........00 wn 44 44, Cloacal canals surround exlensive irregular patches of zooid-free test; systems not crowded; embedded sand not crowded............ ineddniveuaveponcogsresospe son .. A tabascum n.sp, Cloacal panails do not ‘surround extensive ir- regular patches of zooid-free test; systems crowded; embedded sand crowded ...........- 45 45, Cloacal canals surround small circular zooid- FOG ATEAL...csc.cccresserevsvesivese saysteenvereenyereeazare FO Cloacal canals surround narrow zooid-free THO BES vescscsspesecceesventssetsntseree tenures Ay Citteri 46, Larva with median ampullae; southern Aus- UAlIAN .,cycpccrseecerenrersareseeeee. PELFOSUMTLSD. Larvae without median ampullae; not southern AUStraliaty occurs A, solidum Species recorded from waters adjacent to Australia Aplidium cellis Monniot, 1987 has double rows of zooids around raised zooid-lree areas of test as in A, tabascum n.sp., from which the species is distin- guished by its smaller larva (less than 0.6mm trunk) and the sand which is said to completely encrust the surface, albcit it is less crowded over the cloacal canals, Although the colony resembles A, /enticulum n.sp. the zooids differ, having the atnal lip from the upper rim of the opening, similar to the zooids of A. rifteri (> A, lobatun: Monniot, 1987), The species may be conspecific with A. lobalum Savigny (see below). Aplidium flavalineatunt: Monniot 1987, trom the Coral Sea, has zooids (with brood pouch, large thorax and short posterior abdomen), and possibly also the lar- vae (with epidermal vesicles around each adhesive organ attached to the epidermis by narrow stalks) like those of A, altariun, Aplidium altarium has only a limited number of systems in each colony, each consisting of double rows of zooids converging to central or terminal cloacal apertures, The Coral Sea specimen has circular systems and a brood pouch like A. uteule. However, the latter species has more nu- merous lateral vesicles in the larval trunk. Aplidium lobatum Savigny, 1816 from the Gulf of Suez, later (as A, africanum Sluiter, 1905) reported from the Gulf of Aden, is said to be irregularly lobed but smail (5 to 6mm thick), with yellowish white zooids in lines amongst the sand embedded in the transparent test. The upper surface is divided into flat-topped oval cushions which have deep furrows surrounding them. The cloacal canals into which the zooids open are in these furrows. Sand does not THE AUSTRALIAN ASCIDIACEA 3 31 adhere externally, stigmata are in 9 or 10 rows of about 12, the trifid atrial lip is part of the anterior rim of the aperture, The posterior abdomen is short, the male and female gonads are not present in the same zoaid at the same time and the larval unk is 0.5mm long (Michaelsen 1920), In his review of the species, Michaelsen added records from the Mediterranean (through the synonymy of A. tremulum Savigny, 1816), and suggested a range extending to Malaysia and Indonesia through synonymy with Aplidium tre- mulum: Sluiter, 1909, However, this requires confir- mation as most assignations from the western Pacific by Tokioka (1967), Nishikawa (1984a), Monniot (1987) and Monniot and Monmiot (1987) appear to be synonyms of A, ritieri (see below). Aplidium lobaium: Kott, 1963 is conspecific with A. macrolo- batum usp. Aplidium tabascum n.sp. and A. cellix Monniot, 1987, with smaller lobes on the apertures, and elevated cushions surrounded by depressions into which the zooids open, have both colony and zooids like A. lobatum from Suez. Particularly A. cellis, with its embedded sand and small larva, is indistinguishable. The small specimens from the Marianas (A. aff. lobatum: Tokioka, 1967), with the stnal lip separate from the aperture, is not this species. It is similar to A, caelestis or A. nadaense Nishikawa from Japan, Auntie specimens assigned lo A. lobatum (see Van Name, 1945) have fewer rows of stigmata than the types, the colonies are different, and they probably are not conspecific with the Red Sea specimens. Their similarity could well be the result of conver- gence. Aplidiam maru Monniot and Monniat, 1987, from French Polynesia, has a larva with median ampullae and, on each side, a band of lateral epidermal vesicles at the anterior end of the trunk (see Fig. 60), There is no other species in the group with similar larvae which also has circular to oval systems of zooids arranged around a protuberant cloacal aperture, 5 stomach folds and an atrial hp separate Irom the opening. Aplidium merravensis (Brewin 1956) trom the suban- tarctic Chatham 1., New Zealand (sec also Millar 1982) is reported from the tropical New Caledonjan waters by a single colony (Monniot 1987). The col- ony has sand externally but not internally. lis upper surface is lobed, each Jobe containing a single circu- lar system. Zooids have red thoraces, with 8 bran- chial lobes, 5 stomach folds and 5 rows of stigmuta. The occurrence of this species in a tropical Jocation is surprising and the identity of the New Caledonian specimen requires confirmation, especially in view of the possibility of convergence in species with zooids as small as these, The New Zealand specimens have almost twice as many sugmata as A. filiforme n.sp. (sec below). The larvae of both the Chatham [. and New Caledoniin specimens are larger than AL filiforme nsp.. but they arc otherwise all similar to one another. ~ Aplidiam meltipapillatum Millar, 1975 from Hong Kong und New Caledonia (Monniot 1987) is charac- terised primarily by its larvae with 5 adhesive organs. Apliditen aadaense (Nishikawa, 1980), a species prot- ably indigenous to Japun has the atrial lip separate from the aperture, 13 to 15 rows of up to 8 stigmata, and 5 stamach folds. The colony is smooth-surfaces and the systems are-not as conspicuous as they are in A. caelestis. Monniot and Monniot (1987) and Moa- riot (1987) assigned specimens, from French Poly- nesia and New Caledonia respectively. to this species. Aplidium naadaense. Monniot, 1987 from New Caledonia has 14 to 16 rows of stigmata, but branchial apertures are in furrows in the surface, between sinall polygonal arcas, resembling A. seli- dum, which, however, does nol haye the atria) lip separated from the aperture, Aplidium nadwenses Monntiot and Monniot, 1987 has a longer, narrower zooid bul fewer rows of stigmata than the type. Neither the number of stigmata per row nor the form of the systems is reported. The small larva and other features of the zooid are similar to A. lobarum, but ihe long narrow posterior abdomen is noi, Probably neither A, nadaense: Monniot 1987 nor A, nadaense: Morinint and Monniot 1987 are conspecific either with the Japanese species or any known Austratian ones. Aplidium woo Monniot and Monniot, 1987 tnom French Polynesia resembles Aplidiaen ritteré in its long, winding systems, atrial Jip from the upper border of the apertures, 10 rows of about 6 sigmata, 5 stomach folds, short posterior abdomen containing gonads of only one sex at a time, anda larva without median vesicles or ampullac and a band of lateral vesicles, It is distinguished only by tie absence of embedded sand usually associated with A, ritreri, the: band of lateral vesicles is not as wide, and the larval trunk (0.7mm long) is longer than that of A. rilteri. Aplidium acroporum 1.sp. (Fig, 61. Plate 10,6) DISTRIBUTION TYMi Locality: South Austnilia (The Gap, near This tle £. Spencer Gulf, coll, 8, Shepherd 8.4.87, holatype SAM E2534 OM GH4/69; Harseshoe Reef near King- ston, 36°35.0"E, flat rock platform, small overhang, sand patches 20m, coll. R. McCauley 17,2.89, paratype QM GH5443; Kangaroo [, Cape D’Estaing, Eniu Bay north of reef, 33°34.3°S 137°S0.2°E brown algae cov - cred rocks merging lo sand and sea grass 12m, coll. AIMS Bioactivity Group 30,189, paratype QM GHS43 1. PURTHER RECORDS: None, DESCRIPTION EXTERNAL APPEARANCE: The colony forms i 3-dimensional ceticulum of hard, rigid, sandy, MEMOIRS OF THE QUEENSLAND MUSEUM 514 ‘aqeraduray ‘a ‘TeoIdon ‘n toytorg WaiseMy ‘AAA ‘SnouasIpUt “y | Uog WaIsoy, “ LOT “ “ Buoy] ratral “ 0% 01 PS WINgy0D ary ungodo “y sseu Aysary Jooy Joumeg u2aaa JepnsaLt 1D Weypiou Wey soBpNS 60:L-T euou 0) ay] -J994s “ O@-FI‘11-6 “« t= PS WINGyD0D ard unwayjdiyjmu “y snoJowinu stg é{ MoYBnonp —aeJaurey ysudn E 38 ‘Suryouesq SI uelyensny 19 ary una “y JOE Aqyeseq uorysno yoys ra kral - 0z dy wooded ny ‘dsuwniuvsos-y snoJoumnu ‘dsu $09 ymnoySnonp peddoy-ey isudn r rates syerper ral Avg uoja10yy ny wunjo8a18uo02 “y Weng SOE P « “ 871 “ 81 sseg oy Aueqry ary wunaind "y Hog Waysayy, ‘ds‘u 60'r ATuo yTeas “ “ SI-FEST “« 0% FIND sJe0uedg ary suagsasoyful “Y qs uelensny 1D QO:T — moysnomp “ “ Or “ OT 9 PS WINGys0D avy = wndDALDIIAaIq “V uog € 0} T UlaIsa AA 0} I1NSTG S60:E ATUo TeuIa)xe 7 Z ZI-OL'ST ‘sMmol yoTTezed €Z-07 uelensny 19 ay ‘dsu wnjpunuas -y snoyauinu wreyaloys 0} CLT auou poyeis suo] SST ‘sMol [aq[esed OI “Yory ayoreysoy avy asualjo4jsno “Y Ps wingys0) OE euou — suorysno 1y3tdn yoys SZ‘01-6 auo‘yerpel O€-SZ 0} Aeyoryy NdM uiniiDyD “Wy (wu) TSU] eryensny yuny uerpeur punose Spoyeqnoutr pues uawiopge MOI/‘OU'SMOI Joquinu ‘ULIOJ Sploy astmyooponue juonduoseq PIO ‘ou :oBAIR] «=: poppequigy Auojop = JouIaysod “Ou :ByeUISNS rsuaysX§ — YoRUIOIS asuey orydeiZoes01g sotsedg SPO} YoRUO}s ¢ URY} aOUI YIM pure ‘Suruado oy wos payesedas jou dy eine ay wm saroeds :w dnoip Blersny Woy pepiosar wnipydy Jo satoads Jo siojovsreyo Jo ATewUINS */, ATAV.L 515 -ayetoduiay ‘ay ‘yeaidon ‘n ‘oytoeg waysayy ‘dA, ‘ShouadIput *y I THE AUSTRALIAN ASCIDIACEA 3 winpnonar POT Inoysnony JEUOISUSUIp ¢ i L-9'¢ a g vs aty ‘ds u wnsodoson wy snoseumnu Por s]aays 0] UOTYysno - ZI-or's upnods ol OA, ayy ‘ds winypnosiunu -y diy Siaqepung jetne ou SEO’ i syeays uoys ore SNOJOWINU *f, Il 0} "J UOIaHy dM wnssaadap “y saqo] wsudn *] uoray OZ Ayyeseq 0} Sjeoys Buoy ole é ST ©) pg wingys09g ard asuasdauy “y ayf[-129qs ‘| prezry SOE . 0} uoTysno yr 9OL 4 PI-Z 0} '] uoIaH ny ‘ds‘u wnosasonyf -y BULIES * LO:S-Z “ Sysoys yoys S711 “ SE-SC 0} "] WorsH dM anen'y peddoyey yonod pooiq sc'o'9 = noygnomp Wysudn oy aye, ib SL‘OL a Sz Avg uojajopy ayy -dsuwnjnqnoui -y LOZ auou peddorrey iysudn 4 eral 3 St PIO aS ny ‘ds uwnyouo -y UdA8 ddRyINS i = noysnonp 7 ul OZ Ai ral uosyoer WOg avy qosyoul "y Joey Jou « Ob ouou « mn S01 “ 81-91 Wealh WayuON ny ‘ds‘uwnuisvjas -y swaysAs I punouns uosmag 01 14aIg ‘dsu sespu 0:4 Ajuo peusayxa J nx +U9T a +7 unyensny 15 ay — - unjoauyoisod y eruRuise |, ‘ds‘u “ 60% Ajfeseq = worysn zeynom9 “ 91-01 “ O€ ps Wingyo0D, ay winjpaUurnynu “vy SUOIYSNo [eoTU0D PIUPUIST], “s 90° TE “ 0} jeourayds “ TL91-F1 “« 0c 0} Aueqry ay ‘ds‘u wnjsnqos "py uaAo snoaunu Hog Wisse, aoepins L0'9 auou aytpeoys Buoy SNOJSWINU‘OT te[noqo §Z O1F[ND sa0uadg ay “ds'u x1po) "y (Wu) Tsue] eyensny yung uerpeur punor ‘pereqnour pues uawopqe MOI/OU'sSMOI = JaquINU ‘LUIOy plo} astmyooponue —_, uonduiasaq BO ‘OU ‘oRAIRT = pappequiq Auojod = JOLIaysog “OU :EyeWIaS rsulayshg YORTUOIS asuey omyder8oa8org saraadg (quoZ ‘y dno1p) elyensny wo1y pepioser wnipiydy Jo satoads jo siajoevreyo Jo Areumung */ ATAVL MEMOIRS OF THE QUEENSLAND MUSEUM 516 ‘qeoidon ‘n ‘ayrsaduiay ‘af ‘ouloeg Waysam ‘AA, ‘SnouasIpUT ‘Yy - "Sploj YoRUOYS 6 JO g IIH “ds-U wnasT 'y st uondaoxa ay , surajsAs punouins sespu Cg'0'1 ~ wysudn i " TUL “ “ “ Uae aorpns So'O'T is peyyeis 1ysudn 7 Ss0'7 “ pagol wydudn a “ Tel “ “« “ “ OT “ “ “ suorysnd Yortp 10 4 SLUT “ adpam oAIsseul “ suia)sfs punouns saspu LOtl fe a sura)sXs ayeredas sad pir 69'0'T “ a és Spunoul payaso punouins stuaysAs Sg'0'T Noysnonp syeoys Yor Suoy (umn) duay yung uerpaur ‘pereqnour pus uawopqe RYO ‘ou :oRAIe] ~=—s pappaquig Auojoz JOWIaISOg aqoy Jed ¢ JO Z ‘[erpel dnoip OE ov Je[NaM woouded ny ‘dsuwnasus ‘y snorauinu wasrg ‘dsu 07:17 6 uelensny 15 ay wnaippuspuU “y snozawinu ‘yeiper Arg projomy 0} 877 0} Iv[NONS Avg uoist[q ay umniafiuod “y aqoy Jed auo Avg woa10yy Pl‘eI-O1 ‘ye[nogto 0} By]NUOID ay unj2a4Ip “Y ASN snojawnu = Jaany Suruueyy “aou “wou SI-ZI‘sI-O1 saepnomo Q} saquRAIaZ ay uinjosovuny “y snozauunu = Jn ynourxg PI‘IZ-81 taepnoao 0} “] uoIapy dM windafisaipsa “ye snozaumnu ‘| UoIap OF ZTUSI-OT 01 Ma *yerper pur[pay uog any ‘ds‘uwnsoayo -y uog Wasa snozawinu 01 1ysIg Pr:zI-11 ‘Teper uelensny 1D ary wnpoo Gna *Y snoJauinu "| prezrq vI-O'8I-O1 —-1Y8rens Zuo, =o} Avg yey arn‘d AN susajapo “Wy snozauinu "] uoIaH 8:0Z-9T {paains Suc] 0) pg wINgys0D ay ‘ds wnjnoyuay -y eyensny punoze MOI/OU'sMOI —Jaquinu ‘Woy = astMyoofonue =—_-uoNduosaq, “OU :RyRUIsNS :suayshg aduey o1ydersoas01g sotoedg : SP[OJ YOeUIOJs ¢ UTM ATTeNsN pue ‘Suruado ay) wo payeredas diy jeje ay wIM satoads :g dnoip BleNsny Wo pepiosal wnipydy Jo satsads jo siojovreyo Jo Areulwing “8 FIGVL 517 THE AUSTRALIAN ASCIDIACEA 3 "Teordoy ‘Nn tayeraduiay ‘ay oytorg Waysam ‘gq AQ ‘SnoUasIpUr “yz “Spfoy YORWOIS p IPM wmoydsown “y st UoNdaoxXa aYL. | ds'u wnoy s]ass2a onewsnseed wi a y 7 ZUST ¥ *] uoray, ny -pusisvind “y s]assaa onRUIsTsered Atg SIAlaf 0} ‘saqoy TeTyouesq g rants rr “ “ OUST “ Aeg wowst{[y ary wnipydiown “y Jooy Jone g Soop JO snozeumnu 1D Wetpiou saqg] jemoueiq g cxote auou saroyds m OZ+Z-B1 ‘eynoms Ir} Ol MSN Dd A supjzajoid “y £107 dsu U4 oysnosyy Jejnsam Buol TERT ‘Jatrered ‘Buoy eTURUSE |, yyw ‘uinwauynsod -y snoJauinu ‘3u0] “y prezry ‘ds‘u ampade uo sdy ods] S90 x i f SI‘01-6 oq JE[NoI 0} "] UoIapT y uinjoqojossw -y snoyaunu cS OE “ “ “ LZI-10 ‘Buoy ‘] UoIsH dM Hail “Y suorysn3 PS winqys05 LOZ Ayoyed Jo syaoys 2 gist d ‘dp wosude9 vy ‘dsuuinaspqn -y ‘yory saidueq pur Siaqepung SP'0'S-Z in aeyaurey yysudn yoys 8ZI-6 a 0 Aeg stalag arn'y ‘ds‘u wnpijos “y winjnonar snaraunu SOE ae [euorsuawip-¢ o 06 {pacino vs ay ‘dsuunsoyad -y aqoy Jad auo bt ™ pexyeis duoy 9g “eynoso WS ay ‘dsuwnjna9nq -y paddoy-ey aqo] Jad ¢ 10 p dnaip LrO-E'0'€ ke qysudn 7 9°¢ sqeyNoD woonded My ‘dsuauopfipf -y snosaumnu irens sseg 0} ported eyewsns Jo smol TSO moysnoimp Jensen yoys Zp vA Jy s20ueds ay ‘dsuwnydvisip -y (ur) 3a] elyensny yun uerpaur punoze ‘paieqnout pues uauropar MOIOU'SMOI JaquINU SULIOJ «= STAM OOTONUe qosuey BNO “Ou :avAre'T] pappequry Auojod JOLWIa\s0g “ou sess rsuuayskS adury oydersoes01g satoadg SPIO} HoRUIONS ¢ MEM Aqjensn pue ‘omnyade ayy wosy payeredas jou diy pete ay} yim sotoads :- dnoip Byensny Woy papiooas wnipydy jo saroads Jo slajovseyo Jo ATeWILINS *6 ATAVL 518 branching, anastomosing stalks, usually with horizontal terminal branches that lie in the one horizontal plane, The whole upper surface and the sides of each terminal branch are covered with naked, saucer-shaped circular concavities of 2 to 3 mm diameter, separated from one another by raised sandy ridges. The base of each of these concavities consists of a thin layer of test overly- ing a shallow thoracic cloacal cavity with a cen- tral, sessile cloacal aperture. In the paratype colony (QM GH5431) some of the branches of MEMOIRS OF THE QUEENSLAND MUSEUM the stalks, including the terminal ones, are often flattened vertical lamellae. Zooids are in circular systems of about 10 around each cloacal cavity. Qwing to the de- pressed, transparent, clear, naked test over each cloacal cavity the surface depressions appear deeper than actually they are. Sand is crowded throughout the remainder of the colony. The zo- oids, smaller than the sand grains, are embedded in the test between the crowded sand grains. Rigid open compartments in the test, which occur in } "tg DB c o “Up ate i Cera FIG. 61. Aplidium acroporum n.sp. — a, part of colony (paratype QM GH5443); b, surface of colony showing naked areas ayer cloacal cavities with branchial apertures around the margins (holotype QM GH41 69), ¢, zooid, with embryo in atrial cavity and posterior abdomen separately (paratype QM GH5443); d, larval trunk, with large test cells (?) posteriorly (QM GH5341). Scales: a, lem; b, Smm; e,.0.2mm; d, 0.1mm. THE AUSTRALIAN ASCIDIACEA 3 some aplousobranch species to accommodate the zooids, do not occur in the present species. The maximum dimension of the available specimens is about IQem and they are Sem high. However, in no case was the entire colony taken, andit is probable that they are particularly exten- sive in situ. The branches toward the top of the colony have a diameter of Smmi, but the lower branches are narrower, and the basal vertical stalks taper like roots, INTERNAL STRUCTURE: The zooids are particu- turly small, up to 2mm long, and thread Jike. The thorax, abdomen and posterior abdomen are of about equal length, except in those species in which the latter is contracted. The branchial and atrial siphons are short, the former with 6 (some- limes bifid) lobes, and the latter with a single pointed lip from the upper rim of the opening. There are 5 rows of stigmata with 6 or 7 ina row. The small stomach has 8 folds. A short posterior abdomen has 4 or 5 male follicles bunched in it in specimens collected in May. One colony (QM GH543 1) collected in February has mature ova in the short posterior abdomen, and 2 embryos are incubated, one at the top of the abdomen and another completely obliterating the atrial cavity- The larval trunk js spherical, 0,4 mm in diame- ter, and the tail is wound three-quarters of the way around it, There is a Jarge cerebral vesicle with otolith and ocellus. Epidermal vesicles are in a band about 3 vesicles deep to form a conspicuous are on each side around the anterior half of the trunk. These develop from a circle of epidermis around the adhesive organs, subsequently becom- ing detached and scattered in the test. Large morula corpuscles are scattered in the test over each side of the posterior half of the trunk. The 3 adhesive organs are wide but shallow, with short stalks. REMARKS Characteristics of the present species are the small zooids and the form of the colony, with its tigid, narrow, sandy branching stalks, and surface depressions separated by raised ndges which give the colony the appearance of the corallites of an acroporid coral. The anastomosing horizontal terminal branches of this species, with zooid systems opening on the sides and upper surface, probably fonn a mat above the sandy substrate, with the rest of the reticulum (which lacks zooid openings), buried in the substrate. Such a growth form ensures that the colony is firmly anchored, and must add consid- S19 erably to the consolidation and stability of the substrate. The cloacal systems are smaller than similarly Shaped ones of A. Jutacratum nom. nov., from which the species is further distinguished by tts 8 {rather than 5) stomach folds, 5 (rather than 9 or more) rows of stigmata and the atrial lip which 15 separate from the aperture in A. Junacraturn, Aplidium altarium (Sluiter, 1909) (Fig. 62. Plate 10c,d) Amaroucivn ultarium Shuiter, 1908, p.105. Aplidium altarium: ?Millar, 1963, p.€95. Katt 1966, p.282, Not Michaclsen 1919, p.90. Not Millar, 1956, p.914. DISTRIBUTION New RECORDS: Western Australia (Broome, WAM 873.83 OM GH27/8; Ningaloo Barrier Reef, WAM 1029.83, Houtman's Abrolhos, WAM 79.75 189.75 388,75 763.82 754.83 837.83 1030.83 OM GH2i46 QM GH5557; Shark Bay, WAM 871-2.83 OM GH2148 GH2147 1026-8,83 209.88; Cockburn Sound, WAM 75.75 143.75). Queensland (Moreton Bay, QM GH3478-9; Capricorn Group, QM GHS307 GH552!-2 GH5558-9; Broadhurst Reef OM GH5131; Sarina, OM G4996, Lizard f., QM GH330-1 GHS5480). PREVIOUSLY RECORDED: Western Australia (?Cape Boileau — Millar 1963). Northern Territory (Darwin — AM ¥1396 Kott 1966), Indonesia (Sluiter, 1909), DESCRIPTION EXTERNAL APPEARANCE: Colonies are circular to oval cushions up to 3cm in greatest dimension, lightly attached to hard substrates such as coral skeletons as well as weed. In life the colonies are less flattened, sometimes almost spherical or top- shaped, narrowing basally. They are found either in large populations, their margins probably in contact, to form a mosaic-like covering over the substrate; or they are bunched together. In d via servative most of the colonies from Western Aus- tralia collected in 1974 to 1979 are a purple colour. Apparently the colour is progressively lost when in preservative for longer, as some from the Abrolhos collected in 1963 (WAM 754-82) are beige transparent cushions. The colour of living colonies is recorded for only one of the Western Australian specimen lots (AMP1 60 QM GH5307), which is said to have been purple and white, although photographs of it are white: It is possible that the colour notes were taken after the specimen was collected, when some oxidation 420 had taken place. Living colonies from the Capri- corn Group were described as ‘brick’ coloured with white in longitudinal lines down the sides of the colony, or “pansy purple’ (Ridgeway 1886) with beige or yellowish zooids, or ‘brown ball ascidians’. The white lines referred to appear to have been calcareous faecal pellets in the cloacal canals. The specimens from Lizard J. and Myora are reported to have been ‘maize yellow’ (Ridge- way 1886) with a red dot each side of the aper- tures, When present, the purple or brick pigment par- licles are crowded in the surface layer of the soft but turgid gelatinous test of the preserved colo- nies, becoming more sparse below the surface, and absent altogether from the basal test. Between the pigment particles, and after the pigment has faded, the test is clear and transparent. Sand is neither attached to, nor embedded in, the colony. CIN | ao ca uo 0290 5en 00" y \ hy 4s ‘ i — y 7 ’ Con o YS Pl ¥ ; apo ge out a == 7 Tle ,o th SKA Fee an 99 om”, A ar F = i 4 7 ia MEMOIRS OF THE QUEENSLAND MUSEUM Usually a single, central, large, sessile and ir- regular common cloacal aperture is on the upper surface, and branching cloacal canals conyerge to it from the outer margin or the base of the upright colonies. Some colonies have up to 3 systems, Zooids, cream in preservative, are arranged in rows along each side of the cloacal canals. They are tightly enclosed in the firm test and are diffi- cult to remove, Their large, white, atrial lips can be seen stretched out along the roof of the cloacal canals, and inserted around the rim of the cloacal apertures. White opaque vesicles in the surface test around the sides and upper surface are termi- nal ampullae of test vessels, INTERNAL STRUCTURE: The zooids are about 6mm Jong when contracted, the thorax, abdomen and posterior abdomen each being about one- third of the total length, About 20 fine longitudi- nal muscles extend from the thorax along the FIG. 62. Aplidium alrarium — #, colony (QM GH5307); b, zooid, with posterior abdomen separalely (AM Y 1396); c, larva (WAM 79.75). Scales: a, lem; b, Imm; ¢, 0.2mm. THE AUSTRALIAN ASCIDIACE, 3 length of the zooid, and even thotigh all of the examined zooids are contracted, the muscles are inconspicuous. The branchial siphon is short. Only a short imperforate area is postertor to the prepharyngeal groove. The atrial tongue is of variable length, and is undivided or sometimes divided into 2 or 3 points. ft arises from the anterior rim of the atnal aperture. The atrial aper- ture is a vertical slit, bounded on each side by a projecting convex lip which has muscles in its wouter, convex, vertical margin. The homology of these muscles appears to be with the sphincter muscle usually found around the atrial aperture, The unusual configuration of the atnal sphincter in this species results from its component muscle bundle being drawn together by fine longitudinal muscles, one each in the middle of the anterior and posterior mms of the aperture. to form, respec- tively, the anterior and posterior extremities of a vertical aperture. As well as being long. the branchial sac is wide, with up to 25 stigmata per row. However, despite its length, there are only 9 or 10 rows of stigmata. The stigmata are long. In the contracted. pre- served zooids they usually are found folded hori- zontally along the centre of each row. The stigmata are fusiform, pointed at each end. They are only slightly reduced in length toward the dorsal and ventral mid-lines. The oesophagus increases in diameter anterior to the stomach to form a prestomach. The stom- ach is wide, reduced in diameter at the cardiac end, with some oblique folds terminating agamst the suture line. It has 25 to 30 fine folds, some- times interrupted along their length. The duode- nuin is Wide, and the short mid-intestine nurrows abruptly to enter the oval posterior stomach in the bend of the gut loop. Short rectal ogeca are present at the proximal end of the ascending leop. The rectum extends half way up the branchial sac (@pposite the fifth row of stigmata), but the con- traclion of the anterior part of the thorax can draw the anus up behind the atrial aperture. The anal opening is bounded by two large semicircular lips. The ovary is in the relatively short, stumpy posterior abdomen a short distance behind the eur loop, and 12 to 20 male follicles are clustered behind it. There is no constriction between the abdomen and the postenur abdomen Up to 2 embryos are in a brood pouch projecting from the postero-dorsal corner of the thorax in colonies collected in April and May from West- ern Australia (WAM 754.82 763.82 837.83 1030.83 87|-2.83), and from the Capricorn Group in November (QM GH5307) and Decem- ber (QM GH5559). Mature gonads are not pre- sent in colonies collected in August (WAM 1029.83). In the Darwin colonies, collected in October, up to 4 embryos are lined up in the atrial cavity, the largest a tailed larva. Specimens col- lected in October (QM GH5131) and November (Heron L.), May (Myora) and June (Lizard I.) have up to 3 embryos at vanous stages crowded into the atrial cayity. projecting dorsally from it when the thorax is contracted. These embryns distend the whole atrial cavity and are not in a postero- dorsal brood pouch as they are when only one embryo is present. Larvae are short and deep, with atrunkto Immlong. As they mature they become almost spherical. the tunk about [.Omm long with the tail wound about three-quarters of the way around it. Circles of 6 to 8 epidermal vesicles are arqund cach of the adhesive organs and these remain attached by narrow stalks to the epider- mus. Aboul 5 epidermal vesicles are also attached to the trunk ectoderm in the median line ventrally, behind the adhesive organs. The 3 median adhe- sive organs at the anterior end of the trunk are small and shallow. A mass of blue pigment is in the haemocoelic cavity, especially dense in the postcnor homs, although these are largely oblit- erated as the oozooid develops. Mature larvac with a spherical trunk have 4 rows of stigmata, and arelalively short yolk mass beneath the large, upright pharynx, with its vertical endostyle, REMARKS The branching cloacal canals of these colomes resemble some species of Polyclinwm, The masa- ics formed by the closely spaced colonies resem- ble those of Ritterelfa dispar, and some species of Diplesoma (see Katt 1980). Nevertheless, the zooids are characteristic of Aplidium, It is passi- ble that these lightly attached small colonies sub- divide, and move to space themselves evenly over the substrate. The zooid, sometimes with a brood pouch, and with bunched male follicles in a short posterior atxiomen and numerous stomach folds,resembles Aplidium ateute, However. the latter species has zooids with longer, more barrel-shaped stomachs, arranged in circular systems, and the colonies are largerand more irregular than those of the present species, Further, although larvae of A. wrewte have urcs of lateral vesicles, they also have median vesicles which do pol occur in the present species. Aplidiumn muliiplicanen has branching systems and simular larvae to lhe present species. How- ever, A, melriplicatute colonies are larger, softer, the pigmentation is different, thoraces are nar- 522 rower and systems longer and more crowded. Aplidium opacum has more crowded systems, spreading colonies, narrower thoraces, more rows of stigmata and fewer stomach folds. A. flavolineatum: Monniot (1987) from Ches- terfield Reef has 12 rows of stigmata and about 30 stomach folds. Although Monniot does not record the number of stigmata per row the zooids are similar to the present species. However the colonies are sheet-like rather than being cushions or wedge-shaped lobes, the zooids are arranged in circles rather than radiating double rows, em- bryos are incubated in a brood pouch and larvae have median ampullae (like A. uteute, see below). Although the larvae have some similarities, the present species is not conspecific with the South African A. flavolineatum (see Millar 1962) which is sandy (rather than naked) and red (rather than a cream, yellow, purple or brown colour). The most conspicuous feature of the present colonies, viz. the double-rows of zooids converg- ing to the central or terminal cloacal apertures, were not observed either by Sluiter (1909) or Millar (1963) in specimens they assigned to Aplidium altarium. However, the shape of the colonies, the form of the stomach, and the short posterior abdomen without a constriction be- tween it and the abdomen are similar in all de- scriptions. The large semicircular lips of the anal opening, and its unusual, relatively anterior, po- sition are recorded by Sluiter for the type speci- mens, which also have an expansion in the oesophagus as in the present specimens. The 4 to 6 embryos reported to be in the atrial cavity (Sluiter 1909, Taf. V9a) are very much smaller than those in the newly recorded specimen — indeed they are smaller than is known for any species of Aplidium and possibly Sluiter misin- terpreted these bodies. A further discrepancy be- tween Sluiter’s account of the type and the newly recorded specimens is that he recorded only 10 stigmata per row — however there often are discrepancies of this order in Sluiter’s counts of stigmata. Reexamination of specimens from Darwin (Kott 1966) shows that the number of stigmata per row and the shape of the colonies were re- corded incorrectly. The colonies are cushions, rather than sheet-like and the branchial sacs have the same number of stigmata as the present newly recorded ones. These Darwin colonies appear to be conspecific with the newly recorded speci- mens and with the Indonesian type material (Sluiter 1909). MEMOIRS OF THE QUEENSLAND MUSEUM Specimens assigned to A. altarium from Zanzibar and Mozambique (Michaelsen 1919; Millar 1956) appear different from the present species, with more numerous stomach folds (about 40), and small larvae with only 4 large epidermal vesicles and no ampullae (see Millar 1956). Aplidium amorphatum Kott, 1963 (Fig. 63) Aplidium amorphatum Kott, 1963, p.101; 1975, p.6. Aplidium pseudobesum Kott, 1963, p.101. DISTRIBUTION New REcoRDs: Victoria (Deal I., QM GH ; South Gabo I., QM GH5155-6). New South Wales (Jervis Bay, QM GH5617-9). PREVIOUSLY RECORDED: South Australia (Elliston Bay — SAM E2581 Kott 1975). Victoria (Bass Strait — AM U3918 Kott 1963). New South Wales (Eden — AM U3922 U3924 Kott 1963), DESCRIPTION EXTERNAL APPEARANCE: Colonies are soft, ge- latinous cushions, about 8cm in diameter, almost spherical or dome-shaped. The test is translucent and the zooids can be seen through it arranged in circular systems of 9 or 10 in each circle, although in these preserved colonies they are generally withdrawn from the surface and the systems are obscured. Specimens from South Gabo I. (QM GH5155-6) are described by the collector as ‘red rose’ ascidians. INTERNAL STRUCTURE: The zooids are large, the thorax and abdomen together being about 5mm long. In relaxed condition the thorax would be longer than the abdomen. The posterior abdo- men is long, but when contracted is drawn up to the left side of the gut loop. The zooids are mus- cular with about 20 longitudinal bands on the thorax that extend along the ventral border of the abdomen and posterior abdomen. The branchial aperture has 8 pointed to shallow rounded lobes around the rim of the short siphon. The long, robust, tongue-like atrial lip, from the upper bor- der of the opening, has a tridentate tip with each division of equal size. The pharynx has 15 rows of up to 16 long stigmata. Each of the rows is crossed by a parastigmatic vessel. An intermedi- ate dorsal languet associated with each of these parastigmatic vessels alternates with, and is the same size as, the languets on the primary trans- verse vessels. THE AUSTRALIAN ASCIDIACEA 3 The oesophagus is about one third of the length of the abdomen, The large voluminous stomach occupies the middle third. Tt has 4 shallow glan- dular ridges in its internal wall. The usual duode- nal and mid-intestinal regions and a pear-shaped posterior stomach are in the posterior third of the descending limb of the gut loop. A double series of testis follicles is in the posterior end of the av i PO iin a | FIG. 63, Aplidium amorphatum — a, colony outline (SAM E2581): b, thorax and abdomen (holotype AM U3918); e, branchial aperture (holotype AM U3918); d, outline of contracted zooid, showing posterior abdomen pulled up alongside the abdomen (QM GH5617); e, larva (SAM E2581). Scales: a, 2cm; b, Imm, c, e,.0.2mm, d, 2mm. posterior abdomen and the ovary is anterior te em. The colony from Elliston Bay collected in Feb- ruary has a single, large, embryo being incubated in the atrial cavity, almost completely occluding the pharynx, The larval trunk is 1.2mm long, and the tail is wound only halfway around it. Ectoder- mal vesicles are crowded in the test around the anterior part of the trunk, obscuring the 3 adhe- sive organs in the anterior mid-line. They consist of wide lateral bands of vesicles, and some in the median line attached by 2 or 3 long branching stalks in each imterspace between the adhesive organs. The developing adult organs are at the posterior end of the long trunk. REMARKS The distinctive aspects of this species are its soft, rounded colonies, large zooids in circular syslems, 8 branchial lobes, parastigmatic vessels, large stomach with shallow glandular ridges, and the large larva filling the atrial cavity in which it is brooded. The species has a superficial resem- blance to Aplidium protectans which also com- pletely lacks sand, and has a similar rounded shape, circular systems, 8 branchial lobes, large zooids, and shallow stomach folds. However, in A, protectans the test is firmer, the systems are depressed into the surface test, rows of stigmata are more numerous, parastigmatic vessels are ab- sent, and the range is tropical. The wide ventral muscle which draws the pos- terior abdomen up alongside the posterior end of the abdomen is reminiscent of the muscle band in Aplidium circumvalutum which distorts the zooid in a similar way (see Miller 1982), Aplidium australiense Kolt, 1963 (Fig, 64) Aplidium australiensis Kott, 1963, p.111 (part, type specimen from Shoreham). Aplidium colelloides: Kott 1972a, p.15; 1972b, p.176, 1975, p.7, DISTRIBUTION New Recorps: Western Australia (Recherche Archi- pelago, WAM 147,75), South Australia (Eyre Penin- sula, SAM 62557; Great Australian Bight, SAM E2478; Nuyts Archipelago, SAM E2472 E2476 E2558; Hotspot, QM GH1325). PREVIOUSLY RECORDED: South Australia (Great Aus- tralian Bight — Kort 1972b, 1975; St. Vincent Gulf — Kott 1972a; Investigator Strait — Kott 1972b), Victo- ria (Shoreham — AM Y 1398 holotype Kott 1963). 