VOL. 99, PART 1 28 FEBRUARY, 1975
TRANSACTIONS OF THE ROYAL SOCIETY OF SOUTH AUSTRALIA INCORPORATED
CONTENTS
Kott, Patricia The Ascidians of South Australia III. Northern Sector of the Great Australian Bight and Additional Records 1
Bnonaiuto, M. F. Notes on the Genus Pseudomalaxis Fischer (Mollusca: Gastro- poda) and its Fossil Species in Australia - - - 21
Gradwell, N. The Clinging Mechanism of Pseudophryne bibroni (Anura: Leptodactylidae) to an Alga on Glass - - - - 31
Schmitt, L. H. Genetic Evidence for the Existence of Two Separate Populations of Rattus fuscipes greyii on Pearson Island, South Australia - 35
Mawson, Patricia M. Two New Species of the Genus Cloacina (Nernatoda: Strongylida) from the Tammar, Macropus eugenii 39
Bullock, D. A. The General Water Circulation of Spencer Gulf, South Australia, in the period February to May ------43
PUBLISHED AND SOLD AT THE SOCIETY'S ROOMS STATE LIBRARY BUILDING NORTH TERRACE, ADELAIDE, S.A. 5000 THE ASCIDIANS OF SOUTH AUSTRALIA III. NORTHERN SECTOR OF THE GREAT AUSTRALIAN BIGHT AND ADDITIONAL RECORDS
by Patricia Kott
Summary
KOTT, PATRICIA (1975).- The Ascidians of South Australia III. Northern Sector of the Great
Australian Bight and Additional Records. Trans. R. Soc. S. Aust. 99(1), 1-20, 28 February, 1975. An account is given of 58 species of the Ascidiacea from South Australia, of which 7 species are new, including two assigned to new genera in the sub families Euherdmaniinae and Botryllinae. Records of 22 species from the northern part of the Great Australian Bight are the first from that area and suggest that the ascidian fauna there has a considerable endemic component. Many of the species common in other parts of the Flindersian marine faunal Province have not yet been recorded from this location. 1HE ASCID1ANS OF SOUTH AUSTRALIA III* NORTHERN SECTOR OF THE GREAT AUSTRALIAN BIGHT AND ADDITIONAL RECORDS
by Patricia Kott*
Summary
TTI Northern Great KoiT, Patricia (1975). —The Ascidians of South Australia + Sector of the Australian Bight and Additional Records. Trans. R, Soc. S. Aust. 99(1;., 1-20, 2H Feb- ruary, 1975. An account is given of 58 species of the Ascidiacca from South Australia, of which 7 species arc new. including two assigned to new genera in the suh families Tuherdmaniinae and Bolryllinae. Records of 22 species from the northern part of the Great Australian Bight are the first from thai area and suggest that the ascidian fauna there has a considerable endemic com- ponent. Many of the species common in other parts of the Flindersian marine fatmal Province have not vet been recorded from this location.
Introduction All available collection data for the speci- This account of the ascidian fauna of South mens discussed arc given in the Appendix.
Australia is based on the following collections: Order ENTEROGONA ( I ) from Ihe northern part of Spencer Gulf Suborder APLOUSOBRANCHIA in connection with environmental (made Family CLAVEL1N1DAE studies in that area); (2) from the northern Subfamily ciavelininah part of the Great Australian Bight (made in connection with an experimental Prawn Trawl Podnclavelta cylindrica (Quoy & Gatmardj. Survey, Explorer); (3) from Investigator Strait Kott, 1972a: 5 (synonymy); 1972b: 167.
