VOL 96, PART 1 29 FEBRUARY, 1972 TRANSACTIONS OF THE ROYAL SOCIETY OF SOUTH AUSTRALIA INCORPORATED CS eT ee CONTENTS Kott, Patricia’ The Ascidians of South Australia I. ge Gulf, St. Vincent Gulf and Encounter Bay - - 1 Harris, Wayne K. New form species of pollen from southern Australian early Tertiary sediments - - - - - - - - 53 Sa a a I ST SRE EIS RS PUBLISHED AND SOLD AT THE SOCIETY’S ROOMS STATE LIBRARY BUILDING NORTH TERRACE, ADELAIDE, S.A. 5000 TRANSACTIONS OF THE ROYAL SOCIETY OF SOUTH AUSTRALIA INC. VOLUME 96, 1972 PART 1, 29 FEBRUARY Kott, Patricia The Ascidians of South Australia I. BESESETES Gulf, St. Vincent Gulf and Encounter Bay - . x 4 2 a Harris, Wayne K. New form species of pollen from southern Australian early Tertiary sediments - - - : - = = s PART 2, 31 MAY Preiss, W. VY. The systematics of South Australian Precambrian and Cambrian Stromatolites. Part 1 - - - - = 2 f 4 Mawson, Patricia M. The nematode genus Maxvachonia (Oxyurata: Cosmo- cercidae) in Australian reptiles and frogs - - - - Mawson, Patricia M. Three new species of the genus Cloacina Linstow (Nema- toda: Strongylata) from macropod marsupials - - - Schodde, R., Mason, I., & Wolfe, T. O. Further records of the Pitted-shelled Turtle (Carettochelys insculpta) from Australia - - - Wollaston, Elise M. The morphology and relationships of Muellerena wattsti (Harvey) Schmitz (Ceramiaceae: Rhodophyta) - - - PART 3, 31 AUGUST Smith, Meredith J. Small fossil vertebrates from Victoria Cave, Naracoorte, South Australia. Il. Peramelidae, Thylacinidae and Dasyuridae (Marsupialia) - - - - - - - - - Mawson, Patricia M. The genus Acuwaria Bremser (Nematoda: Spirurida) in Australia - - - - - = = = = > Milnes, A. R., & Bourman, R. P. A Late Palaeozoic glaciated granite surface at Port Elliot, South Australia - a - . 3 : * Tyler, M. J., & Parker F. Additions to the hylid frog fauna of New Guinea, with description of a new species, Litoria timida - - - - PART 4, 30 NOVEMBER Kott, Patricia The Ascidians of South Australia II. Eastern Sector of The Great Australian Bight and Investigator Strait - - & + x Dubois, Georges, & Angel, L. Madeline Strigeata (Trematoda) of Australian Birds and Mammals from the Helminthological Collection of the Uni- versity of Adelaide - - +! - - - - - OBITUARY: KEITH RODNEY MILES - = B 2 ¢ 2 2 Annual Report of Council, 1971-72 - = = 2 4 s - 3 Award of the Sir Joseph Verco Medal - - = 7 > 4 a = Balance Sheet - - - - - 2 - : : = 2 5 List of Fellows - : = 2 “ 2 3 = x m 4 ~/ 53 67 101 100 115 119 125 139 149 157 165 197 216 220 221 222 223 THE ASCIDIANS OF SOUTH AUSTRALIA I. SPENCER GULF, ST. VINCENT GULF AND ENCOUNTER BAY BY PATRICIA KOTT Summary A large and representative collection of Ascidiacea from St. Vincent Gulf and adjacent locations is discussed. Fifty-nine species are represented, of which Pyura scoresbiensis and Ctenicella antipoda are new to science. Ascidia aclara Kott, previously known from other Australian locations, and Aplitiium colelloides Herdman, previously known only from South Africa, are recorded from the area for the first time. The fauna of St. Vincent Gulf is typically of the Flindersian marine biogeographic region, but includes several endemic species. Morphological characteristics accounting for the success of certain species and groups of species sharing a habitat are indicated. THE ASCIDIANS OF SOUTH AUSTRALIA IL SPENCER GULF, ST. VINCENT GULF AND ENCOUNTER BAY by Patricia Kott* Sutmnmary A large and representative collection of Ascidiacea from St. Vincent Gulf and adjacent locitions is discussed. Fifty-nine species are represented, of which Pyyra scoreshiensix and Crenicella antipoda are new to science. Ascidia aclara Kott, previously known from other Australian locations, and Aplidium colelloides Herdman, previously known only from South Africa, are recorded from the are for the first time, The fauna of St. Vincent Gulf is typically of the Flindersian marine biogeographic region, but includes several endemic species, Morphological characteristics accounting for the success of certain species and groups of specics sharing a habitat are indicated. Jntroduction This large collection of ascidians, mainly from St. Vincent Gulf, South Australia, was made by Mr. 8S. A. Shepherd of the Depart- ment of Fisheries and Fauna Conservation, South Australia. It is a valuable and represen- tative collection and demonstrates the value of SCUBA collections of this benthic group from otherwise inaccessible localities. Colour notes made by the collector provide most useful data for comparison with the preserved specimens in which colours are generally lost or change completely. The large number of individuals of most species that are available in the col- lection has demonstrated a wide variability in certain characters and some synonymy has been established. Information on the environmental conditions operating in various locations, also supplied by the collector, has been related to the morpho- logy of the species prescnt to contribute to an assessment of selective mechanisms affecting the ascidians. Full station lists of species are also given to facilitate consideration of the faunal associations and their ecological rela- tionships. The specimens are deposited in the South Australian Muscum. The following species have previously been recorded from South Australia (Kott 1952, 1957a, 1962, 1963) but were not in the present collection. Polyclinum nepiunium Polyclinum marsupiale A plidium flavolineatum A plidium australiensis Lissoclinym ostrearium Dideninum turritum Didemnurm angusti Didemnum pseudediplosama Trididenmum nataletise Trididemnum cerebrijorme Leptaclinides imperfectus Syniplewma viride Styela lohata Asterocarpa cerea Pyura stolonifera Zoogeography The fauna is typically that of the Flindersian marine region, together with Distaplia viridis which is also recorded from Part Phillip Bay, Ascidia aclara which has been taken from simi- lar sheltered locations on the Victorian, New South Wales, and Queensland coasts, and Aplidium colelloides, previously recorded from South Africa. The new species, Pyura scores- hiensis and Ctenicella antipoda, may be endemic. The records of A. colelloides from off South Africa and South Australia suggest a circum- polar distribution, as demonstrated for many ascidian species (Kott 1971a). A wide dis- persal of larvae, however, does not provide a satisfactory explanation for this pattern of dis- * Zoology Department, University of Queensland, Qld. Australia 4067. Trans. R. Soc. S. Aust. 96, Part 1, 29 February 1972, 4 tribution since, for successful sexual reproduc- tian, minimal population densities of adults are required. The existence of so many circum- polar species in the extant fauna may be the result of a slow rate of evolution and the per- sistence of relict forms in certain areas. Habit of the Ascidian Fauna In the present collection, ascidians have been taken frum a wide variety of locations, espe- cially in St, Vincent Gulf. The terminology qualifying the conditions encountered is partly that described by Shepherd & Womersley (1970) and Womersley & Edmonds (1958), as fol- lows: (1) “Reugh Coast Subformation” (R.C.S.) refers to coasts exposed to the southern ocean swell (wave periods LO-12 secs.). tere hl tlt MISTRESS ein 5 me Fig. |. Map showing i cape Spences PATRICIA KOTT Water movement resulting from this swell is strong and pulsatile on the surface but decays with depth so that surge is moderate at 15 m and slight at 25 m depth. (2) “Sheltered Coast Subformation” (S.C.8.) (see Womersley & Edmonds 1958) refers to sheltered coasts where there is no swell and the coast is subject to waves of short period (up to 5 seconds) which decay rapidly with depth. Much of the coast-line in both Spencer Gulf and St. Vincent Gulf is of this type. (3) “Offshore Benthic” locations are those away from the shore where water movement results from tidal current rather than wave action. In St. Vincent Gulf tidal currents are generally about | m/sec., except over Tapley Shoal where they ate 1-2 m/sec. a 138° 9° 1% — = we “ , aa? ki An f+. fe ia Bite” .Y : ae aly vald ( 34° vajfera & \ SLE of BY = Snuse la dang . yORN) FINTRSULA a 3 Outer Harkour emaphore as Hoan ‘ « wipregntan 35 getenies Shral mn Or evaneae Sate Habiert Cave TWmarion keer Port Sbanvar te ae Trowbridge Foent Mart Naaeauigs® Eas -* ave * tn syAidinga arickalinga Wea anhais)ig LAUR eR el ght Yeotar nage, Lo] an + yathour & pet Tstand wos a ee locations in and adjacent to Spencer Gulf and St. Vincent Gulf, ASCIDIANS OF SOUTH AUSTRALIA These different locations. provide environmental conditions favouring an ascidian fauna of very varied habit. For each species its shape, or size. or mode of fixation, or growth characteris- tics appear to operate as selective mechanisms contributing to its success in the environmental conditions operating: (1) Jn Rough Coast Subformations, at depths less thin 15 m where surge is moderate to strong (Wright b, West Li, the dominant ascidian fauna is adapted ro the conditions by virtue of their colonial form, their vivi- parous larvae, their usually well-developed ¢loaval systeros, and cither (ad un encrusting habit providing a large surface area for fixation (Atapozoa fanpea- siene, Cyxtodytes deHechiajel, Dideninum crendidum, Leptoctinides rufus, Lixsoclinisanr sf. Oeulinaria australis. — Barrylleides nigrwen); ar (G} small stalks or sessile habit and cylin- drical body form enabling them to occupy sheltered crevices (e.g. on Frklonig hold- fasts. under Jedges, etc.). The stalks of these species are thick, and the colonics da not move freely with the currents (Podectevella cylindrica, Pseudodistomia cereum, Rilterelly herdimania, Syroicium papiliifertum) . (2) In Rough Coast Subformations, at 15 m und greater depths (Weight 1. West T.}, the strong surge at the surface ts reduced to mode- rile te slight water movement. Species with pliable stalks form a daminant component of the 4seidian fauna and are best able to exploit the changing direction of the water movements by moving with the water so that their bran- chial openings ate presented to the oncoming eurrent which thus reinforces the ciliary feed- ing mechanism. Only some ot these species have viyipurous lurvae (Borrylloides magni- coecus, B. leach, Palyearpa pedunculata, P. vlavata, Pyuru auytralis), (3) In Rough Coast Subformations, ut all depths, are large species fixed by a relatively small part of their surface. At shallow depths, they appear to be more offen on vertical rock fuces or in caves, where firm fixation can be uchieved, while at greater depths they are on the bottom (Polyciror giganteunt, Stvela pedata, Cremidecarpa erherldei, Herdntania mamus). (4) In Offshore Benthic locations with mode- rate currents and sandy bottums and some sedi- ment, there are: agnin, stalked species that are raised above che substrate und sometimes, bv we virtue of a pliable stalk. move with the current so that the branchial aperture is presented to the oncoming flow (A plidiuim colellnides, Pely- carpe clavate, Pyura sceresbiensis, P. spinifera. P, australis), (5) In Offshore Benthic Iecations with sluggish 1o slow currents, dhere are: (a) Large species lying on of partly em- hedded in, of fixed to, rocky hottoms or to solid objects in sandy, often mobile. boi toms, These individuals and colonies are oriented. to take maximum advantage of the prevailing current flow by differential growth nf the colony or of the test, especially in the region of the siphons (Syeozan rerebyi- Jarmis, Aseidia spp.. Phallusia lepreysins- eala, Ctenieella antipoda, Herriman miomus), Sycozea cerehrifarmis, which is wbundant on the bottom, especially in upper St. Vincent Gulf, has its “fans oriented to receive maximum current" (S. Shepherd, pers. comin). Its stalk is thick, short. and not pliable, and the species adapts to the direction of prevailing current flow by growth of the colony, In large sessile anil partly embedded species, the orientation of the siphons in relation to the current is effected by their Uifferential growth e.g, Phallusia depressivseulal, R. dispar Kott, 1957a) from Japan and from the central east coast of Austealia (soc Tokioka 1953a; Kott 1957a, 1963). and R. vigillinaides Brewin, 1958u. from Stewart T., have only the 5 primary rows of stigmata und noo parastignyatic yessely, Re asvenerrica Millar has increased numbers of rows of stig- Mata and apparently no parastigmatic vessels. The type species of the genus Exherdmania, E. claviformis (Ritter) (see Van Name 1945), together with &. selidy Millar, 1953 fram the Africeun Gold Coast, E. virrest Millar, 1961 fram Brazil, and E. afgitarad Millar, 1963 from northwestern Australia are easily distinguished by a Jong aesophagus, a large number of rows of stigmata and the absence of parastigmatic vessels and, where ther larvae are known, by the modified adhesive organs as described for this genus and for Placenicla spp. (Rou 1969), Kitherdmunia australis Kott, 1957a, however. fram South Australia, Wietoria and New Sanath Wales, has a short ocsaphagus, 12 10 19 rows of stigmata, purastigmulic vessels, aimst is papilla in the middle of the transverse vessels on cuch side of the body. It as distinguished fram A&. herdmania by the single zooid in each lohe of the colony, the ahsxence of stomach folds. the number of rows of stigmata and the testis fel- licles which wie bunched in the posterior abdo- men. Larvae are known for R. prolifervs and R. herdmania, and are typically palyclinid with ampullary vesicles. Tn the present species and in K. australiy the papillae on the transverse vessels ate remini- seent of Tylobranchion and felated genera, and probably represent a primitive character. Pseudodistoma cercum Michaelsen, 1924: 364. Kou, 1963: 77 and synonymy. Monniot, 1969; 437 New Record: Nora Creina Bay. Previous Records: NSW, (near Eden) —Kote 1963. New Zealand (Stewart I. (Paterson Inlet), Foveaux Strait, Otsgu coast, Lillle Papanui, Great Barrier 1)—Michacisen 1924: Brewin ASCIDIANS OF SOUTH AUSTRALIA i3 1950e, MY58a. Atluntic Ocean ( Dakar}— Munniot 1969. The species t& known inter- tidully and down to 87 om, FIGS, 1k, 19 DPesiprian: Soft. gelulinous, semitesnsparenr, rounded or evlindrical hens of slightly preater diumeter than the more leathery stalk of up to 5 cm leneth. In some specimens the stalk is expanded into yu thick mut from which humerous heads arise. The zooids are numerous and cpen all around the head by separate G-lobed branchidl and atrial openings, The contracted thoray and abdomen together measure only 2 om. Fine jongitudinal muscle bunds on the thorax mumber 20 to 30 and these extend along both sides of the abdomen. There are 15 10 20 pows.of stigmata in each nf the 3 rows, The 4 stomach folds are obscure and may be artefacts resulting from the col- lapse of the stomach. A duodenal awelling and wu rounded posterior stomach are also present. There is a long ovary, with numerous eges more than halfway down the abdomen, but no temis follicles were present in the colonies from these stations, There is a single developing embryo in a brood pouch from the postero- dorsal corner of the thorax. Renurks; The generul form of the colonies. arrangement of body musculsture, the bran- chial sac, gut and the situation of the ovary some distance down the nosterior abdomen, all agrze with the previously described specimens. All other species of the genus have a similar situation for the ovary some distance along the pesterior abdomen: P. efricatigm Millar. 1954, 1%62, P. fragiliv Tokioka, L9SR; P. evrnetsense Péres, 1952: P. antinboia Tokioka, 1949; S. wpaca Brewin, 1950¢, P. Arieni Péres, 1949. The stalked colonies of P. africanum are also reminiscent of the present species in the pre- sence of a single developing embryo in a thoracic breed pouch and ace distinguished only by a smuller number of longitudinal thoracic muscles, As there has been consider- able variation demonstrated in this character, the distinction is suther doubtful, and the spe- cies Of its telatives appear to have a wide cir- cumpalar distribution in the southern temperate region as Monniot (1969) has ulready inei- cated, Subfamily PocycuNtwac Aplidium pliciferum (Redikorzev) Kort, 1963; 106 Amaroticii pliciferim Redikorzéy, 1927: 39D, Tokioky, [953a; 183; 1962: 2: 1947; 32, Aplidine phortax, Millar, 1966: 359. New Records: Troubridge Shoal, Hallett Cove. Previous Records: W. Aust. (Point Peron, Rottnest 1.)—Kout 1963. Vio. (Port Phillip Bay)—Millar 1966, Japan (coastal water of Honshu, Shikoku and Kyushi and the Inland Sea)---Redikorzev 1927; Tokioka 1953a, Hawaiian Is. (Auau Chunnel)- - Tokioka 1967. FIG. 20 Description: Rounded, soft, sessile colonies. 2 cm in diameter. Iq life the colonies are bright yellow. The surface of the colony has deep Jurrows marking it off into extensive rounded areas with up te 2 common cloacal openings from which double row systems muiate Test transparent. zooids orange in the living speci- men. Thorax and abdomen wre of cyual length and together meusure 2.5 mm. The posterior wbdomen is long, up to 8 mm. There sre 4 well-defined branchiai lobes. a strong circular branchial sphincter and & fine longitudinal muscle bunds which extend down cuch side of the thorax, The upper border of the atrial opening is extended inta a small pointed lip sometimes tridentate There are 8-10 rows of about 19 stigmata, The oesophugus is long and the stomach, about half way dawn the abde- men, has 19 to 25 well defined folds. There is w duodenal swelling and a small posterior somuch, Two developing embryos are preseni in i brood pouch formed by the expansion of the distel end of the oviduct ar the postero- dorsal corner of the thorax. The ovary is pre- sent about halfway down the posterior abdo- men anda single series of pyrifarm testis lobes attached to a single duct are present behind the ovary. Larvac have the usual three median suckers with three ampullae between the suckers and many small ampullary vesicles in two rows from euch Jateral line as described previously for specimens from Western Aats- Iralia (see Kott 1963), Remarks: The species is closely related to Apl- iin phertax (Michaelsen) from New Zeo- land, which has a similar number of fine longi- tudinal muscle bands, and stomach folds, and alsa has. a brood pouch, Consequently, there has been some confusion hetween these species. Unfortunately, Michaelsen (1924) did not des- crthe larvac from his species. Aplidiam plici- fern (see Kott 1963) from Western Australia has smaller zooids (therax and abdomen to- gether about | mm long. posterior abdomen 2mm) and are densely distributed in the test, largely obscuring the systems. In Apliditen Pharvay (see Kott 1963) trom eastern Australia PATRICIA KOTT Figs. 13-17. Figs. 18, 19, Fig, 20. Figs. 21, 22. Fig. 23- Ritlerella herdmania, (Port Noarlunga). Fig. 13—Young zooid, contracted thorax, Fig. 14.—Extended thorax of young zooid, Fig. 15.—Zooid with contracted thorax showing parastigmatic vessels, Fig. 16.—Thorax of more mature zooid showing parastigmatic vessels successively subdividing rows of stigmata. Fig. {7:—Portion of colony. Pseudodistoma ceream (Nora Creina). Fig, 18,—Outline of colony. Fig. 19.—Zooid with brood pounch, Aplidium pliciferum, (Hallett Cove, 8 m). Zooid. A plidiumn coleltnlih, (Tapley Shoal, off Trowbridge Light, 17 m). Fig. 21.—Colony. Fig. 22.—Zooid. Synoicium papilliferum. (West 1., sheltered coast, 3 m). Zooid (showing muscles on thorax only). ASCIDIANS OF SOUTII AWSTRALTA is and the Pacific. the larger zooids (thorax and abdomen together 3.5 mm long, and posterior ahdomen 1.5 mm long) are arranged in cit- cular systems. sometumes extending inta more elongate and double row systems. radiating from the common cloacal openings. In all Kott's (1963) specimens the test is gelatinous and semi-transparent with red-purple spherical piement eclls, and the larvae provide the main distinguishing character between the two spe- cies. A, phortaxy has larvae with a limited number of ampullary vesicles: and a complete absence of median ampullae, while the larvae of A. pliciferum retain median papillae and have many small ampullury vesicles from the lateral lines either side of the three median suckers. Millar (1966) described specimens from Port Phillip Bay as 4. phortax. He points out that A, phertax (see in Kott 1963). is not apparently the sam species as his colonies although he cun only distinguish them by the different larval form. He apparently over- looked the similarity in the size and form of the larvae of bis specimen andl of A. pficiferwmn (Redikorvev); Tokioka 1953a: Kott 1963; and based his identification ot the ratia of length to depth of the hirvae of Michaelsen’s species and his own specimens from Port Phillip Bay. However, Kott (1963) has already indicated thit larvae of A. pkortax (Brewin 19463 trom New Zeulund, do have the same rounded form as the larvae of specimens of A. phartax (Kott 1963) from eystern Australia. Tt is apparent. therefore, that specimens from Port Phillip Bav were erroneously identified by Millar. The adul! zooids can definitely be distin- gttishe? hy the longer posterior abdomen, the smaller size, and the greater crowding of zooids of A. plicijerunt The specimen from Hallett Cove was taken with a specimen of Divtaplia viridis in which the zooids wre the same orange colour, The specimen from Troubridge Show! was taken from 4 spiny crab. Aplidium rubricollum Kott, 1963: 103. New Record: Upper St. Vincent Gulf. Pre- vlous Records: W. Aust. (Rottnest T.). S, Aust; (Reevesby 1}. Vic, {Balnarring Beach)—Kott 1963, Description: The single colonv is flattened, thout OF em thick aiid 3.5 cm in maximum diameter, The honiers of the colony are rounded. Sand is present basally and some is enclosed in the common test but the surface is smoath and without sand. The common eloucul apertures with Frilled und protuberant lips ure present on the surface of the colony about 0.3 cm from one another. Spherical pig- ment. cells are present in the test and zovids show as clear points between the pigmented test. In this preserved specimen the pigment cells are pale pink. Zooids are small. up to: 2 mm tong. There are 10 longitudinal thoracic muscles. A short pointed atrial lon- guet arises from the dorsal surface just anterior to the atrial opening which is cenerally on a short protuberamt siphon surrounded by u cir cular sphincter muscle, There are 1) rows af 6-8 stigmata, and 4 stomuch folds. Remurks: The species is distinguished by the form of the atrial aperture and lip. by the nar- Tow branchial suc with relatively few stigmata in each row and by the body musculature and stomach folds, [n the present specimen the test is not so thickly invested with sand as pre- viously described for this species Aplidiam colelluides (Herdman), Millar, 1962; 125, Amarovciam colvlluides Herdoian. 1886: 223, New Recenid: OF Troubtidge T. Previeny Records: South Afries (Cape af Gond Hopel —Herdman 1886; Millur 1942. FIGS. 21. 22 Veseripiion: Rounded velatinous heads on a long hard stalk, The head is up to 4 em in ength and 2.cm in diameter, The stalk, up to 20 em in Jength, is hardened by dense sand inclusion in the surface test which fades out in the test of the head region. ‘lhe stalk i4 branched basally inta short toot-like processes. 7ooids are minute, opening around the surface of the head. Long thread-like posterior abdomina criss-cross in the centre of the head and sometimes extend down jnto the stalk. Some common clowew! apertures are evident around the head and some longitudinal cloacal canals were identified, although the form of Ihe systems is obscure and difficult ta distin- fuish. The thorax and abdomen are of equal length and together measure only about 145 mm, The long, thresd-like post-abdomen is at least four times the combined length of the thorax and abdomen ‘Where are abour 6 deli- cate longitudinal muscles on the thorax. The My PATRICIA KOTT branvhial lubes are cistinet and rounded. The utrial aperture is sometimes produced un a fairly long cylindrical siphon but in another colony is sessile, the upper border of the atrial aperture produced into a pointed languct, There are 18 rows of phour 10 short oval stigmuti. The oesophagus is long, the stomach is present halfway down the abdomen und bas 13 very distinct folds. The gonads are not developed in these speciniens and it is nut known ta what extent they fill the long posterior abdomen in muture zooids, Remarks: This is the only species of A plidium known with a Jong stolk. The size and form of the colony, the size-of zootds and their arrange- ments. in the present colony are identical with the South Africun specimens previously des- cribed, The delicate longitudinal thoracic muscles and the stomach folds are similar The present specimens differ from those des- cribed from South Africa only in the larger number of rows of stigmata. This does not represent a sufficient difference on which to establish a new species and in view of the great similarity in most characters the specimens probably represent one species with a wide cir- cumpolar distribution in the southern colu- temperate region. Synviciuin = papilliferum = (Michaelsen), 1963; 87, Millar, 19A6: 360. Macrollinumn pepilliferion Michuelsen, 1930: 520, Katt, New Records: Port Noarlunga reel, West 1. (neur Penguin Rock), Previony Records: W, Aust. (Bunbury to Nornalup)—Michael- sen 1930; Kotr 1963. Vie. (Nepean Penin- sula)—Millar 1966, The species is known intertidally and to LS m, FIG, 23 Deyeription: Un life the colony is dark red or bright brick red. Flat-topped to rounded colo- nies, narrowing basally Wa common stalk or encrusting. Zooids Jie parallel i the test and open on the upper surfiuce. ‘Vhe colony is firm, gelatinous. There are circular systems arcune protuberant common cloacal apertures. The branchial aperture has 6 small pointed lobes and there is a small circular sphincter muscle at the base of the branchial siphon. The atrial aperture ix opposite the first to second row of stigmata. It is surrounded by a well developed circular sphincter muscle. and is extended into w short cylindrical siphon. The anterior boner ol the atrial aperture is produced into a long mnscular lip, broken into 3-4 minute pointed lobes terminally, There ure !0 very fine longi- tudinal muscle bands on the thorax which is very delicate and transparent. There are {0-12 rows of about 10) stigmata in each row, Che hody wall below the atrial aperture is pro- duced into the small rounded pupillae charac- teristic of Synoiciun spp. The wall of the slomach is riised into faint mulberry-like swel- lings. The posterior abdomen is shart and there is no constriction between it and the abdomen, Rentarks: Both colony and zoaids conform with previous descriptions in all characters except the reduced mumber of rows of stig- mata. The species has been recorded fram south-western Australis along the south coast of Australta ta the Nepean Peninsula in Vice toria (Millar 1966). Tamily DIDEMNIDAE ? ‘Trididemunm spiculatums Kon, 1962: 281. New Record: West 1. (near Penguin Rock), Previous Records: W. Aust. (Rottnest 1., Point Peron), S. Aust. (Quter Harbour) Tas. (Wreck Bay). Qld. (Heron b.I— Kare 1962, Description; Living colonies pile pink, enctitst- ing. Small, almost spherical spicules with up ty 12 points in optical transverse section, evenly distributed throughout the test, and oeeasonally large spicules with fewer rays. There are small thoracic common cloacal cavi- Ges. Zooids are smull with three rows of stig- mata, The atrial aperture is wide. cxposing a lurge part of the branchial suc. Gonads are nut mature in the present specimens, Remarks: Colonies generally conform with specimens previously assigned to this specics, ulthough the proportion of smaller isner-lke spicules lo larger stellate spicules with about & rays in optical section. is greater in the present specimen, Colonies with mature zdutds are desirable for positive identification. Leptoclinides rufus (Sluiter). Tokioka, 1952; 92. Kott. 1962: 286 and synonymy. Eldredge, 1967: 221. Rossenpranate refus Sluiter, $9092 72; 1913: New Records: Off Pore Gawier, Hallett Cove, Port Noarlunga, Rapid Head, West T., Wright F, Previous Records; §, Aust. (Port Noarlunga). View (Shoreham). Tas, (Maurin 1}. NSW, (Port Jackson)—Kott 1962. Old. (Heron T.)—Hustings 1931. New Zea- land {Great Barrier t.. L. ¢luiteri}—-Brewin 1950b; (?Stewart I, L. nNovdezelatidiae}— ASCIDIANS OF SOLTH ALSTRALIA 7 Brewin 1958a, (?Chutham Rise, Lauran ticis)—Brewin 1956: (North t.)—Michgel- sen 1924) Brewin 1958b; Millar 1960, Indo- Pacific fArafura Sea, Indonesia, Hawati}— Tokioku 1952; Stuiter 1909; Eldredge 1947, ‘The spectes is Known intertidally and to 36 m (Sluiter 1909). Deseripiion; Encrusting colonies, Living Specimens: while matrit with grey oar dark animals, or orange ta light fawn (Port Noar- lunga); or dark reddish brown folf Hallett Cove), mottled white to uniform light grey colour (Wright 1.).. In preservative all calonies ire While to orange-white or streaked and blotched with grey. ‘Vhe colonies are investing, sometimes extensive. Cloacal cavities radiate from randomly distributed apertures, Zoids ire sametimes present in the roof of the com- mon cloacal cavity. Spicules ure present in the surface tese but basally the testis jelly-like and transparent. There are 9 tongituclinal muscles ev the thorax. The posteriorly directed atrial siphon has a wide circular sphincter muscle. There ure 4 rows of 10 to 12 stigmata, There is a superficial layer of bladder cells and small oval to spherical pigment cells are present amongst the surface Jayer of spicules. A literal organ is present opposite the middle of the fourth row of stigmata, Cloncal apertures are present, espectully uround the burders of the calony. Canals. at thoracic level radiate from the cloacal apertures between clumps of zooids although sometimes they extend deeper to abdominal level. The cloucal canals around the border of the colony ire often completely sub-abdominal. The ypicules are of the usual stellate form, 0,0/-0.04 mm in diameter. Larvae are present in some colonies from Hullett Cove. They are of usual form, fairly deep with 4 paired ampullue. In one colony from Hallett Cove (dark reddish brown in life) no common cloacal cavities were present and zooids Were not mature, nor were zooid open- ings to the exterior detected, The arrangement of spicules is characteristic of this species and jl a8 probable that the colony is one in which sexual reproduction ts completed and new vege- tative buils are developing. Remarks: The species ix distinguished by the complete absence of spicules from the basal layer of the test, sometimes giving the colony a very fleshy appearance. The characteristic common cloacal system and the distinct mus- culature on the thorax, together with the pos- tetiorly directed atrial siphon and the spherical to oval pigment cells are distinctive, Leptoclinides kingi Michwelscn. Palveyneraron aibiint. Van Nate, (STB: b55, Hartmeyer, 1819: 136, Leplectinidis dublax f. kinet (930: 507, Kou. 1942) 2489 New Record; Upper St. Vincent Gulf. Pre- vious Records: W. Aust. (Fremantle, Albany) —Michaelsen 1930. Old. (Sarlna)~ -Kott Michaelser, 1962, Philippines (dolu Light} —Van Name 1918. The species is known intertidully anu to 18 m. FIGS, 24, 25 Deseripnion: Vhe colony is massive with the surface tuised into mounds und single cloacal apertures ut the apex of each mound. Each mound ts lormed by thickened basal test often with embedded pirasites, Zooids are present in the surface test above the very »xtensive posterior abdominu!l spaces round the centre of each lobe or mound, The zooids ure large with 4 rows of about 12 stigmata, There are $ very fine longitudinal muscles on the thuran, The ‘spicules are very small, (1.01 to 0,02, mm. and are ranged in a shallow laver at the level of the branchial siphons. They are only very sparse elsewherd in the test. There is 4 surface layer of bladder cells. Remarky: The elevation of the surface of this coluny into mounds or lobes with terminal common cloacal apertures characterises this species, which was previously regarded as a form of Leptoclinides dubius (Sluiter). Lepte- clinédes dubius is distinguished from ihe present specics hy its larger spicules and by the utrangement of common cloacal system with openings around the margins of each opley, us in B, rufus. In £. kingi large cloacal sys- tems with terminal openings develop fram the centre of the colany, As both forms have been recorded more of less ever the same geographic unge it as unlikely that they represent geo- graphic subspecies of the ons species, and in view of the different development of the com- mon coacal systems it is probable that they rep- resent different species. The ony gut loop which is bent anteriorly to Form a double loop is a character shared with Leproclinides didits, Posteriorly directed atrial siphons of the zooids open inte the common cloacal cavities and canals, The openings sometimes appear 5 lobed due to the arrangement of spicules around the aperture, The genus Askonides Kott, 1962, therefore cannot be distinguished from Leptociinides and A. imperfectur and A, ceelenterwius are distinguished from other spe- cies of Leptoclinides only by the extent to 1k PATRICIA KOTT which zooids open directly into Ihe common cloacal chamber rather than into cloacal canals. Their relations are set out in the following key: |, Single systems develop around central com- mon cloacal cavities with terminal epen- ings Passeeccietge ie Sade 1, Numerous systems develop around peri- phery of colony 3 2 Spicules accumulated in surface layer o test; spicules 0.0)-0.02: larvae with 4 paired ampullae; most zoaids open into cloacal carvals L. kingi 2. Spicules throughout; spicules 0.04-0.08, lurve with reduced ampullae; most zooids open direct into common cloacal cavily L, covlenreranus and L. imperfectius 3, Spicitles 0.01-0.02; double gut loop... L. dubius 3. Spicules 0,02-0.04; simple gut loop L. rufus Leptoclinides reticulatus (Siuiter}. Kott, 1962: 285 and synonymy. Didemniun refientaive Siuiter, 09> ha. New Record: Tipata Reel. Previous Reiords; Qld. (Noosa to Mackay, Heron 1, Low Is.)—Hastings 1931; Kolt 1962, New Zealand (North 1.)—Michaelsen 1924, Japan—Oka 1927; Tokiokn 1953a, 1953b. Indonesit —Stuiler 1909, 7Philippines-Van Name 1918. Indian Ocean (Ceylon) —Herd- man 1906. FIG. 26 Deseriplion; Young colonies were taken invest- ing Micrecosmus squarmiger and Pytire irregu- lariy. Frequent common cloacal openings are scattered over Vhe surface. There is super- ficial layer of bladder cells with orange and black pigment in stellate cells forming streaks on the surface. Spicules are present benenth this superficial layer and are reduced in den- sity toward the base af he colony. The spi- pules ire sicilate with about 7 conical rays in optical transverse section and from 0,03 to O05 mm ty optical section, The vrimary clovcal canals are deep, but in these specimens. do not extend posterior to the zooids. The znoids are small with the usuul 4 rows of stigmata and a large posteriorly divected atrial siphon. There are 4 testis lobes and 44 coils of the vas deferens. Remarks! This 1s the most southerly record for this: conspicuous. and widespread species, dis- tinguished by its unique stellate pigment cells whicl) form the charucteristic “ogertike” mark- ings on the surface, Didemnaum tambitum (Stuiter), Kou, 1962: 317 and synonymy, 1971: 19%. Didemnoides lanbitem Sluiter, 1900, 18, New Record: Aldinga “drop off. Previous Rerords: N.S.W.---Kait 1954, 1962, New Zealand (Chatham I.. North L.. South [,)— Sluiter (1900; Michaelsen 1924; Kott 1971: and unpublished records from Otage (call R. Ceump) and Stewart I, (coll. . Barham). Description: Two clavate lobes arise from a common base, Maximum diameter 1.5 em and maximum height 3.0 em. There are traces of orange pigmen{ in the surface Lest, but no superficial layer of bladder eclls, There is a layer of spicules in the surfuece lest which ceases abruptly at ocsophugeal level. Thin Jayers of spicules line the common eloacal canal. Spicules. are absent at the abdominal level of the zooids, and in dhe centtul test core. They are 0.01 to 0.05 mm and stellate, Ter- minal cloycal aperture opens into the charac- (éristic common cloacal cavity surrounding the central core of test. Zooids are small and crowded in the surface layer of test, “The atrial aperture is wide and open. There are 8! coils of the vas deferens around a single testis lobe. Didemnum patulum (Herdman), Leptaclinym petuiim Herdman, 1499: 92. New Record: Aldinga, Previous Recerels: Vie, (Port Phillip Bay)}—unpublished regord N.S.W. (Port Jackson) Herdman 1894. FIG. 27 Deveriplions Tough, investing colonies. In preservative the specimens ure white with grey streaks and blotches formed by palehes of stel- late pigment cells in the surface test, especially in the région of the commun cloacal canals The surface of the colony is marked off into alightly raised rounded arews where solid pil- lars of test traverse the coramuan cloacal cavity, Zooids are embedded! in the periphery of these pillars of test and open to the surface around the raised ares, “Whe cloacal cavity is thoracic. The surface layer of test is especially thick and the zooids huve especially long and musculus branchial siphons which estend through this surface layer of test. Spicules often form a plug inside the branchial siphon—possibly caused when the superficial layer of test is pulled down into the aperture as it is Teiracted into the surface of the test. The branchial siphon is almost the same length as the rest of the thorax. The atrial opening is wide, expos- ing a ptt of the dorsal surface of the branchial sue, The anterior border of the atrial opening ASCIDIANS OF SOUTH AUSTRALIA iW \§ produced into a natraw pointed languer, sometinves bidentate at the tip. There ane con- spicuous circular muscles in the branchial siphon, in addition to the usual longitudinal muscles that extend down the length of the thorax and into the test to form a short retrace tor muscle. The abdomen, of the usual form lor this genus, is especially small. Oesophageal buds are present but the gonads are nat mature, Remarks: The grey veins in the surface identify This specimen with Herdmun’s species, The long branchial siphon and atrial lip wre also distinctive. The specres is especially common in Port PRillip Bay, but is not common in St. Vincent Gulf The species also strongly resembles 2, tabulate Sluiter from the East Indies and Aru 1. (sec Sluiter 1913; Kott 1962). Didemoum moseleyi (Herdman). Vat Name, 1918: ISL. Tokioka, 195Sa: 212: }955h; 44; 1959: 226; 1961: 106. Katt, 1957b; 136: 1962: 328 and synenymy, Eldredge. 1967; 212, Leproctinuen moseleve Merdman, 1&6: 272. fupreclinion incanim Hendman, (898° 90, Herdman & Riddell, (813; 888. New Records: Cioose |, Carickulinga Head. West 1. Previous Records: W, Aust, (Rote nest L., Point Peron, Trigg 1). S. Aust. (Reevesby LL). Vic. (Balnarring Beach )— Katt 1962. Tas. (Spring Bay, Maria I.}. N.S,W, (Port Jackson, Port Stephens, Coffs Hatbour)—Herdman 1899: Kott 962. Indiun Ocean {Southern Arabin}—Kott 19S7h. Indonesia (Arafura Sea)—Sluiter 1914, 1913; Tokioka 19553. Pacific Ocean (Patau is. New Caledonia, Philippines. Hawallan ts, Marshall Is.) —Herdman 1886: Van Name 1918; ‘Tokioka (9556, 1961; Eldredye 1967. FIG, 28 Description: Investing sheets, There is a very thin Jayee of surface test which is often tatged imo spicule-filled conical papillac between the branchial apertures, The cloacal cavity is thoracic and the thoraces of szooids are enclosed in an independant test sheath, The atrial opening is wide, in all cases exposing the branchial sac tothe cloacal canal, Spicules are 0,02 to 0.04 mm in diameter with no more than 10 pointed rays in optical transverse section and are densely distributed throughout. Zooids are colourless. They are minute. the branchial sac espycially small with four rows of only & stigmata, The yas deferens coils 64 times around a single undivided testis follicle, In the specimens from West 1. and Carickalingy Head there is a smull laters] organ opposite the last two TOoWs of stigmata. Remurks: Eldredge (1967), discussing the difficulties in distinguishing between the present species and D. candidun, hus suggested that in 2, candidum the surface test is always smooth. the atrial aperture is w small slit and Jateral organs. are always absent. He has not been able to confirm the presence of larger numbers of yas deferens coils for D. cenedidum (Kott 1962) nor is the condition of any of these characters constant in specimens previously ascribed to the species, Only the regularly stellate spicules and dark pigmented zooids of the present specimens uppear to distinguish them from D. movelevi which has a variety of diferent types of spicules. Didemnum candidum Savigny, 18i6; |94, Michaelsen, 1924; 358 und synonymy. Van Name, 1945; 83, Hastings, 1931: 94, Brewin, 19467 98; 1950u: 55: 1950b; 345; 1951: 104: 1952b: 188; 1956: | 22; IYST= S77, 196 14. Tokioka, 1954y; 246; 195Sa: 45. Koti, 1954: 162; 1962: 327. Eldredge, 1967: 23. The above synonymy refers only to Indo- Pacific records. For full jist of synonyms see Fldredge 1967; 213. New Records; West T., Wright lL, Previnviy Records: South-western Australia, Tasmania, north-eastern Australia. the Envlish Channel, Trish Sea, West Africa, South Africa and East Africa, Red Sea, Mediterranean Sen, New Zealand. west and mid-Pucific Ovenn (Marshall is. and Hawaiian ts ), the Carib- bean and West Indics and the est coast of the USA. Records are lacking from the north Pacific and west coast of the Amenecan continent, but elsewhere the species pevurs widely in temperate and tropical regions. FIGS, 29, 40 Deseription: Colonies arc Hat and investing, small and rounded or more extensive sheets. The test has dense spicules throughaut, In preservative the zovids are brown and show through the white spicules. The common cloacal cavity is thoracic but extensive and limited only. by thin layers of surface and slightly thicker basal test in which the abdo- mina of the zooids are embedded, Thoraces cross the common cloacal cavity in an indepen- dent sheath of test, Spicules are dense through- out. They are 0.02 to 0.03 mm in diameter 20 PATRICIA KOTT and demonstrate the same range in form pre-almost cylindrical marginal rims stiffened by viously described for this species with up to 1Sthe dense spicules enclosed in the test. Zooids or more rays in optical transverse section. Con-are very small. There are 4 rows of about 8 spicuous common cloacal apertures present onstigmata. No gonads were distinguished in the the surface are surrounded by protuberant,present colonies. ASCIDIANS OF SOUTH AUSTRALIA 2] Remarks; The present colonial systems are typical of the species although no gonads Appeared to be mature, It was not possible to confirm Eldredge's observations concerning the slitlike atrial opening as in the extended zouus of the present colonies, these were wide open, exposing a great part of the dorsal aspect of the branchial sac. The vartety of spicules. therefore, rémain the principal distinguishing character for this species. Curliste (1954) has characterised specimens of 2D. candida Savignay Fram the North Sea, the English Chan- nel, north-west Afvicu, the Medilerrancan and the Red Sea (type bocality) by the absence of the third adhesive pupilla in the lurvac, and Lafargue (1968) confirms the condition for specimens trom. the French coast. The speci- mens agree in all other respects with those des- cribed from New Zealand, Australia, Malaysia, Japan and the Atlantic coast of America. Carlisle concludes, therefore, that: “D. cendj- dum is a tropical and lemperate species extend- ing from the West Indies to the East Indies, New Zealand and Japan”. However, later workers have not observed the universal absence of a third adhesive papilla in luryae from these localities, while there are the usual three larval papillae in Australian, New Zealand and Japanese specimens. It is possible, therefore. that two separate specics are Involved. Polvsyncraton orbiculum Kott, 1962; 300. New Record; Rapid Head. Previous Recordy: W, Aust, (Rottnest 1). 8. Aust. (Por Noarlungy)—Kott 1962. Peseripion: The preserved colony is light pinkish brown, owing to the darkly pigrnented vooids seen through the single layer of spicules presemt in the thin. surface test. The dark coloured zooids are ulyo seen through the branchial Openings clearly marked on the sur- face test. There aro the usual vesicular cells arranged in « complete circle around the branchial openings. and interrupting the ather- wise even disuibution of the spicules in the surface test. There is an extensive thoracic cloacal cavity, crossed hy the thoraces of the zoulds, exch with a discrete ventral sheath of test. There is a iitteral organ about halfway down the thoracic test sheath, The vooids are small, with 4 rows of stigmata, There is a long retractor muscle, These specimens conform with those described previously (Kott 1962} in all respects; however, the gonuls are not muture mn the present zovids. Remarks, The condition of the cloucal cavity, the dark pigmented zooids, the rather Jarge stel- late spicules and the unique, large transparent vesicles in regular circles in the surface, together, charucterise the species. Echinoclimom $yerrilli Van Name, 1902: 372, Kot. 1962: 312 and synonyms, Diplosoma VLissoclinum) verrillt, Eldredge. 1967: 242, New Recards: Hallett Cove. The species has been observed investing the underside of rocks aj 4 depth of S20 im at many Joca- trons in St. Vineent Gulf where conditions are quiet. The colonics are so fragile, how- ever, that they usually break up when removed (S. Shepherd, pers. comm.) _Pre- vious Recares: Vas, (West Coast)—Kot 1954. America (West Indies, Florida)— Van Name 1902, 1945; Hartmeyer, 19f9— 11: Plough & Jones 1937. Africa (Accra) —Millur 1953. Japan {Sagami Bay)—- Tokioka 1958. FIGS. 31-35 Deveription: Living colony soft. white, jelly- like. In preservative the present colony is deli- cate and soft, It appears to be investing but is, unfortunately, damaged and its exact form could not he determined. Spicules are mostly G-rayed, but there are alse spicules with 4 anu with 3 rays. They form a dense spiny, tough capsule around the abdomina of the zaojds but are sparse in the remainder of the cobony- Zooids ate arranged more or less in the dauble rows previously described (Van Name 1945) although common cloacal openings were not deiected. The cloacal canals spread usit beneath the zonids whith are retained m the Figs. 24, 24, beptoctinides kingi, (Upper St. Vincent Gulf, 10-12 m), Fig. 214.—Spiciles. Fig. 25. Thorax, diagrammatic, showing mus Gut loap, Fig. 26. Leptoclinides: reticulatus, (West 1,, under boulder), Snicules, Fig. 27, Didemnum parnlam, (Aldinga “drop-off, 3-8 m). cutature. Fig, 28 Didermnum moseleyi, (Cavickalings Head, 5-6 m), Spicules. Figs, 29, 30, Bidemunm vandidum, (Wright T. Fig 30—Spicules, Figs, 31-35. rough coast, 10 m), Fig, 28—Diayram of colony Echinoellnum verilli. (Hallet Cove, 8 m), Fig. 31. Spicules. Figs. 32, 33, 34.—Larvae of incyeagiog maturily, Fig. 35,—Matare anterior ampullae of larvac. ps PATRICIA KOTT surface test, Zooids ure small with large lateral organs on each side of the thoras. Larvae fre large with a short tail which. when extended, is only hall the total length of the lurva. There is a large ocellus and an -ote- fith, At jeast one precocious bud is present although the exact number is obscured by the laver of spherical to oval granulate bodies that extend around the posterior half of the body of the lurva. Antenorly there ure the usual three adhesive papillae in the median tine and [4 ampullae from the Imeral lines on either side of the suckers. Ipitially these lateral ampullae are very small and sessile. Subsequently they jncveuse in size und become “tear-drop” in shape supported by very narrow stalks from we lateral line. Remarks Wis unfortunate that the present eolony is so damaged thal its shape cannot be discerned. Although previously deseribed specimens have been clavate (Kott 1954) Van Name [945] the present damaged colony ts investing and living colanjes have been observed investing the under-surface of rocks, It is possible therefore that two distinet species may be involved, characterised by a difference uy the consistency of the text and in the shape of the colony. ‘The soft nature of the colony and its ten- dency to break up hus probably been the cause of the lack of records of this form, which is reported as common in St. Vincent Gulf- Eldredge (1967) has suggested that the genus. is synonymous with Diplexoma (Lisin- clini), due to the similarity of the cloucal systems avid the fact that tetrahedral spicules ure not unique in the family Didemnidac. Eldredve's contention cannet he maintained, Vhe common cloucal cavity in the two genera is extensive and extends posterior to the zooitts which remain connected to the hasul test by strinds of test. However, the cloacal syster in Echinocliniwn differs from that in Diployoma (Livsoelinimy in the absence ol the secondary cloacal spaces around the thoraces of the zaoids which remain connected to and in the surface test in continuous rows. In Diplosema” { Lice. clinuar) the secondary cloacal spaces sepacate cither the thoraces, or the whole zooids, from one another. Further, the spicules in Eehine- elinum are very much larger (0,05—0.1 mm) than those generally found in other genera of the famlly and, in addition co their unusuul form and size, their distribution in the coluny differs. entirely from other venera of the Didemnidac. The capsules formed around the zooids by ihe spicules are reminiscent of the eapsules formed in Cystodyres spp. and in no other venus of the Didemnidue do the spicules remain in such an intimate relationship with the zon. The genus is further distinguished by a Unigue larval form with wv multipleity of narrow-slulked epidermal ampullae and prece- cious. buds. The larvae of P, aspiculatum and D, (Lissoclinnin) spp. show a similiar oirkedt increase in the number of lateral ampulbic, The ampullue in Eehinoctnum wre unique, how- ever, in their distinet “tear-drop” shape, their natrow stalks and their diserete origin from the lateral jine without subsequent stibdivision. Precocious budding generally occurs in the farvae of Diplosema spp. and in D. Lésxa- clinum) spp. However. it alsa occurs in Didemaunt (B, psendodiplasamu— Rott 1962, and D. ternarenuni—Koie L866) and in Paly- syecraton (P. aspiculatim—Koir 1962) so can- not be considered characteristic of any single genus. The zeunular bodies present in the larval test are indeed similar to those found in O. 1 Livsa- clinum | fragile—Eldredge 1967 and D. | Lisso- clingeny) dstrearinmn—-Kort 1962, They do not like up huematoxylin stains (Eldredge 19671 and thus de not appear to be calcareous spi- cules nor their precursors. as Kott (1962) had sutycsicd. However, despite che relationship with D. ( Livsoclinunty medicated by these enclosed granules. the genus is distinct from ether genera in the Didemnidae und entirely justifies. iis Laxonomic position as a monotypic genus in that family- Didemnunt sp. Renard: West I, (near Penguin Rock) Deseription: Living colony “yellow, crusuase". In preservative the investing colony is a light fawn colour. There are common cloacil aper- tures with large spicule-filled lips scattered over the surfuce of the enlony. Zooids are suspended between the basal and surface layers of tesr by connecting columns of test in which the abdomina are embedded in clumps, although the vooids are separated from one another in their own discrete sheath of Lest, open to the vommon cloacal cavity on the dorsum. Stellate spicules are thick throughout the test. The branchial siphons are Carly tong with distinct circular muscles. There are Jatge oval hitecul organs on either side of the thorax. There are four rows of stigmata. ASCIDIANS OF SOUTH AUSTRALIA Reniarky! The gonads are not developed and a definitive identification of the genus is there- fore not possible. The cemditiun of the colony with a well developed posterior abdominal cloacal canal is reminiscent of cerluin species Of Didemnrium, Suborder PHLEBOBRANCHIA Family CORELLIDAE Subfamily rilencsomaTiNar Khadosoma tireicuns (Savigny). Kotr, 1952: 417 and synonyity.. Tokioka, 1952: 111; W953q: 230, Phallisia vcica Saigny, 1B162 102, Rhadoxoma papillosum. Van Name, 1918: 113 and synonymy. Hartneyer. 1919: 99, New Record: Hallett Cove, — Previons Reeords: NW. Aust. (Cape Jaubert) —Harst- meyer 1919. S$. Aust. (Port Nourluinga)- Old.