524 The records suggest that the species occurs only off the southem Australian coast. DESCRIPTION EXTERNAL APPEARANCE: The colonies are long (up to &em), more or less elliptical heads to 6cm diameter on long (to 20cm) narrow, tough, sandy stalks narrowing to a basal holdfast of short. thick root-like branches. The test on the head is soft, gelatinous, translucent. The terminal end of the head sometimes is pointed. In the larger preserved colonies there are 4 to 6 deep, longitudinal furrows or creases formed be- tween deep folds of the surface of the colony. Common cloacal canals are at mid-thoracic level and extend into large cloacal spaces in the edge of the folds referred to above. Three or 4 large cloacal apertures are either sessile or on protuber- ant cones along the edge of the folds. Sometimes they seem to alternate from side io side of the edge of the fold. Branchial openings of zooids are on both sides of these folds, i.e. on their outer and inner surfaces, as well as in the base of the depres- sions between and under the folds. Branchial apertures are absent only from the edges of the folds over the large cloacal canals, Zooids are seen to be in longitudinal rows near the top of the stalk. However, in preserved colo- nies the arrangement of zooids towards the mid- dle and top of the head 1s obscure. Relatively small heads (2cm long) on long (10cm) stalks (SAM E2558) suggest that heads may regenerate on persisting stalks, These smaller heads lack the deep creases of the larger ones, and the zooids are arranged in more or less conspicuous double rows, converging to large cloacal apertures — 3 or 4 randomly distributed around the sides of the head, and 2 or 3 on the top of the head, INTERNAL STRUCTURE: Zooids are particularly long and thread-like, their posterior ends curving down into the stalk, The rim of the branchial apertures is divided into 6 large, rounded lobes. The atrial aperture is a small sessile opening in the middle of the dorsal border of the thorax with a small pointed languet from the upper rim of the opening, About 10 longitudinal muscle bands on the thorax extend the length of the zooid. joining into a single wide ventral band along the long postenor abdomen, Stigmata are in 15 rows with about 15 per row, although both the number of rows and the number MEMOIRS OF THE QUEENSLAND MUSEUM per row are difficult to count owing to contraction of the small zooids. Twelve to 16 deep, longitu- dinal folds are in the stomach wall. The number vanes with the size of the zooids. Testis follicles are in a double series in the posterior end of the posterior abdomen. One large embryo or tailed larya and sometimes also what appears to be an infertile ovum are in the posterior part of the atrial cavity in colonies collected in September (SAM E2557) and tn Oc- tober (Kott 1972b). Specimens collected in March (SAM E2476) have mature male gonads. The laryal trunk is 1.5mm long. Its anterior end is obscured by a cloud of minute epidermal ves- icles along each side of the base of the adbesive organs. The tail is wound three-quarters of the way. around the trunk. REMARKS The colonies of this species, with their long, narrow, hard, sandy stalks and long gelatinous heads with small, crowded zooids and deep lon- giluclinal creases, are distinctive. Despite Herdman’s (1886) speculation that Aplidium colelloides (Herdman, 1886) from South Africa had a short stalk, Millar (1962) established that it has a long (up to 6cm), narrow, hard stalk like that of the present species and reminiscent also of the long-stalks of species of Sycozoa and Pseudodistoma. Kott (1972a,b; 1975) in assigning South Australian specimens to the South African species, ignored the faci that the former have larger colonies with external creases on the head, more numerous stomach folds, and a larval trunk twice the size of the South African specimens. The large colonies that char- acterise the Australian species also have more crowded zooids less conspicuously arranged in long, oval systems (Millar 1962). The larger colonies, deep creases, complex crowded systems as well as the long, narrow, hard stalks help to distinguish the species from A. geminatum n.sp. which has double rows of za- oids, but short and thick stalks, less crowded zocids, more numerous stomach folds, longer, more conspicuous and often 3-lobed atrial tongues, and different larvae, The type specimen of the present species from Shorcham (AM Y¥1398) is a head which has lost its stalk, The other specimens assigned to this species by Kott (1963) are colonies of FIG. 64, Aplidiian australiense — a, Whole colony (QM -GH1325), b, head of large colony (SAM E2478): ¢, thorax and abdomen with embryos being brooded in posterior end of atrial cavity (SAM E2557), d, larva(SAM E2557). Scales: a, b, 2cm; c, 0.5mm; d, 0.2mm. THE AUSTRALIAN ASCIDIACEA 3 525 526 MEMOIRS OF THE QUEENSLAND MUSEUM A. geminatum n.sp., to which the larva figured by Kott (1963, Fig. 25) also belongs. Aplidium bacculum n.sp. (Fig. 65. Plate 10e) DISTRIBUTION Type LOCALITY: South Australia (Yorke Peninsula, Edithburgh, 35°9.7S 137°47.4E, 12m, coll. AIMS Bioactivity Group 6.2.89, holotype QM GH 5434). FURTHER RECORD: South Australia (Thorny Passage, SAM E2568). DESCRIPTION EXTERNAL APPEARANCE: The colony is a com- pact mass of parallel, long (3cm), narrow (to 5mm) sandy stalks, that branch once or twice along their length and adhere to one another. Their terminal free ends are flattened, horizontal, and level with one another to form a more or less even platform. They have a cloacal aperture in the centre of each terminal flat tip. This opens from a shallow, circular, cloacal cavity which is sur- rounded by about 10 zooids. Branchial apertures open around the upper margin of the terminal flat surface, and atrial apertures open into the cloacal cavity. A single cloacal system is in each terminal branch. The specimens from Thorny Passage have the terminal test inflated over each zooid, and the central common cloacal apertures pro- trude slightly. Sand is crowded over the surface of each stalk, and also is embedded throughout the internal test, but is absent from the surface over the terminal end of each lobe. The small thread-like zooids are difficult to remove from their narrow longitudinal spaces in the hard, sandy colony. INTERNAL STRUCTURE: Zooids extend the whole length of the sandy stalk, but the thorax and abdomen are together only about 3mm of which the abdomen is about two-thirds. The branchial aperture has 6 rounded lobes. A fleshy atrial lip, divided into 3 pointed lobes, extends from the upper border of the atrial aperture. Longitudinal muscles on the thorax are moderately strong, but they become finer as they extend back along the ventral surface of abdomen and posterior abdo- men in a wide band of delicate muscles. Stigmata are in 5 rows of about 6 stigmata per row, but these were difficult to count owing to contraction. The oesophagus is relatively long, FIG. 65. Aplidium bacculum n.sp. (holotype QM GH5434) — a, colony; b, thorax and abdomen, Scales: a, lcm; b, 0.5mm. THE AUSTRALIAN ASCIDIACEA 3 the small stomach being two-thirds of the way down the abdomen. The stomach wall has 5 deep folds. The duodenal area reduces in diameter to a short mid-intestine which opens into an oval pos- terior stomach in the descending limb of the gut loop. There is a small rectal valve at the proximal end of the rectum. Gonads are not developed in the available material. REMARKS Superficially, the sandy stalks of this species differ from those of Ritterella spp. and Monniotus spp. (which have no sand internally) principally in having embedded sand crowded in the internal test. Despite the long oesophagus, however, zo- oids are characteristic of Aplidium and open into a well developed cloacal cavity rather than di- rectly to the exterior as in Ritterella. The form of the colony, and the long oesophagus (which prob- ably is associated with the length of the colony lobes), together with the small zooids with only 5 rows of stigmata, characterise the species. Exter- nally, the sandy lobes of the colony, especially the one from Thorny Passage with the test inflated over the anterior end of each zooid, and the pro- truding central cloacal aperture, resemble Synoicum stewartense Michaelsen, 1924 (see Millar 1982). Other Aplidium species with 5 rows of stigmata viz. A. filiforme n.sp. and A. depressum lack the long stalks and long oesophagus of the present species. Aplidium brevilarvacium Kott, 1963 (Fig. 66. Plate 10f,g) Aplidium brevilarvacium Kott, 1963, p.113. Aplidium digitatum Kott, 1975, p.7. DISTRIBUTION New RecorDs: South Australia (Ward I, QM GH2412; Top Gallant I., QM GH953; Investigator Group, QM GH1311 GH2413). Queensland (Heron I., QM GH961). PREVIOUSLY RECORDED: Western Australia (Pt. Peron; Cape Naturaliste — AM U3926 syntypes A. brevilarvacium Kott, 1963). South Australia (Northern Great Australian Bight -QM G7508 AM Y1982 pa- ratypes A. digitatum Kott, 1975). The record from Heron I. is anomalous (see Remarks, below). DESCRIPTION EXTERNAL APPEARANCE: The colonies are rounded heads (0.5 to 1.5cm in diameter) on thick 527 stalks, 0.5 to 6.0cm long, branching 2 or 3 times along their length, and up to about 0.5cm in diameter at the base. These sometimes arise from a basal test mass. Zooids are arranged in longitu- dinal rows along each side of cloacal canals that extend the length of the head, converging to 2 or 3 large, terminal cloacal apertures. The outer sur- face of both head and stalk is even, without wrin- Kles or protuberances. There is a layer of sand in the outer surface that may obscure the zooids and their branchial open- ings. Sand is present also in the internal test. The anterior ends of the zooids are perpendicular to the outer surface and posteriorly they curve down into the stalk of the colony, criss-crossing one another as they go. In life the colonies are white to a sandy pink colour. INTERNAL STRUCTURE: Zooids are robust. When contracted the thorax and abdomen are of equal length and together about 2mm long. About 12 very fine longitudinal muscles from the thorax run the length of the zooid. The branchial lobes are rounded. The atrial opening is sessile, with a fleshy, pointed lip from the anterior rim of the opening which sometimes is divided into 2 at the tip. The pointed dorsal languets are to the left of the mid-dorsal line, opposite the third row of stigmata. Stigmata are relatively short and oval, about 10 per row, and in 14 rows. The stomach, halfway down the abdomen has 10 deep longitu- dinal folds. The duodenum is long, and the oval posterior stomach is separated from the duode- num and rectum respectively by short lengths of proximal and distal mid-intestine. The posterior abdomen is narrow, up to lcm long. In the type specimens from Cape Naturaliste (collected in January) and the newly recorded South Australian specimens (collected in April) there is a large tailed embryo in the atrial cavity and sometimes a large (possibly non-fertile) ovum as well. The larval trunk is about 0.9mm deep, but only about 0.6mm long. It has an ocellus and an otolith, 3 medium stalked adhesive organs, and the tail is wound one and a half times around it. Epidermal vesicles are present along each side of the adhesive organs. REMARKS The colonies from Heron I. appear to have only one or 2 terminal cloacal apertures, instead of the more numerous openings in the South Australian colonies. However the zooids have the same num- ber of rows of stigmata and stomach folds, and are not distinguishable from the southern or west- ern Australian ones. These Heron I. colonies may 528 represent a population at the northern extremity of the range of this otherwise temperate indige- nous species, Exterally, the colonies of this species, and particularly the specimens from Heron I. with one or 2 systems per lobe resemble Distaplia austral- iensis (see Kott 1990). Colonies also resemble Aplidium geminatum n.sp. which, however, has more numerous (20), fine stomach folds, appre- ciably larger colonies, and less branched and more wrinkled stalks. Aplidium caelestis Monniot, 1987 (Fig. 67. Plate 1 la—f) Aplidium caelestis Monniot, 1987, p.517. Synoicum hypurgon: Kott, 1963, p.86, ?Aplidium alf. lobatum: Tokioka, 1967, p.22. MEMOIRS OF THE QUEENSLAND MUSEUM DISTRIBUTION New REcorbs: Western Australia (Shark Bay, WAM 883.83; Mullaloo Beach, QM GH5450; Hillary's Boat Harbour, QM GH5452), South Australia (Edithburgh, QM GHS5437; Beachport, SAM E2566 E2569; Kanga- roo 1, QM GH5428 GH5430). Victoria (Bass Strait QM GH5648). NSW (Norfolk I., QM GH5783). Queensland (Capricorn Group, QM G9474 GH5594 GH5633-40 GH5647-8 GH5650-9 GH5S661-2 GH5665; Lizard I., QM GH5631-2). PREVIOUSLY RECORDED; Western Australia (Rottnest — AM Y1382 Kott 1963). Queensland (Heron I. — AM U3991 Kott 1963). New Caledonia (Monniot, 1987), ? Marianas (Tokioka 1967). DESCRIPTION EXTERNAL APPEARANCE: Colonies form ro- bust, always sandy, investing shects up to 20cm FIG. 66. Aplidium brevilarvacium — a, b, parts of colonies (QM GH953, G7508); ¢, details of a zooid system, with double rows of zooids converging Lo a lerminal cloacal cavity (QM GH961); d, thorax (QM GH2412); e, thorax and abdomen (QM GH953); f, larva, showing arcs of lateral vesicles and median ampullae (syntypes AM U3926), Scales: a, b, 2cm; c, 5mm; d, e, 0.5mm; f, 0.2mm. THE AUSTRALIAN ASCIDIACEA 3 in maximum extent and up to Icm thick. A row of zooids is along each side of the cloacal canals that extend between or around long, narrow areas of zooid-free test, Large sessile common cloacal apertures are at the junction of several canals and they often are elevated by the large cavity beneath them. The surface of the colony is depressed where relatively thin, usually transparent, naked test overlies the cloacal canals. Oval granular bodies about 0.1mm long, and some shade of brown to almost black are in the test and line the sides, base and walls of the cloacal canals, and 529 often are in the surface as well. The grey to brown colour of the colonies results from these bodies and the included sand, sometimes in small clumps. Both the oval bodies and the clumps of sand appear to have been interpreted as faecal pellets. Sand is embedded throughout the test except in the thin transparent surface layer over the cloacal canals. Colonies have been described as ‘sandy grey’ or ‘drab, with fawn zooids’, ‘sandy grey with yellow zooids’, ‘grey encrusting ascidian’, Living zooids are yellow, or orange. The test over the cloacal canals is irridescent in a. aii rt | «(ill y oanaepiit ih wil MIN, FIG. 67. Aplidium caelestis — a, colony (QM GH5437); b, thorax with embryo being incubated (QM GHS150); ¢, thorax and abdamen (QM GH5647); d, whole zooid (QM GH5633); e, larva (QM GH5647). Scales: a, 5mm; b,c, 0.5mm; d, 1mm; e, 0.1mm. 5H) living specimens, A enlony from Heron 1 (QM GH5661) had blue cells in the surface test be- tween the rows of zooids of the living colony — as in the type from New Caledonia. Prochloron is often present on the surface (see Kott er al., 1984; Aplidiiunsp, 1). Two colonies haye red and green symbiotic cells embedded amongst the sand in the floor of the common cloacal canals. These are pink in preservative. Sand always is present in the basal half of the colony, and in the ridges between the double rows of zooids, INTERNAL STRUCTURE: The zoids are Jong and thin (thorax and abdomen together to Smm), but robust, with strong longitudinal muscles that ex- tend their full length, along the ventrum of abdo- men and posterior abdomen, The thorax and abdomen, of about equal length, are together about 3mm, The branchial siphon has 6 rounded lobes and a single conspicuous bulging sphincter muscle at their base. A long prebranchial area is posterior to the tentacles. The atnal aperture, a short distance down the dorsal surface, also has a conspicuous sphincter muscle. A robust, but not very long, pointed atrial lip, usually not divided, arises from the body wall anterior to the aperture. The atrial aperture at the level of the second or third row of stigmata opens into the sides or base of the cloacal canal. Stigmata are in 10 to 18 rows, with 6 to 14 the maximum number per row. Expanses of unperfo- rated pharynx are along each side of the mid-dar- sal and mud-ventral lines. The cesophagus and post-pyloric part of the gut loop are relatively long for this genus, the gut loop being about the same length as the thorax when both are extended. The small stomach, halfway down the abdomen, has 5 distinct folds, although these sometimes flatten and occasionally it is difficult to see traces of any folds tn the stomach wall. The duodenum, mid-intestine and posterior stomach are charac- tenstic of the genus. Gonads are in the centre of the abdomen and consist of an itregular row of testis follicles with an antenor ovary a little dis- tance behind the gut Loop. A single embryo with a trunk 0.65mm long and median ampullae alternating with the adhesive organs is in the atrial cavity of spectmens taken in March and August from Heron I. REMARKS The zooids of this species, with their well- formed branchial siphons. sphincter muscles, atrial lips separate from the apertures, and long and narrow thoraces, resemble A. crateriferum, A. griseum o.sp., A. lenticulin nsp.. A. Janacra- MEMOIRS OF THE QUEENSLAND MUSEUM tum nom. nov. and A. clivesun osp. The long, rather straight double row systems with long ca- nals surrounding ridges of solid sandy test, some- times can be seen in A. ritteri, but its atrial lip is not separate from the aperture. The oval granular bodies are present also in A, /enticulurm n-sp., which has cloacal canals around raised polygonal areas rather than between the long ridges of the present species. The larvae also appear to be similar to those of A. lenticulum n.sp. The type specimen of A. caelestis Monniot, 1987 from New Caledonia has the same almost smooth stomach, narrow thorax, separate atrial lip, long double rows of zooids. The test is entirely en- crusted with sand as it usually is in the newly recorded specimens. Although the transparent test over the cloacal canals is not always sky-blue in living specimens as it is in the type, it some- times is. Colour and the number of rows of stig- mata appear to be variable characters in this widespread species. Kott (1963), interpreting the oval bodies as faecal pellets, assigned specimens of this species to A. kypurgon (Michaelsen, 1924), a New Zea- land temperate species. Aplidium clivosum n.sp, (Fig. 68. Plate 12u-f) Apliditn pantherinuin; Kott, 1963, p. 98; 1972b, p. 176. DISTRIBUTION Tyre Locauiry; South Australia (S.E. Beachport, 6— 7m, on jetty piles, coll. W. Zeidler 20,2.89, holotype SAM E2561; Kangaroo I,, on jetty piles, coll. N. Cole- man 12.3.78 AMPI 220, paratype QM G1197). FURTHER RECORDS: Western Australia (Port Hedland, WAM 1020-2.83; Montebello Is, QM GHS406; Cock- burn Sound WAM 869-70.83, OM GH2149; Bussel- ton, QM GH5469; Green Pools, QM GH5629). South Australia (Topgallant J,, QM GH1287 QM GH2419, Hotspot, QM GH2417; Nuyts Archipelago, SAM 22562; Eyre Peninsular, SAM E2566 E2569; Flinders 1., QM GH2305; Kangaroo 1, QM G11997 GH5432 GH5433, SAM E2564; Ward 1,, Beachport, QM GHS447; West L, OM GH2415), New South Wales (Jervis Bay AMPI 78 QM GHS5628). Queensland (Heron L QM GH5569 GH5593 GH5597 GH3669-70 GHS5675); Westem Australia (Rottnest to Hamelin Bay — Katt 1963), South Australia (Elliston Bay — SAM E2585 Kort 1972b). THE AUSTRALIAN ASCIDIACEA 3 DESCRIPTION EXTERNAL APPEARANCE; Colonies are large, to 4cm thick and to 15cm or more in maximum dimension. Some are of even thickness attached by the whole of the basal surface. Others con- verge toward the basal attachment area, or have irregular processes that may project into the sub- strate or around rocks and rubble. One colony (WAM 1021,83) is upright. rather flattened and divided into 2 lobes, about 15cm high, with sy¥s- tems evenly distributed over the both sides of the upright lobes. The base and sides of the colonies have sand adhering to the surface. Sand usually is present on the zooid-free raised areas of the surface sur- rounding single systems, or groups of systems. The test over the systems usually 1s naked, Em- bedded sand is absent at thoracic level where #ooids curve around the shallow cloacal cavities, butis present throughout the remainder of the test. In some colonies it can be seen projecting in from the periphery of each system as narrow wedges between the double rows of zooids, The cloacal canals converge from the periphery of each system, to the large central cloacal cavity, A cloacal aperture raised on a conical proluber- ance (up to 5mm diameter across the base) is in the centre of each system. Zooids are arranged in double rows, one row along each side of the canals, In preserved colonies the surface is de- pressed over the systems, although it is not un- usual for 3 or 4 systems to be present in one large depression, each system with its protruding com- mon cloacal aperture. Systems are not always circular, often being irregularly shaped, the cloa- cal aperture being excentne with longer canals on one side than on the other, The diameter of the systems tn this species ts variable, ranging trom 0.5 to lem, often in one colony. In some colonies the whole upper surface is horizontal, and depressed within a rounded mar- ginal rim. In others there is no sharp division between the margin and a convex upper surface. In photographs, living colonies are pink, “light wine’ or bluish red, In preservative they are a translucent bluish-red, The colour is conspicuous on the upper surface over the systems and araund ihe cloacal apertures. INTERNAL STRUCTURE: In contracted zooids the thorax and abdomen together are Irom 3 to 6mm long, Long. thread-like posterior abdomina extend down through the crowded sand in the lower half of the colony. About 12 fine longitidinal thoracic muscles extend posteriorly along the whale Jength of the zooids. The branchial siphon is long, with a nar- row, bulging sphincter muscle behind the bran- chtal lobes. Some fine circular muscles are on the remainder of the siphon, but they do not form a sphincter. The branchial tentacles are about hulf- way up the branchial siphon, with a long pre- pharyngeal area between them and the prepharyngeal groove. The diameter of the siphon bulges out in the vicinity of the branchial tentacles between the anterior sphincter and the pre- pharyngeal groove. A single, pointed atrial lip, sometimes bifid or trifid at the tip, arises from the body wall just anterior to the atrial siphon, which, depending on the state of contraction of its sphincter muscle, is either a sessile, circular o ing, of is on. a short, cylindrical siphon. The atrial siphon is well anterior, opposite the second row of stigmata. In the branchial sac are 16 to 18 rows of up to 12 stigmata. A wide ynperlorated band of phanyn- geal wall is present each side of the endostyle. The abdomen ts almost as long as the thorax. The oesophagus is long and narrow, The small stomach has 5 deep folds in its wall. A long duodenum is continuous with a narrow proximal part of the mid-intestine. A small oval posterior stomach is in the distal part of the descending limb of the gut loop. A short, narrow distal section of the mid-intestine is between the posterior stomach and rectum in the bend of the eut loop. A distinct reetal valve is present. The anal open- ing is Opposile the fourth most pastemor row of stigmata. A single of double series of male follicles is in the posterior part of the posterior abdomen. A large embrya (trunk 1.5 to 2.5mm long) isin the posterior half of the atrial cavity of specimens collected in March at Busselton (QM GH5469), and January at Kangaroo L. (QM GH5432). The tail extends only to the antenor end of the trink which has crowded hair-like ampullae obscuring the antenor end. Embryos and larvae are heavily pigmented {red and opaque). REMARKS The species is distinguished by 2 characters which although they occur separately in other species, seldom are found together as they are in the present one — the atrial lip is chose te, byt separated from the aperture, and the double rows of zovids converge ta the cloacal apertire, Ttalso has a group of characters which it shares with A. crateriferue and the temperate and partly sy mpa- ic A. funceraiwin nom. woy., viz. the bulky sandy colonies, protuberant cloacal apertures, narrow MEMOIRS OF THE QUEENSLAND MUSEUM THE AUSTRALIAN ASCIDIACEA 3 zooids with 5 well-formed stomach folds, and the airial lip separate from the aperture. The zooid systems are not always so deeply depressed into the surface as they are in A, crateriferum, and A. lenaeratum nom. nov, has sand in the test over the cloacal cavity and in the protuberant cone of the cloacal aperture. Both the latter species have their zooids arranged in a single circle around each common cloaea. In A, crateriferum the atrial lip is not close to the aper- ture as it 1s in the present species and A. lumacra- tum does not have the same bulging branchial sphincter and long siphon, In A. limacratum the thoracic muscle bands, and in A, crateriferum the number of rows of stigmata are more numerous. The larvae of A. clivosum are in the same size range as those of A. lunacranem but larger than those of A. crateriferum, Larvae of the present species differ from A, /wracratum in haying fewer epidermal yesicles and more ampullae, Aplidium pantherinum (Sluiter, 1898) and its synonyms from South Africa (see Millar 1962) have a smaller larval trunk, and although they have narrow thoraces and about 17 rows of stig- muta like the present species, the atrial tongue is nol separate from the aperture. Aplidium clivosum has possibly more variation in iis external appearance than most other species of Aplidium, The size of the systems varies, as does the number per colony, the amount of em- bedded sand, the colour and the depth of surface depressions. These variations are not geographi- cally based however and the whole range of mor- phological variations occurs in tropical as well as temperate waters. In view of its wide range in Austrahan waters, it is probable that it will be found to eccur in the Westem Pacific. It is possible that some of the specimens assigned to A. craferiferuwm by Van Name (1918) from the Philippines are of this species. Aplidium congregatum n.sp- (Fig. 69) DISTRIBUTION Tyre Locaurry: Queensland (North Suradbroke L., Point Lookout, coll. A, Rozefelds 17.11.78, holotype OM GHS5584), FURTHER RecorDs: None, Specimens are found tucked into narrow rocky crevices at the type locality. DESCRIPTION EXTERNAL APPEARANCE: Sandy flat-topped lobes 1.5 to 2cm diameter around the upper sur- face, narrow toward the base where they are joined through common test. The edges of the flat upper surface fuse with adjacent lobes. A layer of sand adheres to the outside of the stalks and sides of the lobes, but is absent from the upper surface. Internally sand grains are scattered sparsely amongst the zooids. Zooids are crowded and par- allel in the test. They may be in rows along euch side of Jong canals that converge to the sessile cloacal apertures on the flat upper surface, INTERNAL STRUCTURE: Zooids are long and narrow, up to Smm long, of which the thorax and abdomen together comprise about half, and the thorax is about twice the fength of the smill abdomen, About 16 fine longitudinal muscles extend separately along the length of the zonid and are not gathered into bands on either side of the abdomen or the posterior abdomen, A distinet sphincter muscle is at the hase of the 6 narrow, pointed lobes fringing the branchial aperture, The atrial aperture is at the anterior end of the dorsal surface with the forked atrial tongue extending from the anterior nm of the opening. Fine muscle fibres extend along the length of the atrial lip. The stigmata are in 11 rows of up to 12. The oesoph- agus is moderately long and narrow, The stomach is small and narrow with 12 longitudinal folds in its wall. There is the usual narrow mid-intestine, interrupted at ihe end of the descending limb of the gut loop by an oval posterior stomach, Small rectal caecaare at the proximal end of the rectum. A large (4 or 5 oocytes) ovary is at the proximal end of the posterior abdomen and the testis folli- cles in a single or paired series are behind it in the long, narrow posterior abdomen. The holotype has up to 6 embryos being incubated in a devel- opmental series lined up im the atrial cavily, lend- ing to stretch the thorax longitudinally. ‘Tailed larvae have 2 trunk 0.5mm Jong with the tail wound three quarters of the way around it. Coni- cal median ampullae (4) alternate with the mechan adhesive organs. An arc of scattered epidermal vesicles (about 3 or 4 deep) is around the Jateral line on each side of the adhesive organs and extending pasterior to them along the dorsal und ventral berder of the larval trunk. An ocellus and an otolith are present. FIG. 68. Aplidium clivasum n.sp.—a, colony (QM GH5669); b, thorax and abdomen (QM GH5670); c, branchial aperture, atrial aperture and atrial lip showing sphincter muscles (SAM E2564); d, larva (QM GH5469). Scales: a, tem; b, 0.5mm; e, 1.0mm; d,,0.4mm, 534 REMARKS The species is found in similar habitats and in the same geographic location as other flat-topped sandy, lobed species, all superficially resembling one another, viz. Aplidium incubatum n.sp., A. directum and Botryllus stewartense (see Kott 1990). The flat topped lobes of A, congregatum have more complex systems of crowded zooids than the others; Aplidium directum is distin- guished by its tougher colonies with distinct cir- cular systems, only 5 stomach folds and an atrial lip separate from the opening; Aplidium incu- baium nsp. tends to divide into separate lobes MEMOIRS OF THE QUEENSLAND MUSEUM each with a single system, its zooids have 25 stomach folds, and the larvae have numerous vesicles but no ampullae. The larvae of the present species resemble those of A. opacum which has investing, naked colo- nies; and A. geminatum n.sp. which has stalked heads with long cloacal canals converging to terminal cloacal apertures. Zooids of the present species share some char- acters with those of Aplidium fluorescum n.sp., but both larvae and colonies are different. FIG. 69. Aplidium congregatum n.sp. (holotype QM GH5584) — a, b, colonies with positions of zooids shown in part of a colony lobe; e, thorax, abdomen, part of posterior abdomen and embryos in atrial cavity; d, larva, Scales; a, b, lem; ¢, 0.5mm; d, 0.1mm, THE AUSTRALIAN ASCIDIACEA 3 Aplidium coniferum Kott, 1963 (Fig. 70) Aplidium caniferum Kott, 1963, p.102. DISTRIBUTION New RecorbDs: South Australia (Elliston, SA E2580). Tasmania (off Devonport, AM E309; Huon Channel, TM 0D2074). Victoria (Bass Strait, QM G11861 GH5605, MV H462; Cape Everard, AM B6231-2; Gabo [., SAM E2592). PREVIOUSLY RECORDED: New South Wales (Mon- tague North, Twofold Bay, AM U3921 Kott 1963). The species is known from depths to 400m (AM E6231-2). DESCRIPTION EXTERNAL APPEARANCE: The colonies are up- right spherical to egg-shaped lobes, largely naked and without sand in the thin layer of smooth surface test. A basal hard sandy holdfast is prob- ably rooted in sand. Sand is present in moderate concentrations internally. Zooids appear crowded in circular, oval to elongate systems, as well as some in double rows. However, the test is hard, whitish and almost opaque in preservative and it is difficult to determine the exact arrangement of the zooids. The shallow common cloacal cavities are relatively limited. They are surrounded by the fine, thread-like zooids, which are perpendicular at the surface, but cross one another posteriorly in the centre of the colony. INTERNAL STRUCTURE: Zooids are narrow, and long. The branchial siphon is short, with 6 rounded lobes, but the prebranchial area is rela- tively long. The thorax and abdomen are each about 2mm even when contracted, and the poste- rior abdomen is many times that length. The atrial lip is small, pointed and arises from the body wall opposite the first row of stigmata. The atrial ap- erture is a small round opening, well removed from the lip, opposite the fifth or sixth row of stigmata. There 1s a pointed papilla posterior to the atrial aperture, About 2 Jongitudinal muscle bands are on the thorax and continue in wide bands along each side of abdomen and posterior abdomen. Stigmata are in 22 rows of 8 per row. The oesophagus also is long, and wrinkled when contracted. The stomach wall has 5 deep folds. There is a long duodenum and mid-intestine. and a small oval posterior stomach in the descending limb of the gut loop. Rectal caeca are not well developed. A series of male follicles is in the posterior half of the posterior abdomen, and the small cluster of 535 oocytes is just anterior to the male follicles. A large embryo sometimes is found being incubated in the atrial cavity (QM G11861), The larval trunk is 0.65mm long. It has lateral ampullae on each side near the base of the 4 median ampullae, the r, } ad LARS FIG, 70. Aplidium coniferum — a. colony (QM GH797); b, thorax and abdomen (MV 1462); e, larva (QM G11861). Scales: a, 2em; b, 0.5Smm;.c, 0.1mm, 536 MEMOIRS OF THE QUEENSLAND MUSEUM latter alternating with the adhesive organs, The tail is wound about two-thirds of the distance around the trunk. REMARKS The large colony with embedded rather than surface sand and basal sandy holdfast, the long and narrow oesophagus, small and isolated atrial lip, and long thorax, are distinctive. The stomach folds are well formed, confirming that it is a species of Aplidium rather than Synoicum, which it resembles in having a dorsal papilla. The crowded systems distinguish the species from others such as A, crateriferum which, also has a Jong, narrow thorax, and separate atrial lip. A colony from Bass Strait (AM E309) charac- teristic of this species, with sandy basal stalk. naked, upright, ovoid colony, and long narrow posterior abdomina criss-crossing internally, originally was assigned to A. jacksoni by Katt 1963. Despite her interpretation that the zooids were similar to those of A. jacksoni, this was not confirmed on reexamination of the specimen. The zooids are juvenile vegetative ones and neither stomach nor branchial sacs are developed. Aplidium crateriferum (Sluiter, 1909) (Fig. 71, Plate [3a—c) Amaroucium crateriferum Sluiter, 1909, p,103, Not Van Name 1918, p.163. Not Tokioka, 1967, p.40. Aplidium crateriferum: Millar, 1975, p.242, DISTRIBUTION NEW RECORDS: Western Australia (Flinders Shoal near Onslow, QM GH5403; Exmouth, QM GH5398). Queensland (Capricorn Group, QM GHS5158-9 GH5567; central section Great Barrier Reef, QM GH5422), PREVIOUSLY RECORDED: Philippines (Sluiter 1909, Van Name 1918, Millar 1975), DESCRIPTION EXTERNAL APPEARANCE: The colonies are bulky, gelatinous and translucent but firm. The surface is smooth, and raised in high rounded ridges around irregular surface depressions, One to 3 large conical elevations with a terminal com- mon cloacal aperture arise from each depression. Single apertures are sometimes more or less cen- tral but often at one or the other end. A shallow but extensive cloacal cavity is beneath the rela- FIG. 71. Aplidium crateriferum — a, colany (QM GH5158); b, zooid (Heron I.), Scales: a, 2cm; b, 0.5mm. THE AUSTRALIAN ASCIDIACEA 3 tively thin surface test in the floor of each depres- sion. Zooids surround each cloacal cavity, their branchial apertures at the base of the surface ridges, and protected by them. The atrial apertures are directed Isterally to open into the cloacal cavily, In some colonies a little sand is in the surface test around the apertures, but otherwise embed- ded sand is nol present, and the surface is naked. In other colonics, sand is scattered throughout the test, amongst the zooids as well as in the zooid- Iree areas, and is on the external surface of ihe high ndges, although the surface of cach de- pressed area is naked (QM GH5403), Living colonies are said to be grey with red zooids, although photographs of specimens fol- lowing their removal from the substrate ate brown and yellow, In preservative the zooids, test and preservative are bluish-pink, INTERNAL STRUCTURE: The zooids are long and thread-like, with the thorax (to 5mm) twice the length of the abdomen. The posterior abdomen is many times the length of the rest of the zooid. A long, conspicuous branchial siphon has a narrow bulging sphincter muscle at the top, near the base of the branchial lobes. This sphincter appears to have a blood sinus associated with it, and it is invanably pink and translucent. There is a long pre-branchial area between the branchial tentacles and prepharyngeal groove and this effectively increases the length of the bran- chial siphon, The branchial tentacles cause a bulge about halfway down the siphon. Imperfo- rate areas are anterior to the stigmata, and along each side of the endostyle. The atrial lip is usually a Single pointed but quite fleshy lobe fram the body wall anterior to the opening. The atrial ap- erture also has a sphincter muscle, About 15 fine longitudinal muscles are on the thorax and extend the whole length of the zooid_ Fine transverse muscles are in the transyerse vessels, but not in the parietal body wall. There are 18 to 21 short rows of up to 14 oval stigmata on the left and up to 10 on the night side of the branchial sac. The maximum number of stigmata (8 to 14) is in the fifth to eighth rows. Up to 4 more stigmata are on the feft than the nght. On the left the triangular dorsal languets are opposite the third to fourh stigmata i.e. almost halfway across the pharyn- geal wall in the smaller, narrower zooids. The short gut loop has the usual parts. with a slightly longer oesophagus than usual, and a rela- lively small stomach, with 5 distinct, deep folds in its wall. The posterior stomach alsn is shot. Two conspicuous rectal caeca form a valve at the proximal end of the rectum. The posterior abdo- men is long and thin, Two irregular longitudinal series of testis follicles are about halfway down it. Single large embryos are in the thoraces of zooids collected in November, but not in May. The larvae are amongst the largest known for Aplidium, with the trunk 2mm long. The tail winds about three-quarters of the way around if. The long slender stalks of the 3 median adhesive organs aré obscured by crowded epidermal ves- icles which develop from the antenor part of the larval trank. REMARKS The large, firm gelatinous colonies with surface depressions over the systems are one of the most conspicuous aspects of this species. The colonies are reminiscent of the colomes of Lissoclimwn patella (see Kott 1980), Synoicum castellatiun (see above) and Aplidium clivosmn. The sandy species Aplidiunt caelestis, with z0- oid openings protected by sand-filled swellings on the surface test, has similar narrow zooids with their atrial lip separate from the opening but they do not have the conspicuous surface depressions and circular systems of the present species. The mostly temperate Aplidium clivasuin also has similar zooids which open into deep surface depressions, However, it has Zooids in double rows converging to the protruding common eloa- cal apertures, and fewer (14 to 18) rows of stig- mata. Sluiter's type specimen, from the Philippines, 4 large, fleshy mass, maximum dimension 10cm and up to 1.Sem thick with characteristic surface depressions and projecting cloacal apertures, closely resembles the present colonies. Millar (1975) describes similar colonies from this loca- tion, Other colonies from the type locality (Van Name 1918) are regular with deep folds, bul only shallow depressions, large systems with ir- regularly distributed zooids opening onto the flat- tened upper surface of the colony, and the surface thickly encrusted with foreign particles. These do not seem to be conspecific with the present spe- cies and are more like specimens of Aplidinin caelestis (see below), which has similar zooicds, but double row systems and embedded sand. Tokioka’s (1967) specimens from the Palaw fs also appear to belong to some other species. They lack the characteristic surface depressions and the atrial lips extend from the upper rim of the open- ing Tathee than being separated from it as in the present species. 