(collected by J, Watson) and from "West New Records: Tipara Reef (Spencer Gulf T: Island; (4) additional collections from Elliston on reel" NNVV Douglas Bank (upper Spencer Bay at the eastern end of the Great Australian Gulf). Bight (collected by S. Shepherd). 1 lie report is supplementary to previous papers on the South Subfamily HOLOZOINAE Australian ascidian fauna (Kott 1972a, 1972b). Distaplia anstraliensis Brcwin. 1953: 6L Kott. It includes records of 57 species, including 7 1957: 95. that are new to science. Two of these new New Records: upper Spencer Gull (Stn D5). species have been assigned to new genera in the sub-families Kuherdmaniinae and Botryllmae. Previous Records: Tas. (D'Entrecasteaux Four species are newly recorded from South Channel and southern Tasmania). Australia. Description: Colonies consist of a rounded head The occurrence of 6 new species in the on a short cylindrical stalk. There is a single northern part of the Great Australian Bight terminal common cloacal aperture and the suggests an unusually high endemic component zooids are arranged along cither side of the for the ascidian fauna of that area, and its zoo- cloacal canals that radiate from this aperture geography is discussed. and extend down the length of the head, There
Type material is deposited in. Australian are about 12 flue longitudinal thoracic muscles museums as indicated by the abbreviations AM Ten stigmata arc present in each of the four (Australian Museum), NMV (National Mu- rows and these are crossed by fine parastig- seum of Victoria), QM (Queensland Museum), mutic vessels. There are 8 rounded stomach and SAM (South Australian Museum). folds, The gonads extend, from the pole of the
* Queensland Museum. Gregory Tee., Fortitude Valley. Qld, Australia 4006. PATRICIA KOTT gut loop, /olo a short posterior abdominal stilTencd test. 'I here are 9-1 1 rows of 27-2S extension separated from the abdumen by a stigmata, each row crossed by a paradigmatic shoa neck. Seven to 8 elongate testis lobes are vessel. A pointed papilla is present in the nrrnnged in a circle with their long axes paral- middle uf each primary transverse and para- lel to tme another to form a barrel shaped mass. stigmaric vessel oo both sides of the body. The the VftfS deferens, extending from the distal end internal wall of the siomaeh is arranged in of the centre of this mass, passes around it longitudinal and transverse slandulaf ridges into the abdomen. There is also an ovary in rather than folds. the posterior abdomen. RiUerella hcrrimauia Koll; 1972a; II fsyno- ftfnmfa; The colonies arc identical wilb those nymy); 1972f>: 172; 1972c: 24 vesicles that extends postero-ventratly I almost the same diameter as the head. The (Fig. ). colony from Stn Z6 is the largest yet recorded, Remarks: This species has been taken from Elliston Bay measuring 17 cm of which the stalk is ouly 5 (Kott 1972b) in May 1971. and cm, The minute zooids are present in the sur- the present colonies were collected in the pre- face layer of test with long posterior abdominal vious February. Only the latter arc sexually stolons penetrating into the centre of the lobe. mature and contain larvae. It is not clear whether rhe I here is the usual brown pigment patch over colonics taken in May were newly the anterior cn<| of the cndostyle. settled forms, yet to reach reproductive matur- ily, or whether they were older colonics that Family POLYCITOK1DAK reproduced earlier in the year. "However, the Ptilycilor gigantctnn tHerdman). Kou, 1972a; 9 species appears to leproduee sexually at the (synonymy |:. 1972c; 244. vai\ of summer, Collections from I'ort Hacking, Mew Records: northern dear Australian N.S.W. (Kott 1972c) indicate that Iheie. although new lobes Bight; West T. | Amphitheatre RocfcK were being added to ttw colonics at the end of Augusl. the species dis- Family FOLYCLIN1DAK appeared during the summer and did not return Suhfamily uuhurdmanunai-. until autumn. Kecolonising- stock must there Eulierdnifcinia aiisimlfs Kott, 1957: 103; 1972b: lure exist olf Port Hacking, which reproduces 172. sexually at *he end of summer or early autumn. 