—Kon 1952. Indonesia—Sluiter 1904; (Aralura Sea)—Tokioka 1952, Indian OQcean (CeylonJ—Herdman 1906, Pacific Ocean (Philippines. Califarnia)—Wan Name 1918, 1945; (Chile}—Traustedt 882, A835: (China)—Stimpson 1855; {Japan)—Oka 1927: Hartmeyer 1906; ‘Tokioka 1953a. Red Sea—Ehrenberg 1828, Mediterranean —tLacaze-Duihiers 1865. The species is also recorded From the Caribbean region (Van Name 1945) Remarks; Nothing further cun be added to the deseriptian of this cosmopolitan but rare species. His never taken in large numbers, nor is it taken very often, The species is, however, not inconspicuous, It is probable that, with its highly developed closing mecha- nism, it may exhibit a high degree of vivipary. In which case it is probable that rehiytively few larvae are incubated, and that the free-swim- ming time of larvae is shart. “he dispersal of lurvae could be. therefore, limited, and the survival of the apparently small populations of the species enhanced by larval settlement close to the parent zooids, The specics has heen taken from a wide variety of depths. Lnfor- tunately, little is known of the current condi- tions at locations from which the species has been taken, but it is possible that it favours less turbulent conditions where there is misimal current flow so that the larvae would be even lest exposed to dispersal, Only a single specimen ts present in this col- lection, iz “a Subfamily COREBLINAE Corella cumyota Traustedt, (882; 271, Kot, 1969; 84 und synonymy; 1971 > 20 New Records; Hallett Cove, King Beach. Previous Records! W. Aust (Trigg 1.j— Kott 1952. Vie, (Balnarring Beach, Franks- ton) —Kott 1952, Millur 1966, Tas, {D'Entrecasteaux Channel), New Zealand (North and South 1s.)—Sluiter 1898: Michielsen 1922; Brewin 1946, 1948, 1950a. 1957. 19640. South Africu—-Stuiter 1898; Michuelsen 1915; Millar 1955, 1962. ‘The species also has i wide circumpolar distribu- tion in the Antarctic (Kott 1969). FIG, 36 Deseription; The living $pecinens were noted transparent anc’ no colour was recorded. There are both separate individuals and indi- viduals ageregated together more or less in a line. Zooids are generally fixed to one another er to the substrate by almost the whole of the fight side, The atest is thick, gelatinous and semi-transparent. On the right side of the body where it is fixed to the substrate the boily wall is especially’ thin and there are no nuscles except those which radiate a shoarr dis- tance from the branchial siphon. On the upper or left side of the body there are mostly trans- verse muscles branching and ramilying and some short and more regular transverse muscles in a single row extending, sround the ventral border. The branchial siphon is terminal and on a short cylindrical siphon. The atrial aper- ture is sexsile and from the posterior third ef the dorsal border, ‘The branchial sac, gut and gonads are of the usual form characteristic of the genus. Remarks: These specimens do not differ in uny Way from other specimens of this ubiquitous species which has been recorded in very large numbers from open sea locations in ecireum- polar waters of the Antarctle and the sub- Anturetic (Kot 1969, 1971). The vorthern extent of the recorded range is at Trigg I, {Kott 1962) on the western coast of Australia, hut the species has not been taken on the eastern count of the Australian mainland: the most easterly record on the Australian cuast is at Frankston in Victoria (Millar 1966). Family ASCIDIIDAE Phallusia depressiuscula (Heller), Kutt, 1472: 8 and synonymy. Ascidia depressiustula Heller, 1878: = Herd- man, 1906; 305. Ascidia julinéa, Vasseur, 1967; 129. am PATRICIA KOLT New Records: Tapley Shoal. off Part Gawler, off Grange, off West Beach, Hallett Cove, off Port Stanvac, Wright I. Previous Records: W, Aust. (N.W. Aust. Shark Bay, Fremantle)—Hartmever 1919; Michjelsen & Hartmeyer 1928; Millar 1963. N.S.W- (Port Jacksonj—Herdman 1899. Qld. (Great Barrier Reef)—Hastings 1931, Kott 1952, 1946. Bass Strait (Enst Moncgeur 1.) —Herdman 882. Pacific Philippines, Palao Is. New Caledonig)—Vun Name L913; Tokioka 7250; Vasseur 1967, Indo- Malaya (Ceylon, Indonesia, Arafura Sea)— Heller 1878: Hetdman 1906; Stuiter 1919: Tokioka 1952, The species. is krown inter- tidally and to 52 m, Description: Living specimens from off Hallete Cove are noted as large, white or transparent, and cammon on sandy bottom. Many living specimens. however. ore bluixh, with black and yellaw markings. The preserved specimens may be whitish, or blackish grey und may have black spots in the surface test, The test is thick and firm, smooth on the surface with rounded ridges and swellings, The individuals reach a large size. ‘The present specimens exhibit the range of variation described by Kott (1966) for the species, Remurks: The relationship of Phallusia julined Sluiter lo the present species remains in doubt. The specimens in the present collection have the atrial aperture from the anterior third of the body while specimens of P julines have been distinguished by the position of the atrial aperture From the posterior third of the body. Ascidiq sydneyensis Stimpson (?part), 1885: 387, Kott, 1972 and synonymy- New Records; Taplcey Shoal, Hallett Cave, Port Noarlunga, Wright 1, Previous Records: W, Aust (Cape Jaubert to: Albany )—Hart mever 1919; Michaelsen & Hartmeyer 1928; Millar 1963, S, Aust. (Victor Harbor, Port Noarlunga). Vic. (Ralnarring Beach, Point Leo, Port Phillip Bay}—Kott 1952; Millar’ 1960; 1963; 1966, Tas. (Spring Bay). N.S.\W, (Port Jackson)—Stimpson 1855; Herdman 1882, 1899, Qld. (Caloundra to Townsville)—Sehmeliz 1879; Koil 1962, 1966. Indonesia (Arafura. Sca)—Sluiter 1886, 1904; Tokioka 1952, Pacific Qcean— Traustedt 1885; (Palao y., New Caledonia) —Tokioky 1950; Vasseur 1967, Japan— Harimeyer 1906, Tokioka I9S3a, 1954b. Indian Ocean (Seychelles) — Michaelsen 1918; (Zanzibari—Traustedt & Weltner 1R94; (East Africu}—Millar 1956. South Africa—Heller 1878; WHurtmeyer 19t1, 19) 3; Shutter 1898: Millar 1955, 1962. The species is also recomed from. the Caribbean region (Van Nume i435). [tis taken inter- tidally and to 31h m, FIGS, 37, 38 Descriprion; Vhe living specimens are. trans- parent and fleshy. The largest specimens in the present collection are 20 cm long und 12 em wide. The test is thin, but firm and tough, and in larger Specimens slightly leathery. There is sometimes. especially on the larger specimens, a very sparse encrustation of weed and worm tubes. Both the branchial and atrial ipertures. ore on short cylindrical siphons and ure usually about half the body length distant fram one another. Specimens may be fixed ta the substrate by the posterior, ventral, or left side of the body. The branchial siphon is turned away Front the atrial siphon lo varying extents. here is a row of short transverse muscle bands around the dorsal and ventral borders of the right side of the body, The gut is always filled with mud, which appears to accumulate during the life of the individual until in lirger specimens the gut is so swollen with mud that the branchial sac is occluded und confined ta a small area to the right and dorsal to the mud-filled gut, This mud hegins to collect, in smaller specimens, in the descend- ing limb of the primary gut loop, beyond the stomach, and it extends from there into the rectum and continues to accumulate in these sections Of the intestine. Remarks: The physiological significance of the mud-filled gut which appears. to be characteris- tic of this species is nol known. It has been noted in specimens from all parts of the Puci- fic. Abbott (pers, comm, 1955) noted that it appears to be associated with the termination of the typholosole at the top.of the gut swelling instead of extending further down the intestine. The stomach appears to be free from the mud accumulation, but distal to the stomach the gut becomes so distended and the whole body inside the test becomes so compressed by it that it is difficult to imagine normal feeding and respiratory functions proceeding. Some of the mud must be lost through the arws anil ASCIDIANS OF SOUTH AUSTRALIA Ek Fig. 36. Curella eumyota. (Hallett Cove, 25 m). Individual removed from test. Figs, 37, 38. Ascidia sydneyensis. Fig, 37.—Individual from Tapley Shoal, 13 m. Fig. 38.—Individual from Wright I., 10 m. Figs, 39, 40. Ascidia gemmata, (Upper St. Vincent Gulf. 10-12 m). Fig. 39. -Individual removed from test. Fig. 40.—Diagrammatic section through branchial papillae, Figs. 41-43. Ascidia thompsoni. Fig. 41.—Dorsal lamina. Fig. 42.—Individual removed from test (Carickalinga Head, 5-6 m), Fig. 43.—Individual in test (aff West Beach, 8 m). Figs. 44, 45, Ascidia aclara. (Off Seacliff, 16 m). Fig. 44.—Whole individual, Fig. 45 —Individual removed from. test. 6 PATRICIA KOTT uifial opening and until ubseryations are made on living specimens. it must be asstimeéd that the property of the distal part of the gut ta dis- tend ilsell in this way is characteristic af the species and results in the acctimulutions of gut contents al i greater rate than they are removed from the body. Ascidia gemmata Sluiter, 1895: 177. Kou, 1966; 296 and synanymy. “Cokioka, 1967- 149, New Recordy: Upper St. Vincent Gulf, off Port Gawler. off Glenelg. Previous Records: W. Aust. (Cape Jaubert to Albany)]—Hart- meyer 1919: Michaelsen & Hartmeyer 1928: Koil 1952, Vic. (Port Phillip Bay}—Koall 1952; Millar 1966. N.S.W, (Port Jackson, Acruwarra)—Herdman 1899; Kott 1952. Qid. (Hervey Bay)—Kort 1966. [ndo- Pacific (Indonesia) —Sluiter 1904; Tokioka 1952; (Palao Is., New Caledonia, Matianas Is,, Caroline Is. Wake ts.) —Tokioka 1950, 9G. TYaT. FIGS. 39. 40 Meveriptions Externally the test is fairly thin and flaccid and is slightly irregular, The bran- chial aperture is terminal on a short cylindrical siphon, ‘The atrial aperture is on a similar but venecully shorter siphon Tron) the antero- dorsal aspect of the body, Both siphons are regularly prooved externally wlong their length, Individuals are attached by almast the whule of the left side. Internally the atrial siphon arises from half way dewn the body and is especially jong. The branchial siphun is alsa long inter- nally, There are circular and longitudinal museles around both the siphons and these extend only a Short distanee pusterior to the siphonk. on the left side of the body where there is nu musculature, On the right side of the holy the lonyitudinal qiuseles from the siphons mingle with the irregular meshwork of muscles which occupy the whole body wall on the right side. There is only a very narrow prebranchial area terminated anteriorly by very numerous branchial tentacles, and covered with minute papillae, The dorsal tubercle is fairly large circular cushion with a U-shaped slit turned to the fight and with the pos- tering horn turned in, The pentubercular ater is shallow and is completely filled by Vhe dorsal tubercle. The dorsal lamina is a broad, single membrane, strongly ribbed on both sides. The ribs of the dorsal lamina extend into pointed languets on the free margin, There iy a long ocural gland almost one-third of the body distant Fran the dorsal tubercle. The branchial sac is simply folded between each fongitudiqul vessel and has 4 to & stigmata in cach mesh. There are large spatulate papillae at the junctions of the longitudinal and transverse vessel and these ure expanded into rounded expansions on cither side of thelr base, The gut forms a Ueep double loop enclosing the gonads jn the pri- mary loop. The pole of the gut loop in the large specimens available in this collection does not catend anierior to the buse of the atrial siphon und is level with the anus, There ts. however, some variation accerding to the size of the specimens and jn smaller specimens (Michaelsen & Harimeyer 1928: Millar 1966) the gut loop extends anterior to the atrial siphon und occupies u relatively larger portion of the left side Remarks: This species hus been recorded often from) locations around Australia extending north 19 Indonesia and into the Pacific (Tukioka. 1967). The species is distinguished by the absence of intermediate papillac in the branchial sac. hy the heavily ribbed broad dor- sal lamina, and by the origin of the atrial siptraw from the middle of the body. Although in the present specimens the atrial siphon as long and directed anteriorly, in specimens pre- viously deseribed there js a great variation hoth in the length uf the atnul siphon and in its orientation ¢Michaelsen & Hartmeyer 1928)- Specimens have been deseribed with sessile ex- ternal apertures umd it is probable that the pre- sent specimens with short grooved cylinders represent more mature individuals. Exrernully the species resembles bouh 4. svdieyensiv and A. thompyoné and it is probable that in all these species the test is firmer and relatively thicker and the extemal siphons less evident in the younger specimens, while in older speci- mens the test becomes rougher externally and less transparent, and the external siphons dJeve- lop as short grooved cylinders. The body mus- culgture. concentrated on the right and on the siphons, is so arranged that the left side, fixed io the substrate, does not contract over the voluminous gut. In these specics the gut eceu- pies a relatively smaller proportion of the body wall as the individual inereases in size. In A. genimatad growth appears to increase the pro- portion of the body anterior to the gut, and although the point of origin of the atrial siphon remains about onc half ta two thirds of the distance down the body, the gut does not appear DO increase in sive gt the same rate as the rest of the body. The oricntation of ASCIDIANS OF SOUTH AUSTRALIA ar the gectum and the curvature of the gut loop is therefore reduced us growth proceeds. 1 is also possible that this cifferential growth caitses the varigtions that have been observed in the length and orientation of the atrial siphon, although this may also be affected by the oricn- tation of the body on the substrite, Axeidia muilacca ansiteliensis Hartmever, 1928, resembles the present species in the pre- senge of a broad ribbed dorsal laming with the Iree margin produced into pointed projections sorresponding to the ribs, However. the spe- Ges is distinguished by the specially long exter- nal siphons. by the dorsal ganglion which js only one-ninth to ofe-thirleenth of the body length from thé dorsal tubercle, and by the amall stumpy cone-like branchial papillae as opposed to the sputulate papillae of 4. ge mao. Harimeyer's subspecies was recorded from a seasonally hvackish environment in Freshwater Bay, a considerable distance up the Swan River estuary from Fremantle Harbour and he regurded jt as an isolated endemic spe- cies, Ascidia thompsoni Kott. 1952: 312. Now Reeords: OF West Beach. Hullett Cove, Calickalinga Head, Previews Recordy: Tas. (Greai Taylor Bayi—-Kott 1952. FIGS. 41-45 Pescripion: In smaller specimens the test is firm and almost glassy and transparent. Ante- rioly, expanded terminal ampullac of the test vessels are clearly visible through the test. Indi- viduals from 2 to 7 cm Jong are available tn the present collection. Both apertures are sessile. the branchial aperture terminal and the ttrial aperture two-thirds of the distance down the dorsul surface. Most individuals are firmly fixed by the whole of the left side, however the apecimen fron} Carickalinga Head is fixed posteriorly. The body musculature is present only on the right side, consisting af a mesh of dvansVerse and longitudinal vessels Internully the atrial aperture is on a siphon of variable length ising apposite, anterior or pos- terior 10 the external opening. The atrial siphon shows the same vanations in length and orientation as have been described previously for A. geminata (Michaelson & Hartmeyer 1928; Millar 1966). Both siphons are well equipped with circular and longitudinal muscles, There arc about 40 branchial ten- tacles, a papillated prebranchial area, a shallow perituberculir area completely filled by the dorsal tubercle which generally has a sirnple U-shaped opening. In an especially lurge and Opaque specimen from West Beach (at % 1) there is a second opening to the feht ol the larger U-shaped opening. The dorsal ganglion is about half the bouy distant from the dursal tubercle, The dorsal Jamina is 2 wide mem- brane, dowble for about one-sixth of its length, The right section of the double membrane is plain, the left section is ribbed un the left. For the remainder of its fength the dorsal lamina is a single membrane ribbed on the left side, although these cibs do not extend to the outer margin of the membrane. There are minute ind irregular papilla-like expansions from the free border of the membrane tn its posterior extenl. Intermediate branchial papillae are generally present, especially in the posterior part of the branchial sac, The intermediare hranehial papillae are half the size of the pri- mury papillae, and bowh are pointed. ‘The gut is voluminous and forms a deep double loop which varies slightly in relation to the atrial siphon as the individual grows, us in 4. gen)- freilet, Remarks: Ihe dowble dorsal famina with slightly irregular membranous border pos- teriorly and the form of the intermediate and primary branchial papillae distinguish this spe- vies from the very similar A. genceta with whith tty geographic runge overlaps, The origin ind the variable orientation of the atrial siphon are shured by the two species, and in bath, owing to differential wrowth of the body, the gut loop is confined to the posterior half of the left side in latger specimens, Tt is of consider- able interest that the present species has been recorded only from fairly sheltered constal environments (subject however ta some wave action) in the present collections. while 4, geninlase was taken only from Offshore Benthic locations subject to currents in middle and \pper St. Vincent Gult Ascidia aclara Kot), 1952: 309. Millar, 1963: 721; New Record: Off Seacliff. Previous Records: Vic, (Lakes Entrance, Port Phillip Bay)—Kott 1952, Millar L963, Cd, (Moreton Bay)—unpublished recurds, PIGS. 44, 45 Description, There are two specimens in the present collection, maximum length 17 em and 10 cm high The body is slightly darso-ven- irally flattened. The test is rigid anu encrusted with suind and shell particles and is produced into two tigid cylindrical tubes from around as PALRICIA KOTT the branchial and atrial apertures at the anterior end of the dorsal surface and from about onc- third of the distance ulong the dorsal surface respectively, The apertures afte completely sessile und lie at the base of these tubes, The hody nvseulature, within this rigid test, is reduced to strong bands ucross the dorsal sur- face posterior to the atrial aperture ind between the atrial and branchial apertures. Internally the specimens are exactly as pre- viously described with the branchial sue tormm- ing a fold across the dorsal tubercle. The gut forms ihe usual simple open Joop, opening adjacent to the alrtal upercure. Remarks; This unusual species apprars to be highly specialised for an existence on a sandy bottom, with the rigid tubes extending vertically from the apertures forming « permancnily open channel thraugh the layer of sand jo which the species is probably buried. It is probuble that the immediate environment outside the apertures is madified by these permanently open chambers to facilitate a less interrupted feeding process and confer distinct »dvantuges In locations where steady flowing currents and absence of sedimentation pertain, The species is also of considerable interest in that ils records are confined to the semi-enclosed waters indicated above. IL is possible that there Is a Wider, more continuous distribution on the continental shelf or, alternatively. that it represeuts a Telict population of a species which once hud such a continuous distribution on the open coast. Suborder STOLIDOBRANCHIA Family STYELIDAE Subfamily PaLyzornak Stoloniiy australis Michuelsen, Michaelsen & HMartmeyer. Kott, 1952; 253, New Records: Vipara Reef. Por Noarlunga. Previows Records: W. Aust. LAlbuny)— Michaelsen 1927; Michaclsen & Hartmeyer {4928 ‘Tas, {Spring Bay}—Kott 1952, FIG, 46 Dexcriptlon: Rounded, sandy. stalked or sessile dividuals connected to basal stolons, 0,6 to (7 em maximum diameter. The colonies in the present collection are encrusting specimens of Pyne irregularis and Polyearpa peditnen- love, The apertures are both sessile on the upper surface. There ate two folds on either side of the branchial sac with 6 to 9 internal longitidinal vessels. The gut loop is simple 1927; 928: 202, 352. and open with a gastro-intestinul ligument enclosing a rounded endogurp in the pole, The short stomach has about 18 Folds. tt is. reduced jn diameter at either end and his a thick pyloric caecum of moderate length. Gonads are not malure in the present specimens und their arrangement could not be determined. The ligaments anchoring the gut to the body wall! extend in a tow along the lateral aspect of the intestine. There are also large hygaments anchoring the stomach and the pole of the gul loop. Remarks: This species appeyrs to he confined to the southern coast of Australia, but has been recorded only from locdions away from the open coast, It is inconspicuous. however, and iL is possible that its occurrence in protected locations on the open coast has been over- looked. [nthe absence of mature gonads the species miy be distinguished from dmplicarpa dipiycha by the low rounded branchial folds, the presence of a curved pyloric caecum and the less developed musculature. Stolonica carnosa Millar, 1963: 734. New Record: ‘Vipara Reef. Previnses Record; Wo Aust. (Cottesloe), FIG. 47 Description: The colony is oval, 3 em long, 2 em wide and | cm thick and, as in the type specimen, hus developed around iin algal stem. The 4-lobed apertures of zooids are close Togelher an slight swellings all yround the outer surface which is encrusted with sand, There is nO sand inside the colony, Each indivatual ix Jorsa-ventrally flattened and most of ils left side is directed toward the centre of Lhe colony. There ure 2 folds on euch side of the branchial sur with internal longitudinal vessels according to the following formula. E O( 434491 DL, There are only 3 stigmata between the endo- style aod the ventral fold. The cut forms a rounded loop and the rec- tum turns anteriorly and dersally at a sharp angle. The stomach is pyriform, narrowest at the curiae end, has 15 pacrow folds and a very long, curved pyloric caecum in the pole of the put loop. There is a gastro-intestinal ligament and ligaments connecting the gul loop to the body wall as in Distemuty diptyche (see Kort 1952). The anus is 2-ipped. The gonads are in single rows on cach side of the endo- style, The testes arc fMask-shaped and the ovaries contain 3 exes of varying sizes, and a testis and an ovary are generally loosely asso- ciated so that there are 6 to 7 hermaphrodite gonads on each side af the body, ASCIDIANS OF SOUTH ATISTRALIA ay Remarks; Although in Millur's specimen the testes and Ovaries appeared often to be sepu- rate, the condition and the artungement of the gonads in the present colony suggest that this is Mole apparent than real. and may depend on the relative stages of Uevelopment af the ovary In Millar’s specimen the stomach is folded internally but externally the tokds were pro- hubly obscured by the membrane covering them. The course af the reetum in the present specrmen also differs from Millar's specimen and is bent back against the gut loop, prohahly by dorso-ventral flattening of the individual. The extent of vhis dorso-ventral flattening. therefore, is am individual, rather than # speci- fic. character. Oculinaria australis Gray, (868: 564. Katt, 1952: 25) and synonymy. Millar, 1963: 73d; 1956; 369, New Records: West 1. (Scal Rock), Wright I. Previous Recérdy: W, Aust. (Fremantle to Albany )—Gray 1868; Michaelsen & Hart- meyer 1928; Kott 1952; Millar 1963, Vie. (Port Phillip Bay)—Millar 1966. Description: Colonies of the usual form with numerous zuoids closely coalesced, identified only by the paired apertures on wart-like siphons from the anterior surface of each zanid which project slightly from the other- wise campact colony. The test is very brittle and completely (Mpregnaled with sand. There are 4 branchtal tolds on each side of the body with 4+ to 8 longitudinal vessels on cach Fold and about 4 between the folds, The gut loop is as previously described, with about [8 spiral folds in the stomach wall. No pyloric caecum has heen detected. There is an elongute gas- tric gland reservoir extending between the stomach and the intestine, There are up to 9 lonz gonads on the right side of the hady, a lurger number than hus previously been recorded for this species. There is a single row of testis lobes beneath cuch short ovary. Remarks: The species is well adapted, by its compacted form, far the occupation of turhu- lent locations and, in fact, it has been recorded only Erom the exposed open coast, Externally iL resembles colontes of Palyandrocarpa spp. from which it is readily distinguished not only hy the location of the gonads on one side of the bady, but also by the spiral course of the stomach folds, the presence of 4 gastro-jntes- tinal reservoir and the form of tbe gut loop Subfamily moveyoirivar Botrylnides leachl (Savigny), Michaelsen & Huitmeyer, 1928: 341 and synonymy Millar, 1952: 24.1962: 177) Kort, | 952) 233; 1966: 297 Boiryllus leachii Savigny, 18107: 7. New Records: Tipara Reef, Port Noarlunga, West 1, Wright l. Previews Recarey: W. Aust. (Geraldton to Albany) —Michaelsen & Hartmeyer 1928; Kott, 1952, N.S.W. (Port Jackson)—Herdman 1899. Qld. (Moreton Bay}—Kott 1952; (Sarina)—unpublished record, Northern Territory ¢Darwinj— Kott 1946, New Zealand (Hauraki Gulf )— Michaelsen 1921; Brewin 1948; (Stewart |.) —Michaelsen 192[; (French Pass)—-Sluiter 1900; (Otago Harbourj—Brewin 1946; {Auckland 1,}—Bovien 1922. South Africa —Hartmeyer 1912; Millar 1962. The spe cies 2s alsa Known from the North Atlantic. the North Sea and the Mediterranean and Adriatic (see Hartmever 1923, Arnbiick 1923, and Millar L952). Deseription; Living colonies from Ocdipus Point, West T. have u colourless matrix and red zooids, while in those from Port Noarlunga the mulrix (s transparent and the zooids yellow- bright orange. All the colonies have trans- lucent test and purple zooids in preservative Colonies form fMattencd, long, Iohes with a shore stalk. There ate circular to oval svsiems of closely packed zoids. The test i< firm and transparent. The system af zooids are arranged in rows along the Jength of the head, These systems May appear to be confluent and form almost continuous rows, but in fact separate cloacal openings remain in the centre of a limited number of zooids and dis¢rete circular to oval systems are niainivined) There are 9 lo 12 rows of about 20 stigmata. “Phe stamach is long, with 10 folds and u very short caecum, Remarks: The form of the colonies és very similar to those of B, magnicoecuim bul the circular systems and firm test, with common cloacal openings along the sides of the lobes ire distinctive. The shape of the stomach and the form and length of the pyloric caecum is similar to the condition found in B, nigrum. However, the smaller number of rows of stig- Mata with more stigmata in cach row also dis- tinguishes this specics from both B. magai- coecum and from 8. nigrum. Records for this species extend from the North Atlantic to the Medilterrancan and Pucific Oceans, and from All around Australia. It is not known from the 30 PATRICIA KOTT Indiun Ocean beyon the West Australian coast nor is it known fram the South Aflantic. Botrylloides nigrum Herdman, 1386: 50, Van Name, 1845: 227 and synonymy. Katt, 1952: 257 Surcvlniviiaides jacksonianam EHerdman, 1899: 102. penbeyestatat pamasniny Herdman, 1899; New Records’ Port Gawler, elf West Beach- Off Seneliff, Curickalinga Head, Rapid Head, West 1, (near Penguin Rock, Seal Rock). Wright | Previous Recards; W, Aust., 5. Aust. Vie—Kott 1952. N.S.W. (Port Jack- som t— dlerdmarn 1899; Kott 1952. Old— Kote 1952) Indo-Malaya (Ceylon)—Herd- man 1906: (Red Sea)—-Michaelsen 1919. Fast Africa-—Shuiter 189%, Michaelsen 1918. South Atrica—Hartmeyer 1912. The spe- cies is also recorded from the Curibbean region (Van Name 1945), Deyeription: Colonies investing sheets some- times extended into irregular lobes, The zooids wie arranged in long double row systems well separaled from one another with transparent test between In preservative the zooids are purple-lback with the pigment contained in cells in the hady wall. The colour of the pre- served specimens does not reflect the variations in colour of the hving specimens which irc: “dark blue and bright purple” zaoids {West §.): ay “vellow and musturd” (off West Beach), There ave 16 rows of about 12 stigmata with 3 internal longitudinal vessels of each side of the hranechial sac. The atrial opening exposes the anterior half of the dorsi! surface of the bran- chial sac, but the lip from the anterior border of this opening ts not especially pronounced. Ihe stomach is the usual long organ charac- teristic wl this species, with 10 folds, I is wider at the cardiac end and reduced in width at the pyloric end where there ty a very short cuecuin. Retnarks; Aithough the variation in colour and the irregularity of the colunies make this spe- cies difficult io identify in the field, the: shape of the stomach with its shurt caccum and the widely spaced double rows of zooids are dis- linctive. [ts recorded distribulion is wide in the Indian Qceun and from the West Indies. At this stage there is no known character avail- uble (o indieute that all these records refer to more than the ane species with an almost cir- cumpoler distribution. in the southern temper- ate region, absent onty from the middle and eastern Pacific Ocean. Rotryloides magnicwecum Huttmeyer. Kott, 1952: 258, Millyr, 1966: 368. Borylloides nigrum var, magnicoecum Hart- meyer, 1912; 271, Reavllus magnicoecus. Michaelsen, 1923b: 50; 1923: 4, Mivhaclsen & Hurimeyer, 1928; 331 and synonymy. Hustings, J93L. 79. Brewin, 1951: (09. Millar. 1958: [95> |9n2- 175. Tokioka, 1967; 153, Bolryllay auceps Michaelsen & Harimeyer, 1978; 335. Millar, (963: 774. Palvexelas rufus Oka. 1927) 6b8 Betevilius refs. Tokioks. 1943h: 240 New Records: Off West Beach. West b. Wright J. Previews Reecards; W. Aust (Shark Bayj—Michwelsen & Hartmeyer 1928. S. Aust, Tas.—Kott 1952, Vic (Port Phillip Bay}—Millar 1963, 1966 N.S.W. (Port Jackson) -Herdman 1891; Millar 1963. Qld. (Great Barricr Recf}— Hastinus 1991. New Zealand (North b.j)— Michaelsen 1921: Brewin 195). Japan— Tokinka 1952; Oka 1927. Chinu (Hony Kong)}—Michaclsen 19232, Tokioka 1967, tadian = Ovean (Paumbur) — Michuelsen 19235. South Africa—Hartmeyer 1912; Millar 1955, 1962. South West Africu— Hartmeyer 1913; Michaelsen (915. Natal— Michnelsen 1918, 1921. Rurope (Portugal) Tyar—Michaelsen 1923b; (Mediterranean) ‘var. Michaelsen 1923b. Meserprons The living colonics from West 1. are “bright yellow" although other specimens are “oreyish with pale zoojds”, In preservative, however, all the colonies are purple owing to the pigmentatiow of the zovids which shows through the very solt transparent test. The colonies in this collection always consist of soft, long, narrow, Mattencd, stalked lobes with zooids arranged in closcly set double rews running parallel to the Jength of the lobes Zooids are absent from the stalks. In preserved Specimens there is always an accumulation of dark pigment at the top of the endostyle and on either side of the bine of the branchial aper- ture Common cloacal openings are ulwiys present around the free end of the lobe as. in Syvrezoqd spp. There ure |4 rows of stigmata in the present specimens with 3 tw 4 stiymata between the Jongitudinal yessels. The stomach is short and rounded with 9 folds und a long caccum curv- ing into the pole of the gut loop. Remarks: Millar (1963) regards the form of ihe colony of the Australian specimens (long stalked lobes) as providing & character which distinguishes it from the South African forms which are irregularly lobed and tnvesting, as ASCIDLANS OF SQUTH AUSTRALIA 3) are Brewin’s specimens from New Zealand, The closely set double row branching systems are present in all the specimens represented in the synonymy above and all these specimens have the characteristic shart, rounded, stomach with a long curved caecum. distinzuishing them From other spectes of the genus It is possible that the Australiin members of this species may represent a geographic subspecies charac- terised by the particular form of the colony with terminal cloacal apertures and close-set double rows of zooids parallel to the longi- tudinal axis af the head. Bostrylinides leacht colonics are similurly lobed but the cloacal apertures are present along the side of the head hetween the double raw ol zoeids- Botryllus schlosseri (Pallas). Van Name, 1945; 220 and synonymy, Kott. 1953: 259 {purt), Aleyaniumn yehlasseri Pallas. 17662 454. Non Aeeryllus sehlosserd. Kott, 1952. from Hamelin Bay and Green Pools, W.A, New Recera: OF Hallett Cove, Previnne Records: W. Aust. (Shark Bay, Fremantle}— Hartmeyer & Michaelsen 1928: Kou 1952 Vic, (Port Phillip Bay)—Millir 1966. Else- where the species has a wide distyibution from the Faeroe Is. and southern Norway. the British Istes, the North Sea. the Mediter- fanedn, Adrintic and Black Seu; from the eastern and western seaboards of the U.S.A- and from New Zealand (see Van Name 1945), The locul abtindance of this species and its occurrence on wharf piles. ship hulls, buoys, ete. in shallow water has been pointed out by Van Name (1945). ‘This wide cosmopolitan distribution suggests that, like Ciona intosrin- «lis, the species favours sheltered locations and is transported largely by ships. Description. The specimens are delicate and invest the sea vrass Poyidania australis, The lest is almost completely transparent and the zoaids are pale grey, Zooids form srnall cir- euwlar systems which ate crowded close to- gether in the test. The zooids are relatively short. with only about 8 rows of stigmata, The atrial aperture is on a siphon produced to a varying extent and the upper margin of the aperture is produced into a lip. There is a conspicuous pyloric caecum: with a large bulb- like expansion on its free end. The stomuch has about 10 very fine folds. is longer than wide, and is only of slighty greater diameter than the fest of the gut. Developing embryos are present in the peribranchial cavity of some of the zovids, bul on the right side of the hos only. Remarks: The zooids ina colony of the present specimens are identical with those described for Borrvilix gracilis Harimever & Michaclsen 1928; Millar, 1966, from Shark Bay, Western Australia and fram Port Phillip Bay. Millar (1966) regards this type of thin transparent colony us a species distinct from B. sechlosseri. Juvenile colomes of B. schlosseri as described by Verrill (Verrill & Smith 1873) are identical with the presont coluny and the zooids are iden- tical with those previously described for this species especially in regard to the atrial open- ing, stamach anc pyloric caecum. and it Is unlikely that graciliv is distinct from &. schlosseri. Subfamily stveciwae Cnemidocarpa etherldgii (Herdman) Stycle etheridgii Herdman, 1899; 38, Kolt. 1952: 219 and synonymy; (964: 129 (F. per- sonata), Millar, 1966: 370, New Records: Tapley Shoal. off West Beach, West J. (off Oedipus Point), Wright . Pre- vious Records: W, Aust. (Vriga 1). 8. Aust. (Spencer Gulf and St. Vincent Gulf}. ie. (Phillip 1,1 —Kott 1952: (Port Phiflip Bay) —Millar 1966. Tas. (D’Entrecasteaus Channel}—Koatt 1952, N.S.W. (Part Juck- son, Port Stephens)—Herdman 1899. Qlu- (Morcton Bay)-—Kott 1964. 'Uhe species is known intertidally and down to 30 m. It is abundant in St. Vincent Gulf on sanity boltoms at 7-20 in with slow currents, and On Open coasts in decper water of 20-30 m (Shepherd, pers. comm.). FIGS. 48, 49 Description: Individuals are large. up to 11 em high, rounded and of greatest diameter pus- lertorly. gradually reducing in diameter ta the terminal branchial aperture. The terminal branchial aperture is sometimes curved. The atrial aperture ts on a slight rounded projec- tion from about half way along the dorsal surface. Colour of living specimens Varies Fram pale cream to bright vellow (most often the fatter), In preservative the lest is white and Opaque, with longitudinal furrows converging to the branchial aperture on that part of the body anterior to the atrial aperture, The test is thin and leathery, There are up to 25 inter- Nal longitudinal vessels on the folds and up to 7 between, although in some specimens there are as few ys 4 internal longitudinal vessels between the folds. There are 6 stigmata per mesh. 42 PATRICIA ROTT Fig. 4G, Fig. 47. Individuul bisected along the ventral surface, and endocarps in body wall. Fig, 50. Polyearpa clavari. Whole individual, Stalonicu australis, (Tipara Reet), Gut loop. Stolonica carnosa. (Uiparn Reef). Right side of body removed to show organs on left hody wall. Figs. 48, 49. Cnemidocurpa ctheridgii. (Tapley Shoal, 13m). Fig. 48—Whole individual. Fig, 49.—- branchial suc removed, showing gonuds Figs. 51, 52. Polvearpa pupillata. (Of Glenelg, 15m). Fig. 51,—Body wall on left showing gut loop, gonads and endocarps. Tig. $2.—Individual showing gonads on right side of the body. Figs, 53-36. Polyearpa pedunculata, Fig, 53.—Individual from Aldinga (10-25 m). Fig, 54—indivi- dual from Taptey Shoal (24 m). Fig. §5.—Individual from West I. (25 m). Fig. 56—: Gat Joop and endocarp, The gut forms a gently curved, fairly narrow loop across the left side of the posterior end of the body, enclosing a Jong narrow curved endocarp which is continuous with the body will on both the right and left side. The gut loop is almost entirely posterior to the bran- chial sac: the elongate stomuch und proximal part of the intestine forming the proximal limb of the gut loop lie almost in the mid line pos- tero-ventrally and the distal limb of the gut loop passes to the Teft of the posterior end of the branchial sac. The gut loop is almost ASCIDIANS OF SOLTH AUSTRALIA 33 entirely embedded [hn the thickened body wall und i covered by endocatp which encloses. the left gonad (in the curve of the gut) and extends ventrally iacross the pole of the gut loop to join the thickened body wall ventrally and pos- terjorly, The pole af the gut loop thus projects into jt pocket in the thickened body wall, The ovsophagus is short and the stomach is long and elliptical with internal longitudinal glundu- lar folds, The anal opening has a smooth border, There ure one or two flask-shaped gonads on the myht side of the body. On the lelt the gonad is embedded in a single large cndocarpal thickening Of the body wall, Here there may he a single branched gonad with single 3 and ¢ ducts emerging from the endocurp and directed to the utrial aperture “This condition may have resulted from the fusion of two gonads. In another specimen there are lwo discrete yonuds embedded in the teft side of the body with their own sets of cf and © ducts emerging from the endocarp. The testis lobes are enclosed by the ovarian tube as is characteristic of this genus. Remarks: The present farge specimens con- form with those ascribed (Row 1952) to the “erheridgii” condition af this species, It is most probable that this distinction relates only lo the stage oF maturay of the individual, where the “persuran” condition represents Jess mature individuals. Both forms hive been taken [rom the same locations in both cast and western Australia. Polycarpa clavata Hartimeyer. Millar. 1963: 723. Polvearpa mau (Quoy & Gaimardy f. chwaty Hartmeyer, 1919: 40, Michachen & Hartmeyer, 1928: 363. Kott, (952: 23h, Tokiokn. 1961; 123, Vasseur, 1967; 133. New Revoards: “Vapley Shoal, near Marion Light, off Troubridge Light. Previons Reoordys: W. Aust, {Bathurst J, to Rottnest [y—Hartmeyer 1919; Kort 1952: Millar 1963. Pucific (Noumes, New Ciledonia) Tolkinka 1961: Vasseur 1967, FIG, 50 Deserfptlon: Large stalked specimens [rom fawn to reddish-brown The test is very soft and gelatinous and the surface is marked with rounded longitudinal ridges which are some- times interrupted horizontally. The branchial uperture is on a short siphon from the basal une third of the dorsal surface. P. suleata) thar Hiusting’s specimen is identical, A re-examina- (ion of the type specimen of P. atraie var. plana Herdnvan. 1899 from Port Jackson has shown that its gonads are also the usual short polycurps of ?. anrare which is now knoWwo from Port Juckson and the Great Barrier Reet und from the Indian Ocean. Mulaya, and Indo- nesia The ranye of P-avrete, therefore, dors now overlap that of P. efavera, Polycarpa papiliata (S|uiter). Stvela pupitlata Sluiter, 1886: 192. Tokika, 1952: LI7. Vasseur, 1969: 925, Palycarpea intéstitata Kott, 1952: 23k. New Records: Tipara Reet, olf Port Gawler, uff Glenele. Aldinga “drop-off” Previous Records: NSW, (Port Jackson)—Kott 1452. India) Oceut (Madagascar)— Vasscur 1969, Indonesia Sluiter 1886: (Arafura Sea}—Tokioka 1952. FIGS. 5], 52 Descriprion: Small ugeregates of individuals, the posterjor fest sometimes extended into a short stalk, The branchial aperture is terminal, the alrial aperture one third t one half of the distance along the dorsal surface, Both uper- tures are sessile. The test ix tough, rough and wrinkled exterqully, with some sand and algae irregularly adhering but yenerully the surface test is muiked. The body musculature consists of a moderately thick continuous external coat of circular muscles with longitudinal bands internally. The dorsal tubercle is a large blister- like swelling with a simple U-shaped opening; i} completely fills the V of the peri-tubercular area. There are 4 wide overlapping folds with about 15 jaternal longitudinal vessels on ech fold and 3 wo 4 between folds. There are 4 § stigmata in cach mesh. Anteriorly the endo- stvle Follows a winding Goutse, which ix effected by the subdyvision of transverse vessels and multiplication af the puntber of rows of stiz- mata ventrally, in a localised region slong the anterior cxtent of the endostyle. The gut forms a hnrizantal loop in the posterior end of the body. The stemach is olliptical with Jongi- tudinal striations. The rectum extends ante- tiorly toward the atrial opening. The anul border is broken up into 14 long fingerstike lobes, Tall endecarps are present im the vut loop and seatiered over the body wall. Seven ta 12 aval to elongate polycarps pre present in 1 to 2 rows in the centre of cach side of the body, directed lowand the atria] aperture. These polycarps ute fixed to the body wall wong their whole length Ino smaller specimens with smaller immature gonads there are more often 2 rows of polycurps, anu. as the gonads increase in length and the body length increases, (hese rows upper to merge inte au single inreguilier row, while in a single specimen with well deve- loped gonads there is only a single regular row, Remerky: The present species resembles Paly- cerpa clavate (Hartmeyer), Po lavnet/urreis Tokioka and P. attellens Herman, in the tall endocarps enclosed ih the put loep, but (5 dis- tinguished by the rows of recumbent gonads fixed alone their whole length to the body wall. The anal lobes also resemble those of P. atollers and PQ lengiforpis, The form of the body, the position of the attial aperture, che form of the dorsal rubercle. and the form and arrangement of the gonads are similar to P. emrctinurdte (Sluitert, Vasseur, 1967, which is distinguished by its short oval stomach. greater number of rows of gonads und greater number of internal longitudinal vessels between the branchial folds, Cnemidocarpa omiutedagascariensty — plelagas- carteusts Harumeyer from Madagascar and C, moadegascarienyis reguliy Michelsen from New Zealand (see Kott 197]a) also resemble the present Species in external appearance and in the arrangement of gonads, and are cistin- guished principally by the greater length of the gut loap and greater number of internal lonyi- tudinal vessels; between the branchial foldy The papillac on the branchial sac described by Sluier (1886) are not present cither m the South Australian specimens or tn the specimens fram the Arafura Sea (Tokioka 952). Tr ts possible that Sluiter mistook particles adhering to the branchial sac for papillae, ASCIDIANS OF SOUTH AUSTRALIA ss The species has a wide geographical cistribu- con from Indonesiq and apparently around the cast Goast of Australia, from focky subsirates I) sheltered localities, or in Olfshore Benthic locations where there are slight currents. Populations of Lhis species du not appear to be dense and tecards are few, Polycarpa pedunculita Heller, 1878; 106. Kott, 1952; 237 and synonymy. Miliar, 1966: 369, Falscurpa e@bsenra. Kott, (952: 245 und spnanynty, alycurpa steplienensis Herdmun, 1899; 45, Koit, 1952: 232. Millar. 1963, 726, Polyearpu moehii, Katt, (952: 244 and syno- nymy; 1966: 299. Vasseur, 1967: 136, Polycaurpu obrecta. Kot, (52: 242 (nat P. ohrecta Traustedt), New Records: Vipara Reef. Tapley Shoal, near Marion laght upper St. Vincent Gull, off Port Gawler. off Semaphore. off Grange, aff West Beuch, off Glencly, off Broadway, otf Hallett Cove. Port Noarlunga, Aldings, Cinckalinga Heal, Rupid Head. West L., Wright |. Previnns Records: W. Aust. (Cape Ivuhert to Bunhburvi—Hurtmeyer 1919; Michvelsen & Haortmeyer 1928; Katt 1952, S. Aust, (Reeveshy 1). Vie, ( Balnarring Beach )—Kute 1952; (Bass Straiti— Heller 1878; Michaclsen 1905; (Port Phillip Bay) —Millur 1966. N.S.W. {Port Jackson, Two- fold Bay}—Herdman 1881. Qld, (Mereton Bay}—Kotl 1964, The species hus dlsu been recorded fram New Claledonia (Vasseur 1967). FIGS, 53-56 Description. This ts by tar the most common ascidian in St. Vincent Gulf and is very vari- able to external appeatance, The colour of living specimens from Port Nourlunga has been described us “bright to pale yellow”. ‘Vhese specimens are black to greenish in preservative. Most often living specimens are sandy with a “reddish linge” to “reddish brown" becoming brown to purplish brown when preserved in formalin. ‘They are slightly laterally flaiened and almost oval shaped, and are most often 3 to 4 cm long and 2 to 3 cm wide, Larger specimens up to 8 cm long ure usually grecnish- black in preservitive. The apertures are ses- silé. the branchial aperture terminal but direc- ted slightly to the side, away from the dorsul surface, and the atrial aperture one-third of the distance down the dorsal surface, The test. is tirm and gelatinous and the sur- face is generally smnuch and naked, There is often. hawever, 4 light encrustation ol sand or the test may be more heavily encrusted. or may heeome almost brittle with included sand, In larger specimens the test becomes thinner. more flaccid and leathery, Posteriorly the dest may he produced into a narrow atalk up to half the length of the bouy, or the body may taper gradually from a straight upper or anterior surface whene the branchial aperture ts central and the atrialaperture is on the antero-dorsal corner. The posterior end of the body, with or without iw stalk, may be pro- duced into root-like structures, or the incivi- dual may be fixed to the substrate by the postero-ventral surface. Ihe body wall is Jight tu dark brown, hrownish-green. greenish-black, or black. It is not very closely adherent to the test and js thick, firm and very muscular with internul longitudinal bands and a continuwus. thick eXternal coat of circular muscles. Both layers of musculatiire are often embedded in fleshy non-muscwlar tissue and yvencrally spherical vesicles are embedded in the smuscle layers interrupting the regularity and cominwity of the fibres. The hudy wall is more flaccid |W larger specimens. There ate ubeut 100 simple tentacles at al least 4 orders. The prépharyngeal arca has small papiive and is of moderate width, The dorsal tubercle varies and is sometimes small, in the centre of a fairly Large peritubercular area. Wt is sometimes much Jarger but never completely fills the perituberculur area. The opening forms a U with borns turned in or out and directed to the side, antesiorly or poste riurly and in larger specimens may be inter- rupted. The dorsal lamina is a plain edged narrow membrane. The branchial folds are low and rounded with 2 ta 3 thick internal longitudinal vessels between the folds and LL to 13 on the folds. There are f to B stigmara in each mesh between the folds but on the folds the jnternal longitudinal vessels are more crowded together. There are often vesicles, similar to those embedded in the body wall, embedded in the branchial vessels and in the dorsal tubercle, The gut is confined to the pos- terior cnd of the body distal to the atrial aper- ture. ‘The intestine forms a short rounded loop enclosing 4 circular endocarp. The stomach itself is elliptical with pronounced fulds, There may be a second small endocarp separating the rectum from the ocsophagus as the former extends anteriorly toward the base of the atrial opening. In smaller specimens the anal border is broken into 7 sometimes subdivided rounded Ais) PATRICIA KOTI lobes. In larger specimens there are up to 25 lobes. The circular endocurp enclosed by the gut appears to be the miyjor mechanism anchor- ing: the gut loop to the badly wall and is con- fluent with the connective tissue surrounding the gut. There are 20 te 50 short oval poly- carps on the Jef and 25 to 60 on the right These are sometimes, but not always, embedded completely in the body will. Wher completely embedded only the openings of the ducts wre apparent us holes. in che inner surface of the hody wall. Primarily there appear to be about 3 longitudinal rows of polycarps on euch side of the body. As each polycurp increases in length it subadivides and new gonoduols open from the proximal half to form secondary rows of gonads overlapping the primury row closest to the atrial opening. Wi is possible that. this process, resulting in Increases in the number of polcarps present, explains the great variacion in the number recorded for this species. Remerkss Mivhuelsen & Hartmeyer (1928) dew. attention to Lhe similarity between species listed in the s¥nonymy above and suggest that P. obyeura is a vuriely of PL peduneulata CP. viridis). Michaelsen regarded = Polvcurpa moebu, however, as 4 distinct species charac- ierisedl by differences in the gut and gonads. In this collection there are individuals demon- strating every condition previously deseribed tor P. pedtinenlata, P. moebil, P. obseura and P. siephenensiy. There are specamens demon- vtrating every condition from stalked or ranted io sessile individuals; every colour und every condition of the test is found and there ps con- siderable yariation in the number of polycarps and the extent lo whieh they are enthedcled. The gut loop is always constant and encloses the cireulitr endocarp which has. a pointed tip dorsally, The thick internal longitudinal yes- sely of the teanchial sac. their erowding on the nartow tolds. the spherical vesicles embedded in the hranehial sac amd hady wall, the thick layer of circuhir musele, and the pnupillated pre- branchial area can be regarded as characteristec of this otherwise highly Variable single spuctes. The extent to which gonads are embedded in the body wall. and the exient to which the body wall is marked off inte areas probably indieutes more mature specimens. Palycarpa macniara Hartmeyer, 1906, has a similar endocarp enclosed by the gut loop. and ihe sume type of vesicles embedded in the hody wall, Tt es distinguished from the present species, however. hy the weaker musculature which also distinguishes 1 tram the West Indian species PF. ehiecia “Traustedt. PL pedunculata is The most common ascidian in St. Vineent Gulf and generally both greenish and reddish brown specimens occur. Large bluck specimens were also taken from Seal Rock. West t. from Hallett Cove, und from Tapley Shoal. There is no apporent cerrela- lion hetween the Lype of environment and the colour of the individuals wt ¢uch Station. A cuse of genetic polymorphism im Ascidiacea has been described for Boltenia ovifera CL) (Ploush 1969). This dominant in the ascistian population of the Gulf of Maine, has colours ranging From while to crimson red in a single boul. and variations in lest texture aod in muscle band colour and thickness can be related to these coluur variations. It has been suguested that the species demonstrates genetic segregation of the ability of individuals co jiccumulite pizments, The situation in Paly- corny pedunenulata may indicate a similar gene- tie segregation, Family PYURIDAR Pyura scoresbiensis 1.5). Type Location: Off Semaphore: 18 m, in sparse Posidonia, 27.1.69 (Holotype: Soush Australian Museum, registration number E876). Further Recards: Of Tapley Shoal, ifm. 22m FIGS. 57-59 Deseription: Ronnded heads on stalks of vary- ing length, somcumes thick and no Jonger than the head, but sametimes long and. narrow (up to 20 em). supporting a head § em long and 3 cm wide. The heal is more or less cge- shaped with its greatest diameter basally hefore narrowing abruptly te the stalk. The apertures une both sessile, either side of « more ar less pointed projection forming the anterior apex of the head. The atrial aperture is slightly more posterior than the branchial apertare The test is thin, hare and tough with a dense sindy encrustation on the outer surface of the head and the stalk, The body wall is thin and semidtransparent with moderately developed fine and diffuse musculature, with muscle bands most closely placed around the anterior part of the branchial sac and siphon. The branchial tenwacles hive a large Hanged axis. fairly short prinvury branches. stumps secondary branches and minute tertiary branches and are not very bushy. The siphons are lined with long needle-like spines, closely set, up to O<75 mm tong. There are no sm- ASCIDIANS OF SOUTH AUSTRALIA 1 cules in either the test or the body wall. Vhe Horsal tubercle is a simple U-shaped opening with both horns lurned inwards. "The dorsal lamina has pointed languets but is very short owing to the close-set branchial and atrial siphons and contracted dorsum, The branchial sac is delicate with 6 high, overlapping folds on each side of the body with up to 20 internal longitudinal vessels On the olds and only 2 or 3 between, There are 4 to 6 stigmata in cach imesh, There ts a simple and fairly narrew eu Joop enclosing the gonad on the left. The gunad on the right occupies a corresponding position. ‘There are very arborescent liver lobules in the Tegion of the stamach, The gonad may consist fan undulating ovarian tohe with fringing testis follicles along hoth sides with the testis ducts extending along the mesial surfyce of the ovary. In some specimens the undulations of the covarian lube extend out into pinnate branches with tess follicles around their eXtremities. These pinnate branches may sub- sequently separate off into separate polycarp sacs on either side of a ventral duct. The anal border is divided into 3 large shallow Iohes, Hemerks: Specimens demonstrate the develop- ment of the polycurp sacs of the pyurid wonad from the continuous tubular styelid type of vonad. All stages of this development can be observed if the specimens available and it may be that the condition of the gonad indicates the age of the individuyl, The stalk’of this species alsa shows great variation in length and thick- ness. Despite these variations the species is characterised by the relatively smooth west, sand encrusted, bur without twhereles or furrows. und hy the constant position of the apertures. The pasition of the apertures, on the upper end nf the head, fairly close together, with the branchial anal atrial openings on opposile sides of the apex, is unusual in a stalked species of the Aseuliacea. where. more gererully, both apertures are on the dorsal side of the head with the branchial aperture directed uown- wards, and the atrial epefure uppermost and directed upwards. The relationships of this species are indicated by the stphonwl spines, which resemble those described for Pye albunyensis Michelsen & Hartmeyer, 1928, from Oyster Harbour. Albany, Western Australia. In which apertures wre also separated by » cushion of test in the middle of the upper surface and in which the dorsal surface of the hudy is very much con- tractedl and the dorsal! lamina comsequently very short. Pypra albanyensis has. however, charac teristic papillae on the convex bonier of the scubre-shaped stem and primary branches of the branchial tentacles. Pyara curvigona Tokioka, 1967, From the Palao Js. is a similar Closely related 300; 1969: 133. For further syno- nymy wid literature to the species in the Aulanuc and West Indies see Van Name 1945. 