538 The narrow branchial sac, with wide bands of imperforate pharynx dorsally and ventrally is a characteristic of this species referred to by Sluiter (1909) in his description of the type. It should be noted that his figure of the branchial sac of the type (Sluiter 1909, pl.V7b) has been reversed — the larger number of stigmata and the dorsal languets are on the left, not on the right. However, Sluiter appears to have overlooked the position of the atrial aperture and the sphincter muscles of both apertures, and miscounted the number of stomach folds (see Millar 1975). Sluiter has not always been accurate in his count of stomach folds (see A. pantherinum: Millar, 1962). Aplidium depressum Sluiter, 1909 (Fig. 72) Aplidium depressum Sluiter, 1909, p.102. Hartmeyer 1909, p.1499. Van Name, 1918, p.167. Kott, 1963, p.95. Kott and Goodbody, 1982, p. 515. Not Kott 1976, p. 60; 1981, p. 160. DISTRIBUTION NEW RECORDS: Queensland (Capricorn Group, QM GH890 GH916 GH2275 GH5680 GH5682 GH5690 GH5692 GH5693-702 GH5705-6). PREVIOUSLY RECORDED: Queensland (Bargara — Kott 1963). Indonesia (Sluiter 1909; Van Name 1918). Hong Kong (QM G12796 Kott and Goodbody 1982). These transparent, investing sheets, are prob- ably more common than the records suggest. They are difficult to remove from the substrate and possibly are often overlooked by collectors. DESCRIPTION EXTERNAL APPEARANCE: Some colonies are thin, investing gelatinous sheets with a smooth upper surface, or they are irregular with the sur- face divided by shallow creases, or raised into convoluted ridges with common cloacal apertures along the ridges. One colony (QM GH5695) is a small cushion, trimmed to fit the back of a small crab on which it was found. In preservative, the small zooids are irregularly distributed and ran- domly oriented, usually lying at angles to one another. The test is translucent to glassy, firm and colourless. In life the zooids are maize yellow or orange to beige or pinkish beige, but they become white in preservative. Prochloron is found in the cloacal cavity and on the surface of some colonies (QM GH5682; see also Kott et al. 1984). Sand grains are sparsely scattered through the test, occasionally becoming crowded toward the base. MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 72. Aplidium depressum — a, colony (QM GH890); b, e¢, thorax and abdomen, posterior abdo- men separately (QM GH5697 QM GH5688); d, larval trunk (QM GH5693). Scales: a, Imm; b, c, 0.5mm; d, 0.05mm. THE AUSTRALIAN ASCIDIACEA 3 Minute test cells appear as evenly spaced grains in the test. INTERNAL STRUCTURE: Zooids are small, being: ya. more than 3mm overall, The thorax and abdo- men are of about equal length, and together are slightly shorter than the posterior abdomen which usually is relatively slender, tapering to its poste- rior end. The branchial aperture has 6 lobes, The atrial aperture is a single circular opening, some- times ona short siphon, halfway down the thorax. There ts ne atrial lip. Stigmata are in 5 rows of up to 10, their Jength reducing at the dorsal end of each row, The stomach wal] has 1] distinct folds. A:shott thick duodenum and oval posterior stamach are in the post-pyloric part of the descending limb of the gut loop. The posterior abdomen is relatively short with a smail ovary anterior to one or 2 rows of testis follicles. Upto 3 embryos are in the atrial cavily of colonies collected from Bargara in Av- gust and from the Capricorn Group from August to March (QM GH2276 GHS680 GH5683 GH5693-—4 GH5702-3). The Jarvae are amongst the smallest aplonso- branch larvae known, with a spherical trunk only about 0.35mm in diameter. The greater part of the trunk is occupied by the developing adult organs — gut and pharynx. However, the cerebral ves- icle is large in relation to the size of the trunk. A row of 7 or 8 epidermal vesicles is on cach side of the median adhesive organs, Conspicuous test evlls are in the surface of the larval test, tending tw obscure the internal structure of the larva. REMARKS The principal characters of the present species are the transparent gelatinous test, the beige, yel- low or pink zooids, the lack of an atrial lip, the 5 tows of stigmiata, the relatively numerous stom- ach folds (12) and the minute sphencal larval trunk. The type specimen (Sluiter 1909) was gelati- nous, with violet zooids seen through the test. Systems were not observed, although cloacal ap- ertures were, The atrial siphon, without ah atrial lip, is halfway down the thorax. Sluiter reported 9 rows of up to [4 stigmata in the branchial sac and numerous narrow folds m the stomach wall, Thus there is a discrepancy in the number of rows of stigmata and the colour between Sluiter’s de- seription and the newly examined material. There also is a discrepancy in the numbers of rows of stigmata in Van Name's (1918) specimens, with 6 or 7 rows of stigmata. Both Van Name and Sluiter may have had difficulty in determining the exact number of rows of stigmata in these small zooids, for most of the other characters conform with the newly recorded material. The ragged colony from Fiji (QM GH86) assigned to this species by Kott (1981), has been reexamined. ht has, as reported, 8 rows of stigmata and a minule stomach with about 12 folds. There also is a naiow atrial lip from the upper border of the aperture which Kott overlooked. The colony is not conspecific with the present species. The col- ony (QM G12796) from Hong Kong (Kott and Goodbody 1982) appears to be correctly as- signed, Aplidium directum Kou, 1972 (Fig. 73) Aplidium direcium Kott, 1972c. p.246. DISTRIBUTION New ReEcoRDS: Queensland (Dunwich, QM GH5517 GHS5519; Cape Moreton, QM GH5614). PREVIOUSLY RECORDED: New South Wales (Cronulla — AM Y1110-1 Kote 1972c). The specimens are dredged from 6m to 140m from the sea floor. DESCRIPTION EXTERNAL APPEARANCE: Colonies are flat- topped, upright lobes, etther separate or joined to a common base to form irregular, sandy masses up to 2.Scm high. The common base often is thin and keel-like and probably ts buried in the sub- strate. Each separate Jobe 1s about lem diameter, with a raised rim around its upper margin, sur- rounding a central depression containing a com- mot cloacal aperture. Zooids are around the shallow common cloacal cavity beneath this de- pression, Crowded sand particles are all over the surface of the colony and embedded throughout the test. Smaller sand grains only are on the surface in the terminal depression around the common cloacal aperture. Jn one extensive colony (QM GH5517) these smaller grains in the centre of each ternuinal lobe are black, apparently coal particles. Larger siliceous sand particles mixed with sparse black grains to give a grey colour are all over the remainder of the colony. INTERNAL STRUCTURE: Zovids are thin and thread-like. The thorax and abdomen are of about equal length and together about 6mm, The bran- chial siphon has 6 distinet lobes, The atrial lip anises from the body wall just anterior to the aperture. which projects slightly oa short siphon 340 FIG. 73. Aplidium directum (QM GH3517) — a, col- ony; b, top of colony lobe showing depressions into which systems open with smaller grains of embedded sand; ¢, thorax, abdomen and embryo in atrial cavity; d, larval trank, Scales: a, Icom; b, 2mm; c,.0.5mm; d, 0.imm. MEMOIRS OF THE QUEENSLAND MUSEUM with a sphincter muscle. The tip of the atrial lip is pointed or bidentate. There is a long unperforated band of pharyngeal wall both anterior and poste- rior to the perforated area. Short, wide stigmata are in 10 to 13 tows of up to 14 per row, The oesophagus is 4 narrow vertical tube, The stom- ach, also relatively long and narrow, has 5 well- defined folds. In extended zooids the post-pyloric part of the gut loop is relatively long. The de- scending limb has a Jong narrow duodenum and mid-intestine, but a relatively thick and slightly spiralling extent of gut in the lower one-third of the descending limb is in the position usually occupied by the short posterior stomach, well developed rectal caeca are at the proximal end of the rectum. One or 2 embryos are in the posterior part of the atrial cavity in the material from Dunwich (col- lected in August), but not in the other specimen lot from Cape Moreton taken in the same month (QM GH5614). The larval tnink is about 0.5 to 0.6mm long. It has a conical median ampulla in each of the interspaces between the adhesive or- gans. An arc of epidermal vesicles is on each side of the median line around the anterior half of the larval trunk. The tail is wound about two-thirds of the way around the trunk, Ocellus and otolith are present. REMARKS The colony of this species resembles that of Synvdicum obscurum and Aplidiwm filiforme 1.sp. in the quantity of sand present, the size of the zooids, and the circular systems in lobes rising from the upper surface, Aplidium filiforme n.sp. is distinguished by having only 5 rows of stigmata a shorter branchial sac, a longer oesophagus, shal- lower stomach folds, 4 or 5 systems per colony lobe, and Jarvae without median ampullae. The species is found in association with others superficially similar and sandy, viz. Aplidium incubatwn nsp., and Borryllus stewartense (sec Kott 1985). Larvae are not known from the New South Wales population — the type specimens of this species. However the similar morphology of these and the material from Moreton Bay indicate that they are conspecific, Aplidium distaplium n.sp. (Fig. 74) DISTRIBUTION TYPE LOCALITY: Victoria (Bass Strait, off Ninety Mile Beach, 10 km SW Scaspray, 560m off shore, calce- THE AUSTRALIAN ASCIDIACEA 3 narite reef 12m, fast currents, coll. J.E. Watson 21,3.80, holotype QM G12728; paratype QM G12729). FURTHER RECORD; South Australia (Reevesby 1, QM GH5602), DESCRIPTION EXTERNAL APPEARANCE: The colonies are solid, sandy irregular masses. Sand is crowded throughout, obscuring the smal! zooids and their arrangement. Zooids are tightly enclosed in the sandy test. anaes ) yy, FIG, 74. Aplidium distaplinm n.sp, — a, 2ovid, show- ing small posterior abdomen, long oesophagus (holo- type QM G12728); b, thorax (paratype QM G12729); ¢, larval trunk (holotype QM G12728). Scales: a, b, 0.5mm; c, 0,1mrm, 541 INTERNAL STRUCTURE: Zooids are minute, the thorax, abdomen and posterior abdomen together being about 4mm long, of which the abdomen is about half the length, the thorax and posterior abdomen being relatively short. The 6 branchial lobes are triangular and the siphon is short. A flat atrial tongue, undivided, bi-, or tripartite, extends out from the upper rim of the small aperture, About 15 muscle bands extend from the thorax, although they are fine, and the zooid is delicate, Stigmata are in 4 rows of 12, and these rows are arranged in 2 pairs as in Sycozea (see Kott 1990), The oesophagus is an unusually long vertical tube, and the 5-folded stomach is in the posterior half of the abdomen. The post-pyloric part of the descending limb of the gut loop (with the usual duodenum, mid-intestine and posterior stomach) is not as Jong as the oesophagus, Sometimes the posterior abdomen is short, with only an ovary developed, In the male phase it is longer, with a double row of male follicles, Embryos (up to 4) are incubated in the atrial cavity, The larval trunk is 0.52mm long. A large conical median.ampullae with epidermal vesicles branching off it is in each interspace between the adhesive organs. Also there is a median dorsal ampulla and a ventral one. Epidermal vesicles are also scattered in an are in the test around the anterior half of the trunk on each side. The tail is wound about three-quarters of the way around the trunk on each side of the median line. REMARKS The species resembles A, soliduin in its solid hard sandy colony, although the larval trunk of the present species is longer. The presence of 4 rows of stigmata rather than 5, is unusual in this genus. In the related genus Synogicum only &. howerbanki has a similar number of rows of stigmata (see above). Aplidium elatum Kott, 1972 Aplidium elatunt Kou, 1972b, p.177 DISTRIBUTION NEW RECORDS: None. PREVIOUSLY RECORDED: South Australia (Elliston Bay — SAM E905 E906 Kott 1972b). DESCRIPTION EXTERNAL APPEARANCE: Colonies form tall, undulating fan-shaped lamellae, Jem thick to 6¢m high, Sand is present throughout, and the colony is hard. Common cloacal apertures are randomly 542 distributed over the surface and zooids are along each side of the cloacal canals converging to them. The surface test is raised slightly over the anterior ends of each of the zooids. INTERNAL STRUCTURE: The anterior part of thoraces and the endostyles are orange in preserv- ative. Zooids are about 3mm long, the thorax and posterior abdomen approximately equal in length and slightly longer than the abdomen. The bran- chial lobes are sharply pointed. The atrial aper- ture, opposite the fourth to sixth row of stigmata has a large lip divided into 3 large, muscular, leaf-like lobes from its anterior rim. The muscles in the atrial lip are longitudinal, extending down the centre of each lobe. About 12 longitudinal thoracic muscles extend posteriorly in 2 bands along each side of the zooid. Stigmata are in 8 rows of about 8. The stomach is short with about 15 longitudinal folds. The usual duodenum and mid-intestine interrupted by a posterior stomach comprise the post-pyloric part of the descending limb of the gut loop. Gonads are in 2 series in the posterior abdomen. Larvae are not known. REMARKS The sandy colonies of this species resemble those of A. petrosum n.sp. and A. solidum, al- though they are neither as branched, nor as exten- sive. The zooids, with their large atrial lips from the upper border of the opening, and 15 stomach folds resemble A. opacum, as does their arrange- ment in double rows of zooids converging to the common cloacal apertures. Aplidium filiforme n.sp. (Fig. 75) Aplidium thomsoni: Kott, 1963, p.97 (part, specimens from Cockburn Sound). Not A. thomsoni Brewin 1950a. DISTRIBUTION Type LOCALITY: Queensland (Heron I. Blue Pools, coll. D. Parry 9.11.86, holotype QM GH5527; 9.11.85, paratype QM GH5528). FURTHER RECORDS: Western Australia (Cockburn Sound — Kott 1963). Queensland (Capricorn Group, QM GH5511 GH5515 GH5523 GH5554—5). DESCRIPTION EXTERNAL APPEARANCE: The colony consists of upright, flat-topped sandy lobes that rise from the surface of a common basal mass, and aggre- gate together along their length through the sand attached to the surface test. They form a sandy MEMOIRS OF THE QUEENSLAND MUSEUM platform up to 4cm in maximum dimension and about Icm high. Groups of 4 or 5 systems open onto the upper surface of each lobe. In some colonies the tops of a number of lobes seem to fuse to form a cushion-like platform, with crowded systems opening on its upper surface. Each system consists of a wide, sessile cloacal aperture in the centre surrounded by branchial apertures obscured by the sand which is embed- ded in the test. Sand is not always present around the cloacal openings. The systems are usually quite conspicuous, even when sand is present on the upper surface, for the branchial apertures have small sand particles outlining them, while large sand grains are embedded around the outside of each system and in the centre around the common cloacal aperture. Living zooids have orange tho- races. INTERNAL STRUCTURE: The zooids are minute, thread-like, about 0.3mm thick and about 7mm long. The thorax and abdomen together are about 3mm long and the posterior abdomen is at least twice that length. The branchial siphon is short although the branchial sphincter is large, bulging and conspicuous. The atrial aperture, with a short, stumpy tripartite lobe from the upper rim of the opening, is antero-dorsal. The 5 rows of stigmata have no more than 6 per row, although these could not be counted accurately. The oesophageal neck is longer than the post-pyloric part of the gut loop. The stomach has 5 distinct folds. Ovary and testis are not mature in the one zooid at the same time. The ovary, when present, is a short distance from the posterior end of the gut loop — in the middle of the anterior third of the posterior abdomen. When the testis follicles are mature they are in a single row in the middle third of the posterior abdomen. In the holotype colony 3 embryos are lined up in the atrial cavity. The most anterior of these is a tailed larva. The trunk length is surprisingly variable being 0.32 to 0.6mm. The adhesive or- gans are in the anterior mid-line. Epidermal ves- icles, scattered 2 or 3 deep, in a long narrow arc along each side of the median line, branch off common stalks. Median vesicles also branch off one or 2 common stalks in each interspace be- tween the adhesive organs. The arc of lateral vesicles extends well postero-ventrally and pos- tero-dorsally. REMARKS This species, with its unusual lobed colony, small zooids with 5 rows of stigmata, 5 stomach folds, orange thorax and colourless abdomen and THE AUSTRALIAN ASCIDIACEA 3 543 posterior abdomen resembles Aplidium mer- oids in each lobe of the colony, and 8 branchial nooensis (Brewin, 1956) from Chatham I., New _ lobes. Although Brewin records the height off the Zealand and one colony from New Caledonia substrate of A. mernooensis as only 2mm, its (Monniot 1987). Like the type, the specimens zooids are said to be 8mm long. Thus it appears from New Caledonia have single systems of zo- that the base of the colony was embedded in the SIC RIS SOS poe ie races 22 ae Sed mS SASS Rn ak B : PD ch apt eI SO a COL d FIG. 75. Aplidium filiforme n.sp. — a, whole colony (QM GH5523); b, upper surface of colony lobes showing systems (QM GH5515); ¢, thorax with embryos in atrial cavity (QM GH5515); d, outline of whole zooid (QM GH5515); e, larval trunk (QM GH5555). Scales: a, lcm; b, 5mm; c, 0.5mm; d, 1mm; e, 0.1mm. 544 substrate, and thar it is much the same size as the present species. However, the present species has more than one system per colony lobe to distin- guish it, and although A. merneoensis has sand externally, it lacks the embedded sand of A. fili- forme, There also are more numerous stigmata in the Chatham f. colonies than in A. filiforme. Lar- vae from the New Caledonian colony (A. nter- nocensis. Monniot, 1987} have a trink 0.65mm long with a wide lateral band of vesicles obscur- ing the anterior end of the trunk similar to the vesicles in A. caelestis, A. clivosym, A. craterif- erum, and A. griseum n.sp,. The vesicles im the narrow lateral arc of the present species are less numerous, and are spread over a smaller area at the anterior end of the trunk, The large bulging branchial siphon known in A, crateriferum and A. ellvosum is an unusual char- acter not usually present in species with zooids as smal] as those in the present species. Monniot (198?) records a strong branchial spincter in the New Caledonian specimen. The larvae and zooids of the present species are among the smallest known in the genus. Aplidivm bacculum also has only 5 rows of stigmata, but its terminal heads, onto which zo- olds open, are naked, its systems less regular, its stalks longer and the oesophagus abou! two- thirds of the length of the abdomen. Aplidium fluorescum n,sp. (Fig_ 76) Aplidium multiplicatuar Kovt, 1963, p.103 (part speci- men from Sarina). DISTRIBUTION Tyre LOCALITY: Queensland (Heron f., NE Point, rubble fauna, low tide, coll P. Kott 4.5.85, holotype QM GH5?243; Heron I., NE reef, rubble fauna, coll, P. Kott 14.11.85, paratypes QM GH5244; coll. P, Kott May 1987, paratypes OM GH5245: NE Heron reef, coll. P_ Kon 2.11.86, paratypes QM GH5246; Heron 1., The Canyons, coll. P. Kott 13.11.85, paratypes QM GH5247), FURTHER RECORDS: Queensland (Sarina. AM ¥1448; Lizard 1, QM GH5630-; Capricorn Group, QM GH5318-24). DESCRIPTION EXTERNAL APPEARANCE: The colonies are small, rounded to dome-shaped cushions, to MEMOIRS OF THE QUEENSLAND MUSEUM about Icem in maximum thickness. Only the col- ony from Sarina (AM Y1448) forms extensive investing sheets, Living test is translucent. or opaque-whitish, with green fluorescent particles around the branchial apertures and in the surtace test. These particles pive an azure-blue sheen to the colony. Zooids are ‘buff pink’, or ‘maize yellow’ (Ridgeway 1886), or pinkish white, wath an orange stomach, In preservative the zooids are beige to white and the test is almost opaque, or translucent to glassy. One of the living specimens (holotype) was described as u “translucent fluo- rescent potato’. The testis firm and naked, and there is neither adherent nor embedded sand. The systems are citcular, although often they are obscure in pre- served specimens, With the zooids lying at van- ous anges to one another in the firm test. INTERNAL STRUCTURE; Zooids are about hmm long when relaxed, the thorax and posterior abda- men of about equal length and the abdomen only a fifth of the total length, The branchial lobes are small and pointed and the branchial aperture al- most sessile, The small and undivided atrial tongue arises from the upper border of the open- ing, which has 4 sphincter muscle. Fifleen ta 20 longitudinal muscle bands are on each side of the thorax, and extend the length of the zooid, evenly spaced around it rather than forming a band on each side of the abdomen and posterior abdomen. The branchial sac has 10 rows. of up to 16 stig- mata, The moderately long stomach has 12 to 14 deep folds in its wall. The duodenum and proxi- mal sections of the mid-intestine are wide, the latter being only slightly constricted from the oval postenor stomach, Gonads are absent from the anterior half of the posterior abdomen, which is relatively short and thick and only occasionally larger than the thorax. The ovary is about halfway down the posterior abdomen, with the testis follicles in 2 iregular series posterior to il. With only one exception (May 1987) the cojo- nies from the Capricorn Group, all collected in April, May, August and November, have zooids with 2 or 3 embryos lined up in the atrial cavity. The larvae are small, the trunk being about 0.5mm long. They have the tail wound slightly more than halfway around the trank. An ocellus and an otolith are present in the cerebral vesicle. There are 3 median adhesive organs on narrow stalks at the anterior end of the trunk, and up to 9 particu- MG, 76, Aplidium fluorescum n.sp.— ab, zovids (paratypes QM GH5247 GH5244), c, larva (paratype OM GH5247). Scales: a. b, 0.5mm; ¢, O.imm. THE AUSTRALIAN ASCIDIACEA 3 i Meth Se 1 000 00 \ ruc 0000, pa on 545 546 MEMOIRS OF THE QUEENSLAND MUSEUM Sodqgoggu0e AOgquag (nn, larly large, spherical ectodermal vesicles separate from the lateral line on each side. Up to 3 embryos are in the Sarina specimens collected in August. Their larvae differ from the Capricorn Group ones only in having one more (total 10) epidermal vesicles in each of the lateral arcs. Ectodermal ampullae are never present. REMARKS The species is distinguished from others with naked colonies by its small zooids with the small atrial lip from the anterior rim of the opening and by the number of stomach folds and rows of stigmata. Aplidium gelasinum n.sp. is distin- guished by its narrow thorax with only 8 stigmata per row. Its small larvae are also distinctive, and although similar ectodermal vesicles occur in other species (e.g. A. multiplicatum), there are fewer in the present species than in any others known. Aplidium gastrolineatum n.sp. (Fig. 77) Aplidium flavolineatum: Kott, 1972b, p.177 (part, specimen from Waldegrave I.). DISTRIBUTION Type LOCALITY: South Australia (1km NW of Walde- grave I., Anxious Bay, 33°33’S, 134°46’E, rocky bot- tom, slow current, 23m, coll. S. Shepherd 11.5.71, holotype SAM E2584). FURTHER RECORDS: South Australia (Pearson I. QM GH1292; Flinders I., QM GH2305). DISTRIBUTION EXTERNAL APPEARANCE: The holotype colony is an almost spherical cushion, 2.5cm diameter and 2cm high. It is fixed basally by a relatively broad sandy area. A thick layer of sand is present around the outer surface, being absent only from the circular depressed areas into which the sys- tems open. Protruding common cloacal apertures are in the centre of each of these depressions. Zooids are in circles around each common cloacal aperture. The colony from Pearson I. lacks sand on the upper surface although it is present around the sides and base. It has the same circular sys- tems and protuberant cloacal apertures. Sand is not embedded internally. The test is soft, white and translucent, and both colonies are distorted. FIG. 77. Aplidium gastrolineatum n.sp. (holotype SAM E2584) — a, colony; b, thorax and abdomen; c, stomach from the left side. Scales: a, lem; b, 0.5mm; c, 0.5mm, THE AUSTRALIAN ASCIDIACEA 3 The zooids are tightly embedded in the test and a sheath of it, containing white opaque cells, clings to the zooids when they are removed. The living colonies are described as having a pink matrix with white zooids. INTERNAL STRUCTURE: The zooids are robust, and the thorax and abdomen together are about 4mm long. The relaxed thorax is longer than the abdomen. The 10 fine longitudinal muscle bands on the thorax extend the length of the zooid in 4 Wide band on each side. The 6 rounded branchial lobes are distinct. The atnal lip extends out from the upper nm of the aperture at the anterior end of the dorsal surface, It is thick and fleshy, with its tip divided into 3 rounded lobes. The stigmata are in 16 rows of up to 14, The stomach is rela- tively smail, its diameter not markediy greater than that of the duodenum. Its wall has 24 mostly oblique folds. The stomach wall is more or less divided into 4 quarters with the folds in each quarter running obliquely to meet those of the adjacent quarters dorsally and ventrally, and in the middle of the mesial and Jateral aspects. Three folds on the lett side of the mid-dorsal line of the stomach, and 3 on the right of the mid-dorsal line do not reach the postenor end of the stomach, and further oblique folds in the middle of each side do not reach the anterior end. The duodenum and proximal end of the mid-intestine are wide. Rectal cucea are at the proximal end of the rectum. Larvae are present in South Australian speci- mens collected in October (Kott 1972b, p.174 Fig. 30). They have a larval trunk 0.8mm long, and median and lateral ampullac between the adhesive organs, They lack epidermal vesicles. REMARKS The naked circular depressions cach with a central protruding cloacal aperture, into which the systems open, are reminiscent of the surface of Aplidium lunacratum nom. nov. However, the latter species has only 5 stomach folds, and its atrial lip arises from the body wall anterior to the aperture, rather than from its anterior rim, Aplidiwn incubatum n.sp. from Queensland also has circular systems opening into conspicu- ously naked surface areas. However, it has the surlace of the colony partially divided. each divi- sion containing a system, and it has paralle) lon- vitudinal stomach folds, Kott (1963, 1972a-c, 1975) erroneously as- signed various specimens to the South Afncan species A. flavolineatiim Sluiter, 1893 on the busis of the number of guste folds, Re-examvina- tion of these specimens has shown that they have 347 other characters which distinguish them from one another, and from the African species (see A. multilineatum 1.sp., A. robustum n.sp., A. multi- plicatum, A. uteute and the present species). The present species is distinguished from the African A, flavalineatum by its oblique stomach folds and more numerous rows of stigmata. Aplidiun robustum nsp., which has similar numbers of stigmata and stomach folds, has more crowded systems and sessile cloacal apertures. Aplidium gelasinum n.sp. (Fig. 78) DISTRIBUTION TYPE LOCALITY: Queensland (far Northern Great Bar- ricr Reef, Tydeman Reef, coll. AIMS Bioactivity Group 23.4.87 Q66 B2089, holotype QM GH5333). FURTHER RECORDS: None. DESCRIPTION EXTERNAL APPEARANCE: The colony is a flat- topped cushion; 7em in maximum dimension and icm high. In preservative, it has a white opaque deposit in the surface test, which emphasises the circular zooid systems, being more opaque around the outside of each system where the surface is raised into a rounded rim and the test is thicker — not being interrupted by either zooids or cloacal cavity, Each sysiem is about 3mm diameter. The cloacal cavity is circular with a central protuberant aperture. White deposit is concentrated in the surface layer of test. Internally the test is gelatinous and translucent. The zooids are firmly embedded and the sheath of test imme- diately surrounding each contains opaque, white cells. The specimen is reported to have been orange with orange zooids in life, INTERNAL STRUCTURE: The zooids have a long posterior abdomen, although ihe thorax and ab- domen together are only about 3mm. The atrial lip, from the upper rim of the opening, has from one to 3 points. Stigmata are in 16 rows of only about 8 per row. The small, relatively short stom- ach has 16 to 18 Jongitudinal parallel folds, Duo- denum, mid-intestine and posterior stornach are of the usual form. When contracted the posterior abdomen is pulled up into acoi] behind the abdo- men. Gonads are not mature in this specimen, and larvae are nat known, REMARKS Superticially, the regular circular systems in this opaque, white colony resemble Distaplia prolifera Koil, 1990 and Morchel/ium pannesum. 548 MEMOIRS OF THE QUEENSLAND MUSEUM The crowded small, regular circular systems also resemble those of A. lundecrarum nom. nov., al- though the latter species is distinguished by its external Jayer of sand between the systems and its fewer stomach folds. Aplidium protectans has circular systems. but 8 branchial lobes, more nu- merous rows of stigmata, and fewer stomach folds. Aplidium fluorescum and A, uteute have naked cushion-like colonies with circular sys- tems, and numerous stomach folds like the pre- sent species. However they are softer colonies, have more stigmata per row and lack the surface depressions of the present species, Aplidium geminatom n.sp. (Fig. 79) Aplidium australiensiy Kott, 1963, p.111 (part, speci- mens from Port Lincoln and Western Port). DISTRIBUTION TYPE LOCALITY: South Australia (Price [., Avoid Bay, coll. 8S. Shepherd 9.4.87, holotype QM GH4175; Port Bonython jetty, 6m, coll. G.A. Powell, paratype QM GH4229). FURTHER RECORDS: Tasmania (Bass Strait, SAM £2593); South Australia (Port Lincoln — Katt 1963); Victoria (Westernport-— AM U3914 Kott 1963). Although other records are from relatively shal- jow water, the specimen from Bass Strait is from 92m. DESCRIPTION EXTERNAL APPEARANCE: Colonies consist of spherical to oval heads (3 to 6cm diameter), each on a thick, sandy stalk, occurring singly or in clumps. Long, more or less parallel, sometimes branching cloacal canals are along the sides of the heads. Zooids are arranged in double rows, a row along each side of the canal. The canals open to a number of large cloacal openings around the up- per margin of the head, or to a few terminal apertures. A wide zooid-free strip of test is be- tween adjacent double rows of zooids. A layer of sand is present on the outer surface of the colony, including the stalk. It is absent only from a bare area around the cloacal apertures on the top of the head. Sand is not present internally, The con- tracted, preserved colonies have a conspicuously transversely wrinkled sandy stalk. FIG. 78. Aplidium gelasinum n.sp. (holotype QM GH5333) — a, colony from upper surface; b, zooid. Scales: a, lem; b, 0.5mm. THE AUSTRALIAN ASCIDIACEA 3 The anterior ends of the zooids are perpendicu- lar to the surface, but the posterior ends curve down into the stalk and cross one another. INTERNAL STRUCTURE: Zooids are robust, the thorax and abdomen together being about 3mm long when contracted. The posterior abdomen is longer and varies in length. About 12 longitudinal thoracic muscles extend posteriorly, forming a wide band along each side of the abdomen and posterior abdomen. The 6-lobed branchial siphon is short, The atrial aperture is a conspicuous, anteriorly oriented opening with a large, con- bo | Shi ite _ ain: \ & - Oy Day a Siritpe 549 spicuous atrial lip, deeply divided into 3 long pointed lobes, produced from the upper rim of the aperture. In some zooids 5 or 6 conical points project from the body wall in a transverse row just behind the atrial aperture. Longitudinal thoracic muscles lie between these projections. Fifteen rows of 10 to 12 large, oval stigmata are present in the wide pharynx. The relatively long barrel-shaped stomach has 20 to 23 parallel longi- tudinal folds. The duodenal area is relatively long and roomy, and an. oval posterior stomach is in the descending limb of the gut loop near its bend, FIG. 79. Aplidium geminatum — a, b, colonies both showing thick sandy transversely wrinkled stalk (a, QM GH4175, indicating arrangement of zooids in vertical rows with a number of terminal cloacal apertures; b, SAM £2593 with 2 terminal cloacal apertures); c, d, thorax and abdomen (e, SAM E2593 showing transverse row of dorsal papillac; d, QM GH4175); e, Jarval trunk (AM specimen from Port Lincoln). Scales: a, b, lom; e¢, d, 0.5mm; e, 0.2mm. 550 Gonads are in the long posterior abdomen. Lar- vae are not present in the newly recorded speci- mens taken in February and April. However, they are present in the type material collected in Sep- tember and November (sce Kott 1963), The larval trunk is 0.9mm long with a band of numerous epidermal vesicles from the antero-lateral ridges along each side of the 3 median adhesive organs and the median ampullae that alternate with them. REMARKS The stalked colonies with parallel, double row systems down each side resemble Sycozea and some Distaplia species (c.g, D. australiensis). Amongst Aplidium the species has some charac- ters it shares with A, australiense and A. brevilar- vacium, notably the stalked colonies with double No ee nny r ee ce ree MEMOIRS OF THE QUEENSLAND MUSEUM row systems converging to relatively few com- mon cloacal apertures. Of these two species, A. brevilarvacium, with its common cloacal aper- ture on the upper surface of the head, and rela- tively few cloacal canals, most closely resembles the present species. However, the zooids of both A. australiense and A. brevilarvacium are smaller than in the present species and they have fewer stomach folds, and fewer stigmata. There are other differences in the size and shape of the colonies (see species descriptions below). A con- spicuous character of the present species is the large atrial lip with its 3 long pointed lobes. The protruding dorsal papillae in a transverse row behind the atrial aperture are at the level of, and may be homologous with, the dorsal median papillae present in species of Polyclinum and Synoicum. FIG. 80. Aplidium griseum n.sp.— a, colony (QM GH5171);.b, ec, zooids (holotype QM GH5212 GH5172):; d, larva (QM GH5213). Scales: a, 2mm; b, ¢, 0.5mm; d, 0.2mm. THE AUSTRALIAN ASCIDIACEA 4 The larvae resemble those of Aplidium opacum with a band of numerous epidermal vesicles along each side of the median adhesive organs, similar numbers of stomach folds, and a similar large atrial lip, The form of the colonics and systems differ from those of the present species, and the zooids have fewer rows of stigmata. Aplidium griseum n-.sp. (Fig, 80) DISTRIBUTION TYPE LOCALITY: Queensland (Heron 1, under rubble, car reef edge, low tude, coll, P. Kott May 1985, holotype QM GH5212; May 1985, 1987, paratypes OM GH5214 GH5215). FURTHER RECORDS: Queensland (Capricorn Group, QM GHS5171-3 GH5213 GH5216-7 GH5529-31 GH5613 GH5668). DESCRIPTION EXTERNAL APPEARANCE: The colonies are small, upright (to 1.0cm high), almost cylindrical {to Icm diameter), flat-topped lobes. Embedded sand is crowded in the lower half of the lobe and becomes less crowded in the top half, Sand is present also on a ridge around the periphery of the upper surface of each lobe, and is crowded into a line around each system where the branchial ap- ertures open at the base of marginal ridges. As well as the penpheral ridge around ihe upper surface of each lobe the marginal ridges are on the upper surface separating the systems from one another, The thin threadlike zooids are in irregu- lar or circular systems around 2 or 3 sessile com- mon cloacal apertures on the upper surface of each Jobe. Radial extensions from the peripheral fest extend in toward the centre of a system and have zooids along each side. The regular circles of zooids are thus interrupted and the number of zooids in a system ts increased. Living colonies are ‘Indian-’, or ‘prune purple’ ta ‘blackish’ or ‘light translucent Javender’, or “Prouts brown’ extemally and “poppy red” inter- nally. Zooids are ‘vermilion’ (see Ridgeway L886). In life, red pigment in the atrial lips can be seen through the translucent test. The preserv- ative sometimes is staimed reddish-orange. Brown spherical cells are in the test of preserved specimens, INTERNAL STRUCTURE: The delicate zooids have fine Jongitudinal muscles extending their whole length. The branchial lobes are rounded and petal-shaped, The atrial lip from the body wall anterior to the small rounded aperture is 55) narrow and pointed. Opaque brown pigment par- ticles are around the zooid openings and along the endostyle. These particles probably cause the red colour ia the living zooids. The thorax, is longer than the abdomen with 13 rows of up to 16 stigmata per row. The small stomach in the mid- die of the abdomen has 8 or 9 deep folds in its wall. Of these 2 or 3 do not extend the whole length of the stomach. Caeca at the proximal end of the rectum form a rectal valve. The posterior abdomen is long and thin, with an anterior ovary and double row of male follicles in its posterior two-thirds. In specimens collected in May, a tailed larva and a second less advanced embryo are in the atrial cavity. The larval trunk is 0.85mm long, with small, pointed median ampullae in each inter- space between the 3 adhesive organs. The whole anterior end of the trunk is ebscured by an arc of scattered epidermal vesicles in the larval test along each side of the mid-line, REMARKS The species is distinguished by its upright colo- nies, small stomach with & folds in the stomach wall, relatively wide thoraces with numerous (16) stigmata per row, and delicate musculature in the body wail. Incomplete stomach folds that do not extend the full Jength of the stomach ate known in other species of this genus e.g. Aplidiwmn multipapilla- fam Millar, 1975. The latter species also has a similar number of stigmata in each row, but it bas more stomach folds than the present specics, Colonies resemble those of Aplidium craterif- erum and A, clivesum, wilh systems opening into depressions on the surface of the robust coln- nies. However, the present species has more stomach folds. The larva is like that of Aplidiwn ritteri with scattered yesicles around the anterior end, al- though the trunk of the present species is larger. Aplidium incubatum n.sp. (Fig. 81) DISTRIBUTION Tyre LOCALITY: Queensland (Point Lookout, in crey- ices on rocky platform, high tide, coll. P. Kott 12.5.87, holotype QM GH5309; paratypes QM GH5310). FURTHER RECORDS: Queensland (Point Lookout, QM GH5311-2; Cape Moreton, QM GHSG615). The records are from 0 to 10m, from rocky substrates, 552 DESCRIPTION EXTERNAL APPEARANCE: The colonies vary in shape from extensive plates up to 2cm im maxi- mum diameter to upright mushroom-shaped more or less flat-topped lobes (up to Icm high). Sometimes the base of the colony is drawn out into a long, flattened extension which fits mto narrow crevices and sometimes narrow stolons join the separate lobes of a colony to one another, or function as root-like processes fixing the col- ony in the substrate, Small circular systems of up to 10 zooids, and Smm diameter are crowded in the colony and open on the upper surface. Large, sessile common cloacal apertures are in the centre of each system. The test above the systems is naked, pinkish-white and translucent. Sometimes MEMOIRS OF THE QUEENSLAND MUSEUM sand is on the surface test separaling single, or groups of, systems from one another, Sand also is present on the surface test around the sides and base of the colony and yarying amounts are embedded in the test amongst the zooids, The test is firm and transparent in preserv- ative. In life, colonies are pink around the penph- ery of the systems, or red-orange with orange or salmon coloured zooids, INTERNAL STRUCTURE: The zooids are robust, but the thorax and abdomen together are only about 2mm long, the thorax slightly longer than the abdomen. The posterior abdomen occupies about half the length of the zooid. The 6 branchial lobes are large and fleshy, with a small, pointed papilla in the centre of each lobe, sometimes FIG.81, Aplidium incubatum n,sp,—a, colony (holotype QM GH5309); b, whole zouid (paratype QM GH5310); ¢, anterior part of zooid with embryos in brood pouch (holotype QM GH5309); d, branchial lobes (paratype QM GHS310); e, larva (holotype QM GH5310). Scales: a, 5mm: b-d, 0.5mm; e, 0.2mm. THE AUSTRALIAN ASCIDIACEA 3 55 slightly depressed into its surface. The atrial lip is produced from the upper rim of the opening and is single or divided into 2 or 3 lobes. Twelve thick, conspicuous muscles are in the thoracic wall and extend the length of the zooid in a narrow band on cach side. Broad, rectangular stigmata are tn 9 or (0 rows with up ta 15 per row. The siomach has 25 folds in its wall. The oe- sophagus and post-pyleric part of the descending limb of the gut loop are of moderate length, an ovury 9s in the anterior part of the abdomen behind the gut Joop, and the male follicles are in a long, irtegular and rather loose bunch occupying most of the posterior abdomen, Zwoids of the holotype each have a large brood pouch, constricted off from the postero-dorsal comer of the thorax, with up to 6 embryos in a developmental sequence, lined up in it. The most advanced tailed larva ts uppermost. It has a trunk 0,55mm long, and a tail wound three quarters of the way around it. About 16 epidermal vesicles are in a single arc on each side of the mid-line around the anterior half of the trunk. A total of 7 epidermal vesicles im the interspace between the upper and middle adhesive organs branch off 2 common basal stalks, and 5 branch of 2 basal stalks in the lower interspace. REMARKS A, uteute and the present species have circular systems, a brood pouch and similarly arranged epidermal vesicles. However, the larvae of the Present species lack the median vesicles, the zo- oids have fewer stomach folds and fewer stigmata per row than A. Wtewte, and the test has embedded sand, which A. utente does not have. Aplidium ornanen n.sp. has sand around the border of the colony, like the present species, but is distin- guished by its more numerous and scattered larval epidermal yesicles and lack of a brood pouch, Aplidium inflorescens n.sp. (Fig. 82) ” Aplidium opacam Kott, 1963, p.108 (part, Jobed specimen trom Balnarmg Beach). DISTRIBUTION Tyrt LocaLity: South Ausiralia (SW Point, Grindal 1, NNE Cape Catastrophe 18m, coll. L. Hobbs, H. Brandon on RV Neeria 29.9.89, holotype SAM E2588), PurTHER ReoorD: Victona (? Balnarring Beach — AM Y1401 Kot 1963) Ww DESCRIPTION EXTERNAL APPEARANCE: The colony consists of multiple stalked heads branching from a sandy common base, Heads are hemispherical to 2.4cm diameter, and the thick stalks are of slightly less diameter and up to 1.5cm Jong. Sparse sand is embedded in. and is present on the surface of, the stalk, increasing in abundance toward the base. Sand is absent from the head. Zooids are arranged in double rows converging to a few common cloacal apertures on the terminal surface of each lobe. INTERNAL STRUCTURE: The zooids are of mod- erate length, the thorax and abdomen together being about 4mm. The posterior abdomen is Jong and thread-like, The atrial lip is from the anterior tim of the aperture, It is undivided and usually its median longitudinal