1 i.e. a similar seasonal cycle to that New Rec/nds Elliston Bay (outside bar>-; occurring at Elliston Bay. Investigator SlraiL (Sin YS), Di'stripriim; The colonies are iormed of the MATRIDIUM n. gen. usual long club-shaped lobes joined basully. /.ooidv completely embedded with both 1'itch lobe is composed o\ a single zooid ripcrturcs opening separately to the exterior covered by its own separate sheath of sand- and withuut colonial systems. Internal lonei- ASCIDIANS OF SOUTH AUSTRALIA 1TI Fig. U Ritterelta hcrdmunia. Larva. Figs 2-3. Patridhim pulvinawrfu Fig. 2.—Portion of branchial sac showing internal longitudinal vessels. Fig. 3; —Thorax and abdomen of adult zooid. pigs 4_6. Aplitlium foliorum. Fig. 4.—Dorsal aspect of interior portion of thorax showing tripartite atrial lip. Fig. 5* — Stomach. Fig. 6.—Larva. Fig. 7. ApJidittm pronum. Thorax and abdomen of adult zooid. Fig. 8. Aplidium disihuum. .Portion of colony. Figs 9-10. Leptoclinides voh'Us, Fie. 9.—Colony. Fig. 10.—Thorax, tudinal vessels are present in the branchial sac. Remarks: Only two genera of the subfamily The stomach is folded. Gonads are present in Euherdmaniinae are known in which either the thrcad-lrke posterior abdomen, the testis longitudinal vessels or their vestiges are re- follicles arranged in a double row and the tained in the branchial sac, These genera are ovary present just anterior to the testes. The Tylobranchion Herdman, a monotypic endemic heart is a U-shaped tube at the distal end of the antarctic genus (see Kott 1969), and Protopo- posterior abdomen. lycUnum Millar, in which 3 species are known, PATRICIA KOTT viz. P, pedunculatum Millar. I960, Irojn Now both sides nl the zonid for its whole length. Zealand; P. sabutoxa (Millar, 1963), from Port I here arc about 25 rows of 16 short, oval, stig- Phillip, Victoria; and P. claviforme Kolt. 19i>>. mata; rather tall papillae are present on Ihe from Eden, rV.S.VV. 'I ylohrtwrliwn retains some liarisverse vessels and these support longitu- primitive characters in the presence of the dinal vessels running the whole length of the heart half way down the posterior abdomen, branchial sac. The longitudinal vessels are and a large ovary posterior lo the hunched crowded, being .separated from one anolher by testis follicles In ProropolycUmtm the stomach an interval equivalent, to Ihe width of about does iiol folds testis have longitudinal and the one anil a half stigmata. The oesophagus is follicles are bunched in a short posterior abdo- J'airly long and there is a voluminous stomach men as in Polyclinum spp. "I he present genus. about halfway down the abdomen with 25 con- bears the same relationship lb ApUdtum us spicuous longitudinal folds, The proximal pari Protopolyclbutm hears to Polyclinum. It differs ol the posterior abdomen does not contain front both Protopofyclbwm and Pofyclintim in gonads bin this region is often contracted. The ihe presence of stomach folds and id its long ovary b present just anterior lo the double tow thread-like posterior abdomen in which the of testis follicles that occupy the greater part ot testis follicles arc arranged in tows; it is these the posterior abdomen. 'I 'he heart is a wide U- characters that relate it to ApUdtum. Jt differs shaped tube in the distal tip of the posterior from Ritterelhi, also in the subfamily Eubcrd- abdomen. maniinnc, in the presence of ?hc longitudinal Pseiulutlixfurmi cereuna vessels, and the absence of parasUgmutic vessels Michaelsen. Kolt, 1972a: 12 isynonyroy); ly72b: 173. in Ihe branchial sac, Riueretlo is usually further distinguished by the presence of 5 primary rows New Record: Marxarel Brock Reef (Cane ofstigmala, although these arc often subdivided Jaffa). by paradigmatic vessels. The restricted num- Remarks: Specimens in the present collection ber of primary rows of stigmata suggests that measure up to 12 cm of which the pointed or Rittcrella may be more specialised than Pntri- roimdcd head represents half of Ihe total tliiwi, which demonstrates primitive affinities length. The zooids, opening all around the j r the presence of it large number or rows, of head, are small, the contracted thorax and stigmata a^ well as in the retention ot the inner abdomen together measuring only 2 mm (Koti longitudinal vessels, 1972a. 2 cm ste), Subfamily r»oj,YcuNJN.