32). Cynthia vittata Simpson, 1852) 230. Pyura jacatrensis. Kou, 1952: 273; 1954- 127. Millar, 1960: 125. New Records: Tapley Shoal, off Trowbridge Light. Previous Records: W, Aust. (SW. Aust.)—Kott 1952, Tas.—Kott 1954, Qld, -Kott 1964, 1966, Pacific (Arafura Seca) ~-Tokioka 1952: (Palao Is.}-—-Tokioka 1967; (Japanj)—Tokioka |953a; Van Name 1945, Atlantic—WVan Name 1945; Pérés 1949) Millar 1960, Sub-antaretic (Mac- quarie I,)—Kott 1954, 1969; (Kerguelen 1.) —Kott 1954; (Mation I.)—Millar 1966. The species has a wide circumpolar distribu- tion in the southern hemisphere and extends north through the Indo-Malayun region to Japan, It is also found in the Auantic and in the Cartibhean (see Van Name 1945). FIG. 40 Description: Only a single individual is avan- able, 3 om long with a terminal branchial aper- ture and the vtrial opening half the distance alang the dorsal surface. Both apertures are almost sessile. “The external surface of the testis rough and has sand and foreign pirticles adhering. ‘The siphuns ure lined with tong needle-like spines, 0.1 mm to 0.2 mm long, overlapping. These extend onto the outer sur- face of the siphons, cover the lobes bordering the siphons and extend ome the outer layer wf test, The spines have a slight widescence ao VATRICLA KOLT which confers on this ouler siphonal urea a arcenish tinge. The siphon is lined with red siripes i the preserved specimen, The test is thin, leathery and firm, The dorsal tubercle is a rounded cushion filling the peritubercular aren with u simple U-shaped slit With both horns turned in. The branchial tentacles are nor bushy and have only primary briunehes und very shert secondary branches The internal siphons are fairly long. Longi- tudinal muscle bands radiate fram both siphons but do not extend very far down the bady on the Jeft. Circular muscles torn a Fairly irre- gulor network over the right side of the body. hecoming more sparse posteriorly. They are practically absent from the posterior hall of the hody on the lefe side, over the gut loop, The branchial suc is fairly delicate. It has 18 tnternal Jongitudinal vessels on cach folk! and 4 between. There ate 6 stigmata per mesh Phe gut forms the usual logo enclosing the lett gonad, The anal border is smooth and bi- labiute, The gonads consist of the usual cen- trol ovarian tube with pinnyte branehes on both sides terminating in polyearp-like sacs, Rndo- carp-like tissue 3s present on the free surface of the gonads where it is broken up into lobes. Remarks: Vhe synonymy of this widespread species has been very confusing owing ts) the variation in the length of the siphonal spines and the variation in the condition of the anal border, 11 appears. however, that Sluiter’s spe- cies from Indonesiit ind Northern Australia (P. jacutrensis). with very much smaller siphonal spines that do not extend onto the outer surface of the apertures, may he a distinct species Jes- pile the spines of intermediate length that are present in specimens from the Palau Istunds {Tokioks 1950; see Kott L9711. Pyura cvrvi- noma ‘Vokioka, 1967, trom Palao Is., 1s un wither closely related species in which the very long (2.75 mm) siphonul spines extend onto the outer surface of the apertures, In Pynra alhanyensis Michaelsen & Hartmever. 1928, and Pi xcoresbiensis nap. the siphonal spines extend up to 0.275 mm, only slightly Jonger than the present species. However, these siphonal spines do not extend onto che outer surface uf the apertures. Hyura icregularis (Herdman), Kott, 1952: 271 Millar, 1963; 739: 1966; 370, Cwuthia ireecularis Herdman, 1881: 60; 1882+ 141 New Records: Tipara Reef, off Beach Hut, T ke off Port Vincent, upper St. Vinoent Gulf. off Grange, off West Beach, off Glen- elz, Port Nogtiunga, Aldinga “drop-oll". Cunchalings Head, Previous Records: S, Aust. (Quter Harhour), Vic, (Port Phillip Ray)—Millar 1963. 1966. Tas. (D’Entre- casteaux Channel)—Kott 1952. NWSW | Port Jackson)—Herdman 1852. The spe- cits fas nol prewously bean taken ih waters of less than 25 m in depth Deseviptior; Living specimens are red. orange tu light fawn. Externally the test is very hurd. leathery and wrinkled and thickened inte small oelugonul plates. There are also wart- like protuberances, especially anteriorly. Individuals are usually cluniped together in light aggregates und the shape of the body is consequently very icmegulan The maxim body length is about 2.5 em. Both apertures afe present af the end of EGuurly long siphons which are zenefally oriented away from one another. The test is very strong with internal longitudinal art outer circular muscle bands ws in all species of Pyuridue. Delicate cup-shaped scales, 0.02 mm long. line the siphons, There ure 15 branchial ten- tacles with short sparse primary branches wnd minute secondary branches. The primary opening from the neural gland is U-shaped with horns turned i er out. The dursal tubercle is blister-like and there is often an uccessory openiny [rom the neural gland, The tubercle is not always longitudinally attenua- ted. however the peritubercular are 15 always a very deep V-shape und generally the tubercle does extend dawn into it. The neural ganglion is especially long, extending most of the dis- tance atone the dorsal lamina, The dorsal Juminu has a dowble row of Languets, These are fine anil pointed. closely set on the left. und on the right they are stouter and more sparsely urrunged. ‘There are from & to 10 branchial folds un each side of the body with ahout 12 longi- tudinal vessels on the folds and 2 between. There are 6 to 8 stigmata in euch mesh crossed by parastigmatic vessels, The gut loop is simple and curved and encloses the left gonad which is subdivided into 15 1 20 separate polvearp sacs arranged on either side of central male and female ducts, There is a corres- ponding gonad on the right. Remarks: This species resembles very closely the Antarctic species Pyura discoveryi Herd- man (see Kott 1969). ‘Che tough, wrinkled external test with embedded polygonal thicken- ings is alsa reminiscent of the Antarctic Pyare squsniata Herdman although the polygonal ASCIDIANS OF SOUTH AUSTRALIA 3y seules und the body shape of P. squameta are more highly specialised than in either P. ass. covers? or in the present species The bran- chijil tentacles with their sparse branches und the long siphons ure alsa similar to those of P. discoveryi and it is possible that the protec- lion afforded the individual by these Jone siphons may be assncigied with the absence of the more bushy tentacles usually found m this uchus. The individuals are never very farve and their leathery test and hahit of occurring in ageregates suggests a species udapled Jor yery turbulent conditions. Che present records do not support this, however as they are vither fram Offshore Benthic locations in St. Vincent Gulf. ar from reefs in sheltered coastal loca- ions. Pyora australis (Quoy & Gaimard) 5.sp. ausira- fis Quoy & Gaimard. Aseidia vastralix Quoy & Guimard, 1234: 614, Priva australis €. rypieq, Kou, 1952: 268 and synonymy, Pivre australis, Millar, 963° 739. New Records: Vipata Reef, Tapley. Shoal. near Murion Light, off West Beach, off Rroadway. off Hallelt Cove, off Yankalilla Baw West LL, N.W. of Robe. Previous Records: W. Aust. (Geraldton to Albany) —Quoy & Gaimard 1834; Michaelsen & Hartmeyer 1928: Kou 1952; Millar 1963. Vic, (Weslermport, Flioders}—Quuy & Gatmard $8342 Koi 1952. Tus. (D’Entre- casteaux Channel, Tinderbox)—Kott 1952. FIG. 61 PDescripiion: Specimens of all sizes up tu a maximum of 4 cm long head with a stalk of 30 cm. ‘The lest is usually without foreign bodies adhering, though in one specimen there are some cirtipedes growing on the stalk. The surface of the test is marked with variable Jongitudinal furrows pnd ridges but is some- times almost smooth. In preservative the specimens are pinkish-fawn, although living specimens. ure usually dark red and, Occa- sidually, yellow. Both apertures are close together on the dorsal surface, the atrial uper- ture directed upwards and the branchial aper- ture directed bisully.. The Jobes of the atrial uperture wre clearly continuous with the ridges in the dorsal part of the test There are stellate spicules of about 0,02 mm diameter with 6 rays i optical transverse sec- tion in the body wall, and the siphons are lined by conical spines of U.02 mm maximum height from buse to apex, The bronchial sac, gut loop and gonad ure as previously described and there arc 18 long fisttened characteristic lobes fringing the anul border (see Kott 1952). Remarks: Nothing can be added to previous descriptions of this constant species which appears. to occupy a wide Tange of conddirions in exposed to sheltercd locations from Geralu- ton, in Western Australia, to Flinders in Vic- loria. It is common in wave besten areas from the low water mark to 22 nv. Pyura spinifera (Quoy & Gajimard). Kot, 1952, 269 and synonymy. Aridio spintfera Quoy & Gaimard, 1834; 6) 7_ Cyethie multiradicata Herdman, 18¥9: 30. Now Records: Upper St, Vincent Guif, off Hallet. Cove, Aldinga. Previous Records: Wo Aust. (Albany)—Quoy & Gaimard 1834. Vie. (Buss Strait)—Michi#elsen 1905; Heller 1878. NS.W, {Port Jnckson. Port Hacking)—Werdman 1891. 1899; Kott 1952. FIGS. 62, 63 Description. Specimens with head to 8 cm tong am 4.5 em wide. Stalk is of very variable length. maximum 20cm. Externally the test is smooth without longitudinal burrows. but with characteristic tubercles, varying in their density, and sometimes. especially in Jarger specimens, absent ultogether, The head is often completely enveloped by an investing sponge which in specimens from olf Hallett Cove hus been noted in the field as yellow. Minute overlapping scales, 0.05 mm tmani- mum length from posterior patt of the base to their apex. line localised areas where thick- ened lobes of the test project into the siphons. Otherwise. there ure no spicules in the test or in the hody wall, There sre 7 branchinl folds on cither side of the body wall in the larger specimen but only 6 on cach side in averuge- sized to smaller specimens. There are abou 25 branchial tentacles alternating with rudi- mentary tentacles. The larger tentacles have regular pinnate primary branches with seoon- dary branches and minute tertiary branches and are very bushy. The dorsal tubercle has a double coiled opening. both horns coiled inwards anc the inner spirals of cach coil are slightly convoluted. There is & 9 Eudistoma renieri - \ op tee ae WG Halocynthia hispida ar, _ Al Herdmania momus - ; 41 Leptoctinides kingi ere saeli> adalat LY Leptoclinides reticulatus _. 18 Orr YANKALILLA Bay, St, Vincent Gulf: in Amphiholiy community. sandy bottom: slight surge; depth as indicated: 18.11.67. Pyuraanustraliy 20m Halocynthia hispida 20m Crenicella antipoda 15m Page Leptaclinides rufus ; tie tam esp Microcosmus helleri 43 Microcosmus nichollsi 42 Microcosmus squamiger - - = 43 Microcosmus stolonifera = ef “43 Molgula mollis ; ; 45 Oculinaria australis 29 Phallusia depressiuscula ., 2... 4... 23 Podoclavella cylindrica .- 1. te Podoclavella moluccensis : 5 Polycarpa clavata . ee . 33 Polycurpa papillata .. t Sie som 4 Polycarpa pedunculata —- _f.. Pegs Polycilor giganteum tenet nie she Polysyneraton orbiculum - = 21 Pseudodisioma cereum =... 4, 4.) «612 Pyura australis = - et «se Pyura irregularis ee ee a nee Pyura scoresbicnsis 1.sp. a 36 Pyura spinifera : , 39 Pyura vittaia |) 4. -. + Rhodosoma turcicum . 23 Ritterella herdmania — ee. Pk Stolonica australis _. He tte Stolonica carnosa a 28 Sycozoa cerebriformis Sycozoa tenuicaulis... ny Synoicium papilliferum . ; oe ate | 16 Trididemnum spiculatum ih tte ee DG NEW FORM SPECIES OF POLLEN FROM SOUTHERN AUSTRALIAN EARLY TERTIARY SEDIMENTS BY WAYNE K. HARRIS Summary Sixteen new form species of dispersed pollen grains; Sparganiaceaepollenites barungensis, Amosopollis dilwynesis, “Triorites” psilatus, Tricolporites valvatus, Triporopollenites gemmatus, Ericipites crassiexinus, Sapotaceoidaepollenites rotundus, Proteacidites confragosus, P. tripartitus, P. kopiensis, P. tortuosus, P. clintonesis, P. fromensis, P. varius, P. wilkatanaensis, and P. concretus and one new form genus, Gambierina, are described from early Tertiary sediments from southern Australia. NEW FORM SPECHFES OF POLLEN FROM SGUTHERN AUSTRALIAN EARLY TERTIARY SEDIMENTS by Wayne K. Harrts* Summary Sixteen new form species of dispersed pollen grains: Sparganiaceacpallenites hearungensin, i a Anwxopollis dilwynensis, “'Triorites" psilatus, Priculporites valvatus, Triperapollenitey cemmatus, Eri- cipites crassiexinus, Sapotacevidacpollenites kopiensis, P. tartuasus, P. elintonensis, P. fromensis, P. varius, P. wilkarandensis, rotundius, Proteacidites. confragasus, P. lripartitus, Py and #.. comerepirys and one new form genus, Garmhierina, are described from early Tertiary sediments from southern Australia, Introduction This paper describes several new species and anew genus, Gambrerina, that were mentioned 4S Manuscript names by Harris (1971) in an account of the palynology of Tertiary sedi- menis in the Otway Basin. These forms were considered to have some biostratigraphic sig- nificance. Previous taxonomic studies of Tertiary apgiosperm pollen from Australia are limited ta the works of Coakson (1947, 1950, 1953, 1954, 1957 & 1959), Cookson & Pike (1954) and Harris (196Sa), Dettmann & Playford (1968) described four new angiosperm pollen species from Upper Cretaceous sediments from eastern Australia and some of these probably extend into the carly Tertiary. The preparation technique is that outlined by Harris (1965a) und the descriptive termino- logy is largely adapted from Erdtman's glossary (1952). Dimensions are based on fifteen or more specimens. Biostratigraphic data are ‘based in part on unpublished studies by the author and on Harris (1971) and McGowran, Lindsay & Harris (1971). Sample data are presented jn the appendix. All co-ordinates are from the Leitz Ortho- plan (715494) microscope in the Palynology Laboratory of the Geological Survey of South Australia and Holotypes (catglogue numbers prefixed Py) are deposited in the Geological Survey Palaeontological collection, Systematic Palynology Genus SPARGANIACEAEPOLLENITES Thiergart 1937 Type species: Sparganiaceaepollenites polyga- nealis Thiergart 1937: 307. Sparganiaceaepollenites barungensis Sp. tov. FIGS. 1-3 Pollen monoporate. sphucrojdal to slightly bilateral. Pore circular (3-4 pin diam.) with incrassate margin 1-1.5 ym wide. Exine 2 ~m thick, sexine as thick as nexine, reticulate. Reticulum undifferentiated over the grain, lumina 1-1.5 pm diam, Dimensions: Equa- torial diam. {8 (22) 25 ym. Holotype: Preparation and slide number— $T325/15, 42.1: 110.8. Py 195. Figs. 1, 2 Type localify; Hd. Bartinga Bore 4 at 65.5 m. Clinton Formation, 7Lower Miocene. Distribution: This species first occurs in the Upper Eocene and continues through to the Upper Tertiary. Comparison and affinity: The pollen figured by Couper (1960, pl. 9, figs. 21, 22) as Typha 8p. appears to be very similar to §. barunvensis. S. harungensiy differs from Aglaoreidia Erdt- man in not having a differentiated reticulum. “Monoporitey” subreticulata Cookson has a wider rim to the pore, The specics here des- eribed appeared under this generic name in Harris (1971) and McGowran, Lindsay & Harris (1971). S: magnoides Krutzsch (1970) ST a ee * Geological Survey of South Australia, 169 Rundle Street, Adelaide, S.Aust. 5000 Published with the permission of ihe Acting Director of Mines, Trans. R. Soc. S, Aust, 96, Part 1, 29 February, 1972, 54 WAYNE K. HARRIS approaches S. barungensis in size but hax a wider meshed reticulum. S. batiurgensiv 1s very similar to pollen of Typha and. Sparge- niu, Genus AMOSOPOLLIS Cookson & Balme 1962 Type species: Amoxepollix cruciformis Cook- son & Bulme 1962: 97, Amosopollis dilwynensis sp, noy. FIGS. 4. 5 Synonymy. Amosopollis erucifarmiis sensu Harris 19650: 97, Ph 29, fig, 26, Pollen grains in rhomboidal tetrads. Indi- vidual grains prolate to sub-prolate, [xine 2 am thick, psilate to seabrate and finely granu- Jale, except near the margins of the aperture where grana 1=1,5 jm diam. are present. Aper- ture ix a long gaping sulcus extending the full length of the grain, Margins of sulcus not ragged, Dimensions: (10 specinrens) Overall diam, of tetrad 50 (60) 68 pm. Individual arains 22 (34) 40 ,.m diam, Noloiype: Preparation and slide number— §T209/ 2, 39.3; 100.7. Py O15. Figs. 4. 5, Type locality: Ditwyn Bay, Victorian, Pebble Point Formation, Paleocene. Distribution; A. dilwynensis is a rare specics hut has been observed in Paleocene sedi- ments fram the Murray and Otway Basins, and a Similar forny hus been reported (Harris 1965h) from Queensland in sediments of similar age. Comparixon and affinity: A. dilwynensis is in general Jarger than the genotype but can also be distinguished by the psilate-scabrate sculp- tre und more importantly by the straight mar- fins of the sulcus. Genus TRIORITES Cookson ex Couper 1953 Type species (hy subsequent designation of Couper 1953, p. 60): Triorites magnificus Cookson 1950, ‘“Triorites” psilatus sp. nov, FIGS. 6, 7 Pollen radiosymmetric, isopalar, oblate, triorate. Amb sub-triangular. sides straight to slightly convex. Ora sunken, 2-4 pm wide, circular Exine 2m except around apertures where it thickens to 3 or 4 pm. Enxine psilale- Dimensians: Equatorial diam, 24 (32) 40 pm. Holotype; Preparation and slide number— 5964/1. 32.8: 100.6. Py 411. Fig. 6. Type locality: Polda No. | Bore at 55.1 m. Poelpena Formation, Middle Bocene. Distribution: ‘Vhis species is a very common form throughout the Lower Tertiary in southern Australia. ft first appears in the Princetown Member and ranges through to ihe Lower Miocene. The upper limil has nor been determined, Camparivon and affinity. “T" psilaris is com- parable and may be conspecific with "7." yewbratas Couper, "The ornament on the latter however is scubyate. This species would more appropriately be placed in a new genus (see section on Trieritey below). Genus VRICOLPORITES Cookson 1947 Type species: Tricolporites sphacrica Cookson 1947; 195; genus monotypic when proposed, Tricolporites valyamms sp. nov, FIGS, &, 9 Pollen radiosymmetric, prolate tricolpurpie Amb in equatorial view ellipsoidal. Apertures compound, colpi reaching to within 3 or 4 jn) of the poles, margins strongly invaginated to about § pm. Equatorial aperture orate, 5-8 pm diam, Kxine 2-3 am thick unornamented. Dimensiony: Pole diam. 45 (52) 55 jum, eque- torial diam, 30 (35) 39am. Holoiype: Preparation and slide nurmber— ST241/12, 35.1; 98.1. Py 176, Fig. 9, Type docality: Lake Torrens Bore 3A at 247.8 of “Wilkatana Formation”, Middle Focene. Distribution: Common in the “Wilkatana Formation" but less common in other Middle Kocene (Proleacidites confragosus Zonule) assemblages, Camparison und affinity: The strongly. invagi- nated colpi and psilate exine make this a very distinctive species, Its natural affinilies are un- known, Genus ERICIPITES Wodehouse 1933 Type species: Fricipites longisitlearis Wode- house 1933: 517. Ericipites crassiexinus sp. nov. FIGS. 15, 16 Pollen united in tetrads. Individual grains indistinctly tricolporate, tetrahedral in shape FARLY and strongly united in the tetrad. Exine 2.5- 3.5 ,m thick, sexine as thick as nexine psilate. Aperttires complex, colpi about 14 pm long and 1.5 pm wide, Pores indistinct and difficult ta detect, 2 wm diam, Dimensions: Overall diam, 35 (42) 33 am. Individual grains 24 (3 35 pon diam Holarype, Preparation and slide mumber—- S660/ 1, 52.9: 96.9. Py 415, Fig, 15. Type locality: Bore, Hd, Cummins at 35,7- 43.30 om. Wanilly Formation, Middle Eocene. Distribution: Often wu yery coramon. form in middle and upper Eocene sediments. Compariven and affinity: The psilate nature of the exine and the larger size of this species cis- tinguishes it from A. scabrases Haris. Pollen of this type characterise the Order Ericales. Genus TRIPOROPOLLENITES (Pflug) Thomsen & Pfluz 1953 Type species! = Triperopollenites Pflug in Thoms. & Pf, 1953: 84. Triporopollenites gemmatus sp. nov. FIGS, [0, 11, 13, 14 Pollen occasionally free but most commonly united in tetrads. Tetrads 34-40 2m in overall diam. Individual = pollen = radiosymmetric, oblate, sub-isopolar, triorate. Amb sub-trian- gular with straight to convex sides. xine 4—5 am thick (including ormament), Sexine and nexine difficult to separate but nexine appears to be thicker than ‘sexine. Exine covered with verrucae 2-3 «wm wide, sphacrical and 2 ym high. Verrucue separated from each other (by 2-3 am) by granulate ornament. Apermres obscured by ornament, porate or orate open- ing 1.5-2.5 pm wide, Dimensions; Individual pollen, equatorial diam, 25 (29) 31 jam. Holotype; Preparation and slide number— $547/1, 31.7: 98.4, Py 720. Fig. 11. Type locality: Lake Cootabarlow Bore 2 at 163.4 m. Great Artesian Basin. Murn- peowie Formation, Upper Eocene. Disrriburion:; Appears to be testricied to Middle and middle-upper Eocene sediments from the Pirie-Torrens and Great Artesian Basins and Eyre Peninsula, Comparison and affinity; T. 2zemmatus is simi- lar to 7. bullis Gruas-Cavagnetto (1966) from the Sparnucian of the Paris Basin but this coryloides JERTIARY POLLEN SPECIES 55 species is more of less circular and appears to have a more strongly thickened sim to the aperture: CGienus GAMBIERINA gen, nov. Type species: Triorites edwardsit Cooksoly & Pike (in part) 1954: 214, pl. 2, figs, TOT, 105, 106, Diagnosis; Pollen raciosymmetric, oblate, lohate. angulaperturate, triorate. Apertures sunken. Sexine imperforate tectate, thinner than nexine, the two separated by a faintly dis- cernable baculate layer, which forms a “nick” point in the apertural region, Aperture fortyed by sexine larger than that of the nexine. Nexine thickens more rapidly than sexine about the apertures. Exine psilate. Figured specimen: Fig. 12, Remarks: The characters of the exine, the apertures and general shape distinguish this venus from Triorites. As Dettmann & Play- ford (1968, p. 86) have pointed out, the species figured by Cookson & Pike (1954, par- ticularly Ags, 104 and 105) as 7, edwardsii is distinct in being unthickened about the aper- tures, Dettmann & Playford (1968) summarised the present status of ahe genus Triarites but chose to continue using the diagnosis of Couper (1953) pending a review by the present author. Potonié (1960) clearly indicated that the two species T. mugniftcuy Cookson and T. efavatus Cookson were morphologically com- parable and distinct from other torms allocated to the genus. However, Potonié gave no indi- cation as ta where these other forms should be placed. It ig clear that T. magnificus and T. clavatuy are very closely related morphologically and perhaps phylogenetically, Indeed Cookson (1957, p. 49) voes so far as to state thar “there is little or no doubt that they were produced by closely related plants. Both species have the same shape, type of ora and exine stratifi- cation, and structure...“ Thus these two species form a natural grouping and all other species assigned to the genus ate better acco- modated elsewhere. Couper’s (1953) diagnosis is too broad and suggestive of a suprageneric category. Mildenhal) & Harris (1971) have reached similiar conclusions. Genus SAPOTACEOIDAEPOLLENITES Pot. Thoms, & Thierg. 1950 Type species: Saparaceoidacpollenites. (al. Pal- lemlex} vratttfesnes (Potonié) 1931: 3. Sapataceoidaepollenites rutundus sp. nay. FIGS. 17, 18 Pollen radiosymmetric, subsphaeroidul to sub-prolate, four and less frequently three apertures. Apertures compound. Colpi 2/3 length of polar axis, 2-3 ym wide. Equatorial aperture more or less circular, S=6 pm diam. and slightly elongate 0 an equatorial direction, Apertural margin prominently rimmed ancl thickened, xine 2-2.5 pm_ thick, nexine ubout as thick as sexing, Sesine psilate to finely scabrate. Dimensions: Polar diam, 30 (36) 39 am, equatorial diam. 26 (33) 35 pm, Holotype: Preparation and slide number ST241/3, 45.7: 102.8. Py 167. Fig. (7. Type locality: Lake Torrens Bore 3A ut 247.8 om. “Wilkatuna Formation, Middle Focene. Distribution: The species Orst appears in the. Middle Hocene and continues on Into the mid-Tertiary. Comparison and affinity: The closest resem- blance of this species is with Tricalperopollen- ites latizonatuy MeIntyre 1968, which is most commonly 3-aperturale, has a longer polar/ equatorial axis ratio and has a broad thickened zone of the exine in the equutorial region, The species ts very similar to pollen of the Sapo- taceae, Genus PROTEACIDITES Cookson ex Couper 1953 Type species: Proteacidites adenamhotdes Cookson (950: 172, designated by Couper 1953; 42, Remarks, The genus Proteaciditey wecomodales ul present. a wide variety of forms described fram both the Southern and Northern Hemis- pheres. Some from the latter clearly do not belong in this genus but until a review of the Australian forms by the author is complete (and on present evidence the genus will be split into three or more genera) comment on these is reserved. Although the following new spe- cics show a wide variation in form with regard to aperture construction and exine stratification and ornamentation they ‘will be described under this genus but will be further reviewed in a forthcoming paper. 56 WAYNE K. ITARRIS Proteaciites coufragosus sp. nov. FIGS, 19-22 Pollen sub-isopolur anguluperturate, oblate. tiporate. Arub triangular with slightly conves sides. Pores simple subcireular 6-7 pm diam.. obscure. Exine 4-5 pm thick, Sexine three limes us thick as nexine, heavily ornamented with a dense reticulum, lumina 3-4 pm diam.- polyvonal wind made up of single rows of dis- tinct bucula 1-15 um diam. Dimenstons: Eyuatorial dian, 34 (60) 69 pm, Halorype: Preparation and slide number— Sr24t/9, 41.4: 105.7, Py 173, Figs. 19, 21, 22. Type locality: Lake Torrens Bore 3A. aol 2478 m “Wilkaturm Pormation’. Middle Eocene. Distribution: An index form for Middle Eocene sediments, PF, confrugosus has been recorded from the North Maslin Sands, the Renmark, Poelpenu and Wanilla Formations and the Burrungule Member of the Knight Formation, Comparivan and affinity: This is 4 striking spe- cies and is ¢learly distinct from any other known in the genus. Proteacidites tripartitus sp. nov, FIGGS. 23 25 Pollen sub-isopolar, oblate, angulaperturate, triporate, Amb triungular with more or less straight sides, Apertures sub-circular, simple but obscure 2-2.5 ym wide. Exine 2.5-3 pm thick, Sexine half as thick as nexine, fove- alate, Lumina ca. 1 am diam., slightly smaller at the poles and towards the apertives. Muri 2-3 ym wide. Nexine thickens. to 5 ym at 10 pm from the apertures. Pore “canal” 7-8 yam long. Déinensions: Equatorial diam, 27 (30) 34 pitt, Holoi\pe: Preparation and slide nuniber — $650/1, 32.2: 99.8. Py 406. Pigs. 24, 25 Type locality’ Hd, Cummins Bore at 114- 116.4 im. Wanilla Formation, Middle Eocene, Disiribution: The species first appears very high in the Princetown Member of the Dil- wyn Formation but does not became com- mon unfil the Miidle Eocene. Compurixon and affiniry: The detail of the apertures closely resembles that found in P. latrohensts Harris and P. concretus but is dis: linguished bY the characteristic ornament, EARLY TERTIARY Proteacidites kopiemsis sp, nov, FIGS. 26, 27 Pollen sub-sopolar, oblate. angulaperturate, Iriporate, Amb triangular, sides straight or nearly so. Apertures subcircular. simpic. 7-8 pity diam. Exine 2m thick and slightly thicker in the equatorial inter-aperturate regions. Sexinc about half as thick as nexine, orma- mented with a reticulum, Muri 1-15 pm wide, Lumitta 2-3 am diam, at the equator and decreasing gradually to 1 um towards the apertures and polar regions. Dwnenstons: Equatorial diam, 36 (40) 47 ym, Hololype: Preparation and slide number~- S560/1, 26.1: 106.9. Py 393. Fig, 26, Fiipe tocality: Pala No. | Bore at 37,5 m, Poelpena Formation. Middle Eocene. Distribution: The species is present in the uppermost section of the Princetown Mem- ber of the Dilwyn Formation and continues in all basins into the middle-upper Eocene. IL does not appear to range higher than the Triorites magnificuy Zonule, Comparison and affinity: This species is teaditv Wistinguished from other Proteacidires spp. by the characteristic ornament pattern. Proteacidites tortuosus sp, nov, FIGS, 28. 29 Pollen sub-tsopotar. oblate, anguluperturate. triporate. Amb rounded triangular, sides con- vex. Pores simple 13-15 »zm wide. Exine 5 pm thick. Nexine thicker than sexine. Sexine ornamented with scattered verrucae, 2 «um wide and up to 6 xm long, rounded in optical section and 2 ym high. Areas between these elements psilute. Dimensions: Equatorial diam. 53 (55) 58 pm Holorype: Preparation and slide number— $563/2, 32,2; 99.8, Py 409. Figs, 28, 29, Type locality: Polda No, | Bore at 55.1 mm. Poelpena Formation, Middle Eacene, Distritiaion: This species has heen recorded from Middle Eocene sediments on Eyre Peninstila, Poelpena and Wanilla Porma- tions. Comparison and affinity: The large distinctive verrucae, thick exine and rounded sub-triangu- lar shape separate this species from other spe- cies described here. P, tortuasus differs fram P. nibereudatuy Cookson in being smaller. The vermicae are not arranged in u reticuloid pat- tern and are not confined to a sphencal shape POLLEN SPECIES 57 Proteacitites clintonensis sp. nov. FIGS. 30-34 Pollen sub-isopolar, oblate angulaperiurate. triporate, Awb more or less triangular with eoncave sides, Pores circular 2435S ym in diam, Exine 3 jm thick, sexine slightly thinner thin nexine. Capita of bacula coalesce to form groups Up to 7am wide and shmw an LO pattern, Elements rounded in optical section, Nexine in region of pores, alternately thick and thin, The sexine is readily lost by corrosion, Dimensions: Equatorial diam. 62 (75) 98 yum. Hoelompe; Preparation and slide aumber— $705/1, 31,3: 105.2. Py 405. Figs, 32-34, Type fecaliry: Poyniz Bore, Hel. Ettrick at 94.5 om. Renmark Beds, middle-upper Eocene. Distribution: The species is almost ubiqui- tous in Eocene sediments and is particularly common in the Priarites staynificuy Zonule, {| ranges from Middle Eocene to at feast Lower Miocene, Comparison and ajfinity: This species is similar to FP. rectomerginis Cookson but has much larger apertures and strongly concave sides. Figure 30 more closely resembles P. reeto- mmarginus with its finer ornament, larger size and straighter sides. It is possible that the two forms intergrade, Cookson’s figure (1950. fig. 27) of P. rectomarginuy appears ta show some thickening of the nexine about the apertures. The species is distinguished from. P. jncurvelius by the nature of the sculpture and the charac- teristic aperture. The exine dacs not rhin markedly near the apertures as it does in P, irearvatus, Proteacidites fromensis sp, nov. FIGS. 35-38 Pollen sub-tsopolar, oblate, angulaperturate. triporate, Amb triangular, sides strongly con- cave, Pores simple, circular, 5 ym in diam. Exine 2,3-3 »m thick. Sexine slightly thinner than nexine in the inter-angles and thins to- wards the angles, evenly granulate to scahrate. Nexine thickest im the inter-angles, Désrten- sions: Equatorial diam. 61 (65) 70 um, flolotype: Preparation and slide number S17/2. 35.1: 1014, Py 408. Figs, 35-37. Type locality; Lake Eyre Bore 20.at 73.2 m. Murnpeowie Formation, Paleacene. Distribution: P. fremensis is restricted to and characteristic of Palacocene sediments and is most cormmmon in the Murray and Great Artesian Basins, 38 WAYNE K. HARRIS Comparison and affinity: The strongly concave sides of the amb, size, and the scabrate orna- ment separate this species. fram others in the genus, It differs from P. granoraius Couper ive that the ornament does not become coarser around the apertural region. Proteacidites varius sp. nov. FIGS. 39-42 Pollen small, sub-isopolar, peroblale, angu- japerturate. Amb triangular with straight or slightly Concave sides. Apertura] pores, three, 2.5 »m diam. Exine 2-2.5 pm thick. Sexine much thinner than nexine and thins markedly near the apertures, Necxine thins toward the apertures with loss of Jendonexine elements near ihe wperture. Ornament 0.5 pm high, consisting of fused groups of bacula ca. 1 pm djam, Groups becoming smaller to absent near apettures. Dimensivns: Equatorial diam. 20 (25) 37 am, Holaryye: Preparation and slide number— §705/1, 37.2: 111.0. Py 399, Figs. 39, 40. Type locality: Poyntz Bore, Hd. Ettrick ut 94.9 m. Renmark Beds, Upper Eocene. Distribution: A common species in middle and upper Eocene assemblages in the Murn- peowie and Poelpena Formations in parti- cular, Comparison and affiniry; Vhe nature of the ornament distinguishes the species from P. reliculainsy Cookson and P. xymphyonemoides Cookson, The churucteristic nexine structure (see particularly Fig, 41) is distinctive, The relitionship of this species. lo P. obscurus Cookson is not clear. Her figured specimens (Cookson 1952, figs. 30, 31) have lost most of the sexine, The specics described here shows thinning of the exine about the apertures rather than slight thickening and is not “lanect- shaped” as described for P. ebscurus. Proteacidites wilkatanaensis sp, nov. FIGS, 43-47 Pollen sub-isopolar, oblate, angulaperturate, triporale. Amb triangular with straight lo con- cave sides slightly bulging about [0 ~m from wpertures. Apertures circular 4-6 pnt diam, Exine 4 um thick, thinning rapidly near the Apertures. Nexine about 3 times as thick as sexine, thinning and apparently losing the basal layer near the apertutes, Sexine consists of a thin baculate layer and an ectosexinous layer formed of united baculate elements. giving a fuw rugulate ornament. Rugulae L-3 ,«m long, lesa than 1 pm wide. Dimensions: Equa- torial diam. 51 (55) 67 2m. Holotype. Preparation and slide number— $2273/'4, 44.1: 106.7. Py 720. Figs, 43. 44. Type locality: Bore near Ediacara at 280,4— 283.5 m, “Wilkatana” Formation, middle- uppet Eocene. Pistribuion: The species is commonly ob- served in miiddle-upper Eocene sediments in most basins. Comparison and affinity: The species differs from P. ineurvans Cookson and P. clintonensis in not being puhcti-tegillate. The ornament is similar to that of P, varius bul the species is much larger and does not show the charac- teristic structure of the nexine wround the aper- tures, Proteacidites concretus sp. nov. FIGS, 48, 49 Pollen sub-isopolar. oblate, ungulaperturate, Iripotute. Amb triangular with straight sides, Apertures circular 1.5 ,m diam, Exine 2 um thick but thickens in the region of the aperture to.4 am and forms a pore “canal” 5 pm long. Exine faintly and evenly scabrate to finely spinulate. LO pattern distinct. Dimensions: Equatorial diam. 25 (28} 32 ym. Holatype: Preparation and slide number— $360/2, 35.8: 105.5. Py 404. Fig. 48. Type locality: Kopi Anomaly KR9 at GT om. Peolpena Formation, Middle Eocene. Distribution: A common species in most Locenge sediments, Comparison und affinity: This. species is most closely similar to P, farrobensis Harris, parti- colacly in the nature of the aperture. It differs, however, from this. species by the nature of the ornament. (P. farrabensis has a scrobiculate pattern.) EARLY TERTIARY POLLEN SPECIES 59 References Baime. B. E. ((962).—Amuosopollis crucifarmis gen. el sp. noy., a pollen tetrad from the Cre- taceous of Western Australia. JR. Soc. West. Aust. 45, 97-99, CooKson, [2 C. (1947).—On fossil leaves {Oleaceae) and a new type of fossil pollen grain from Australian brown coal deposits. Prac, Lint, Soc. N.S.W. 72, 183-197. Cooxson, I. C. (1950).—Fossil pollen grains of proteaceous type from ‘Tertiary deposits in Australia. Aust, J, Scient. Res., B. 3, 166-176. Cookson, 1. C. (1953).—Difference in microspore composition of some samples from 8 bore at Comyum, South Australia, dusr. J, Bor, 1, 462-472. Coonseon, I. C. (1954).—The Cainozoic occur- renee of Acacia in Australia, Aust. J. Bot. 2, 52-59, Cookson, LC, (1957),—On some Australian Ter- iiary spores and pollen grains that extend the geological range and geographical distribution of living genera. Prov. R. Soc, Vict, 69, 41-53- Cookson, I, C, (1959);—Fossil pollen grains of Nothofagus from Australia. Proc. R. Aac. Viet. 71, 25-30. Cookson, I. C.. & Pike, K, M. (1954).—Some dicotyledonous pollen types fram Cainozoic deposits in the Australian region, Ausr. J, Bot. 2, 197-219, Coueer, R. A. (1953)—Upper Mesozoic and Cainozeic spore und pollen grains from New eetand. N.Z. geol, Surv. paleoni, Bull, 22, Covuprr, R. A, (1960).—New Zealand Mesozoic and Cainozoic plant microfossils. N.Z peel. Surv. paleont, Bull, 32, 1-87. DrtrMann, M. £.. & PLAayrorp, G. (1968).— Taxonomy of some Cretaceous spores and pollen grams from eastern Australia. Proc, R. Soc. Viet. 81, 69-94, EroTMAN, G. (1952)—"“Pollen morphology and plant taxonomy: Angiosperms.” (Almavist & Wiksell: Stackholm, } GRUAS-CAVAGNETTO, = C, {(1966).— Complexes sporopolliniques du Sparnacien du phare @AUly (Quest de Dieppe, Seine-Maritime). Rev. Micrapalévntol. 9, 47-67. Harris. W. K. (1965a).—Basal Tertiary micro- floras from the Princetown area, Victoria. Australia, Palaeontographica B. 115. 75-106. Harris, W. K. (19656).—Tertiary niucrofioras from Brisbane, Queensland. Geol. serv. Queensland. Rept. 10, t-7. Harws, W. K. (1971).—Tertiary stratigraphic palynology, Otway Basin. Ch..4 in ~The Otway Basin of south cast Australia.” Spec, Bull. Geol. Survs, S. Aust, & Vici, 67-87. KrutzscH, W, (1970).—“Atlas der mittel—-und jungieriaren dispersen Sporen—und Pollen sowie der Mikroplankionformen des nordlichen Mitteleuropas. Lieferung VII,” (Veb Gustay Fischer Verlag: Jena.) McGowran, B., Linpsay, J. M., & TIArnis, W. K. (1971), -Attempted reconciliation of ‘Tertiary biostratigraphic systems, Otway Basin. Ch. 14 in “The Otway Basin of south east Australia.” Spec. Bill. Geol. Survs, 8, Aust, & Vict, 273- 21. McIntyre, DB. J, (1968)—Further new pollen species from New Zealand Tertiary and Bppermass Cretaceous deposits, N.Z, J/ Bot. 6. 177-204. Mitoengari, D. C., & Harris, W, F. (1971).— Status of Jlaloragacidites (al, Trivrites) harrisii (Couper) Harris comb. nov, and Haloragacidites triorauts Couper, 1953. N.Z. Jl Bot. 9, 297-306. Potonte, R. (1931 ),—Pollenformen aus tertiaren Braunkohlen, 3. Jb. preuss. geol. Landesanst. RergAkad, 52, 1-7. Potontz, R. (1960)—Synopsis der Gattungen der Sper dispersse, UI Teil. Beth geal. th. 39, 1- . Potons:, R.; THomson. P. W., & Tinercart. F. (1950).—Zur Nomenklatur und Klassifiky- tion der néogenen Sporomorphae (Pollen ond Sporen). Geol. Jb, 65. 35-70. THIERGART, F. (1937).—Die Pollenfora der Niedetlausitzer Braunkohle, besondets im Profil der Grube Marga bei Senftenberg. Jh- Preis. geol. Landesanst. BereA kad. 58, 222- S56. Tuomson, P. W., & Prruc, H. (1953).—Pollen und sporen des mittelcuropiischen Tertiirs, Palacontographica B, 94, 1-138. Wooexouse, R. P. (1933),—Tertiary pollen JI. The oil shales of the Eocene Green River ternation, Bull, Torrey Bot. Club, 60. 479- 524, 60 WAYNE K. HARRIS APPENDIX SAMPLE DATA Depth in metres Type of Sample Bore name (or outcrop) (feet in parenthesis) Formation Basin Lovality Sample No, Nullabor No. 6 105-122 (344-400) Pidinya Eucla Lat. 31°09'00"S Percussion $623 Long. 131°12’30"E sludge Polda No. | 37.5 (123) Poelpena Polda Lat. 33°33’00"S Core SSal? Long. 135°20’00”E Polda No, t 51.8 (170) Poelpena Polda Lat. 33°33/00"S Core $562 Long, 135°20’00”E Polda No. | 55.1 (ABU) Poelpena Polda Lat. 33°33/00"5S Core 5863 Long. 135°20'00 TE Polda No. | 57.9 190) Poelpena Poida Lat. 33°33/00"S Core S564 Lone. 135°20’00” E Kopi Anomaly K.R.9 61 (200) Poclpena Unnamed Lat. 33°24'10"S Care S360 Long, 135°44’45"E Hd. Cummins (W.Con.Res. 35,7-43.3 (117-142) Woanilla Cummins Lat. 34°15710"S Percussion $660 adj. Sec. 16) Long. 135°40’45"E sludge Hd, Cummins (W,Con.Res. 114-116,.4 (374-382) Wanilla Cummins Lat. 34°15°10"S Percussion 5650 adj, Sec. 14) Long. 135°40°45”"E sludge Cummins school residence 32-39 (115-128) Wanilla Cummins Lat. 34°15/50"S Percussion S74! Long. 135°43'20"E sludge Lake Torrens Bore 3A 247.8 (813) “Wilkatana” Piric- Lat. 31°14’00"S Core S241 Torrens Long. 138°01’45’"E Near Ediacara 280,4-283,5 (920-930) ‘'Wilkatana’™’ Piric- Lat. 30°48’34"S Cuttings $2273 Torrens Long. 138°07/30"E Hd. Barunga Bore 4 65.5 (215) Clinton St, Vincent Lat. 33°45'°55""'S Core $325 Long. 138°13'35”"E Lake Cootabarlow Bore 2 163.4 (536) Murneowic Great Lat. 30°16730"S Care 8547 Artesian Long. 140°08'30” E E. A. Rudd Bore § 116.1 (381) Murneowic Great Lat. 31°13’00"S Core $1986 Artesian Long. 139°52/50"E Lake Eyre Bore 20 73,2 (240) Murneowic Great Lat, 28°48’00"S Core SIT Artesian Long. 137°30’20”E Payntz Bore, Hd. Ettrick 94.5 G10) Renmark Beds Murray Lat. 35°O00°307"S Percussion $705 Long, 139°31’45”E sludge S.E. side of Dilwyn Bay 1.8 m ahove buse Pebble Point Otway Lat. 38°44/00”S Gutcrop S208 of formation Long. 143°10°30"E SE. side of Dilwyn Bay 1.2 m above base Pebble Point Otway Lai, 38°44/00"S Onterop S209 of formation Long. 143° 10730" E EARLY TERTIARY POLLEN SPECIES 61 Note: Unless otherwise specified the figures are X500 in normal transmitted light. NDIC refers to Nomarski Differential Interference Contrast. Figs. Figs. Figs. Figs Figs. Fig. Figs. Figs. Figs. Figs. Figs. Figs. Figs. Figs. 3 Figs. Fig. Figs Figs Figs 1-3. 4,5. 3 06,. 7. . 8,9. 10, 11. 12 Figs. 1-12 Sparganiaceaepollenites barungensis sp. nov., X 1250, Figs. 1, 2.—ST 325/15, 42.4: 110.3. Fig. 1.—High focus. Fig. 2.—Mid focus. Fig. 3.—S 325/3, 39.7: 96.8. Pore in N.W. quadrant. Amosopollis dilwynensis sp. nov. Py 015, 38.7: 100.1. Fig. 5, X 1250. Notice the granulate margin of the sulcus. “Triorites” psilatus sp. nov. X 1250. Fig. 6.—S564/1, 32.8: 100.6. Fig. 7.—S562/1, 37.7: 104.9. Tricolporites valvatus sp. nov. Fig. 8—ST 241/12, 39.0: 98.2. Fig. 9—Py 176, 35.1: 98.1, X 1250, NDIC. Triporopollenites_gemmatus sp. nov. Fig. 10.—S650/1, 43.4: 104.5, single grain. Fig. 11.—S547/1, 31.7: 98.4, tetrad. Gambierina edwardsii (Cookson & Pike) Harris comb. nov. ST 208/2, 32.1: 103.7. SSS Figs. 13-25 T. gemmatus sp. nov. Fig. 13.—S547/1, 31.7: 98.4, X 1250 high focus. Fig. 14.— S741/2, 38.6: 106.8. Ericipites crassiexinus sp. nov. Fig. 15.—S660/1, 52.9: 96.9. Fig. 16.—S560/1, 19.2: 107.2. Sapotaceoidaepollenites rotundus sp. nov. Fig. 17.—Py 167, 45.7: 102.8, X 1250, NDIC. Fig. 18.—S564/1, 29.5: 106.4. Proteacidites confragosus sp. nov. Figs. 19, 21, 22.—ST 241/9, 41.4: 105.7. Fig. 21, 22, X 1250. Fig. 9 focused on ornament, fig. 10 focused on apertural region. Fig. 20, —ST 241/4, 35.6: 104.7. Proteacidites tripartitus sp. nov. X 1250. Fig. 23.—S560/1, 27.7: 100.8, NDIC. Figs. 24, 25.—S650/1, 32.2: 99.8, sectional and high focus respectively. SSS Figs. 26-37 Proteacidites kopiensis sp. nov. Fig. 26.—SS60/1, 26.1: 106.9, X 1250, NDIC. Fig. 27.—S623/1, 16.6: 109.3, NDIC. Proteacidites tortuosus sp. nov. $563/2, 32.2: 99.8. Median and high focus respectively. Proteacidites clintonensis sp. nov. Figs. 30, 31.—S741/2, 47.8: 96.3. Fig. 31, X 1250, high focus on polar region. Figs. 32-34.—S705/1, 31.3: 105.2. Fig. 33, X 1250, high focus on polar region; fig. 34, focus on apertural region. Proteacidites fromensis sp. nov. §17/2, 35.1: 101.4. Figs. 36, 37, X 1250, Fig. 36, high focus on polar region; fig. 37, median focus on interapertural region. eee Figs. 38—49 Proteacidites fromensis sp. nov. §1986/2, 27.0: 99.3. Proteacidites varius sp. nov. X 1250. Figs. 39, 40.—S705/1, 37.2: 111.0. High and mid focus respectively. Fig. 41.—S547/1, 22.7: 96.0. Fig. 42.—S705/1, 98.8: 44.3. Proteacidites wilkatanaensis sp. nov. Figs. 43, 44.—S2273/4, 44.1: 106.7. Mid and high focus respectively. Fig. 45—S705/2, 22.8: 105.9. Figs. 46, 47.—S2273/3, 33.2: 107.1. High and mid focus respectively. Proteacidites concretus sp. nov. X 1250. Fig. 48.—S360/2, 35.8: 105.5. Fig. 49.— $705/3, 26.9: 105.5. WAYNE K. HARRIS EARLY TERTIARY POLLEN SPECIES 63 f4 WAYNE K. HARRIS EARLY TERTIARY POLLEN SPECIES 6 an VOL 96, PART 2 31 MAY, 1972 TRANSACTIONS OF THE ROYAL SOCIETY OF SOUTH AUSTRALIA INCORPORATED CONTENTS Preiss, W. V. The systematics of South Australian Precambrian and Cambrian Stromatolites. Part 1 - - - - - - - - 67 Mawson, Patricia M. The nematode genus Maxvachonia (Oxyurata: Cosmo- cercidae) in Australian reptiles and frogs - - - - 101 Mawson, Patricia M. Three new species of the genus Cloacina Linstow (Nema- toda: Strongylata) from macropod marsupials - - - 109 Schodde, R., Mason, I., & Wolfe, T. O. Further records of the Pitted-shelled Turtle (Carettochelys insculpta) from Australia - - - 115 Wollaston, Elise M. The morphology and relationships of Muellerena wattsii (Harvey) Schmitz (Ceramiaceae: Rhodophyta) . - - 119 PUBLISHED AND SOLD AT THE SOCIETY’S ROOMS STATE LIBRARY BUILDING NORTH TERRACE, ADELAIDE, S.A. 5000 THE SYSTEMATICS OF SOUTH AUSTRALIAN PRECAMBRIAN AND CAMBRIAN STROMATOLITES. PART 1 BY W. V. PREISS Summary The methods of field study and detailed morphological analysis using three-dimentional reconstructions and thin sections, developed by one Russian school, were applied to the abundant Precambrian and Cambrian stromatolites of the Adelaide Geosyncline. Although other schools either demand formal taxonomy for algal remains only, or use informal descriptive nomenclature of morphologies which they believe are determined entirely by environment, it is concluded that valid and consistent stromatolite form-taxa can be distinguished by these studies. The recognition of stratigraphically restricted taxa suggests biostratigraphic subdivision and intercontinental correlations. New forms Acaciella angepena. A. augusta, Baicalia burra and Boxonia melrosa, and an indeterminate form of Acaciella, are described. THE SYSTEMATICS OF SOUTH AUSTRALIAN PRECAMBRIAN AND CAMBRIAN STROMATOLITES. PART L by W. V. Preiss“ Summary The methods of field study snd detailed morphological analysis using threc-dimentional secon- structions anid (hin secliuns, developed by one Russian school, were applied to the abundant Precambrian and Cambrian stromatolites of the Adeloide Geosyncline. Although other schools either demand fortnal taxonomy for algal remains only, or use informal descriptive nomenclature of morphologics which they believe are determined entirely by environment, it is concluded that valid and consistent atroma- tolitc form-taxa. can be distinguished by these studies. The recognition of Stratigraphically restricted taxa suggests biostratigraphic subdivision and intercontinental correlations. New forms Acacielia angepena, A. augusta, Baicalia burra and Roxenia melrosa, and an indeter- mitiate form of Acaciella, are described. Introduction Stromatolites are laminated structures formed in sediments, mostly carbonates, by the tapping and precipitation of sediment by mats of algae and bacteria. They are known through. out the sedimentary record, and are particu- larly abundant in otherwise unfossiliferous Pre- cambrian sequences. This observed abundance and the diversity of forms have made stroma- tolites potentially useful as index fossils, pro- vided that taxa can be defined which have stra- lgraphically restricted time-ranges, A group of Russian stromatolite specialists has been en- gaged in the systematic description and elassifi- cation of stromatolites for the past fifteen vears and their results stimulated this study of South Australian stromatolites in an attempt to apply biastratigraphic methods to the problems of the age and correlatron of the Precambrian sequence in the Adelaide Geosyncline. This paper is hased on the systematics sec- tion of a thesis submitted for the degree of Ph.D., University of Adelaide, It is necessary here to briefly discuss the taxonomy of stroma- tolites and particularly to formalize several new tixa meeded for subsequent discussions of bio- stratigraphy and palacoccology. This paper. the first. of three, will include yn outline of pre- vious studies, consideration of some taxonomic pringiples and problems, and descriptions of four new forms and one indeterminate form. The other parts will comprise descriptions of further forms and a discussion of the strati- graphic distribution of stromatolites. Stroma- tolite forms will be described in alphabetical order, Background This history of the early study of strama- toliles Was comprehensively reviewed by Masloy (1960), Although most researchers prior to 1914 sought an animal origin for these structures, for example Hall's Cryptezoon (1883) and Steinmann’s Gyrmrosalen (1911), Walcott'’s (1914) discovery of filamentous microfogsils in Precambrian stzomatolites from the Belt Series of Montans paved the way for the understanding of stromutolite formation by algae. Later workers clarified the role played by the algac. In particular, Black (1933) established that the algal mats of the Bahamas are poly- specific and that the mucilaginous filaments of the blue-green algae present trap detrital grains. Pia (1926) recognized the rock-building pro- perties of modern blue-green algac. Algal fila- ments were also found by Bradicy (1929) to aceur in stromatalites of the Eocene Green River Formation of Wyoming, In Australia, Mawson (1925) recognized stromatulites in the Flinders Ranges, and started a collection which was partly used in this study. During the 1930's fossil stromato- lites were described by numerous authors, the most. important being Young (1933a, (933b, 1935), Fenton & Fenton (1931, 1933, 1936, 1937, 1939), Johnson (1937, 1940) and Maslov (19374, 1937b, 1938, 19394, 1939b). The work of others were reviewed by Masloy : 5 A aad ial Survey, South Australian Depariment of Mines, Rox 38, Rundle Street UO, Adelaide Trans, R. Soc. 5, Aust, Vol, 96, Part 2, 31] May 1972, 68 W, OV. PREISS (1960), Mast of these uuthors tacitly accepted the validity of a formal binomial nomencta- ture Por stromatulites. Cloud (1942) was the first to question the validity of such uw classification, arguing that stromatolites are built by associations of alyal species. Similarly, Johnson (1966) has more recently rejected the use of this nomenclature, and suggested rather that only actual algul spe- cits should be named. if they are present. Nevertheless, Rezak (1957) found it useful to retain a binomial nomenclature and used the defined tuxa successfully for intrabasinul corre- lution. Since the controversy regarding the classi- fieation of stromutolites arose, ut Jeast three schools al thought have evolved, Firstly, a small group of Russian students (c.g. Vologdin 1962), like Johnson, considered that. oniv actual ulgul remains can be validly named, But algae are very rarely preserved in Precambian stramutolites, and mast of the micro-structures referred hy Vologdin to fossil algae are very doubthully of organic origin. A second proup rejects the concept oF bio- logical comirol over stromutotite morphology, and uses purely descriptive classifications to aid environmental interpretations. For example, Masloy (1960) used “generic” names such as Collenia, Conpphyton and Glebrilella, but he modified these by a series of descriptive Latin auljectives. Logan, Rezak & Ginsburg (1964) used symbols, and formulae to deseribe various features of stromatolites, which they showed to be influenced by the local environment. Both Maslov's multinomial nomenclature and the variable descriptive formulae of Logan ef al, tend to be cumbersome, and cannot in them- selves describe all the useful characters of stromatolites. Some of the simpler formulac are. hewever. very useful in routine lichd des- criplions, Hofmann (1969a) found difficully in upplying a binomial nomenclature to the surno- matolites of the Gunflint Iron Formation, and later (Hofmann 1969b) discussed the signifi- eance of various characters of stromutolites, concluding that dilferent morphologies are moze likely to be environmentally than bio- legically controlled, Tloffman (1967, 1969) gave an outstanding example of the use of stromatolites in palaeocurrent determination The third school is a Russian group which describes and classifies stramatalites on tbe Basis of morphology and microstructure. and uses them for blostratigraphy, Their first results were reported by Keller et al, (1960). Despite differences Of emphasis today between dilferent workers, all usc a binomial nomenclature with the form taxu “group” (analogous ta genus) und ‘form™ (analogous to species), They have found that the time ranges of the defined taxa are restricled, and this allowed them to sub- divide and correlate Late Precambrian sections throughout much of the USSR. The biosirati- graphy was supported by numerous radiometric datings, both K-Ar determinations on glaueo- nites and K—Ar. Rb-Se and U-Th-Pb deter- minattons on intrustves. The subdivision is ws follows: Cambrian $70 = LO my. Vendian ———— —- 6380 = 20 my Late Riphean -—- = 950 = 50.m,¥, Middle Riphean 1,350 = 50 my. Early Riphean 1,600 = 50 mv. The approach of this group was applied to Austruliun strontarolites, and it was found that many of Ihe Russian taxu do occur here, in a similar order of succession (Glaessner, Preiss & Walter 146% Preiss 1971). The resulting correlations with the daicd Russian sequences were in agreement with most of the radiometric evidence available for the Australian Precam- brian. The successful use of stromatolites in bto- striligruphy implies that their morphotogy a al least partly controlled by genctic characters of algue which evolve in time. The concept of biological control is supported by some studies of madern algal mats (Eardley 193%; Hom- meril & Rioult 1965; Monty 1967). Each of these suthors. has shown the partial dependence of mat type on the predominating algal spe- cies present. This in turn affects the miicrostrue- ture and lamina shupe of the stromatolite, und indirectly, the pross morphology. Thus, decid- ing which characters are genetically determined and which are directly shaped by locul environ- mental fuciors becomes the major difficulty in classifying stromatolites. The Russian work of recent years has showa that it is mainly the columnar stramatolites which are of value in biosirutigraphy, Only Komat (1966) has given 4 detailed account of laterally linked stromatolites but their useful- ness has not been confirmed io the extent of thar of columnar forms. In this study, atten- SOUTH AUSTRALIAN STROMATOLITES 6y Fig, DIAGNOSTIC TERMINOLOGY MODE OF OCCURRENCE BIOHERMS: SUS, Cow Aa Kean SN = (a — _————————— on ES SUBSPHER ICAL DOMED TABULAR NNN rs ete Pips 2 mS PEAAANIMEO GLAD Dg eter IL Fonte Oe TE > OORT A ere all) Y wy Ree TT ARE TTT S S a anual BOMED EUAN DURUCUTE Lt a URAC CAR ASLAN UA cb cd AAA COTTA OTT BIOSTROMES: TABULAR BRANCHING AND COALESCING nay Gel alpha beta gammo SLIGHTLY PARALLEL DIVERGENT MARKEDLY DIVERGENT COLUMNS COLUMN SHAPE AND MARGIN STRUCTURE — NICHE AND PROJECTIONS PROJECTION RIBS WALL m LAMINA SHAPE pn GENTLY CONVEX RECTANGULAR “WAVY /\ EX STEEPLY CONVEX ‘RHOMBIC —- WRINKLED UNCONFORMITY esta ESA CONOPHYTONS NON-COLUMNAR STROMATOLITES SS RES an SS ERBReESe ee aa fs ge oN Ne oo OO Pe FLAT-LAM INATED ASS SRS SE Zeer ae oN el NNN = = = SOS SSS OAS ENS Undulatary Pseudocelumnor oS CUMULATE ; LATERALLY -LINKED COLUMNAR-LAYERED Fig. 1. Diagnostic terminology found useful in the description of stromatolites, The diagrams illus- trate features discussed in the Appendix. 