muscles are contracted caus- ing the edge to be frilled. Stigmata are in 15 rows with 14 or 15 per row. The oesophagus has u slight prestomach enlargement. The stomach wall has 20 folds, Up to 4 embryos are crowded in the atrial cavity of the type specimen. In the contracted zooid this causes the dorsal surface to protrude. Larvae are large, the trunk almost 0,9inim long, with median and lateral ampullae alternating with the adhesive organs. There are no epidermal vesicles REMARKS The colony of this species with its thick basal stalks and longitudinal tows of zooids converging to terminal cloacal apertures, resembles Aplidiuni geminatum, which also has a similar number of stomach folds, and is recorded from South Aus- tralian waters. The present species lacks the layer of extemal sand characteristic of A. geminatum. and its stalks are shorter, thicker, more branched and not so wrinkled. Further, the larvae of the present species lack the arc of crowded epidermal vesicles found in A. geminatum. They haye, in- stead. well-formed lateral ampullae. Aplidium brevilarvacium has similar cloacal systems but the heads are smaller and sandy, the stalks thinner and longer, and the stomach has only 10 folds, thus is readily distinguished from the present species. The larvae of the present species are similar to those of A. coniferum, which is distinguished by its large undivided upright colony, and its zooids with only 5 stomach folds. Aplidium lodix n.sp, has a similar larya without vesicles, and with median and lateral ampullae, but its colonies are extensive and investing with branching rows of 554 MEMOIRS OF THE QUEENSLAND MUSEUM Pgonoponaa® Hi = ; X ash \\ oN } . ie / 2001) 00) 2000 diwudd zooids converging to the randomly distributed common cloacal systems. The colony assigned to A. opacum by Kott (1963) is in vegetative condition, and although the colony resembles that of the present species, zooids are not available to confirm its identity. Aplidium jacksoni Kott, 1963 (Fig. 83) Aplidium jacksoni Kott, 1963, p.110 (part, holotype). DISTRIBUTION New Recorps; None, PREVIOUSLY RECORDED: New South Wales (Port Jackson — AM U3917 Kott 1963). DESCRIPTION EXTERNAL APPEARANCE: The colony fs a cir- cular slab, about 12cm diameter and 2cm thick. Sand is embedded thickly in the surface layer of test, but is not present externally adhering to the outer surface. Sand is sparse in the lower half of the colony, Circular cloacal spaces are just below the surface, covered over by a thin layer of test, and with large, irregular, sessile cloacal apertures in the centre. Zooids are in wide circles of about 8 to 12 surrounding the cloacal cavities. Each branchial aperture opens on or inside a small upstanding papilla, which has one or more of the yentral branchial lobes embedded in it. INTERNAL STRUCTURE: The zooids are robust, with a short thorax and a long posterior abdomen, The atrial aperture has a small pointed lip from the upper border of the opening, Branchial lobes are small and pointed. Stigmata are in 12 rows of 16. The wide stomach is relatively short with about 12 regular, parallel longitudinal folds. Two rows of male follicles are in the long, robust posterior abdomen, Larvae are not known. REMARKS Although it is not impossible that A. jacksoni occurs in Bass Strait, a naked, stalked, colony from off Devonport (Tasmania), assigned to this species by Kott (1963), is not conspecific. It is in the vegetative phase, and appears to be a colony of A. coniferum (see above). FIG, 82. Aplidium inflarescens n.sp. (holotype SAM £2588) — a, colony; b, thorax showing embryos being incubated; ¢, larva. Scales: a, 2cm; b, 0.5mm; e, 0.2mm. THE AUSTRALIAN ASCIDIACEA 3 The present species is characterised by its distinct circular systems, with the position of the branchial openings indicated by a papilla on the surface of the colony, The zooids have some characters in com- mon with A. triggsense which, however, has more crowded zooids and sand embedded in the basal layer of test rather than the surface. FIG. 83. Aplidiim jacksoni (holotype AM U3917) — a, surface of part of colony showing large sessile common cloacal apertures surrounded by the bran- chial apertures on small papillae; b, zooid. Scales: a, Smim; b, 0.5mm. Lay mal wh Aplidium lenticulum n,sp. (Fig. 84. Plate 13d-f) DISTRIBUTION TYPE LOCALITY: Queensland (Heron I. under coral slab 3m, coll. N. Coleman 26.11,74 AMPI 141, holo- type QM GH5060. Heron I., coll. P. Fredrickson May 1975, paratype QM GH5061. FURTHER RECORDS: Western Australia (Carnac L., QM G9475; Rockingham, QM GH5457), South Aus- tralia (Topgallant I, QM GH941), Queensland (Capri- corn Group OM GH1497). DESCRIPTION EXTERNAL APPEARANCE: The colonies are ro- bust sheets or slabs up to 20cm in maximum extent and to 2cm thick. The upper surface is divided into flat-topped, circular, oval or polygo- nal zooid-free elevations of varying diameter up to lem. These are separated from one another by narrow depressions. In the base of each depres- sion is the thin layer of test forming the roof of the common cloacal canals. The zooids open along each side of these common cloacal canals in the angle between the base and sides of the depression. The ventral surface of each zooid is in the solid test beneath each elevated area, atrial apertures are in the base or sides of the cloacal canals and atrial lips are in the test above the canals. Sand is packed in the lower half of the colony and around the outer margin, but the upper half of the colony is completely or partially free of sand. The sand occasionally spreads from the lower half of the colony into the test around the zooids and above them, into the outer margin of each clevation, thus protecting the branchial openings. Sometimes some small clumps of sand are scat- tered in the surface of each elevation and these may also be present in small papillae projecting from the upper surface. The sand is also occasion- ally present in the surface test above the cloacal canals, and around the large common cloacal apertures which appear to oceur randomly at the junctions of cloacal canals. The cloacal apertures are conspicuous, projecting from the surface of the colony, with a large cloacal chamber beneath therm. Brownish oval bodies (about 0.1mm long) are in the test especially crowded around the cloacal canals, in their base, walls and roof. They become more sparse toward the base of the col- ony. These contain granular material, but they do not appear to be faecal pellets. In life colonies are said to haye been black, preenish-black, red, aqua-blue (South Australian 556 specimens) translucent gels, although all photo- graphed specimens appear to be cream and red. INTERNAL STRUCTURE: Zooids are robust, The thorax and abdomen are together about 3mm long, and of about equal length, but the posterior abdomen is at least twice the length of the rest of the zooid, The branchial siphon is long with a large sphincter around the base of the lobes. Fine lon- gitudinal muscle bands number about 12 on the thorax. The atrial lip arises from the body wall opposite the first row of stigmata, and the atrial siphon surrounded by a sphincter muscle, is op- posite the fifth or sixth row, The tip of the atrial lip is invanably entire and pointed. The atrial lip is particularly long and broad. Zooids located in the vicinity of a cloacal aperture have their atrial lips inserted into the test around the rim of the opening. MEMOIRS OF THE QUEENSLAND MUSEUM An extent of unperforated pharynx is between the tentacular ring and the first row of stigmata. Also, bands of unperforated pharynx lie along each side of the endostyle, the perforated part being a relatively narrow part of the dorsum of the pharynx. Stigmata are in 16 to 20 rows of up to 8 per row. The stomach has 5 folds although when inflated only slight anterior and posterior protrusions re- main. The posteriar abdomen is long and thin, the test follicles in one or 2 series. are usually separated from the posterior end of the abdomen by an interval of various lengths. The ovary is at the anterior end of the male follicles. It increases in $1Ze as it moves up into the atrial cavity from 0.7 to Imm, It is a long, pointed, egg-shape, rounded at one end and tapering to a blunt point at the other. Embryos are present in the atrial cavities of specimens from Rockingham (collected in FIG, 84. Aplidium lenticulum (paratype QM GH5067) — a, part of colony surface; b, zovid. Scales; a, 5mm; b, Imm, THE AUSTRALIAN ASCIDIACEA 3 March). Tailed larvae are in a Heron 1. specimen collected in May (QM GH5061). They have me- dian ampullae alternating with the adhesive or- pans, and a mass of epidermal vesicles obscuring the anterior end of the Imm trunk. Only one embryo is incubated al a time in the posterior half of the atrial cavity. REMARKS As well as the narmw zooids with long thora- ces, long branchial apertures and atrial siphons separated from the atrial lip by 3 or 4 rows of stigmata, the gelatinous, fat-topped elevations on the surface, surrounded by the cloacal canals and zooid openings, are characteristic, The species is the opposite of such specics as A. clivosum and A. lunacratum nom. nov, which, although they have similar zooids, haye zooid- free surface elevations of the test that surround the systems rather than being surrounded by the systems as in the present species. A line of sand outlmes the zooid-free areas of test where the branchial apertures open to the surface also in Aplidium tabascum n.sp., although in the present species the sand grains in the mar- gins of the elevated areas form a crawded band yather than a single line, Also the cloacal cavities of A. fabascum are more extensive, and the atrial lip is from the nm of the aperture and not separate from it as it is in the present species. Aplidium solidum has similar, though smaller systems, more embedded sand, and a different zooid. The surface elevations and similar systems are found also in A. cellis Monniot 1987 from New Caledonia but its atrial lip is not separate from the opening, and the larvae of the 2 species differ. Aplidium caelestis has similar zooids and oval bodies embedded in the surface test as the present species. However its long, crowded meandering systems separated by long, thin ridges, rather than the circular double rows surrounding polygonal areas, and the sandy colonies with surface as well as embedded sand, distingutsh it. The nature and role of the oval bodies is not known, although it is possible that they have sometimes been mistaken for faecal pellets (see A, caelestis, above). Aplidium lodix n.sp. (Fig. 85) DISTRIBUTION TyPE LOCALITY: Victoria (Western Pon. Crawfish Rock, coll. J.E. Watson 14.11.78, holoytpe QM G12712), 357 FURTHER RECORD: South Australia (Perforated 1, QM GH4144). DESCRIPTION EXTERNAL APPEARANCE: The colony is a large sheet investing pebbles on an irregular substrate. It is uneven in thickness from 0.4 to Icm, The maximum dimension of the colony is 6cm, and it is irregularly indented and Jobed around the mar- gin and smooth and even on the upper surface, The test is firm and translucent, without sand on the upper surface, although there is some on the base. There is no embedded sand. Moderately crowded zooids are arranged in circular to stellate systems, In preservative it is greyish, with whitish zooids. INTERNAL STRUCTURE: Zooids are robust, to 6mm ia length, with thorax, abdomen and poste- nor abdomen each about one third of the total length. The 6 branchial lobes are conspicuous and rounded, The atrial tongue arises from the ante- tior rim of the aperture. Numerous stigmata are present m 10 rows, although the exact number was impossible to count, The stomach has ahout 25 longitudinal, paralle] folds. Up to 6 crowded embryos are incubated in the atrial cavily. Larvae from the holotype have trunks 0.65 to 0.75mm long, with the tail wound half-way around. Ocellus and otolith are present. A lateral ampulla is on each side of each of the 4 median ampuilae which alternate with the adhe- sive organs. A ventral median ampuila projects posteriorly, but there are no epidermal vesicles. REMARKS Superficially the preserved colony resembles Aplidium opacum. However, the latter species has more crowded zooids arranged in double branching rows, and its larva differs from the present species in having epidermal vesicles and lacking lateral ampullae. Aplidium robusium 1.sp-is another temperate species with a similarly shaped and naked colony, circular systems of varying size, and robust zooids with about 20 stomach folds, It has up to 16 rows of stigmata, its thorax is longer than that of the present species and its larva has clumps of dorsal and ventral vesicles which are not present in A. lodix. Aplidium inflorescens has lateral and median larval ampullae. lacks epidermal vesicles, but is distinguished from the present species by the form of its systems with apical common cloacal apertures, 558 MEMOIRS OF THE QUEENSLAND MUSEUM Aplidium lunacratum nom. nov. (Fig. 86. Plate 14a-c) Psammaplidium ordinatum Herdman and Riddell, 1913, p.885, Koitt, 1963, p.99. Not Psammaplidium ordinatum S\uiter, 1906, p.22. Not Aplidium ordinatum Sluiter, 1914, p.35, Hart- meyer, 1909-1911, p.1471. Van Name, 1945, p.46. DISTRIBUTION New Records: Western Australia (Cervantes, WAM 237-9.87); South Australia (Great Australian Bight, SAM E2563; Kangaroo I, QM GHS5429; Beachport, QM GH5448; Cape Jaffa, SAM E2553 E2560 E2565, QM GH5446). Victoria (Portland Harbour, MV F59370), PREVIOUSLY RECORDED: New South Wales (Manning River — AM G12216 holotype A. vrdinatum Herman and Riddell, 1913). The discontinuity in the records probably re- flects collecting effort. DESCRIPTION EXTERNAL APPEARANCE: Colonies are irregu- lar sandy mats, or upright wedges and lamellae up to lem thick with zooids opening on both sides, or cylindrical lobes of various sizes arising from common basal test with zooids opening only on the upper surface. Zooids are in evenly spaced circular systems of 5mm diameter. Branchial ap- ertures are in a circle around the periphery of each system and a circular cloacal cavity is in the centre beneath the naked concavities on the sur- face of the colony. Crowded and relatively large sand grains adhere to the ridges between the systems, and usually to the outside of the conical common cloacal protuberance that rises from the centre of each surface concavity. A little fine sand is embedded in the naked test in the concavity around the base of each cloacal protuberance. Radial bands of red pigment from the cloacal aperture to the circumference of each depression are. in the surface test, Sand is crowded through- oul the remainder of the test, Zooids lie amongst the embedded sand. They are perpendicular at the surface but more posteriorly they cross one an- other, In thin lamellae or sheet-like colonies they bend at right angles so that the long narrow pos- terior abdomina extend through the fine sand in the middle layer of each colony lobe toward its point of fixation, or to where it branches off a FIG. 85, Aplidium lodix n.sp, (holotype G12712) —a, colony with systems indicated on part of the surface; b, thorax and abdomen; ¢, larva, Scales: a, lcm; b, 0.5mm, ¢, 0.2mm. THE AUSTRALIAN ASCIDIACEA 3 Ln WN Go FIG, 86. Aplidium lunacratum nom. nov.— a, surface of part of colony (QM GH5446); b, cross section through ihe upper part of a zooid system (diagrammatic); ve, thorax and abdomen (QM GH5429); d, extended zooid posterior part of posterior abdomen not shown WAM237.87); e, larval trunk (SAM E2563). Scales: a, Smm; b-d, 1mm; e, 0.2mm. 560 basal plate. When such lamellae are broken across, the posterior abdomina can sometimes be pulled out in tufts like fine hair-like threads. INTERNAL STRUCTURE: The zooids are muscu- lar and often are contracted, although the poste- rior abdomen always is relatively long (up to lem or more). When relaxed, the thorax is 3 or 4mm long, and the abdomen about half of that length. Dark pigment cells are in the tip of the atrial siphon and in the branchial siphon of preserved zooids. Longitudinal musculature is strong with about 25 bands on the thorax. The branchial siphon is short with 6 relatively shallow lobes around the aperture. It has circular muscles in a wide band along its length, and there is no narrow, bulging sphincter muscle. The atrial tongue is fleshy and pointed to long, wide and strap-like, with parallel longitudinal muscles. It arises from the body wall opposite the first row of stigmata, anterior to the atrial siphon which is opposite the second to third rows. A ring of muscles is around the forward pointing atrial aperture on its short siphon which opens into the base of the cloacal cavity around its outer margin. The prebranchial area between the prepharyngeal band and tentacular ring is wide, as is the imper- forate pre-stigmatal and post-stigmatal parts of the pharynx. Also a wide band of unperforated pharyngeal wall is along each side of the en- dostyle. Stigmata are in 10 to 15 rows of 12 to 15 per row. The stomach is small with 5 shallow folds, sometimes (if the stomach is flattened or inflated) difficult to distinguish and reduced to longitudi- nal glandular areas internally. The anal opening is opposite the seventh row of stigmata. Spherical to oval male follicles are in a long single or double series in the middle third of the long posterior abdomen. The ovary is just anterior to the testis follicles. A single embryo is in the posterior half of the atrial cavity of specimens collected in January (SAM E2563). The larval trunk is 2.2mm long, more or less cigar-shaped, with the ocellus and otolith near its posterior end. Two median ampul- lae, one in each interspace, alternate with the adhesive organs. The anterior end of the trunk is obscured by numerous ectodermal vesicles that extend in bands, one along each side of the ventral and dorsal mid-lines from outside the base of the ventral and dorsal adhesive organs, respectively. The larva is pigmented red and is opaque. MEMOIRS OF THE QUEENSLAND MUSEUM REMARKS Like Synoicum castellatum, the present species has large circular sytems, protruding cloacal ap- ertures, atrial lips separate from the apertures, stomach folds flattened, and occasionally the stomach in the ‘atopogaster’ condition — with horizontal folds. However the internal glandular areas are all parallel and longitudinal and none are irregular as they are in the large stomach of S. castellatum. The present species also has close similarities with other Aplidium spp. It has the same long larval trunk (in the vicinity of 2mm) and a long narrow zooid with the atrial lip sepa- rate from the opening as in A. crateriferum and A. clivosum but it lacks their narrow, bulging branchial sphincters, has fewer rows of stigmata, and its systems are smaller. Aplidium lunacratum has circular systems (as in A. crateriferum) rather than stellate ones (as in A. clivosum). Despite the circular systems of the present spe- cies being different from those of the sympatric A. clivosum, in which the double rows of zooids converge to the cloacal apertures, preserved colo- nies are readily confused with one another. Both species have naked surface depressions with cen- tral protuberant cloacal apertures and red test. In both the colony shape varies, and variable quan- tities of sand are on the upper surface. In A. lunacratum there seems always to be a single system per depression, as in some specimens of A. clivasum (SAM E2564—S). As well as the form of the cloacal systems the present species can be distinguished by its larger zooids, with more mus- cles, and fewer rows of stigmata but more per row (10 to 14 rows of up to 15). Aplidium acroporum has similar systems but smaller zooids, with only 5 rows of stigmata, 8 stomach folds and the atrial lip arising from the anterior rim of the aperture. Aplidium protectans has similar circular systems, a large roomy stomach, and larvae with median ampullae and dorsal and ventral vesicles, but its zooids differ in other respects, and its colony is naked and gelatinous. Examination of the type specimen (AM G12216) of A. ordinatum Herdman and Riddell has shown that previous reports (see Kott 1963) of 8 folds in the stomach wall are incorrect. The zooids are contracted, and some of their details are obscured. Nevertheless, the relatively large stomach is seen to have the characteristic 5 shal- low folds, the atrial aperture is on a short siphon just posterior to the muscular atrial lip, and each protuberant common cloacal aperture, in the cen- tre of acircle of zooids, has sand embedded in it. THE AUSTRALIAN ASCIDIACEA 3 The name Aplidium ordinatuwm is a junior sec- ondary homonym of Psamnplidiunt ordinatum Sluiter, 1906 (sce also Van Name 1945) from the Antarctic and the name Aplidium funacratum num. nov. is accordingly proposed for this spe- cies. Aplidium macrolobatuns n.sp. (Fig. 87) Aplidium lebatum: Katt, 1963, p.97 (part, colonies from Heron J. and Sarina). DISTRIBUTION Tyre LocaLiry: Queensland (Heron 1., north eas, low tide on under surface or rubble, outer edge of reef, coll. P. Kott 2.11.86, holotype QM GH516¥; 14.11.85, pa- ratype QM GH5170). FURTHER RECORDS: Queensland (Capncom Group, AM Y1404, QM GHS5166-8 GH5219 GH5276 GH5526 GH5544-9 GH5595-§ GH5606—7 GH5660 GH5663; Bntomart Reef, QM GH380; Sarina — AM ¥ 1445 Kott 1963, Lizard 1, OM GHS550-1). One colony trom Heron L (QM GH5663) was found on a small crab. DESCRIPTION EXTERNAL APPEARANCE: The colonics are in- vesting sheets up to 0,5cm thick, or irregular masses with the upper surface and margin extend- ing into rounded or flattened lobes. Sand is em- bedded throughout the test, although often it is relatively sparse and the colonies are predomi- nantly gelatinous, Sand ts less crowded mntemally than just below the surface, and fs absent from the smooth outer surface of the colony. Branchial apertures on the upper surface have each of the 6 lurge triangular branchial lobes. outlined in fine sand grains. The cloacal apertures are large, ses- stle or protruding from the surface, They vary in srze and shape, some being itregular openings bul others regular and circular and often they are particularly extensive long or crescent-shaped Openings exposing an extent of the cloacal cavity ur canal directly to the exterior, The systems are either circular, occasionally separated from one another by slight depressions i) the surface test (holotype); but in other colonies crowded double rows of zooids converge to ir- regular and randomly scattered cloacal apertures. {n several well developed colonies (QM GH380, AM Y1445) the common cloacal apertures are along the upper rim of flattened colony lobes and double rows of zooids extend down each side of the lobes, almost parallel to ane another, The test 561 usually is firm, gelatinous and occasionally glassy in preservative. In life the colony is cloudy, owing to the em- bedded sand. When living, the thoracic body walls of the zootds are mauve, purple, raspberry, ‘pink’, ‘hazel’, ‘rose-red’. or ‘ruby’ (Ridgeway 1886) but other pasts of the zooids are colourless, Colonies, with pik or red colour seen through the embedded sand, have been described as “turkish delight’. ‘pink lozenge* or ‘pomegranate’. Zonids are pale pink jn preservative. Prochloron is on the surface of some colonies and often large red and green spherical algal cells, 0.1 to 0.2mm in diameter are in the test between and around the zooids and in the floor of the cloacal cavities especially when these are ex- posed to the exterior (QM GH380 GHS606 GH5660 AM Y1445)}. INTERNAL STRUCTURE: Zooids are small, 2mm long when contracted, the thorax and abdomen of about equal length and the posterior abdomen slightly shorter. Branchial lobes are large trian- gles with a narrow muscle band in the centre of each lobe, reaching to its pointed tip. The atrial lip from the upper rim of the opening is exvep- tionally large, and is divided inte 3 to 5 long, pointed or straight-edged, leaf-like lobes with a narrow strap-like band of muscle fibres in the centre of each lobe. Both atnal and branchial lobes have a similar pointed tip into which the central muscle band is inserted. This tip some- limes appears as a pointed papilla projecting from the end of the lobe, When the muscles in it are contracted jt is drawn back into the lobe, subdi- viding its terminal tip into 2. Six to 1 strong longitudinal muscles are on the thorax of contracted specimens. These continue along the abdomen and posterior abdomen in 2 strong Ventral bands. Nine or 10 rows of up to [5 stigmata per tow are in the branchial sac, Stig- mata are long and narrow, and are folded up when the delicate thorax is contracted. The stomach wall has 5 deep folds. Sometimes the posterior abdomen is short, with up to 10 or more pear-shaped male follicles bunched jn i. In other zooids, apparently in the female phase, a 3 to 10-ege ovary is about halfway down a long posterior abdomen, possibly after the testis falli- eles are spent (holotype). Male and female gonads are never present in the one zooid at the same time. Up to 7 eggs and embryos are packed in the atrial cavity of the holotype and other colonies collected in August and November. The most advanced embryo is anterior. The larval trunk is MEMOIRS OF THE QUEENSLAND MUSEUM ?). Kott 1963, p.103. Not Amaroucium multiplicatum: Tokioka 1953, p.180. Nishikawa and Tokioka, 1976, p.379. Nishikawa, 1990, p.89. Aplidium opacum Kott, 1963, p.108 (part, from NSW and SW Australia). Aplidium flavolineatum: Kott, 1963, p.105 (part, speci- mens from NSW and AM U3956 from Rottnest I.); 1972c, p.246. Aplidium californicum: Monniot, 1987, p.518. ? Aplidium multiplicatum: Renganathum and Monniot, 1982, p.259, DISTRIBUTION New RECORDS: Northern Territory (Darwin Hbr., QM GH5361). Western Australia (Cockburn Sound, WAM 46.72 22.75 29.75 143.75 227.82). South Australia (Yorke Peninsula, SAM E2540 E2587; St Vincent Gulf, SAM E2594). New South Wales (Port Hacking QM GH31).Queensland (Capricorn Group, QM GH1359 GH5108-27 GH5470 GH5218 GH5224 GH5241-2 GH5275 GH5281 GH5532-8 GH5540-1 GH5646; Whitsunday Is, QM GH5366; Lizard I., QM 567 GH5539; far northern Great Barrier Reef, QM GH5337). PREVIOUSLY RECORDED: Western Australia (South- western — AM Y1394 Kott 1963). New South Wales (Jervis Bay — AM Y1439; Port Hacking — AM Y815 Y823 Kott 1972c; Port Jackson — AM Y1423 U3931 U3956 Kott 1963). Kiribati and the Palau Is (Tokioka 1967); Phillipines (Van Name 1918); Truk and Ponape Is, Majuro Atoll (Nishikawa 1984a). Hong Kong (Kott and Goodbody 1982). Despite confusion in the definition of this spe- cies which resulted in its being placed in synon- ymy with eastern Pacific and S. African species, the range appears to be Indo-West Pacific. It is especially common in tropical, usually reefal, habitats of the western Pacific. It extends into the temperate waters of south-western Australia, but not across the southern coast of the continent. DESCRIPTION EXTERNAL APPEARANCE: The species forms cushion- or plate-to spreading sheet-like colo- nies, to 7cm in maximum dimension and Icm thick, with irregular borders. The upper surface is often even and horizontal. A colony from the Great Barrier Reef (QM GH5470) is a glassy dome, 6cm high, with the surface deeply divided into lobes of varying size and shape; and the specimen from Darwin Harbour (QM GHS5361) has the upper surface raised into high, zooid-free round ridges between the double rows of zooids. In other parts of the colony, however, systems are more crowded. Zooids are crowded, along each side of canals that converge to a few large, sessile common cloacal apertures with irregular borders. The living specimens have translucent, slightly cloudy test coloured orange, pale apricot, ‘poppy red’, ‘salmon’ ‘pinkish buff’, ‘yellow-buff’, “‘cream-buff’ (Ridgeway, 1886). Zooids are crim- son, ‘scarlet vermilion’, ‘geranium red’, ‘scarlet’, ‘ruby’, ‘coral reef (Ridgeway Joc. cit.) brilliant or bright orange, or pink. When embryos are present they are a dark maroon colour. In preserv- ative the test is translucent but cloudy, cream with pink zooids and mature eggs and embryos (but not tailed larvae) are a translucent brownish col- our. Opaque white (possibly blood) cells are in the sheath of soft test around the zooids, and are crowded together to form a white band around the branchial siphon. The test is particularly soft, jelly-like, is easily torn, and the surface is smooth. In some colonies green Prochloron cells and diatoms are in the common cloacal cavity, in the canals converging to them and on the outer sur- face of the colony. The species is one of the 5 or 568 MEMOIRS OF THE QUEENSLAND MUSEUM THE AUSTRALIAN ASCIDIACEA 3 6 Aplidium spp. which sometimes have Prachio- ron in the cloacal cavity (see Katt ef al, 1984; Aplidium sp.3). INTERNAL STRUCTURE: Zooids are small. the contracted thorax and abdomen together bemg about 2mm long. The posterior abdomen is short (about one-third of the total zooid length). The opaque white corpuscles that are such a feature of this species are in the body wall of the pharynx and atrial lip as well as around the base of the branchial siphon, and im the test around the zooid. The body wall is delicate wiih about [2 fine longitudinal muscle bands. Transverse muscle fibres are in the transverse vessels berwen the rows of stigmata. The 6 branchial lobes are variable, but they usually are long, and are either pointed or rounded. The atrial lip, from the upper rim of the opening contains Jongitudinal muscles and is of various lengths, Its tip either is pointed and undi- vided, or is divided into 3 or 4 separate points, There are 9 to 1] rows of 14 to 20 stigmata. Twenty-two longitudinal folds are in the wall of the small stomach which is about halfway down the abdomen. The stomach folds are sometimes irregular, interrupted, and slightly oblique on the mitside or ventral border of the stomach, but elsewhere they are longitudinal and parallel. Bunched testis follicles occupy about three- quarters of the length, of the posterior abdomen, leaving the anterior quarter free of male follicies. In relaxed zooids the follicles are spread out into 2 regular rows. A small ovary 1s behind the gut loop, anterior to the testis. Up to 2 embryos are in the atrial cavity of zooids collected in August from Darwin, May from the northem Great Bar- rier Reef, October from the Whitsunday Is, and January, April, May, August, October and No- vember from the Capricorn Group. Some speci- mens from Port Hacking (AM Y815) have up to 7 embryos packed into the atrial cavity of speci- mens collected in June, When zooids are particu- larly contracted the embryos project out from the boxy wall in what could be mistaken for a brood pouch, The larval trunk is up to 0.9mm long, and the tail is wound about two-thirds of the distance around it, On each side of the 3 median adhesive organs, is a single are of up to 30 large epidermal vesicles. Small conical median ampullate alternate with the adhesive organs in all except some of the smaller (0.6 to 0.7mm long trunk) and alniost spherical larvae from the Capricorn Group, which also have fewer (14 to 20 rather than 20 to 30) epidermal vesicles in each lateral arc. The sume opague white corpuscles are in the larval test as those in the test sheath around the zooids, and in a band around the branchial siphon. These some- times obscure the laryal organs, especially those at the anterior end of the trunk REMARKS The characteristic of this specics are the opaque white corpuscles clustered around the branchial siphon and the zooid, and scattered in the cloudy, soft gelatinous test, as well as long, double row systems, numerous stomach folds, the moder- ately wide thoraces (with up to 16 stigmata per row), and larvae with lateral arcs of numerous (14 to 30) epidermal vesicles. The discrepancy in larval size, and in the num- ber of epidermal vesicles, between smaller almost spherical larvae from the Capricorn Group and the larger ones from the same as Well as all other locations is puzzling. The only other difference observed is in the size (and probably the age) of the colonies. These Capricorn Group specimens, taken at low tide from rubble near the edge of the reef, are smaller than the colonies taken from other locations by SCUBA divers. Sluiter’s type specimens of A. multiplicaten are irregularly lobed plates 3cm in maximum diameter and 5 to 6mm thick, glassy and transpar- eni, with the yellowish-white zooids showing through the test. Systems were not identified. Sluiter records delicate muscles, 7 rows of 10 stigmata, 18 longitudinal stomach folds and a shert posterior abdomen with hunched male fol- ticles, Van Name (1918) reported larger specimens from the Philippines with more complex systems and 7 to LO rows of up to 16 stigmata per row, with the same number of stomach folds (18-20) and double series of testis follicles. As in speci- mens of this and other species, the arrangement of the gonads and the length of the posterior abdomen changes with development of the zonid and with its state of contraction. Aplidiien multiplicatum: Kott_ 1963 came from 2 locations in Queensland. Zooids from both lo- cations are smaller than those of A. nm/tiplicatum with only 6 or? rows of stigmata. Colonies fom Sarina (AM Y1448) are specimens of Aplidtum fivorescum. The colonies from Currumbin (AM 3994), with a layer of sand on the surface, FIG. 90. Aplidium multiplicatum — a, part of small colony, indicating zooids nlong each side of the clemenl a): canals which converge tO the cammon cloacal apertures (QM GHS11 b. zooid (OM GHSTIOY, ¢, stoniach, showing interrupted folds; d, larva (QM GH536)), Scales: a, 2mm; b, Imm; ©, 0.5mm; d, 2mm, S70 uppear to be colonies of A. ornatunr with circular systems and sand around the margin of the col- ony. Aplidjum mudsiplicatum: Millar, 1963 is also wrongly assigned, The 3 specimens have circular systems, about 25 stigmata per row and 2] stom- ach folds. The larvae have lateral ampullae but no vesicles. It is possible that more than one species is involved. The number of rows of stigmata and number per row suggest A, wfeite is one of the species, but the larvae and the number of stomach folds do not support that view, Specimens assigned to A. multiplicatum by Tokioka (1967) also may include more than one species. In specimens only 2.0cm in maximum dimension, and not more than Icm thick, there are some with circular. and others with more complex systems, a range of colours from “pale brown’, ‘reddish brown’. to ‘purplish brown’ (Tokioka 1967). 