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


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


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... 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... .- ees eee ce ee cececcecce. nb ds sath « 302



The Australian Ascidiacea part 3, Aplousobranchia (2) KOTT, P.

The Australian Ascidiacea, Supplement 2 ...... 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)


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



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.


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... .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



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) 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


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-


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.


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).


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


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


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


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-


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


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