70 W, V. PREISS Hon was also concentrated on columnar forms since these have the most characters allowing them to be classified, Therefore the binomial nomenclature has been applied onty to these. The terms used here to describe stromptolite characters, ate largely based on translations of Russian lerms, with minor alterations and addi- wens. Mast of the new terms introduced by Ilofmann (1969h) are unnecessary from the point of view of this study, The diagnosti¢ ter- minuloxy proposed by Glagssner er al, (1969, Fig. |) bas been expanded (Fig, J), and the terms used in the descriptions ure defined in the glossary, Appendix I, ‘Taxonomy Ia general the methods of stromatolile study and classification used by the Russians. Krylov (1963, 1967), Semikhatov (1962), Nuzhnoy (1967) und Komar (1966) have been applied here, inclucling hinomial nomenclature. Al- though muny of the group names have been accepted! tn pulaeohotanical literature os penera, of. Baicelid, Conophiyron and Gyiino- yolen by Andrews (1970), it is considered that retention of the terms greup and ferm empha- sizes the distinction between stromatolites asx organo-scdimentary structures and actual [rag- mentary plant remains to which the terms form genus and ferm species may be applic- able. While similar groups of characters are studied Cor cuch stronvitolite, The relative 1ako- none significance attached to any particular Character muy vury from luxon to ison, tle- pending on its diagnostic value, Mostly. groups are defined an the basis of gross morphology. column shape, branching and margin structure, Lamina shape and microstructure are fre- quently useful in the distinction of fornrs But sometimes these features are diagnostic at group level—Conopiyion, lor example, is diag- nosed by its lamina shape, and is characterized by purecuhirly distinet Januinulion, Baicalia tends Lo he characterized by banded laminating, except where altered by cliagenesis, Although the presence of a wall is a dingnostic difference between sone eroups, Walter (1970) 7 has des- etihed single stromatetiles which are unwalled in their flower parts and walled at the top. in which case other features are diagnostic. No similar situation is known from South Austra- lias, IL is considered unnecessary to use categories higher than the group, as Raaben (1964, 1969a, 1969b) has done. Her higher taxa are somewhat arbitrary and several alternative ones could be proposed, but all are equally question- able. On the other hand, the wodery asa sub- division of the fornt is Gseful in cases where finer subdivisions can be made, and is there- fore retained for Coaaphyton garganicnm. It could be argued that a single name would be sufficient ta characterize a particular stroma- tolite, but the value of a binomial nomen- elature is that it mudicates real similarities and differences between various forms. Thus groups contain one or more forms which all share a number of characters considered diagnostic for that group. Forms are distinguished within a vroup whenever there are suflictent gross or microstructural differences, But the essential compunson between closely related Turns would be lost without a binomial nomenclature. The chief difficulty in the Waonomy of sino matolites is the isolation of discrete character combinations, where intergradution is commean- Thus, subjective choice may he required in some causes. Where the morphology temsains uniform Vhreughoul a particular occurrence, & single name can easily be applied, but if there is Varialion within the occurrence, the delini- ton must be broadened, Whether or not the morphology of 9 stromatoltte [rom another egcurrence falls within this range of variation is dificult wy decide. Coaversely, if a signili- cantly dillerent morphology occurs as a disercte portion of un deecurrence, is this to be classi- fied separately? Examples of stromatoliies with a broad cunge of variation ure Puagassta etina and Linelln muanyallina, Toth of these show a spectrum of intergrading branching types and colunin shapes, even within single outcrops, se that the range of variation between specimens of different areus lies within the range of varia on in one locality, and these are therefore included in the one form. Tr has been found that many characters over- lap. and distinctions must be made even al group level Gn the most Commonly occuring expression, ie. the mode. of each character. This is especially true of branching, Bexania is characterized by «-paralie] and some §-parallel branching, while ysparallel is rare. In Gymno- sole +-parallel predominates, but not to the total exclusion of the other types, Similarly, there is overlap between the branching styles of Burealia {the forms of which show a Cromen- Unpublished PAD. thesis, University ef Adelaide. SOLTH AUSTRALIAN STROMATOLITES 7 dous vanation of branching, as shown by Krylov 1967) and that of Tungussia, Bul while Baicalia has predominantly slighthy to mode- rately divergent branching, in Twngessia mae- kedly divergent branching predominates. Although it is often easy to recoynize groups on the basis of even limited reconstructions and lonvitudinal sections, the identification of forms is more difficult and subjective, Forms are dis- tinguished orn minor features of column mor phology, lamina shape, or microstructure. Microstructure is the most difficult chataciter la use, partly because different types intergrade to some extent and partly because it ia so easily altered by diagenesis. The distinctive lamina- tion. of Conopiryron is somewhat ¢xccptional, and is amenable to statistical analysis. Although Raaben (1969s) has attempted similar studies on Migeria and Semikhatovy, Komar & Serebrya- ‘kov (1970) have measured the sizes of clots and pelleis in Bexonia, it is uncertain Whether or not the structures measured are primary. The laminae of most South Australian columnar branching stromatolites are too diffuse and variable to allow a detailed statis- tical study, although the well-preserved repre- sentatives of the banded microstructure of Baicalia burra might be amenable, Stromatolites can be classified only on the basis of combinations of characters, and, as Walter (1970, unpublished) has also con- cluded, the classificatory significance of charac- ters must vary to some extent from taxon to taxon, The classification has been found em- pirically to be useful in that the resulting taxa are lemporally restricted. The questian arises as to the fundamental meaning of these taxa, and why they wre sa restricted, Several possi- bilities. exist: (1) Each form is built by a particular asseo- ciation ef algal species, and forms change as the content of the associations changes. Each form is built by a dominant algal Species, in association with other species that have Jittle effect on stramatolite morphology. (3) The environment. and not the algal composition, entircly controls the stro- matolite morphology. ff (3) were true, we should expect 4 tem- poral restriction of forms only if the environ- Ment has systematically evolved in time. Lt is difficult to sce how local factors such as current activity or sediment accumulation which could (2) conceivably control stromatoliie morphology, can exhibit continent-wide, if not world-wide, Unidirectional change. On these grounds, this possibility must. at present be rejected. If (i) were true, Wwe could expect the morpholazy te change gradually as the overall algal coniposi- tion changes. one species replacing another in the association. On the other band, if one specics controls the morphology, a rapid change would be expected. At present, itis not possible to fell which of (1) and (2) is correct and possibly both upply,; the first possibility may explain the intergradations sometimes observed between taxa, When classificahoa becomes difii- cult. Although Hofmann (1969h) regarded Cone- pPhyren as apatt from other stromatolites, recent work has shown that conuphytons pos- sibly intergrade With columnar branching forms such os Baicalia (Shapovalova L968). Similarly Bertrand (1968) described intergra- dations of Conephyren and branched forms. While the taxonomic significance of these changing morphologies and theit relationship to environmental factors has not. been fully determined, it is clear that Conophyton is not fundamentally different from other stromato- lites, It is concluded that stromatolites must be defined on combinations of characters, the sige nificance of each of which may vary in differ- ent taxa. The fact that some taxa have much broader ranges of variation than others results from the necessity of grouping intergradina morphologies present in single stromatohte occurrences, Methods Stromatolites were studied both in the field and in the laboratory, but ficld observations were often limited by outcrop conditions and lichen cover on tock sutfaces. Where possible, the mode of cecurrence, column shape and umangement and branching were observed in order to vjin an impression of the tutal yari- ability, The variable nature of stromatolites neces- silates sampling of sufficient material to dcter- mine the modal expressions of churacters pre- sent. Depending on the size of columns, large specimens weighing from 4-70 kg were col- lected, and the relative position and orientation were noted, Ideally, bioherm centres and mar- gins were both sampled. The diagnostic gross features of colummar stromatolites can only be determined from a 72 W. V. PREISS three-dimensional view of the stnicture, This is achieved by the method of “graphical recon. struction” described by Kryfov (1963), A series of LO to 15 serial Jongitudinal slabs 2 to 6 mm wide were cut on an oil-cooled 60 cm diamond saw with a saw cut about 2 mm wide. ‘The columms were outlined jn pencil on the slabs and traced on to a block diagram frame- work on tracing paper. each Jongitudinal sec- tion heing parullel to the front. face of the block. The reconstructions were retraced with shading to show surface morphology and finally redrafted by stippling. Lamina shape, margin structure and micro- structure were studied in large, longitudinal, thm sections, up to 20 em long, Thew thick- ness varies with the nature of the rock, hut in general they must be thicker chan petrological sections to preserve the distinctness of the structures. Carbonates were mostly identified by staining with Alizarin Red §, but were [re quently checked by X-ray diffraction powder photographs, Systematics For each group from which forms are des- cribed, 4 diagnosis, a list of the known cansti- tuent forms and the stratigraphic and geogra- phic distributions are presented. Fotms are diagnosed only if described here for the first ume. Descriptions are given under the headings made ef occurrence, colunu shape and arrange- ment, branching, margin structure, laenine shape and microstructure, The interspace sedi- ments and the nature of secondary alteration aré wiso described since they provide importunt clues to the depositional environment and dia- genelic history, The distribution of forms refers to both their geographic distributions and te the rack-strati- graphic units (Thomson er «il. 1964) in which they occur. Reference ts made for each locality and stratigraphic unit to the relevant geological sheet (either 1:63.3A0 er 1:250,000 map sheets, Geological Atlas of South Australia). Tt was found convenient (Preiss 1971) to subdivide the Adelaidean into two time units: the Barly Adclaidean, represented by all seci- ments up fo the pre-tillite unconformity, and the Late Acelaidean, represented by sediments from the base of the lower tillite to the base of the Cambrian. This subdivision reflects both @ climatic change and a major change in stro- matolité assemblages. Ages of stromatolites will be referred to as Early or Late Adelaidean, but the probable corrélations with the subdivi- signs oF the Riphean will he noted in each case. Type specimens are kept in the Department of Gealogy and Mineralogy, University of Adec- vide, catalogued under numbers prefixed by S, Group ACACIELLA Walter Walter has supplied the group name Avca- ciella and the following diagnosis, “Type Form: Acaciella australica (Cryptozeen dusivalicum Howchin 1914). Diagnosis; Nearly straight, parallel or radially arranged sub-cylindrical columns with a-, f- and rately y-parallel and very slightly divergent multiple branching. On column margins are numerous low bumps and occasional small cor- nices ond peaks; small areas Of wall occur in- frequently, Laminae dominantly are rect- angular. rhombic of gently domed and are not markedly wavy or wrinkled; the microstructure is streaky.” Content: Acaciella australica Walter, A angepere f. nov. aid A, aneusta f. nov. Age and Distribution: Adelaidean to Early Cambrian: Loves Creek Member of the Bit- ter Springs Formation, Central Aust.; the Lower Cambrian of S. Aust.) the Wundowje Limestone and Brighton Limestone equiva- lent. Umberatana Group. S. Aust. and as erralics in the lower (Sturtiqn) elaciily, 8. Aust, Acaviella angepena f, nov. FIGS. 2 10, lla Material Forty-seven specimens from ie localities. Nolorype; S460 (Pigs. 2a, 10c), Lower Cambrian, | km south of Angepena MS., Northern Flinders Ranges, Name After the type locality, Diagnesis: Acaciella with vertical or radially arranged columns. or pseudocolumus, which may branch upwards from either flat-laminated or small cumulate stromatolites.. Columns may branch upwards into minute, irregular columns, Bridging 1s. extremely common. Microstructare is regularly banded, with thin continuous Jaminae. Verniform microstructure may be de- veloped, Description Mode of oecurrence: Cambrian stromatolites were studied in outcrop only in the Angepena aren, where lenticular stromatolite beds consist of closely spaced ellipsoidal and domed bio- herms 3 to 50 m wide. These overlie flaggy, SOUTH AUSTRALIAN STRGMATOLITES am] laminated, dark grey limestones with irregular efosional contacts. Cumulate or pseude- columnar stromalolile individuals commence growth upon the erosional highs, aod pays up inte radially arranged or parallel short columns with very numerous bridges, At bioherm mar- gins, columns and psetidyceltimns become hori- vontal. and laminae are deflcxed parallel to the overhanging sides of the bioherm. su thut here growth actually proceeded duwnwards (Fig. 1Ou & b, showing longitudinal sections of a bioherm margin), Where adjacent biaherms become contiguous, they are overlain by a damed biostromal layer of columnar, pseudo- columnur und columnar-layered stromatolites, similur to those of the bigherm. Ac the edge of a stromatolite bed, the terminal hioherm has an abrupt vettical margin and the laminae bend downwards only slightly. The surrounding sedi- ment of dark Jime mud accumulated synchran- ously with stromatolite growth, and vecasional lgul Jaminae are intercalated with it: the bia- hero probably never hod more than 10 em of relicf over the surrounding sediment surface. Also, there is evidence of contemporanedus compaction of the time mud the bicherm resis upon (Fig, 10ce). the lower layers of the sur- rounding muds ate depressed, while the upper ones simply abut agalnst and cover the bio- herm, Column Shape anid Arrangement: Column shape is highly variable in single bioherms, mainly due to different degrees of coulescing and brilging. The structures vary from laterally linked pseudocolumns with some discrete small cumuli (Fig. 10d) to frequently bridged and coalescing calumas (Fig 11a), to discrete, parallel subcylindrical columns (Pigs. 2d, f, a: 10f). The latter chiefly make up Muawson’s (1925) callection from Italowie Ciorge. In all specimens where columns are reasonably dis- erete, they are smooth to slightly bumpy, some- limes wilh pointed terminations (Fig. 2b, j), while others branch into minute columns 1 to Jom wide. Columns are commonly Jess than ! em diam., but brogd, cumulate columns up to 10 cm diam. have heen observed. Transverse sections of columns are round. reunded poly- sonal or lobale (Fig. 2d, f. 2). Columns may he verticul or radially attunged, especially on dhe margins of contiguous bioherms. Dolomitiza- tion of interspaces frequently obscures the ori- ginal margins of the minute columns sq that their shape cannot be accurately devermined_ Branching; Branching is most commonly g- of B-parallel, vccasionally y-parallel, Columns fre- quently branch into narrower columns which do not regain the former diameter. Some branches ure in the form of thin pointed pro- jections (Fig. 2j) Ac bioherm margins, branching may remain parallel] (Fig, 10¢) or hecome radial (Fig. 100), but here the stronia- tolites are Jargely pseudocolumnan Margin Srracture; Colum matgins are rarely preserved intact, Commonly they are corroded by dolomite thombs, if the interspaces are dolo- milized; otherwise very fine stylolites may be developed. Bridging is extremely common in all specimens except those from Matvsan's cal- Iection from Ttalowie Gorge. in which the columns are mostly disctete. These alsa have the smoothest margins, with only slight, acca- sional bumps and ribs. Columns are always unwalled. the laminae thinning only slightly near the columo murgin, Laminse may slightly overhang the margin, but long peaks and ¢or- nices are absent, Lonine Shape: Fig. 8a illustrates common lamina shapes: most ire gently convex. Of 101 Jamina measured, 69% have height to diameter ratios (b/d) beneath 0.2 and 0.4, only 7% have ratios greater than 0.6 (Fig, Say, Laminue ure smoothly domed, without sharp changes in shape from lamina to Jaminoa, A few of Mawson's specimens from Itulowie have wavy laminae, Of Wavelength 3 tc 10 mm, amplitude 1 10 3 mm (Fig, 10f). Microstructure; Microstructure in all spect mens is regularly, thinly banded, with contin. ous Jaminac of uniform thickness across & colunin width. In most specimens there is little contrast between dark and light laminae, except in the amount of organic pigment. Some speci- mens, especially fram Angepena, have ifregu- larly tubular, sinuous, agastamosing. vermi- form spurry patches, O05 to 0.1 mimi thick anc up to 0.6 mm long, crossing the dark Juminae. Dark laminae, varving in thickness from 0,03 to 0.07 mm, consist of xenotopic calcite of grain size varying from 0.003 to 0.01 ming, stained With vroy orgunic pigment, bul in some specimens, subhedral dolomite rhombs of grain size O=.1) to 0.02 ram are interspersed, Minor subangular quartz, silt may be present, Indivi- dual laminae ure continuous, anc af constanr thickness across the colunin width, but may be markedly wavy, In specimens with veentiform microstructure, dark laminae are generally thicker. up te 0.3 mm, but remnunts of finer luminution ut ullen preserved, The hwundaries of the sparry patches are often irregular and their orientation varies from petpendichlsr to TA W. V. PREISS gently inclined to the lamination, but is com- monly at a high anule to it. The vermiform microstructure may be consistently developed preferentially on one side of u column Trans- verse sections of the tubules ate found to elon- gated, irregularly Oriented and anastomoxing. The tubules may be interpreted as algal borings in the fine, lime mud laminae, but not the whole sediment was alfected. since homo- geneous ahd bored! laminae occur side by side This fact also makes it unlikely that they arc casts of actual algal filaments. Whe distribution of borings on one side of columns may be environmentally determined. Bathurst (1466, p. 20) illustrated a sequence of events Invalved in boring by algac; if the process were stopped ul stage 12), and the borings infilled’ with sparry calcite, a structure similac tw the vermi- forn) microstructure olf A. angepena would result, Light laminae are 0.03 to 0.1 mim thick, frequently indistinct, but continuous ucross a colimm width. They are especially poorly dif- feruntiated in specimens with vermiform micro- structure, Where the tubules may pass ocross the light-dark lamina boundaries. Light laminae consast of xenolopic calcite, often with inter- locking ervstals G.015 to 0.03 mm in diameter. Subhedral to) cuhedral UO. mm dolomite rhombs are scattered throughout the light linninac in some specimens, Jrevspacey> Tnterspaces are filled either with Altered micrite or fine sandy and silly micrite Specimens from Jralowie (Mawson's collee- tion) have very narrow interspaces Filled with sparse, qngular quartz silt. supported by a Micrite matrix (Fig. 1Of), somelimes exten. sively dolumitized, with inequigranular hypidio- topic dolomite ringing in gruin size from O.005 10 O.L mm. Extremely finely disseminated hac- mutité may be present tn interspaces. Strom.- lolites from Angepena also have sandy inter- spaces, byt these are more frequently inter- rupted by bridging laminae, Subangular to sub rounded quartz grains vary in diameter from 0.08 to .S tom, and may be partially or wholly replaced hy calcite, Qpids and small intraclasts occur very rarely. Secondary Alteration: Dolomitization is com- mon jn all specimens, and is probably of late diagenetic origin. Within columns, rhombs post- dule the vermiform microstructure and have also formed in the micrite of interspaces. anc in places, interspaces may be totally dolo- mitized, Here sparry calcite occurs as irregular patches between dolomite rhombs perhaps fill- ing @ secondary porosity, The grain size and density of dolomite rhombs decrease murkedly across the ealumn margins: perhaps interspaces were uriginally more porous, lo cause the pre- ferential dolomitization (in Fig, MWe¢ nore the dark calcite columns and the white, dolomine interspaces), StvJolites may follow column niat- gins. or may be grossly cross-cutting. Haema- tiie dispersed through carbonate a4 probably sccondaty. In fealowie specimens, it is concen- trited in interspuces which pass inte ine stylo- lites. Minute irregular cal¢ite veins cut the whole rock, apparently predating the major dolomirzation. Large patches of course sparry cnleite wire bounded by markedly lobate fine atvlolites, suggesting their origin as solution cavities, Compariseny In sross morphology (mode of occurrence, column shape, branching and margin structure) the strumatutites from Ttalowie are similar to AcecieHa Waller, Columns are less discrete in uther aureus, due to frequent bridging and coulescing, but their columnar portrons ure similar to those of Ttalowie specimens. Micro- structures are uniform, except fur the local ver- mniform structure interpreted as ulgal bering, Madiyonites nuiwsont Walter, from the Middle Cambrian Jay Creck Limestone of the Amua- deus Basin, also has vermiformy microstrucnure, but here the tubules ure more consisiently deve- loped, and wre complesty intertwined. the inter- yening micritic areas being reduced to clots. The gross form of Muarlivanites anivsonl is similar to same Acuciella aaxepena in having numerous ireegular frequently bridged columns und pseudocoluniis, however, it lacks the sub- eylindrical, parallel branching, discrete columns Yourd at Tralowre. Acacrelle — ueyeprentiea resembles Verella svehhasiea Krylov in having evenly banded lamination and wide columns branching into nurrow columns, bul has ragged eclumo margins and Jacks the wall of P- uethbayitd, Mlicta composite Sidoroy is siniilue jn also possessing vermifarm microstructure, hut is distinguished by its very smooth, walled. ehlurins. At thes stage uf is difficult to be cer- tun of the content of the form Aeuetella ange- pena, Despite some variation nf column shipe (specimens frony [talowie have predominantly suhevlindricail, discrete columns, while those from Angepensa have numerous bridges and less recular column margins, all the specimens studied are included in the one form, since these column morphologies intergrade and the Microstructures rempin constint. The stremp- tolites are assigned to the group Acetyl on dratielle angepena, from Lower Carshrian Timestancs, !'lliiders Ranges, 14) —Molptype, 5460, | km $. of Angepena H.S,) (b)—S458. 1 kin) S. Of Angepena ALS.y (c). (2) & (i }—S489. 1 kin S, of Angepena WS; (0) & tz) SB. 4.8 km W, of Itatowie Gorge (oallecred hy Sir Douglas Mawson}; (f)—S44, 4.8 kin W, of Nalowie Gorge (collected by Sit Douglas Mawsan?; (h)—Probable 4. angepena, near Old Wirrealpa (collected by Mr, P, G, Haslett): (i)—Possihle 4. dagepena, 4.8 km WL af Tiulnwie Gorge. V6 WwW. ¥. the basis of gross morphology, They are dif- ferentivied Trom other forms of the group by their thin, continuously banded microstructures and by frequent. development of bridges and pseudocolumns. The very narrow, minute columns into which broader columns branch are abseac in other forms A ferrupinous specimen from Old Wirrealpa is problemutical. Its dark laminae are strongly haematiti¢, the hacmatite being in part distri- buted inte minute dendrites. The small columns branch from basal cumuli, the interspaces being filled with recrystallized biomicrite fhyolithius, spange spicules. archacocyathan and brachio- pod fragments may be recognized). Although the grass morphology resembles that of Aca- cielld ungeperia (Piz, 2h), the extremely regu- lar lannnation is atypical of stromatolites, and the possibility of an inorganic orgin for the structure cannot be excluded Divrifucion: Widespread in the dark lime- stones of the Lower Cambrian at Angepena, Old Wirtealpa, near Point Well, at Mern Merns, Beliana Hill, Chace Range, near Narina HS,, Moro Springs. south of Balca- noona, and 4.6 km west of Ttalowie Gorge; Flinders Ranges, South Australia. (COPLEY and PARACHILNA 1:250,000 map sheet areas. } Age: Early Cambrian. Acaciella angusta £. noy. FIGS. 3a-m, 11d-f, 12 Material; Thirteen specimens. trom two loca- lities plus eight spécimeéns: of Uncertalyy iden- tificulion From a further two localities. Ffoloiype, S401 (Figs 3c. e 120), Brighton Limestone equivalent, Depot Creek, South- crn Flinders Ranges. Name: After the city of Port Augusta, 32 km south of the type occurrence. Diagnesiy: Acuciella with extremely frequent coalescing and bridging of columns at all levels, and with broad und narrow columns closely associated, Colutin margins bear short ribs, low bumps and short cornices. Laminae are geutly to moderately steeply convex or rec- angluar, and of distinct, revulerly streaky Microstructure. Description Made of OQecurrencée: The stromatohtes form lenticular and tonguing bicherms (Fig, 12a) varying in thickness from 3 m fo 50m, and cx- tending laterally for up to nearly 2 km, inter- calatéed at varying siraligraphic levels within PREISS the Brighton Limestone equivalent, Most com- monly. growth commences on a substrate of ooid wod intruclust grainstones, as laterally linked stromatolites, up to 3 m thick; these gra- dually develop interspaces to form broad, bridging and coulescing columns (Fig. 11d). At various levels, these columns branch into harrawer columns 1 to 3 cm wide, frequently with parallel hasal and slightly divergent upper branches (Fig. 12c). Occasionally, narrow columns anse directly from an undulatory or fat-laminated base (Fig. 11f). Columns repeatedly alternate with continuous undula- tory Or flat-laminated stromatolires, which com- monly intertongue with the adjacent sediment; they apparently mark periods of reduced influx of coarse sediment. At bioherm margins, columns bevome slightly inclined. Rarely. there are hemispherical bioherms with columns strongly inclined at their margins, Column Shape and Arrangement: Basal columns are up to 20 cm wide, of ipegular shupe, with frequent coalescitig and bridging, Their margins are frequently inclined. although laminae remain subhorizontal. The narrow columns are 1 ta 3 cm wide, and up to 10 cm long between branches (Figs. 3a-j; | 2b-d). Transverse sections are round, rounded poly- agonal, clongated, or complexty lobate, At least some of the clonggtion is of tectonte origin Columns are Straight of gently curved, with slight swellings and constrictions (Fig, 4a-j); a few are short and marrow, and terminate their growth after a few centimetres (Fig. je). Coalescing is an Tréquent thut almost all columns are interconnected! one specimen con- tains numerous irfegular, short, frequently bridged and coalescing columns. Branching: Branching is frequent at all levels, and generally multiple (Fig. 3a-j). Broad basal columns divide by a-parallel branching into narrower columns, which frequently branch ugain at intervals of less than 10 em} this branching is usually q- or 8-parallel, occa- sionally y-parallel, or slightly divergent (Fig. 3e-c, g). Near points of coalescing, branching tends to be more irregular; gamma-purallel or divergently branched columns approach each other and coalesce (Fig. 3d). Margin Structure: The \ateral surfaces of all columns bear relatively low bumps, short dis- continuols ribs, and a few peaks and comices (Fig. 3a-j). In places. bridges, varying from delicate bridges only onc or two laminae thick to massive, thick bridges (Fiz. 3g), are very frequent; in other places, columns remaln rela- SOUTH AUSTRALIAN STROMATOLITES 77 uvely unilfected by bridging throughout most of their jength. Columns are unwalled, und be- tween bridges and cornices their margins are relatively smooth. Depending on the degree of convexity. Jaminac approach the margin at various angles. Lamina Shape: Lamina shape varies according to columin diameter; narrow columns have moderately convex, Or sometimes steeply con- vex, laminae. h/d greater than 0.6 being rare. Broad columns have very gently convex to rect anvular laminae (Fig. 8b). Of all laminac mea- sured, 70% have h/d between 0,2 and 0.6 (Fig, 9b), Laminae are most frequently smooth, but sometimes brojdly wavy, ¢spe- cially before branching. Laminse frequently become donbly-crested before branching (Fig. \2b-d), but the interspace so formed may be heidged over, in Which case the colunin resumes its former growth pattern. Mirrasiructure: I the best preserved speci- mens, distinct, regular light and dark (gzeen) laminae, and in places macrolaminae up to 4 mm tnck, alternate, forming a regular streaky microsiructure (Fig. 12). Durk larmincde, vary- ing In thickness from 005 mm to 2 mm, are smooth to gently wavy, occasionally wrinkled, and have parallel ipper and Jower boundaries. Single laminae have relatively constant thick- ness across the column width, but frequently lens out They consist chicfly of hypidiotopic to idiotopic dolomite, of grain size ranging from 0,005 to 0.02 mm. The crystals are equi- dimensional, commonly euhedral, and stained pale green, which gives the laminae. their colour. Dolomite crystals are densely packed in the dark luminae. leaving only occasional irre- gular undolomiuzed patches. vonsisting of xenotopic calcite, ranging if grain size from 0.003 to 0.01 mm. Light lamina: vary in thick- ness [rom (07 te 2 mm, single laminae having constunt thickness. They are sparsely deolo- mitized, and consist of xenotopic ta hypidio- tipic calcite, varying from 0.01 to 1.02 mm in grain size, with scattered euhedral dolomite rhombs, 0.005 to 0.04 mm long. Laminae are frequently grouped into broad mucralaminae, up to 4 mm thick, in which very thin, lenti- cular’, erther light or durk laminae predominate. In places, laminae are slightly wrinkled, or uraped over underlying irregularities; in one case laminae are domed over Jenses af spatry calcite, probably open space fillings (Fig. 11e). In a few places small scour structures up to 2 mm decp are cut into the tops of dark laminae. Occasional euhedral to subhediral red- dish hrown limonite peains of 1.01 to 0,02 mm diameter (possible pseudomorphs after pyrite) eceur in both lamina tWpes, Qiterspaces: The distance between columns vanes from | to 14) mm. Interspaces are filled with banded limestone, layers of micrite 1 to 6mm thick alternating with thicker intervals of partially dolomitized intramicrite, Laminag im the interspace commonly abut against the column margins, having accumulated after the arowth of that part of the column (Fig, 12c) The micrite laminac, consisting of xenotopic calcite of grain size varying Sram 0,003 to 0.01 mm. are frequently silty, and slightly graded, Senerslly with sharp upper boundaries, and are overlain by matrix-supported intramicrites and some aomicrites, This sediment may originally have been more porous, as It is extensively dolarmitized; the dolomite is of similar texture Lo that in columns, All remnant calcite is re- crystallized to a hypidiotopic sparry masaic; no micrite matrix remains. Alternatively, this cal- cite may fepresent infilling of volds left by Solomitization, Intraclasts, which may bea pre- served as undolomitized micrite, or entirely idiatopic dolomite, are from 1 co 10 mm long, and up ta | mm thick, and may represent eroded fragments of algal mat- Strongly reerys- tallized dolomitized oolites are actasionally present. Tnotraclasts. which communtiy [te at a high angle to the bedding, muy have fine erained laminae draped over them. Coarse sedi- ment mux was periodic; columns may have had up to 2 cm of relief over the interspace sediment or a bridge, then the interspaces were filled rapidly with intraclasts and finer ¢al- ca/cous ‘sediment. During periods of |elative quiescence, lime mud accumulrted to form thin layers. In some specimens, bridging is very fre- quent, so thal there never was more than ahnut a centimetre of relief. Secondary Alrereasian; Liile is preserved of the primary difference between the light and the dark (green) laminge, which now differ in the extent of dolomitization, The dolomite is equi- granular, idiotopic, and probably secondary, although a detrital origin cannot he miles? ot fi the dolomite originated by replacement of calcite, the preferential dolomitization of dark laminae may indicate that they were oristhally more porous, Smull irregular patches of coursely crystalline sparry calcite within both columns and interspaces post-date dolomitiza- ton, and are associated with fine calcite veilis. Stvlolites aze very fare, being restricted 16 a few which afe concordant with ihe liminaiion or 74 column margins. The grec staining of dolo- mite crystals oxidizes under subaserial weather- ing to form finely disseminated limonite, which may be concentrated along colunin margins or slylolites. Colamas are commonly slightly fiat- tened parallel to an axial plane cleavage, which is better developed south of Depot Creek and at Munialilo Creek. ‘The cleavage is an irre- gular fracture which passes around, not through, slromatalite columns and is commoaniy expressed as stylolites in the carbunate rucks. A specimen from Mundallio Creek contains light laminae wiih prominent radiating, struc- tures; these consist of dolomite crystals aligned in Tows almest perpendicular to the Juaminution (Fig. 12d), and may represent a dolomitized, earlier ueicular texture. In specimens from the Wundowie Limestone (Wundowice Bore and Copley), column margins have been ulmast completely removed by stylolites, Icading to unceruinty of identification (Fig. 3k. 1 & m). Comparisons The predominantly parallel branching (o- parallel at hase, then 8- or rarely y-parallel) and almost total absence of a wall, identify the stromatulites as Acuerelle. 4eagiella qugusta is distinguished fron A. oumniica by the rarity of discrete, brogu, basal culumos. by its mode of oecurrence (lenticu- Jar and wonguing bioherms instead of tabular tnd domed hiostromes), by its extremely fre- yuent coalescing and bridging, and by its very distinct mierostructure, The mode of ogcur- renee and microstructure abe distinguish it from A dnewpena, A. angusta has many wavy, sometimes Tenticalar laminae, and prominent macroluminac, the dark laminas being pre ferentially dolomitized, A, augusta is very simi- lar in gross morphulugy to Eucapsiphora para- disa Clond & Semikhatoy, from the Puradise Creek Formation neu Mt, Isa, N,W, Queens- and. &. paradisa is difficult to distinguish on the basis of the published description, but apparently has a patchy wall. Specimens of poorly preserved stramutolites from the Wondeowie Limestone near Copley and Wundowie Bore. originally tentatively identified as Linella munyallina (Preiss unpubl.) are belter assigned to Acacielfa augusta on the basis of column shape, branching and micro- structure. Where column murgins are not re- moved by stylolites. they arc unwalled, Diswibuyion; Brighton Limestone, Depot Creek ard Mundallio Creek. Southern Flin- ders Ranges, and possibly the Wanclirwie limestone near Copley and Wundowie Bore, wlsy.: PREISS Northem Flinders Ranges (PORT AUGUSUA und COPLEY 1:250.000 map sheet areas), Age: Late Adelaidean, correlated with the Late Riphein of the USSR, Acaciella. form indet. FIGS. 3n-y; 11b, ¢) Material: Two specimens from one locality. Description Mele of Occurrence: Both stromatolite speci- mens are erratic boulders in the lower (Sturtian) glacials; their provenance is un- known. Column Shape and Arrangemens One speci- men (S509). eunsisty of pseadocolumns ancl frequently bridged columns, oriented sub- parallel to shebtly rachialing, wn passing tuter- ally inte flat-laminated stromatolites (Fig. | le). The uther specimen (8339) consists of rather smooth, erect, parallel. cylindrical, discrete columns, I-5 cin wide. Transverse sections are round ar rounded polygonal {Frg. 3n-p). Brunching: Hraneching ix commonly a- or p- parallel; columns either retain their width or widen gradually before branching. Axes of branching columns mav be very slightly diver- pent (Fig. 11b), Specimen §539 shows only dichatomous branching, but S509 has sume multiple, o-parallel branching. Murgin Stractvre; S539 hos wo ruther smtoth margin structure. with low bumps and a few very shorr peaks and overhanging laminie (Fig, 3n-p). There is no wall; laminae simply tefminute, withoul appreciable thinnmg, at the column margins. Bridges are extremely fre- quent in SSOQY, but otherwise column: margins are similar to 3539, Few columns in $509 are entirely discrete. Tamina Shape; All laminae are gently conven (Fig. 8c). hid never exceeding 0.5. und 84% of Jaminue measured have h/al between, 0.2 and 4 (Fig, Yel. Laminae are smoothly curved, rarely rectingular, and without wrinkles or sharp Nexurcs. Occasionally Inminae are slightly: wavy, and belere branching always develop multipte crests (Fig. 11b, ¢), Laminae are not normally deflexed at rhe column margins. Microstructure: Microstructure consists of very smooth or broadly wavy, light und dark. dolo- mitic, striated ta banded, laminae. ‘There is little conttust between laminac. Dark laminae are 0,05 jo 0.5 mm thick, and commonly pinch and swell slightly across the column, and in places they ate lenticular, but otherwise, they have smooth, parallel boundaries. They consist (a)-(); Acactella augusta, Brighton Limestone equivalent, Umberatana Group, Flinders Kanges: fa), (bo, (d¢) & (1) S404, Depot Creek, Southern Minders Ranecs; (cl & fey— Holotype, 5401, Depot Creek: (£) S538, Mundallio Creek, Southern Flinders Ranges; (2) & (h)—S396, Depot Creek: (j)—8537, Depot Creek: (Kk), (1) & (m)—Paossible 4, augusta, Woadowie Limestane Member, Woudowie Bore. Northern Flinders Ranges; (n). fo) & (p} —S539, Aenelella F, indet. from an erratic in the Sturtian glacials, N.E. of the Pnorams Diapit, (q)—S509, Acaciella F. indet,, from the same locality. Sketch traced from a thin sec tion, 80 W. V. PREISS ot pale vrey stained hypidiotopic dolomite of gruin size varying from 6.003 to 0.015 mim. Light laminae vaty in thickness from 0.05 to 1.0 mm, generally with little change across a colummn. thinning only shghtly towards colunia margins. They. consist of hypidictapic to idio- topic transparent, unstained dolomite of grain size varying from 0.015 to 0.06 mm. Very characteristic of $539 is the presence of very fine, limonite-rich solution surfaces, concordant with the laminac, Although these are probably stylolites, surfaces with lithe or no wrinkling, which follow the fine-scale structure of laminae exactly, are especiully common (Fig, 11b). In places these are only about 0.5 mm apart, and light Jaminae muy be separated by them, with- out intervening durk laminae. Interspaces. Interspace sediment 1s completely dolomitized, consisiing of equigranular, idio- topic dolomile of grain size ranging from 0.01 to 0.05 mm. There is little contrast between columns und interspaces, but small amounts of subangular quartz silt ure present in the inter- spaces, Fraginents of slightly darker stained dolomite, of similar texture to the matrix, pro- bably represent original intraclusts. The nature of the matrix cannot be determined, bul the sparsity of intraclasts stiggests that these were mud supported, Intraclusts are better preserved in S509, Secondary Alteration: Dulomitization af the siromatolites and interspaces is clearly second- ary, as indicated by the general idiotopic,, equi- granular texture, and poor preservation of the finest structures, Stylolites are of at least two generutions: the earliest stylolites are almost perfectly concerdani, without lobes or wrinkles: these possibly predate the dolomite euhedra, which in places cut inte them, and certainly predate 2 relatively coarse grained dolomite vein (grain size up to 0.1 mm). The vein is itse]E cut by more prooounced, slightly diseord- ant stylolites. Occasionaly cross-cutting srylo- hites cut interspaces and columns, some follow- ing column margins. Dotomitization almost certainly predates the erosion und depasition of the clasts into the glacial sediments. Comparisons The straight, chielly 4- to f-parallel branch- ing unwalled columns allow assignment to the group Acaciella, They are clearly distinguished from Acueella uvgasia by the discrete, rather smooth, more cylindricul columns; although bridging and coalescing oceur in S509, this specimen is considered to represent the basal purt of the stromatolite bed. The distinct, sub- evlindrical columns with telatively smooth mar- gins and gently convex laminae are similar to A. awsiralica Walter, but the specimens are inadequate for identification, Distribution, As clasts in the lower (Sturtian) glacials, on the Aynks of Enorama Diapir, 6.4 krn North of Orgparinna HS., Cep- tral «ss Flinders Ranges {PARACHILNA £:250,000 map sheet area). Age: Probably Adelaidean, but net younger than the Sturtian glacials Group BAICALIA Krylov Bailie Krylov 1943+ 64. Semikhatoy 1962: 198. Komar 1966: 82. Krylov 1967: 25. Nuzhnev 1967: 135, Cloud & Semikhatay T1969: 1035. Type Form: Baicalia baiealica (Muslov) Krylov, from the Cluntuy Suite of the Pri- baikalye [based on Collenia baicalice Maslov 19374, 287]. Diagnosix: Tuberous, bumpy, swelling and von- stricting. parallel to markedly divergent branch- ing columns, genetally withour wall, with fre- quent overhanging laminae. Lamination is dis- tinctly banded. Coment; B, baicalica (Maslov) Krylov, &, Kirgiste Krylov, B. rata Semikhatav. &. unea Semikhatov, B, prima Semikhatov. B. ampla Semikhatov, A, meilensis Nuzhnov, 8. maica Nuzhnov, 8B. ainica Nuzhnov, B- minnie Komar, 8. eapricornia Walter and B. burra fF. nov. Age: Middle Riphean to carly Late Riphean. Baicelia barra f. nov. FIGS. 4, 5, 6, 13, 14, LSa-c fietcalit spp. Glacssner, Preiss & Walter 1969-1056. Marerial: Thirtythree specimens [rom ten localities. Holotype: S222 (Figs. 3a & d), Skillogalee Dolomite 3.2 km west of Yatina, Southern Flinders Ranges, Nanie: From the Burra Group in which the stromatolites occur. Diaginus: Baicalia vith moderately frequent slightly to markedly divergent branching, irte- gular, coalescing columns with highly variabJe lamina shape and continuous, distinctly bandetl microstructure, Description Mode of Occurrence: Two modes of ocvur- tences have been noted: biostromal and bhio- hermal. the latter occurring only at one locality SOUIH AUSTRALIAN STROMATOLITES $] (Yatina), Hiustromes vary in thickness Jrom 0.3 ta 2m, the stromutolites being eventy dis- tributed throwehoul their extent; they have been followed for |00 mm or more, without lensing, before the outcrop disappears under soil cover. Biostromes are frequently interbedded in green shales (e.g. Myrtle Springs, Willouran Ranges), platy dulomiles (e.g, Arkaroola, Worumba) or massive dolomites (e.g. Burra). The bicherms at Yatina afe restricted to (wo thin beds; they are small lenticulac stromatolitic mounds, approximately 20 to 30 em thick and up ta 1m wide (Fig 130), interbedded with and surrounded laterally by platy and shaly dark grey dolomites, The overlying sediment is draped over the mounds, showing that the stromalolites had at least 10 cm relief over the surrounding surface. Colunins arise from sub- strates in several ways: (1) Flat-laminatesd slromufelite passes gradually up inte undula- tory and pscudocoluimnac stromatolites, then Info discrete, vertical to inclined columna, often with steeply domed lieminac fe.g, Burr, West Mount Hut); (2) Colurnns urise directly fram eroded surfaces of laminated or intractastic dolumites (e.g, Yatina, Fig. 44))3 (3) Columns arise from flat-laminated stromatolite wine broad cumuli (e.g. West Mount Hut), The degrees of discreteness of columns varies greatly; in some beck. columns are almost immediately bridged over by Jaterally linked stromatolites, but usually columns remain discrete for 20 lo 30 cm. In some arcas new sets of columns may arise from pscudocolumns. The upper surfaces of biosuwomes vury From flat (¢,g, in the Wil- louran Ranges, Burra) to broadly undulating (e.g. Worumba). Column Shape end Arrangement: Columns are tubctous, varving From subeylindrical ta irregu- lar, with round, oval and irregular cross sec tions (Figs. 4, 5, 6), Elongated or Aattened colunms are varionsly onented. The diamerer of colarns varies Erom J to 10 ¢m, most com- monly 3-5 em, with rapid swellings and con- striclions, Coltinins aré 2-15 cm high between branches, Some but not all columns are con- stnicted at the point of branching (Figs. 4c, f; Sc, d). The onientation of columns varies greauly From vertical to inclined, and is some- times sub-horizontal for short distances (Figs, 4e, 5)). Column axes vary from. straight to strongly curved. In some specimens, the upper- most columns swell matkedly upwards und be- come bridged over by laterally linked stroma- tolites. Adjoining columas coalesce very fre- quently, even in the discrete portions, but speci- mens from Burra show the least coalescing and bridging. In the Willogran Ranges, column arowth is frequently mterrupted by penecon- [éemporaneous erosion; columns may grow over hroken-off fragments of eurlier columns, con- tribuung to the irregularity of the structure. Branching: The most common form of hranch- ing is moderately divergent (Fig. 4a, i, 6F) though same sub-parallel branching (Fig, 4e, fg, m) and some very markedly divergent branching occtirs (Figs. 4a, Sd & 7)- In same specimens several] branches arise from nearly one point (Fig. 4a). Branching is moderately frequent, the length of column between branches commonly being only a few centi- metres; bed at any one point of branching it is usually dichotomous of less offen trichoto- mous, In some specimens branches arise at a high angle to the main columns, and then tumn shurply upwards, Some columns arise from the side of a main column (Fig. 14d). Great varia- tion is seen even in single outcrops, Murgin Srreciure: The lateral surface varies fromm smooth to very irregular, laminae ap- proaching the column margins at Yaridus angles. Some specimens have very patchy walls, while the intervening unwalled areas are smooth or only slightly fringed with smull peaks and cornices, for example those from Burra (Fig. 14d), Yatina (Fig. 4a}, River Broughton (Fig. Sc), Arkaroola (Fig. Sd, ¢). Willouran Ranges specimens contin both smooth and highly irregular edges, with large overhanging peaks composed of one of niore laminae (Fig, 6b & c}_ Frequently large swell- joys are composed of numerous Jaminae over- hanging a constricted portion of g column (Fig. 4¢), Bridges between columns are espe- cially common ncar the tops and bolioms of biostromes (Fig, te), Lamina Shape: The lamina shape is most com- monly genily convex, but varies in single speci- mens from very gently convex to pearly cons cil; nutny laminae jire sleeply convex, Micro- unconformities are espectally prominent in specimens from the Willouriun Ranges, bul occur ta some extent in alf areas. In places, branching cammences upon a partly eroiled column surface (Fig, 15b). Fig. $d illustrates the more commenly occurring lamina shapes; 92% of lamina have h/d between 0.1 and 0.6, the mode being h/d between 0.3 and 04 (28% 1 (Fig. 9d). Generally, the widest columns have the most gently convex laminae, while strongly elongated columns hive laminue gently convex in the sectinn parallel to the long eo W. V. PREISS Fig.4 Fig. 4. Baiculia burre. Skillogalee Dolomite. Burra Group, Southern, Flinders Ranges: (a) & (c)— Holotype. S222, 3.2 km W, of Vatinn: (b)—S218. same locality; (d) SIS{, 13 km S.W, of Werumba H.S. (e) & (f) S151. same Jocalityy ¢g), ( & (()—S221, Dutton’s Trourh H.S,, i¢ km S. of Burm; (h) & (1) —S314, same locality; (kJ—S534. sume Jocality; (o1)—float specimen, River Broughton, W, of Spalding. as. 0 Baicalia burra, Skilogalce Dolomite, Burra Group: ().