7 to 10 rows of stigmata, 10 stigmata per row Gn zooids from Kinbati) up to 30 (larger zooids from Palau Is), and 20 to 33. stomach folds. The range in these characters is greater than could be reganded as intraspecific Vanation and greater than could be regarded as the effects of growth, especially when the range in colony size is tela- tively small. The larvae of the small colony from Kiribati, and the zooids figured from both Kinbali and the Palau Is (Tokioka 1967, Figs 8a—d, 9a,b) appear to be A. multiplicatum. Colonies with more numerous stigmata and stomach folds could be other species such as A. ufeute, which resem- bles the present species in the colour of its zooids and embryos. However, A. utewte has median as well as lateral larval epidermal vesicles, mere stigmata In cach row and more stomach folds than A, multiplicetunt, Nishikawa and Tokipka (1976), Nishikuwa (1990), Tokioka (1953) and Rho (1975) have assigned specimens from Japanese waters with 7 to 1 L rows of stigmata, and 21) ta 25 stomach folds to A. snultiplicatim. The number of stigmata per row, and the arrangement of larval epidermal vesicles and ampullae do not fall within the ranges recorded for the present westem Pacific Species, Tokioka (1953, 1967) proposed that the eastern Pacific Aplidiven californicum (Ritter and Forsyth, 1917) is a synonym of the present spe- cies. However, the Californian species has sand embedded in the test, an atrial lip separate from the opening, fewer stigmata per row, and lacks the characteristics pigmentation of the present spe- cies (see Kott and Goodbody 1982, Nakauchi 1987, Millar 1988). Aplidium californicwn: Monniot, 1987 from New Caledonia has the char- MEMOIRS OF THE QUEENSLAND MUSEUM actenstic naked colony, pigmentation, stomach wall, branchial sac and larval form of A. mulripli- catum. In Australia Aplidiur opacum is the species most closely related to the present one and speci- mens have been confused. [t has similar colonies with crowded zooids in long branching double- row systems, Zooids are sheathed in a layer of soft test containing opaque white cells which are simi- lar to those in A, multiplicatum, and are crowded around the branchial siphon in much the same way. The zooids also are similar, although the Present species usually has more stigmata perrow but fewer rows than A. opacum. Aplidium opacun has firmer test, and the larvae have long finger-like median ampullae that project between the adhesive organs, as well as a band of numer- ous lateral vesicles (rather than the single series are thal occurs in A. mulliplicatum). Re-examina- tion of some of the specimens from New South Wales and south-westem Australia assigned to.4- opacum by Kott (1963) has shown them to belong to the present species.. Larvae of the south-west- em Australian specimen are characteristic of A, multiplicatum (see Kott 1963, p.110, Fig, 22). The ranges of A. mulriplicatum and A. apxicum overlap on the eastern and western coast, but the former is not recorded from southern Australia, and the latter species does not extend into the tropics (see A. opacim, below). Kott (1963) wrongly assigned colonies of the present species and others (viz. A. gastrolinea- ruin, A. multilineatum and A. robustum asp.) to the $8. African A. flavolineanun, Aplidium opacum Katt, 1963 (Fig. 91. Plate 15a-c) Aplidium upaciwn Kort 1963, p.108 (part, holotype AM 133929), Aplidiim pliciferunt: Kott, 1963, p, 106; 1972a, p. 13; 1976, p.62, Aplidium feliorum Katt, 1975, p35. DISTRIBUTION New Recorbs: Western Australia (Rockingham, QM GH5461). South Australia (Great Australian Bight, SAM E2591, Port Bonython, QM GH4234-4, Perip- tated 1., QM GH4194; Spencer Gulf, QM GH5441-1; Yorke Peninsula, SAM E2540 E2571 E2587; St. Vin- cent Gulf, SAM E2484 E2586 E2594), Tasmania (SE Fluted Cape, SAM E2589). New South Wales (Port Hacking, QM GH31). PREVIOUSLY RECORDED; South Australia (Great Aus- iralian Bight — Kolt 1975; St. Vincent Gulf — Kott THE AUSTRALIAN ASCIDIACEA 3 1972a), Victoria (Balnarring Beach — AM U3929 holotype Kott 1963; Westernport — Kott 1976), DESCRIPTION EXTERNAL APPEARANCE: Colonies are large and fleshy sometimes almost spherical or flat- tened upright masses to 8cm high and 6cm diame- ter. More often they are flat-topped, extensive cushions often with the surface partly divided. Crowded zooids are perpendicular to the surface, but posteriorly criss-cross one another in the test. In preservative the test is soft, but firm, gelati- nous and translucent white with minute opaque cells, possibly blood corpuscles, in a soft layer of test that sheaths each zooid. These opaque cells are especially crowded around each branchial aperture, In photographs of well-developed living colo- nies zooids are seen to be in crowded, branching double rows, with large branching primary cloa- cal canals separating areas packed with zooids and secondary cloacal canals, Large, sessile com- mon cloacal apertures are scattered over the sur- face at the junction of the primary canals. Zooids are cream (QM GH5441) or pink (QM GH5440). \ \\ 571 One colony (QM GH4194) is described as an ‘orange morph’: and one is bright yellow (SAM E2586). At least 2 colour variants occur, both translucent, One pink or orange and one yellow or yellowish cream. INTERNAL STRUCTURE: Zooids are relatively long and robust. The thorax and abdomen even when contracted are together up to Smm long and the posterior abdomen is from twice to many times that length. About 15 strong longitudinal muscle bands extend from the thorax to the pos- terior end of the abdomen. The branchial siphon is short. often with well-developed bifid lobes. The atrial aperture, beside the third row of stig- mata, has a fleshy muscular atrial lip, usually deeply divided into 3 equal, pointed lobes, from the upper border of the opening, Muscles are along the centre of each lobe, and when these are contracted the lobes are frilled and crinkled along the edges, The atrial lip is often (but not always) large and conspicuous, Stigmata are in 12 rows with up to 12 in cach row. The stomach is of moderate length with 20 narrow, sometimes interrupted folds. The thick duodenum, mid-intestine and oval posterior FIG. 91. Aplidium opacum —a, zooid, showing anterior part of posterior abdomen and embryos being incubated (QM GH5441); b, larva (QM GH4234), Scales: a, 0.5mm; b, 0,1mm, stomach are characteristic of the genus, and there are 2 pronounced rectal caeca, Gonads are in the posterior past, or the whole of the narrow paste- rior abdomen, with the small ovary anterior to the long double series of male follicles. One to 3 embryos are incubated in the top of the abdomen and in the atrial cavity, but only one at a time develops toa tailed larva. They are present in specimens from South Australia collected in February (QM GH5441 GH4234 SAM E2586). April (QM GH4236 GH4194), May (QM GH4235) and November (SAM E2571). Larvae have a trunk about 0.7mm long and a sturdy tail reaching to the anterior end of the trunk. Ectoder- mal yesicles (about 35) are crowded in an are along each side of the adhesive organs. These vesicles are not in a single series. and clumps of them branch off a single stem from the larval ectoderm. Narrow, pointed, median ampullae al- ternate with the adhesive organs. REMARKS Both the zooids and the large gelatinous colo- nies, with randomly scattered, sessile common cloacal apertures, and crowded branching sys- tems of zooids in the present species are similar to Aplidium multiplication, These species also haye in common the cloudy opaque cells in a sheath of soft tést around each zooid, and about 20 stamach folds. The zooids of A. opacum are larger than those of A. multiplicatum however, and although they have the white opaque cells in a sheath around the zooids, they do not form as wide a band around the branchial aperture as 1 A, multiplicatum-The branchial sac of the present species 1s relatively narrower than that of A, niul- iiplicatem, Which has up to 20 stigmata per row but fewer than 12 rows. Larvae differ from those of A. muliplicatum, having jong median ampul- lae and clumps of lateral vesicles in lateral bands (tather than a single arc of vesicles on each side). Kott (1963) assigned seyeral colonies to this species which have subsequently been found not conspecific with the holotype. A colony from Balnarring Beach (AM Y140)1) made up of sev- cral lobes joined to a common base is probably a specimen of A. inflorescens {see aboye). The colonies from westerm Australia and Port Jackson (AM Y1423 Y1439) are specimens of A. multi- plivatum as 1s the larva figured by Kott (1963, Fig. 22), Colonies from Tasmania with zooids in which the atrial aperture is separate from the atrial lip are also erroneously assigned to this species and appear to be colonies of a Polycliniuwmn sp, MEMOIRS OF THE QUEENSLAND MUSEUM Kott (1963) assigned specimens of the present species from Cockburn Sound to Aplidium plicif- erum (Redikorzev, 1927). The latter species, known from coastal waters of Japan, including the Inland Sea (Tokioka 1953), has large colonies with branching double row cloacal systems as in the present species, and zooids and laryae which also ar to be similar to the Australian species. Only the stomachs are different, having from 25 to 32 fine pleats in the Japanese species, while the Australian one has 20 to 24. Thus the number of stomuch folds appears to be the principal charac- ter separating these species, which are indigenous to temperate Australian and Japanese waters re- spectively. Despite their resemblance to one an- other, they apparently have different geographic ranges, and are separated from one another by the tropics. Aplidium folierum Kott, 1975 was thought to be distinguished by its large tnpartite atrial lip- An identical lip now is known to be characteristic of the present species, although its length yaries to some extent according to the position of the zooids relative to the cloacal apertures and ac- cording to their state of contraction. Aplidiwm macrolobatunt has similar large lobes around the apertures, but it has sand embedded in the test, and only 5 stomach folds. Aplidium ornatum n.sp, (Fig. 92) ? Aplidium multiphieatum: Kott, 1963, p.J03 (part, specimens from Currambin). DISTRIBUTION TYPE LOCALITY: Queensland (Wistari Reef, NW cor- ner, low tide under nibble near reef edge, coll. P. Kott 3.11.86 holotype QM GH5157). FurRTHER Recorb: ? Queensland (Currumbin — AM Y3994 Kott 1963). DESCRIPTION EXTERNAL. APPEARANCE: The colony is small (Smm high), mushroom-shaped, with a flat top about Icm in diameter, The diameter reduces toward the base. Sand is present around the out- side of the colony but is absent from the upper surface and internally, Zooids. are arranged in circular to elongate systems opening on the upper surface. The 3 or 4 common cloacal a res are Jarge and protruding, sometimes in tbe centre of a circle of zooids, but sometimes at one end of a THE AUSTRALIAN ASCIDIACEA 3 double row of zooids. In life the zooids are ‘ruby’ and the test ‘isabella’ coloured (Ridgeway 1886). INTERNAL STRUCTURE: The zooids are vertical, with the long posterior abdomina extending into the base of the colony, Thorax and abdomen are together about 3mm long. The atrial tongue from the upper tim of the opening is undivided or divided into 3 pointed Jobes, There are 12 rows of stigmata, and the maximum number per row is about 10, although the holotype is contracted and the exact number difficult to determine. The stomach wall is folded longitudinally into 25 folds, Two large embryos are in the atrial cavity of the holotype. The larval trunk is 0.66mm Jong, In each interspace between the middle and the dorsal and ventral adhesive organs respectively, is a median ampulla with a bi-lobed lateral ampulla 573 on each side. Dorsal and ventral clusters of nu- merous epidermal vesicles extend posteriorly from the anterior end of the larval trunk. The tail is Telatively short, extending about two-thirds of the way around the trunk. REMARKS The distinctive characters of this species are its relatively narrow thorax, numerous stomach folds, and larvae with epidermal ampullae and numerous vesicles. Aplidium incubatum has a similar colony to the present species, but its cir- cular systems are more regular and its larvae have median and lateral vesicles rather than the ampul- lac of the present species. Aplidium multiplicaium, which it otherwise re- sembles in the colour of the zooids, and the bran- FIG, 92. Aplidium ornatum n.sp. (holotype QM GH5157) — a, colony; b, tharax and abdomen; c, larval trunk. Scales; a, 5mm; b, 0.5mm; ¢, 0.1mm: 574 chial sac and stomach folds, is distinguished by having single arcs of epidermal vesicles in the larval trunk, sand absent from around the border of the colony, a softer test, opaque white pigment particles around the branchial apertures and long branching double-row systems. The specimen from Currumbin assigned to A. multiplicatum by Kott (1963) has sand around the base and sides of the colony, but not in the upper suface or embed- ded. The zooids are small, contracted, and appear to be arranged in circular systems. The atrial lip is from the upper rim of the aperture, and there are 18 stomach folds. Other details of their mor- phology are obscured. Aplidium uteute has numerous stomach folds, but is distinguished by its smaller circular sys- tems, completely naked colonies, more numerous stigmata per row but fewer rows, single arcs of larval epidermal vesicles, and, usually, the ab- sence of red pigment from the thorax. Aplidium altarium (Sluiter, 1909) has numerous stomach folds and colonies of similar dimensions to the present one, but the 9 or 10 rows of stigmata have about 25 per row, the posterior abdomen is short with bunched male follicles, and unlike the pre- sent species, the colony is transparent. MEMOIRS OF THE QUEENSLAND MUSEUM Aplidium paralineatum n.sp. (Fig. 93) DISTRIBUTION Type LOCALITY: Tasmania (45 nautical mls NNE of Sister I.,39°00’S 148°25’E, 92m, sand bottom, epiben- thic sled, CSIRO Statn 22, coll. W. Zeidler 14.10.84, holotype SAM E10648). FURTHER RECORDS: None. DESCRIPTION EXTERNAL APPEARANCE: The specimen is a long (3cm), narrow cushion, broken at one end. It probably is a piece from a longer colony. A coni- cal prominence about lcm high with a terminal common aperture is at the other end. Zooids are in parallel double rows, a row along each side of the long, straight cloacal canals that run the length of the colony, converging to the large, sessile cloacal aperture. Long ridges separate the double rows of zooids from one another. The test is relatively soft, although sand is embedded throughout. Along the under surface the test is produced into fine root-like processes, which, with their adherent sand, create a felt-like mass holding the colony in the sandy substrate. The colony appears to have been organised in a single FIG. 93. Aplidium paralineatum n.sp. (holotype SAM E10648) — a, colony; b, thorax and abdomen. Scales: a, Smm; b, 0.5mm. THE AUSTRALIAN ASCIDIACEA 3 cloacal system, although there may have been a second cloacal aperture at the mutilated end. INTERNAL STRUCTURE: The zooids are robust, However, the thorax and abdomen together are only about 3mm. The posterior abdomen is long and threadlike. The atrial tongue is relatively small, pointed, and produced from the anterior rim of the aperture. There are 15 rows of stigmata although these zooids are too contracted to deter- mine the number in each row. The stomach is small with 5 deep folds in its wall. A single series of testis follicles are in the posterior end of the long posterior abdomen. REMARKS The species is distinguished by its long double row systems and sessile colonies, Similar long systems are 1n the stalked colonies of A. austral- iense which, however, has more numerous stom- ach folds than the present species. Aplidium parastigmaticum n.sp. (Fig. 94) DISTRIBUTION Type LOCALITY: Queensland (Heron L., reef slope 10m, coll. §. Cook 28.4.88, holotype QM GH5603). FurTHeR Recorp: Queensland (Heron [., QM GH5604) DESCRIPTION EXTERNAL APPEARANCE: The colonies are ses- sile spheres about 3cm in diameter. The test is firm and translucent, with some sand on the ridges between the systems. Living zootds are vermilion with buff coloured branchial apertures, [n pre- servative both zooids and test are pinkish. Zooids are arranged in circular to star-shaped systems, about 7mm in diameter. The cloacal cavity is large, with some peripheral extensions to reach the zooids in the outer periphery of each of the systems. The test is soft, and the zooids all lie parallel to one another. Sand is embedded in the basal test, but not elsewhere, INTERNAL STRUCTURE: Zooids are large and tobust, about 7mm in overall length even when contracted. In the contracted zooid the posterior abdomen is about half and the thorax is slightly more than one quarter of the total zooid length. FIG. 94, Aplidium parastigmaticum n.sp, (holotype QM GH5603) — a, colony; b, zooid: e, portion of branchial sac showing muscle fibres in primary trans- yerse and parastigmatic vessels, Scales: a, lem, b, Imm, ¢, 0.5mm. = 576 The longitudinal muscles tend to form a band in the dorsal half of each side of the thorax, and then continue in a wide band along the ventral part of the abdomen and posterior abdomen. Their contraction causes the thorax to curve around the shortened dorsum. Strong transverse muscle fi- bres are in the primary transverse and parastig- MEMOIRS OF THE QUEENSLAND MUSEUM matic branchial vessels. They separate into short terminal branches on each side of the endostyle. The branchial aperture is 6-lobed, and the atrial aperture has a fleshy lip from the upper border of the opening. The tip of the atrial tongue is either undivided, or divided into 2 or 3 lobes. The for- ward pointing atrial aperture opens from a pouch FIG. 95. Aplidium paryum — a, colony (AM Y¥1425); b, thorax and abdomen (AM Y 1425); ¢, mature larva (AM Y 1418); d, immature larva (AM Y1418). Scales: a, 5mm; b, 0.5mm; ¢, d, 0.2mm. THE AUSTRALIAN ASCIDIACEA 3 in the atrial cavity, which does not contract with the contraction of the dorsal part of the zooid. The stigmata are in 18 rows, with about 12 per row. Each row is crossed by a conspicuous parastig- matic vessel. A dorsal languet is on each paras- tigmatic, as well as on each primary transverse vessel. The stomach at mid-abdominal level, has 5 distinct longitudinal folds. The male follicles are in two longitudinal series. REMARKS The red zooids in the translucent test resemble A. uteute which, however, has close circular sys- tems (rather than the star-shaped ones of the present species) and more numerous stomach folds. The parastigmatic vessels, the atrial lip from the upper border of the opening, the pouch that forms beneath the atrial aperture when the zooid is contracted. The large stomach with 5 folds, and the parastigmatic vessels all occur in the temperate Aplidium amorphatum which is distinguished by its 8 branchial lobes, distinct circular systems, and poorly developed gastric folds. Aplidium protectans, which extends further north than A, amorphatum but has the same large zooid and atrial and branchial apertures, has dis- tinct circular systems opening to depressions in the surface, and lacks parastigmatic vessels. Some features of the present colony, such as the stellate systems, and surface sand between them, are reminiscent of A. clivosum. However, the atrial lip is separate from the aperture in A. clivo- sum, the zooids are larger than those of the present species and there are no parastigmatic vessels. It appears, therefore, that the colony is a member of a formerly undescribed species. Aplidium parvum Kott, 1963 (Fig. 95) Aplidium parvum Kott, 1963, p.112. DISTRIBUTION NEw REcorDs: None. PREVIOUSLY RECORDED: Western Australia (Albany — AM Y1425 Kott 1963). Victoria (San Remo — AM Y1418 Kott 1963). DESCRIPTION EXTERNAL APPEARANCE: Colonies are small spherical transparent heads on short stalks to lcm high overall. The zooids are arranged in longitu- dinal rows, converging to a common cloaca on the top of the head. INTERNAL STRUCTURE: Zooids are small, the length overall extending to about 6mm. The pos- terior abdomen is up to twice the length of the remainder of the zooid. The short branchial si- phon has 6 lobes around the aperture. The atrial aperture has a short, undivided lip from the ante- rior rim of the opening. Stigmata are in 12 rows of about 8. The short stomach has 18, mostly parallel, folds. The oesophagus has a pronounced prestomach enlargement at its distal end. Three embryos are incubated in a brood pouch that projects from the thorax in the syntypes (AM Y1418) collected in October. The larval trunk is 0.8mm long. Anteriorly, single median ampullae are in the interspaces alternating with the adhesive organs. These me- dian ampullae have columnar cells on them, and when well developed they become flat-ended rather than conical, the stalks of the adhesive organs become shallower and saucer-shaped, with long, slender stalks. REMARKS The small colonies, zooids and larvae are all distinctive. Although the systems resemble those of A. altarium, A. geminatum, A. inflorescens and A. brevilarvacium, the colonies are smaller, and stomach folds and stigmata are fewer. Like the present species the larva of A. inflorescens lacks epidermal vesicles. However, its trunk is about twice the length of that of the present species, and is further distinguished by the presence of lateral as well as median ampullae. Aplidium gemi- natum, Aplidium altarium and Aplidium brevilar- vacium have different larval epidermal vesicles from the present species, and larger larval trunks (about Imm long). Further, Aplidium altarium has appreciably more stigmata per row than the present species, A. brevilarvacium has fewer stomach folds (10), and in A. geminatum the stomach folds are more numerous. Aplidium petrosum n.sp. (Fig. 96. Plate 15d-f) DISTRIBUTION TyPE LOcALITY: South Australia (Cape Jaffa, Mar- garet Brock Lighthouse, coll. AIMS Bioactivity Group 18.2.89, holotype QM GH5454). FURTHER RECORDS: South Australia (The Gap, QM GH4165-6; Hopkins I., QM GH4190). DESCRIPTION EXTERNAL APPEARANCE: Colonies are rigid, sandy, flat lamellae, about Icm thick, which di- 578 vide and curve to form a large 3-dimensional maze. Zooids open on both sides of the lamellae. They are arranged along each side of cloacal canals that surround circular to elongate zooid- free areas. Common cloacal apertures are at the junctions of the cloacal canals. The test is slightly depressed over the cloacal canals in the preserved specimens. Sand is densely packed in the test and obscures the systems. Zooids, lying parallel to one another, are tightly enclosed in the test. INTERNAL STRUCTURE: Zooids are small and slender, the thorax and abdomen together being about 2mm long. The branchial siphon is short and the aperture small with 6 short, rounded lobes. The small atrial lip is pointed or bipartite and projects from the anterior rim of the opening. MEMOIRS OF THE QUEENSLAND MUSEUM Stigmata are in 9 rows with only 6 per row, in the narrow thorax. The stomach is long and narrow with 5 longitudinal folds. The posterior abdomen is about twice the length of the rest of the zooid, with male follicles in double rows; or when the zooid is contracted, drawn up into a bunch behind the gut loop. Three embryos are being incubated in the atrial cavity of specimens collected in April (QM GH4165). The larval trunk is about 0.8mm long. Single large, conical, median ampullae alternate with the adhesive organs; which have slender stalks that expand into a cone at the base. A wide band of epidermal vesicles is along each side of the median line around the anterior half of the trunk. FIG. 96. Aplidium petrosum n.sp. — a, part of colony (QM GH5454); b, thorax and abdomen (QM GH5454); c, larval trunk (QM GH4166). Scales: a, lcm; b, 0.5mm; ¢, 0.1mm. THE AUSTRALIAN ASCIDIACEA 3 REMARKS Colonies forming stiff sandy lamellae are known also for A. solidum from the eastern and north-western coasts. They also have similar zo- oids, arranged in similar systems. The present species is distinguished by its more branched colony (A. solidum having upright lamellae rather than a 3-dimensional maze); and by the larvae which are larger than those of A. solidum (0.5 to 0.6mm long trunk) and have median ampullae (not present in A. solidum). Aplidum ritteri have similar zooids with the atrial lip from the upper rim of the opening, but also have smaller larvae than those of the present species and long straight cloacal canals. Aplidium protectans (Herdman, 1899) (Fig. 97. Plate 16a,b) Amaroucum protectans Herdman, 1899, p.73. Aplidium protectans: Kott, 1963, p.102. Aplidium longithorax Monniot, 1987, p. 525. DISTRIBUTION NEW RECORDS: Queensland (Capricorn Group, QM GH5562 GH5574; central Great Barrier Reef, QM GH5362 GH5378; Lizard I., QM GH332 GH5553 GH5611-2; far nothern Great Barrier Reef, QM GH5336). PREVIOUSLY RECORDED: New South Wales (Mon- tague South — Kott 1963; Port Jackson — Herdman 1899). New Caledonia (Monniot 1987). The species is recorded from 0 to 100m depth. DESCRIPTION EXTERNAL APPEARANCE: Colonies are firm, gelatinous naked domes, spheres or cushions 2.5cm to 6cm in maximum dimension. The test is translucent both living and in preservative, and the zooids are pink, yellowish or orange. Circular systems, each of from 10 to 16 zooids are evenly distributed around the outer surface of the colony, and are slightly depressed into the surface in preserved colonies. The common cloacal aper- tures are in the central of each system. Systems are well spaced and never crowded. They appear as regular daisy-like patterns over the surface of the colony. INTERNAL STRUCTURE: Zooids are large and robust, the thorax and abdomen even in con- tracted specimens together being more than 4mm long, and the posterior abdomen about the same length. The branchial siphon has 8 rounded lobes, the atrial siphon is produced out toward the cen- tral common cloacal aperture and an undivided, 579 fleshy atrial lip arises from just above the sphinc- ter muscle in the rim of the opening. In some zooids the atrial lip appears to be part of the rim of the siphon, but in others the atrial siphon is close to, but separate from its lip. The blunt tip of the atrial lip is divided into 3 to 6 charac- teristically regular, short, finger-like rounded lobes. The body wall is muscular with about 20 longitudinal bands on each side of the thorax, extending along the length of the zooid in a wide lateral band on each side. Stigmata are in 18 to 24 rows with about 20 per row. The oesophagus is relatively long, opening into a long barrel-shaped stomach with 5 shallow folds about one third of the way down the abdomen. When contracted the stomach wall tends to be drawn into horizontal folds. There is a short thick duodenum, but the remainder of the post-pyloric part of the descend- ing limb of the gut loop is long, curving around in acorkscrew spiral, which may be a homologue of the posterior stomach, although it is longer than usual. A narrow, short section of mid-intestine intervenes between this thick distal end of the descending limb and the rectum. The rectal caeca are not well developed. Two or 3 large eggs are present just behind the gut loop, separated from the pear-shaped male follicles which are in one or 2 longitudinal series in the remainder of the posterior abdomen or, if the longitudinal body muscles are sufficiently contracted, bunched. Up to 3 embryos, brown in preservative, are lined up in the posterior part of the atrial cavity of specimens collected in August (QM GH5362) and tailed larvae are present in colonies from Lizard I. collected in June (QM GH5553). The larval trunk is 0.8-0.9mm long. It has single median ampullae between and dorsal and ventral to the adhesive organs, and 4 clusters of epidermal vesicles, one on each side of the dorsal and ventral mid-lines behind the adhesive organs. REMARKS The large zooids in conspicuous circular sys- tems, larvae with median ampullae and ventral and dorsal clusters of vesicles, and fleshy colo- nies are characteristic. Kott (1963) also noted the tendency for the stomach to collapse in horizontal folds in contracted zooids from a specimen from NSW waters - which was found on a crab. Polycitor giganteum (also found on crabs) has similar large, firm, gelatinous colonies which may be a convenient consistency for the crab to hold in place and trim to an appropriate regular size and shape. 580 MEMOIRS OF THE QUEENSLAND MUSEUM Colonies and zooids resemble Aplidium amor- Aplidium indicum (Renganathan and Monniot, phatum in having & rather than 6 branchial lobes, 1984) from India has a similar fleshy colony to The present species has firmer colonies alonger the present species and equally shallow stomach thorax and abdomen, and it lacks parastigmatic folds. However, the small, extended zooids of the vessels. Larvae, although smaller, have median Indian species were not seen to be arranged in ampullae and dorsal and ventral vesicles like A. circular systems, and have fewer rows of fewer lunacratum. stigmata than the present species. FIG. 97. Aplidium protectans —a, small colony (QM GH5329); b, thorax, abdomen and anterior part of posterior abdomen showing digitiform tip of atrial lip and large stomach (QM GH5553); e, branchial aperture from above THE AUSTRALIAN ASCIDIACEA 3 Aplidium ritteri (Sluiter, 1895) (Fig. 98) Amaroucium ritteri Sluiter, 1895, p. 10. Aplidium lobatum: Tokioka, 1967, p.22. ? Nishikawa, 1984a, p.110. Monniot, 1987, p.525. Monniot and Monniot, 1987, p.73. DISTRIBUTION NEW RECORDS: Queensland (Capricorn Group, QM GH5649 GH5660 GH5685 GH5704 GH5779-8 1). PREVIOUSLY RECORDED: Queensland (Torres Strait — Sluiter 1895). Western Pacific (Palau Is, New Caledo- nia — Tokioka, 1967, Monniot, 1987; Truk, Ponape — Nishikawa 1984a). French Polynesia — Monniot and Monniot 1987). The species is a tropical one. DESCRIPTION EXTERNAL APPEARANCE: The colonies are in- vesting sheets or lumpy and massive with a rela- tively even surface and sand embedded throughout. The ridges that separate the long dou- ble rows of zooids are relatively inconspicuous, and the zooid openings along each side of the cloacal canals between these long ridges often are obscured by the embedded sand. Common cloa- cal apertures are at the junctions of 2 or more canals. Living colonies, are described as ‘sandy with pink’, and ‘solid, sandy’. INTERNAL STRUCTURE: Relaxed zooids are about 3mm long. The thorax is about half the total length and the abdomen and posterior abdomen each half of the remainder. The 6 branchial lobes are well-formed and triangular. The atrial aper- ture is small and rounded with a short, pointed atrial lip from the anterior rim of the opening. A conspicuous bulging sphincter is behind the bran- chial lobes. The body muscles are strong and the short posterior abdomen is often drawn up into a short clump behind the abdomen. Stigmata are in 11 or 12 rows of up to 8 in the narrow pharynx. The most ventral and dorsal in each row are short. The oesophagus is moderately long, and the stomach is divided into 5 by the deep folds in its wall. There is a long, narrow duode- num, narrow proximal part of the mid-intestine and an oval posterior stomach in the end of the descending limb of the gut loop. The posterior end of the posterior abdomen is curiously flattened and straight edged. Up to 8 male follicles alternate with one another, or they are in a single series in the relaxed posterior abdomen; when the zooid is contracted they are bunched together. Male and female gonads are 581 not present together in the one zooid. Embryos are incubated in the atrial cavity of specimens collected in November (QM GH5649) the tailed larva is small, the trunk 0.47 to 0.57mm long, with the tail wound three-quarters of the way around it. The adhesive organs are relatively deep FIG. 98. Aplidium ritteri — a, zooid (QM GH5779); b, larva (QM GH5649). Scales: a, 0.5mm; b, 0.1mm. 582 axial cones in epidermal cups. The base of their stalks expand into wide cones. Short median am- pullae alternate with them. The anterior end of the larval trunk is obscured by a long, wide lateral arcs of epidermal vesicles. REMARKS The specimens described above have all the characters of the type from Torres Strait, with the atrial lip from the upper rim of the opening, about 12 to 14 rows of 6 or 7 stigmata, 5 stomach folds, and furrows and ridges on the upper surface of the colony. The species has much in common with the Red Sea Aplidium lobatum Savigny, 1816 with which it often has been confused. The prin- cipal differences between these species are the larger branchial sac of A. lobatum, with up to 12 stigmata per row, and the division of the upper surface of the colony into more or less regular flat-topped, oval cushions, separated from one another by narrow furrows (see Michaelsen 1920). Aplidium macrolobatum n.sp. has a longer atrial lip, more stigmata per row, and median vesicles in the larva as well as lateral ones. In Australian waters the temperate and east- coast species Aplidium solidum has much in com- mon with the present one, having sandy colonies, the atrial lip from the upper rim of the aperture, 12 rows of up to 7 stigmata, 5 stomach folds and bunched male follicles in a short posterior abdo- men with a flattened straight-edged tip. A. ritteri has straight rows of zooids in contrast with the curved ones surrounding circular zooid-free ele- vation of the test in A. solidum. Although this appears to be a minor difference, the larvae con- stitute more compelling evidence that the species are separate — the larvae of A. solidum lack median ampullae but their trunk length (0.4 to 0.5mm) overlaps the range in the present species. Aplidium caelestis has similar sandy colonies, but more distinct long, double-row systems and zooids with the atrial tongue separate from the aperture. Aplidium robustum n.sp. (Fig. 99, Plate 16c) Aplidium flavolineatum: Kott, 1975, p.5. DISTRIBUTION TYPE LOCALITY: Western Australia (King George Sound, Albany, small breaking reef, south of Mt Mar- tin, 35°0.8’S 117°57.0’E, 2.0m, in small crevices in flat base rock, coll. AIMS Bioactivity Group 27.3.89, holo- type QM GH5467; paratype QM GH5506). MEMOIRS OF THE QUEENSLAND MUSEUM FURTHER RECORDS: South Australia (Great Australian Bight — SAM E2595 Kott 1975). Tasmania (Dunnal- ley, TM D1866). DESCRIPTION EXTERNAL APPEARANCE: Colonies are sessile, spherical or conical to 8cm high and 6cm in greatest (basal) diameter. The test is soft and gelatinous, but turgid and opaque, cloudy beige in preservative. In life the colony is pink around the common cloacal apertures and white over the zooids. Zooids are in circular, oval or elongate systems around and converging to the common cloacal apertures, which are numerous, large, ses- sile, and scattered randomly over the surface. They are said to ‘turn to slits’ when the colony is touched. Sand is present on the basal surface of the South Australian specimen, which is smaller than the others. The zooids are perpendicular to the surface but posteriorly criss-cross one another in the centre of the colony. A sheath of soft test containing opaque white corpuscles clings to the zooids. INTERNAL STRUCTURE: Zooids are robust, the contracted thorax and abdomen together being about 3mm and about equal in length. When relaxed the thorax probably is longer than the abdomen. About 15 longitudinal muscles are on the thorax and extend the length of the zooid. The 6 branchial lobes are small and round. The atrial lip from the upper rim of the aperture has a fleshy lip divided into 2 or 3 pointed lobes, or undivided with or without smaller lobes on each side. Stig- mata are in 14 to 16 rows of up to 12. The oesophagus is moderately long, and the relatively small stomach, in the middle third of the abdo- men, has 20 longitudinal, parallel folds in its wall. The duodenum, mid-intestine and posterior stom- ach are in the posterior third of the descending limb of the gut loop. There are rectal caeca at the proximal end of the rectum. Up to 3 embryos are in the atrial cavity of the type material, of which one may be a well-devel- oped tailed larva. The larval trunk is long (1mm), and almost cigar-shaped. The tail reaches to its anterior end, A lateral ampulla is on each side of the conical or cylindrical median ampullae that is in each interspace between the adhesive organs. The median ampulla between the middle and ventral adhesive organs is divided into 2. A clus- ter of epidermal vesicles is along each side of the mid-dorsal and mid-ventral lines, but not around the anterior end of the trunk. Up to 6 embryos are lined up in the atrial cavity of the South Australian specimen. The tailed lar- THE AUSTRALIAN ASCIDIACEA 3 vae are smaller (trunk 0.8mm long) and the lower median ampulla is not subdivided but otherwise they are similar to the larvae of the type speci- mens. REMARKS The species resembles A. opacum in having a sheath of soft test containing opaque white cor- puscles clinging to the zooids, large fleshy colo- nies, and the same number of stomach folds. FIG, 99. Aplidium robustum n.sp. (holotype QM GH5467) — a, thorax and abdomen; b, larva. Scales: a, 0.5mm; b, 0.2mm. 583 However zooids of the present species are more robust than those of A. epacum, with more rows of stigmata. They also have a larger larva with lateral ampullae and fewer vesicles than A. opacum. The present species has larger opaque white cells around the zooids and firmer test than A. multiplicatum, and it lacks the characteristic sin- gle arc of lateral vesicles in the larval trunk. In the distribution of epidermal vesicles this species resembles A. ornatum from tropical wa- ters, although the latter species is smaller than the present one and is a different shape. The smaller colony (A. flavolineatum: Kott, 1975) from South Australian waters with a layer of sand on its under surface and smaller larvae than the type may be found not to belong to the present species. However, at this stage it is pos- sible that its differences represent no more than intraspecific variation. It was assigned to A. fla- volineatum by Kott (1975) on the basis of its large number of stomach folds. However, the South African species has more numerous larval lateral ampullae and a brood pouch. Aplidium gastrolineatum has many characters in common with the present species, but its com- mon cloacal apertures are protuberant rather than sessile, and its larvae are different. Aplidium rosarium n.sp. (Fig.100) DISTRIBUTION TyPE LOCALITY: Queensland (Heron I., Blue Pools, rubble fauna, low tide, coll. P. Kott 8.8.82, holotype QM GH 5224), FURTHER RECORDS: None. DESCRIPTION EXTERNAL APPEARANCE: The firm, gelatinous colony is wedge-shaped, with a little sand embed- ded basally. When living the holotype was rose- purple, but in preservative it is white and cloudy, with beige-pink zooids. Zooids are along each side of long canals converging to the cloacal apertures. INTERNALSTRUCTURE: The zooids are robust, with up to 4 embryos crowded in the atrial cavity. The longitudinal thoracic muscles are strong and the thorax and abdomen of contracted zooids are together only 2mm. The 6 branchial lobes are triangular and conspicuous. The atrial lip is fleshy and arises from the upper rim of the aperture. It has a central band of longitudinal muscles that 584. extend into the tip, which is pointed or divided into 3 small terminal lobes. The stigmata are in 12 rows of about |2 per row. The short stomach wall has 20 folds, The poste- rior abdomen is about twice the length of the rest of the zooid. Larvae are of moderate size, the trunk being 0.6mm long, with 2 rows of stigmata in the larval pharynx, A single arc of about 20 vesicles is along each side of the median line around the anterior nse! SOR GSCOOE FIG, 100: Aplidium rosarium n,sp. (holotype QM GH5224) — a, zooid; b, larva. Scales: a, 0.5mm; b, 0.Jmm. MEMOIRS OF THE QUEENSLAND MUSEUM end of the trunk. In immature larvae, a few me- dian vesicles branch off the tip of the median ampulla which is in each interspace between the adhesive organs, The ampullae subsequently dis- appear and the stalk of each median vesicle ap- pears to be independent, and attached directly to the larval ectoderm. REMARKS Aplidium ornatum has a similar gelatinous col- ony, butits branchial sac is narrower and its large larvae have posterior vesicles and median and Jateral siripilise, and lack the anterior arc of lat- eral vesicles that is present in A, rosarium. The larvae resemble those of A. filiforme, but colonies differ, and the zooids have more rows of stigmata, The present species has some affinities with A. aliarium but its larvae are smaller, the pharynx narrower and more muscular, and the stomach smaller with fewer folds. Aplidium uteute has circular systems, a brood pouch and more numer- ous stomach folds distinguishing it from the pre- sent species. Some aspects of the larvae suggest a relationship with A. incubatum, however the larval trunk of the latter species is larger, has more median vesicles and, the zooids have more stig- mata and stomach folds, and are arranged in cir- cular systems. Thus, despite the fact that only the holotype is available, the species is readily distin- guished from other known species. Aplidium rubricollum Kott, 1963 (Fig. 101. Plate 16d) Aplidium rubricollum Kott, 1963, p. 103 (part, not specimens from Western Australia AM Y1403 <7? A, solidium), 1972a, p.15; 1972b, p.176. DISTRIBUTION New Recorps: South Australia (Avoid Bay, QM GH4191; Port Noarlunga, QM G9304). PREVIOUSLY RECORDED: South Australia (Reevesby !. — AM Y1417 holotype Kott 1963; Upper St. Vincent Gulf — SAM E2579 Kott 1972a; Pearson I. — Kott 1972b). Balnarring Beach — AM Y1415 Kott 1963). DESCRIPTION EXTERNAL APPEARANCE: The colonies range from small cushions inyesting weed or rubble to regular oval cushions to Scm in maximum extent but never more than 1em thick. Sand is embedded in the internal test, becoming sparse in the upper layer around the thoraces. Sometimes it is com- pletely absent from the upper surface of the col- ony, but in other colonies only the test above the THE AUSTRALIAN ASCIDIACEA 3 systems, including the protruding common cloa- cal apertures, is naked. Zooids are crowded along each side of relatively short canals that converge to the protruding cloacal apertures which are scat- tered over the surface, about 5mm apart. Red pigment cells are said to have been present in the surface layer of test in the living and freshly preserved specimens, and the colourless zooids are conspicuous, interrupting these. In the long- 585 preserved specimens a brownish deposit in the surface test may be the remains of these red cells. INTERNAL APPEARANCE: Zooids are small, with 10 fine longitudinal muscles. The atrial lip has an undivided tip which becomes bidentate through the contraction of a median band of mus- cles. The lip arises from the body wall close to, but separate from the opening, which is on a short protruding siphon, often flattened antero-posteri- Wy Bn oo ath mu | \ Swag } wilh / _ FIG, 101; Aplidium rubricollum —a, surface of colony showing 2 systems (QM GH4191); b, zooid (holotype AM Y1417); ¢, larva (holotype AM Y 1417). Scales: a, 2mm; b, 0.5mm; c, 0.2mm. 386 orly. Stigmata are in 11 or 12 rows of up to 14. The stomach is short with 5 deep folds, One specimen collected in February (QM G9304) and the holotype collected in December contain single, large embryos in the atrial cavity. The laryal trunk is 0.7mm long. It has median and lateral ampullae, but no epidermal vesicles. REMARKS ‘The systems and the colonies resemble those of A. clivosum except that the canals converging to the cloacal apertures are not so numerous, the zooids and systems are smaller, and the larvae Jack the vesicles that obscure the anterior end of the trunk in A, chivesum. The zooids are smal] and. like those of A. clivosuim, have the atrial aperture separate from the opening, but the present species has relatively shorter and wider thoraces with up to 12 rows of 14 stigmata while A. clivesum has up to 18 rows of about 12 stigmata. Colonies assigned to Aplidium caelestis (sec above) often superficially resemble those of the present spe- cies, However, A. caelesris has a narrower thorax with fewer stigmata per row, the systems have longer double rows of zooids and the Jarvae con- tain epidermal vesicles. Aplidium solidum (Herdman, 1899) (Fig. 102) Pecmneaplidions soliduen Herdmun, 1899, p85. Psammaplidium lobatumn: Herdman, 1899, p85, Psammaplidiam fragile Herdman, 1899, p.86 Psammaplicdium incrustans Herdman, 1899, p.87, Aplidiam solldum Millar. 