\i: PatrMiiim puWlnatuw n. sp I'oJvclinnm neptunium Hartmcycr Kofi. Type Uyrmum: northern Uteat f Australian I /72b; 175 (synonymy). v Bight t$t*WSk 133 10'Fh 42 ru deep. Ti& Record: Invexiiguior Strait (Stti VII, 5.v.ry73. P S\woruL Httto/vpe: SAM. Description: E ID35. The present colonies are small. with rounded heads only about 3 mm in dia- FIGS 2. 3 meter on thin, branching stalks. Each head Description: The hololype o«Uy is available II contains about 6 zoo ids surrounding a central is cushion, h a circular cm in diameter and 2 common c'oacal opening. The test h very deli- cm high, mure or less flat topped and with cate. There iv no sand inlernally but externally rounded borders. Ir appears to have been ses- there is a heavy encrustation. sile and attached by a small area jn the centre The zooids are minute. The atrial lip is of the basal surface although there could have typical of the genus (Koti 1963) and has A been a short stalk in this position. The test is longitudinal muscle hands It arises from above very soft and semi-iransparcnt, generally with- the upper dm of the opening and appears to out sand or other adherent foreign particles close down over the aperture which is produced except for a small sandy urea at one side ot the to point directly anteriorly. There arc 7 muscle basal surface. The zooids are thread like, the bands radiating from the hranchial aperlurc thoiax and abdomen together arc 1.5 mm long but these do not extend onto the posterior part and the posterior abdomen about A mm They of the thorax. There are 5-8 small oval bJis- open all around the upper surface and the pos- mata in each of the 7 rows, and papillae on ihe lerior abdomina ptojeei down into the centre transverse vessels coincide with the sligmata. of the colony. The apertures are both 6 lobed The gonads ufj! nol developed The stomach is Fine longitudinal muscle bonds extend along smooth. ASC1DIAN5 OF SOUTH AUSTRALIA III Remark*. Although Lhe number of muscle alternating with two median ampullae. Numer- lateral bands in the atrial Up, and the number of rows. ous ampullary vesicles rise from the of stigmata and the number in each row. arc ridges extending anteriorly along both sides or very much less lhan that usually reported for the cudostyle and to the post ventral aspect of this species. The arrangement of the branchial the larval body. papiHue is the same as that usually rcpoUcd Henmrkx: The species is distinguished from (Kotl 1972b}. and it is possible that The colo- Aplidiufn plidierum by the larger size of the nies arc juvenile. colony, the very distinct radiating double rows Of yno'uU which comprise the systems, and by A pit (Hum foliomm n. sp. the characteristic foliaccous muscular atria* hire Loatrion; northern Great Australian C lobes that are present on many of the zooida. U3':V>'U» 42 m deep Bight (3Z 24'S. ( ft The colony does resemble thai of A. aitsrra- Sytnond. 5.V.1973. lioloiype. SAM, h 1036. {ienxix which has similar systems and in which FIGS 4-b the branchial sac is the same. In A. utistw Dei(ripfiorv The colony is a circular cushion Uensis\ however, there is a lesser number of high, 6 cm in diameter anil 2 cm forming a low stomach folds ami they ate sometimes irregular sljohilv conceive baldly where the zooid dome, and oblique, while the zoo ids lack the distinc- hearing surface layer of the test on one side has tive ainal lobes of the present species. The around onto the hasal surface to form grown larvae of A plidferunu A, anxi rathnxis and the crescent shaped pocket invaginated towards a present species arc, however, identical. There the border of the colony where the