—S333, Dutton’s Trough H.S., 14 km S, of Burra; (4)—S534, same locality, (cj S383, River Broughton, W. of Spalding; (d)— §456, 6.4 km S, of Arkaroola; (¢)—S457, same locality: (f)—S491, 2-4 km E. of Myrtle Springs H-S. (upper member of Skillogslee Dolomite); (g)—S489, same locality; (h)— $490, same locality: ((}—S488, 1.4 km E. of Myrile Springs HS. (lower member of Skillo- ealce Dolomite); (j)—S487. same locality; (k)—S319, the Avoodale Mine, Lyndhurst (col- lected by Mr, P. J. Binks); (1)—S302, West Mount Hot, Willouran Ranges; {m)—S99, same locality (collected by Mr. C, R. Dalgarno): (m)—S97. near Chintapanna Well, Willouran Ranges (collected by Mr. C. R. Dalgarno). Pig, #- Baicalla burra, Skillogalee Dolomite, Burra Group, north-western part of the Adelside Geo- synctine: (a), (b) & (f)—S96, Chintapanna Well, Willouran Ranges (collected by Mr. C. R. Dalgarno}; (c)—S98, West Mount Hut, Willouran Ranges (collected by Mr C. R. Dulgarna); (t)—S496, 4.8 km W, of Copley; (2)—S301_ West Mount Hut, Willouran Ranges, SOUTH AUSTRALIAN STROMATOLITES Ha axis. nd steeply convex ut right angles to it, Rarely co laminue turn over sharply and thin al the column margins, to form a wall. Geoe- rally, where a patchy wall Js present, ii is formed by the edges of steeply convex or paru- holic laminae (Fig. 14c), Frequently, laminae develop two e¢rests, anticipating branching immediately above (Fig. 4a, m). On a smaller scale, lamina shupe varies from smooth and regularly curved to slightly wavy, with discon- tinuous curvature and sharp crests. Both types occur in single specimens (Fig. 14). Microstructure: The microstructures and tex- tures Ohserved tn the clifferent areas vary con- sidvrably depending oo the deyree of recrystal- lization. In the best preserved specimens, the layering comprises alternating relatively thick, contmuous, very distinct. light and dark laminae, giving a banded appearance. Some are single homogeneous thick layers, while others are macrolaminac consisting of several very thin light-dark Jamination pairs, Most commonly single laminae traverse the whole eolunia width, excepr where cul by micra- unconformities. Onids or other detrital grains may be included in the laminae, Upper and lower boundaries of laminae are usually smooth and even, sumetimes wavy or broadly wrinkled, bul always more or jess parallel. Fxeéeptians occur only where erosional scour has taken place during growth, Rarely, lenti- colar swelhngs occur. Aight Javinae vary in thickness from .02 ro OS mm, very rarely to 1.0 mo Mast light laminae thin towards the column edges, but rarely Jens out. In the best preserved specimens, the sparry dolomite forn- ing them is ineqnigranular, xenotopic, and of vrain size ranging from ().005 to 0,06 mm. With greater recrystallizatlon an equigranular mosaic of 0,05 to 0.2 mm grain size results fee. Burra), The light laminae usually have sharp and smooth upper boundaries, hut some- limes grade down into grumous textured lami- hac, consisting of irregular and interconnected mivritic patches vp to 0.1 mm diam,, set in xenolopice =equidimensional sparry dolomite wilh a grain size of about 0.01 to 0.03 mim, ic. partially recrystallized’ dark laminac. In some specimens (2.g. Yatina, West Mount Hut, Worumba}, the light [aminae contain detriral granules. including small fiat inteaclysts, up to 0.5 mm tong, and rare onids up to 0.3 mm in diameter, Overlying laminae are draped over the larger detrital grains, Laminae in the Cop- ley specimen may be pelletal (Fig. !4e}. Dart faminae occur either singly, alternating with light Jaminae, or in dark macrolaminae, Thin dack laminae are commonly 0.04 lo 0.3 mm thick, but macrolaminae range up ta 2.5 mm in thickness, generally constant across the column, or thinning slightly towards the mar- gins. They are either continuous, or consist of a-series of aligned lenses, each up to 0.2 mm long. In well preserved specimens the dark laminae have smooth, sharp, parallel boun- daries; rarely, single laminae may be wrinkled, suggesting intrafarmational crumpling during growth. Well preserved dark luminac consist of dense, brownish-pigmented xenotapic dolomite, of equidimensional grains 0.003 to 0.0L mm dism,, but vertical and lateral gradations from unaltered to erumous textures are commen. Where dark laminae are grouped into miacto- laminae, they alternute with very thin, discon- tinuous light laminae, and {requently fuse to form solid. thick dark Taminac. Interspaces, A tow specimens have interspaces filled predominantly with bedded dolomite mnd (e.g. Burra), but generally the sediment is unhedded intraspucite or oosparite, less com- monly intramicrite, Frequently, intraclasts are derived from the erosion of stromatolitic columns; in places, 9 large fragment torn from a column has acted us a base for new growth. Tntraclasts ate flat to gently curved tabular dolomite pebbles up to 3 cm long, | lo 2 mm thick, and only slightly rounded. Many are Fragile and could have survived very litle transpor!, They contiin the typical internal laminations of the associated stromatolites, and are probably derived directly from them. Gcca- sional flat pebbles stand vertically, bur ene- rally they lie fiat or imbricated, Ooids yary in shape from round to oval, 0.2-[.0 mm diam, und consist of one dark-rimmed sparry Liver coating a micritic core, or Jess commonly. sevetal sparry layers. Most commonly, allo- chems are closely packed and cetnened by a clear, sparry dolomite cement, Some specimens contain significant amounts of dolomite mud, variously recrystallized. forming a macrex be- tween allochema; in these cases the seciment is poorly laminated. Secondary Alteration; Secandury alteration has extensively modified the textures and often the microstrictures af stromatolites from many drews. The following four stages of alteration miy be fecognized: (1) Peneconemporaneous. The fact that dolo- mite consistently constitutes the whole rock to the exclusion of calcite, while still preserving fine structures, suggests very early dolomitiza- 86 W. V. PREISS tion, during the growth of the stromatolites, It is also possible that penecontemporaneously Uolomitized lime muds were reworked and trapped in the algal mats. During growth, cro- sion by strong currents scoured the living sur- faces of columns, creating micro-unconformi- ties. In some specimens (e.g, West Mount Hut), laminae may be separated by lenticular vughs, later filled with sparry dolomite (Fig. 12c), These voids were probably formed by arching up of laminae, perhaps due to lateral expansion in growth of the algal mats building the siromatolites, of 1 partial desiccation. (2) Early Diaveneric. Black chert very com- monly replaces porlions of slromatolites and interspace sediments, Sometimes dark laminae are preferentially silicified, perhups during growth, but more commonly, silicification post- dites the growth of the colamns (e.g, one side of a column may be replaced). In places, silt- cified laminae are brokea by minute dolomite filled cracks, (3) Late Disyvenetic. Dark Jaminae and macro- laminae may be recrystallized tay grumous tex- lures, Consisting of patches of dark, dense micritic dolomite (remnants. of the original cutbonute) varying greatly in size from 0.005 to 0.1 mm, set in a matrix of xenotupic sparry dolomite, of cquidimensional grains ranging in size from 0.01 to 0.03 mm. Light laminae are commonty slightly recrystallized and sparry, consisoug uf hypidiotopic to idiotopic. equi- dimensionul dulomite grains of similar size to (hove of the sparry matrix of the grumous tex- lures, Coarsely recrystallized laminae also occur, i, places Cutting ucross. the fine stmuec- ture of primary laminar and corroiling their boundaries, They consist of idiotopic tran- sparent. dalomele of grain size up to 0.1 nun. (4) Yeerwnte. The only specimens affected by tectonic deformation are from the Burra region, Here columns are slightly Nuttened and laminae ate erenulated along a slight tectonic foliation, These are alsa the must highly meta- morphuscadl, cisplaying the greatest degree of recrvstallization, Tensional joints filled with toursely Crystalline Uolumile are commnan it most areas. Conpartrons The stramatolites are assigned to the group Buicalia on the basis of their taberous, swell- ing and constricting, humpy, variously oriented columns general absence of wall, numerous overhanging peaks and short cornices, and gencrally divergent branching. Some specimens have horizontal colomns for short distances fesombling Fusigussia, but are distinguished by the absence of the multiple horizontal branch- ing characteristic of Tunpussia, and hy their generally oainre mgged column margins. Baicetia busra is distinguished from B. printa Semikhatov, 8. aimica Nuzhnov, and 8. cupri- cornia’ Walter, by fits Frequently divergent branching and general complexity of columns, and from &, mimeéa Kormar by its larger size and more complex structure, Some specimens resemble &. baicalir (Mastov) Krylov, but most have more inclined and irregular columns. 8. lacera, B. rare, B. ample ans B. wit Semi- khatoy are not adequately ijlustrated for reli- yble comparison, and the illustrated micro- structures are badly altered; single specimens of B. burre may show microstructures similar to B, wnea, B. lacera, and especially the pelletal laminae of B. rare, Some specimens have long overhanging peaks and thus resemble A, ingilenvis Nuzhnov, but are distingunshed by more frequent and divergent branching. &. hurra most closely resembles B. rare Semi- Khatey and 8, maica Nuzhooy) it is distin- suished from &. rara in that neither pellectal tuminse nor knee-shaped bends in columns ure consistently developed, and from B. meied by its more inregwlar and coalescing columns, and its more continuous kiminae, Disteibution: Widespread in the Skillogalee Nolomite, Burra Group: Yutton’s Vrough H.S., 16 km south of Burra; Scrubby Raoge, 27 km south of Burra; 3 km west of Yatinat River Broughton, 8 km west of Spatding: 11 km south-west of Worumba; 11 kn south of Arkaroola; 3 km west of Copley; 3 km east of Myrtle Spnngs HLS. near Leigh Creck: West Mount Hut, 27 km west of Witchelina H.S. wod Chintapanna Well, abouc If kny west of Witchelina DLS. Possible 8. burr oceyrs also in the Skillogalee Dolo- nite, Depot Creek, but chesé have not been studied in detail. Specimens from possible River Wakefield Group, Carnieton (Fig. 15c) ate inadequate for identification, but are possibly to he inchided (BURRA, ORRO- ROO, PARACHILN’. COPLEY. ANDA- MOOKA and CURDIMURKA 1:250.000 map sheet areas), Age. Early Adetaidean, correlated with the younges| Middle Riphean of the USSR- Group BOXONIA Korolyuk Boxonia Korolyuk 1960-129 Komar 19Af: 7¥. Cloud & Semikhatov 1969:1036- Glaess- ner, Proiss & Walter 1969: 1056 SOUTH AUSTRALIAN STROMATOLITES 87 Type Ferm: Boxenia grorilis Koarolyuk. from the Bokson Suite, Eastern Sayan. Diagnosis; Straight, subevlindrical columns with moderately frequent g- to @-patullel branching and smooth, walled margin strac- ture, Content; B. gracilis Keorolyuk, 8B, lissa Komar, B. krasivica Golovanoy. B. allab- jueice Komar & Semikhatoyv, B. ingilice Komar & Semikhatov, B, hiunca Raaben and B. perruknarre Walter. Raaben {1969a) places B. erumulosa Komar into partial synonymy with & gracilis Koro. lyuk, &, diverata Sidorov has only 2 patchy wall and may therefore be exchided- The South Australian form is Bexenia melroxa. Age; Late Riphean and Vendian. Boxonia melrosa f nav, FIGS. Fah. |5d-f Meaverial: Four specimens from one locality. Nelotype: S503 [| Figs. 7b, e & dy 15e), 16 km west of Melrose township, Southern Flinders Ranges, Nunes After the type locality. Diagnesix: Baxania with long, narrow, ¢lasely spaced columns, a- and A-parallel braiching, without very broad basal columns, with occa- sional rounded projections, and with indis- tinctly banded. muderitely convex, Ipminae lacking pelletal microstructure. Description Mode of Occurrence: ‘The stromatolites are relatively poorly exposed in a faulted area, so that relationships are not clear, At least two bioherms occur, preserved as grey or pale buff dolomite. ‘I'he beds are overturned, dipping south at about 40°, The narrow, parallel columns arise difectly from futetally linked stromatolites. partly pseudocolumnar, the base of which is not exposed. The overlying columnar portion ts approximately 6 m thick and constats of vertical columns neal the centre of the bioherm, and inclined colunms at the mareios, where they pass laterally into pseude- columnar stromatoliies. Columns are overlain hy wavy laminated stromatolites, which cover the whole bioherm, Bioherms are of cumplate shape, broadly domed, up to 60 m long, and are surrounded by flut-bedded dolomite, Column Shape and Arrangement: Columns are straight, erect, subcylindrical. smooth to gently bumpy, with circular or slightly lobale, rounded polygonal cross-sections, 1-5 cm diam. (Pig. 7a-h). The diameter of 2 single column generally remains constant throughout its length. Columns may reach a length of up to 20 cm between branches, bul some columns are only a few centimetres high, occasionally in the form of rounded projections Branehing: Branching varies from a- to f- parallel; y-purallel branching is tate (Fig. 7h). Commonly a 3-5 cm column divides into two or three narrower, parallel, very closely spaced columns, I-2 em diam, (Fig. 7e,d,f), Ooca- sionally, two narrow columns may coalesce {Fiv, 7c). Not all branches develop into long columns; some terminate their growth only a few centimetres above branching (Fig, 7d), Mirgin Structure: The lateral surface is even, smooth ar with low, broad bumps, ap to several centimetres wide. with a relief of I—5 mim. Peaks anc cornices are entirely absenr, bur very rarely bridges up to 1 cm thick occur between adjacent columns. A multisaminate wall is almost ubiquitous, At the margins of columns Jaminae are poorly preserved, bur in plices up to 10 laminae may be seen to com- prise the wall. Single laminae gencrally extend for au distance of 1-2 em down the column margin (Figs. |Sd—f). Camina Shape; Laminae are most commonty moderately convex, hemispherical, in places approaching rectangular (Pig. 8e). Frequently they are slightly asymmetrical, especially in in- clined columns, Belore branching, lammae usually develop two crests, The degree of con- vexity, h/d, is moderately conslunt, even in columns of differing widths, Of laminac mea- sured, 91% have h/d between 0.3 und 0.7, the mode (39%) being 0,5-0.6 (Fig, 9¢). The shape of crests varies from tightly urenate to gently rounded (Iig. Se). Most laniinae are broadly wavy (wavelength up to 8 mm, ampli- tude 1-2 mm) but not wrinkled, Microsinneture: Microstructure is poorly pre- served in both pale and dark specimens; laminac are broadly continuous, with smooth, patullel upper and lower boundaries, but niwy he broken into a series of clots and lenses by recrystallization, and even where their con- tinuiry is preserved, they are extensively em- bayed by recrystallized carbonate. Microstrac- ture is indistinctly banded with alernating darker and lighter laminac. Light jane vary in thickness from 0.08-0.4 mm, but usually thin towards column margins. Continuity és usually retained across a column, although the finest Jaminac: {frequently Jose their identity by 88 recrystallization, The laminae consist of tran- sparent, slightly incquigranular (of grain size 0.01-0,04 mm) cguidimensional dolomite of polygonal, hypidotopic fexture. Within this occur irregular 0.05-0.1 mm segregations of darker, greyish pigment, with no relation to grain boundaries. These ate apparently reni- nants of pigment left by partial recrystallization, W. ¥, PREISS as they may grade into more or Jess continuous laminae, Distinct round to oval pellets (as in Russian Boxonia) are ubsent. Dark laminae are less continuous, and often diffuse. Their thick- ness vaties from 0,08-0.3 mm; towards the margins they trequently thin or lens out com- pletely, and do not take part in the formution of the wall. (The Jayering in the wall is Fig.7 Fig: 7. Koxonia melrosa, Brighton Limestone equivalent, Umberatuna Group, 1.6 km W, of Melrose: (a), Ce), (2) & (h)—S502; (b), (ce) & (d)—Holotype, $503; (1)—S504. % a} ? Dy») fm Representative examples of lamina shape: (4)—Acuciella angepena: (b)—A. augusta: (cj— Acaciella f. indet.. (d)—Haicalia barra, (€)—Boxonia melrasa. SOUTH AUSTRALIAN STROMATOLITES ay extremely indistinct). In places, they lens out also within the central patt of a column, Dark laminse are composed of équidimensional, Renotopic, equigranular dolomite (of grain size ranging from 0,003—0.01 mm), and in places, ure disrupted into a series of irregular clots and lenses separated by sparry dolomite. Interspaces: Interspaces between columns are extremely narrow (usually Icss than 5 mnt), und are filled with partially recrystallized dola- mite mud, now largely of fincly grumous tex- ture, containing in places, round or ovoid clas- uc pellets, 0.2-0.7 mm diam. Much of the sedi- ment ts vaguely Jumingted, the laminae abutting against the walls of columns, which they post- date. Secondary Alteration: Stromatolite columns and interspaces consist of dolomite, considered to result from the replacement of original cal- cium carbonate, Most fine structure has been lost; dark laminae are outlined mainly by seg- regations of dark pigmented dolomite, but re- crystallization has partly embayed and partly obliterated the fine dark laminae. The irregular distribution, of pigment is due to recrystalliza- tion. In places, coarser, sparry laminae of grain size up to 0.08 mm occur, and may contain dismembered remnants of dark laminae. Stylo- lites are moderately frequent, and usually dis- cordant to the Jamination. In places they follow Fig.9 column margins for short. distances, removing the wall. Occasional thin dolomite veins follow the path of stylofites. Some stylolites are parallel to overall bedding, and displace column axes slightly (Fig. 7c). Comparisons The stromatolites are assigned to the group Boxonia on the basis of their long, smooth walled columns with moderately frequent a- and B-parallel brunching. Katavia Krylov and Acaciella Walter have similar gross structure; Katavia is distinguished by its very prominent humps, while Acaciella generally lacks a wall. Minjaria Krylov also has parallel straight columns but is distinguished by its less frequent branching. Most other described forms of Boxonia have well defined pelletal microstruc- tures; forms are largely distingutshed on the basis of the size of the pellets, A specimen of B, gracilis sent by M. A. Semikhatoy and I. N, Krylov, has pellets consisting of rounded car- bonate grains with dark, fine-grained rims. These are absent in B. mielrosa, which also has less wrinkled laminae. 8. melrosa is distin- guished from B&. ingilica Komar & Semikhatov by its ubiquitous wall and straight columns: B. allahjunica Komar & Semikhatov apparently has some complex branching, B, lisse Komar, B. gracilis Korolyuk, B. grumulosa Komar, B. bianca Raaben and B. krasivicu Golovanoy Histograms of Jamina convexities. The convexity of a lamina is ihe ratio of the height of that lamina to its diameter (h/d). Histograms are plotted for each stromatolite form at inter- vals of 0.1; # is the number of measurements made for each form: (a)—Acaciella angepena: (b)—A. augusta; (c)—Acaciella £. indet.: (d)—Baicalia burra; (e)—Boxonia metrose. 9D W. V. PREISS may all be synonymous. & melrose is distin- guished from 2B. pertaknurra Walter (in press), which also. lacks a pelletal microstructure, by its more steeply convex laminae, its occasional short, projection-like columns and by the absence of well defined broad basal columns. B. melraxa most resembles B. fissa, from which it is distinguished by the absence of pellctal microstructure, and by the presence of some short, projection-like columns. Distribution: Brighton Limestone equivalent, 1.6 km west of Melrose (ORROROO 1:250,000 map sheet areu). Age: Late Adelaidcan, correlated with the late Riphean of the USSR, Acknowledgements Tam indebted to Prof. M. F. Glavssner for supervising this study al the University of Ade- Inide, and to Dr. B. McGowran for his critical reading of the manuscript. Many of the con- cepts considered arose from discussions with ny colleague, Dr. M. R. Walter, now of Yale University. The figures were drafted by Mc. B. Frost, Department of Mines, and by my wife, The paper is published with the permission of the Director of Mines. References Arrken, J, D. (1967). Classification and environ- mental significance of cryptalgal limestones and dolomites, with illustrations from the Cambrian and Ordovician of southwestern Alberta. J. sedim. Petrol, 37, 1163-1178. ANDREWS, 8S, H. N. (1970).—Index of Generic Names. of fossil plants, 1820-1965. Bull. U/.S- geol, Surv. 1300, 1-354. Barnursr, RB. G. C, (1966).—Boring algae, migcite envelopes and fithifaction of mal- aca biosparites, Lpool Manchr geal. J. 5, BERTRAND, J, (1968)—DLes Guifices stromatoli- tiques. précambricns de la “Série & stroma- tolites” du Nord-Ouest de I" Ahugear (Sahara) Bull. Soc. véot. Fro 7010), 168-178. Brack, M, ((933)—The algal sediments of Andros Island, Bahamas, PAil. Trans. R. Sec., wer, A. 222, 165-192. BrapLey, W. H- (1929)—Algae reefs and oolites of the Green River Formation, Prof. Pap. US. geal. Surv. 154, 203-223. Croup, P. CE. 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(1937b).—O rasprostranenil kar- honatnykh vodornsley v vostochnay Sibiri: (On the distribution of the caleareous algae in eaeca Siberia.) Problemy Paleont, 2-3, 327- Masitov, V, BP (1938).—0) plirode Siromatolita Conepinten. (Om the nature of the sizoma- folite Conuphytor,\ Problemy Paleont. 4, 4329-331, Masinv, V. PL (1939a)—Popytka vozrastnogo opredelen|ya nemykh tolshch urala, pomo- shel’yu stromatolitov. (An attermpi of the age detennination of unfossiliferous beds of the Urals with the atd of stromatolites.) Problemy Puleane. S. 277-281, (Eng. transl 281-284.) Mastov, V. P, (1939%b).—Rod Collenix. (The genus Collenia) Problemy Paleont. 5, 297- Mascov, V, P, (1960).—Stromatoliy (ikh gene- zis, metckd izucheniya svyaz's Eutsiyami j geo logicheskoe znachenie na primere ordovika sibiraskoy plutformy). (Stromatotites; their genesis, methods of study, connection with facies and geologic significance in the example of the Ordovician of the Siberian Platform, ) Trudy. Geol. 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Akad, Nauk SYNR Trudy. 210, (-207. SnArovatova, I. G. (1968)—O novoy gruppe stromatolitoy Jacntophyion iz verkhnego pro- terozoya Vostochnogo skiona Aldanskoy onte- klizy. (On a new group of stromatolites Jacutophytan from the Upper Proterozuic of the ecastorn slopes of the Aldan antectize.) "Tektonika, strutigrafiya i litologiya osadach- nykh formatsiy Yakutii, Dokl. 17 nawch. Sevs. Yakur: Fil. Sibir. etd. Akad. Nank SSSR 97- 103. Steimmann, G, (1911).— Ober Gyrinasalen ram- sayi, eine Coclenterate von der Halbinsel Kanin, Ball, Sec. gewgr., Finland 31, 18-23- Thomson, B. P., Coats, R, P., Minams. R. C.. Fonnes, B. G.. Darcamno, CR, & Jores- 90N, J. E. (1964)—Precumbrian rock groups in the Adelaide Geosyneline. A new subdivi- sion, Quart. geal. Notes, geol, Jury. S. Anst, ' VoLoapin, A. G. (1962).—"Drevacyshie Yaodo- rosli SSSR". (The most ancient algae of the USSR.) (Ted. Akad, Nauk SSSR. Moscow.) Warcorr, C. BD, (1414).--Cambrian Geology and Patacontology Tl: Precambrian Algonkian algal Nora. Smithsun. misc. Collins, 64, 78-156. Warren, M. R. (1972).—Stromatolites amd the hiostratigraphy of the Australian Precambrian amd Cambrian. Palaeontelegy, Spec. Paper (in press), Youne, R. B. (19334).--Vhe occurrence of site- matolithic or wgal limestones in the Camp- bell Rand Seri¢s, Griqualant! West. Trans. geal, Sac. §, Afr. 35, 29-36. Yours, R. B. (1933b) —Conditicns of deposition of the Dolomite Series. Trans. yweol. Sov, 5, Afr. 36, 191-135, Yours, R, B, (1935). A compurson of certain siramatolichic rocks in the Dolomite Series uf South Africa, with modern algal sediments in the Bahumus. Trans. peal. Soo, ¥. Afr. 37, 153-167, Appendix 1.—Glossary Axis: The centre-line of a eoluoim, Bioheren) A circnmsctibed organo-sedimentary Structure whose minimum width is less than or ¢qual to One hundred times ity maximum thick- ness, embedded in rocks of different lithology. Note: the definitions of the terms Aloherm und Biostrome pve based on those piven by Nefsan, Brown & Brineman (1962) but since, at Jeast in stromatolites, the two. integrade, an arbitury limit must be set. Tabular bieberms have parallel upper and lower surfaces, while deemed bioheems have gently convex upper surfaces. Subspherical hivherms had the highest growth relief relative to their width, Toasuing dbioherme are bioherms which had little or no growth relief, and therefore inter- tongic at their margins with the surrounding sediment. Blastveme: A stratiform organo-scdimentary arie- tire whose minimum width is more than one hundred times ity thickness, Note; jo practice if is rarely possible to see the Uiee dimensional shape of the strueture in outcrop. The distinction helween, bivherms and biostromites must there fore be based on the dimensions visible in out- crop. Ef the outcrop is inadequate, the informal tern) “hed” is used. Tabular biestremes have parallel lower and upper surfuces. Domed biestromes ether may cunsist of jietaposed domed bioherms or may be continuous with jirxtapased domes on their upper surfaces. Branching? The’ division of a column into new, diserete coltimmns. The columns became discrete when they are first separated by an interspace. In parallel branching. the aves of the new columns ure parallel (most commonly they are also parallel to the axis of the original column). a-puradlel branching is parallel branching in which the widih of the individual remains con- stant, Io a-paralle? branching the original column wklens geadually before branching, while in y-parallel branching, it widens abruptly before braaching, In sliphrly divergent branching. the nxes of the new columms diverge at less than 45°. while in markedly divergent branchins they diverge al mure Man 45°. Dicharorioey branching (§ branching ips more than twer columns at approximately onc level, Bridge: A stronvatolitic lamina or set of laminne linking adjacent columns. Bump: A low, rounded pratrasion on the ade of a column. Coaleseing columas: Adjacent columns which joi und continue arowth as one column. Column; A discrete stromutolite structure, with the dimension in the direction of growth greater thum at least one of the transverse dimensions. Column shupe and arrangement offen vary ac- cording to the position in the bioterm, Columnur-layered strometelite: A stromatalite in which short columnar and laterally Unked fusually pseudocalumnar) portions ullernate. Cornice: Peripheral overhanging portion of a Jamina of set of laminae. elongated transversely to the column axis; Crest: The summit of an upward-convex Janina. Creéral line: The line jointing, the crests of succes- sive laminae Cresrat zene: The environs of the erestal line, In Conephyran, the crestal ome is specifically the yone of thickening and contortion of the laminae; the width of the crestal zone is the width of the thickened and/or contorted por- tions of Jaminac, Three types of crestal zones of Canophyion were distinguished by Komar et af. (1965), SOUTH AUSTRALIAN STROMATOLITES 93 Cumiulate stromatelite: A rounded protruding nan- columnar stromatolile. Domed: With approximately constant radius of curvature, Flat-laminated strometolite: Non-cotumnar stro- mutolite with flnt continuovs laminae. Aitken (1987) has proposed the term cryplalgalaminate for stromatolites with planar lamination. Gently convex lamina, A, lamina whose ratio a hejght to diameter iy less than or equal to 0,3, Meusuremenis of this. ratio are best treated statistically by plotting on a histogram. Grarled columa: A column with large bomps Grumois: A mineral texture in which fine-grained Palehes ave surrounded by coarser grains, inter- preted jo have formed by partial reerystulliza- lion. Nypidiotopic: A mineral texture intermediate be- tween xenotopic and idiotopic. Miotapie: A texture in which the mineral grains ure hounded hy crystal faces. Individual: A single discrete stromatolite within which either the laminac are continuous or which comprises @ proup of columns arising from ia single basal column. Tiderspace: The space beuveen columns, usually filled with sediment. Lamina: The smallest unit of layering in a stroma- tolite. Lanceolare: An clangate Lransverse section of 4 column. tapering at both ends, Laterally linked stromatalite: Stromatolite with wavy laminae which are continuous between crests. 4 Mutralamina: A distinct set of laminae, Microstructure: Uhe fine-scale structure of the stromaltolite Jamination, in particular the dis- linciness, continuity, thickness and composition of the laminae. Banded microstrucnire is characterized by very continuens laminae with sharp, distinct, more or less parallel boundaries. In. streaky mticrosirue- ture less distinct and continuous laminae fre- guently grade into one another. The darker faminae are usually more distinct. Sttiated = micrystruciure consists of primary chains of Jonses, oriented parallel to the lamina. tion (this excludes cases where originally con- tinuous faminae are disrupted by recrystalliza- tion). Fermiform microstricture consists of narrow, Sinniows, pale calaured areas (uswally of sparry carbonate) surrounded by darker, usually finer grained areas. Micra-unconformity: Surface of lamination dis- cordance due to penecontemporancous erosion within a stromatolite. Niche; A deep indentation in the side of a colomn, Parabolic lamina; A lamina whose axial longitu- dinal section approximates a parabola. Peak: Overhanging portian of a lamina or set of laminae with a small dimension transverse to the column, Pellet; Ovoid to sub-ovoid micritic carbonate grain of silt or sand size, lacking internal structure Piginent: Organic or inorganic colouring matter. Plary celienn) A strongly transversely elongated eoluimn. Projection: A small columnar or conical out- erowth from the side of a column. Psenducelumnar: stramatolite: laterally linked stromatolite in which Successive crests are superimposed, forming calumn-like structures Cpsendocolumns), Rectangular lamina; Lamina which in a longitu- dinal section of a column js flal-topped with edges deflexed at about 90°. Rhombie lemina; Lamina which in a longitudinal section of a column is flat-topped but has sub- parallel edges not perpendicular to the top. Rib+ A low, rounded protrusion which ts elonguted transversely to the column on which il ozcurs, Sefvage: An unlaminated coating on colum mur- zing, Possible explanations for this include (a) micritization by algal boring: (b) inorganic pre- cipitation of lime; (c) a thin algal film an column margins during growth. In some forms 4 selvageé-like structure is probably ihe result of differential recrystallization of a wall, Steeply Convex latina: A lamina whose ratio of height to diameter is greater than 0.5. Tuberous coliana, A coltimn with prominent ex- pansions and constrictions, Wil: Structure at the margin, of a column formed by one or more laminae from within the colunin bending down and coating the margin for at least a short distance Wavy lamina: A lamina with flesures of wave- length greater than 2 mm. Wrinkled lamina: A lamina with flexures of wave- length greater than 2 mm. Undulatory xtremurolite: Lateralty linked Stroma- tolite in which successive crests are not super- imposed, Xenolapic texture: A texture in which the mineral grains are anhedral or irregularly shaped, ie. net bounded by crystal faces, 4 Fig. 10, Fig. 11, Figs 12 Fig, 13. Pig. 14. Fig. 15, W. ¥. PREISS Acaivila angepena, Irom Lower Cambrian limestones, Flinders Ranges: sections perpendicu- liar to bedding. showing mode of occurrence and microstructures; (a) —Marginal section of a bioherm, Note thut the laminae are completely recurved wnver the biohetm evige. The speci- men is in sifu. Ube ball-point pen is 16 cm Tong, Angepena; (b)—Etched section of Sab, the recurved margin of the bioherm in (a) cur at right angles (o bedding, Nate that here growth partly proceeded downwards, Specimen is 15 cm wide; it was collected from the surtcrop shown in (ay; (co) —Laleral termination of a bioherm, which partly sank into the soft substrate during growth, The white areas are dolomitized. Width of specimen (S460) is 20 cm. Ange- pent; (d)—Pseudocolumns with rare interspuces, Note the domed Liminae grown upon partly Kiril intvaclasts, and the extremely continuaus lamination, Uhin section. Angepena (8462); (ej—Evenly laminated ferruginous structure, probably the stromatolite A. aneepena affected by secondary ferruginization, Thin section, from near Old Wirrealpa, The durk laminae are outlined by finely disseminated hematile. (S364, collected by Mr. P. G. Haslett), (f)—Dyenly lanimated discrete columnar form from halowie Gorge (Sir Douglas Mawson's specimen), (a j—Arociely dagepenc; irregular coluoins freny ihe marginal portion of @ small bioherm 15458 Anwepena), Note the vermiform microstructure within parts of columns, here inter- reted us duc to algal boring, disrupting the normally very even, continuous lamination; (b) » de)—aAeaciella f, indet, Both specimens are erraticy from the Storiian glacials north of the Enorama Dispir. Thin seaions. Note the very numerous concordant stylolites in (bh). (S539 and $509 respectively; S$39 was collected by Dr. B. Daily); (dl. Ce) & CE) —Aceetella wngustu, Brighton Limestone equivalent, Depot Creek, Vertical sections showing mode of Ocuurrence aml microstructure; (dJ—-Details of transition from broad, frequently bridged basal colunmms to upper nurrow, discrete columns. Broad coltimns in lower right-hand corner five inclined margins and subhoriznntal laminae: tel--Lenticular open spaces between uminue, possibly representing origina) gay vesicles (S163). (f)—Portion of a bioherm show- ing the inlercalytion of columnar snd laterally linked siromatolites. Avueiella wigusta, Brighton Limestone: equivalent, Depat Creek, showing mode of wocurretice and ticrosiructures. (9)—Margin of a bioherm (pale colonsed at right of photograph) inter. tonguing laterally with massive ooypuciie (at left); (hb) & (c)—S4U4 & S401 respectively. The gross shape and branching of columns, The interspuces ure filled wit inteclayered micrite and intramicrite, In U5 (o L.Weem bands. te) Is natural size. In (b), laminae become daubly Crestcd before branching, but in the centre of the photograph (c) is an example of a short interspace beiween crests bridged by the overlying laminae: the voltimn then resumes ils former growth pattern; (d)—Recryslallized specimen from Mundallio Creek (3538). illustrating ra- isting recrystallized acicvlor textiires im the Lower part of the pholagraph. Ralrwia Narra, Skillogalee Dolomite; Sections perpendicular to bedding. showing the mode of focurrence and microstructure: {a)—Small lenticular biohermes interbedded in thinly bedded dolomites, Yalina; (b)—Portion of a biostrome interbedded in massive, fine eiained dolo- mites, Dutton’s Trough H.S. Longitudinal section of partially: silicified columns. The section is parallel to the tectonic cleavage, in the plane of Mattening of the columns; (c)—trregular eolumnas with numerous mickruncontormitics and highly variable lamina shape, West Mount Hur td)—Moderarely divergent branching columns, with some pelletsl laminde, Thin séc- tion, Yutina ($222, holotype); (c)—Stightly divergent branching in regilar, sub-cylindrieal collimns. Thin section, $533, Duiton's Trough HS, the specimen is taken from {he biostrome shown in Fig. 13 {h)} Reivwlia barra, Skillogalee Dolomite: (a)—Tuberouvs and inclined columns with evenly handed microstructare and high-angle micro-unconformities, Thin section, S487, Myrtle Springs; (b) BR. barra with minor pelleial laminae. Thin section, S150, Worumba, (c)—Sub- evlindrics) columns with steeply domed, evenly banded laminae, Thin section, $302, West Mount Hut: (d3—Branching of narrow columns from the sides of a main wide column, Cut slab, 5534, Dutton’s Trough H.S. The specimen is taken from the blastrame shown in Fig. 13 (6b): fev}—#, burre with predominantly pellets! luminue. Thin section, $496, Copley: (EJ —Conplex branching of columns [ram Arkaroola, Thin section, $457. (aj—Baicalio Suera with finely silicified laminue. Thin section, natural size, S151, Warumba. Note the vertical tectonic dolomite veins; (b)—#. burra. Cut slob tliustrating sub-parallel bronehing columns with high-angle micro-unconformilies and bunded tamination. 396, near Chintapanna Well. specimen collected by Mr. C. R. Dalgarno, Note the overgrown siroma- tolite fragment in the lower left quadrant, and he branch arising from an_eroded column ih the Upper right; fe}—Indcterminate siromatolite, possibly Baroulia barra, Thin section, $322, near Cartieion: (4), fe) & (£)—Boxonia melrose, Brighton Limestone equivalent, Melrose: fd)—Hand specimen iflustrating longitudinal sections of columns: (¢)—Thin section of halo- type. 5503, Che lamination is indistinctly banded, and becomes diffuse in the wall zone, (f) —Thin section illustrating Jamination and wall structure, S177, natural size, Note that the upper left and lower left corners of the thin section ure composed ot highly weathered rock, YS SOUTH AUSTRALIAN STROMATOLITES 10 “1G, "6 W. V. PREISS | 7 PON ™. ae a She SAAS a ; FIG. 11 97 SOUTH AUSTRALIAN STROMATOLITES 12 FIG. 98 W. V. PREISS SOUTH AUSTRALIAN STROMATOLITES 94 FIG. 14 PRETSS wv 100) abe te x Lj “a THE NEMATODE GENUS MAXVACHONZA (OXYURATA: COSMOCERCIDAE) IN AUSTRALIAN REPTILES AND FROGS BY PATRICIA M. MAWSON Summary The genus Maxvachonia Chabaud & Brygoo, 1960, previously known only from reptiles in Madagascar, is now recorded in Australia and New Guinea. New species described are M. chabaudi from 7 species of skinks, 1 species of gecko, and 1 species of snake (? from food); M. brygooi from 5 species of agamid lizards; and M. ewersi from a frog, Litoria nasuta, from New Guinea. M. ftindersi (Johnston & Mawson) [syn. Aplectana flindersi J. & M.], 1s recorded from 5 species of Australian frogs and one introduced species, Bufo marinus. The genus Austrocerca Inglis, 1968, is regarded as a synonym of Maxvachonia. THE NEMATODE GENUS MAXVACHONIA (OXYURATA: COSMOCERCIDAE) IN AUSTRALIAN REPTILES AND FROGS by Patricia M, Mawson* Summary The gétnus Maxvachonia Chabaud & Brygoo, 1960, previously kaown only from reptiles in Mada- Rascar, 8 now recorded in Australia and New Guinea. New species described are M. chabauedj Irom 7 species of skinks, | species of gecko, and 1 specics of snake (7 from food); M, brygeoi from 5 species of agamid lizards; and M. ewersi from a Frog. Litoria nasuta, from New Guinea. M_ flincersi (Johnston & Mawsun) [syn. Apleetana flinderst J. & M,), is recorded from 5 species of Australian frogs and -one introduced species, Bufo marines. Che genus Anstrocerca Inglis, (968, is regarded as a syno- nyin of Maxvachania. Introduction Maxvuchonia dimorphe Chabaud & Brygov (1960, p. 129) was first described from Chamaeleon pardalis, and tater also from C. australe (Chabaud, G. R. Caballero, & Brygao 1964, p. 846}, in both cases from iw small island. Nossi-Bé, about 20 km from the main- land of Madagascar. It has since been recorded from one chameleon and two otler specics of hzards (Zonesauras maxinus and Mebuie gravenhorviii) on Madagascar itself. (G. Cuballero 1963, p. 192.) Although the genus was net recognised until recently, and docs not appear to be common in any host species, it is surprisingly wide- spread. The seventeen species of Australian lizards from which Maxvachonia spp. are recorded in this paper belong to the families Scinctdae, Agamidae, and Gekkonidae, and they come from a wide geographical range. One collection was. made from the stomach of 4 snake. but as this also contained some semi- digested skinks, the snuke may not be a true host record. The gents is not confined to reptiles, A ustra- cercu Inghs (196%, p. 164) appears to be a synonym of Muxvachonia. Inglis recorded 4. finders? (lotision & Mawson) (syn. A plectana finders!) from three frog species in Western Australia, Tt has now been recognised from five mute frog species from various parts of Aus- tralia, and from a twad, Bufo marirtus, intro- duce? inte Queensland sugar cane fields in 1934, Another species is recorded from a frog from New Guinea, The males and females of Marvachenia spp. we very different in size, but the morphology of the anterior end is similar in the two sexes. Both males and females are easily distinguished from other cosmocercoid genera, the Female by the great distance of the anus from the pos- terior end of the body, znd by the shape of the eggs, und the tale by the shape of the guber- nuculum, which is very large and bears two prominent projections near its proximal end. The differentiation of species within the genus is rather more difficult. The presence or absence of lateta] alae on the anterior part of the body in the female appears to be a specific character. There is a wide varislion in the body length of the female within a species, although fully adult specimens from the same host ant- mal are usually about the sanic size, The ratio of the body length to that of the oesophagus varies considerably, possibly due at Jeast in part to the degree of contraction of the body in different collections. The ratio of body to tail length in the adult is more constant, and may be of specific significance. The em size is simi- lar in all specimens available, but there is some vanation in the shape of the projection on the egg shell and of the envelope which surrounds the egg in the vagina, and these appear to have specific. value. The male worms are rare compared with the female. so thal it is even harder to assess the specific value of any character. The body length and that of the ocsophagus are very similar among all the specimens examined. ‘There is some vuarigtion in the iengths of spi- cules and gubernaculum but even these vary almost as much between two specimens from the same host animal (in the only case where two males were found in one host) as among all the males collected in Australia. * Zoology Department. University of Adélaide. Adelaide, S$. Aust. 5000. Truns, R. Soc, S. Aust. Vol, 96, Part 2, 3! May 1972, 1bz PATRICIA. M. MAWSON On theve slender ctiterin, three species have heen distinguished from Australian reptiles, one (M. flinderné) fron Australian frogs, and onc from a New Guinea frog. The general body form is similar in all species, and agrees gene- rally with the descriptions of Chabaud & Brygoo (1960) and Chabaud et al (1964) and o£ Inglis (1968), Some additionul observations and westinguishing characters are noted under the species. Measurements are given in Tables lanst Hl. Pype specimens will be deposited in (he South Austtalian Museum, Maxvachunia chabaudi nap. FIGS. |-6 Hosts and lucalities: Murerhia {(Dumerit & Bibron}, iwpe host, Leriva hoacaimillii (Gray), Crenoras — leae (Boulenger), Psendonaja ? affints Ciiinther, all from Eyre Peninsula, S. Aust.; Crenofus labillurdlevi (Gray) from Pemberton, W. Aust; Afeviergis peronil (Fitzinger) from Pemberton and Esperance, W. Aust.; Spivrte- niorplius chxtralis (Gray) from Wilgarup, W. Ausut S. koscinsko’ (Kinghorn) from the New England district, MUS.W.; Egernia whirel (Lacépie) from Penola, S. Austy Phyllurus silit (Bory) from Kangaroo 1), 3. Aust, Must of these collections consist of female worms. adult and/or juvenile (i.e. with or Without embryonated eges). There arc five males, two from Crenotux leae, one from Leriste lineoecellatuy and one in cach of two H. pevonii, Tn these last two there were no Temules, and as this species of Maxvuchonic is separated from others by chiuructers of the female. the inclusion of the males js arbitrary. Alac ure present in both sexes. There are three lips, the inner border of each projecting as a cuticular lamella. The mouth is triangular, or triradiate, Euch Jip is strengthened by it chitinous bar, the three bars mecting to form a triangle around the anterior end of the buccal cavity, The short triungular buecal capsule rests against the anterior end of the eesophagus Three well-defined teeth project fram the oeso- phageul lining into a depression in the anterior end of the oesophagus. Female: Lateran] alge extend from the level of the nerve ring to about the mid body. The pos- terior end of the hody ends in a more or less distinct mucro, which is rugose, The vulva, a transverse slit, lies at about the level of the isthmus of the oesophugos. The two ovartes commence shortly in front of the anus, pass lineoocellara backwards nearly to the posterior end of the body where each enters a short oviduct, lead- ing to a slightly wider, sometimes almosr spherical, thicker-walled sectiun {? seminal receptacle) from which the uterus Ieuds for- ward. The two uteri pass forward sido by side. uniting to form the vagina at about » quarter of the body Jenpih Lrom the anterior end, or a little in front of this. Eggs in the anterior parts of the uteri cach contain a coiled Jarva The eyes are rouchly sphericul, slightly longer in the axis through the knob on the shell, In the vagina, where they are less crowded, they ire seen to be surrounded by 4 spongy or reticulate material which Lorms a loose envelope attached to the shell by, or iat, the apica) knob, mare or less open at the oppu- site pole (Fig, 5) «ind often trailing two ribbon- like pieces from the open end. This envelope was noted in the ariginal description of Mutva- chania dimorpha, Male: The lateral alae extend for most of the body length, from the Jevel of mid-oesophagus to shurily in front of the anus, The posterior end of the body is strongly curved ventrally. The gubernaculum is Jarge and heavily built, With q pair of lateral processes neat the proxi- mal end. The spicules are slender. well chiti- nised, sind blunt-tipped. The cloacal opening is on un clevation of the body wall. ‘The thirteen pairs of caudal papillac are arranged as shown in Pig. 6. The species is distinyuished irom M. di- morphea chiefly because of the presence of lateral alae in the female. The females ave all shorter, and the males about the same size, as those of ML dimaerpha bot the spicules and gttbernacuilum are larger, Muavachonia brygoot nap, PIGS, 7-10 Hosts ind localities: Amphibolurus. decresti (Dumeénil & Bibron), type bust; A. nracrelarns (Gray), both from Eyre Peninsula, S. Aust.; A. inermis (De Vis) izam Yuendunu, North- erm Territory; 4, muricatus (Shaw) and A, herbotay (Cuvier) from N-S.W~ Only femples have heen taken from these agamid lizards. All of them, however, differ from those from skinks in the absence of lateral alae. In other respects they are very similar. Althotigh this distinction is slight, it is con- stant Notwithstanding the fact that agamids and skinks aceurret in the same tocality in Hincks National Park an Eyre Peninsula, Afiex- vachowa spp. from the agamids were always Figs. 1-6. MAXVACHONIA IN AUSTRALIAN REPTILES AND FROGS 103 Maxvachonia chabaudi. Fig. 1.—Oesophageal region. Figs. 2, 3.—Lateral and en face views of head. to same scale. Fig, 4.—Posterior end of femalc. Fig. 5.—Egg. Fig. 6— Posterior end of male. 104 PATRICIA M. MAWSON without alae, while those from the skinks had alac. In view of this it is thought safer to regard the two groups as separate spccics, at least until more specimens, especially males, are found. Maxyachonia sp, Host and locality: Morethia taeniopleura, Mornington I, Gulf of Carpentaria. Only one female was collected from this host; it is very similar to females of M. chabaudi but the ratios of oesophagus and tail {o the body length differ markedly (Table 1). Maxvachonia filndersi (Johnson & Mawson) FIGS. 11-13 Aplectana flindersi Johnston & Mawson, 1941: 148, from Liroria ewingi (syn. Ayla jervisensix) from Kangaroo L, 8. Aust. Figs. 7-10. end of female. Fig. 10.—Egg. 200. um Ausirocerca flindersi (Jobnston & Mawson) Inglis, 1968: 165, from Literia coelo- rhyncha, Heleioporus barycrugus and A. psrammophilus, from W. Aust. Host and localities: Bujo marinus Linn. from Queensland: [.imnedynastes dorsalis (Gray) from Adelaide, $. Aust.; Heleioporus ittor- naatus Lee & Main, Litoria moorei Copland, L. adelaidensis. (Gtay) from near Perth, W. Aust.; L. caerulea (White) from Alice Springs, N.T. All the hosts listed above are new records for M. flindersi. The new male specimens agree closely with the earlier descriptions, both in size and appearance, but the females are dis- tinctly larger; even those from related hosts in Western Australia. Through the courtesy of Maxvachonia brygooi. Figs. 7, 8.—Lateral and en face views of head. Fig. 9—Posterior Figs. 11-13. M. flindersi. Fig. 11—Anterior end of female, Fig, 12.—Posterior end of female. Fig, 13.—Egg. MAXVACHIONIA IN AUSTRALIAN REPTILES AND FROGS LOS Dr, W. G. Inglis and of the Western Australian Museum it has been possible to compare all the known specimens, and no significant differ- ence other than size was observed. The details of the female reproductive system have now been studied, and these agree generally with the form in other species of the genus, The ovaries begin shortly in front of the anus, The eggs in the uteri are enclosed in the characteristic outer envelope, which in some specimens is very dark. The envelope is in the form of a bell atlached to the knob of the shell at its apex and open at the other end; from the open end come two long ribbons of material similar to that of the envelope. In one case an egg lying fast outside the body of the female was still attached by one of these ribbons, which passed into the vulva. Maxvachonia flindersi differs from MM. di- morpha in the presence of well developed lateral alae in the female, and from both M. dimorpha and M, ewingi (see below) in the shape of the seminal receptacle. The size of the spicules and gubernaculum vary greatly in the few male specimens known, but the gubernacu- Jum is always distinctly longer than the spicules, 14 BOG LTT E a I9s 60 Lin Maxyachonia ewersi 1.sp. FIGS, 14-20 Host and locality: Lireria nasuta (Gray) from Brown River, New Guinea. The material consists of three female and one male worms. The genera! body form is very similar to that of M. flindersi and other species of the genus; measurements are given in Table Hs The churacters distinguishing this species from M. flindersi are the following: 1. The oesophageal teeth are much smaller (Fig. 15). 2. The spicules are distinctly longer than the gubernaculum. 3. There are only two pairs of preanal papil- lae in the male. The other caudal papillae are arranged as in M. flindersi, 4. In the female the posterior end of the hody uppears rounded, becuuse the extreme tip is slightly withdrawn forming a dimple. 5, ‘The artaiigement of the reproductive organs in the female is slightly different. The ovaries start much further forward at about Pigs. 14.20, Maxvachonia ewersi. Fig. 14.—Oesophageal region, female. Fig. 15.—Lateral view of head, Fig. 16—Posterior end of female, Fig, {7.—Fge. Figs. 18, 19—Two views of apical extension of egy shell, to same scale, Fig. 20.