1963, p-1. Aplidium arboratum Kott, 1963, p,96. Syroleam investi Kort, 1963, p.90, Aplidium lobaium; Kott, 1976, pel. ? Aplidium rubricallum Kott, 1963, p.103 (part, speci- men from Rottnest 1. AM Y1403). Not Amaroucium solidur Ritter and Forsyth, 1917, p.486; Van Name, 1945, p.49, The species name is a secondary homonym and accordingly a nent. nov. is required for this species. DISTRIBUTION New RECORDS: Western Australia (Dampier Archi- pelago, QM GH5411). New South Wales (Jervis Bay, QM GH5778; Batemans Bay, AM ¥2278; Bass Point, QM GH5620; Arrawarra, QM GH5777). Queensland (Point Lookout, QM GH5782). PREVIOUSLY RECORDED: New South Wales (Botany Bay — AM 3941 Herdman 1899; Millar 1963). Queensland (Bargara — hololype AM U3928, paratype QM G4937 S. investum Kott, 1963). MEMOIRS OF THE QUEENSLAND MUSEUM DESCRIPTION EXTERNAL APPEARANCE: The colonies form extensive rigid, sandy sheets, lobes and Jamellae from about 3mm to 2cm thick. These branch and fuse with one another in a vertical plane, Along their upper border the lamellae terminate in rounded margins. The upright Jobes and lamellae form when the sheet is folded, fuses back to back, and extends up in the vertical plane. Sand is embedded throughout, Zooids appear to be fairly evenly placed and clase ta one another, along the sides of narrow, shallow canals. Circular or oval, raised zooid-free anrcas sometimes are between (or surrounded by) the double rows of zooids. Common cloacal openings are scattered over the surface of the colony at the junction of the canals. The colour of the colonies is dominated by the sand, although the zooids are pinkish in preservative. INTERNAL STRUCTURE: The zooids are small, seldom more than 4mm evenin relaxed condition. Thorax, abdomen and posterior abdomen are of about equal length. Fine Jongitudinal muscle bands extend the length of the zooid, A sphincter muscle is at the base of the 6 pointed branchial lobes. The small, circular atrial aperture has a small lip, undivided or divided into 2 or 3 pointed lobes, from the anterior rim of the opening. Stigmata are in 9 to 12 rows of about 6. The long, narrow stomach has 5 folds. The posterior stomach is large, oval, at the distal end of the descending limb of the gut loop. The posterior abdomen is relatively short, with a single series of up to 7 male follicles in a single longitudinal seres of sometimes bunched up behind the ante- ner ovary. However, even when it is not con- Iracied, the extremity of the posterior abdomen has a truncated flat appearance. A small ovary is often present anterior to the testis follicles, Two to 5 embryos are crowded into the atrial cavity of specimens from Botany Bay from Jervis Bay in February (QM GH5778) and from South Australia in April (QM GH4165), Larvae are small, the trunk 0,4 to 0.5mm Jong with the tail wound half to three- quarters of the way around it. A mass of epidermal vesicles surround the 3 median adhesive organs and obscure the anterior end of the trunk, but there are no median ampul- lac. The base of staiks of the adhesive organs expand into a cone. REMARKS Like the present species, Aplidivin deroporuin n.sp. has small! zooids wath the atnal lip from the rim of the opening. However it has 8 stomach THE AUSTRALIAN ASCIDIACEA 3 folds (rather than 5), and circular systems with zooids surrounding the cloacal cavity (rather than along each side of narrow canals), The specimen from Rottnest 1. appears to have been wrongly assigned to A, rubricollum by Kott (1963). Both colony and zooids resemble those of 587 the present species except in the colour (pink with red zooids), which they have retained over a long time (40 years) in preservative. The colour of the living colony is not known. Although brown to orange living specimens of A. solidium are known the colour of the embedded sand dominates their x Ny, Mi} 5 OO NM a ay Wy Te Hoy n0dy jong 0010 on 000g FIG, 102: Aplidium solidum — a, b, parts of colonies (QM G4937, AM U3941); ¢, surface of colony showing cloacal apertures at junction of the cloacal canals that surround the small zooid-free test areas (QM GH5782); d, e, zooids (QM GH5777 GH5782); f, larval trunk (QM G4937). Scales: a, b, 2em; ¢, 2mm; d, e, 0.5mm; f, 0.1mm. 588 MEMOIRS OF THE QUEENSLAND MUSEUM THE AUSTRALIAN ASCIDIACEA 3 colour in preservative. Therefore, the specimen from Western Australia is unusual and despite similarities in its morphology il may not be con- specific with A, solidus. The sandy investing colonies resemble some of Aplidium caelestis, although cloacal canals of the present species surround circular areas rather than the long ridges of A. caelestis, which also has colonies that are thicker and do not fold back (0 form upright lamellae as they do in the present species. Further A. caelestis, has zooids with longer branchial siphons, the atrial lip separate from the opening, and testis follicles in a double series in a long (rather than short, truncated) posterior abdomen. The species resembles Aplidium ritfert in its sind-filled colonies, small zooids with narrow thoraces, small atrial lip from the upper nim of the opening: and short, truncated and flattened tip of the posterior abdomen, The systems of A. ritteri are in long and relatively straight double rows, and the larvae have a trunk 0.5 to 0.4mm long, with median ampullae as well as lateral vesicles, Aplidium petrosum from South Australian waters hus similar systems as well as similar zooids to the present species, but is distinguished by its more complex colonies and larger larvae with median ampullae, Aplidium tabascum n.sp. (Fig. 103. Plate 1 6¢,f) DISTRIBUTION TYPF LOCALITY: Queensland (Heron L reef, 15m, coll. N. Coleman 5.8.77, AMPI 207, holotype QM GH5208; Heron I, reef, 9m, coll. N. Coleman 16.7.73, AMPI 85, paratype QM G9726: coll. P. Fredrickson, paratype QM GH5542; Erskine I. 5—10m-. coll. Roche (SLY 12) 21.3.76, paratype QM G9727). FURTHER RECORDS: Queensland (Capricorn Group, QM G9725 GH5513 GH5543; Swain Reefs, MV). DESCRIPTION EXTERNAL APPEARANCE: The colonies form large fleshy mats or sheets to 7cm or more in maximum dimension, The surface has a ‘mar- bled appearance owing to irregularly shaped zo- oid-free areas of test, between and around which are slightly depressed areas where a relatively thin layer of surface test covers the cloacal canals. Sparse but evenly distributed sand grains are in the zooid-free solid parts of the test and sand 589 grains are crowded inte a line along the margins of these areas where zonids line the cloacal canals and the branchial apertures open to the exterior. Sand is also crowded around the margins and in the base of the colony, but is not present on the upper surface either over the common cloacal canals or the zooid-free areas they surround, Large common cloacal apertures are in the de- pressed areas over the common cloacal spaces and appear to be randomly distributed. Sometimes the solid test areas are isolated from one another, forming islands surrounded by com- mon cloacal cavities and zooids, In other speci- mens they are long and curved and meander through the colony between and around the cloa- cal spaces. The colonies are a bright red-orange colour in life. Although a trace of the orange pigment at first persists in the preserved colony, sth- sequently they are white. In fresh material, the red igment is in minute crowded spherical cells in the test and the zooids but these are not present in preserved specimens, INTERNAL STRUCTURE: Zooids are small and narrow, 2 or 3mm Jong, although when contracted they are shorter with the posterior abdomen bunched und drawn up alongside the gut loop. The 6 branchial lobes around each branchial ap- erture have bands of muscles which sometimes are drawn in to subdivide the tip of each lobe. The atrial ip from the upper rim of the aperture is undivided or divided into 2 cr 3 pointed lobes of varying size, Fine longitudinal muscle bands are om the thorax, and extend the length of the abdo- men and posterior abdomen in a wide ventral band. The stigmata are in 15 rows with up to 8 per row, The stomach wall has 3 folds. The gut loop is relatively shor. The posterior abdomen is club- shaped when the zooid is relaxed, with a long neck anteriorly and about 8 large male follicles in a single series in the expanded posterior part. When contracted, however, the testis follicles are bunched, and the posicrior abdomen is drawn up alongside the gut loop. An ovary with 2 or 3 large oocytes is anterior to the testis follicles. One or 2 embryos are in the atrial cavity of zooids collected in July and August (QM G9726 GH5208 GH5513) but none in March or Decen)- ber. The larval trunk is oval, about 0.6 to 0.8mm long. with the tai] wound the whole way around it. On each side of the median line dorsal and ventral clusters of epidermal vesicles extend pos- FIG, 103: Aplidium tabatcunt n.sp, — a. surface of colony showing almost continuous line of sand particles around each area of zooid-free test containing sparse embedded sand (QM GH5513); b, zooid (gut loop twisted to the left) (QM GH5208); c, sexually immature zooid (QM GH9725); d, thorax showing large bilobed atria! lip (QM GH5208); e, larva (QM GH5208), Scales: a, lem; b-d, 0.5mm, e. 0.2mm. 590 teriorly from opposite the base of the dorsal and ventral adhesive organs, respectively. These dor- sal and ventral bands of vesicles are not continu- ous at the anterior end of the trunk. Neither vesicles nor ampullae are present in the median line between the adhesive organs. One larva found in the holotype had 5 adhesive organs instead of the usual 3. REMARKS The present species, based on a significant number of specimens, has a striking appearance with apparently stable characters showing little or no variability. It is distinguished by its double rows of zooids opening into depressions sur- rounding cushion-like elevations on its naked upper surface, the capacity to sort sand particles and arrange them around the margin of the zooid- free surface elevations, its red-orange colour, nar- row thoraces, and the interruption of the band of lateral larval vesicles. The same sort of cloacal systems, and the ca- pacity to sort embedded sand particles and ar- range them in the colony are properties of other Aplidium spp. (e.g. A. caelestis, A. lenticulum n.sp.). Aplidium lobatum Savigny, 1816 from Suez and A. cellis Monniot, 1987 from New Caledonia also have similar cloacal systems. Aplidium caelestis and A. lenticulum differ in having an atrial lip separate from the opening but the Aplidium cellis and A. lobatum have similar colonies, zooids and larvae to those of the present species, differing only in their embedded (and sometimes encrusting sand), smaller larval trunk (0.56mm long), and colour (white rather than red). Despite the description of A. cel/lis, the re- ported incrustation of sand on the upper surface (slightly less densely distributed over the cloacal canals) is not apparent in the published photo- graph (Monniot, 1987, Pl. 2B). Further, it is not clear whether the white colour recorded for the 2 New Caledonian specimens is their living colour, or their colour in preservative, or merely the colour of the sand encrusting them. Thus, in ad- dition to the possibility that the New Caledonian species is a junior synonym of A. lobatum, A. tabascum could be as well. Aplidium triggsense Kott, 1963 (Fig. 104) Aplidium triggsensis Kott, 1963, p.104; 1976 (A. trig- giensis) p. 61. Monniot, 1987, p.531. MEMOIRS OF THE QUEENSLAND MUSEUM DISTRIBUTION: NEw RECORDS: Western Australia (Cockburn Sound WAM 23.84). South Australia (Pearson I., SAM E2574). Queensland (Heron I., QM GH5305). PREVIOUSLY RECORDED: Western Australia (Triggs I. — AM U3923 holotype, Y 1408 Kott 1963; Rottnest I. — AM Y1421 Y1428 Kott 1963; Nornalup — AM Y 1405 Kott 1963). Victoria (Balnarring Beach — AM Y1427 Kott 1963; Westernport — Kott 1976). New Caledonia (Monniot 1987). The Western Australian records are all from coastal habitats subjected to surf and turbulence, and the Westernport location is one where strong tidal currents prevail. A continuous temperate to tropical range is suggested by the record from Heron I. DESCRIPTION EXTERNAL APPEARANCE: One colony from Heron I. (QM GHS5305) is an upright lobe, but usually the colonies are firm, gelatinous, invest- ing sheets 0.5 to 0.6cm thick and 4 to 5cm in maximum extent, Varying quantities of sand are present in the basal half of the test but not in the transparent upper half. This strengthens the col- ony. The margins are irregular and reduced in thickness. Root-like projections from the base of the colony extend into and around particles of the substrate helping to anchor the colonies, which are found in rigorous, turbulent habitats. Zooids are crowded, conferring a reddish-pink colour on the preserved colonies. The living colony from Heron I. is reported to have been orange buff, with translucent test. Circular cloacal apertures are randomly distrib- uted over the surface. It is probable that the zooids are in double-row systems but this could not be confirmed in these crowded colonies. The poste- rior parts of the zooids criss-cross one another in the basal half of the colony. INTERNAL STRUCTURE: Zooids are small, the contracted abdomen and thorax together being less than 3mm, although the thin posterior abdo- men is up to twice that length. The branchial aperture is small, with a short siphon. The atrial aperture is a small, rounded, sessile opening with a small pointed or tridentate tongue from the upper border of the opening. The thorax has 12 wide longitudinal muscles and these extend the length of the abdomen. Stigmata are in 8 to 10 rows of up to 15. The stomach is short with 15 longitudinal folds. Testis follicles are in 2 longi- tudinal series, each of about 6 follicles. One or 2 embryos are in the atrial cavity of specimens collected from Cockburn Sound in THE AUSTRALIAN ASCIDIACEA 3 January, Pearson I. in November, Heron I. in No- vember, and Westernport (Kott 1976) and Western Australia in December. The larval trunk is 0.6mm long, and the tail is wound almost halfway around it. From each location larvae have from 7 to 14 small adhesive organs. These are on fine stalks expanding into cones at their base, and are in the mid-line 591 around the anterior half of the trunk. Scattered epidermal vesicles arise in an arc from the lateral line on each side of the adhesive organs. REMARKS The species is distinguished by its firm colonies with sand in the basal half, its small zooids, 15 FIG. 104: Aplidium triggsense — a, b, zooids (WAM 23.84, QM GH5305); c, larva (AM ¥1428). Scales: a, b, 0.5mm; ¢, 0.2mm. 592 stomach folds and remarkable larvae which, with 12 adhesive organs, seem adapted to make a firm adherence to the substrate in the stringent habitats from which it has been taken. The colonies from 8cm in the lagoon, New Caledonia (Monniot 1987) are flattened lobes, joined basally, similar to the newly recorded up- right lobe from Heron I. This was previously regarded as a temperate species, but tropical records demonstrate a wider geographic range. The alternate possibility that different species are represented is not supported by the morphology of either colonies, zooids or larvae, which are all similar, with only small intrapopulation variations in size, number of stig- mata, stomach folds, and adhesive organs. Variation from the usual number (3) of adhe- sive organs occurs in A. multipapillatum Millar, 1975, from the China Sea, which has 7. It is probable that the large number in this apparently unrelated species is a convergent character, se- lected for in response to environmental pressures. Aplidium uteute Monnniot and Monniot, 1987 (Fig. 105) Aplidium uteute Monniot and Monniot, 1987, p.79. Aplidium phortax: Kott, 1963, p, 109. Aplidium latusexitus Monniot, 1987, p. 523. DISTRIBUTION New RECORDS: Queensland (Capricorn Group, QM GH1342 GH5132-48 GH5220-3 GH5313 GH5510 GH5524—-5 GH5598-601 GH5610 GH5621-6). FURTHER RECORDS: Queensland (Sarina — Kott 1963). Western Pacific (Solomon Is — Kott 1963; New Caledonia — Monniot 1987; French Polynesia — Monniot and Monniot (1987). DESCRIPTION EXTERNAL APPEARANCE: The colonies form flat-topped cushions, to sheets fixed by a large part of the under surface, or they reduce in diame- ter toward the base to form flat-topped mush- room-like colonies about lcm in maximum diameter and 0.5cm high, fixed by a small part of the under surface. The test is firm, clear and transparent in life, honey-coloured, ‘rufous’ (Ridgeway 1886), or colourless. It is only rarely cloudy, although it usually does become so in preservative. The living zooids, seen through the clear, transparent test, are red, ‘chrome scarlet’, ‘buff-yellow’, ‘orpiment orange’, “salmon-col- oured’, cream with orange stomachs, or with ‘or- ange chrome’ thoraces and flesh-coloured MEMOIRS OF THE QUEENSLAND MUSEUM branchial apertures, with ‘geranium-coloured’ or ‘dragons-blood red’ abdomina, and often a ‘drag- ons-blood red’ patch each side of the base of the atrial lip or between the apertures (Ridgeway 1886). Sometimes 4 lines of dark orange pigment radiate out from the branchial aperture, including one along each side of the endostyle. Embryos when these are present, are ‘saturn red’, “burnt carmine’ (Ridgeway loc. cit.) or maroon. In pre- servative the test is yellowish and translucent and the zooids white or cream. The zooids are ar- ranged in conspicuous circular systems of 7 to 10 zooids. Sand is often present around the outer margin of the colony, but is never embedded internally. INTERNAL STRUCTURE: Zooids are up to 8mm long in semicontracted condition. They have large, rounded, sometimes subdivided branchial lobes and a large, fleshy, sometimes 2-lobed atrial lip from the upper border of the atrial aperture, which also has lobes around its posterior margin. Longitudinal muscles on the thorax are fine, and numerous (about 20), extending the whole length of the zooid in a wide ventral band. There are 9 to 11 rows of up to 25 stigmata in the wide thorax. The thorax is large probably exceeding the length of the posterior abdomen when it is not con- tracted. In preserved specimens there often is a crescent-shaped patch of greenish coagulated blood near the top of the abdomen, probably an artefact associated with the removal of the colony from the substrate. The stomach is large and cylindrical with from 25 to 35 fine parallel longitudinal folds. The duodenal area and mid-intestine are continuous, and open into the oval posterior stomach at the base of the descending limb. A rectal valve is at the junction of the distal part of the mid-intestine and rectum at the end of the abdomen. An ovary with up to 5 oocytes is about one-third of the distance down the posterior abdomen, The pear-shaped male follicles are posterior to the ovary, sometimes bunched but in other specimens spread out in an irregular longitudinal series. The posterior abdomen in this species is relatively short, often stumpy and only occasionally more than twice the length of the rest of the zooid. Usually one or 2, but occasionally up to 5, embryos are in a brood pouch that projects out from, and is constricted off from, the posterodor- sal corner of the thorax in zooids collected at Heron I. in May to August. The tailed larvae have a 0.65 to 0.8mm long trunk, and are deep some- times almost spherical. The tail is wound two- thirds of the way around the trunk. Three or 4 fine, THE AUSTRALIAN ASCIDIACEA 3 593 OOM 0009009 &VIQdY FIG, 105: Aplidium uteute — a, surface view of colony indicating systems (QM GH5134); b, zooid with embryos in brood pouch (QM GH5145); e, larva (QM GH5623). Scales: a, 5mm; b, 0.5mm; ¢, 0.1mm. 5394 sometimes branched, stalks with lerminal ec- todennal vesicles extend out from the anterior mid-line in each interspace between the adhesive organs; and 20 vesicles develop in the same way from each lateral line around the anterior end of the larva. There are 3 rows of stigmata and a large ocellus and otolith in the cerebral vesicle. REMARKS The characteristics of this species are its trans- parent test, conspicuous circular systems, wide thorax, relattvely Jong cylindrical stomach with numerous folds, postero-dorsal thoracic brood pouch, deep larval trank with median epidermal vesicles alternating with the adhesive organs, and arc of about 20 vesicles along each lateral line. Stomachs are orange, some red pigment in the zooids, embryos and laryae are maroon-coloured, The Australian material appears to be conspe- cific with Aplidium ureute Monniot and Monniot, 1987 from French Polynesia which has a similar stomach, large atrial aperture with the upper rim produced into a large lip, embryos incubated in a brood pouch constricted off from the thorax, red pigment each side of the neural complex, and apparently circular systems, The number of rows of stigmata are the same but unfortunately the number per tow is not recorded for the Polynesian material, The number of stomach folds. of the Polynesian type is said to be 50, but the number shown in Fig. 28 (Monniot and Monniot 1987) appears to be no more than 35, Monniot and Monniot (1987, Planche V-H) compared the spe- cies with a range of probably unrelated specimens assigned by various authors (Tokioka 1953, 1967a; Kott 1972, 1975; Nishikawa 1984a) to A. pliciferum Redikorzey, 1927 from the north- westem Pacific. Aplidium ureute is readily distin- guished from these by many characters. However, there are others that appear to have closer affinities with which it was not compared. Aplidium latusexitus Monniot, 1987 has circu- lar systems, and, like A. wtewte, red pigment patches each side of the neural gland, a large atrial aperture with a large lip, 10 rows of sligmata with up to 28 per row and lateral as well as: median epidermal vesicles in the larva. Monniot (1987) MEMOIRS OF THE QUEENSLAND MUSEUM has discussed its relationship with A. mudriplica- tum ( A. californicum: Monniot 1987), but not with A. uteure with which it appears to be synony- mous. Aplidium uteute is readily separated also from A. multiplicatum by its larvae, the form of the systems, the number of stigmata per row (i.e. the width of the thorax), and the consistency of the test. Kott (1963) assigned specimens of the present species to Aplidinm phortax (Michaeisen, 1924) from New Zealand. However, despite similarities in the colonies and brood pouches, A. ureute has a wider thorax than the New Zealand species which has only 9 or LO stigmata per row, The larva of A. phortax is not known. Aplidium flavolineatum (Sluiter, 1898) from South Africa (see also Millar 1962) has zooids arranged in conspicuous round to oval systems, and embryos incubated in a brood pouch slightly constricted off from the thorax. Michaelsen (1934) thought it was synonymous with A, phor- tax, however A. flavolineatum differs from both A. phortax and the present species by having embedded and encrusting sand, generally more stomach folds (although the numbers overlap, and the stomachs are a similar shape) and by usually being a ted colour m life. Also, A. fla- volineatum incubates up to 4 embryos in the brood pouch and has a Jarval trunk of 1mm with median ampullae and some large lateral vesicles or ampullae (Millar 1962); while A. utente has a larval trunk less than 0.8mm long with more numerous epidermal vesicles. Aplidium flavolineatum: Monniot, 1987, has circular systems, brood pouch, and stomach re- sembling those of the present species. However the Jarvae, found in the single colony that Mon- niot described, have fewer epidermal vesicles than A. ureute. These vesicles, like those of A. altariam remain attached to the trunk epidermis. Aplidium altarium also has a large number of stomach folds, but is distinguished by its con- verging double row systems. Aplidium ornaivm is another species with large numbers of stomach folds, but its larvae lack the long ares of eyenly spaced epidermal vesicles that charactense the larvae of the present species. THE AUSTRALIAN ASICIDIACEA 3 595 PLATE 1: a, Monniotus australis (QM GH4147 Price I., SA). b-d, Condominium areolatum n.gen. n.sp. (b, QM GH4161 Taylor I. SA; c, d, QM GH5418 central Great Barrier Reef Qd 20m). e, f, Ritterella asymmetrica (e, QM GH4195 Hopkins I. SA; f, QM GH5445 Cape Jaffa SA). Photos: a, e, S.A. Shepherd; b, W.H. Sasse; c, d, f, AIMS Bioactivity Gp, Q66-C1723 -C2457. 596 MEMOIRS OF THE QUEENSLAND MUSEUM f PLATE 2: a, b, Ritterella compacta n.sp. (a, QM GH2399 holotype Flinders I. SA; b, QM GH4177 The Gap SA). ¢, Ritterella cornuta n.sp. (QM GH4176 holotype Price I. SA). d, Ritterella dispar (QM G12001 Lord Howe I.). e, Ritterella multistigmata n.sp. (QM GH5463 holotype Rockingham WA). f, g, Ritterella peduncu- lata (f, QM G10156 George’s Bay Tas. 5m; g, QM G10160 Phillip Bay Vict. 10m). Photos: a, N. Holmes FL975-PE0030; b, c, W.H. Sasse 9.4.87 7.4.87; d, f, g, N. Coleman AMPI 250 211 170; e, AIMS Bioactivity Gp Q66-C2822. THE AUSTRALIAN ASICIDIACEA 3 597 PLATE 3: a, Euherdmania translucida n.sp. (QM GH2310 Flinders I. SA). b, Pseudodistoma acuatum n.sp. (QM GH2323 holotype Ward I. SA). c, d, Pseudodistoma australe (QM G92 Exmouth Gulf WA). e, f, Pseudodistoma candens n.sp. (QM GH5462 Rockingham WA). Photos: a, b, N. Holmes FL985-PE0064; c, d, N. Coleman AMPI 76 77; e, f, AIMS Bioactivity Group Q66-C2798. 598 MEMOIRS OF THE QUEENSLAND MUSEUM PLATE 4: a-e, Pseudodistoma gracilum n.sp. (a, QM G1 1989 Kangaroo I. SA; b, QM GH4158 Cathedral Rock SA; ¢, Kangaroo I. SA; d, QM GH4167 paratype Grindal I. SA; e, ? SA). f, Pseudodistoma inflatum n.sp. (QM G9472 paratype Byron Bay NSW). Photos: a, ¢c, f, N. Coleman AMPI 227 223 158; b, d, W. H. Sasse 23 19; e, R. Kuiter 47SS. THE AUSTRALIAN ASICIDIACEA 3 599 PLATE 5: a, Pseudodistoma oriens n.sp. (Port Phillip Heads Vict.); b-d Pseudodistoma pilatum n.sp., showing colour variants (b?; c, SAM E2409 paratype Cathedral Rock SA; d, QM GH4140 paratype Golden I. SA). e, f, Pseudodistoma pulvinum (e, QM GH2396 holotype Ward I SA, in caves; f, QM GH2300 Flinders I, SA).Photos: a, J. Watson April 1977; b, R. Kuiter 54SS; ec, d, W.H. Sasse 20 10; e, f, N. Holmes FL979-PE0012 FL964-PE0060. 600 MEMOIRS OF THE QUEENSLAND MUSEUM PLATE 6: a, b, Polyclinum incrustatum (a, QM GH4141 Perforated I SA; b, The Gap, QM GH4163). ec, d, Polyclinum marsupiale (ec, QM G10138 Kingston Jetty SA; d, QM G10175 Deal I. Bass Strait)). e-g, Polyclinum nudum n.sp. Coffs Harbour NSW, showing colour variants (e, QM GH5387; f, QM GH5388; g, QM GH5390). Photos: a, b, W.H. Sasse 34 3; ¢, J.E. Watson September 1977; d, N. Coleman AMPI 114 6.5.74; e-g, AIMS Bioactivity Gp Q66-C1108 -C1129 -C1131. THE AUSTRALIAN ASICIDIACEA 3 601 PLATE 7: a, Polyclinum tenuatum n.sp. (QM GH5444 Cape Jaffa SA). b, ¢, Polyclinum terranum n.sp. QM 5460 holotype Rockingham WA (b, distant view showing top of colony exposed on surface of substrate; ¢ close up). d, Aplidiopsis mammillata n.sp. QM GH1468 holotype Cathedral Rock SA). e, Aplidiopsis sabulosa n.sp. (QM GH4153 holotype Avoid Bay SA). f, Sidneioides tamaramae (QM G11859 South Ulladulla NSW). Photos: a, b, c, AIMS Bioactivity Gp Q66-C2455 -C2794; d, e, W.H. Sasse; f, I. Bennett 27.1.87. > 602 MEMOIRS OF THE QUEENSLAND MUSEUM g | PLATE 8: a, Synoicum bowerbanki (Blacknose Point Portland Vict.). b, Synoicum buccinum n.sp. (QM GH5045 holotype Marion Reef, Coral Sea). c-e, Synoicum castellatum n.sp. (¢, QM GH5401 Exmouth Gulf WA; d, Great Keppel I. Qd 10m; e, QM GH5322 Lizard I. Qd). f, Synoicum citrum n.sp. (QM GH2294 Port MacDonnell SA). g, Synoicum concavitum n.sp. (QM GH5465 holotype King George Sound WA). Photos: a, J.E. Watson June 1979; b, d, N. Coleman AMPI 209 243; c, e, g, AIMS Bioactivity Gp Q66-C1362 -B2069 -C2889; f, N. Holmes PM1193-PE0073. THE AUSTRALIAN ASICIDIACEA 3 603 PLATE 9: a, Synoicum macroglossum (QM GH5421 Mackay Qd). b, Synoicum prunum (QM GH5394 Arrawarra NSW). c-e, Synoicum sacculum n.sp. (ce, d, QM G10163 holotype Flinders Vict. 5m; e, Tipara Reef SA). f, g, Synoicum suarenum n.sp. (QM GH5377 Hardy Reef Qd). Photos: a, b, f, g, AIMS Bioactivity Gp Q66-C1880 -C1175 -C0872; c, d, N. Coleman AMPI 201; e, S. Shepherd 13.5.82. 604 MEMOIRS OF THE QUEENSLAND MUSEUM PLATE 10: a, b, Aplidium acroporum n.sp. (a, QM GH5443 paratype Kangaroo I. SA; b, SAM E2554 holotype The Gap SA). c, d, Aplidium altarium (c, QM GH5557 Houtman’s Abrolhos WA; d, QM GH5522 Heron I. Qd). e, Aplidium bacculum n.sp. (QM GH5434 holotype York Peninsula SA). f, g, Aplidium brevilarvacium (f, ? SA; g, QM GH2412 Ward I. SA). Photos: a, e, AIMS Bioactivity Gp Q66-C2445 -C2304; b, W.H. Sasse 14; c, d, N. Coleman AMPI 60 150; f, R. Kuiter 43SS; g, N. Holmes WD894-PE0020. THE AUSTRALIAN ASICIDIACEA 3 605 PLATE 11: a-f Aplidium caelestis (a, b, whole and part of colony respectively QM GH5430 Kangaroo I SA; ec, d, whole and part of colony respectively QM GH5437 Edithburgh SA; e, QM GH5428 Kangaroo I. SA; f, Grindal I. SA), Photos: a-e, AIMS Bioactivity Gp Q66-C2186 -C2317 -C2184; f, W.H. Sasse 6. 606 MEMOIRS OF THE QUEENSLAND MUSEUM PLATE 12: a-f, Aplidium clivosum n.sp. (a, QM GH2417 Hotspot SA; b, QM GH5469 Geographe Bay WA; e, QM GH5447 Beachport SA; d, ? South Australia; e, QM GH5628 Jervis Bay NSW; f, QM GH5305 Flinders I. SA).Photos: a, f, N. Holmes HS1035-PE0008 FL792-PE0050; b, c, AIMS Bioactivity Gp Q66-C2928 -C2493; d, R. Kuiter SS51; e, N. Coleman AMPI 78. THE AUSTRALIAN ASICIDIACEA 3 607 PLATE 13: a-c, Aplidium crateriferum (a, Heron I. Qd; b, QM GH5406 Montebello I. WA; c, Heron I. Qd). d-f, Aplidium lenticulum n.sp. (d, QM GH5457 Rockingham WA; e, QM G9475 Camac I. WA; f, Denmark WA). Photos: a, ¢, e, f, N. Coleman AMPI 146 197 46 119; b, d, AIMS Bioactivity Gp Q66-C1548 -C2780. 608 MEMOIRS OF THE QUEENSLAND MUSEUM PLATE 14: a-c, Aplidium lunacratum nom. nov. (a, QM GH5429 Kangaroo I. SA; b, c, QM GH5446 Cape Jaffa SA). d-f, Aplidium multiplicatum (d, QM G10151 Wistari Reef Qd, with Prochloron on surface and in cloacal canals; e, Heron I. Qd; f, QM GH5361 Darwin Harbour NT). Photos: a, b, ¢, f, AIMS Bioactivity Gp Q66-C2185 -C2458 -C0635; d, e, N. Coleman AMPI 160, Nov. 1987. THE AUSTRALIAN ASICIDIACEA 3 609 PLATE 15: a-e, Aplidium opacum (a, QM GH5440 Edithburgh SA; b, e, Port Hacking NSW 3m, 5m). d-f, Aplidium petrosum n.sp. (d, e, QM GH5454 Cape Jaffa SA; f, QM GH4165 The Gap SA). Photos: a, d, e, AIMS Bioactivity Gp Q66-C2376 -C2728; b, c, N. Coleman AMPI 255; f, W.H. Sasse 32. 610 MEMOIRS OF THE QUEENSLAND MUSEUM PLATE 16: a, b, Aplidium protectans (a, QM GH5378 central Great Barrier Reef; b, QM GH5336 far northern Great Barrier Reef). ¢, Aplidium robustum n.sp. (QM GH5467 holotype King George Sound WA). d, Aplidium rubricollum (QM GH4191 Golden I. SA). e, f, Aplidium tabascum n.sp. (e, QM GH5208 holotype Heron I. Qd; f, QM G9726 Heron I. Qd). Photos: a-c, AIMS Bioactivity Gp Q66-C0897 -B2177 -C2914; d, W.H. Sasse 14; e, f, N. Coleman AMPI 207 85. THE AUSTRALIAN ASCIDIACEA 3 LITERATURE CITED ABBOTT. D.P. and TRASON, W.B. 1968. Ritterella rubra and Distaplia smithi: wo new colonial ascidians from the west coast of North America. Bulletin, Southern California Academy of Sci- ences 67(3): 143-154. BERRILL, N.J, 1935. Studies in tunicate development IV, Asexual reproduction. Philosophical Trans- actions of the Royal Zoology London, B 225: 327-379. 1950, The Tunicata. Ray Society Publications 133: 1-354. BREWIN, B.1. 1950a. Ascidians of New Zealand, Part 4 — Ascidians in the vicinity of Christchurch. Transactions of the Royal Society of New Zea- land 78(2—3): 344-353. 1950b, Ascidians of New Zealand, Part 5 — Ascidi- ans from the east coast of Great Barrier Island. Transactions of the Royal Society of New Zea- land 78(2—3); 354-362. 1951. Ascidians of New Zealand, Part 6 — Ascidi- ans of the Hauraki Gulf, Part 2. 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Bulletin of the American Museum of Natural History 84: 1-476. 616 MEMOIRS OF THE QUEENSLAND MUSEUM INDEX TO TAXA (Taxon descriptions and figures in bold) Adagnesia, 385 386 Agnesiidae, 379 386 Amaroucium, aequalisiphonis, 400) altarium, 519 argus, 504 crateriferum, 536 multiplicatum, 567 Anadistoma, 375 387 389 390 422 440 attenuatum, 440-441 Aplidiopsis, 375 380 382 386 387 389 390 399 442 443 445 467-468 472 473 474 507 amoyensis, 445 468 470 atlantica, 468 confluata, 468-469 discoveryi, 468 469 helenae, 468 mammillata, 468 469-470 471 pannasa, 468 470 pyriformis, 468 sabulosa, 468 470-471 stellata, 468 tokaraensis, 468 471 Aplidium, 375 380 381 382 383 384 385 386 387 390 393 399 400 406 424 442 443 445 447 474 483 497 504 507-513 527 537 560 sp. 3: Kott et al, 1984, 569 acroparum, 384 442 512 513 515 518 560 586 604 africanum, 512 altarium, 376 381 384 386 387 388 511 512 514 §19-522 574 577 584 594 604 amorphatum, 380 383 509 512 517 522-523 577 580 arboratum, 442 586 australiense, 383 494 511 514 523-525 550 575 australiensis, 548 bacculum, 497 512 517 526 544 604 brevilarvacium, 511 514 527-528 550 553 577 604 caelestis, 384 509 512513 516 §28-529 530537 544 557 582 586 589 590 605 caeruleum, 504 californicum, 567 570 594 cellis, 486 512 513 557 590 circumvolutum, 523 elivosum, 485 512 516-530 532 537 544 551 557 560 577 586 606 colelloides, 523 congregatum, 511 514 533-534 coniferum, 512 516 535 553 554 564 craleriferum, 381 384 442 484-485 49] 495 497 507 512 516 530 531 533 536-538 544 551 560 607 depressum, 511 515 527 538-539 564 566 digitatum, 527 directum, 312 516 534 539-540 distaplium, 512 517 540-541 elarum, 511 514 541-542 563 Jiliforme,376 497 512 513517 527 540 542-544 584 flavolineatum, 512 522 546 547 565 566 567 570 582 583 594 Jluarescum, 511 515 534 544-546 548 569 Joltorum, 570 372 gastrolineatum, 511 515 $46-547 567 570 583 gelasinum, 511 515 546 547-548 geminatum, 384 494 511 514 524 526 528 534 $48-551 553 577 griseum, 512 516 530 344 550 551 hypurgon, 530 incubatum, 380 511 515 534.540 547 551-553 573 384 indicum, 580 inflorescens, 376 494 511 514 553-554 557 572.577 Jacksoni, 511 515 536 554-555 latusexitus, 592 594 lenticulum, 312 516 530 555-557 590 607 lobatum, 442 491 495 497 507 510512513 528 561 363 581 582 590 lodix, 511 515 553 557-558 longithorax, 579 lunacratumn, 388 390 512 516 519 530 531 533 547 548 557 558-561 580 608 macrolobajum, 380 384 389 512 513 517 561-563 572 582 magnilarvum, 512 516 563-564 maru, 513 mernovensis, 509 513 543 544 mintseulum, 511 515 564-566 multilineatum, 388 390 51] 515 547 565 566-567 570 multipapillarum, 389 513 551 592 multiplicatum, 511 514 521 544 546 547 567-568 572 573 574 583 594 608 nadaense, 513 opacum. 511 514 522 534 542 550 553 554 557 563 567 570-571 583 609 ordinatum, 558 560 561 ornatum, 388 390 511 515 553 567570 572-573 583 584 594 pantherinum, 530 533 538 paralineatum, 494 5\2 517 $74 parastigmaticum, 383 512 517 575 parvum, 511 514 576-577 petrosum, 384 512 517 542 577-578 589 609 phortax, 592 594 pliciferum, 570 572 594 THE AUSTRALIAN ASCIDIACEA 3 protectans, 380 388 390 509 512 517 523 548 560 577 579-580 610 pseudobesum, 522 recumbens, 478 ritteri, 512 513 517 530 55] 579 581 589 robustum, 388 390 511 515 547 557 567 570 582-583 610 rosarium, 511 514 583-584 rubricollum, 512 516 584-585 586 610 solidum, 384 512 517 540 542 557 579 582 584 586-587 tabascum, 384 388 390 486 512 513 517 557 588-589 610 tremulum, 513 563 trigesense, 389 511 515 555 590-591 triggsiensis, 590 uouo, 513 utente, 381 386 388 511 512 515 521 547 548 553 566 567 570 574 584 592-593 Aplousobranchia, 375 376 378 379 380 384 387 390 442 Ascidiacea, 375 378 379 384 386 442 Ascidiidae, 379 386 Atopogaster, 474 504 507 trepicum, 503 Botryllinae, 379 384 442 Batryllus, 385 schlosseri, 385 stewartensis, 534, 540 Ciona, 381 384 intestinalis, 385 386 Cionidae, 379 382 385 386 387 444 Citorclinum, 375 390 391 422 laboutei, 422 Clavelina, 387 421 cylindrica, 387 dagysa, 42) dentatosiphonis, 416 meridianalis, 42) moluccensis, 387 ostrearium, 421 Clavelinidae, 379 380 381 383 387 388 389 390 392 Condominium, 375 381 382 383 390 392 393 396 504 areolatum, 384 397-399 400 403 406 413 595 Corellidae, 379 Corellinae, 379 Cystedytes, 388 Diazonidae, 379 380 382 383 384 385 386 390 391 444 474 505 Didemnidae, 378 379 380 381 383 385 442 475 Diplosoma, 521 listerianum, 385 617 Distaplia, 380 381 382 387 391 505 550 australiensis, 494 528 550 prolifera, 547 Distoma pulchra, 400 416 Dumus, 375 380 382 390 398 399 400 401 412-413 416 areniferus, 384 399 400 413 dumosus, 412 Ecteinascidia turbinata, 385 386 Enterogona, 378 Eudistoma, 382 388 405 422 440 angolanum, 405 442 474 503 carnosum, 405 reginum, 405 Eugyra, 385 386 Euherdmania, 375 380 382 383 386 387 389 390 392 393 398 400 412 413 415-416 417 420 421 505 areolata, 416 australis, 396 416 clavifarmis, 393 399 415 416 417 419 claviformis group, 416 dentatosiphonis, 381 386 415 416-417 419 421 digitata, 376 386 415 416 417-420 digitata group, 416 dumosa, 416 Jusciculata, 390 415 416 gigantea, 415 416 417 gigantea group, 416 morgani, 414 416 rodei, 415 416 solida, 415 416 translucida, 381 386 415 416 417 420-421 597 vitrea, 414 415 416 Euherdmaniidae, 375 379 380 381 387 388 390 391 399 413-415 504 Euherdmaniinae, 375 390 441 Exostoma, 442 Hexacrobylidae, 379 Holozoidae, 379 380 381 382 383 387 388 389 390 391 392 399 442 Holozoinae, 475 Homoeodistoma, 392 longigona, 392 401 michaelseni, 392 omasum, 392 397 399 Hypodistoma, 388 442 Hypsistozoa, 391 505 Lissoclinum patella, 484 537 Macroclinum, arenaceum, 478 hypurgon, 490 499 618 macroglossum, 494 papilliferum, 497 vilreum, 467 Molgula, 386 calvata, 385 ellistoni, 385 manhattensis, 385 sabulosa, 385 Molgulidae, 379 384 Monniotus, 375 380 381 382 383 384 387 390 392 393 398 399 400 403 412 415 416 442 504 527 australis 393 394-396 412 595 radiatus, 384 393 394 396 397 ramosus, 393 394 Morchellinm, 375 380 386 387 390 399 443 504-505 albidum, 474 505 argus, SDS intercedens, 491 pannosum, 474 484 505 506 547 partitionis, 505 Neodistoma, 391 Nephtheis, 381 Perophora, 386 Perophoridae, 379 384 Pharyngodictyon, 375 391 399 400 bisinus, 399 cauliflos, 399 elongatum, 399 magnifili, 399 mirabile, 399 Phlebobranchia, 378 379 381 384 444 Placentela, 375 387 390 391 392 400 areolata, 392 396 397 398 crystallina, 391 392 ellistoni, 397 translucida, 392 Placentelidae, 375 379 380 381 382 391 399 504 Pleurogona, 378 Plurellidae, 379 Polycarpa, 384 385 386 intonata, 385 tinctor, 385 linctorella, 385 Polycitor, 382 384 388 390 415 giganteum, 579 Polycitorella, 388 Polycitoridae, 379 380 381 383 387 388 390 391 392 422 474 504 505 Polyclinidae, 375 378 379 380 381 382 383 384 385 387 388 389 390 391 399 441-443 446 Polyclininae, 375 390 Polyclinum, 375 380 381 382 383 385 386 387 388 389 390 393 399 400 413 441 442 443 445-447 448 MEMOIRS OF THE QUEENSLAND MUSEUM 455 460 467 468 471 472 473 474 495 497 504 507 509 521 550572 aurantium, 459 circulatum, 445 452 complanatum, 445 constellatum, 443 445 449 459 467 crater. 