surface are slight variations in we (e.g. larvae from layer of test has grown to over- zuoid bearing the holotypc of A, australicttitx arc 0.9 mm lap it. The test is soft, gelatinous and semi- long) and in the length ot the tail which about i< ctoacal transparent. There arc common extends* from half to the entire distance around of the apertures scattered over the surface the body. However, the relationship of the colony, about 1.5 to 2 cm apart. Canals radiate length of the tail to the larval body does not tijit from lined each side by lhe opening* on appear to be constant for any single species. r.onids. These radiating canals sub- rows of The characteristic atrial lobes of the present divide rimes zooid* lining them on many and form are similar to those described for A. the Zounds are at right each side crowd test datum Koit. 1972b, which however differs con- lhe surface. angles to upper siderably from the present specimen in colon} aperture is with The branchial terminal the form, opening surrounded by a circular sphincter. The atrial lip rising from the upper border of Aplidiurn flavolineaturn iSluiter) Kott. JV72h: the aperture is very variable and may be simple 176 (synonymy). or tripartite, while the lobes may be large and New Record- northern Great Australian foliate or small and pointed There is a band Bight. lobe. ot muscles down the centre of each atrial Description' The colony is mushroom shaped, There are 14 rows of about 15 siigmnta. The 4 cm in diameter across the flat upper surface stomach is large with 18 to 25 narrow longi- and 2 cm high. The flattened znoid-beanng tudinal folds. head narrows very suddenly Jo a snort stalk The zuoid* are long and thread-like, the pos- from the centre of the under surface, Sand is terior abdomen comprising the great part of present on the stalk and, to a lesser extent, on rheir length while the thorax and abdomen To- the upper surface. The test is clear and glassy gether are only 2 mm long. There arc one or hut soft. The zooids are crowded in the lest ot two embryos at very different stages develop- and it was not possible io distinguish the form ment in a brood pouch that is formed by an of the systems. Zonids open only onto the expansion of the distal end of the oviduct at upper surface. They are 6 mm long, of which the postero-dorsal of the thorax. Dense end the thorax is 2 mm and abdomen only 1 mm. lestis follicles are present in two rows m the 1 he posterior abdomina cross one another in posterior .abdomen. The ovary is present ante- the internal lest although the Iho/accs arc rior to the testis (in tbc usual position for this parallel ai the surface. The atrial lip is divided genus). into 3 very pointed lobes from the upper border / iflwaw The larvae are 0.7? mm long with a of the opening. There are 12 fine Jongiludinal long tail that completely encircles the body. muscles on the thorax. There aic 12 rosvs of There are the usual three anterior papillae 10 long rectangular stigmata. The stomach is PATRICIA KO'I I especially small with 17 tlistinct rnngitudinal There are 12 rows of about Hi stigmata. Each folds, there are 2 rows of testis follicles in the row is crossed by a paradigmatic vessel. The posterior abdomen. stomach is small with 5 folds. Larvae; Up to 5 developing cmbryus sit pre- Larvae: The atrial cavity is occupied by a single sent in (he atrial cavity, They arc 0.75 mm birgc embryo. I mm long. It has the usual 3 long, have 3 median papillae alternating with median, stalked, papillae and numerous ampul- Single median ampullae and corresponding lary vehicles ate developed from the attterioi lateral ampullae develop from the lateral ridge. part of ihe body. The median ampullae arc narrow and in some Remarks: The specimen is identical with that cases appear to be bifurcated. 'I here are also previously described for the species, ihe larva clusters of ampullary vesicles both above the fe the same as that of W. pantiles htiwi (see KoU endo.stvlc and ventral to the body of the larva. 