—Posterior end of male. o. ovary; Tr, seminal receptacle; u, uterus, 106 two-thirds the body length from the head, and the seminal receptacle is not so much wider than the uterus and ovejector, 6. The shape and size of the eggs are different. The knob on the shell is shorter, and appears conical on one axis, but hroad and grooved on un axis at right angles to this; the egg itself is slightly flattened in this latter view. The coils of the larva lie in the PATRICIA M. MAWSON plane of the wider diameter. The envelope surrounding the egg is thinner than in other species, although it is dark in colour, and forms a bell, attached at knob end of the egg, similar to those of M. flindersi, but more definite in shape (in these specimens at Icast). In eggs furthest from the vulva (bul in the vagina) the mouth of the bell is open, but in those nearest to the vulva it is closed. Acknowledgements Several of the collections examined were made by Dr. John Hickman of the Zoology Department, University of Tasmania; speci- mens from Literia nasuta were sent by Dr, W. Ewers of the University of Papua and New Guinea. To both these helpers I am most grate- ful. 1 also wish to thank Dr, W, G. Inglis, then Direcior of the South Australian Museum, because I would not have examined. any frog material for Maxvachonia sp. had he not pointed out that his genus Austrucerca is 2 synonym of Maxvachonia. I am also indebted to officers of the South Australian Museum, Mr. M. Tyler, Hon. Asso- ciate in Herpetology, and Dr, T. Houston, Curator of Amphibia and Reptiles, for infor- mation on the nomenclature of the hosts. References CABALLERO, G. R. (1968)—Contributions @ la connaissance des nématodes da sauriens Mal- gaches. Annis. Parasit hum. comp. 43, 149- 200. CHazaup, A. E. & Bryaoo, EB. R. (1960). Neéma- todes parasites de caméléons malgaches. IL Mem, Inst. Scient. Madagascar A. 14, 125- 159, Cuasaup, A, E., & BryGoo, E. R. (1962)—Néma- todes patasites de caméléons malgaches. Deuxiéme Note. Annis. Parasit. hum. comp. 37, 569-602. Cuapaup, A. C., CABALLERO, G, R., & Brycoo, E. R. (1964).—Affinilés entre les genres Skrjabinelazia (Ascaridida Seuratoidea) et Maxvachonia (Ascaridida Cosmocercoidea). Bull, Mas, Nat. Hist. Nar., Patis 36, 844-848. Incuis, W. G. (1968)—Nematodes parasitic in Wester Australian frogs. Bull. Brit. Mus. nat. Hist. Zoology. 16, 163-183. Jounston, T. H., & Mawson, P. M. (1941).— Some nematodes from Kangaroo Island, S.A. Ree. S. Aust. Mus. 7, 146-148. 107 MAXVACHONIA IN AUSTRALIAN REPTILES AND FROGS —_—_————————_::.:.’?Rk axXKkeee eee se Os Cr-6'E obs’ v6 Vr-“e cr SE6E Ts-s'p Tr OE THE qsuay et /7 Ve OT Pe-v]e GT-ET c60 6C-TT Ses Tt Ceo £c-0'% T-Sh 97-6 (uw) [rey 082 °— 00s —O06P ose oso-des 009-005 001-009 009-07S OO8-O8E OOS—~O8E BATA — oss *— OCR-OSE “OLE Ove 0S¢—-00P 009-00P 09S—-0S+ OOS-OIP O£9-D0E OLEOLT ‘dxa— ost *— Occ-O1e OLT-OST Oot OOE-OFT —0zE- OLT-07e OLC 0¢7 O8C-)9T O8lVsI lu—Vv OTL 9OL L9I-s'Et £T-€'01 O'LT sTI-Ol GLI-9YCTL PFPRI-VEL ScI-OIl gol-c6 OLI-cOr yysua] -ydoseo/J O80T “OSL 022-009 006-008 ets 068-02L Oz6-OLS 0°6-068 088-008 086-025 008-07L sndeydosag 611 ‘8 6'C1-8'3 80I-€'°6 8 ETL OPI-GL Lel-Ltt = €0l-9"°6 ¢'sI-0'9 LS-ek (uw) misuse] OST ‘SE O9T ‘O9T OET uinjnoeitiaqn£) O€I OZ] = SZ ‘OZT ra | sopnords OLE ‘OOE 00d ‘007 Ofe ‘d-xe— — ‘sor 00£ “OLE svt iu—vy EL ‘E'S ors ‘T's 9 yidug] “s9Q/7 OLE ‘OSP =. ONS ‘OTS O8E sndeydoseg) _ = — = — — =) Lt PZ SZ'ET Sz (una) yysueT en” a “EN “MSN “NY "S I BNW 'S ‘ny '$ “ASN “SRY “M ‘yty S$ “sty 'S Aged] uo BUINIOPY ‘SI2UIO ‘soy adA yz ‘PIO suoseig yuys ayeusg soqaen SYULYS dnoaz soy ooskKg jy “ds Jy ipnpgpya ‘jw sataedg ‘Md Ul UaAIS a4v StuauiainsDelt ‘patDIS asnsayzo ssajun ‘sajiidad unyoisnp wo ‘dds Bmoyseaxey fo sjuausainsvapy } FTV PATRICIA M. MAWSON 108 pS SP-L'P 6'S-8'P Le-l? SPO Fr 0'S-6'F I's-S'F SS-SF Pe/1 EEE T e791 Ti-sT 2-8 SI-LT CZ-ET StH 1 (urur) [ey OFS-OSP OL9-OES OSs-OLP 079-085 069-009 069-0rS OTL-OLS BapnA— OTP-OZE OLS-00F OTP-OSE QOS-OSF OrS-OSP Oss-OtPr as¢s-OEr ‘dxa— O£E-OST 067-002 O1Z-00T 067—0¢7 062-087 OLT-O¢T OLZ-OET iu—vy CLI-L +I 6 EI-e'LT Vtl-s8 LsI-o bl rii-+'ol V'é1-6'8 cI-6'6 “qdose0/T 006-002 008-089 O¢SL-O0L O8L-07TL Oc8-O1L OL8-OSL 006089 sndeydosec £9I-O'TT V'TI-8°8 0'6-79 CM-CUl 06-18 86-L9 T'T1-6'8 (uw) ysue] ¢ OOl O81 0zT um[noedagney 07 OST oor aqnords 06 OET ‘d'xa— OL1 O06! iu—y 69 Z9 6's Ipsusy ‘seo /"} OSE ‘ O8E OLE sndvydoseo are a SET CZ — - “= (mu) Yue P pe ce ee SS —E NS EO Ee eS DUISOU DONT Snujinu ofig vajndaoa-J ss SisUapywjapy "T 19400 SHJOUAOUI SYDSLOP 4soy MUOTT sniodolajapy = Sajspuspouiwty 1 8? | SS See, eee ‘ON (peonporur) “PIO "TN “ysnly “AA ISN “AA “ISN'Y “AA ‘sny ‘'S AYTeIO] IT 1sdaMa "JA isdapuapl “Jr satoadg a ‘ur Ut UadIS alD Squaluainsvaue “PatvoIpUr asiam4ayjo ssaqun :sdo4f upyosisnp wos dds BIUOIeAXRIA, fo Siuatuadnsvay ¢ ATaVL THREE NEW SPECIES OF THE GENUS CLOACINA LINSTOW (NEMATODA: STRONGYLATA) FROM MACROPOD MARSUPIALS BY PATRICIA M. MAWSON Summary Three new species of Cloacina are described: C. mundayi from Macropus rufogriseus, from Tarraleah, Tasmania, characterised by the presence of a dorsal buccal tooth associated with the duct of the dorsal oesophageal gland; C. clarkae from M. eugenii, from Kangaroo Island, South Australia, characterised by the shape of the cephalic papillae and the structure of the oesophagus; C. edward,vi from M. bicolor, from Sunday 1, Victoria, characterised by the presence of oesophageal ‘plumes’, very short spicules and very short vagina. THREE NEW SPECIES OF THE GENUS CLOACINA LINSTUW (NEMATODA: STRONGYLATA) FROM MACROPOD MARSUPIALS by Patricla M, Mawson” Summary Three new species of Clog¢ina are described: ©. miadayt front Macrepus rifegrivens, Trom Vactaleah, Tasmania, characterised by the presence of a dorsal buccal tooth associated with the duct of the dorsul oesophageal gland: tC. clarkae from M. eugenii, from Kangaroo Island, South Australia, characterised by the shape of the cephalic papillae and the structure of the oesophagus; C. edwardsi from Af, hicéfeor, from Sunday U., Victoria, characterised by lhe presence of oesophageal ‘plumes’, very short spicules and very short vagina. Cloacina mundayi n.sp. FIGS. 1-7 Host and locality: Macropus rufogrised, from ‘Varcaleah, Tus. This species is u relutively short stout nema- lode. The material consists of four males and four females. The submedian papillae are small, with the distal segment much smaller than the proximal. The long threadlike cercival papillae are close to the anterior end. The shallow buccal ring is somewhat hexa- gonal in shape, and is uneven in depth, A snl! dorsal ocsophazeal tooth projects into the buccal cavity, and is traversed by a duct from the dorsal oosophageal gland, The oesophagus is cylindrical, widening only slightly at the pos- Tertor und. No teeth ure present in the lumen. The nerve ring is at about the mid-length of the oesophagus, and the excretory pore shortly behing it. The posterior end of the female tapers from in front of the vulva to the tip of the tail; the distance from the anus to the vulya is ubout equil to the tail length, The yiagina is slightly longer than the tail, The eggs are 110 by 65 phy The spicules aré ahout 1/2.5—-3,0 of the bady lcngth.. The dorsal lobe of the bursa is unusu- ally Jong for the genus. and the ventral lobes are united, The bursal rays are as shown in Figs. 6 and 7, The genital cone is short and conical, and on either side of it there is a cuticuJar mflation. No accessory cone can be seen, Measurements are given in Table 1, This species is distinguished from all others so far described in the shape of the bursa and in the presence of a dorsal tooth in the buccal capsule. In describing C. dahl’, Linstow (1897, p. 287) mentions the presence of a gland (seen in T\S,) in the dorsal wall of the oesophagus, with «a duct opening dorsally (presumably into the lumen of the oesophagus or into the mouth). Such a distinct gland has not been noted in descriptions of other species of the genus, nor in re-examination of fresh material of various species, Tl is not present in €, clarkue ov CG. edwerdsi. In the deseription of ©. daidi there ix no indication of a dorsal touth associated with the gland. Cloacina clarkae nsp- FIGS, 8-13 Host and locality; Muacropuy eudgenii, from Kangarou T This is a large worm from the stomach of the host. The submedian papillae are Jong and slender, and the distal segment of euch is dis- tinctly longer, but not wider, than the proximal one. The cuticle is thickened just behind the cervical grouve, The threudlike cervical papillae are Telatively close to the anterior end, The buecul ring is deep, its walls relatively. thin, and sloping outwards towards the anterior end; the unteriar muryin is lobed. The oesophagus is Jong and slender, except for a distinct terminal bulb, It is clearly divided mio four remions—(1) in the anterior half there are 9-11 distinctive equidistant places where the lining appears to he creased; (2) in the rest of the cylindrical part of the oeso- phagus the cuticle is more or less featureless; (3) just below the terminal bulb, the oeso- phagus and its lumen are slightly wider, and in this region about 8 well deyeloped teeth project * Deparment of Zoology, University of Adelaide. Adelaide. S. Aust. 5000. ‘Trans. R. Soe. 8. Aust. Vol. 96, Part 2, 31 May 1972. 110 PATRICIA M, MAWSON 50 j Figs, 1-7, Cloacina mundayi. Fig. 1.—Oesophageal region. Fig. 2.—Head, lateral view. Fig. 3— Head, en face, Fig. 4—T.S. anterior end shortly behind buccal ring. Fig, 5.—Posteriar end of female. Figs. 6. 7—lateral and ventral views. of bursa. Figs. 8-13. Cloacina clarkae, Fig. 8—Lateral yiew of oesophageal region. Fig. 9- Head, ventral views. Fig. 10.—Part of oesophagus showing teeth in tumen. Fig. 1].—1],ateral view of bursa. Fig. 12.—Dorsal ray and genital cone. Fig. 13.—Posterior end of female. Figs, 14-22. Cleacina edwardsi. Figs. 14, 15—Lateral and en face views of head. Figs. 16, 17. Oesophageal regions of male and female respectively. Fig. 18.—Part of oesophagus shaw- ing plumose structures, Fig, 19,—T,.S. oesophagus in region of plumose structures. Fig. 20,—Posterior end of female. Figs. 21, 22—Dorsal and laieral views of bursa. CLOACINA FROM MACROPOD MARSUPIALS iT] Figs. 8. 13, 16, 17 and 20 to scale beside Fig. 13. Figs. 14, [5, 18 and 19 to scale beside Fig. 19. Figs. 9 and 1() to scale beside Fig. 10. Figs. 21 and 22 to scule beside Fig. 22. We PATRICLA M. MAWSON into the fhumen (Figs. 8, 10); (4) the terminal both. Vhe nerve ring lies at about a third the length of the oesophagus from the head, and the excretory pore wt the level of the anterior end of the oesophageal bulb. In the feniale the tail is conical and pointed, and the yulva is about half the tail Jongth in fron! of the ants. The yagina is rather longer than the distance from the vulva to the tip of the tail, and is somewhat convoluted, The eggs are about 173 x 30 am, and contain a “tad- pole’ stage larva, The bursa ts much shorter ventrally than dorsally. The arrangement of the rays is shown in Figs, |) amd 12. The genital cone is well deycloped and bears dorsally a short pair of appendanges, The spicules are a little more than a quarter of the body length: a gubernaculum is present. The species is among the medium-large sized Cloacina spp.. and can be distinguished by the characters of the head and ocsophagus. It is perhaps closest to C. communiy Johnston & Mawson (1939), which however ts larger, and in which the oesophageal teeth are arranged ditterently (Mawson 1961, p, 196). The specific mame is given in recognition of the work of Miss Helen Clark who tsolated the worm, and who included a study of the eurly stages af the life history In work for an Honours Degree in this Depnriment. Cloacina edwardsi asp, FIGS, 14-22 Hast and locality: Wallibia bicolor, trom Sun- day '. Vie This apparently new species of the genus Cloncina belongs to the group in which the dis- tal segments of the submedian cephalic paptilae are much larger than the proximal segments. The cuticle behind the head is thick, becoming less su towards the base of the gesophagus, The long threadlike cervical papillae lie shortly in front of the level of the nerve ring. The buceal ting is short, wide and stoutly built, the walls gyal to triangular in section, The oesophagus is short, more or less cylin- drical, with a small terminal swelling in the female, but not in the male. ‘There are no teeth in the lumen, but there are three very distinct “nlumose"” areus, one on each of the three walls of the lumen (Figs. 18. 19). in the region just unterior to the nerve ting. These areus arc formed by confluent ridges on the cuticle lining the himen. ‘They appear to be similar in form to such structures figured and described for some Marshidia spp. from elephants and rhina- ceros, but have not previously been described from Australian trichonematines, The nerve ring surrounds the o¢sophagus al or just behind its midlength. and the excretory pore is jist post-oesophageal. The posterior end of the femule tapers gra- dually from about the vulva, enditg in i slen- der pointed tail; the vulva is tather more than a tail length in front of the anus. The vagina is very short, The eggs are 83 4 50 aim. The spicules are Unusually short for Cloacini spp., about 1/14 of the body length; a guber- miculum is. present. The genital cone is well developed, conical, with two small projections forming the accessory cone. The form and arrangement of the bursal rays are shown in Figs, 21 and 22. This species is. distinguished from any pre- viously described by the presence of the plu- mose structures in the Gesophagus, as well as by the unusually short spicules and the very shorl vagina. Acknowledgements Lam very much indebted to the people after whom the species are named, who. have col- lected These and other nematodes and sent them to me—Mre. Barry Munday of the Mt. Pleasant Luboratories of the Tasmanian Department of Agriculture, Launceston, Tasmania, Miss Helén Clark of Adelaide, and Mr. Geoff Edwards. a post araduate student (1969) in the Depart- ment of Zoology, Monash University, Vic- torin. References Jounsvon, T. HL. & Mawson, P. M_ (1939) — Strongyle nematodes from Central Australian kangaroos and wallabies, Trens, R- Sor, S- Anat, G2, 263-28, Linstow, QO. vow (1897),—Nemuathelminther gesummele von Herm Praf. Dr. F, Dahl im Bismarck-Archipel. archiy, & Marturges. 63, 281-291, Mawson, P. M. (1961).—A note on the occur- rence of oesaphogeal iceth in species of the seniis Cfloatina (Nematoda: Strongyloides), Trans, R, Soc, S$, Aust, 85, 85-89. CLOACINA FROM MACROPOD MARSUPIALS 113 TABLE 1 Measurements of Cloacina edwardsi, C. mundayi and C. clarkae; unless otherwise indicated, measure- ments are in um C. edwardsi C. mundayi C. clarkae male female male female male female Length (mm) 6.3 6.5—7.7 3.1-3.4 4.45.0 8.8-9.2 12.3-13.6 Oesoph. length 480 470-520 470-490 470-570 1100-1220 1250-1350 Antr. end—nerve ring 260 250-290 250-260 250-320 300-350 300-350 —cervical pap. 220 170-200 100 80-90 115-160 110 (3x) —excret. pore 550 480-610 250-340 330-350 890-900 930-1020 Spicule length 430 —_ 1100-1150 — 2400-2600 _ Tail length — 290-300 —_— 210-250 — 260-350 Vulva postr. end — 680-730 —_— 410-500 — 430-500 Length/oesoph. L 13 13-15 6.5-7.1 8.6-9.3 7 2-8.5 9,5—-10.7 Length/spic. L 14.6 — 2.83.0 = 3.43.7 — FURTHER RECORDS OF THE PITTED-SHELLED TURTLE (CARETTOCHELYS INSCULPTA) FROM AUSTRALIA BY R. SCHODDE, I. MASON AND T. O. WOLFE Summary Further records of the Pitted-shelled Turtle, Carettochelys insculpta, including the first breeding record, are reported from northern Australia. It is concluded that the species occurs in river systems right around the landward margins of the Sahul Shelf. The stone fruits of Pandanus, whenever they are available, appear to comprise an important item of the turtle's diet. FURTHER RECORDS OF THE PITTED-SHELLED TURTLE (CARETTOCHELYS INSCULPTA) FROM AUSTRALIA by R. Scuoppr,* L Mason,* and 'T. O. WoLre* Summary firther records of the Pitled-shelled Turtle, Caretlechelys inseulpia. including the first breeding record, ace reported from northern Australia, 1 is concluded that the Species occurs im river systems vight around the landward margins of the Sahul Shelf. The stone fruits of Pandanus, whenever They are available. appear ja comprise an important ilem of the lurtle’s diet. The Pitted-shelled Turtle! Caresoehelys insclilpre Ramsay, sole living species oF a family, Curettochelyidae, that apparently vecurred widely in palaearctic and nearctic revions up until early Tertiary times, was known only from the river systems of southern New Guinea until 1969 (de Reoij 1915, 1922: Wermuth & Mertens [961). In that year, the first specimens were recorded for Australia (Cozver 1970). Ten specimens were captured. all in the Daly River, Northern Territory, in freshwater teaches about 13 km above tidal influence. Because the single specimen exa- mined did not appear to differ significantly from New Guinean specimens, and na evi- dence of breeding was found, Cogger (\.c.) speculated on whether or not the Duly River turtles represented merely uw non-breeding out- lier Of a parent Papuun population. AL speci- mens were relatively small, ranging in curapace length from about 26 to 38 erm, It Was also delermined from faeces that food ingested by one of them comprised figs and freshwater snails. A seeond verified record now cones from the South Alligator River system. approxi- mately 400 km eust of the Duly River site. There, in’ Yellow Waters billabong on Jim Jim Creck, a single female (CSIRO R. No, 320) was ecuught by uw CSIRO fauna survey team on 35 November 1971. The specimen (Fig. 1). deposited in the museum of CSIRO's Division of Wildlife Research, Cunberra, is large, having a carapace Jength of 45.6 cm. This compares with ca 48-50.5 cm for the Jargest New Guinea specimens (Walther 1922: Schultze-Westrum) 1963). Other dimensions, taken from life, are: carapace breadth (includ- ing marginals) 36.5 cm: total height (carapace + plastrog) 14.8 em: head length (to base of crown) 15 cm; head width 8.1 cm; heqd + neek length (to gular shields of plustron) 18.5 en Tore-limb length (posterior margin of flipper) 26.4 em: hind-limbh length (anterior margin of flipper) 25.3 em: tail (to base of anal shields of plastrow) 16.9 em long. with 12 or 13 dorsal scule bands. Soft part colours: shell. limbs. head and tail, mid to durk olive- brown dorsally, prading to cream ventrally (fleshy cream on plastron), iris mid blue-green- grey, Except for the reduced number of caudal scutes, quoted ut 14-16 for New Guinean specimens by de Roo (1915). the South Alfi- gator specimen appears to be identical with New Guinean forms. The condition of the reproductive tracts showed the female to be in the process af lay- ing: a large number of enlarged megalecithal ova Were present in the ovaries (Pig. 2). Many follicles also appeared ta have ruptured recently, and both oviduets were murkealy swollen. The large unshed Ova were jlmost the size of shelled eggs according to the dimensions for the latter illustrated by de Rooij (1915. fig. 102). Because Yellow Waters billabong. c@ 30 km up stream above tidal influence on the South Alligator system, had been landlocked between April and November during the mon- soonul dry season, there can be no doubt that egus had been deposited somewhere ulong the billabong, This represents the first evidence that the species breeds. in Australia. While kept alive for a lime in water. the turtle defaecated large quantities of partly digested husks of the stone Fruits of Purteinis, as Well ds a few shoot leaves (Melaleuied and * Division of Wildlife Research. CSIRO, P.O, Box 84, Lyneham, A,C.T., 2602, Ausiratia, ! Known locally in the Northern Territory by the appropriate vernacular of “pig-nosed turtle” (J. Cin. pers. comm. }. Trios, R. Suc, S, Aust. Vol, 96, Part 2, 31 May 1972. 116 R, SCHODDE, I. MASON, AND T. O. WOLFE 1OcntT Fig. |. The Pitted-shelled Turtle. Caretiochelys insen/pta (CSIRO specimen R, No. 320) - 7 . [een See Oe ee 7 = Fig. 2. Reproductive tract of ovulating Carettochelys insculpta (CSIRO specimen R. No. 320). PITTED-SHELLED TURTLE FROM AUSTRALIA W7 Leguminosac spp,), seeds, roots. picces of aerenchymatous plunt slem, and traces of ani- Mal matter. ‘The animal matter, comprising ca | of the defaccated matertal, included fresh- Water snails (Thiaridae sp.), water-hoalmen (Corixidag sp.), the water beetles Hosheod view scutellaris Germ, (Dytiseidae) and A ydro- philus fatipalpus Cast. (Hydrophilidac). and ants (fridantyrmesx sp.). Upon dissection, the eolon and Juwer intestine of the turtle were found to be packed with Pandenus fruit husks. Pertunth segments remained attached to many. of the husks, indicating that the turtle had presumably -broken and euten hurd, green, fruiting cones with its juws—no mean feat. T G. Schultze-Westrum (pers. comma has also Observed the species feeding on pandan fruit in New Guinea. Thus, although turtles bave proved to be somewhat omnivorous {Schultze-Westrum 1963; Cogeer 1970; J. Cann, pers, comim.}, the fruits of Pandanus, whenever falling from trees of the Various species of the genus that commonly line and overhang the estuarine and lower freshwater reaches of rivers in both southern New Guinea und northern Australia, would appear to con- stitute a rather significant item of their diet. There have been numerous other records of “freshwater turtles’ in northern Australian rivers jn recent years (cf, Cogger 1970) and. though in-one case unsupported by specimens, at least two appear to be of authentic Carerro- chelvs insculpta. Qne is of specimens examined hy J. Cann (pers. comm.) from the upper teaches of the Daly River about 140 km above tidal influence in the Northern ‘Territory. A caripace of one of the specimens has been CReg. No. R.31717), The other, published inci dentally by St, John (1967, p, 527) and Parker (1971), is of observations made by the lute A. de Lestang who observed “herds of turtles” devouring the fallen fruits of Pundanus (P. de- lestangii St. John) in perennial rivers south- west of Burketown in north-western Queens- lund and/or eastern Northern ‘Territory. As all other freshwater chelonians in the region are curiivorous, it seems probable that de Lestung'’s observations cefer to Carerrochelvs. These records suggest that the Pitted-shelled Turtle occurs in the lower (to upper) reaches of major river systems across northern Austra- lia, at least from the Victoria River District af the Northern Territory to as far east us the Gulf country of Queensland and perhaps western Cape York Peninsula. From local in- formation. J. Cann (pers. comm.) believes the lure to Be rather common in most coastal rivers flowing into the Joseph Bonapatie and Van Diemen Gulfs, In averall distribution, then, the species apparently occuts in river systems along the landward margins of the Sahul Shelf, Little is yet known of its oecur- rence or movernents in the intervening Arafura Sea. Acknowledgements We are indehied to Messrs. W. Vestjens and I. H. Calaby, CSIRO Division of Wildlife Research, for assistance with measurements and examination of the CSIRO specimen and in the preparation of the text of this paper; and to Dr. T. G. Schultze-Westrum, and Mr. J. Cann of Yarra Road, Phillip Bay, Sydney for additional observations on Cerettachelys in- senipia. Mr. &. Slater, CSIRQ Division of placed in the Australian Museum, Sydocy Wildlife Research, took the photographs. References Coccrr, H. G. (1970)—First recard of the SeuunTze-Westaum, T. (1963).—Die Papua- Pitted-shelled Turtle, Caretrochelys prsculpta, schildkrote aus Neuguinea, Natur nad from Australia. Search 1, 41, De Koop), N. (1915)—"The reptiles of the tndo- Australian archipelago, 4, Lacertilia, Chelonia. Emydosauria.” (Brill; Leiden.) De Ron, N. (1922), -Reptiles (Lacertilia, Chelonia and Emydosauria), Neva Giined 13, 4001... 133-153. Parnsik, S. A. (1971) —Association hetween the Sulphur-crested Cockatoo and Pandanus. WA. Naturalist 12, 23. Museunt 93, 119-127, St. Jonn, H. (1967)—Revision of the Genus Pandanus Stickman, Pt. 23, Three Australian Species of Pandanuyy, Paci{. Sei 21, 523-530. WALTHER, W. G, (1922)—Die Neu-Guinea- Schildkrote Carettachelys inseulpia Ranwsay. Nova Guinea 13, Zool,, 607-704. Wermurs, H., & Merrens, R. (1961 ),—“Schild- kroten, Krokodile, Bruckenechsen.” (Gustav Fischer: Jena.) THE MORPHOLOGY AND RELATIONSHIPS OF MUELLERENA WATTSII (HARVEY) SCHMITZ (CERAMIACEAE: RHODOPHYTA) BY ELISE M. WOLLASTON Summary The morphology and life history of Muellerenu wattsii (Harvey) Schmitz is described and its relationships discussed. It is recognized as belonging to the tribe Crouanieae (Ceramiales, Rhodophyta) on the basis of thallus morphology and stages in development of the procarp and carposporophyte. Features including regularity of branching pattern and elaboration in development of the involucre surrounding the carposporophyte suggest a probably phylogenetically advanced condition. THE MORPHOLOGY AND RELATIONSHIPS OF MUELLERENA WATTSII (HARVEY) SCHMITZ (CERAMIACEAE; RHODOPHYTA) by Fitse M, Woitaston* Summary The morphology and life history of Muellerena wattsit (Harvey) Schmitz is described and jts relationships discussed, It is recognized asx belonging to the tribe Crouanieae (Ceramiales, Rhodo- Phyta) on the basis of thallus morphology and stages in development of the procarp and carposporo- phyte, Featares including regularity of branching pattern and elaboration in development ot the involucre surrounding the carposporophyte sugeesr a probably phylagenctically advanced condition, Introduction Magllerena waist CAHarvey) ‘Sehmitz in Schmitz & Hauptfleisch was described as Crowenia watsii by Harvey (1863), who con- sidered it closely allied to C'. agardhiana [now Ptiloecladia upardhiana (Harvey) Wollaston 1968). Schmitz (TASD) listed the species as helonging to a new genus Muellerella which, however, he formally described (in Schmitz & Hauptfieisch 1897) as Muellerena with M. witht as the type species. Muellerend is a monotypic genus scemingly quite distinct from other closely related genera. Schmitz (1889) placed it in the Dasyphileue but Kylin (1956) suggested that tt was pro- hubly more closcly related to Cronania and considered it to be insufficiently known for correct placement. Hommersand (1963), after cxamining specimens of M. warnsii in TCD, concluded that it was probably correctly placed in Dasyphileae, Elowever, detailed study of both veyetitive and reproductive features show thal it is best placed io the Crouanicae, Material used for investigation has been muinty drift plants collected at Stinky Bay, Nora Creinu, §. Aust. (Wollaston, 14.xi.1955; ADU, A20004: Wollusten, 19.v,1964; ADU, A27924) and at Seal Bay, Kangaroo T., S. Aust. (Womersley, 211.1965: ADU, A2ZB819), These collections inchided both tetrasporangial and carposporangial plants. Muellerena wattsii (Harvey) Schmitz in Schovitz & Hauptfleisch 1897: 496, De Tont 1903: 1388; $924; 490, Lucas 1909; 50. Lueas & Perrin 1947; 344, Mazza 1911: Na, 397. Crouania wartsii Harvey 1863; synop. No. 637, pl, 291, J. Agardh 1876: 86, Tisdall T&98: 503. Muellerelle owatisii Schmitz 1889; (nomen nudum), Kylin 1956: 397. Challus to 13 cm high with terete, sparingly- branched axes beuring alternate. distichous lateral branches up to scveral em long und usually pinnately branched in the outer part (Fig. 1)% laterals borne from allernate axial cells, occasionally with # shorter branch (less thar | cm long) opposite or between the longer laterals (Fig. 2}. These shorter branches are initivted on the basal cell of an omginal whorl-branchlet and develop in its place. Axial cells aro 1-LS times as long us broad with cells of the central mature thallus usually 350-400 ym long, Each axial cel) bears trom its upper part a whorl of 5 whorl-branchlets (Figs. aii, 4), with the exception that those cells which bear lateral branches often produce only 3 whorl-branchiets and | lateral branch (Fig. 31, ini). Growth takes place by transverse divisions of an apical cell and whorl-branchlets are ini- tiated usually on the sub-apical cell (Fig. 5) with the first-formed initial of each whorl in lateral position, the second and third to the right and Icft of it respectively and those last- formed oppasile the first one (Fig. 3ii).. During early development, lateral branches are charac- teristically curved due to their having the first- formed and hence the longest whorl-branchlets borne on the abaxial (outer) side, while the shortest most immature ones are adaxial in posilion (Fig, 5), However, after initial elonga- tion of 1—-several mm, each young lateral pro- duces at its tip further alternate. distichous 451 ? Deparimenit of Botany, University of Adelaide. &. Aust. 3001. Trans, R. Soc. S, Aust. Wot. 96, Part 2, 31 May 1972. (20 ELISE M. WOLLASTON 1 2 Plant habit (Stinky Bay, Nora Creina, S. Aust. drift, Wollaston, 19.v.1964. ADU A27924), Type specimen, TCD (Warrnambool, Vic., cast ashore, Marts, Sept. 1860, Harvey 221), Regular. distichous arrangement of short lateral branches and occasional shorter branches 4 *\ “ Fig. 1, Fig, 2. opposite or between the longer ones. lateral branches on alternate axial cells, On each of these axial cells the lateral branch is initiated first followed by whorl-branchlets to the right and left of it and the last formed one opposite the lateral branch (Pig. 3i, ili, iv). Most lateral branches cease growth early bul ou few continue to elongate and form indeter- minate thallus branches (Fig. 2). Young cells enlurge rapidly and gland cells and. tetras- porangia may occur very close to branch tips, Mature whorl-branchlets consist of several consecutive die or tri-chotomous whorls of cells, and terminate in short, 2- or 3-celled chains of small cells, cach up to 7 yam diam, often terminated by a slender, elongate hair to [80 pm long (Fig. 4). Cells of whorl-branch- Jets ure 1-2 times as long as broad and up to 60 pm Jong in the central part of mature whorl-brunchlets. Whorl-branchlets are com- monly lost trom older axes and particularly from the upper and lower axial face between the distichously-arranged branches, Axes ure corticated, except when very young, by descending, branched filamentous rhizoids of elongate cells which arise from the basal cells of whorl-brunchlets (Fig. 6) and intertwine to form a dense anial covering with short, horizontal branches, composed of a chain of several small cells, projecting out- wardly. In older parts of the thallus the axl cells become very thin-walled aod may be almost indistinguishable within the cortical cylinder. Ovoid to pyriform ghind cells up to 16 am long, evch within a thick gelatinous sheath, occur as homogencous, refringent structures horne in place Of outer branches of whorl- branchlets and scattered, sometimes bun dantly, over the thallus (Fig. 7). Cells of the thallus appear to be uninucleate although pro- perly fixed material has not been available for study with specific nuclear stains. Rhodoplasts vary from small and rounded in young cells lo reticulate and finally to elongute in mature cells (Fig. Si-iii), MORPHOLOGY AND RELATIONSHIPS OF MUELLERENA 21 Procarp and Carposporaphyte—Carpogonial branches, 4-celled when mature, are initiated sinaly on uw supporting cell which ts one of a whorl of 4 cells borne from the upper part of the terminal cell of a short 2 (-3)-celled spectul fertile branch (Fig, 9). The fertite branch is produced at the outer end of the basal (or stcond) cell of a whorl-branchlet and repluces one branch of the normal di- or trichotomy. The sub-apical cell of the fertile brunch also bears a whorl of (4-)5 cells (Fig. 9). Bach fertile branch is initiated near the tip of a branch axis, so that a suceession of maturing procarps and carposporophytcs is produced as the branch axis elongates, Celle of the carpo- gonial branch are formed by transverse divi- sions of an initial cell which is cut off out- wardly from the supponing cell (Figs. 10-13). The lower three cells stain densely and appear homogeneous in structure while the carpo- gonium is smaller, often with a densely-staining protoplast concentrated im one portion of the eell, and bears an elongate trichogyne. Io 90 pm long, usually swollen at its base and its tip (Figs, 12-14). A sterile cell is formed on the upper side of the supporting cell after ini- tiution of the curpogonial branch and is usually well-developed by the time the carpogonial branch is mature (Figs, Fl-13). Following fertilization. the carpogonium en- larges jind becomes rounded in form while the uichogyne degenerates and an auailiary cell develups from the upper part of the supporting cell (Fig. 15). At this stage the three stevile cells, making up the whorl which includes the supporting cell of the carpogonial branch, conimence to enliurge und ench becomes roughly triangular in shape; the sterilo cell berne on the supporting cell divides to form a chain of several cells (Fig. (5) while the cells forming the whorl ov the sub-apicel wxial cell elongate and produce terminally the first cells of branched involucral filaments (Fig. 16), Fusion takes place by means of a connecting cell between the carpogonium wand the upper part of the auxiliary cell, leaving only one or two small cell fragments in place of the carpo- gonium on the degeneraling carpogonial branch (Fig. 17}. Branched involuctal filaments formed. from the enlarged sterile cells on the apical and sub- apical uxial cells of the fertile branch develop rapidly, and loosely surround the developing cntposporophyte (Fig. 17). The hhasal cells of the upper whori of flaments remain charac- teristically triangular and larger than other cells of these branches (Fig, 17). The auxiliary cell cuts off a gonimoblast cell fromi its upper side and simultaneously forms 2 pil-connection with the apical cell of the fertile branch axis (Fig. 18). Through this connection mutriment ix. pos- stbly conveyed more directly to the carpo- sporophyte, while the old supporting cell acts as the basal cell of an involucral filameni. Gonimolobe initials, which each give rise to a rounded group of curposporangia, develop suc- cessively with the first one or two gonimalches produced in a lateral position, Further goni- molobes are produced without regular order 30 that a total of 6 of more groups of cgarpo- sporangi wt various stages of development may be present at the one time (Fig. 19), As the first carposporangia mature, the newly formed pit-connection between the axial cell and auxiliary cell gradually widens and the connection between the lower part of the aus liary cell and the supporting cell remains small and peobably non-functional or is finally broken (Fig. 19), The involucral filaments, each branched several times, curve upward and loosely surround the mature carposporophyte. Spermarangia—not recorded, Tetrasporangia— Sphencal, tetrahedrally-di- vided tetrasporangia, seldom greater than 25 pm diam, are home on the outer cells of whorl-branchlets in place of vegetative branches {Fig, 20), in a similar position to glund cells. They may occur of aity pal't of the thallus but are usually most abundant on young brunches. Type Locality—Warrnambool, Vic. ( Wars, Sep: 1860). Holotype—TCD, Harvey Alg. Aust. Exs. No. 221. Disiribeion—From West I, and Kangaroo I,. 8. Aust.. to Warrnambool, Vic. Diseyssion Muellérene wellsit is characterized hy the following vegetative and reproductive features: (a) a consistent branching pattern and short Iwterul branghes developed regularly from the outer end of axes, whorl-branchlets in whorls of 5 on each axial cell, except on those which bear short laterul branches, (c) fusion between the lower part of the auxi- liary cell and the fertile axial cell during carposporophyte development, (d) a distinct filamentous involuere surround: ing the carposporophyte and involying the th) 122 ELISE M. WOLLASTON original supportilig cell of the catpogonial branch, Vegetative features such as the alternate-dis- ichous srrangement of shurt Literal branches at the tip of each prevyiously-formed lateral, the form of axial corti¢alion, und the consis- tency in arrangement and number of whorl- branchlets on @uch uxial cell, suggest a telu- tionship with the Prifocladia group of the Ciousniexe. These features were tegarded by Wollaston (1968, p. 404) as indicative of a phylogenetically advanced thallus form. Hom- mersand (1963) stated that Muellereng wattsii was quadriverticillute und he considered the order of initiation of whorl-branchlets in a thodomeluceyn sequence (the first abaxial, the next two to the right and left of the first and the fourth one adaxial) to be a significant taxo- nomic feature characteristic of the Dasy- phileae. However, M. waftsii has in fact 5 Whorl-branchlets per whorl except where short Jateral branches are produced, and in these Whorls, the order of development of the 4 ifitixls could have adsen from a crowanioid seyuence in which the second branchlet is formed opposite the first and the third and fourth at right angles lo them, Suppression of the adaxial branchlet when adjacent to another axis is commonly tound in a number of taxa, for example, in species of Plaryrhanuion J. Agandh and Amoenothamnion Wollaston (Wollaston 1968). In Muellerena waersit this could explain development of the intermediate whorl-branchlets of cach whorl prior to initia- tion of the adaxial whorl-braunchler{s), which al fimes is completely lacking at the base of lateral branches. M4. wenesii clearly evolved a stable pattern of branching and on this basis is probably vegetatively advanced. Several other vegetative features of M. warrsil also suggest telationship with the Crouaniewe group. Gland cells, not previously recorded for A¢_ wartwsii, aré similar in form to those found in Prilucladia australis (Plarv.) Wollaston, P. vestita (Hary.) Wollaston, and Gulvonia annulate Harvey, although they lack position of fusion with connecting cell from carpo- 1 place of carpogonium on degenerating carpogonial Tig. 3. tiv. Arrangement and sequence of initiation of short lateral branches (Lj and whorl-branch- lets in whorls on successive axial cells near the tips of axes. (Diagrammatic.) i, ui, iii repre- sent in transverse section the 2 celis shown in iv. Fig 4, ‘Vransverse section of axial cel! bearing a whorl of 5 whorl-branchlets with tetrasporangia. Fig. 5, Tip of branch sxis showing altcrnale, distichous arrangement vf youne lateral branches (Ly-Ly) on alternate axial cells and abaxial initiation of first-forroed whor)-branchlets on cells of Jateral branch ates. (Whotl-branchicts on faces of axcs omitted for clarity.) Fig. # Benanched, descending cortical rhizoids borne on basal cell (6) of a whorl-branchlet. Fig. 7, Gland-cells borne im place of branches of whorl-branchlet. Fig. 8 Rboduplast structure (i) young cell with rounded chodoplasts, (it) enlarging cell with reti- culate rhodoplasts, (iii) mature cell with elongate rhadoplasts. (Diagrammatic. ) Tig. 9. Carpogonial branch on special 2-celled fertile branch borne in place of a whorl-branchict brinch on basal (or second) cell of whorl branchlet. Fig, LO. Carpoyonial branch initial eut off ounyardly fram supporting cell, Fig. |]. Young curpogunial branch, 3-celled stage, on supporting cell Which also bears a small aterile cell (5), Fig. 12. Curpogonial branch, 4-celled, with developing trochogysuc. Fig. 13. Carpogonial branch with fully elongated trichogyne. Fig, 14, Fusion of spermatium (sp) with mature ctrichogyne Fig. 15. Auxiliary cell (4) formed from upper side of supporting cell; curpogonial branch with enlarging carpogonium (c) and terminal remnant of trichogyne; sterile cell (s) bearing first cells of involucral filament. Fig. lé. Carpogonium (c) enlarged just prior to fusion with auxiliary cell (a); involucrsl filaments commencing to form from cells, including the supporting cell, of whorls an axial cells (axy, ake) of the fertile branch, Fig, 17. Pratrusion on auailiary cell (a) markin gonium; small cell fragment remaining braach: miurked increase. in develupinent of involucral filaments. Fig. (2. Formation of pitconnection between lower part of auxiliary cell (a) and axial cell (ax) prior to breaking of connection between quxiliary cell and supporting ecll; initials of 2 lateral groups of carposporangia formed on gonimoblast cell (2). Fig. (9. Enlarged fusion between lower part of auxiliary cell (a) and axial cell (ax,); succession of carposporangial groups forming on gonimoblast cell; supporting cell free from carposparo- phyte. bearing involucral filament. Fig- 20. Tetrasporangia and gland-cell borne in place of vegetative branches of whorl-branchlet MORPHOLOGY AND RELATIONSHIPS OF MUELLERENA “o mm ul nS ~ SS & ur ou a E a 2 3 2 2 a re ! 2 wo g = 2 2 124 ihe crystallike inclusions recorded for these species. Branched cortical filaments bearing short ollwardly-orientated chains of cells are similar to thase found in Prilectadia palchra Sounder and a tendency towards distichous branching nf the thallus, well defied in wellerend watisi, is also characteristic of the Crouanieae group and is best developed in species considered to be phylogenetically ad- vanced, Development of the procanp and carpo- speraphyte also biusically resembles that found in gerera of Crouanieae. The 4-celled carpo- gontal branch is borne on a special fertile branch as in Gilsonta. The connecting cell in- volved in fusion between the ¢arpogonium and auxiliary cell is much larger in Muellerena watesii than in genera of Crouanieue, but yub- sequent development of the carposporophyte with futeral initiation of the two first-formed groups of carposporungia is similar ito that seen in species of Plilecladia, Gulsonia and Euptilocladin Wollistan. Mitellerener wattsii differs, however, in elaboration of the involucre which surrounds the carpasporophyte and in the secondary development of a pit-connection linking the lower part of the auxiliary cell ro the axial cell upon which the procarp was developed, Following this fusion, the original connection between the supporting cell and the lower part of the auxiliary cell is ustially broken so that the supporting cell functions as an eilarged basal cell of an involucral filament similir lo its sister-cells of the whorl. The in- yolucral lament bome on the supporting ceil ts Inithved as a sterile cell on the supporting cell during enlargement of the carpogonial brunch amd elongates at whout the same dime as the other involucral branches commence to ELISE M. WOLLASTON develop. These events probably allow a better nutritional supply to the carposporophyte while at the same time providing for development of the filamentous involuere. Although Mfiel- lerena watts differs from) species of Prilecladia in having a more consistent branching pattern, § whorl-branehlets pee whorl, fusion between the auxiliary cell and fertile axial cell and a more claborare involucre surrounding the carpasporophyte, the two genera mre basically similar in both vegetative and reproductive features, This similanty was noted by De Toni (1903) when he placed two specics now recog- nixed as Prifocladia pulchra Sonder and P. agardhiana (Harvey) Woll. in the genus Muel- lerena. M, watisit is also samilar lo Gulsonia in the presence of gland cells and the develop- ment of 4 special Fertile branch bearing the procarp and, later, the carposporophyte. Thus it seems likely that Priloctadin, Gulsonia and Muellerena are closely related and MreJleresa, showing greater consistency in vegetutive feu- tures. and elaboration in carposporophyle organization, is phylogenetically the most highly advanced. The singe of Features already known for gencra of the Crouanieac covers 3 possible evolutionary sequerice leading to the increased organization and stability of thallus features characteristic of Muellerena. No simi- lar relationship can be traced in the Dasy- phileae or other group of the Cerannacese and it thus seems logical to include Muellerena in the tribe Crouanieae of the Ceramiacewe. Acknowledgements Tam grateful for the loan of specimens Crom the National Herbarium, Victoria, and for technical assistance provided by a grimt from the Australian Research Grants Committee. References Acarpu, J, G, 19876),—Species, Genera et Ordines Algarum 3 01), pp. 1-724. Eperisis Systemulis Plorideurum. (Lund.) De Tost, J. B, (1903) —Sylloge Algarom omaiim hucusygue Cognitarum 4. Florideac, Sect. 3, p. 775-1521. (Padua, ) De ‘emt, J.B. (1924).—Sylloge Alvarum omnium hucusque Cognitarum 6. Floridese. (Padua) Harvey, W. H, (1863) —Phycologla Australica 5, Plates 241-300, synap. pp. [-73. (London.} Hom™erasann, M, H. (1963).—The marpholosy ant classification of some Ceramiuceac and Rhodontelaceae. Univ. Catfif. Publ. Bot. 35 (2), 165-366. Kris, H. (1956)—Die Gattungen der Rhode phycesn, (Lund.) Lucas, A. H. S&S (1999).—Revised list of the Foucoideae ond Florideac of Australian. Proc, Litin, Sac. NSM. 34, 9-40. Lucas, A. HH. S.. & Prerin, F, (1947).—The Sea- weeds of South Australia. Part I The Red Scaweeds, pp 107-458. (Govi. Printer: Ade- laide.} . . Mazza, C, GRRU Booapurie. A. anthuriv (Rud Key for identification of male specimens nf Acuaria spp. The descriptions of A, gaywnyiy Bisseru and A, iwashkini Brhardoya are not available to mes u full description has not been seen of 4. eremophila Erkulov. A. denuiy Duj. has been omitted because the cordon length and the number and arrangeroent of the caudal pupillac are not known, it falls among species below choice 14 in the key, Species from crows, 4 altenmeta (Rad.j}, Al oernata (Gendreb, A. longicaudata Hoeppli & Hsti, and A, seutara Maplestone, und synonyms of these, have becn assigned to ane group, the “A. anthuriy come plex”. Ic is probable that examination of the Lypes of all deseribed species attributed to Acwearia would show considerable synanymiy, unl might also indicate more important differ- ences helween some species than ure revealed by existing descriptions. J. Left spicule longer Iban 190 em. on yD 1, Teft spicule shotter than 190 em 9 2. Spicule ratio L.1-14 3 2. Spicule ratio 1.5 ar more ees 3, Cordons yery long, extending well past o¢sa- phagus “A anthuris Complex" 3. Paatons very ‘short, not t extending much past excrolory pore ov... 4 4, Body length 4-6 mm, spicule ratip 1.4 A. muyvori Lem, Freitas & Proenga 4, Rody Jengih 10-11 mm, spicule ratia 1.1- 14, _ A, vordata (Mueller) S. Teft spiciile Texs. than 230 vim long & 5S. Lett spicule more than 250 em long 7 6. Cordons end ubout midlength of muscular cesophagis A. subula (Duj,) 6. Cordons nearly as long as oesophagus A. colluricinelae 0, sp. 7. Lett spioule 262 “im A. turdi (Wang) 7. Left spicule over 300 am... x & First pair of postanal papillae ubutil 2 third tail length fronr second pair A. cyanavilla (Boyd) 6. First and second pairs of postanal papillae not much separated 4. streperina J..& M 9. Spicule ratio 1.5 or over 10 9. Spicule ratio Jess than 1.5... 0... 14 1). Cordons reach to end of muscular oeso- phagus a al, conica Maplestoene LU, Cordons very shorl, not much past exerclory pore... tl 11. Six pnirs. of postanal papillae ... 12 11, Seven pairs of postanal papillae ... .. ... 13 12 Left spicule 165 wan long ., A. sialia Williams 12. Left spreule 150 “em long A. puyittitfera ‘Lins, ACUARIA (NEMATODA: SPIRURIDA] IN AUSTRALIA 14} 13, Left spicule 140 #m long . A. paragalliardi Ch, & P, 13. Left spicule 170 4m Jong . A, parorioli Ch & PB. 14. Cordons more or less to end of glandular oesophagus 1s 14. Gurdon harelly longer than rouycular oes phagus _. a (§ Four pairs of preanal papillae aes n 1S. Fewer than four paira of preanal papillae is to. Left spicule 129 um long ' 4. orem) Rasheed 16, Left spicule longer than 150.am . | 17 17. Left spicule slightly grooved near Gp A. partonl Williams 17. Left spicule deeply grooved throughout length A minor Wiliams 8. Three pairs of preanal Papillae .. A_hrevispicula Maplestone 18, Two pairs of preanal papillae . . 19 19, Seven pairs of postanal papillac — - A, alii Rasheed 19. Six pairs of postanal papillae .. A. stn Rasheed 20. Spicule ratio close lo 1.0 . at ap Gs pl 20, Spicule ratio t4I-14 - - 25 21. Spicule Jength Jess thaw 130am........, 22 2), Spicules longer than 130 4m 24 22. Six pairs of postanal papillve A. cremopiila Erkuloy 22. Seven pairs of preanal papillae in 23 23, Cordons not much past nerve ring .. di. Keng Singh 23, Cordons reach aboul to end of muscular oesophagus _ .,,_ ,_ . A. micraccoe n. sp. 24, Cordons not past nerve ring ., A. martinagliai Le Roux 24. Cordons nearly to end of muscular deso- phacus |, , A upupa Rasheed 25, Six pairs of postanal papillae... ... 26 25, Seven pairs. of postanal papillae 31 26. Postanal papillae in two groups of three pairs... _ 27 26. Postanal papillae not in 1wo distinct BrOUPS ae 29 27. Spicule ratio about |.] A. enter itae mn, sp, 27, Sptcule ratio |.3-1.4 . 28 28. Caudal alae widen al. mmidllengti A. giisenfe Williams 28, Caudal alae about same width throughout .. ..., A, dolifasi Ch, & Petter 29, Cordons reach onty to cervical papillae... .., A. cettiag Heit 29. Cordons reach further than cervical papillae 0. 30 30, Right spicule grooved for n most bof its Tength A. gracilis (Gendre) 30. Right spicule ‘simple - A. dierura Rashecd 31. Three pairs of preanal papillac ... A, brumpti Ch, & Petter 31, Four pairs of preanal papillae 32 32. Left spicule Jess than 125 4m iaae ‘ . A. galliard’ Ch. & Fetter 32, Left spicule more than 135 #¢m long .,,. 33 33. End of right spicule enlarged . A. skriablad Ozerska 33, Tip of night spicule without prominent enlargement 34 34. Cordons reach past excretory pore, and more than half distance between bead and posterior end of muscular oesophagus - A. butinerae Ch. & Petter 34. Cordons shorter, less a half Wis Mistames ay _ 35 35. Parasitic in African oriate ‘ A. orioll Ch. & Petter 35. Parasitic in Australian passerines .... ..., A. petterae n. sp. Descriptions of Species The general morphology of Acuaria spp. is so similar that only the special features of each species Will be described, Measurements are given in Table 2; those of paris of the ocso- phagus are taken from the anterior end of the body to the end of the organ in question; the spicules are meagured in lateral view (often very different from those taken in ventral view), Acuaria anthuris (Rudolphi, 1819} FIGS, 1-3 Hosts and Jocalitics: Corvus coronaides fram Adelaide and Pt, Augusta, S, Aust; ©, mellort from Balgowan, S. Aust. and Laun- ceston, Tas,; C. bennetti from Lock, 8. Aust and Erldunda, NT; C. ofre feom Plenty River, N.T.; C- sp. from Pearson 1, S. Aust, Acuaria cnthuris has been recorded many times from different parts of the world; refer- ence lists and discussion of its synoymy may be found in Skrjabin et al (1965) and Chabaud & Petter (1961). The present study deals only with the variations observed in the Australian specimens. The species is quite common in Australian crows and ravens. Measurements are given in Table 2. The general appearance, except where noted below, agrees with descrip- tions given by Singh (1948), Rasheed (1960) and Chabaud & Petter (1961). The cordons extend well past the oesophagus in both sexes, reaching a little under a third of the bady length in the male and a little more than this in the female, but never quite reach- ing to the vulva, The cordon structure (Fig. 1) is different from that figured hy Rasheed, The papillae on the male tail are usually more or less symmetrical, comprising four pairs and one median preanal paplilae, and six paits of postanal papillac, arranged as three pairs on the anterior half of the tail and three pairs of rather smaller papillae on the last PATRICIA M. MAWSON 142 O8r—-OST (00F8) 0076-0095 0091 00ST (0s) M061 ——-OSF-0OZ yBuay wops0g U's OE (LOE) eFI-Ls gs LSI-8 Tl (29) YL-6F yyaua] “ydosao/"] OTE ces (%F' 8) 8'0S-8°9r PES YES PE (%9 CP) YOS-EP yrsuay 9% sb ‘ea[NA—IIUY 601 99 (S01) = SZI-£'8 88 6¢FI-TIl (Slt) = 6 I-88 (uuu) Bayna— OLE Ocr (Té€) O00r-OtE asy OLE-09T (799) O08-OSS ‘d -19xXa— 06L 067 (997) 00-002 Ott OPzZ-O08T (itr) 009-0Z£ ‘ded *araa— ara OTT (Fit) =00£-097 QOL Orc-O8T (807) O£S-ODE SU] cAlau— pus “uy agT O&e (617) O€Z-OIZ OTS QO7T—-O9T (ZOE) OSE-OST arnqussA, 006 OOS (798) 086-008 O£8 O0TI-O09 (16z1) OOSI-O0OL “ydosao ‘osnyy v¢ st (9607) 0097-029 | ooze NOOZ-OOST (007) DOOS—ODE sndeydosag : —= Lie rel (SIZ) 6 FT-0'9T SOL — 87-6 ET (y's) =e 0E-l'6I ysuaTy ayeuoy Cr SP OE 67 (LF) to-0F Oo-CF (et) Se-8c yysuay “ydosac/T OLE ‘OLZ O°7 0007 (L61) $6c-0T1 OEEOOT (OOLF) OOSF-DOTZE yj8u3] UOpIOD UE'TT OT ol (er) rI-el LI-¥'l (TT) PI-lt ‘ards “yf /aids “J STLOET OOT OCT (O€l) OFI-OTT OFI-OLL (927) OL7-061 gmaids "y Ort ‘OPT OOT Ole (TZ1) = S8I-OFI 007-06 | (pit) = OTE-NET aynaids “7 ORZ OL? 09Z O€e (942) OZE-OFT OvE-OZZ (19) OES-ORE ‘d -t9xa— 002 ‘SLT OLI O61 (£61) O1Z-09T OOZ-OFL (O€£) OFP-OLT ‘ded *Alga— OI? Sst Sor 00z (Z0t) = $@Z-SLI 12-051 (€Ze) O9€-00E SUI aAJou—pua “UY OLI ‘OST OZ OT (S91) O8T-OST OLT-OfT (8pZ) O8Z-00Z aNqusaA 0729 ‘08 OOF OOL (PIS) Of9-00r OLL-O¢Pr (¥Z6) + O80T-O00L ‘ydosaa ‘asnjAL O9LI ‘OEET OOST O0ET (6671) OOSI-O00T 0061-0071 (0067) OOFF—-00SZ snaeydosag oF 39 (6S) 69-TS — O6-Ve (vIl) wv tI-L'6 (mur) ysueT ae aapegul MOOG ET, SEP uvaul) duel Dulsadasy mignlsys ues) asuel pony ed pint ae pavoaee, DLN puDnap sh Senpue pupnoy ‘utd U1 BAD STUAIMAINSDAUL J]7 Palys asimaayso ssajusy “dds eirenoy fo siuatwaimspa fy! c a 1avL ACUARIA (NEMATODA: SPIRURIDA) IN AUSTRALIA 143 quarter of the tail, as well as a pair of yery small phasmids almost terminaliy. The mem- hers of a postanal pair are not always strictly opposite to one another. Individual variations from this occur, some specimens having one or two papillae missing from one side or the other. Of 85 male worms examined, 18 showed some abnormality in the caudal papillae. Most of these were one papillae more or less on one side or the other; in a few there was one papilla more or less in the terminal group of postanal pupilluc. In three specimens there were six pairs in the preanal group, the most posterior of these lying just posterior to the anus, so that they could be regarded as an extra postanal pair except that they continued as a closely spaced line of small preanal papillae on each side and were quite separated from the larger papillae of the anterior group of postanal papillae which were further apart. Except for these three specimens, all had six pairs of post- anal papillae, of which the anteriormest lay 4 toe lo =————— —— 3 \\ 1 nm Segk= =06c | 3o0F 2 Soo = f eq 7 =3eE8a— Figs. 1-3. female, Figs, 455. Figs, 6-9. _ = le 1 Soge mS == ——T > S03 = SHe= 13 6 1S pint _ os a 5S (c= oe set 2 a JS( tC S34 E= Acuaria anthuris. Fig, 1,—Part of a cordon, Fiz, 2.