446 depressum, 446 Jfestum, 441 445 464 467 fungosum, 376 380 445 446 447 452 gelidus, 445 glabrum, 444 445 446 448-450 hospitale, 445 incrustatum, 445 446 448 450-452 454 458 461 464 600 laxum, 463 464 macrophyllum, 452 454 464 467 macrophyllum phortax, 464 macrophyllum typicum, 467 marsupiale, 388 445 446 447 448 452-454 455 458 461 600 neptunium, 450 452 nigrum, 447 nudum, 445 446 452 454-455 600 orbitum, 383 445 446 454 455 456 467 473 474 pute, 463 sabulosum, 458 459 saturnium, 445 446 448 452 454 455-458 459 463 464 solum, 444 445 446 447 455 458-459 460 464 sundaicum, 447 fenuatum, 445 446 454 459-461 497 600 terranum, 384 388 445 446 460 461-462 601 tralatica, 447 tsutsuti, 388 445 446 448 455 457 458 463-464 vasculosum, 381 389 443 445 446 447 450 455 458 463 464-467 472 499 Polyzoinae, 379 384 386 442 Prochleran, 538 56) 567 569 608 Protoholozoa, 399 Protopolyclinidae, 375 379 380 381 382 386 387 388 389 390 391 392 393 399 401 442 504 Protopolyclinum, 375 382 390 391 392 393 Psammaplidium, Jragile, 586 incrustans, 586 lobatum, 586 ordinatum, 558 560 pedunculatum, 410 prunum, 498 solidum, 586 Pseudodistoma, 375 381 383 387 389 390 392 422-425 428 440 441 444 446 524 acuatum, 422 423 424 425-426 431 439 597 africanum, 425 430 431 438 THE AUSTRALIAN ASCIDIACEA 3 619 antinbaja, 425 arborescens, 422 425 428 436 aurea, 426 aureum, 387 422 423 424 425 426 436 australe, 376 423 424 425 428-430 438 597 australis, 428 brieni, 422 424 425 candens, 386 389 423 424 430-433 438 439 480 597 cereum, 390 422 424 425 426 428 430 431 433 4346 439 440 eyrnusense, 424 425 430 433 fragile, 384 424 425 433 434 gracilum, 384 386 422 423 424 425 433-435 440 598 inflatum, 423 424 428 435-436 598 kanako, 422 424 425 novaezelandiae, 422 425 428 436 opacum, 422 425 428 436 439 ariens, 389 390 422 423 424 425 439 431 436-439 599 pilatum, 423 424 426 431 436 439 599 pulvinum, 422 423 424 425 439-440 599 Pseudodistomidae, 375 379 380 381 382 386 387 388 389 390 391 399 421-422 442 504 Pycenoclavella, 380 384 389 393 420 421 422 Pycnoclavellidae, 379 380 383 386 387 388 390 391 415 422 Pyura, 386 littoralis, 385 Pyuridac, 379 385 Rhodosomatinae, 379 Rhopalaea, 384 Ritterella, 375 381 384 386 387 388 389 390 392 393 398 399 400-402 405 407 409 412 422 442 474 504 527 aequalisiphonis, 399 400 402 408 aequalisiphonis group, 400 arenosa, 392 400 401 413 asymmetrica, 381 382 383 384 398 399 400 401 402-403 406 409 410 412 413 595 cireularis, 400 401 compacta, 381 382 383 400 401 403-405 410 442 596 cornuta, 381 382 383 393 398 399 400 401 403 405-406 409 410 412 442 596 dispar, 389 399 400 401 406-408 421 glareosa, 400 401 herdmania, 410 413 multistigmata, 382 383 389 390 399 400 401 408-409 442 596 papillata, 381 382 383 384 399 400 401 406 409-410 412 442 pedunculata, 380 381 383 384 399 400 401 402 403 406 409 410-412 413 596 pedunculata group, 400 prolifera, 400 401 408 proliferus, 406 pulchra, 389 rubra, 400 sigillinoides, 400 401 402 tokioka, 400 413 vestita, 400 402 yamazii, 400 402 413 Ritterellidae, 375 379 380 381 382 383 387 388 389 390) 391 393 399 400 442 505 Sidneinides, 375 381 386 387 389 442 443 467 471-472 474 japanense, 472 snamoti, 472 ftamaramae, 471 472-473 601 Sidnyum, 390 443 509-510 appendiculatum, 309 elegans, 509 indicum, 509 pentatrema, 509 turbinatum, 509 Sigillina, 381 382 388 389 422 424 440 447 504 grandissima, 428 myjébergi, 388 nigra, 447 Sigillinaria, 392 400 arenosa, 392 aurea, 426 clavata, 391 392 novaezelandiae, 392 pulchra, 400 416 Stolidobranchia, 375 379 384 442 Stomozoa, 415 421 Stomozoidae, 379 381 Styelidae, 379 Styelinae, 379 Synoicidae, 442 Synoicum, 375 381 382 385 386 387 388 390 399 400 402 442 443 448 460 467 472 473-477 505 507 510 541 550 angustum, 475 476 477 apectetum, 475 483 appendicularum, 509 arenaceum, 475 476 477 478 483 atopogaster, 386 474 475 476 478-479 504 506 507 560 bowerbanki, 474 476 480-481 541 602 buccinum, 475 476 478 481-482 602 castellatum, 382 389 442 475 476 483-484 537 560 602 chrysanthemum, 475 476 486 495 citrum, 475 476 486 495 602 clavatum, 501 503 620 MEMOIRS OF THE QUEENSLAND MUSEUM concavitum, 474 475 476 480 487 506 602 durum, 475 476 488 497 erectum, 475 476 478 489 501 galei, 475 476 490 haurakiensis, 475 herdmani, 475 478 hypurgon, 447 475 490 491 528 intercedens, 474 476 491-492 497 503 investum, 586 kuranui, 475 478 489 501 503 longistriatum, 475 476 493 macroglossum, 474 475 476 493 494 497 603 obscurum, 475 476 496 540 occidentalis, 477 otagoensis, 477 papilliferum, 459 460 474 475 476 491 495 497 499 partitionis, 490 pererratum, 477 prunum, 475 476 487 490 498 499 603 rubrum, 474 sacculum, 474 475 476 490 491 493 497 499-500 603 stewartense, 455 477 527 suarenum, 442 475 476 489 493 497 501-502 603 suesanum, 491 493 tropicum, 475 476 488 493 503-504 507 turgens, 473 Tunicata, 375 Tylobranchia, 393 THE AUSTRALIAN ASCIDIACEA, SUPPLEMENT 2 PATRICIA KOTT Kott, P. 1992 10 01: The Australian Ascidiacea, Supplement 2. Memoirs of the Queensland Museum 32(2): 621-655, Brisbane. ISSN 0079-8835. Information from newly examined material and recently published reports supplements that in the continuing review of the Australian Ascidiacea (Kott 1985, 1990a, 1990b, 1992). Recorded ranges of Sigillina fantasiana, Hypsistozoa distomoides, Polycitor obeliscus, Polycitorella coranaria, Exostoma ianthinum, Botryllus stewartensis, and Pyara rapafar- mis are extended; the known depth range of Pseudodiazona claviformis increased, Pycno- clavella aurantia, P. elongata, Distaplia prolifera and Eudisioma aureum are now recorded from outside their type Jocalities; new colour patterns and morphological variations are recorded for Clavelina psendobavdinensis, Pyenoclavella diminuta and P. elongata: and larvae of Pyenoglavella auraniia and Palyciter calamus, described for the first time, are characteristic of theirrespective genera, A new species of the deep water genus Protoholozou is from 7 to 15m off the southern Australian coast. A new species of Cystadytes has unusual spicules in a layer beneath the surface test, and one in Polyandrocarpa, indigenous to the central eastern Australian coast, forms colonies to a metre in maximum extent. A new Cnemidocarpa species has a large number of gonads, and another has only 3 branchial folds on each side. Western Pacific species of Ascidia (1), Ecteineascidia (2), and Pyure (1), in the Australian fauna are found to have representatives in the component of the New Caledonian fauna previously supposed to be indigenous. The relationships and geographic range are discussed of a further 13 stolidobranch and phlebobranch species, including 4 believed to be pantropi- cal, and abyssal Asajirus indicus and Oligotrema psammitey of the Hexacrobylidac. 0 Ascidiacea, Aplousobranchia, Phlebobranchia, Stolidebranchia, Pratehotozoa, Hexacrebylidae, Western Pacific. Patricia Kott, Queensland Museum, PO Box 3300, Seuth Brisbane, Queensland, 4101, Australia; 13 May, 1992. Australian ascidians in the suborders Phlebo- branchia, Stolidobranchia and Aplousobranchia (excepting the family Didemnidac) have been reviewed by Kott (1985, 1990a and 1992), Mor- phological, taxonomic, phylogenetic and geo- graphic information additional to that in the first part (Kott 1985) is presented in an initial supple- ment (Kott 1990b). This, the second supplement, complements parts 1 and 2 (1985 and 1990a). This information is presented only when it ex- pands known geographical range, the known morphological parameters of the (axon, or where new phylogenetic insights are available from newly examined material or recently published accounts of the western Pacific fauna, as in Nishikawa (1984, 1986), Monniot and Monniot £1987, 1990, 1991), and Menniot, C. (1987,b, 1988, 1989, 199] a,b). A number of Australia’s tropical species are discussed here, specifically in relation to their affinities. with New Caledonian populations. The ascidian fauna of the westem Pacific has been relatively well studied as evidenced by the works ~ vited below and in Kott (1985, 1990a, 1990b) — of Sluiter, Van Name, Tokioka, Nishikawa, Kott, Vasseur, Millar, and C. and F, Monniot. From these works it 1s clear that Fiji, the Tokhara Is in southern Japan, Indonesia, other western Pacific Jocations and tropical Australia share a common fauna which sometimes extends into the Indian Ocean (see Kott 1985, 1990a, 1992). In particu- Jar, the north eastern Australian coast and the Great Barrier Reef can be said to be part of the western Pacific region. ‘The reports of C. and F. Monniot (1987-91) on the ascidians taken in the French ‘vaste pro- gramme d’exploration’ (Monniot, C. 1987a, p. 3) around New Caledonia propose new species sug- gesting a larger indigenous fauna than has been found previously in this or any other western Pacific location. Comparison of a number of these species, however, with related ones in tropical Australia and other western Pacific locations has shown them not to be distinct. Differences re- ported between New Caledoman and other popu- lations lie within the range of intraspecific variability that is a property of all biological material, This variubility 1s associated with age differences, intraspecific population differences, differences between genotypes, ecological differ- ences in growth form or pigmentation, and differ- ences due to artefacts associated with the collection and preservation of the material. The status of morphological differences has some- times been difficult to determine where neither intraspecific variation nor the number of speci- mens examined is indicated. Thus, in some, but not all, cases intraspecific differences can be con- fused with the genetic differences indicating iso- lation and spectation. A further problem in resolving some of these problems is the relationship between apparently conspecific populations in the tropical eastem Atlantic and western Pacific — where distance and other geographic barriers would appear to preclude gene flow (see Ascidia archaia, Pero- phora multiclathrata, Pelycarpa aurita, and Cnemidocarpa areolata, below). Museum registration numbers are given for all examined material, Abbreviations used are AM (Australian Museum), SAM (South Australian Museum), OM (Queensland Museum). MV (Mu- seum of Victoria). For detailed data on distribu- tion of each taxon the registers of relevant Australian Museums must be referred to. The following taxa are discussed below: DIAZONIDAE Rhopalaea crassa (Herdman, 1880) Pseudodiazona claviformis (Kott, 1963) CLAVELINIDAE Clavelina pseudobaudinensis (Kott, 1976) PYCNOCLAVELLIDAE Pycnoclavella aurantia Kott, 1990a Pyenaclavella diminuta (Kott, 1957) Pycneclavella elongata Kott. 1990a HOLOZOIDAE Sigillina fantasiana (Kott, 1957) Sigillina grandissima Kott, 1990a Distaplia prolifera Kott, 1990a Hypsistozoa distameides (Herdman, 1899) Protoholozoa Katt, 1969 Protoholozea australiensis tsp, POLYCITORIDAE Cystodytes ramosus 0.sp. Polycitorella coronaria F. Monniot, 1988 Polycitor calamus Kott, 1990a Polycitor obeliscus Kou, 1957 Eudistoma aureum Kott, 1990a MEMOIRS OF THE QUEENSLAND MUSEUM Exoastoma ianthinum (Sluiter, 1909) ASCIDITIDAE Ascidia archaia Sluiter, 1890 Ascidia liberata Sluiter, 1887 PEROPHORIDAE Perophora multiclathrata (S\uiter, 1904) Ecteinascidia diaphanis Sluiter, 1885 Ecteinscidia nexa Sluiter, 1904 STYELIDAE Cnemidocarpa amphora n.sp. Cnemidocarpa areolata (Heller, 1878) Cnemidocarpa intestinata Kott, 1990 Cnemidocarpa tribranchiata, n.sp. Asterocarpa humilis (Heller, 1878) Polycarpa contecta (Sluiter, 1904) Polycarpa aurita (Sluiter, 1890) Palyandrocarpa colemani 0.sp. Botryllus stewartensis Brewin, 1958 PYURIDAE Pyura albanyensis Michaelsen, 1927 Pyura curvigona Tokioka, 1950 Pyura rapaformis Kott, 1990b Herdmania moms (Savigny, 1816) Microcosinus tuberculatus Kott, 1985 HEXACROBYLIDAE Asajirus Kott, 1989a Asajirius indicus (Oka, 1913) Oligotrema Bourne, 1903 Oliogtrema psammites Bourne, 1903 Suborder APLOUSOBRANCHIA Lahille, 1887 Family DIAZONIDAE Seeliger, 1906 The family is distinguished from Cionidae by its relatively smal! thorax, Jong, straight, vertical gut loop, and epicardial sacs isolated from the pharynx. Further, although only 2 species of Rho- palaea are known to form colonies of not more than 2 zooids, other genera of the Diazonidae are colonial, replicating as do some other aplouso- branchs by horizontal strobilation of the abdomen across the epicardium. Lahille (1887) proposed acommon ancestor for taxa with complete internal longitudinal bran- chial vessels, and accordingly included Diazoni- dae and Cionidae in the suborder Phlebobranchia. Most authors accepted this phylogeny until Kott (1969, 1985, 1990a) drew attention to the rela- THE AUSTRALIAN ASCIDIACEA 2. SUPPLEMENT tionships between Cionidae, Diazonidae and other aplousobranch families, based on the pres- ence of entire epicardial sacs and their regenera- live role in the process of replication. In Phlebobranchia (but not in Diazonidae or Cioni- dae) the epicardial sacs are divided into minute vesicles that serve an excretory rather than regen- crative function. Further, in phlebobranch ascidi- ans (including Diazonidae and Cionidae) the gut loop is folded up alongside the pharynx while the aplousobranch gut loop, although sometimes horizontal and twisted, is always behind the phar- ynx, As pointed out by Kott (1985), the loss of internal Jongitudinal vessels occurs in parallel in all sub-orders of the Ascidiacea as zoojds reduce in size with the evolution of replication and a colonial habit (see Perophoridae and Polyzoinac), The morphological relationships between Dia- zonidac, Cionidac and other aplousobranch (rather than phlebobranch) families is confirmed by their chemistry (see Hawkins #7 al, 1983). Genus Rhopalaea Philippi, 1843 C. Monniot (1991a) believes that Diezana and Rhopalaea are indistinguishable, However, these genera ate readily separated by the relanvely primitive characters of Rhopalaea including its particularly large branchial sac, and relatively short oesophagus. Although the epicardium is used to regenerate parts of zooids, apparently the process is not one of spontaneous replication, for only 2 species of Rhopalaea are known to form colonies (see Kott 1990a). On the other hand, Diazona species have smaller thoraces, a gener- ally longer oesophagus, and a prolific replicative capacity resulting in large colonies of numerous zooids. Rhopalaea crassa (Herdman, 1880) Ecteinascidia crassa Herdman, 1880, p,723, Rhopalaea crassa Kott, 1990a, p.26 and synonymy. Nishikawa, 1991, p.25, Rhopalaea respictens Monniot, C. 199Ta, p.494. REMARKS Monniot (1991a) has suggested that species of Rhopalaea can be separated by the colour of the living test, the musculature and relative size of the thorax and abdomen. The significance of the col- our differences that occur in living specimens of this species is not resolved, However Kott and Goodbedy (1982), Kott (1990a) and Nishikawa 623 (1991) have reported on a wide range of speci- mens of R. crassa and its synonyms from a wide range of locations, including type locations and have demonstrated similar variations in the mus- culature associated with contraction, and vati- alions in the relative size of thorax and abdomen, apparently the result of growth, and regeneration of the thorax which appears to occur from time to time. Colour differences appear partly regional, bluc being the usual colour recorded for living speci- mens in the Philippines, while specimens wath colourless. transparent thoraces with yellow, white and black markings, becoming yellow and opaque when the test is thicker, are recorded fron eastern Australia (Kott 1990), Hong Kong (Kott and Goodbody 1982), New Caledonia (Monniat 199 1a) and Japan (Nishikawa (1991), The colour of all specimens is lost in preservative. Rhopalaea respiciens Monniot, 19914 falls well within the range of variation recorded for even single populations of K. crassa, e.g. from Heron I, (see Kott 1990s), Genus Pseudodiazona Millar, 1963 Pseudodiazona claviformis (Kott, 1963) (Fig. 1) Proropalyclinum claviforme Kott, 1963, p.72. Pseudodiazona claviformis: Kot, 1990a, p31 and synonymy. DISTRIBUTION New ReEcoRDS! Victoria (off Cape Everard, SAM £2460). South Australia (Spencer Gulf, SAM E2547), RECORDED RANGE! The specics is known from New South Wales and the eastern end of Bass Strait to the northern Great Australian Bight. The newly recorded specimen from Cupe Everard increases the known depth range from 100 to 446m. DESCRIPTION The newly recorded specimens confirm the morphology of this seldom recorded species. The specimen from Spencer Gulf retains a pat- tern of pale grey pigment in vertical petal shaped patches forming a ring around the base of each of the siphons, In one specimen (SAM E2460) collected in October, the numerous male follicles are bunched in the posterior end of the zooid and eggs are lined up in a single series i the oviduct, In the other specimen gonads are not mature. MEMOIRS OF THE QUEENSLAND MUSEUM 624 THE AUSTRALIAN ASCIDIACEA 2. SUPPLEMENT Family CLAVELINIDAE Forbes and Hanley, 1848 Genus Clavelina Savigny, 1816 Clavelina pseudobaudinensis (Kott, 1976) Oxycorynia pseudobaudinensis Kot, 1976, p.54. Clavelina pserdobaudinensis: Kott, 1990a, p.58, and synonymy. DISTRIBUTION New ReEcoRDS; South Australia (Nuyts Archipelago, SAM E2434 E2471). RECORDED RANGE: The known range is around the southern half of the Australian continent, from Hout- man’s Abroihos to Jervis Bay (NSW). DESCRIPTION One of the newly recorded specimens from South Australia (SAM E2434) has a Jong (3cm) narrow (1.3cm diameter) stalk similar to that of C. baudinensis (see Kott 1990a), Otherwise the zooids are characteristic of C_ pseudobaudinen- sis, with median pigment patches — one between the siphons and one at the antenor end of the endostyle, Larvae are large, with the trunk 0.9mm long (0.9cm sic, Kott 1990a). Family PYCNOCLAVELLIDAE Kott. 1990 Genus Pyenoclavella Garstang, 1891 Pycnoclavella aurantia Kott, 1990 (Figs. 2,3) Pycnoclayella aurantia Kott. 1990a, p.7 1, DISTRIBUTION New Recorb: South Australia (Thorny Passage, SW Hopkins §., SAM E2422). RECORDED RANGE: The holotype from Franklin [. Nuyts Archipelago was the only previously recarded specimen (Kott 1990a). DESCRIPTION Unlike the type specimen, the vertical stalks of the newly recorded one form a sandy aggregated 625 basal mass similar to that observed previously in Pycnoclavella tabella Kott, 1990a. Zooids are a gold colour in preservative. The thoraces are turned at right angles to the long axis of the abdomen, and have a terminal atrial aperture and the branchial aperture on the side of the transparent head. Stigmata are in 6 instead of the 8 rows in the type specimens. The newly recorded specimens, collected in October, have up to 12 embryos in the atrial cavity at the terminal end of the zooid. The larvae are large, with an almost spherical trunk about 0.9mm long with the tail wound almost once around it. There is an ocellus but no otolith, 4 rows of stigmata, and 3 characteristic deeply in- vaginated adhesive organs. REMARKS Pycnoclavella arenasa (Kott, 1972b) also has 6 rows of stigmata, bul its thoraces are not turned at right angles to the long axis of the zooid, the abdomina are not free of the central common test, and the thorax is narrow (with only 20 rather than 50 stigmata per row). The tropical Pyenoclavella detorta (see Kott 1990a) resembles the present species in the orientation of the thorax, and in having 6 rows of stigmata. There is a difference in the number of stigmata, however, P. detoria having only about 24 per row. The larva is also similar to that of P. dererta, having the characteristic 3 inverted tubular adhe- sive organs and Jacking an otolith. No more than 4 embryos at a time have been found in the atrial cavity of P. detorra, while the newly recorded zooids have up to 12, Also the larvae of P. detorta have 1.7mm Jong trunks, longer than those of the present species (1.1mm). The newly recorded specimens confirm the separate status of the related species P. aurantia and P. detorta. Pycooclavella diminuta (Kott, 1957) (Figs. 4-7) Clavelina diminute. Kott, 1957, p.89, Pycnoclavella diminuia; Kot, 1990a, p.73 and synonymy. FIGS. 1-11: Pseudodiazena claviformis (SAM E2547) — |, contracted zonid. Pycnoclavella aurantia (SAM E2422) — 2, zooid in test showing embryos in atrial cavity (Jem excised from oesophageal neck); 3, larva. Pyenoclavella diminuta — 4, section of colony stalk showing vesicles and hair-like test processes (SAM E2473); 5, part of colony (SAM E2473): 6, thorax showing pigment patches around apertures (SAM E2492): 7, detail of pigment patches around atrial (left) and branchial apertures, Proroholozoa australlensis n.sp. (holotype SAM E2423) — 8, head of colony showing zooid arrangement; 9, part of colony; 10, zooid from ventral surface, L1, zooid, dissected down endostyle showing transverse vessels in thorax. (Scules: 1,4,10,11 — 0.5mm; 2,5,8 — 2mm; 3 — 0.25mm; 6 — Imm; 7 — 0.2mm; 9 — 5mm). 626 DISTRIBUTION NEw RECcorDs: South Australia (Nuyts Archipelago, SAM E2473; Lincoln National Park, SAM E2492). RECORDED RANGE: The species is often encountered in benthic collections from all around the Australian coast and in the Western Pacific. DESCRIPTION The newly recorded colonies (SAM E2473), have their stalks aggregated together into a loose sandy mass that readily distintegrates. This mass is formed by hair-like test processes from along the stalks of adjacent zooids adhering to sand particles. The usual oily-looking spheres charac- teristic of this species are in the test of the stalks. It is probable that the basal sandy mass was buried in the substrate, and that only the thoraces pro- jected above the surface. The specimen lot SAM E2492 has unusual comma-shaped pigment patches around each smooth-rimmed aperture. In preservative these patches are a brown-orange colour. Around the atrial aperture 3 of these patches are arranged symmetrically in front of the aperture (between the siphons) and one is behind the aperture. Five patches, each with its point directed posteriorly, are in a row around the ventral border of the branchial aperture. These 5 separate patches sometimes are confluent. Despite this unique pigment pattern, the zooids are characteristic of P. diminuta. Similar (but not identical) pigment patches are around the bran- chial apertures of specimens from Exmouth Gulf (QM GH4083: Kott 1990a, Plate 6d). Pycnoclavella elongata Kott, 1990 Pycnoclavella elongata Kott, 1990a, p.76. DISTRIBUTION New REcorD: South Australia (St. Vincent Gulf; SAM E2418). RECORDED RANGE: The species was previously known only from type localities amongst rocky outcrops around the Nuyts Archipelago. The newly recorded specimen is from a boat ramp at 3 to Sm. DESCRIPTION Thoraces are dark coloured in preservative, and naked. The stalks are aggregated together to form a basal mass of parallel, vertical sandy stalks. Zooids have 11 rows of stigmata, while the type material has 14 rows. Specimens otherwise con- form with the previous descriptions of this spe- cies (Kott 1990a). MEMOIRS OF THE QUEENSLAND MUSEUM Family HOLOZOIDAE Berrill, 1950 Genus Sigillina Savigny, 1816 Sigillina fantasiana (Kott, 1957) Eudistoma fantasiana Kott, 1957, p.76. Sigillina fantasiana: Kott, 1990a, p.92. DISTRIBUTION New RECcoRDs: Tasmania (Break Sea I. Port Davey, QM GH5504). South Australia (Spencer Gulf, SAM E2505 E2518-9; Yorke Peninsular, SAM E2485 E2488; St. Vincent Gulf, SAM E2483 E2494; Pearson I., SAM E2537). RECORDED RANGE: The species is known from Cock- burn Sound (WA) and across the southern coast of the continent to Gabo I. off the south eastern coast. The newly recorded specimen from Port Davey, southwest- ern Tasmania, suggests a range around that island. DESCRIPTION Colonies from Tasmania are small cushions with evenly distributed blue zooids and a cloudy test. Larger sheet-like colonies from South Aus- tralia, growing along Zostera stalks, have almost glassy test and blue pigment around the apertures. Zooids have a characteristically short abdomen up to 3 times the length of the thorax. A relatively long vascular stolon projects from the posterior end of the abdomen. Three embryos are lined up from the top of the abdomen to the postero-dorsal part of the thorax, where the most advanced pro- jects from the body wall. Tailed larvae are present in February (SAM E2519), July (SAM E2494), September (SAM E2483), and November (SAM E2488 E2518) Sigillina grandissima Kott, 1990 Sigillina grandissima Kott, 1990a, p.93. DISTRIBUTION New REcorDs: South Australia (W. Great Australian Bight, SAM E2452 E2457 E2479-81 E2545). RECORDED RANGE: The species is known from Dam- pier Archipelago to the Great Australian Bight, al- though, so far there is a gap in the records between Cockburn Sound and Eucla (SA). DESCRIPTION The specimens are fleshy lobes with thick fleshy stalks about the same length as the oval head. They are a very soft, pale lemon in preserv- ative with patches of dark pigment over the neural complex and at the anterior end of the endostyle. 628 be incorrect, although the type location of the Australian species is off the most southerly part of the Australian continent. The family Holozoidae is represented in Aus- tralian waters by all its known genera, including 2 indigenous ones (Polydistoma Koit, 1990a and Neadistoma Kott. 1990a). Sigillina Savigny, 1816, Hypadistoma Tokioka, 1967, Hypsistezea Brewin, 1956, Sycozoa Lesson, 1830 and the present genus (with the exception of P_pigra) are known mainly from the southern hemisphere (see Kott 1990a), Only Distaplia is a cosmopolitan genus. Protoholozoa australiensis n,sp. (Figs. 8-11) DISTRIBUTION Tyre LocaLiry: South Australia (approximately Iki NE Margaret Brock Reef off Cape Jaffa, 7 to 15m dropoff, overhangs, caverns, coll. W. Zeidler, K. Gowlett Holmes 17,2,89, hololype SAM E2423, pa- ratype SAM E2422). FURTHER RECORDS: None. DESCRIPTION EXTERNAL APPEARANCE: The colony consists of almost spherical heads up to 5mm diameter, each at the end of the terminal branch of a branch- ing stalk upto lcm Jong. The combined length of primary, secondary and terminal branches of the stalk are about 2cm, However the available speci- mens are only part of a colony and the total length of the colonies could be greater. Stalks are nar- row, cylindrical, about Imm diameter, firm and translucent, with a stiff outer cuticle on the lower part. Zooids are in a single circle, parallel with one another, around the perimeter of the head of the colony. The abdomina of the zooids are upper- most around the top of the head. The atrial aper- iures are at the posterior end of the thoraces opening to the exterior in a circle around the outer margin of the head just behind its terminal free end. The antero-dorsal branchial apertures are in a circle at the other end of the head, around the top of the stalk, Posterior abdominal vascular stolons extend parallel to one another along the length of the stalk, running back past the en- dostyle from the centre of the gut loop to enter the top of the stalk, Vegetative zooids are in the stalk and some- times in the head in a circle around the top of the stalk near the antenor ends of the adult zooids. However, there is no sign of more than a single MEMOIRS OF THE QUEENSLAND MUSEUM circle of zooids in each head, and the vegetative zooids formed in the stalk may move up to in- crease the diameter of a head, and/or to form systems in developing terminal branches of the stalk, INTERNAL STRUCTURE: Zooids are delicate and transparent, 2 to 3mm long with the gut bend slightly to the left agaist the posterior end of the thorax. The ventral border of the thorax is longer than the dorsum so that the short branchial siphon turns dorsally. It has fine crenellations around the nm but no lobes. Circular muscles are around the branchial and atrial apertures and short bands extend laterally from each end of the atrial aper- ture which 1s a transverse slit at the posterior end of the thorax, Twelve, short, stumpy branchial tentacles are ina circle at the base of the branchial siphen. The branchial wall consists of 4 trans- verse vessels attached to about the ventral one third of the parietal body wall, and free for the dorsal two-thirds of their length. They are at- tached to the dorsum where they cross the dorsal sinus as wide transverse membranes, but there are no dersal Janguets. A few longitudinal connee- tives run between the transverse vessels each side of the mid-dorsal line. The oesophagus is short, and turns to the right to open into an almost horizontal smooth walled, oval stomach. The post-pyloric part of the gut forms a smooth curve from the distal part of the stomach to the anus near the atrial aperture. As in other genera of the Holozoidae (see Sycozoa: Kou 1990a) a spherical gastric reservoir is in the gut loop, together with 4 or 5 immature ova. How- ae the gonads are nol mature in the type mate- rial. REMARKS The type species of this genus, P. pedunculara Kott, 1969 (from the Scotia Sea, the South Pacific Basin and the Antarctic Peninsula at from 3000 to nearly S000m) differs from the present species in having 2 rather than 4 transverse vessels, an internally ridged stomach, a larger head! with a thick fleshy stalk and zooids in rows of about 3 down the length of the head. Protoholozea anthos Monniot snd Monniot, 1991, and P. Jilivm: Monniot and Monniot, 1991 also have a smooth stomach, but 3 rather than 4 transverse vessels, and, in the former species, a long anterior atrial lip (an unusual feature in this genus), Protoholozea cantrella Monniot and Monniot, 1985 differs in its internally ndged stomach and 3 transverse vessels. THE AUSTRALIAN ASCIDIACEA 2. SUPPLEMENT 29 Yamily POLYCITORIDAE Michaelsen, 1904 Genus Cystodytes Drasche, 1884 Cystodytes ramosus n.sp. (Figs. 12-16) DISTRIBUTION Tyre Loca.ity; Queensland (Central section Great Barrier Reef, Haslewood 1., in bay, 20m. coll, AIMS Bioactivity Group 17.10.87, holotype QM GH5374). DESCRIPTION EXTERNAL. APPEARANCE: The colony is a knobbly mass, basically cylindrical with side branches. Each cylindrical branch is about one to 2em diameter. to 2cm long. The colony is about Sem in maximum extent overall. Zooid systems open into furrows and depres- sions on the outer surface and circular to oval or imegular taised zooid-free elevations are scat- tered between the surface openings of the sys- tems, Each system consists of a circle of up to 20 zooids, surrounding a depression (rudimentary common cloacal cavity) in the surface. This de- pression contains the circle of atrial apertures. The circle of branchial apertures is slightly Iess than lcm in diameter. In life, photographs show the specimens to have a smooth, translucent colourless surface layer covering brownish-pink test, The surface 2mm of lest is a translucent, firm and gelatinous layer of bladder cells. It covers a layer of calcareous plate- like spicules, lying parallel to the surface of the colony. The layer is only 1 spicule thick, but the spicules overlap one another slightly around their thickened, scalloped edges. These spicules are relatively pliable, consisting of deposits of granu- lar calcareous material, rather than the hard rigid spicules of Cystedytes dellachiajei. Minute brown pigment cells are scatlered through the test, especially in the surface layer. In the holo- type the zooids are withdrawn from the surface into the central layer of test through the layer of spicules. INTERNAL STRUCTURE:Contracted zooids’ are one to 2mm long. Both the branchial and atrial apertures are 6-lobed, on cylindrical siphons, the atrial siphon larger than the branchial siphon. Strong longitudinal muscles are in the body wall. Stigmata are Jong, arranged in 4 rows of about 12. The gut loop is vertical, with a large, smooth- walled stomach. About 5 pear-shaped male folh- cles are in the gut loop REMARKS The species is distinguished from Cystodytes dellachiajei by the layer of spicules beneath the surface bladder cells, and the absence of a capsule of spicules enclosing the abdomen of each zooid. Genus Polycitorella Michaelsen, 1924 Polycitorella coronaria F. Monniot, 1988 (Figs. 17-22, 43) Polvejtorella caronaria F. Monniot, 198%, p.228. Katt, 1990a, p. 184 and synonymy, DISTRIBUTION New R&coRD: Queensland (Heron I. QM GH5616). RECORDED RANGE: The species is recorded froni 3 to 190m from Northwest Cape to Port Phillip Bay. The new record extends this range to the southern up of the Great Barrier Reef. DESCRIPTION EXTERNAL APPEARANCE: The colony is 4 small thick cushion, about |.Scm diameter and lem deep. Zooids are arranged in 7 circular systems of about 6 per system. Spicules are crowded in the lower half of the colony, at abdominal level. In the upper half, at thoracic level, they are present only inthe centre of each circle of zovids, forming a pillar of spicules that reaches to the upper sus, face, The branchial apertures open around the outside of the upper margin of this white pillar, and the atrial apertures project through the upper edge and open just inside 11s upper margin. The external surface of the colony, both around the sides and on the flat upper surface is covered with a thick layer of bladder cells, free of spicules and pigment cells. Pigment cells are black 1m the living specimen, but brown in preservative. They are crowded in the test lining the zooid compartments, and are evenly distributed elsewhere except where they are excluded by either colourless bladder cells or white spicules. Spicules are 0.025 to 0.07mm. The smaller ones are most plentiful at abdominal level around the zooids. Moderately sized ones are most frequently encountered, and the largest ones are rare. They have about 30 short pointed rays in optical section, although the smaller ones often have rounded rays. The living colony was black and white, the white circular upper surface of the pillars of spicules in the centre of each system contrasting with the rest of the upper surface where spicules are not present to diluic the black pigment cells. 630 MEMOIRS OF THE QUEENSLAND MUSEUM (TV ern, THE AUSTRALIAN ASCIDIACEA 2. SUPPLEMENT INTERNAL STRUCTURE: Zooids are contained narrow vertical canals in the test, and in the pre- sent colony seem always severed somewhere along the Jong, narrow oesophageal neck. The rounded, slightly expanded posterior ends of the abdomina appear to have remained in place in their test compartment in the base of the colony and are not very much contracted. The anterior part of the zooids also has remained in place at the surface. However, both the thorax and the most of the oesophageal neck is very much contracted, and it is this that appears to have caused the zooids to break apart, the abdamina remaining in the base of the colony and the apertures at the surface, The rim of the branchial aperture is divided into 6 smull lobes. The atrial aperture is on a moderately long siphon. The upper rim of the opening is produced into a tongue-like lip. its lip divided into 3 round lobes, The posterior rim of the aperture also is divided into 3 lobes. The thoracic musculature consists of about 16 narrow longitudinal bands, with very fine trans- verse muscles overlying them to form square meshes. Around the anterior part of the abdomen the longitudinal muscles reorganise themselves into about 8 wider bands on each side, Further posteriorly 4 wide longitudinal muscle bands en- case the abdomen, but break up into separate fibres anterior to the gonads and stomach. Fine circular muscles are in one or 2 bands around the anterior part of the abdomen. These appear to be a continuation of the layer of transverse thoracic muscles onto the abdomen. Their arrangement in several bands may be due to the contraction of the longitudinal muscles, These transverse bands do not appear to have become embedded in the test, and there is not 4 constriction of the test between the thoracic and abdominal compartments, as there ts in the large colonies of this species from southem and western Australia (see Kott 1990a). The branchial sac is long with 13 rows of about {6 stigmata. The stomach is amooth, and located in the posterior end of the abdomen. Stig obscured hy the large testis consisting of numerous crowded pyriform follicles covering the dorsal side of the pole of the gut loop, and partly covered by a single large ovum. 631 REMARKS Although it has been taken from Northwest Cape (Western Australia) this species has not previously been recorded from the eastern tropi- cal coast of Australia or from any other location in the Western Pacific, The species appears to flourish in temperate Australian waters, where large colonies often are taken. This small speci- men from the southem tp of the Great Barrier Reef may represent a population at the northem limits of its range, The single specimen does not appear juvenile. despite its small size, having large, mature male follicles and a large ovum. The gonads, branchial sac, circular systems. spicules, pigment cells and other features of the zooid and colony are characteristic of the species, The only significant difference appears to be the absence of a test constriction with a bundle of transverse muscle bands embedded in i between the Tho: racic and abdominal zooid compartments. Haw- ever the transverse muscles are present outside the longitudinal ones, behind the thorax, and the fact that they have not become embedded in the fest may be assowiated with the small size of the colony. Genus Polycitor Renier, 1804 Polycitor calamus Kott, 1990 (Figs, 23-25) Polycttor calamus Kou, 1990a, p.167- DISTRIBUTION New Record: South Australia (Spencer Gulf, SAM E2530). RECORDED RANGE: The Species is recorded from the Great Australian Bight and Spencer Gulf, South Aus- tralia, and from the Kurnell Peninsula (Botany Bay) New South Wales. DESCRIPTION The newly recorded colony is characteristic, with its long cylindrical stalk, divided into basal holdfasts that are spread out over sea grass leaves (Posidonia), A few epibionts are on the stalk. The expanded head has scattered spots in the surface of the preserved specimen. FIGS, 12-22; Cystodytes ramosus n.sp. (holotype QM GHS374) — 12. colony; 13, section through upper part of colony showing contracted zooids beneath a layer of disc shaped spicules; J4, systems; 15, dise shaped spicules; 16, zooid. Palycitorella coronaria (QM GH5616) — 17, colony; 18, system from above: 19, thorax and anterior part of abdomen; 20, thorax showing atrial lip; 21, zovid showing muscle bands un abdomen, 22, gut loop and gonads, (Scales: 12 — 2em; 13, 17 — 2mm; 14 — 5mm; 15, 16, 20-22 — 0.5mm; 18, 19 — Itnm). 632 The zooids have the usual long thorax and abdomen. In this colony (collected at the end of September). There are up to 9 embryos in a de- velopmental series in the top of the abdomen. In some zooids the 2 best developed have moved up into the posterior end of the atrial cavity. The larval trunk is up to 0.9mm long, with the tail wound about three-quarters of the distance around it. Half of the length of the trunk is taken up with a large frontal lobe which, at its anterior end, supports 3 triradially arranged, large adhe- sive organs. Each consists of a small everting cone ina depression at the end of a thick cylindri- cal stalk. The cerebral vesicle contains a large ocellus and otolith. There are 3 rows of stigmata in the larval pharynx. REMARKS Larvae of this species previously were un- known. Kott (1990a) suggested that since larvae were not present in the known specimens (col- lected in January and April) the breeding season could be restricted. The newly recorded speci- MEMOIRS OF THE QUEENSLAND MUSEUM mens contain large numbers of relatively large embryos which appear to have been fertilised at the base of the long abdomen, and to have been developing as they move up the long stalk of the colony. This could be a lengthy process, and it suggests that sexual reproduction in this species could be an event that occurs once only during the year, The larvae, with their triradial adhesive organs and long frontal lobe, resemble the larvae of Polycitor giganteus, the tropical P. annulus, and P. circes (see Kott 199Qa). Polycitor obeliscus Kott, 1972 Polycitor obeliscus Kott, 1972b, p.171; 1990a, p.175. DISTRIBUTION New RecorbDs; South Australia (Nuyts Archipelago, SAM E2543-4). Victoria (Bass Strait, MV H454), RECORDED RANGE: The species previously was known only from its holotype from Investigator Strait. Its range is apparently restricted to the southern coast of FIGS. 23-26: Polycitor calamus (SAM E2530) — 23, zooid showing embryos in abdomen; 24, 25, larvae. Eudistoma aureum (SAM E2538) — 26, zooid with two embryos in alrial cayity, (Scales: 23, 26 — Imm; 24, 25 — 0.2mm). THE AUSTRALIAN ASCIDIACEA 2. SUPPLEMENT Australia from the Great Australian Bight to Bass Strail, DESCRIPTION EXTERNAL APPEARANCE: The newly recorded colonies are of various shapes. The specimen from Bass Strait (at 70m) is upright, egg-shaped and fixed by its narrow base. The other colonies are fixed by a broad base (to 3cm diameter), One is a Wide cone on the upper surface, but the other has a vertical margin and is flat-topped. Sand is present internally. itis absent from the surface of much of the upper part of the colony although occasionally, especially around the margins, or toward the base, some sand is present on the surface around the zooid openings. Atrial and branchial apertures open separately to the surface. In one colony the zooids seem arranged in pairs, but there are no systems, The test is white and the zooids are reddish-orange to flesh-coloured in preservative. INTERNAL. STRUCTURE: Zooids are large and robust, with almost continuous coat of external longitudinal and intemal circular muscles. The longitudinal muscles continue in 2 bands along each side of the abdomen. Stigmata are in 6 or 7 Tows of about 20. The stomach, in the expanded posterior end of the abdomen, is smooth-walled. The testis follicles are clustered in a large mass on the dorsal side of the gut loop, REMARKS The present species is distinguished mainly by the number of rows of stigmata and the smooth- walled stomach. Genus Eudistoma Caullery, 1909 Eudistoma aureum Kott, 1990 (Pig. 26) Eudistoma aureum Kott, 1990a, p.199 and synonymy. DISTRIBUTION New RECORD: South Australia (off Bucis, 170 to 190m, SAM B2538-9). RECORDED RANGE: The species was known only from ats Lype locality in Posidonie beds in St. Vincent Gulf. DESCRIPTION EXTERNAL APPEARANCE: The largest colony (SAM E2538) consists of upright lobes, possibly spherical in life, but shghtly laterally flattened in preservative, about 4cm diameter. It is fixed to the 633 substrate by a narrow basal part. Internally the colony contains grey sand throughout. Sand alsa is present in the surface test on the lower half, which may be buried in the substrate. In preserved colonies, the surface test on the upper part of the colony, which is free of sand, is translucent and greenish black with black zooids showing through it. The arrangement of the zooids is ob- scure in these colonies, INTERNAL STRUCTURE: The zooids are robust, with conspicuous muscles. about 50 internal transverse bands and 30 external longitudinal ones. Apertures are both 6-lobed. Stigmata are in 3 rows of 25 (middle row) to 30 (anterior) in each row, The dorsal end of the anterior row extends forwards along each side of the neural complex. In the large colony taken from off Eucla, up to 2 large embryos (trunk 1.25m long) are being incubated in the atrial cavity. They have 4 lateral ampullae along each side of the median adhesive organs. However, these are crowded, the larvae are opaque and other details could not be deter- mined, The tail is wound two-thirds of the way around the laryal trunk, REMARKS The pigmentation in the large colony is nore intense than in previously recorded specimens, which were said to have red-brown zooids in grey colonies (Kott 19902). Possibly the colour fades in long-term preservation, for other features of both zooids and colony such as the muscular zoojds, and the large number of stigmata per row, with the anterior row inclined along the mid-dor- sal line, are identical. Previously recorded colomes were said to have been yellow in life. Unfortunately the colour of the newly recorded ones is not known. Genus Exostoma Kott, 1990 Exostoma ianthinum (Sluiter. 1909) Polycitor ianthines Sluiter, 1909, p.20. Exostoma tanthinum, Kott, 1990a, p.234 and synonymy, DISTRIBUTION NeW RECORD: Queensland (Farnorthern Great Barrier Reef — Great Detached Reef, QM GH5726). RECORDED RANGE: The southern limit of the recorded range of this species was previously thought to be Matupore f£. New Guinea. It is known also from Palav Is, Philippines, and Bunda (Indonesia). 634 Suborder PHLEBOBRANCHIA Lahille, 1887 Family ASCIDIIDAE Adams & Adams, 1858 Genus Ascidia Linnaeus, 1767 Ascidia archaia Sluiter, 1890 Ascidia archaia Sluiter, 1890, p.346. Kott, 1985, p.26 and synonymy. Monniot and Monniot, 1987, p.92 and synonymy. Monniot, C. 1987a, p.18. DISTRIBUTION RECORDED RANGE: The species is known from Nor- folk Is, from north of Moreton Bay and northwards up the Great Barrier Reef to New Caledonia, the Philip- pines, Indonesia, French Polynesia, Fiji, the Marshall Is, and the southern islands of Japan; and from the northwest coast of Australia and south to Shark Bay. The tropical Atlantic species Ascidia corelloides (Van Name, 1924) appears to be inseparable from it (see Van Name 1945, Tokioka 1953). Nevertheless, the Western Pacific and Atlantic populations apparently are isolated from one another, neither having a range into temperate waters. REMARKS Monniot and Monniot (1987) do not accept the view (Kott 1985) that A. aperta Sluiter, 1904 is a synonym of the present species on the grounds that it is large, up to Scm long, with an open gut loop and with muscles on the right better devel- oped than in the smaller specimens they have assigned to A. archaia. None of the characters invoked by Monniot and Monniot are outside the range of variation observed in A. archaia, which is one of the most common species on the Heron I. reef flat, under rubble. It is known to reach a size of 5cm in less than 6 months (see Kott 1985). Ascidia liberata Sluiter, 1887 Ascidia liberata Sluiter, 1887, p.251. Kott, 1985, p.43 and synonymy. Ascidia solomonensis Nishikawa, 1986, p.61. Ascidia dorsalis C. Monniot, 1987a, p.9. Not Ascidia liberata: Sluiter, 1904, p.32. C. Monniot, 1987a, p.8. DISTRIBUTION RECORDED RANGE: The species is recorded from Nor- folk I. and the Great Barrier Reef, Indonesia (Sluiter 1887; Bay of Batavia, ZMA TU244), New Caledonia and Solomon Is. MEMOIRS OF THE QUEENSLAND MUSEUM DESCRIPTION The species is characterised by the naked, trans- parent test, pointed papillae on the siphons, the fringe of minute tentacular projections along the edge of the lobes surrounding the apertures, and tranverse muscles across the dorsal surface of the generally laterally flattened, short, often triangu- lar body, extending about half to two-thirds of the way across the right side but only about one-third of the way down the left. Short longitudinal mus- cles extend from along each siphon, those from the branchial siphon crossing the transverse mus- cles. The red colour at the anterior end of the body in the living specimens has been referred to by Kott (1985), Nishikawa (1986) and Sluiter (1887). REMARKS Monniot (1987a) refers to the poor condition of the type specimen (ZMA TU244). Nevertheless it was this type specimen that Kott (1985) exam- ined before assigning this common species to it. At the time she examined it, the characteristic papillae on the siphons, the tentacular fringe around the apertures, and the transverse muscles on the dorsal part of the right side of the body were all clearly visible. The general shape of this individual, which was figured by Sluiter (1887), closely resembles many of the more recently ex- amined specimens. Monniot, C. (1987a) examined and figured a specimen from the Siboga collection assigned to A. liberata by Sluiter (1904), Although Kott (1985) did not include it in the synonymy of A. liberata, Monniot believed the Siboga specimen to be identical with the type of A. liberata, and since it has characters inconsistent with A. lib- erata: Kott, 1985 he assigned the latter to a new species, A. dorsalis Monniot, 1987a. However, the Siboga specimen Monniot examined is not conspecific with A. liberata Sluiter, 1887. Its muscles, both on the right and left sides, as well as the swollen gut, the remnants of a fringe around the apertures and the foreign particles attached to the test, suggest that the Siboga specimen Mon- niot examined is A. sydneiensis. Kott’s specimens, for which A. dorsalis was erected, being conspecific with A. liberata Sluiter, 1887, A. dorsalis becomes a junior syno- nym of A. liberata. Similarly Ascidia solomonen- sis Nishikawa, 1986 has the characteristic features of A. liberata Sluiter, 1887. THE AUSTRALIAN ASCIDIACEA 2. SUPPLEMENT Family PEROPHORIDAE Giard, 1372 Genus Perophora Wiegman, [835 Perophora multictathrata (Sluiter, 1904) Ecteinascidia multiclathrata Siuiter, 1904, p.12. Perophora mutlticlathrata: Katt, 1985, p.106 and synonymy (part, not P. bermudiensis: pers. comm. L. Goodbody). Nishikawa, 1986, p.30 and synon- ymy; 1991, p.38.C, Monniot, 1987a, p.&8. DISTRIBUTION RECORDED RANGE: The species has a wide range in the Western Pacific from the Japan Sea (Sado), in both Micronesia and Melanesia as well as the northeastern coast of Australia, the Great Barrier Reef and Norfolk Istand. Perephora multiclathraia also occurs on the western coast of Africa (> Perophera africana Millar, 1953; fide Nishikawa, 1986) and in the West Indies (although itis nota synonym of PF. bermudiensis). The record from the Japan Sea is |he only one outside tropical locations. REMARKS It is of interest that a possible junior synonym of P. bermudiensis (P. fascia C, Monniot, 1991) oecurs in New Caledonia. Although Monniot has discussed the differences of P. fascia from the jemperate P. Autchisoni and tropical P. name, both with jointed stolons (see Kott 1985), he has not compared the morphology of his new species with either P. multiclathrata or P. bermudiensis, species with which it has more in common. If P. fascia is a synonym of P. bermueliensis, both P. mualticlathrata and P, bermudiensis are sympatric in the Western Pacific and the Auantic. Genus Ecteinascidia Herdman, 1880 Ecteinascidia diaphanis Sluiter, 1885 Ecteinascidia diaphanis Sluiter, 1885, p.168. Katt, 1985, p.90, Ecteinascidia koumaci C. Monniot, }987a, p,3. Ecteinascidia ndowwe C, Monniot, 199 1a, p.505. DISTRIBUTION RECORDED RANGE: The species is known trom north- em New South Wales, Greal Barrier Reef, Northen Australia, Palau Is, Indonesia, and New Caledonia. REMARKS Monniot has compared the New Caledonian material with E hedwigiae Michaelsen, 1918 from the coast of Natal, and with the Western 633 Pacific &. diaphanis. The latter, on geographical grounds is a more likely taxon to occur in New Caledonia. Differences between E. diaphanis and the 2 species from New Caledonia, viz. E. kow- maci and E. ndouae, appear to be either the result of intraspecific variation, or artefacts such as muscle contraction. They include variations in the red or pink-orange pigmentation at the anterior end of the zooids, which forms rings around the siphons, and sometimes spots as well (see Kott 1985). In E. ndeuae C. Monnivt, 199 1a, the col- our is described as yellow-gold. Ecteinascidia kounaci Monniot, 19874 ts dis- tinguished from E. diaphanis by the interval be tween the band of transverse muscles behind the atrial siphon and the siphonal muscles. This is affected by contraction of the longitudinal mus- cles alog the length of the siphon. The mem- brane between the dorsal languets, also said to distinguish the species, is not always present (Monniot, C. 1987a), There seems no valid justi- fication for the assignation of E. koumaci and £. ndouae to species distinct from E, diaphanis- Ecteinascidia nexa Sluiter, 1904 Exieinascidia nexa Siuiter, 1904, p.11, Kott 1985, 9.94 and synonymy. Nishikawa, 19286, p.42. Ecteinascidia aeywale C. Monmot, 1987a. p.5. Ecteinascidia imperfecta; Nishikywa 1984, p.42. DISTRIBUTION RECORDED RANGE: The species is recorded from Lore Howe L., the northeastern Australian coast and the Great Barrier Reef, Solomon 1s, Indonesia, Philipines, Fiji, Tokhara 1s, the Indian Ocean (Sri Lanka), and now from New Caledonia. REMARKS C. Monniot (1987a), comparing E. aeguale with E. nexa, was unable to reconcile the descrip- lions of Kott (1985) and Nishikawa (1986). How- ever, there are not significant differences in these accounts. The species is readily characterised hy its muscles, cloudy body wall in preservative, smooth stomach, duodenal and posterior stomach enlargements in the gut loop, hemispherical mass of male follicles and usually yellow colour in life. All of these characters are referred to by both Nishikawa (1986) and Kott (1985). Ecteinascidia nexa has a range of orientations of the sessile to stalked zooids and a variety of plimary and accessory stolons attaching them to the basal stolons or to one another. Apertures are, respectively, terminal and halfway along the 636 bedy, sometimes sessile, but usually on conical siphons when living. Kott (L985) describes the preserved zooids as flaccid and distorted, cloudy in preservative, with numerous blood cells. Nishikawa (1986) refers to the ‘lustreless’ free surface with pale yellowish spherules densely distributed in the body wall. Similar body mus- cles are described (in sore detail) by Kott (1985) and Nishikawa (1986). Stigmata are in 19 (Kott 1985) or 20 (Nishikawa 1986) rows, and there are from 14 to 24 Jongitudinal vessels on one side of the body. Both authors, independently, examined the types of P. hornelli Herdman, 1906, and proposed its synonymy, Nishikawa (1986) also examined the type of E. nexa. The branchial processes on the prepharyngeal band figured by Kott (1985, fig. 39h) only occur in the most robust zooids and are not charac- tenstic of the species, nor Was il suggested that they are. Ecteinascidia aequale C, Monnict 1987a from New Caledonia appears to fall within the range of yanation reported for E. nexa. Suborder STOLIDOBRANCHIA Lahille, 1887 Family STYELIDAE Sluiter, 1895 Subfamily STYELINAE Hentinan, 1881 Genus Cnemidocarpa Huntsman, 1912 Cnemidocarpa ampbora n.sp, (Figs, 27-31) DISTRIBUTION Tyre Locaury: South Australia (Sir Joseph Banks Group, North Point, Maram 1. 200m offshore in sand, rubble, Posidenta, 7m, coll. W. Zeidler 13.1.84, hoig- type SAM E2531; Sir Joseph Banks Group, W. Partney L., Pariney Shoal Reef, rubble, sand. Posidenia, 5— 1Sm, coll, W, Zeidler and K. Gowlett Holmes 218,86, paratypes SAM E2536, FURTHER RECORDS: None. DESCRIPTION EXTERNAL APPEARANCE: Individuals are club- shaped, with an expanded body, rounded anteri- orly and posteriorly continuous with a thick, cylindinical stalk about the same length as the body. The branchial and atnal apertures are di- rected away from one another in oppasite sides of the anterior end of the body, the branchial siphon wie terminal, and the atrial one on the dorsal side. MEMOIRS OF THE QUEENSLAND MUSEUM The available specimens are contracted, and behind the large, naked lobes of each aperture Is acollar or band of wrinkled test, When the animal is relaxed it is probable that these disappear, and that the siphons are long, projecting out like homs from the anterior end of the body. Sand adheres to the anterior part of the body around the base of the siphons, and to the fine hairs around the base of the stalk. The hairs ag- gregate together to forma sandy holdfast. The test is transversely wrinkled over most of the surface. {t is particularly thin on the base of the stalk where it is surrounded by the sandy holdfast, INTERNAL STRUCTURE: The body wall is red, shiny and muscular, Posteriorly it is extended into an inflated stolon which projects down into the stalk. Anteriorly the two siphons. are relatively long, the branchial siphon directed obliquely away from the substrate, the atrial one curyed over to the side toward the substrate, Attenuated body muscles are confined lo the anterior part of the stolon, which js thin-walled and hollow jn its distal part. The hollow may be part of the haemo- coele, as it does not appear continuous with the atrial cavity, being separated from it by a thin membrane, Longitudinal stripes line the siphons, and nu- merous fine branchial tentacles surround the base of the branchial siphon. The dorsal tubercle is a particularly large cushion with a double spiral shit, both homs rolled in, and directed either to the left or anteriorly. A large dorsal ganglion occupies the space between the base of the 2 siphons in these contracted specimens. The branchial sac has 4 low folds on each side of the body, with crowded internal longitudinal yessels arranged according to the formula DLO(18)2(10)3(12)3(10)2E. About 4 stigmata are in meshes in the interspaces, and more are between the endostyle and the first longitudinal vessel on each side. The gut loop is simple, and vertical, its pole compressed into the ¥V of the posterior end of the body where, in these contracted specimens, it projects slightly into the top of the stalk. The oesophageal opening is halfway down the dorsal side of the pharynx. The oesophagus is short, the stomach long with parallel internal longitudinal folds. The stomach and post pyloric part of the intestine (proximal to the pole of the gut Joop) are of about equal length. The rim of the anal opening (at the base of the atnal aperture) is deeply di- vided into 16 rounded lobes, On each side, up to 10, crowded, parallel gonads lie horizontally (at right angles ta the long axis of the body). They THE AUSTRALIAN ASCIDIACEA 2. SUPPLEMENT consist of a double rows, each row of about 10 large testis follicles, beneath a long ovarian sac. A very short oviduct is at the dorsal end of each ovary. The short vas deferens opens on top of the female duct. Irregular, upright endocarps are crowded along the dorsal one third of each side of the atrial cavity (between the dorsal ends of the gonads, and dorsal to them). REMARKS Both Cnemidocarpa floccosa (Sluiter, 1904) and C. completa Kott, 1985 have a similar num- ber of cnemidocarp gonads on each side of the body as the present species. The former tropical species is rounded posteriorly, while the present one narrows to its stalk; the gonads lie vertically, parallel to the long axis of the body in C. floccosa, rather than at right angles to it as in C. amphora; and the gut loop in the present species lies verti- cally, extending back from the oesophageal open- ing, while that of C. floccosa curves anteriorly around the ventral margin of the body. Cnemido- carpa completa Kott, 1985 is known from tem- perate waters, although not from South Australia. It also is distinguished from the present species by the curve of the gut loop around its posterior ventral border, the absence of a dorsal band of endocarps, the long stomach, and the loose at- tachment of gonads to the body wall. Cnemidocarpa areolata (Heller, 1878) Styela areolata Heller, 1878, p.26. Cnemidocarpa areolata: Kott, 1985, p.122 and synon- ymy. Not Monniot, 1983, p.451 ( P_ vittata: Monniot, 1989, 1991) only by the presence of the swollen recturn in the latter species. He remarks that “tous les autres caractéres sont peu importants et les différences observées entre les espéces pourraient cotrespondre 4 une variabilité’. However, there ire significant differences in a number of con- spicuous and stable characters between these two species. The principal difference is the form of the siphonal spines (Kott 1985). In P. curvigona these have a flat scale halfway along them, which is not present in P. albanyensis, In P. albanyensis the needle-like spines are of two sizes. Further differences are the short dorsal lamina, the long, deeply curyed U-shaped gut loop and the distinct longitudinal stripes down the outside of the si- phons in P, albanyensis. The species are similar only in the crowded endocarps on gut loop and gonads, in the almost pinnate form of the ovarian tube, each short branch with a cap of male folli- cles, and in the distribution of the siphonal spines, which continue over the outer surface from inside the siphon. Monniot (1989) assigned 2 specimens from New Caledonia, one 5cm and one 1.2cm long, to P. albanyensis. In neither were the small spines between the larger ones reported, nor can they be detected in Monniot’s scanning electron mi- crographs. It is possible that these have been obscured by the larger spines, although this does not seem to be the case. The smaller spines, like the larger ones, are on the surface of the test both inside and outside the siphons, and should be apparent if they are present. Further, although the gut loop and gonads have crowded endocarps on them, and the long ovarian tubes have pinnale branches, each with a cap of terminal testis follicles, the New Caledonian specimens differ [rom P, albanyensis in having a long dorsal lamina and a narrow but gently curved gut loop. The form of the siphonal armature needs to be established by light microscope examination of cleared preparations in order to accurately assign the New Caledonian specimens, Monniot(1989) drew attention to the fact that the name Hartmeyer applied to the holotype of this species is preoccupied by the Antartic Pyura obesa Sluiter, 1912. Pyura curvigona Tokioka, 1950 Pyura curvigona Tokioka, 1950, p.147. Kou 1985, p.296 and synonymy, ?Pyura viltala: Monniot, 1989. p.490. 2Pyurg watio Monniol. 199 1b, p.33, DISTRIBUTION RECORDED RANGEL: The specics is known from Shark Bay (Western Ausiralia), Heron and Lizard |. (Queens- Jand), the Palau Js, Indonesia (sce Kott 1985), and possibly from New Caledonia. 646 REMARKS Monniot (1989) has synonymised this species with the Atlantic Ocean Pyura vittata. He appears to have overlooked the opalescent scale which is such a feature of the siphonal armature of the present species (see Kott 1985, Figs 5k, 145a,b). Since the New Caledonian specimens assigned to P. vittata have the enlarged rectum, pinnate go- nads, and endocarps of the present species it is possible that they are conspecific. Examination of cleared preparations of siphonal armature by light, rather than by scanning electron, micros- copy, would, if they are present, display the scales of the siphonal spines which are just under the surface of the test. The green liver and pink body wall of this species is found also in P. uatio from New Cale- donia which, however, lacks an expanded rectum, and may be a different species. Pyura rapaformis Kott, 1990 (Figs 45, 46) Pyura rapaformis Kott, 1990b, p.287. DISTRIBUTION NEw RECorD: South Australia (Beachport Jetty piles, SAM E2510). RECORDED RANGE: The species, previously recorded from the upper surface of a reef at Cottesloe, probably has a continuous range between Cockburn Sound and St Vincent Gulf. DESCRIPTION The present specimens are as previously de- scribed, with both apertures on the upper, more or less flattened, surface of a turnip-shaped body. The surface test has irregular extensions and pa- pillae to which sand adheres, and sand also is embedded in the test. The body has a posterior stolon projecting down into the test. The siphonal spines are about 0.08mm long, with a long narrow opening and 4 rounded knobs at the posterior end of the shaft. Branchial tentacles are feathery, with long primary branches. The branchial sac has 6 wide, overlapping folds on each side with 7 to 18 internal longitudinal vessels on the folds and up to 3 between. Leafy, irregular endocarps are on the gut and gonads. REMARKS Externally the species resembles individuals of Pyura isobella Kott, 1985 from Ulladulla (NSW) and internally it has similar endocarps and gut loop and gonads. The siphonal spines lack the MEMOIRS OF THE QUEENSLAND MUSEUM posterior expansion, and the small opening of spines of P. isobella, and more closely resemble the spines of P. tasmanensis Kott, 1985 (as ob- served by Kott 1990b). Genus Herdmania Lahille, 1888 Herdmania momus (Savigny, 1816) Cynthia momus Savigny, 1816, p.143. Herdmania momus: Kott, 1985, p.338. Pyura momus: Monniot, 1989, p.498. DISTRIBUTION RECORDED RANGE: The species is pantropical, extend- ing into temperate waters of southern Africa and south- ern Australia. REMARKS Monniot (1989) believes the genus Herdmania to be a junior synonym of Pyura. In addition to the characteristic long barbed spines, Herdmania has acontinuous ovarian tube surrounded by male follicles, rather than being broken up into separate gonad blocks as in Pyura; and the liver lobules are embedded in the body wall, rather than lying free in the atrial cavity as in Pyura (see Kott 1985). Genus Microcosmus Heller, 1877 Microcosmus tuberculatus Kott, 1985 Microcosmus tuberculatus Kott, 1985, p.361. ? Mon- niot, 1991b, p.31. DISTRIBUTION RECORDED RANGE: The species is known for certain only from its type locality in the Swain Reefs. REMARKS The species is distinguished from M. exaspera- tus by its large conspicuous conical spines around and lining the apertures (see Kott 1985, Fig. 179b) and the deeply incised anal border. The specimen from New Caledonia (Monniot 1991) has neither of these features. The thick endocarp- like layer of the body wall Monniot observed over the gonads and gut occurs variably in most Microcosmus species, including M. exasperatus, to which the New Caledonian specimens very likely belong. THE AUSTRALIAN ASCIDIACEA 2. SUPPLEMENT 647 FIGS. 45, 46: Pyura rapaformis (SAM E2510) — siphonal spines (Scales: 45, 0.05mm; 46, 0.1mm). Family HEXACROBYLIDAE Sceliger, 1906 The family was reviewed by Kott (1989s), and later, by Monniot and Monniat (1990). The family is characterised by its large buccal cavity, surrounded by 6, usually large, branched branchial arms, a large kidney, and a reduced perforated pharynx, limited to a short band at the proximal end of the gut. Ciliated stigmata are small and rounded. The gut is well developed with a wide, and often long oesophagus, and there is evidence of adaptation for a carnivorous habit (see Kott 1989a). Although, successively, a new pleurogonid suborder, Aspiraculata Sccliger, 1906, and a new tunicate class Sorberacea Mon- niot, Monniot and Gaill, 1975 haye been pro- posed for the family, there uppears to be no justification for cither of these higher taxa {sec Kott 1989a). The Hexacrobylidae have adaptations for deep water habitats, but their morphology is homolo- gous with that of Molgulidae, with which they appear to have closer affinities than the latter family has with other extant ascidian taxa, even within the Pleurogona (see Kott 1989a). The view (Monniot and Monniot 1990) that the family name Hexacrobylidae should be aban- doned in favour of Oligotrematidae (sic. Oligotremidae Monniot and Monniot, 1990) is incorrect (ICZN Article 40). Further, the type genus is Hexacrobylus Sluiter, 1905a, a junior objective synonym of Oligotrema Bourne, 1903, Although Kott (1985) thought the family was nol represented in Australian waters, Asajirus indicus Oka, 1913 and Oligotrema psammites Bourne, 1903 have since been recorded (Kott 1989a). Genus Asajirus Koll, 198% Type species: Hexacrobylus indicus Oka, 1913. Kot (1989a) examined and redescribed the type specimen (ZMA TU564) of Hexacrobylus psanmmatodes Sluiter, 1905a, and found it to be a species of the genus Oligatrema Bourne 1903, possibly conspecific with its type species 0, psanmnites Boume, 1903 from New Britain. Thus was corrected a belief that Hexacrabylus pserm- matodes and AH. indicus Oka, 1913 were congen- eric, and A. indicus Oka was found to be a member of an undescribed genus. Hexacrobylus indicus Oka, 1913 was therefore designated type species of a new genus Asvjirus Kott, 1989a (of which Heyaductylus Monniot and Monniot, MEMOIRS OF THE QUEENSLAND MUSEUM 1990, with the same type species as Asojirus, is a junior synonym). Monniot and Monniot (1990) recognise 12 spe- cies in the genus Asajirus, distinguished from one another primarily by differences in the length of the vas deferens and in |he number and length of testis lobes. It may be difficult to sustain this number of species, in view of the probability that there is some intraspecific variation in taxa which. in many cases, have vast geographic ranges. The differences are leas compelling than the similarities between most of the species Monniot and Monniot (1990) have documented, and many are sympatric in at Jeast part of their range, while some are over their whole range. For instance, A. longitestis (Monniot and Monniot, 1990), A. oyi- rarus (Monniot und Monmiot, 1990), A. hemi- sphericus (Monniot and Monmot, 1990), A. gulosus (Monniot and Monniot, 1984b) and A. indicus (Oka, 1913) are all recorded from the vicinity of New Caledonia, Asajirus millari (Monniot and Monniet, 1991) also is from the West Pacific (the Bali Sea); A. seeligeri (Monmot and Monniot, 1990) and A. gulosus are from the Comoro Is in the Indian Ocean and some records of A, indicus are also from the Indian Ocean; A, ledanoisi and A. ewauchus ave fron wide ranges in the Atlantic, and A. arcticus is trom the Nor- wegian Sea; while A. antarcticus is recorded from avound the Southern Ocean and A. dichotomus is from the southeastern Atlantic (the Cape Basin). It is not intended to review this genus in the present work, rather attention is drawn to some of the problems of species definition in this family. These will not be resolved if descriptions do nol take account of the homologies of organs. of similariues in morphology as well as morpho- logical differences, of probable artefacts resulting from contraction and distortion, and of differ- ences that most likely arise from growth. and miaturily. Asajirus indicus (Oka, 1913) Hexucrobylus indicus Oka, 1913, 9.6. Millar, 1959, p.203; 1969, p.97; 1970, p,146; 1975, p.332. Mon niot and Monniot, 1968, p.32; 1970, p.334; 1979, p.564; 1984b, pp.198 (figs 1,2) 203; 1987, p.4l. Monniot, F., 1971, p.458. Asajirus indicus; Kott, 1989a, p.521- Hesxadacivlus indicus: Monniot and Monniot, p.27l, 1990, THE AUSTRALIAN ASCIDIACEA 2. SUPPLEMENT Aexacrobylus arcticus Hartmeyer, 1923, p.133, Man- niot and Monniot, 1984b, p.203 (specimens from Norbi cruise). Hexadaetylus arcticus; Monniot and Monniot, 1990, p.261, Hexacrobylus ewnuchus Monniot and Monniot, 1976, p.658. Hexadactylus eunrchus: Monniot and Monniot, 1990, p.265, Hexacrobylus sp. Monniot and Monniot, 1982, p,12& (part, Eltanin cruise 5, station 258). Oligotrema psammites: Kott, 1969, p.168. Aexadactylus antarcticus Monniot and Monniot, 1990. p.259. Hexadactylus longitestis Monniot and Monniot, 1990, p.275. Hexadactylus millari Monniot and Moniot, 1991, p.277. Mexadactylus seeligeri Monniot and Monniot. 1990, p.281_ DISTRIBUTION New Recorps: Coral Sea (12° 05° 5S 146° 24'E, Cidaris 3 site 6.1, 16.2,92, 3319-3105m, QM Gi00891, 13° 29°S 147° 13°E Cidaris 2 Site 15.1, 5.9.88, 2542-2457, QM G300892. RECORDED RANGE: The recorded range for A. indicus and its probable synonyms is: A. longitestis — New Caledonian (both sides of the Norfolk Ridge); A. in- fens —the Indian Geean, New Caledonia, off casicra Australia from 151°27' to 148°38"E and 12° to 42°S from the Coral Sea to eastern Tasmania; A, seeligeri — the Indian Ocean and Comoro fs.; A. niillari — the Bali Sea (western Pacific); A. eunuchus — southwest- em and northern Atlantic; A. arcricus — northeastern Atlantic; A. antarcticus — South Indian and Pacific Oceans. The species is found at depths from 1000 ta 5000m. The record for the specimen AM Y2125 Irom off the New South Wales coast, is from 151°27°E, not §1°27'E ay reported by Koti (19894). DESCRIPTION The newly recorded specimens are rounded posteriorly, 1.2 cm long, but contracted anteri- arly, causing the fine hairlike projections mixed with foraminifers and other paricles to appear more crowded than they are posteriorly, Inter- nally the 2 ventral muscle bands are short and broad, the rectum is crowded with hard particies including foraminifers, polychaetes, ostracods, cladocerans. isopods, gastropods and what ap- pears to be particles of ophiurid (including spines), The remainder of the gut (the long oc- sophagus and stomach) are empty. 49 REMARKS Specimens from the northern Atlantic (A. ledanoisi) and from the Cape Basin, south-east- em Atlantic (A. dichotomus) with long sperm ducts ate not included in the above synonymy of A. indicus, although A. eunuchus (Monniot and Monniot, 1976) from the south-western and northem Atlantic, A. arcticus (Hartmeyer. 1923) from the Norwegian Ses, A. antarcticus (Monniot and Monniot. 1982) from the southern (polar) Indian and Pacific Oceans and A, millari from the: Bali Sea have shor! male ducts as in A, indicus, and are here included in its synonymy, as ts A. seeligeri (with both short on long male follicles) from the Indian Ocean, and A. longitestis {with long male follicles) from New Caledonia. Ase- Jjirus indicus (Oka, 1913).as well as being taken from off the eastern Australian coast, is from the Indian Ocean and New Caledonia. Also from New Caledonia, and, for the present, excluded from the synonymy of A. indicus, arc Asajirus hemisphericus and A. ovirarus, with more or less entire testis; and A. gulosus with a long vas deferens like A. ledanoisi and A, di- ehotanwass. Genus Oligotrema Boume, 1903 Type species: Oljgotrema psammites Bourne, 1903, Although not designated a neotype, Monniot and Monniot (1990) redescribed the species Irom a Specimen taken from near the type locality. Monniot and Monniot’s definition of the genus omits certain significant characteristics described by Bourne, which also are present in the type specimen of Hexacrobylus psammatodes Sluiter, 1905a (see Kott 19892), Monniot and Monniot (1990) refer to the widely separated apertures but notto the posterior position and orientation of the atrial aperture; they refer ta muscles extending the length of the body wall, but fail to indicate that these are circular muscles, sometimes inter- rupted; and that longitudinal muscles are very limited; nor do they refer io the flagellated spi- thelial covering of the branchial tentacles de- scribed by Bourne (see also Kott 1989a), Monniot and Monniot (1990) indicate, as Kott (1989a) had suggested, that the branchial stig- mata ate circular perforations, rather than the rectangular stigmata described by Bourne. This removes the principal restraint to the synonymy of H. psammatodes Sluiter, 1905a with the pre- sent species. Other characters, viz. the branchial 650 tentacles, the large endocarps and the large atrial cavity, are as described by Bourne, and by Kott (1989a) for the type specimen of H. psammatodes Sluiter, 1905a (ZMA TUS564). There is no additional evidence regarding the proposed synonymy of Gasterascidia, Sorbera with Oligotrema (see Kott 1989a). Nor is there any clarification regarding the apparent degen- eration of the rectum in Gasterascidia lyra, which, if present, appears to be non-functional (see Monniot and Monniot, 1990 Fig. 5B). Oligotrema psammites Bourne, 1903 Oligotrema psammites Bourne, 1903, p.233. Monniot and Monniot, 1990, p.253. Hexacrobylus psammatodes Sluiter, 1905a, p.135; 1905b, p.135. Hexacrobylidae juv., intermediate between Sorbera and Gasterascidia: Monniot and Monniot, 1984b, p-212; 1990, p.257. ?Sorbera digonas Monniot and Monniot, 1984b, p.209. Oligotrema psammatodes: Kott, 1989a, p.529. Mon- niot and Monniot, 1990, p.258. DISTRIBUTION RECORDED RANGE: The species is recorded from New Britain (Bourne 1903; Monniot and Monniot 1990), Indonesia (ZMA TU564, Siboga, Station 211, Banda Sea, Sluiter 1905a), west of New Caledonia (Monniot and Monniot 1990); New South Wales coast (Kott 1989a) and the south-east Atlantic (the Cape Basin, Monniot and Monniot 1984b). Sorbera digonas is recorded from the tropical Indian Ocean. REMARKS The specimens all have the shallow ventral groove in the pharynx; vertically flattened bran- chial tentacles with one or 2 short, pointed, pro- jections anteriorly, an otolith-like black granule in the neural gland, and the parietal wall of the atrial cavity raised into long endocarps on each side of the oesophagus. These endocarps some- times cover part of the gonads. In the thick pha- ryngeal wall the interconnected chambers (that the Monniots in 1984 and 1990 refer to as branch- ing tubules) eventually open to the atrial cavity by small, rounded ciliated stigmata which vary in number according to the size of the specimen. The ligament attaching the oesophagus to the body wall (Kott 1989a) appears to be the ventral sep- tum dividing the atrial cavity into two sections (Monniot and Monniot 1990). In fact, the atrial MEMOIRS OF THE QUEENSLAND MUSEUM invagination is always from the dorsal surface and the cavity always is continuous over the dorsum. It never penetrates ventral to the phar- ynx, past the ventral sinus. The ligament or sep- tum is merely the ventral body wall where it is not interrupted by the atrium. LITERATURE CITED BREWIN, B.I. 1956. The growth and development of a viviparous compound ascidian, Hypsistozoa fasmeriana, Quarterly Journal of Microscopical Science 97: 435-454. 1958. 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Bulletin of the American Museum of Natural History 84: 1-476. 654 MEMOIRS OF THE QUEENSLAND MUSEUM INDEX TO TAXA (Taxon descriptions and figures in bold) Amaroucium distomoides, 627 valborg, 637 Aplousobranchia, 621 623 Cynthia momus, 646 Asajirus, 648 Cystodytes, 629 antarcticus, 648 649 arcticus, 648 649 dichotomus, 647 649 eunuchus, 648 649 gulosus, 648 649 hemisphericus, 648 649 indicus, 648-649 ledanoisi, 648 649 longitestis, 648 649 millari, 648 649 ovirarus, 648 649 seeligeri, 648 649 Ascidia, 634 aperta, 634 archaia, 622 634 corelloides, 634 dorsalis, 634 liberata, 634 solomonensis, 634 sydneiensis, 634 Ascidiidae, 634 Aspiraculata, 648 Asterocarpa, 640 humilis, 640 Botryllinae, 643 645 Botryllus, 643 stewartensis, 643 645 Cionidae, 622 623 Clavelina, 625 baudinensis, 625 diminuta, 625 pseudobaudinensis, 625 Clavelinidae, 625 627 Cnemidocarpa, 636 amphora, 636-637 639 areolata, 622 637-638 completa, 637 floccosa, 637 aff. humilis, 640 intestinata, 638 irene, 637 legali, 637 lobata, 640 pedata, 640 stolonifera, 640 tribranchiata, 638 639 tripartita, 640 dellachiajei, 629 ramosus, 629 630 Diazona, 623 Diazonidae, 622 623 Didemnidae, 621 Distaplia, 627 628 prolifera, 627 Ecteinascidia, 635 aequale, 635 636 crassa, 623 diaphanis, 635 hedwigiae, 635 imperfecta, 635 koumaci, 635 multiclathrata, 635 ndouae, 635 nexa, 635-636 Eudistoma, 633 aureum, 632 633 fantasiana, 626 Exostoma, 633 ianthinium, 633 Gasterascidia, 650 lyra, 650 Herdmania, 646 momus, 646 Hexacrobylidae, 648 650 Hexacrobylus, 648 arcticus, 649 eunuchus, 649 indicus, 648 psammatodes, 648 649 650 sp., 649 Hexadactylus, 648 antarcticus, 649 arcticus, 649 eunuchus, 649 indicus, 648 longitestis, 649 millari, 649 seeligeri, 649 Holozoidae, 626 627 Hypodistoma, 628 Hypsistozoa, 627 628 distomoides, 627 THE AUSTRALIAN ASCIDIACEA 2. SUPPLEMENT 655 Microcosmus, 646 exasperatus, 646 tuberculatus, 646 Molgulidae, 648 Neodistoma, 628 Oligotrema, 648 649 650 psammatodes, 650 psammites, 648 649 650 Oligotrematidae, 648 Oxycorynia, 625 pseudobaudinensis, 625 Perophora, 635 africana, 635 bermudiensis, 635 fascia, 635 hornelli, 636 hutchisoni, 635 multiclathrata, 622 635 namei, 635 Perophoridae, 623 635 Phlebobranchia, 621 623 634 Pleurogona, 648 Polyandrocarpa, 641 abjornseni, 643 colemani, 641 642 643 644 colligata, 643 maxima, 643 robusta, 643 simulans, 643 Polycarpa, 640 aurita, 622 640-641 circumarata, 640 contecta, 641 insulsa, 640 Polycitor, 631 annulus, 632 calamus, 631 632 circes, 632 giganteus, 632 ianthinus, 633 obeliscus, 632 Polycitorella, 629 coronaria, 629 630 631 644 Polycitoridae, 629 Polydistoma, 625 Polyzoinae, 623 Protoholozoa, 627 anthos, 627 628 australiensis, 624 625 628 cantarella, 627 628 Tilium, 627 628 pedunculata, 627 628 pigra, 627 628 Protopolyclinum, 623 claviformis, 623 Pseudodiazona, 623 claviformis, 623 624 Pseudodistomidae, 627 Pycnoclavella, 625 arenosa, 625 aurantia, 624 625 detorta, 625 diminuta, 624 625-626 elongata, 626 tabella, 625 Pycnoclavellidae, 625 Pyura, 645 646 albanyensis, 645 curvigona, 645 isobella, 646 momus, 646 obesa, 645 rapaformis, 646 647 tasmanensis, 646 uatio, 645 646 vittata, 645 646 Pyuridae, 645 Rhopalaea, 622 623 crassa, 623 respiciens, 623 Sigillina, 626 627 fantasiana, 626 grandissima, 626 Sorbera, 650 digonas, 650 Sorberacea, 648 Stolidobranchia, 621 636 Stylea, areolata, 637 aurita, 640 contecta, 641 humilis, 640 Styelidae, 636 Styelinae, 636 Sycozoa, 627 628 CONTENTS KOTT, P. The Australian Ascidiacea part 3, Aplousobranchia (2) 375-620 KOTT, P. The Australian Ascidiacea, Supplement 2 “621-655