— Right spicule. Fig, 3—Tail of A, strepering, Fig. 4.—Part of a cardon. Fig, 5—Posterior end of male. A. skrjabimt. Fig. 6.—Part of a cordon. Fig. 7,—Anterior end of male. Fig. §—Pos- terior end of mule, Fig. 9—Tail of female, Figs. 10-13. A petrerge. Fig, 10—Anterior end of male. Fig, 11.—Parl. of a cordon. Fig. 12.— : Posterior end of male. Fig. 13.—Tail of femalc. Figs. 2, 5, 7, and 13 to scale beside 2; figs, 8, 10, 12, and 13 to scale beside 12. 144 PATRICTA M. MAWSON some distance hebind the anus. ‘The spicules are grooved (as described by Singh 1948) and alate (Fig. 2), The Jeft spicule is larger thai the right except al the lip und the expanded pirts of the alac are wider. Chabauu & Potter (1961, p. 210) report A. anthuris of two types; the first (from Gurritlus glandarius and Piew pica), rather smaller, with six pairs of postanal papillae in the male; the second (from Corvus corone). larger and with seven pairs of postanal papillae, in addition te the phasmids, The only measurement given is (hat the males of the smaller specimens are less than 12 mm long. The smaller specimens agree with Rudolphi’s specimens selected from material (apparently containing more than one species), by Schneider (1866) as the type for A, anthuriy, Vhe Australian material, though perhaps a little longer, agrees wiih these types Acuarla streperina Johnston & Mawson, 1941: 154, FIGS. +5 Host und locality: Sirepera versicolor melan- optera from Waikerie, S. Aust, The type specimens, of A streperina have been re-exuimined and the onginal description must now be amended; they are old specimens, poorly fixed and much contracted. The length given for the oesophagus, 700 ym in the mate and 800 ym in the female, is that of the mus- cular part of the organ; the end uf the vlandu- lar part is 2.1 mm from the head in the female, which is strongly contracted, and 1.4 nim in the male, which is less so. The cardons reach nearly to the end of the oesophagus in the Female, andl to the end of the muscular oeso- phagus in the male, There are six (not five) puirs of posteloacal papillac in the male, atringec! with three well spaced pairs on the proximal two-thirds of the tail and three pairs. closer together, on the distal third. The papillae of the fatter yroup are much smaller and harder to find. The spicules each have an enlarged proximal end, which is less heavily ghitinised and was apparently not included in the original messurements. The spicules ure IO am and 120 um dong, with a ratio of 1:1.7, The largest eggs are 45 x 28 ym, A single female worm from the type host species is referred ti A. streperina. [It was col- lected and fixed after death and so is in a relaxed condition. fis measurements are cilfer- ent in those of the type [cmale largely because of this, Eges in this specimen are mot em- bryonued and are thin-shelled, Measurements are given in Table 2.. The species is very close to A. cranecitia (Boyd, 1950) bat is distinguished hy the avangement of the pastanal papillae in the male Acuaria skrjabini Ozersha, 1926; 103-1117 vide Skrjabin v7 ai. 1965: 114. FIGS. f--9 Hasts and locality; Exotic aviary finches from New South Wales: Tiuriv canora, Lonchura maleces and Kstrilda melpada, These specimens oceutfed in large numbers in many specimens of the finches and ware con- sidered by the owner of the aviary to be the cuuse of the death of the birds. They degree generally with the figures and description of A. skrjabiné by Ozerska and also by Siagh (194874, the principal differences being [hat there are 7 pairs of postanal papillae in the male. as described by Singh. mot six ay shown by Ocersky; the spicule ratio is nearer that in Ozerska’s specimens than those of Singh. Where isa distinct enlargement al. the distal end of the Tight spicule, The comons in the male reach to, and usually beyond, the excretory pore, and those of the female are longer. reaching to about half the distance from the head to the end of the muscular oesophagus. The caudal alae of the male are distinctly wider anteriorly, There is only a slight distine- tion in spacing between the first four pust- cloacal papillae and the Jast three. In some specimens the postcloacul pairs are not arranged symmetrically and ina few one mem- ber of a pair is absent, Both spicules are in- dented at the tips and this is clearer in the right spicule as i ends more broadly, The cag size is 40-43 by 23-24 ym: this is rather shorter than Oxerski’'s measurements, and clistinetly larger than thase of Singb. Acuaria petterae 1, sp. FIGS. 10-13 Hosts and loculitiess Lalage lencomela Srom Katherine Gorge, N T., type host; Meliphuge virescens, M, pluntinia and Cractieus nigro- eularis from the Pelermann Ranges, N.T.; Arianius melanops from Alice Springs, N.T.; Cinclesoma cinnummameum trom Tobermory Sta., N-Tuy Myiaera inquieta and Drymades hrunneopysie from Blanchetown, S, Aust. Probable hosts and localities (only females present): Acanthogenys rufogularis trom Blanchetown, 8. Aust; Anéhochuera carun- ACUARIA (NEMATODA-: SPIRURIDA) IN AUSTRALIA 145 culata From Verran, S. Aust; Arvarruty melinepy from Port Angusta, S$. Aust. Oregica gutiuralis from the Petermann Ranges, N.T. Although the hosts listed above cover a wide range of bird groups, and a wide geographical Tange, there appear to be no specific differences among the specimens from each. Although there is some variation in the position of the cervical papillac, and in the length of the cor- dons fn the male, there is often as much varia- tion betwoen specimens from one host as be- Iween specimens fiom different hosts. The cordons are short. They do not extend as far as the nerve fing in the male, or forther ihan the excretory pore in the female, The vulva is at about the mid-body, just in front of or just behind this, The vagina passes hackwards. Eggs are 38-39 by 21-23 jam, The caudal alae of the male are slender and only slightly wider in their anterior halves. There are typically four pairs and one median preanal papillae, seven pairs of postanal papillae and a pair of very small phasmids, The postunal papillae are not atranged in two groups, hut lic progressively closer together towards the tip of the tail, In some specimens there are more or fewer papillae on one side or the other, bul these appear ta be ybnor- malities, The spicules are unequal; the tips of both are blunt and rounded. The species appears to be very close to 4. orieli Chabaud & Petter (1961), based on specimens from an vniole from Dahomey, which had been placed (with reserve) by Gendre (1912) in his species A. gracilis, from Recharge otra from the same locality. Gendre stules that the specimens from the onole were in nearly all points similar to those from the drongo, distinguished only by the number of postanal pupillae in the male, andl the shape of the tip of the male tail. The cordons of A. orioli are longer in both sexes, than those of the Australian specimens. In the absence of more information about A. orioli, the Avstralian specimens are regarded as a dis- linct species, In some ways it resenibles A, skrjebini tut differs from this species in the more slender build of the spicules, the un- enlarged tip to the right spicule, the shape of the caudal alae, and the detailed structure of the cordons, Acuaria colluricinclae n. sp. FIGS. 14-16 Hest and locality: Coluricincla rifivensris from Eyre Peninsula, S. Aust, The material consists only of one male art one fomale specimen, but these differ distinctly from A. pelteras which appears to be the com- menest species of the genus jin AUstealian passerines. Mcasurements are given in Tuble 2. The cordons extend nearly to the posterior end of the glandular oesophagus in both sexes, at dittle hearer in the female. Detail of the cor- don structure ure shown in Fig, 14. The spicules are unequal in length; the right spicule ends in a swollen tip. There are four pairs and one median preanal papillae, 6 pairs of postanal papillae, and one pair of phasmids. The postanal papillae are asymmetrical { Fig. 15), presumably an abnormal condition; the first 3 pairs are well spaced and spread over the anterior 220 ym of the 280 ~m long tail, while the Jast 3 pairs are Smaller and Jie on the ter- minal 50 jum, The species is distinguished from other Aus- tralian ones. by the ratio of the spicules, the structure of the right spicule, the grouping of the postanal papillae, and the cordon length. It is distinguished From other close species us shown in the key to species. Acuaria microecae n, sp. FIGS. 17-20 Host and locality: Microeca leucophaea from Waikerie, S, Aust. The measurements of this species. of which onty [ male and | female are present, are given in Table 2. The cordotis of the male reach to the end of ihe muscular oesophagus; those of the female to about halfway between the head and the posterior end of the slandular oesophagus. There are four pairs and one medign pre- anal papillae, scven pairs of pustanal pupillae and a pair of subterminal phasmids. The post- anal papillae on each side are inure or less evenly spaced along the tail, the posterior anes slightly closer (ogether. The spicules are oqual in length and similar in build; each has 9 pair of short alae towards the distal end, and the rounded tip is bent ventrally, The vulva is slightly behind the midbody; the eggs gre 35 x 27 ym. The species is distinguished [rom others from Australia by the presence of equal spi- cules, It differs from other species in which the spicules are equal and in which there are 7 pairs of postanal papillae, in having longer cerdons and in the very short spicules 146 PATRICIA M. MAWSON um — 12) oO B m 100 Figs. 14-16. Acuaria colluricinclae. Fig. 14.—Part of a cordon. Fig, 15.—Posterior end of male. Fig. 16.—Tail of female. Figs. 17-20. A. microecae. Fig. 17.—Anterior end of male. Fig..18—Part of a cordon. Fig. 19.— Posterior end of male. Fig- 20.—Tail of female. Figs, 21-23. A. mirafrae. Fig. 21..-Anterior end of male, Fig. 22.—Parl of a cordon. Fig. 23. — Posterior end of male. Figs. 15, 19, and 21 to scale beside 21; figs. 16 and 20 to scale beside 20; figs. 17 and 23 to scale beside 23: figs, 18 and 22 to scule beside 22. ACUARTIA (NEMATONA: SPIRUREDA) IN AUSTRALIA Acuasia mirafrae n, 4>2>1. The terminal discs are large, the diameter of the disc of the third finger being twicc the diameter of the pen- ultimate phalanx. 2 7 ROld Zim Wipim_e Kuru Collecting sites in the Western Distrizt, Papua. The hind limbs are long and slender with a TL/S-V ratio of 0.605, and toes in decreasing order of length 4>5 — 3>2>1. The webbing between the toes is reduced, reaching the sub- articular tubercle at the base of the penultimate philanx on the third and on the fifth toe, There is a prominent, elongate imner metatarsal tubercle and a prominent raund ouler meta- tarsal tubercle. The dorsal surfaces of the head, body and limbs bear scattered and poorly developed tubercles. The throat and chest ure smooth, and the abdomen and posterior ventral sur- faces of the femora granulur. This male specimen has a single, subman- dibular vocal sac and glandular, but unpig- mented, nuptial pads. The dorsum is dull brown and mottled with slightly darker and irregular patches. ‘The ventral surfaces are very pale cream with a brown reticulum on the throat and chest. There is fine light stippling on the ventral und pos- terior surfaces of the thighs, Dimensions: Snout to vent length 21.5 mm; tibia length HYLID FROG FAUNA OF NEW GUINEA 159 Fig. 2. 13 mm; head length 8.6 mm; head width 6.9 mm; eye to naris distance 2.9 mm; internarial span 1.9 mm; eye diameter 2.7 mm; tympanum diameter 1.5 mm. Variation: The paratype series consists of sixteen adult males and one gravid female: M.C.Z. 82380-90, S.A.M. 11659-61, collected at Menemsorae with the holotype M.C.Z. 82400, Derongo; M.C.Z, 82377-78, Yongtau. The series forms a remarkably homogeneous group, the snout to vent length range of the sixteen males being only 21.3-23.9 m. The snout to vent length of the female is 26.3 mm. Similarly, there is scant variation in their pro- portions: TL/S-V 0.605-0.647 (mean 0.627), E-N/IN 1.381-1.722 (1.577), HL/HW 1.114- 1.348 (1.245), HL/S-V 0.376-0.413 (0.399). In all specimens the canthus rostralis is con- sistently a prominent feature, the eyes are large and protruding and vomerine teeth are present. With two exceptions the paratypes are brown dorsally with or without faint and irregularly shaped paler markings. The excep- tions are grey; one uniformly so, whilst the Litoria timida, paratype (M.C.Z. 82390). other (M.C.Z. 82400) bears two very broad and clearly demarcated, pale, dorso-lateral stripes extending from the eye to the groin. Pigmentation of the ventral surface of the throat and body varies from a very light stip- pling to extensive dark brown reticulations on the throat, chest and anterior half of the abdomen. In life the dorsum is brown with paler mark- ings, and the posterior surfaces of the thighs are dark brown with or without a few yellow spots. The labial stripe is white, and the iris a reddish bronze, being palest around the pupil, The flanks vary from brown to pale yellowish, and the groin is bright yellow. The ventral sur- faces are yellow and the ventral markings re- ported above are brown. Ten additional specimens have been col- lected from localities to the east of the Western District: M.C.Z. 82379, Oroi, Purari River [Lat. 7°237S., Long. 145°11’E.], Gulf. of Papua (obtained by F.P.): S.A.M. 10656-8. 11657; U.P.N.G, 2510-11, 2581-3, Brown River Forest Reserve, north of Port Moresby (collected by J. I. Menzies). They differ from 160 M. J. TYLER AND F. PARKER he type series in the following respects: the posterior surfaces af the thighs and the groin bear a striking pattern of yellow markings on a dark brown background, and several exhibit a very narrow, light, mid-vertebral stripe, Comparisan with other species: Literia timida is @ member of the Literia dorsaliy group, de- fined by Tyler (1968a). which contains L. dorvaliy of New Guinea and L. rmierobelos ot northern Queenslind. The members of the group shure long, slender, unwebbed fingers, reduced interdigital webbing of the toes and extremely small adult size. Litoria timide is larger than L. dorsaliy and L. microbelos (maximum adult size of males and females: dersalis 21 mm and 22 mm; microbelos 194 mm and 18.9 mm), The feature by which these species can be ¢dis- linguished most readily is the canthus Tostralis: strongly curved and sharply defined in rina, but straight or slightly curved and not a prominent feature in L. dorsalix and L. niiero- belos (Fig. 3), From £. nicrobelos the new species is Further distinguished by its coloura- tions examples of L, micrabelos are usuidly only very slightly stippled and there is @ ten- dency for the dorsodateral surfaces ta be durker than the dorsal, In £. tinder plgmenta- tion is more extensive, and in the only speci- nen in which the dorso-lateral surfaces dilfer fran) the dorsal, the dorsal portion is darkest, Litera = mitrobelas lacks yomerine teeth (present in £, Unda) and has much shorter hind fimbs, as indicated by a comparison of the ranues of the TL/S-¥ ratios: 0.605-0.647 in L. Moda, 0.500-0.579 in a series of thirteen 1. micrahelos from Cainns—S.A.M_ 12571-4, M.C.Z. (Uneat.). | / ( UH A B Heads of A, Litoria timida and B, L. dorsalis, demonstrating the difference in ihe shape of ihe canthus rostralis. Both specimens are males and there is no evi- dence of sexual dimorphism. Literia’ dorsalix resembles L. pierahelas more closely than it does L. /imida, Pattern of pigment on the dorsum of L. dorsaliy most commonly tends to form broad and slightly contrasting, longitudinally vrranged. light and dark stripes. Womerine teeth are present but diffieult ta detecL when elevations of the vomerine banes are absent. The range of TL/ §-V ratios is intermediate belween those of L. timida and L. inierebelos (0.541-0,621). The only other hylid species recorded trom the Southern Lowlands of New Guinea (recognised as a faunal unit by Tyler (19683), and including the Western District of Papua) which approximate the size of L. rinida are 1. hicolor and associated species. Such species are usually uniformly bright green dorsally. and can be further distinguished from L. tide by possession of webbing between the fingers. Jateral fringes on the fingers. and fully webhed toes The high TL/S-V raho ot L. tinida ips prouches that of Lo meyuta, Because the first couplet in the relevant key to hylid species compiled by Tyler (1968u) involves separation of species on the basis of 9 TL/S-V_ ratio higher than or lower than 0.65, comparison with L. nasura is mecessary. In habitus these Species are quite distinet, L. nasa being much larger and having o very long. slender head wd body Adult L. nasare attain 45 om (males) and 55 mm (females), juveniles (und adults) can be readily distinguished by their possession of longitudinal skin folds on the dorsum, Haliui: The type locality of Menemsorae and Derongo and Yongtau ute in the viemity of Ningerum at the headwaters of ihe bly River in the extreme northwestern part of the West- ern District (Pig 1), and are within dense rain forest. The lerrain is gently undalating to the south of Ningerum and hilly to the north. The series from Menemsorac were collected on the leaves of plants adjacent to, or overhanging, the water of a permanent swamp in the forest, Oroi is situated in tropical rainforest partly cleared to form garden areas, It is believed tht the specimen obtained there had been collected in the extensive adjacent saga swamps. The Brown River Forest Reserve similarly provides an extremely moist habitat. Call: Calling males were observed occupying a horivontal position on leaves close to the water. We lack tape recordings of the calls, but note RYLID FROG FAUNA OF NEW GUINEA 16] that audibly they are feadily distinguishable, The call of L. timida was likened to two short clicks, each followed by a pause, and then a sequence of tapid notes: “tik... . tik. 2.4, Lindiititi’. In contrast, the call of £. dorsalis comprises a series of from six to twenty short, “woodeny buzzes”, and that of ZL. microbeloy a low, imsect-like “creeeek”, becoming pro- gressively higher pitched. Notes: It was noted that indigenees of the Ningerum area did not recognise the new ‘specics, They consistently considered specimens to be juveniles of larger species such as L. thesaurensis or L. amboinensis, The specific name proposed for the species alludes to the timid nature of the animal. When disturbed it rapidly attempts to escape- ADDITIONS TO THE PAPUAN HYLID FROG FAUNA Litoria nigrofrenata (Gunther) Until recently this species was known solcly from the types collected at an unspecified locality on the Cape York Peninsula, re- examined by Moore (1961) and Tyler (1968b), Additional specimens were sub- sequently collected by the second author on the Cape York Peninsula at Endeavour River, Leggits (Jones) Lagoon and Big Tableland (all within a 15 km radius of Cooktown). The occurrence of this species in New Guinea is based on the following nineteen specimens: Boze, Binaturi River: M.C.Z. 81097; Kuru: M.C.Z. 81094-96, P.N.G.M. (F.P.) 438 (2), §.A.M. 11408; Morehead: M.C.Z, 84549-50, S.A.M. 11316-19, 10660, [1467 (3}: Old Zim. Oriomo River: M,C.Z. 79723-24, S.A.M, 10621 (2); Weam: M.C.Z. 79725-26. These localities are all situated to the south and south- west of the outlet of the Fly River and are within 60 km of the coast. The diagnostic characters of this species arc as follows: Size moderate {maximum snout to vent lengths; males 41 mm, females 42 mm). Head elongate with the snout projecting in profile; head consistently longer than broad. Fisst finger longer than second; fingers without webbing; fourth toe webbed to base of penulti- mate phalans; terminal discs small, Dorsal sur- Faces of the head, body and limbs uniform pale hrown, with a broad, dark brown stripe extend- ing from the tip of the snout to a position posterior to the iympanum. There is a hori- zontal white bar anterior to the cye partially bisceting this dark lateral band (Fig. 4). Because few detailed comparisons of such disjunct populations have been undertaken and none previously published, measurements and proportions of specimens of L. nigrofrenata Fig. 4, 5mm Lateral view of head of Litoria nigrofrenata: 1a M_ 1. TYLER AND F. PARKER Fram New Guinea and Australia are presented below. AUSTRALIA NEW GUINEA Number 12 5 S-V 37,2441 36,9--38.8 (nsales) (mean 38.2) (mean 38.0) SV 38.9- 41.3 414 (females) (mean 40.1) (mean 41.8) Th /S-¥ 0,.541-0,697 0,584-0.666 (pooled) (mean 0,629) (meun 0,632) FN/IN 0.925-1.162 0.975-1.083 {pooled} (mean 0.999), (mean 1,007) HI. /HW 1.078-1.309 1481--1.262 (pooled) (mean 1.180) {nyean 1.230) HL/s-V 0,360,398 387 0.404 (pooled) (mean 0.376) {ime 4.396) In life the dorsal surfaces vary from a pale yellowish-brown ta reddish-brown; adults are immaculate, but juveniles are speckled wath darker pigment. Liroria nigrofrenaa exhibits considerable variation in leg length and individuals with TL/S-¥ ratios equal to or exceeding 0.65 key out in Tyler's (19684) key of lowland species lo L. nasutea. Specimens with lower TL/S-V ratios key out to L. vagabuneda. Lueria nasuta is similar in habitus to LZ. nivrofrenate but is distinguishable by possession of longitudinal skin folds on the dorsal surface af the hody and by lack of a lateral head stripe, In contrast, L. vegabanda is a broad- headed species with short limbs (TL/S-V of allolype — 0.513) (Tyler L968a), Literie werge- bunda is dark blue in preservative and so pro- hably green in life, Litevia nivrofrenate was collected in sparsely forested areas and savanna, and purticularly in qpen patches of grass on damp or swampy eratnd, Same specimens were taken beside air- strips and others in grass-covered clearmgs bordering tracks through the forests, His an extremely timid nocturnal species, leaping rapidly towards the cover of dense vegetation when disturbed, By day it behaves similarly when found sheltering in tall grass. Litoria rothi (de Vis) Lirorle rorhi is a member of the L. perorit species graup. Until recently it was regarded as identical with £. perent whieh, bn reality, it replaces in north-east Australia (f. R. Straughan, personul communication). The New Guinea material consists of the following specimens: Gnao, Pahoturi River: M.C.Z. 84551; Old Zim, Oriomo River; M.C.Z, 79727; Wipim: S.A.M. 10623, Boze, Hinaliirt Rivers M.C\Z 79728, S$1613-15_ PING M. (F.PL) 3350; Weam: M-C.Z. 79729, S.A.M. 13113; Morehead: M.C_Z%. (1.P.) 37605 (3), PNLGM. (F.P.) 561, SAM, 13111-12. This arboreal species is characterised by the following features: size moderate (maximum stioul to vent length of males = 42 mm, females 44 mm); fingers approximately one- Murd Webbed with flattened, dilated terminal discs;, tympanum partially hidden by pro- nounced supratympanic fold; dorsal surfaces of preserved specimens dull brown, sometimes with poorly defined, irrepulir patches; back of Ihighs black, with two or three uniformly shaped, pale cream markings, ‘There ate black markings in the inguinal and axillary regions, and a black line wlong the inferior margin of the supratympanic dermal fold: ivis (in tife) dull ted above the pupil, and pale etey or pale geld below i. The only member of the F, perany spectes group formerly known to occur in the southern lowlands of New Guinea is 2. anthoinensty, reported by Tyler (1968a) from Mabilahol on the Ok Sih River in West Friaa, and more recently found by one of us (F.P.) to coexist with L. rathé in the Western District. Liveria amboinensis is a much larger species wilh fully wehhed fingers, and lacks the black thigh markings of J... reathi. Litoria roi was found in sparsely wooded areas and in monsoon forest. It was commonly collected at night on leaves of plants within 2m of the ground; the series collected al Rue were on the leaves of an ornamentel Cretan near village houses. Durmy daylight at was tuund within curled leaves, under bark anid in similar tefirges- Litorin rnbella (Gray) Litoria rubella is a wide-ranging species. in Australia, oceurring in diverse habitats over the entire northern half of the continent. Fts presence in New Guinea hus been established by the collection of the following 77 speci- mens: Daru Island: M.C.Z, (F.P.) 30592: Weam: M.C.Z, (VP. 31467, 81609-12, §1099-81100, Morehead: A.M.N AL 84527, SAM. LOK3O (2), 13720-23, (3125-29, P.N.G.M, (F.P.) 334, (F.1) 362, M.CZ (F.P.) 30936 (2), (F.P.) 30937 (2), (F.P.) 30938 (14), (RP) 37606, (FP) 37607 (301, ALM. 32709-L4 HYLID TROG FAUNA OF NEW GUINEA 163 The characteristic features of the species are: small size (maximum S-V¥ of males 32 mm, females 34 mm) and squat habitus with very short hind limbs; expanded digital discs; fingers approximately one-third webbed and toes ex- tensively webbed; colouration brown with broad, darker brown dorso-lateral stripes on the body. In life the dorsal surfaces are brown, suffused with a greyish, yellowish or reddish tinge. All specimens taken ot Morehewd were densely marked with smyll black flecks, but such markings were absent in those from other localities. The posterior surfaces of the thighs are either immaculate yellow or unpigmented but for a fine brown stippling. Of the species occurring in New Guinea and recognised by Tyler (1968a), L. rubella is most closely related to L. congertita, from which it is readily distinguished by its possession of the dark dorso-lateral stripes, and by shorter hind limbs (TL/S-V range for rubella — 0.335- 0.432: 0.477-0,520 for congenita, vide Tyler 1968a, p. 72). Litoria rubella was collected mainly in open, man-made grasslands within lightly forested country. Most specimens were collected on air- strips. It is a nocturnal species, hiding by day in situations such as cracks in fence posts and amongst plant Icaves. At night it was found within 2 m of the ground, and consistently did not make any attempt to escape when illuminated by a spot light, Acknowledgements We are greatly indebted to Dr W. G. Inglis, Director of Environment and Conservation, South Australia, for constructive criticism of the manuscript, and to Mr J. 1. Menzies, Depariment of Biology, University of Papua and New Guinea for specimens of L. timida from Brown River Forest Reserve, References Loverioce, A. (1956)—Rediscovery of Hyla dorsalis and Leckrivdus papuanus in New Guinea. Breviora ($5), 1-4, Macieay, W. (1878).—The batrachians of the “Chevert™ Expedition. Proc, Linn. Sac. N.S.W. 2 (2), 135-138, Moore, J. A, (1961).—The frogs of Eastern New South Wales. Bull Amer. Mus, Nat. Hist. 121 (3), 153-385, Parker, F. (1970).—Collecting reptiles and am- phibians in New Guinea, Aust. Nat. Hist. 309-314. Rocx, J. (1920) —Note sur la présence du genre Crinia, amphibien cystignathide, en Nouvelle- Guinée. Rev. Suisse Zool. 28 (5), 115-117, TYLer, M. J. (1968a).—Papuan hylid frogs of the genus Ayla. Zool. Verh. (96), 1-203, Tyrer, M, J. (1968b).—A taxonomic study of hylid frags of the Ayla lesneuri complex occurring in norih-western Australia. Ree, S. Aust, Mus. 15 (4), 711-727, Tyter, M, J, (1971).—The phylogenetic signifi- cance of vocal sac structure in hylid frogs. Univ, Kansas Publ. Mus. Nat. Hist. 19 (4), 319-360. Tytmr, M. J. (in press) —An analysis of the lower vertebrate faunal exchange between Australia and New Guinea, Jn D. Walker (Bd.), “Torres Strait: disjunction or no barrier’. (Australian National University: Canberra.) VOL. 96, PART 4 30 NOVEMBER, 1972 TRANSACTIONS OF THE ROYAL SOCIETY OF SOUTH AUSTRALIA INCORPORATED a a CONTENTS Kott, Patricia The Ascidians of South Australia II. Eastern Sector of The Great Australian Bight and Investigator Strait - = =a 65 Dubois, Georges, & Angel, L. Madeline Strigeata (Trematoda) of Australian Birds and Mammals from the Helminthological Collection of the Uni- versity of Adelaide - - OBITUARY: KEITH RODNEY MILES - Annual Report of Council, 1971-72 - - Award of the Sir Joseph Verco Medal - - Balance Sheet - - - - - ~ List of Fellows - - 2 5 = 2 - perl be PUBLISHED AND SOLD AT THE SOCIETY’S ROOMS STATE LIBRARY BUILDING NORTH TERRACE, ADELAIDE, S.A. 5000 THE ASCIDIANS OF SOUTH AUSTRALIA II. EASTERN SECTOR OF THE GREAT AUSTRALIAN BIGHT AND INVESTIGATOR STRAIT BY PATRICIA KOTT Summary Seventy-two species of ascidians from South Australian waters are discussed. Fourteen of these are new and 42 have previously been reported from South Australian gulf waters. Morphological convergence to exploit the environment is evident in many of the forms present. The data support the existence of a marine faunal boundary at the eastern end of the Great Australian Bight. It is suggested that there are ecological factors responsible for the difference between the South Australian gulf and open coast ascidian fauna. THE ASCIDIANS OF SOUTH AUSTRALIA II, EASTERN SECTOR OF THE GREAT AUSTRALIAN BIGHT AND INVESTIGATOR STRAIT by PATRICIA Ko1tT* Summary Seventy-two species of ascidians from South Australian waters afe discussed. Fourteen of these are new and 42 have previeusly been reported fram South Australian gulf waters, Morphological conver- gence to exploit the environment is evident in many of the forms presént. The data support the exi{s- tence of a marine faunal boundary st the eastern end of the Great Australian Bight. It is suggested thut there are ecological factors responsible for the difference between the South Avstralian gulf and open coast ascidian fauna. Introduction Collections of uscidiuans have been previously made from St. Vincent and Spencer Gulfs {Kort 1972a) and from Port Phillip Bay and other locations on the Victorian coast (Millar 1966). The present collections are the first reported on from the southern coast of Aus-~ tralia between Middleton Beach. Albany (S.W, Aust.) and Spencer Gulf. Seventy-two species of the Class Ascidiacea from South Australian locations (Fig, 60) are recorded. Of these. 42 had already been re- ported From St, Vincent Gulf, Spencer Gulf and Encounter Bay (Kott 19728). Fourteen new species are described, The association of species in the areas being considered differs From that in the gulf areas (Kott 19724). Colonies im which each zooid maintains independent openings to the exterior are upparently favoured and many species demonstrate convergence in their adaptations to exploit the environment. especially in re- yard to their siphonal apparatus. The Jarge number of specics taken. and especially the large proportion of new species, teflects the new habitats that ate now being explored by SCUBA diving. Type specimens are deposited in the South Australian Museum (SAM) or the National Museum of Victoria (NMV}, Order ENTEROGONA Suborder APLOUSOBRANCHIA Family CLAVELINIDAE Subfamily CLAVELININAR Clayelina mirabilis n.sp. Type Location: Waldegrave I.: in gravelly sand, attached to limestone, 23 m, Shepherd. Holotype; SAM, E902, Paratypes; SAM, E903. FIGS 1, 2. Deycription: The colonies consist of a spheri- eal, sand covered, base from 2-5 cm in , Family ASCIDJIDAE Ascidia thompsoni Kou, 19729; 27, New Recerd: Elliston Bay. Recverds: sce Kott 1972a- Peseriprion: Individuals are the usual aval, laterally flattened shape, lying mostly on the lefi side. The body is pinkish and shows throwgh the firm translacent testi, There are nO muscles on the left side of che bowky. Body musculature ix present as an trrégular network on ihe right or upper side of the bouy, stopping abruptly at the endoslyle, and no muscles were detecied on the left side of the body. There is not a row of short parallel bands around the ventral and dorsal border as in A. sydneyensiz, The peritubercular area is very shallow. The dersnl lamina is double in its anterior part and is ribbed on the left. side only, The branchial sic is simply folded between each tongitudinal vessel and there are simple papillae at the june- hon of the longitudinal and transverse vessels, Previous ASCIDIANS OF SOUTH AUSTRALIA IL 181 Fig. Fig. Fig. Fig. Figs. Figs. Fig. Fig. Figs, Figs, 33, 34. 35, 36, 37, 38. 39-41. 42. 43. 44-46, 47, 48, Tridideninum cerebriforme, Spicules. Trididemnum spiculatim. Spicules. Polysyneraton magnilarvum, Colony. Didemnunr ternatanum, Spicules, Leptoclinides fungiformis, Pig. 37.—Calony. Fig. 38—Spicules. Metandrocarpa indica. Fig. 39.—Individual, Fig. 40—Mesial aspect of stomach. Fig. 41. —Later aspect of stomach. Symplegma arenosa. Individual. Stolonica azsiralis. Gut loop and stomach. Stolonica truncata, Fig, 44.—Iindividual. Fig. 45 —Body removed from test to show. body musculature. Fig. 46—Gul, gonads. Polyandrocarpa simulans. Fig. 47.—Calony- Fig. 48.—Gut, gonads, 1&2 Remarks; The present species it distinguished by the coudition of the dorsal lamina and the body musculature, Axcidia sydneyensis Stimpson: Kott, 1972a; 24 and synonymy. New Records: Elliston Bay. Investigator Strait (Sin. ¥18). For Previour Records, Description, sec Kott 1972a, Order PLEUROGONA Suborder STOLIDOBRANCHIA Family STYELIDAE Subfamily povyzoinae Metandrencarpa indica n,sp, Type Leecation: Investiguter Serait (Stn. ¥6), 23 m, scattered low recf{ with shell sund patches and strong surge, Watson, Helorype: NMV, HI59. Paratype: NMV, HIS8. FIGS. 39-41 Descriptions The colonies cansist of sessile, round, laterally flattened, sandy Individuals 5 mo m diam,, fixed to a common basal stalk which sometimes expands into a wider mem- hrane. The present colonies invest an algal stom. The zooids are close to one another but each ff entire and separate and the tests do not adhere. The test is thin and fairly brittle with embedded sand, Both apertures are an- terior and fairly close together. The branehial aperture is sessile and the atrial aperture is on a pointed siphon directed away from the hranchial aperture, The body wall is very delicate and closely adherent to che test. It has a fairly close mesh of very fine muscle fihres continuous all over the body. The branchial sac has 4 infernal longitudinal vessels on each side With 6-8 stigmata per mesh, crossed by paras- tizematic vessels, and there are 9 such rows af stigmata, The gut forms u simple open loop across the pesterior end of the body and the rectum ts produced anteriorly toward the base of the atrial siphon. The stomach is large and has about 14 conspicuous stomach folds. There is a longitudinal ridge along the lateral aspect of the stomach, which continucs to form a leng curved caecum im ihe gut loop, On the posterior side of this longitudinal ridge, the somach folds are parallel to it, "The folds an the anterior aspect of the stomach, however, extent! more obliquely and terminate against this caecal ridge so that they do not extend continuously from the pylone to the cardiac PATRICIA KOTT end af the stamach, “The anal border is smooth. The dorsal lamina ts plain. Unfortunately no gonads were detected, Remarks: Specics of the genus Metandrocarpu Michaelsen, 1922, have heen described with 5-10 internal longitudinal vessels, und male and female gonads on both sides of the body. Although the present specics has only 4 interhal longitudini#l vessels und the gonads ure nol developed, its stomach is similar to that found in M. dura (Ritter) and M michaelsen? (Ritter & Forsyth) both from California (sce Wan Name 1945), In both these species the stomach has a longitudinal ridge canlinuous with the pylori¢ caecum and against which the stomach folds terminate. In other gebera of Polyzoinue, eg., Allecorarpa, Theodorelia and Palyzoa, although there may sometimes be u small number of internal longitudinal vessels, the stomach is barrel- shaped with parsllel longitudinal folds. Symplegma arenosa n.sp. Type Locations Off Waldegrave [., Shepherd. Holoivpe: SAM, H904, Paratype: SAM EYRS. FIG. 42 Description, The colonies are formed of sessile, sundy individuals, more or Jess pillar- shaped but laterally flattened, the zooids taller than their width. They are fatrly closely placed but are entirely separate and arise from a sandy basal plate formed of a tangle of basal stolons. Both apertures are close together on the upper free end of the body in a circular, sand-frec, area of verv thin test. The rest of the test is encrusted with sand, When the zooids are contracted, the stiffer, sand- encrusted, lest on either side of the apertures is drawn together across the openings which then appear to be depressed in a longitudinal slit on the supper surface. The body wall has strong muscles vround the anterior part of the body, consisting of longitudinal bands raciat- ing from both the siphons and short trans- verse bands extending across the mid line dorsal and ventral to both of the siphons, These short transverse bands (as in Agnesia eleciata; Kott 1969h) are instrumental in drawing together the protective test across the aperture. There are 4 internal longitudinal vessels on each side of the body and 15 rows Of stigmata, “Uhese are nol. crossed by paras- ligmatic vessels. There are 6-8 stigmata in each mesh, The longituding! muscles extend only about a third of the distanee down the ASCIDIANS OF SOUTH AUSTRALIA IL body, The gul torms a very short loop across the posterior end of the body and the rectum is long ind extends anteriorly toward the hase of the atrial siphon. The stomach is short and barreleshaped With about 14 longitudinal folds and there Is a short straight pyloric caecum connected to the anterior limb of the gut loop by a divided ligament and blood vessels (see Synpleenia oceania Tokioka, 1961), Vhere Is also i connective between the stomach snd ihe intestine. Unfortunately no gonads ate developed in the present specimens, Remarks: Although the present species resembles Metandrocarpa indica in the presence of 4 internal longitudinal vessels in the branchial sac und in the shape of the body, it differs In the body apertures, the body mus-. culalure, the shape of the stamach and pyloric caecum, arid in the length of the gut loop, In the present species, the stomach, as well as the number of internal longitudinal vessels, resemble the condition in the genus Syin- Plegnias and in Swnplegma vecania Tokioka, 1961, from Noumea, the arrangement of the pyloric caecum and ifs ligaments and the con- nective between the stomach and the intestine are identical with the present species, Although Symplegni spp. are not usually upright as in the present species, they are joined (o basal stolons which form a mesh und eross one another and an upright form has been des- cribed once previously, viz: Syrmplegma viride stolonica Verrill (see Wan Name 1945), The condition of the apertures is. very specialised and the disposition of the body musculature to serve the protective device described above fas not previously been re- corded tn the family Styelidae, although similar mechanisms are developed in the families Agnesidae (see Kott 1969b) and Molgulidue (see Kot 1972b), Stolonica australis Michaelsen, Kott, 19724: 28 and synonymy, New Record: Elliston Bay, Previous Records; see Katt }972a. FIG, 43 Description; In colonies recently collected from Tipara Reef (Spencer Gulf), there is a single row of hermaphrodite gonads anteriorly on both sicdes of the body, but posteriorly there are tather irregular scattered series of simple testis follicles. There is some variation in the apparent shape of the stomach, from pyriform to barrel-shaped dependine on the condition 3 of the body wall cavenng the cardiac end, The caecum is only of moderate size and curved Stoloalea truncata f.sp. Type Location; | koe north-west nf Walde- grave Lon rocky bottom with sund patches, 23 om, ILwI971, Shepherd. Helarype: SAM, E893. Paratyipess SAM E894, E909, FIGS, 44-46 Description: Colonies comprise raunded, sandy, individuals joined by basal stoluus. Each individual js spherical, sessile, and about 1 cm in diam. The apertures are both anterior on the frec end of the body, and are present m conspicuous transverse interruptions to the father solid sand encrusted test, so that cach aperture is surrounded by delicate, thin test Which can be withdrawn and covered over by protective sand-strengthened lips, The body wall is Eairly muscular, with longitudinal bands radiating from both siphons, extending over the rest of the body, There is a broad band of circular fibres around the anterior end of the budy and associated with the protective clos- ing mechanism described shove. There ate also circulac fibres around dhe posterior end of the body, There are 3 branchial folds on the tight and 3 on the left, although they all tend to fade out posteriorly, especially the most dotsal folds, The internal longitudinal vessels are arranged according to the following formula: DL 0(10)5(9)5(6)6 E There are 14-2 stigmata per mesh, although these are set slightly obliquely in relation to the internal longitudinal vessels. The gut forms 4 simple transverse loop with the rectum (lirn- ing slightly anteriorly. The stomach has a large number of very fine longitudinal folds, and the whole extent of the anterior border of the stomach is produced into a tube of peradually decreasing diameter, lined internally with glan- Jular folds in paralle] with those lining the rest of the stomach wall, This tube forms a curve in the gut loop and terminates in a pointed caecum, The anus is bilabiate, On the left hand side of the body, anterior to the put Joop, there may be 5-6 elongate and sometimes branching testis follicles, terminating distally in the long slender duct directed toward the atrial aper- ture. On the right hand side of the body, there are similar testis follicles arranged around the posterior half of the ventral border and around the posterior border of the body. Same of these testis follicles are also associated with an ovary 184 of 2-3 crys. lying on the mesial surface of the distal end of the testis and opening by a short wide oviduct, while the seminal duet curves uround and for most of ifs length lies free in the peribranchial cavity, There are up to 6 hermaphrodite gonads of this type in the. centre of the row of gonads on the right side of the body, Larvae are present in the right peribranchial cavity. They have uitudiate papillae andl appear to be of dhe type generally associated with the Polyzoinuc (see Millar 1960), Remarks: The most unusual production of the anterior aspect of the stomach to form the long curved caccum: in the gut loop is quite dis- linctive, The long branching testis follicles are also unusual; while cxternally the highly specialised device to protect the apertures, each located in « transversely onented strip of unmodified! test, by withdrawing them beneath the firm sand-hardened test is unique. Closing mechanisms as described above for Svenple gira aréfosd tsp. and for specics in the families Molgulilae and Aynesiidae, usnally involve both apertures simultaneously. The test is black un can be seen through the encrusting sand. Ocolimuria australis Gray. Kott, 1972a: 29 and synonymy. New Record; Oll Waldegrave [, Elliston Bay, For Previors Recards, Description, see Kou (972g. Polvandrocarpa simulans n,sp. Type Location: Investigator Strait (Stn. Yo. ¥19), 23 m,.on scattered low-ceel with shell sand patches and strong surge, Watson. Further Reeordy: Anvestigntor Strait (Stn, Y19), Elliston Bay, St. Francis I. Holorype! NMY, HI62. Paratypes: NMY, TITAQ, H161, HL63. FIGS. 47, 48 Deseriprion; The colonies form tight agere- gutes of Individuals about 1 cm in diam, The living specimens are reddish brown, tipped with black, although preserved specimens are black. The test is thick with 4 slight sand enerustation, although this as mul present internally where the test of adjacent zooids is confluent. Both uperlures ure sessile. The branchial aperture is terminal and the atrial aperture antero- dorsal. The body wall is very muscular, with ulmost continuous layers of tongitudinal and circolar muscle bands. The branchial sac has 4 low fulus with crowded, very thick. internal longitudinal vessels. The transverse branchial PATRICIA KOTT vessels aire alsa thick so that the branchial sac is a Very tough organ in this species, Longi- twidinal vessels are arranged according to the following formula} E 1l6)1O9)1T(8IT(9I0 DL The body musculuture is especially thick around the apertires, The gut forms a small ¢losed loop in the posterior end of the body and the rectum extends forward to the atrial aperture to form a secondary loop. A rout, Nt-tupped, enlocarp completely occupies the pomury gut loop as in Polycarpa pediunculate. The small, pear-shaped stomach has internal glandular folds. "The anal border has 4+ some- times indented rounded Johes. ‘The gonads are Nal, Nask-shaped ovaries opening by a short, wide, oviduct directed towards the trial aper- ture and overlying two tows of testis follicles with 5-6 follicles in each row. The ducts of each destis [ollicle join a common duet that runs along euch side of the avary to join logether and open into the peribranchial caviy on the mesial aspeet of the shurt oviduer. ‘Whe gonjds are embedded in thick body wall, There are up to S of these gonads on the Jeft side of the body, usually in a single Tow anterior fo the gut loop. On the right, there gre up to 9 gonads usually arranged in 2 rather irregular rows, Remarks: This species is very reminiscent of Polycurpa pedunculata. In view ef the tight ugyregates and confluent test and the absence of sand between adjacent mdividuals. how- ever, il is clenr that colonics form by vegetative reproduction rather than by aggregation of a number of solitary individuals. The process af vegetative budding tu form these colonies is probably associated with the small size of the mature individuals, thus limiting the number of imternal longitudinal vessels in the brarchial sac and the number of gonads on the body wall, Polyandrocarps tapidosy = (Herdman). 1952; 250. Millar, 1963: 730, Ciendyiric: lapidosa Herdman, 1899: 99. New Recered: Investigator Strait (Stn. Y19), Previews Records: Vio Port Phillip Heads, Westemport)—Millar 1963. NS.W, (Port Jackson) —Henlman 1899; Kott 1952. Deseriprian. The present specimen is a large, sandy. plate-like colony with the upper surface slightly concave. 9 em in diam, with a maxi- mum thickness of 2 cm, fixed by a large pan of the basal surface. The surface of the colony is smooth und sandy wathout conspicitous Kott, ASCIDIANS OF SOUTH AWSTRAITA Th swellings or furrows. The zooids are long, but both openings are on the upper surface, There are 4 long branchial folds of very varying height and internal Jongitudinal vessels are artanged according to the following fornmta: DL (10)1(3)1(7)1(4)1 EB The gut extends in a simple are from the postenor end of the body to the anterior atrial aperture, There are 12 internal longitudinal stomach folds. The gonads are elongate and arranged along either side of the endostyle. Subfamily BoTRYLUINAE Botrylloides Jeachi (Savigny}. Kott, 1972a: 29 and synonymy. New Records: Elliston Bay, off Waldesrave [. For Previous Records, Descriptian, sec Kon 1972a, Botrylloides magnicoecum Jtarimeyer, Kott, 1972a; 30 and synonymy. New Records: Waldegrave 1,, Pearson I. For Previous Records. Deseriprion see Katt 1972, Rotryllus schlosseri (Pallas). Kotl, 19724: 31 and synonymy. New Record: North of Waldegrave [, For Previous Records, Description, see Kott 197 2a, Subfamily STYFLINAE Styela plicata (Lesueur), Kott, 1972b: 239 and synonymy. Ascidia plicate Lesweur, 1823: 5. New Record: Coflin Bay. For Previous Reéeords, Description, sec Katt 1972b, Styela pedata (Herdman). Polycarpa pédata Herdman, -188l: Tl, Tokioka, 1958; 322 and synonymy, Kott, 1964; 137, Pardocia pedata Hartmeyer, 1360, Stvela whitelegit Kott, 1952: 213. Telhyuent whitelegii Hartmever, 1909-112 1364. New Recordy: Off Waldearave I,, Pearson I, Preyious Records: N.S.W, {Port lackson, Port Stephens, Port Cnrtis)—Herdman 1899; Kott 1952. Qld. (Moreton Bay, Great Barrier Reet|—Hastings 1931; Kott 1964, Indonesia—Pizon 1908. Phitlipines—Herd- 1909-11: 184 man 188l: Van Name 1913. Tokioka 1958, FIGS. 49, 30 Description: The preserved specimen is orange, the test ix tough and leathery, with longitudinal ridges without any foreign bodies attached. The body is of characteristic shape. expanded postero-dorsally, Both apertures: are directed upwards and the branchial aperture, on a siphon continuous with the upright ventral surface, is often recurved, The atrial aperiure iv also on a short siphon above the postero- dorsal expansion of the body. Basally the test Is extended into prop-like roots. The body wall is muscular. The dorsal tubercle iy oval with numerous separate circular openings giving it ®& porous, sponge-like appearance. Up to 21 internal longitudinal vessels are evenly spaced on the folds, and 3-7 internal longitudinal vessels are present between the folds. There are 6-8 stigmata per mesh, The wut forms a fairly wide loop with tall endo- carps enclosed by the loop. The stomach 3s ciliptical with longitudinal internal glandular folds, The anal border is lohed. There are 3 beanched and ramifving gonads on the left ubove the gut loop und up to 2 on the right. The gonads are embedded in and eccupy most of the body wall cxcept where the gut is present on the Jeft. Tall endocarps are present over the body wail between the branches of the gonad. The yvonads have testis follicles closely applicd to either side of the ovary. Remarks: The gonads and endocarps of this species are very similar to the much branched and ramifying gonads of S. ramiificata Kott, 1952, which has also heen recorded from Moreton Bay. S. ramificara is, however, a very much smaller species heavily encrusted with sand and shell, with a continuous V-shaped Opening on the dorsal tubercle, a narrow put logp and a longer rectum than in the present species. A similar dorsal tubercle is present in Polyearpa aurata (Quoy & Gaimard) (see Van Name 1918) which is also similar to the present sptcics im other characters. It is dis- tinguished mainly by its short, typically poly- carp gonads in contrast to the long ramifying stvelid gonads of the present species, Sryele pedata, therefore, has a wide distribution from the Phillipines and Japan and around the custern seaboard of Australia to the Great Australian Bight. It overlaps with P. ayrerfe in the Phillipines and on the east coast of Aus- tralia. Japan— 186 Polycarpa tinctor (Quoy & Gaiimard), Kutt, 1964: 134 and synonymy. Ascidia tinctor Quoy & Guimard, 1834: 608. New Record: Off Waldegrave T. Previouy Records: see Kott 1964, Description, A single specimen only is avail- able. tt is large and shyhtly damaged, The test, however, has the usual hard, brittle, sand- encrusted lorm characteristic of the species, The specimen tg laterally flattened, with the wiritl aperture hall way down the dorsum, The apertures are sessile. The dorsal tubercle is Jarge with a complementary slit, The branchial sac has 4 very narrow folds. Gonads were not detected in the present specimen, Remarks: The present specics had not pre- viously been taken further south than Port Jackson. It does occur commonly on the north- West and north-east Australian coast, in the East Indies, and off Japan. The form of the dorsul tubercular slil, broken into several parts, has previously been described in speci- mens from Japan and. the East Indics. Polycarpa pedunculata Heller. Kott, 1972a: 35 and synonymy, New Records: Blliston, N-W of Waldegrave I., 22 m off Waldegrave L, Pearson 1., Inves- tigator Strait (Stns. X9, 15, 19, 21, 25, 27; YG, 12, 23; Z9, 11), Emu Bay {Kangaroo L.). For Previous Records, Description, see Kott 1972a. FIG, 51 Remarks; Specimens in this collection demon- strate the full range in external uppearance, from sessile to stalked individuals, from brown- ish to black individuals, In general, the stalk of specimens froni Investigator Strait is longer than that found mn specimens from ‘St, Vincent Gulf, and many of the specimens are super- ficially very similar to specimens of Pyure scoresbiensis with which they occur, demoan- strating convergence in their external appear- ance relalecl to the environment, Polycarpa clavata Hurlmeyer, Kott, 1972a; 33 und synonymy. New Retwray: Walldegrave 7, 22 m off Waldezrive L, Pearson T., Investigator Strait (Sta. Y¥21). For Previous Records, Description, see Kott 19728, Family PYURIDAB Pyura spinifera (Quoy & Gaitiatd), Michael- sen, 1922: 390 (part: Port Jackson speci- tens}, Katt, 1952: 269; J972a: 39. PATRICIA KOTT Ascidia xspinifera Quoy & Gaimard, 1834: 617. Boltenia australiensis Carter, (885; 197, Bolrenia niberculata Herdman, 189: 511; 1899; 17, Cynthia multiradicara Herdman, 1899: 30. Bolrenia spinifera Michaelsen, 1YO5! 72 (part: not B, gihbosa) , Bolrenia spinesa var. tntermnedia Michael- sen, 1908: 390, Pyura glbbasa var. intermedia Michelsen, 1922; 390, Pyura australiensis 7. tvpica Michaelsen & Hurtmeyer, 1928; 419, New Records: Off Waldescave b., Investi- gator Strait (Stns, YU9, Z?). Previous Recards: W. Aust. {Albunyi—Quoy & Guimard 1834. S. Aust. (St. Vincent Gulf) —Kott [972a, Vie. (Port Phillip Heats, Portland Harbour, Cape Woolami, Bass Strait)—Carter 1835; Michaclsen 1995. NSW. (Mort Jucksonj}—Herdman 1899; Kott 1952. FIG, 52 Remarks: Minute scalelike spines are present on localised ridges in the branchial siphon. The branched keratose fibres (see Carter 1885) and Michaelsen & Hartmeyer 1928) are pre- sent in the body wall and stalk. They are similar to spicules found in Pyura stolorifera (see Millar 1962), The specimens are invar- iably covered with a sponge (Halisarca; sec Carter 1885). Pyura australis (Quoy & Gaimard), Pyurq australis subspecies. auxtraliv Kel, 1972a: 39 and synonymy. Pyura «australis var, parvispinalis Kott, 1952; 26%. Baltenia gibbasa Herdman, 1899: 19. New Records: Off Waldegrave 1, NW. of Waldegrave I., St. Francis I, Pearson I., Investigator Strait (Stux. X27, ¥21), Emu Bay (Kangaroo 1.). Por Prevlous Records, Descripiion, see Kott 1972. FIG, 53 Remarks: The principal character on which the distinction between the two subspecies, P. aius- tralis typica and P. australiy purviypinatis, was based is: the condition of the anal border. A careful ¢xamination has shown that in this group of species Lhe lerminal part of the rec- tal wall always has flat folds projecting into the ASCIDIANS OF SOUTH AUSTRALIA lumen as describe for Po australis parvise pindtiy Kott, (952. These folds are usually extended beyond the anal border mto rounded Jabes which are often Jong and finger like and sometimes subdivided, Occasionally. however, they do not extend oitside the anal tim. There is no constant condition observed for any group of specimens in regard to the anal hor- der and it is concluded that subspecies ara wot indicated. The slight difference in the length of the branchial spines ¢(Kott 1952) is ot signifi- cant, These are conical and only slightly curved and arise from a long oval hase. 0.02-0.05 min Jong, and their length along the spine ts 0,020,04 mm. The largest spines are hearest to the upertures, Typical stellate spicules, 0.02 mm in dium, wre always present in the test und siphonal lining and distinguish the species. Pyora pachyderinatina (Herdman) $,sp, draschii nom, nov, Boltenia pachvdermating Herdman, 1881; 81. Drasche, | 884: 370. Herdman, 1899: 16. 7Herdman & Riddell, 1913: 875. Pyura pachydernmutrina var, gibbosa Kott. 1952: 265. New Records: Waldegrave L. Elliston Bay. Previows Records: W. Aust (Cottesloe to Albany}, Vic, (Julia Percy 1., Flinders, Walkerville)—Kott 1952. NISW. (Kiuma, Port Jackson )—Drasche 1884; further speci- mens in Austedliun Museum: FIGS, 54, 55 Bescripion: Typical specimens with dumb-bell shaped spicules and fringed anal lobes, The siphonal spies are produeed into a long pointed hase which js distinct from the rounded base in P. australis, or the flattened scaletike base in P. xpinifera, Michaelsen (1905, 1908, 1922) and Michaelsen & Hartmeyer (1928) utiempted rationalisation of the relationships of the spegics Pye pachydermatina, P. gibbose and P. spinifera and their synonyms, on the basis of external appearance, condition of the Salk and of the dorsal tubercle, and the presence or absence of spicules. A study of the group in Australia has shown that the form of the spicules and siphonal spines, supported, withio certain Jumits, by the form of the anal border, provides. the only reliable character on which to distinguish the species, Michelsen & Harnmeyer (1928) characterise FP. pachy- dermatina by the presence of a smooth anal border, complicated dorsal tubercle, and dumb- bell shaped spicules. This is anly true, how- ever, for the New Zealand subspecies FP. 187 pachydermatina typica since although the duorb-bell shaped spicules ure wlways present, the Australian forms never have a smooth anal border. The dorsal tubercle may be a Jess com- plicated double spiral cone. Thus Pyara gib- hose; Michaelsen & Hartmeyer 1928 (type speciinen: Cynthia sibbosa Heller, 1878, from Bass Strait) with anal Jobes and dumb-bell shaped spicules falls within the definition of P. pachydermatind and Heller's specimen be- comes the type of the subspecies P. pactiveder- nuninde epibbosa (net P. pechydermartine var. gibhosa Kott, 1952), Pyura gibbose intermedia Michaelsen, 1922 (>P spinifera intermedia Michaelsen, 1905) trom Backstairs Passuye, S, Aust, together with P. spinifera typica (part: P. tuberculaia Herdman) from New South Wales, neither of which have dumb-bell shaped nor stellate spicules, anc consequently syn- onyms of P. spiniferd, rather than af P. pacliy- dermatina gibbova, Pyura pachydermatina draschii is distinguished from P, pachiyder- matine gihbosa (>P; pachydermutine inrer- viedia; Katt 1952) (which has shallow anal lobes) by its fringed anal border. P. Pachyeer- marina gibhasa overlaps the present subspecies from Bass Strait to Port Jackson, and extends Further to the north. The western limit for P. pachydermatina draschit is not at Walkerville, Victoria, as Kott (1952) had suggested, since it extends to Western Austialta. Pyura scoreshiensis Kott 1972a: 36, New Records: Pearson L, Investigator Strait (Sens. MM it. 15, 17, VIA) Previous Records; see Kott |972a, Description; Specimens available show a wide range in the length of stalk and in the develop- ment of the rooting processes at the base of the stulk. These sometimes form large sandy lamellue as the roots spread and sand adheres to them, The apertures are always close together on the upper surface, although the pronounced ridge between the apertures. pre- viously descrbed (Kott 1972a) is not always present, Pyora irregularis (Herdman), Kott, 1972a: 38 and synonyrny. New Reeord: Investigator Strait (Sen. V6) For Previous Recerds, Description. see Kott 1972a, Pyora tendata asp. Type Lucation: ltwvestigator Strait (Stn, ¥21), 30 m, scattered, low, on pebble reef 1X8 Figs. Fig. Fig. Fig. Figs, Figs, Fig. Fig. 49-50. 51. 52, 53. 34, 55, 56, 57, 38. 59. PATRICIA KOTT We Danie Styela pedata. Fig. 49.—Individual. Fig. 50—Gonads from left side of body. Polycarpa peduncalata, Gonad showing flask shaped ovary and testis follicles. Pyura spinifera, Fig. 52a—Branching spicules from body wall. Fig. 52b—Branchial spines, Pyura australis. Spicule and branchial spines. Pyura pachydermarina. Fig, 54.—Dumbell shaped spicule and branchial spines. Fig_,55. —Anal border showing internal folds from rectal wall. Pyura tendata, Fig. 56.—Longitudinal section to show sandy coaling around body.-—Fig,. 57—Body opetied around ventral surface, branchial sac removed to show branchial velum, tentacles, dorsal ganglion, gut and gonads: branchial sac (bs); body muscles (bm); branchial velum (v), branchial tentacles (bt); dorsal tubercle (dt); dorsal gang- lion (dg); dorsal lamina (dl); intestine (i); gonad (g); base of the atrial siphon (as); anal opening (a). Herdmania momus. Individual. Malgula ellistoni, Gonads, gut loop and kidney on inner surface of body wall. ASCIDIANS OF SOUTH AUSTRALIA II with sand patches, J. Wain. Holotype: NMV, H1L56, FIGS, 56, 57 Desevipitons The species appears to occur in aggregates. The holotype, however, is the only complete specimen available and only small parts of at least two others were obtained when the specimen was broken [rom its substrate, The outline of the individual iy not obvious superficially, since jt js completely surrounded by a sandy cout 1 cm or more thick. The test is verv thin, without wrinkles, and minute hair-like extensions of the test are scen extend- ing across a narrow space belween the sandy coating and the test, It is probably these hairs which enmesh the sand forming the thick coal ing around the body, The narrow space between the outer surface of ihe test and the sandy coating is occupied by various commen- sal worms and echinoderms, The animal itself consists of a club shaped hody narrowing to a long terminal branchial siphon, The atrial siphon fs twice the length of the branchial siphon and extends forwards from the dorsal border of the body in the anterior part of the posterior third of the body length and opens level with the branchial open- ing, Excluding the sandy coat, the body is 1 cm deep from its dorsal border at the base of the atrial siphon to its ventral border. The length of the atrial siphon is 2 cm, Owing to the gradual narrowing of the body to the base of the branchial siphon, the body appears to have two long diverging siphons of equal length, The body wall is very muscular, Branchial tentacles are present at the base of the branchial siphon half way between the exter- nal aperture and the hijse of the atrial siphon. Just anterior to the ring of branchial tentacles, which are 3 times branched, there is a large muscular velum protuding into the lumen of the branchial siphon. The dorsal tubercle is in the base of a very long warrow peritither- cular area extending half way down the branch! sac, Tt is u very small, simple UL The clongate dorsal ganglion is associated! with the base of the atrial siphon just posterior to the dorsal tubercle. There are 9 branchial folds with internal longitudinal vessels arranged as follows: E (4)0(7)0013)1(15)2(1.6)3(2)1016)2013 07) DL The gut forms a simple loop in the posterior end of the body. There are minute, branched, liver tubules in the pyloric region and the body wall covering the distal portion of the 189 Tectum is produced into a pronounced non- muscular arial velum so that the bilabiate anus actually opens into the base of the wtrial siphon beyond this velum, The branchial velum bas very strong circular muscles in its basal hall, The body musculature i very strong, with outer circular bands forming an almost con- tinuous coal external to strong well-spaced longitudinal muscle bands. The gonads consist af about 7 or 8 pairs of polycarpike sacs vither side of central ducts in the gut loop on the left amd in a corresponding position on the right side of the body, Remarks: This quite extraordinarily modified species is in most essential aspects typleally pyurid and resembles Pyara cancellaia Brewin (see Kott 1971) in the thick sandy coating created by sand enmeshed by the lony test hairs all around the bacdy, Its most conspicuous character is the very long atrial siphon and the numowinyg of the anterior part of the hody so that the branchial aperture is also pro- duced upwards to u level with the atrial siphon, so that the incurrent ciliary slream is not oh- structed by the sand being accumulated around the body, The thick, sandy coating is so dense and so rigid that it is hard to imagine how the animal is able to increase in size. It is probable, however, that the commensals present between the sandy coating and the test constantly irtt- gate this ares and thus maintain the spice into which the animal can expand as it grows These commensals are therefore probably essential to such sand covered species. (see also Pyvra cancellata; Kot 1971). Halocynthia hispida (Herdman). Kotte 1968; 76 and synonymy; 19772a; 4]. New Record: Investigator Strajl (Stn. X19). aoe Previous Records, Description, see Kot 1963_ Herduiania momus (Savigny). Koti, 1972a; 41 and synonymy. New Records: Off Waldegrave I., St, Fran- cis T,, Pearson L, Investigator Strait (Sto. X17). Spencer Gulf, For Previouy Records, Description, see Katt 19724. Reewrks: There is the usual great range In size ot individuals, Smaller specimens are mere or less upnght with transparent test and short furrowed siphons. Larger specimens become faccid and opaque. Ctenicella antipoda Kott, 1972a: 44, New Record; Investigator Strait {lubcl Hlegble). For Previews Recards, Dexerip- tien, see Kott 19724, 7490 Remarks: The distal part of the gut loop is distended with mud. In ong large specimen the fohuds on the left side of the body are enclosed in the gut loop. In view of the agreement with the type specimen in all other aspects, this must be regarded as. an individual variation. Mokguta eflistoni n.sp. Type Loedion: Elliston Bay, in caves out- side bar, subject ta strong swell, 14.v.L971. Holowpe and Purai'pest SAM, E907. FIG, 59 Description: Small sandy spherical individuals were found adhering to Evlerdmania ausrreis und sometimes forming aggregates. Both aper- tures are present fairly close together on the upper suttace and are directed away from one another. VYery delicate muscles radiate out trom the siphons. There are internal circular muscles around the branchial and atrial siphons. ‘There aro 7 branchial folds on each side of the body, with f meshes along each fold. Each mesh contains a primary spiral in- Fundibulum, which subdivides into two half way up into the fold. here are 3 internal longitudinal vessels arranged along the fold, There is the usual long gut loop open at the pole and a short curved kidney on the right side of the body. The gonads consist of a cireular ovary with a short, wide, duct directed dor- sally. Testis follicles are present along the proximal border of the ovary and join into a very shore vas déferens which opens into the penbranchial cavity on the mesial surface of the ovary. ‘the right gonad is in the middle of the hudy wall and the left gonad is in the secondary gut loop. The specimens are abour )}S cm in diam, Renwrks: The position add form of the gonads with the very short vas deferens opening on the surface of the ovary are distinctive. Anural embrycs qre present in the peribranchial cavity, Molgula subolosa Quoy & Gaimard 1834: 613. Koll, 1972b: 248 ancl synonymy. New Revord: Elliston Bay, For Previous Records, Deyerintian, see Kotl 1972b. CharucterisGes of the Faun Afi outstanding feature of the fauna is the large number (23) of species in the relatively primitive Aplousobranchis. Polycieridae, Clayvelininge und Euherdmaniimae, in which PATRICIA KOTT common cloucal systems ive not developed. A further 7 species ure secondarily colonial species of Styelidue; one is 4 colonial phicbo- branch species: three ure aggregated pyurids: and there are two aggregated spccivy of the family Molgulidae. Thus about half uf the species present, although colonial in habit. pre- serve theit own independent. openings and do not form cloacal systems. They clo, however, demonstrate rematkuble morphological adap- tations in the arrangement, orientation and operation of them apertures ta maximise their Teactions. with the environment. Colonies of Ritterella herdmania, Plucentela ellisien’ n-sp., and the new colonial phlebobranch species are of special interest in that the “Zooids ure arranged in parallel, so that their branchial and atrial apertures are respectively loeatecl on different sides or at different ends of the ealony. Some benefit can accordingly be derived from mutual reinforcement of feeding currents. One would expeet that the exact erientalion of these colonies would be such that prevailing currents could also reinforce the incurrent ani excurrent ciliary streams, These species and many of the Clavelininac. together with the new species Merandraecgrpa indice, Syinplegma arenesa and Stolenica trim- cata, all have specially adapted body muscu- Jature to operate sophisticated siphonal uppara- tus, Adaptation ol the nervous system. to serve the specialised musculature can algo be expected, In fact, im both Pyura fendata and in the new colonial phlebobranch species, the feucal gland anid ganglion ure localed pos- teriorly in association with an unusual posterior position of the alnal siphon. None of the species discussed ubove ure enefusling ant most favour habitats under ledges and in crevices or caves. There are anly 18 species with highly evolved cloues! systems that have jichieved a degree of inde- pendence of the enviranment and, with only 4 eavenlions (Didemanm cendidum, D. moselesi, Trididemnum spiculatuoc and Bowryllus schloy- sere), these jire jilso butky or stalked (rather than enerusting) and favour habitats under ledges or in crevices or caves. Phe majority of the solitury stolidobranch specics that are present are stalked and able fo maximise their seactions with the environ- ment hy moving on their stalk with the current. There are only few solitary botton-living species (Asetdia spp., Polycarpa rnetor, Crenicella antipoda) or large fixed species (Merdmiania mamus, Styela pedata), ASCIDIANS OF SOUTH AUSTRALIA II yi39 GHENT AUST HALL AN FLAT ize 3350 1? SOUTH AUS RAL i Fig, 60. Map of South Australian locations from which specimens are recorded. Zoogeography The known ranges of Atapozoa marshi and Pseudadistoma australis have heen extended from Western Australia to the eastern end of the Great Australian Bight where they overlap with Distaplia distomoides, Euherdmania aus- tralis, Pseudadistoma cereum and Polyandro- carpa laupidosa, whose ranges have been simi- larly extended {tom the easi. The ranges of Aplidium rubricellun: and Palyclinum nep- tunium from the west, and Ascidie thampsoni, Pyura irregularis and P. stolonifera (see Kott 1952) from the east. are also known to extend into either Spencer Gulf or St. Vincent Gulf. Thus the distribution of these ascidian species across the south coast supports the existence of a marine faunal boundary at the eastern end of the Great Australian Bight, separating the Flinderstan marine faunal Province from the Maugean Province to the east (Knox 1963). There is less eyiderice however for a western boundary of the Flindersian Province further south than Cockburn Sound in Western Aus- tralia. It is possible that the 15 new species described herein will subsequently be found to have wider ranges across the Australian coast und provide further data relating tao the western boundary of the Flindersian Province ‘to help resolve the question of the existence of a Western Australian Province (Baudinian of Kott, 1952; see Knox 1963). The ascidian fauna of Gulf waters of South Australia includes only a limited number of the species that occur along the open coast. Of the species discussed. above which terminate their range off the eastern South Australian coast, only Aplidium rubricollum, Polyelinum neptunium, Aycidia thompsoni, Pyura irregu- laris and P, stolonifera extend into guilt waters, Podoclavelia. meridionalis, Distuplia ytylifera, Didemnum ternaranum, Styela pedata, Poly- carpa tinctor, and Pyura pachydermutina are also apparently absent from gulf waters although their geographic range around the Australian coast is wide, It is probable, there- fore, that some ecological factor inhibits the bye spread ot the ascidian fauna Erom the open const Inte gulf waters. It is possibly the same factor Usar favours the occurrence af Distaplig viridis, Aplidium pliciferum, Bideninum lani- bitin, B. psendodiploxoma, Leptoclinides rufus, f, kKingt, Eehinaclinum verrilli, Asetdia aclara, Palvearpa papillata, Pyura cataphracta and Microcosmus nichollsi in St. Vincent Gulf and other eimbayments to the east and nerth around the Australian coast, although they have not been recorded from the open coast. The ascidian fauna of gulf waters of South Australia is therefore distinet from that of the adjucent cogstal waters and has no special zoogeo- graphic affimty with either the Flindersian or the Maugean Provinces. There are & further 42 species discussed herein that are recurded from both coastal and eulf waters. The majority of these have a Wide cusmupulitan distnbution or an extended PATRICIA KOTT range from Western Australia to cither the houndary of the Maugean marine faunal revion in Bass Strait (Hedley 1904, Knox. 1963). or further up the castern Australia coast. Also included in this group of species, however, are Crenicella antipada, Pyura scoresbiensixs and Atapezoa fcititeustana, previously thought en- demic t St. Vincent and Spencer Gulfs, and Aplidiuin colelloidey (see Kot 1972a)- Acknowledgements Grateful thanks are due lu Mrs. }) Watson, who collected the material from Investigator Strait for the Nutional Museum uf Victoria, and Mr. S$. A. Shepherd. of the South Austra- ian Department of Fisheries and Fauna Con- servation, who made the collections from mest other locations. ‘The work was done while the author was the recipient of A.R-G.C. Grant D65/ 15386. References Beowin, Breve |, (1950)—Ascidians of New New Zealand, Part V. 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Senckenh. aaturforseh. Ges 24, 64)—-666 TakiMerver. KR. (1909=11).—Aseidien, (Continua- tion of work by Seeliger). 7 H. G. Bronn, “Klassen und Ordnaungen des Tier-reiches.” 3, suppl (89-98), 1281-1772, (Leipzig. [Ab- stract, repeating Iists of species by A. Schenotieff in Areh. Naturgesch,, (911, 6, 3-271. Hawimerer, R. (1912).—Die ascidien der deut- sehen liefsee Expedition, Wiss, Ereehn. dr, Tictsee-Exped. ‘Vaktivia® 16 (3), 223-392. HartmMever. BR. (19133,—Tunicata. Jn L, Schultse, Zool u. anthrop. Ereebnissee Forsehunpsteise In Stidnfriks. Rd 5, L.ft 2. Denksehr med. ralarw. Ces, Jena VF, 125-144, Aanimeven, R, 1 1919).—Ascidien. Results of Dr EB. Mjébere’s Swedish scientific expediliuns to Australia I91(E13. K_ svenska Verensi-Akud, Hondl, 6) (4), 1=150. Hasiinus, Anna B. (1931).—Tunicata, Sevens, Rep, Gt Barrier Reef Exped. 4 (A), 69-109. Hiotey, C2 1190d). —The effect of the Bitsian Isthmus upon the eaisting maine fauna; a sludy in ancient geography. Pree. Linn, Soo, NSW, 38. 876. Hreooce. C. (1878) —Beitrage zur niihern Kennt- nisé der tudicaten. Sher, Akad, Wise Wien T7 (1), 835-110. HERDMaN, W. A. (1881.— Preliminary report on the Tunicuta of the Challenger Txpedition. Cynthiidae. Molgulidae, Prec Ro Sew. Rilinh, WV (3). 52-88: (4), 233-240. Hrruman, W, A., (1891)—A tevised classifien- tion of the ‘Tunicata, with definitions of the orders, sub-orders, fumilies, sub-families ind wonera, and analytical keys to the species, 3. Linn. Soc, 23, 558-625. HERDMAN. W. A, (1899)—Descriptive catalogue of the Tunicala in the Aastrilian Museum. Aystallon Musenm, Sydney. Catslogie 17, f-139, Hrepman, W, A, and Rippris, W. (191399.—The tunicula of the “Thetis” expedition. fe Scien- tific Resulls of the Trawl Tixpedilion Thetis. Mem. Aust. Mis, 4, 873-889, Kasox. G. A. (1963).—Biogeography and inter- tidal ccology of the Australasian coasts. Ocetimogr. mar. Biol. ann. Rev. 1 341 404. KoTT, Pataicta (1952)—The ascidians of Aus- tralia, t. Stolisiebranchialta and Phlebobran- chlata. Aus. J. mar, Fresh, Kes, 3 (4), 206-333. Rott, Patricw (1957),—The ascidians of Aus- tralia Tl, Aplousobrunchiala Lahilte; Clave- linidae Forhes and Hanly and Polyclinidac seit Aase J. our. Beeshw, Hee BOOED, 64-1Tf- ASCIDIANS OF SOUTH AUSTRALIA Tl 193 Kory, Parrter (1962).—The ascidians of Aus- tralia 111, Aplousobranchiata Lahille: Didtm- nidne Giard, Aust J. ear. Freshw, Res, 13 (3), 265-334, Kort. Patricia (1963).—The axcidians of Aus- tralia JV. Aplousobranchiata Luhille: Poly- clinidue Verrill (continued), sfast. J. mar. Fyeshw. Res, 14 41), TO-118. Kotr, Patrician ('964),—Stolidohyanch and phlebobranch sscidians of the Queensland coast, Pap. Dep. Zak Univ. Ga 2 17), 127-182. Korr, Parricia (1966)—Ascidians of north Aus- tralia. Pay. Dep. Zool, Univ, Ged 2 (19), 279-304, Korr, Patric. (19671..-dtupeged (sic) deerara (Sluiter): A discussion of the relationship of the genus und species. Prac. Linn, foe, N.S.W. 9D (3), 185-188, Kort, Patric (1968) —A veview of the genus Alaloeyethia Verrill, '879. Prav, Linn, Sac. N.S.W. 93 (13, 76N9, Karr, Patricia (19692).—Anturctic Ascidigces, A monographic account of the known species based of specimens collected under 1S. Government suspices 1Y47 to 1963. Anturct, Res. Ser. 13, cexv, 1-239, Kort, Patricia (1969b),—A review of the family Agnesiidac Huntsman 1912, with spacial reference to Aunesia glaciata, Proc. Linn. Sac. N.S.W’. 93 (3), 4444456, Kot, Parraicrs (1971 }—Antarctic Ascidiacea It. Callections made south of 40° south latitude 1963/67, principally by the USNS, Truain. Abtaret. Res. Ser. Biology 16 (4), i-iti, 1-60. Kort, ParnicrA (1972a).—The ascidians of Sonth Allstraha I; Spencer Gulf, St. Vincent Gulf, and Encounter Bay. Trans. Ro Suc. $. Aus, 96 (1), 1-52. Kort. Patricia (1972b),—Some sublittoral ascid- jans in Moreton Bay, Queensland, Mem, Mfrs. Gd, WG (2), 233-260, Kort, Patricia (in press. | )—Plurellidse, o new phtebobranchiate family of the Ascidiacea. Proc. Linn. Sec. NS. Kory, ParriciA (in press, 2),—Notes on some Ascidians from Port Jackson, Botany Bay and Port Hacking, New South Wales. Proc, Linn. Sov. NUS W, Kors, Parricté (in press, 33,—Aaciians from Westernport and adjacent locations. Mem. nate. Was, Vier 34. Kowarrysky. A. OL (1874),—Ueher die Kags. pung der Ascidien Arety. mikrosk, Anes. Earw. Mech. 1 (1), 444-470, Lesunur, C. A_ (1823 3—Deseripilans of several new species of Ascidia. J. Ace jar. Sei, Philad, 3, 2-8. MICHAPLSEN, W, (1905) —ReVision yon Heller's Ascidien-Typan aus dem Museum Godoffrai. Zool. Jb. (suppl,) 8, 71-120. MICHABLSEN, W. (1908).—Die Pyuriden (Hale- cynthiiden) des Natirhistorischen Museums zu Hamburg. Jb, hamb. wiss, Amt, 25 (24, 227-287. MICHAELSEN, W, (1920).—Die krikwhranchen ascidien des westlichen Wndlisehen ozeans: Didemniden. Jb, har, wiss. Anst, 37, 1-74, MICHSELSEN, W. (i922) —Ascidiae Ptycho- branchiae und Jiktyobranchiae van Neusee- land und dem ChathamtInseln. (Papers fram Dr. Th. Moretensen’s Pacific Expedition |914- 16, No, XL). Fidensk. Meddr dausk spareri, Foren. 73, 359-498. MicHaeceen. W. €1924).—Ascidiae Kriky branchiae von Newseeland, den Chatham und den Auckland Insetn. (Papers from Dr, Th, Mortensen's Pacific Expedition 1914-16. No. XXIE) Fidensk, Meddr. dunsk — retarh. Poren, 77, 263-434. MicHABLSEN, W. (1930).—Ascidiae — kriko- branchiae, Faunu Stdwest-Aust, 3 (7), 463- S58. MichHaetsen, W. (19341—The ascidians of the Cape Province of Sotth Africa. Vrans R, Suw. Afe. 22 (2), 129-163. MICHAELSEN, W., and Hantmeynr, R. (1928 )— Ascidiae — diktyobrunchiac und — ptyche- branchiae. Fauna Shdwest-4 rest, S, 251440, Muzak. R. H. (1995),—On a collection of ascidians fram South Africa. Pram, zaal, Sac. Lona, 125 (14, 169-221. Mittar, R. H. (1960) —Asvidivcaa, 'Diseovery' Rep. 30, 1-60. Mrizar, R- H, (1962.)-—Further descriptions of South African ascidians. Ana. S. Afr Mus, 46 (7), 113-221. Mitcan, R. H, (1963),—Australian ascidians in the British Museum (Natural Histery). Prac. zool, Sac. Lond. 141 (4), 689-744. Mactan, R. H. (1966).—Ascidiacea, Part Philip Survey, Mem, nata. Muy, Viet. 27. 387-375 Nast, J. T. (1892).—O the composite ascidians of the North Shore Reef. Trans. NZ. tease. 24, 305-334, Peres, J. M. (1952) —Ascidies de Ia réche [it- torale Corse. Rec. Star. mar, o'Endawme 6. 35-44, Pizon. A. (1908) —Ascidies Suisse Zoal. 16, 195-248, Quor. J. & Gaimarp. A. ¢)934),—Vayuges de découvertes de VAstrafabe 1826-29, Mollusques, Zoolozie 3, 359-626; 4, 304306. Saviany, 3. C, ¢1816)—*Ménioires sur tes animaux sang vertebres,” Pt 2 (Pans, 4 Srurer. C. PL (1898),—RBeitriige zur kenntniss die Fauna von Sidafrica Il. Tunicaten. Zeal Jb, (Systematik) IL, 1-64. Stuer. C, P, (1909),—Die tunicaten der Siboga Expedition Pt. 2. Die merosoynen ascidien Sihaga Exped, 56%, 1=112. Trkioka, T. (1949).—Contributions 40 the Japanese ascidian Fauna Ti, Notes on some ascidians collected chiefly along the coasl of Kii Peninsula. Prhls, Seto mar, bel, Lak 1 (2), 39-64, TOKIOKA, T. (1958)—Contributions to Japanese ascidian fauna XTI. Sporadic memoranda, Publy Sete: mar. hiol, Lah, 6 (3), 31325, Voxions. To (1959) —Amuareucium ervthrueum Michaelsen, a comoound Ascidian from the Cune Province, (Blolozical Resilts of the Jupanese Antarctic Research Expedition 5}. Spec, Puhbls Seta mer, Diol. Lah. (329). d'Amboine. Rev 194 Tokioxa, T. (1961).—Ascidians collected during the Melanesia Expedition of the Osaka Museum of Natural History. I, Ascidians pre- sented by Dr, R. L. A. Catala of the Aquarium of Noumea. Publs, Seto mar. bial, Lab. ¥ (1), 104-138, Toxtoxa. T. (1967)—Pacific Tunicata of the United States National Museum, Bull. 1.8. nuin. Mais. 251, 1-242. Trason, W. B. (1963).—The life cycle and affinities of the colonial ascidian Pycno- PATRICIA KOTT clavella stanleyi. Univ. Calif. Publs Zoal, 65 (4), 283.326, Van Name, W. G. (1918).—Ascidians from the Philippines and adjacent waters, Bull, OS, nain, Mus, 100 (1), 49-174. Van Name, W. G. (1945).—The North and South American ascidians. Bull. din, Mus. nat. ITist. R4. 1-476. VassseuR, P. (1967).—~Ascidies de Novwvelle- Calédonie. Editions de la Fondation Singer- Polignac, 127-146, 2 pls. Appendix I—Station List Stations are listed in the order north to south and west to east. DENTAL BAY. (Coll. I. Thomas). Lat. 32°13’S; Long, 133°38’E, Near Ceduna; in Posidonia community; depth 10m; 1965. Mfapuzoa fantayiana Sycacoa cérebrifermix OFF ST. FRANCIS 1. (Coll. 8. Shepherd), Lat. 32°31'S: Long. 133°15'R. Rocky bottom; slight current; depth 55 m; 8.1971, Pyendodisiorma cereum Palyandrocarpa simulans Pyura australis Herdmania momus WALDEGRAVE ISL. BND. eo $. Shepherd). Lat, 33°36'S; Long. 134°46 Strong surge; depth 20 m; a iv.1970, Distaplia distomoides Synaicinm papilliferum Botrylloides magnicoccum Batryllus schlosseri (22 m) Pyura australis Rocky bottom; 23.x.1970, Enxherdenania australis Aplidium colelloides Oculinaria australis Botrylloides leachi Styela pedate Palycarpa tinctor Pynra australis Pyura spinifera (covered with sponge) Pyura puchydermatina Herdmania momius strong surge; depth 22 m; 1 Km N.W. OF WALDEGRAVE T, (ANXTOUS BAY). Lat. 33°33'S; Long. 134°46'E. Rocky bottom; slow current; depth 23 m; 11.¥,1971, Glaveling mirahiliy (attached to limestone). Distaplia distamaides Distaplia stylifera Syinplegsma arenosa Batryllus schlosseri Sloleonica truncata Holozoinae sp. (grawmg on red algae) Polycarpa clavata {growing on brown or red algae: Sargassum or Osmundaria) Herdmania montis (with red algae ajtached) Rocky bottom with sand patches; slow current; depth 23 m: 11.v.197). Podoclavella cylindrica Synoiciunt papilliferum Stolonica truncata Polycarpa pedunculata Pyara australis Racky bottom; 23.x. 1970, Cluvelina nodyla Polyeitor giganieum Pyendodistoma anstraliy Psendodistoma cereum Aplidium flavalineatymn Symplegnia arenosa Polycarpa clavata (attached to rock) Palycarpa pedunculata slight surge, depth 22. m; WATERLOO BAY (ELLISTON). (Coll, S. Shep- herd), Lat. 33°38'S; Long. 134°51'E. Roof of caves; strong surge; depth 6 mi; 13.v.1971. Clavelina bhaudinensis Padoclavella molaccensis Distaplia distomoides Endistoma renieri Pseudodistama cereum Aplidinen lobatium Polysyneraton paradoxtm Didenripn candidum Didemnum maselevi Ayctdia thampsoni Asctdia sydueventsis Polvandrocurpa sitiulans (growing on Ascidia avdueyensiv) Chitside bar; very strong surge; depth 17 mm 12v.1971. Sycozaa cérebrifarmis Polvelinum nepluninm Aplitinm elatum Ocnulinaria australis Pyura pachydermazina Outside bar, in caves: strong swell, depth 17 m; 14.y.1971,. Pyenoclavella diminuta Fuherdmania rnustralis Ritferella herdmania Placentela ellistont Trididemnaum cerebrifornte diminita) Stolonica australis (around base of FEuherd- mama australis) Molgula ellistoni (adhering to E. Australis) (investing P. ASCIDIANS OF SOUTH AUSTRALIA II 15 Inside bar: depth 6 m; 12.v,1971, Palycitor giganteum Psendodistoma australis Didemnum moseleyt Palycarpa pedurrcilata Inside bar; strong surge; on vertical face; depth 6G m: 12.v.1971, Aplidium pantiterinim Near entrance to bay, in caves; depth 3 m:; l4.v. 1971, Pseudodistona exrnusense Palyelinam neptuniaum Motlenta sabulosa Floor of cave; depth 16 mz: 12.v.1971. Atapozoa mirahilis Centre of bay; depth 11 m; 12.V,1971. Atapezoa mirabilis On root of caves: depth 3-5 m: 14.v.1971, Aplidium flavalineatum Flliston Bay Cystidyles dellechiajei Trididemnum spiculatum Didermmaum ternatanam Oculinaria australis (with EB. M. ellisteni) Ratryllaides leachi australix and PEARSON ISLAND. (Coll. S. Shepherd). Lat. 33°56'S; Long. 134°15’F, Rough-water coast, 400 m offshore on gravelly battom, attached to shell or rock fragments; moderate surge; depth 30 m: 97.1969. Leptoclinides fungiformis Péalycarpa clavata Pyura australis Herdniania mamus Rough-water coast; in caves; moderate surge; depth 35 m; 101.1969. Poadoclavella meridionalis Aplidium rubricoltum Stvela pedata Polveurpa peduncelata Herdmania momus Sandy beitom hetween Dorothee and Veteran Is, Lat. 34°1I'S; Long, 134°15’E. Slight surge; depth 70 mz 11.1,1969. Palycitor giganieum Batrylloides magnicoecum Polxcarpa pedunewlata Pyurg scoreshiensis Outside Pearson 1; depth 35 m, Padoaclavella meridionalis Leptoclinides faungiformiy Styela pedata COFFIN BAY. (Coll. 8. Shepherd). Lat. 34°38'S; Long. 135°30' F. Oyster Trays, at low water; slow currents; no sediments; October 1979. Sivela plicata SPENCER GULF. (Coll. S. Shepherd). Lat. 34°2'S; Long. 137°23'E. ‘Tipara Reef: depth 11 m; 20.vii.1971., Pycnecluvella diminwiv Herdmania mamus INVESTIGATOR STRAIT (Coll. J. Watson). Station X7: depth 30 m: 101,1971. Lat. 35°16’S; Long, 137°30'E. Didemnunt moseleyi Station X9; depth 31 m; 19.j.1971. Lat. 35°17°S; Long. 137°30'R. Palycarpa pedunculata Pyura scoresbiensis Station X11, depth 30 m: 35°19" S; Long. 137°30°R. Pyura scoresbiensis Stalion X15; depth 32 m; 35°23'S: Long. 137°30’E. Polyeitar gigantelm Polysyncraton magnilarvim Polyearpa pedunculata Pyura scaresbiensis Station X?, 19.1.1971. Palvandrocarpa simuyltans Station X17; depth 35 m; 35°24'S; Long. 137°30'E. Trididemaum cerebrifarme Pyura scoreshiensiy Herdmiania momuy Station X19; depth 34 m; 35°26’S; Long. 137°30'F. Ruherdmania australis Colonial stolidobranch—Gen. Axcidia s¥dnevensis Polycarpa pedunculata Haloeynthia hispida Station X21: depth 34 m: 35°28’S; Long. 137°29'E. Polycitor giganteum Polycarpa peduncufata Station X25; depth 35 tn; 35°31'S; Long. 137°29'R, Polycarpa peduncnlata Station X27; depth 31 m: 35°33'S; Long. 137°29'F. Didemaum moseleyi Palycarpa pedunculata Pyura australis Station Y6: depth 23 my; 28.1.1971. Lat. 35°17'S; Long. 137°16'E, Leptoclinides reticulatits Polysyncraion magnilarvum Metandrocarpa indica Polyandrecarpa simulans Polyvarpa peduncilata Pynra irregularis Station Y12; depth 33 m; 35° 23'S; Long. 137°17"E. Palycarpa pedunculate Station Y14: depth 32 m; $5°25'S:, Long, 137°17’E, Alapozou marshi Pyura scoreshicusis Staton Yil6; depth 35 m: 35°26'S; Long. 137°17'E. Sycozea peduncalata Statton Y17; depth 34 mm; 35°27’S8; Long. 137°18'E. Atapozoa marshi Station Y18; depth 31° m; 35°28'S; long. 137°18'E, Oxycorynia arenosa Polycitor obelixcaum Colonial stolidobranch—Gen. & sp.7 Ascidia sydneyensis T9i1971, Tat. WDG1971. Lat. 19.1,1971. Lat, 20.4;1971, Lat. and sp.? 17.i,1971, Lat. 175,197], Lat, 17.41.1971. Lat. 24.0.1971. Tat 233.1971, Lat, DLIO7L. Lat. 205.1971. Lat. 23.1,1971, Lat. 196 Station Y19; depth 33 m; 20.1.1971. 35°29'S; Longe. 137°18'E. Lat, PATRICIA KOTT Station 2? Colonial stolidobranch—Gen. & sp.? Polyandrocarpa lapidosa Polyandrocarpa simulans Pyura spinifera Station ¥21; depth 32 m 35°32'S: Long. 137°18'E. Aplidium colelloides Polyedrpa clavata Pyura australis Pytra tendata Station Y23: depih 32 m; 35°33'S: Tong. 137°18'E. Polycarpa pedunculata Station Y?; 207.1971. Didemnum niaseleyt 201.1971. Lat. 17.4,1971, Lat, Pyura spinifera Stations Z9, 11; depth 38 m; 25.11.1971. Lat. 35°30'S; Long, 137°8’E. Polycarpa pedunculata ? (Lahel illegible) Colonial stolidobranch—Gen, & sp.? Ctenicella grtipoda KANGAROO ISLAND, (Coll. I. FE. Watson). Lat. 35°35’S; Long. 137°31'E. Off Emu Bay, 221,197], Didemnum moaseleyt Didemnum patulum Polycarpa pedunculata Pyura australis Index to Genera and Species Page Aplidium colelloides 176 Aplidium elatum nsp. _ - 177 Aplidium flavolineatum 176 Aplidium fobatum ,, : 176 Aplidium pantherinum .. 176 Aplidigm rubricollum 176 Ascidia sydneyensis 182 Ascidia thompsoni .,,, 180 Atapozoa fantasiana 168 Atapozoa marshi : 168 Atapozoa mirabilis n.sp. 168 Botrylloides leachi - Sete 185 Botrylloides magnicoecum __ 185 Botryllus schlosseri 185 Clavelina baudinensis ,, . 166 Clavelina mirabilis n.sp. 165 Clavelina nodula n.sp. .... T66 Ctenicella antipoda 189 Cystodytes dellechiajei ..., 172 Didemnum candidum ..., 179 Didemium. moseleyi 179 Didemnium patulum 179 Didemnum ternatanum 179 Distaplia distomoides 170 Distaplia stylifera ..,, 170 Budistoma renieri _ - : 171 Euherdmania australis ... 172 Halocynthia hispida 189 Herdmania momus cant 189 Leptoclinides fungiformis n.sp. 180 Leptoclinides reticulatus 180 Metandrocarpa indica n.sp. ..., 182 Molgula ellistoni n.sp. _ 190 Molgula sahulosa . 190 Oculinatia australis 184 Oxycorynia arenosa nsp. 167 Page Placentela ellistoni a,sp. 173 Podoclavella cylindrica 167 Podoclavella meridionalis 167 Podoclavella moluccensis 167 Polyandrocarpa lapidosa 184 Polyandrocarpa simulans n.sp. 184 Polycurpa clavala - - 186 Polycarpa pedunculata ..,, 186 Polycarpa tinctor 186 Polycitor giganteum 171 Polycitor obeliscum n.sp. 171 Polyclinum neptunium . 175 Polysyncratan magnifarvum _ (78 Polysyneraton paradoxum .., 178 Pseudodistoma australis (72 Pscudodistoma cereum 173 Pseudodistoma cyrnusense 173 Pycnoclavella diminuta 170 Pyurg australis 186 Pyura itregularis _.. 187 Pyura pachydermatina 187 Pyura scoresbiensis 187 Pyuri spinifera 186 Pynra tendata n.sp. 187 Ritterella herdmania 172 Stolonica australis 183 Stolonica truncata n.sp, 183 Styela pedata , 185 Styela plicata. ., : oF 185 Sycozoa cerebriformis .... 170 Sycozoa pedunculata 170 Symplegma arenosa n,sp, 182 Synoicium papilliferum . _ 177 Trididemnum cerebriforme ..., 178 Trididemnum spiculatum 178 STRIGEATA (TREMATODA) OF AUSTRALIA BIRDS AND MAMMALS FROM THE HELMINTHOLOGICAL COLLECTION OF THE UNIVERSITY OF ADELAIDE BY GEORGES DUBOIS AND L. MADELINE ANGEL Summary An important collection of Strigeata of birds and mammals has been made by the Department of Zoology of the University of Adelaide. The present work records thirty species, of which seven are new. Descriptions of the new species with additional information on some of the others are given. The new species from birds are: Apatemon (Apatemon) vitelliresiduus (from Biziura lobata), Cardiocephaloides ovicorpus (from Phalacrocorax melanoleucos brevirostris and P. varius), Cotylurus (Cotylurus) magniacetubulus (from Cygnus atratus), Diplostomum (Diplostomum) parvulum (from Hydroprogne caspia and Pelecanus conspicillatus), Neodiplostomum (Neodiplostomum) lanceolaturn (from Ninox novaeseelandiue). The new species from mammals are: Neodiplostomum (Triloborchidiplostomum) diaboli (a new subgenus, for which the diagnosis is given) (from Sarcophilus harrisii) and Pharyngostomoides dasyuri (from Dasyurus viverrinus). STRIGEATA (TREMATODA) OF AUSTRALIAN BIRDS AND MAMMALS FROM THE HELMINTHOLOGICAL COLLECTION OF THE UNIVERSITY OF ADELAIDE by Georces Dusots* and L. MaveLine ANGEL] Summary An important collection of Strigeata of birds and mammals has. been made by the Department of Zouloxy of the University of Adelaide. The present work records thirty species, of which seven are new. Descriptions of the new species with additional information on some of the others are given, The new species from birds are: Apateron (Apatemon) vitelliresidues (from Bittura lobar), Cardiavephaloides avicoraus Grom Phalacrocerax melanoleuces. brevirestris and P. varias), Cotyluras (Corylarus) magniacetubulus (from Cyenns atratus), Diplostomum (Diplostomum) parvalum (from Hydroprogne caspia and Pelecanus conspicillatus), Neodiplostomum (Neodiplostomum) lancealatum {from Ninax navaeseélandiue ). The new species from mammals are: Nerdiplostamum (Triloborchidiplostemum) diaboli (a new subgenus, for which the diagnosis is given) (from Sarcephilus harrisii) and Pharyngostomoides dasyuri (from Dasyvurus viverrintes). Resume Une importunte collection de Sirigeala d°Otseaux et de Mummifares a été constituée au Deéparte- ment de Zoolagie de Université d’Adelaide, Le présent travail comprend Ja description ou ta mention de 30 espéces, dant 7 sont nouvelles: Apateman (Apaternen) vitelliresiduus, Cardiocephaloides ovicorpus, Cotylurus (Cotylurus) magniaceta- Inilus, Diplostomum (Diplostomum) parvelum, Neodiplastomum (Neadiplostomtm) lanceolatum, Ned- diplostamum (Trilehorchidiplostamum) diaboli (n. subgen., dom la diagnose est proposée) et Pharyn- rostumoides dasyuri. Ces deux dernigres espéces sont parasites de Marsupiaux (Dasyurinés), respec- tivemenot de Sarcophilus harrixil et de Dasyurus viverrinus. Introduction An important collection of Strigeata from birds and mammals has been made by the De- partment of Zoology of the University of Ade- laide. It consisted of 92 tubes with spint specimens, and 3 slides, Collections and identifications made before August, 1951, are the valuable contribution of the late Professor T. Hurvey Johnston, to whost memory this work is dedicated. Since that time Mrs. P. M. Thomas (Patricia M. Mawson) has done most of the collecting. One of us (L.M.A.) collected the remainder. Dr. J, C. Pearson, of the University of Queens- land, Brisbane, contributed 5 specimens (Pharyngesromoides dasyuri) from Tasmania. Some of the specimens are valueless, either because of poor preservation or because young stages cahnot be identified with adults. ‘The holotypes of all the new species des- cribed in this paper have been deposited in the South Anstralian Museum (SAM), Para- types, where »vailahle, are in the South Aus- tralian Museum, and in the Helminthotlogical ~ . Collection of the Institute of Zoology, Univer> sity of Neuchatel (G.D_). Preparations of the rest of the material are deposited in the Uni- versities of Adelaide and of Neuchatel. The present publication is a continuation of 3 previous accounts of Australian $Strigeida (Dubois. 1937; Dubois & Pearson 1964, 1967). Thirty species are described or recorded. Seven, one of which belongs to a new sub- genus. are new. Twenty-seven are recorded as u¥ian parasites anc the other three are from Taammals. Family STRIGEIDAE Riuillict Subfumily STRIGEINAE Railliet Apharyngostrigen simplex (S. J. Johnston, 1904). Dubois, 1968: 35, figs, 19-21. Dubois & Pearson, 1965: 79, figs, 1-3, S. J. Johnston, 1904: 112, pl. 7, figs. 1-3- Host and origin; Ardea novachollandiac Latham, from Tailem Bend, River Murray, 5. Aust., 1.v.1940 and 9.xi1,1940 (2 specj- tens); from the Australian Museum, date? {one specimen). *Tustitut de Zoologic, Université de Neuch&tel, Suisse, | Department of Zoolozy, University of Adelaide, Adelaide, S Aust. 5000 Trans. R. Sor. S, Aust. Vol. 96, Part 4, 30 November 1972 198 GEORGES DUBOIS and L- MADELINE ANGEI Hahitat: upper intestine. Deseription® These specinens, taken from. the type-host, measure 2,8-3.0 mm; eggs 92-99 by 61-68 em. Parastrigea, repens (Chase, 1924), Dubois, 1968; 68, fig, 51, Chase, 1Y21; 500, fig. 1 and pl. 26 figs. 1-5. FIGS, |, 2 flast and wviging Circuy approximans Peile, from Tailem Bend, 8. Anst.. 1Qav. 1950 (18 adult and 1 immature specimen) ind Dee. 1938 (10 young specimens), Neahitat; duodenum. Chase (1921) found three specimens of this strigetd in the intestine of Neropheyx novere- hollanediae (Lathany) Srom Terrigal, N.S.W. The holotype, which is registered in the Aus- tralian Museum (W544), was reexamined by Mr, J, C. Pearson (see Dubois 1968, p68, foot- note 1). On the basis of this examination, the: species owas removed from the yenus A phuryngostrigea Ciurea tu Perastrigea Sziduat. Nescription: The smallest specimens: with few eges in the uteri measure 1,6-2,2 mm. Fully mature worms are 5-6 mm long. Suckers weakly developed: oral sucker marginal, 60- 115 pm in average diameter, ventral sucker 92- 165 yan, near oral sucker, Average ratio of the oral to the ventral sucker nearly 2? =: 3. Length of forebody Sram 9-19 faverage 14) times that of oral sucker, Protealytic gland elongated, oval or fusiform, 190-220 by sO- 110 ,»m, composed of closely averegated lobules and lying between the two cencentra- tions of the vitelline follicles, Ovary kidncy-shaped and testes multi-lobed, occupying second hall of the hindbody. Vitel- Jariy of forcbody extending dorsally up to ventral sucker, concentruted in Jateral semi- cordifarm expansions of dorsal lip of tribocy- tic organ. (No follicles tn ventral lip, which is as long as forehody), Scattercd follicles in wall of segment, extending further forward dorsally than ventro-laterally; In hindbody, virclline follicles concentrated in front of ovary, absent dorsally Over the gonads, and extending ventro- laterally to the bursa copulatrix, Bjaculatury duct joming with alerus at entrance to genital conc. Eggs numerous. 90-105 hy 60-68 am. Reluionships: Parastvigea repens, P. inrer- inedta! Tubangui, 1932, and P. flexitis (Dubois, 1954) are closely related apharyngeal strigcids; in the uhsence of pharynx they are distinguish- able from all known members of the genus, Their normal hosis are birds of prey {Falconi- formes). P. iniermecdia (feam the Philippines) differs from P. repens in that the small suckers are subequal, and in the size of the eggs (100 112 by 71-79 nm), PF. flexilis is distinguished from che Australian species by having fewer ceys, and by «a geographical distribution festeicted Lo the holaretic zone of Europe and Asia Parastrigea sp, Host and origin: Uhreskiornix moluece (Cuvier), from Queensland, 26vi.191L (4 contracted speciinens. collected hy A. Breinl), Rauhite: unknown. Desenipiion! Body fength |.4-2.4 mm-. Fore- body 0.88-1.08 by 0.94-1.15 mm (dorso-ven- tral diam.), with two well-developed lateral expansions. Hindhody 0.75—1.40 by 0.81--1,05 mm. Bursa copulatrix small. ‘These worms exhibit a striking similarity to BR. robusta Szidat. A specific diagnosis is reserved until better specimens are obtained. Strigea baylisi Dubois. 1937; 1968: 82. figs. 6-61, Hoss and orivin: Theeskiornis meolucea (Cuvier), from Tailem Bend, 8, Aust, 28.41.1942. (one specimen), Platelea flavipes Gould, from Toilem Bend, 24.17,1943 (23 specimens) and 10.x11.1947 (6 specimens). Habirar. intestine: Relationships: Strigea baylixi, which appears to be @ parasite of Platalcidac, is distinguish- uble from S. promiscua Nicoll by its smotler size, the smallness and the weakness of the phurynx (30-60 by 37-47 um), the extension of the vitellaria to nearly as far as the posterior extremity, and the absence of follicles from the ventral wall of the anterior scgment, Strigea glandulosa Dubois, 1947: 244. fic 9; 1968) JOL, figs. 82-84. Dubois & Pearson, 1965; 82, figs. 4—5. Strigea falconixy Dubois, 1937: 247, fig. 10 (not Szidat, 1928). lavix and origin: Circus approxinums Peale, from Tailem Bend, 8. Aust., 15.xii 1938 (2 young specimens), MValiastur spherurus (Vicillot), from Tailem Bed, Dec. 1958. (1 One of us (G.D-}, having examined three syalypes of P. intermedia, has not found any trace of 7 pharynx, STRIGEATA OF BIRDS AND MAMMALS 199 specimen} and 18.¥i,1941 42 macerated specimens). Faloo submiger Gray from Men- ingic. S. Aust. 6.v.1945 (4 specimens y- Habitat: intestine. Deseripliian; Body ip to 2.8 mm in length (when extended). The smallest specimen, with eggs in the uterus. sacasures only 1.4 mm, Oral sucker terminal. often prominent, 120— 140 by 90-125 ,m, pharynx rounded. very muscular, 95-105 hy 90-105 jm; ventral sucker 160-190 by 150-175 ,m: proteolytic gland well developed, oval, strongly lobulated, 145-210 by 170-250 yim, lying wt base of fore- body, Ovary reniform. Mehtis’ gland intertesticular. well developed. Fggs 89-102 by 57-69 ym, veTy NUMETOUS i Muture specimens, in which the uterus may he distended into sinuotis or lortugus curves and even into Joops, Relavonships: Sirigea plaidulose differs Erom 5. foleenis Szidat in its small size and in the greal development of the proigalytic pland relative lo the length: of the body, Strigea nicolli (Dubois, 1937), Dubois, 1968: 114, figs, 101-102. Sirigea sutiont Dubois, 1937: 237, figs. 5— 7. Hass and origin: Gyninorhina ihicen fLatham) from Canberra, A.C.T., April 1969 (4 specimens) and 27.vii,1960 £1 specimen; collected by R. Mykytowyez), Flebirar: duodenum and intestine. Descriptian: Length 1.40-1.62 mm. Oral sucker 121-12 by 195-177 pm: pharynx §5- LOS by 75-90 ym; ventral sucker 190-230 by 215-230 pm, Eggs 108-115 by 65-72 jm. Relarionships: Strigea nicolli resembles S. bay- fis! Dubois in general anatomy, but differs in the sizes of the suckers and the pharynx, which are definitely Jarger, and in the minor develop- ment of the atrial ring-shaped musculature. Srigea promiscua Nicoll, 1914: 347, Dubois, 1968; 119. figs. 107-1083. Dubois & Pear- son, L967: 1R6, fHlaxts and origins Ninox jevaextelandiac (Gmelin} (syn. N. boofeek) from Yatkuri. S. Aust. 24.viil. 1987 (1 specimen, deserihed below). N, strenaa (Gould) fram Bidsvold, Qid,, 1vil9L9 (22 specimens, very con- tracted; collected by M. J. Bancroft). Habiraty intestine, On June 9, 1965, Dr. I. C, Pearson collected the species from the smal} intestine of Ninox Hovaeseelandiae jn Brisbane, Qld, The species has now been found four times, always from the same host genus. Description; Length 2 mm, Body very con- tractilo, nearly as wide as long when. strongly contracted (as in 1,¥7,1919 material), Oral sucker 160 by 185 ym, pharynx 115 by 105 ym, ventral sucker 230 by 210 ym; proteoly- tic gland 190 by 215 ,m, multilobed. Ovary renitorm, 170 by 260 «mm. Testes roughly lobed. the anterior measuring 285 by 390 ym, the posterior 320 by 400 xm. Vitel- lara extending from cephalic margin to level of equator of genital conc. The latter is robust, well differentiated. larger than the ovary, 260 zm in diam. when retracted, Genital atrium spacious, L80 to 240 jum in depth, with ring- shaped musculature well developed. Eggs (04— 115 hy 75-80 pm, 6 in number. AS at present known, it seems that Sinigea premiscud 1 restricted 4o the Singiformes. especially to the arientul type Nivea Apatemon (Apatemon) vitelliresiduus n.sp- FIG, 3 Host and origin: Biziura lobata (Shaw) trom Tavem Bend. S. Aust., 10.x11.1937 and 9.x. 1940 (35 specimens); Sandgate, Qld. 22.18.1918 (1 specimen); Pumong and Caloot, R, Murray, S. Aust, 20, vi l958 (10 specimens, obtained from two hosts) (type material). Habitare intestine. Holotype: length 2.6 mm, SAM, E927 [with 3 paratypes. E928, on same sliile). Descriptien: Body cambered dorsally, 2.2=2.4 mm long, Forebody ovoid or cup-shaped, 0.71-. 0.90 mm in Jength by 0,51-0.64 mm in dorso- ventral diam., delimited by constriction from hindbody. Hindbody twice as long, banana- shaped, with small bursal region slightly de- lineated, 1.48-1.70 mm long by 0.50-0.56 mm in diam, at the Jevel of the tesies, Ratio of hindbady to forebody ringing from 1.9-2.1, Oral sucker 115-160 by 110-127 «am, terminal in position: pharynx smaller, 85-93 by 70-75 pmMe ventral) sucker postequatorial, relatively large, 190-210 by 165-180 um, Proteolytic gland small, rounded and lobed, situated dor- sally near base ef forcbody, 63-80 by 60-70 pM, Ovary 140-150 wm Jong, 180-190 am thick. located at anterior ihird (27-36/100, average 33/100) of hindbody, Testes latge, upproxi- mately equal in size (first, 260-330 by 240- 300 wm, second 300-370 by 260-300 gm), ovoid, roughly lobed, obliquely orientated with GEORGES DUBOIS and L. MADELINE ANGEL a4 =o ae STRIGEATA OF BIRDS AND MAMMALS lobes directed forward. Vitellaria are confined ventrally in hindbody,, especially well developed in front of gonads, extending to near posterior extremity of Worm, but not masking bursa copulalrix laterally; erratic follicles extend more or Jess far into ventral wall of forebody, but do pot go beyond acetabular level, Vitelline reservoir lies in intertesticular space. Mebhlis” Slabd is situated dorsally and a litle anteriorly. Genital cone of mediam size, 190-200 by 150— 180 nm, not well differentiated, Vesicula sem- inalis S-shaped and voluminous, lying postero- dorsal to second testis. Sinuoux ejaculatory duct opens into terminal part of uterus at entrance lo conc. to form a rectilinear, not pleated, hermaphroditic canal. Eggs 87-95 by 58-67 pm, average 92 by 63 pi. Relationships: This new species closely resembles Aparemon (.4.) fuliguiae Yamaguti, 1933, in geeral morphology, but differs essen- tially in the presence of erratic vitelline follicles in the ventral wall of the anterior segment, This femnant appears to constitute an archaic character. In 4, fitliguiae. the eggs are larger (100-220 by 60-66 um). Apatemon (Australapatemon) — intermedius {$. J. Johnston, 1904), S. J. Johnston, 1904: 109, pl. v, figs. 7-10. Dubois, 1968: 169. figs. 162-164. Dubois & Pearson, 1965: 85, figs, 6-7. Johnston & Angel, 195i; 66, figs, 1-28, Hosis and origin: Oxyure australis Gould, from Tailem Bend, S. Aust., 15.1.1941 and 28.1941 (11 specimens), Accipiter fas- ciatus Vigors & Horsfield from Mallala, 8. Aust., March [965 (20 small ovigerous specimens). (fohnston & Angel (1951) found this species in 5 of 11 Cygnus atrartr (Latham) from Tailem Bend). Habitat: unknown. Description: Body Jength 1.5-2.8 mm (speci- mens from Oxyurer wustraliz), This species is characterised by the structure and size of the genital cone; this organ measures 240-320 by 160-200 ,m when retracted, thus being ahaut one fifth the total length of worm, A wide strongly folded hermaphrodite duct passes through the cone. Laurer's canal descends from 20) oviduct where the latter leaves the ovary, and Teaches the dorsal surface on a level with an- terior leslis, Eges 94-99 by 60-68 am. The specimens from