1963). The shape and consistency of the 'I he tail winds about three quarters of the way colony differ, however, and tremble A. pw- around the body of (he larva. tmems (Hcrdinan); Kou IM63, (n the latter Remarks: The colony and zooids arc similar of species, however, the zooids arc larger* the form to tho.se described previously for /T flavo- branchial sac larger, there are more thoracic IffUititWn with the exception of the stomach in muscle bands and the paradigmatic vessels are which there only nre 17 folds. Ihe size and not present. shape of the stomach and the course of the longitudinal folds are similar to ihnsc described Aplidiuin proiium n. sp. previously for (his species. Type Location: Investigator Strait (.Sin XI). ( Aplidium tonifenim Korr, 1963; 102. J in deep, Watson, Jan 197 J. Hoiotvpe: NMV. H 2s7_ New Records. Ellision Reef. Pn-vlatL\ Records: N.S.VV, (near Twofold Bay, 10-70 FIG. 7 deep: Montague North, I m 3 m deepj —Koit Description; The colony consists of small, Hal- 1963. topped lobes united basally. The lest ts-*ofc and Liescripiion; is Sessile* rounded lobes about 4 cm there very little sand internally. There ft ;i ii greatest diameter, the lest is sandy inter- single common cloacal aperture in the centre nally, but the external layer of tesi is tree pf of each fobe. The zooids arc more than 1 cm sand and is smooth and gelatinous. Zooids are long and thread-like. The thorax and abdomen long and narrow and open all around the head. me of equal length, and about one third of the There is a small pointed atrial lip from the total length of the zooid, The atrial lip is bifid body wall anterior to a muscular siphon Irum is or trifid terminally and extends from the upper present aboui one third of the distance down border Of Ihe atrial opening which is on a short siphon, Ihe dorsal surface. The thotax is long and There are 1 I rows ot 12 .stigmata and narrow with about 15 rows of 10 stigmata. 8 very weak stomach folds. There arc 20 longi- There arc 5 stomach folds. tudinal thoracic muscles. Remarks; The specimen agrees with those pre- Remarks: The colonics resemble those of A> viously described. The clear external layer of novaezehmdiue lire win and A. cotfrelli (Brewiti) test without sand hm] the form of the colony from New Zealand, although the Hal-topped -arc apparently characteristic of the species. lobes Biewin (1952, 1957 respectively) des- cribed for both these species have several sys- Aplidium amorphatum Kott, 1963; tfll. tems, only .1 stomach folds, and the atrial New Record; Klliston Buy. *rt viatts P Record: siphon is not B produced. The atrial siphon of A. Vic. <38"5l'S, U6 55'E1—Koit 1963. hiaritinttHH (Brewiti, 1958a) is. in fact, pro- D?sen prion; The colony is soft, sessile and duced in the same way as in the present dome shaped. The test h scmi-trarispajeiu. species, and ihe stomach has the same ill- Without sand Zooids open all around the upper derined folds. However, the longitudinal tho- suiface and no systems arc evident. The zooids racic muscles are more plentiful in Ihe present are very small and iirc-gulurly oriented in the specimens, there arc fewer stigmata in each test so that they cross one another. There are rnw, the posterior abdomen is longer and mure 10 longitudinal thoracic muscles. The atrial thread-like, and there is only a single system in jperturc is on a short siphon. The upper rim each lobe. Further. In Brcwin's species the of the apcttUie i*. produced into a pointed Up. Jobcs arc separate and do not appear tO be con- ASCTDIANS 0¥ SOUTH AUSTRAUA III 7 cloacal jvvstem consists of narrow liituous In their ba*ui hall as in the present common level, radiating from species. canals at oesophageal common cloacal apertures and lined on either \ptiilium dtgitatum n. sp. side by zooids. There are no spicules. The Type Locatfon: northern Great Australian zooids arc of moderate -size, with 4 rows of l C Hight ('32 74S, 133 3(VE), 49 m deep, stigmata and a long hifiiTcaicd atrial tongue s v. 1 973. /'. Sytnond. Holotype: SAM, E The oesophageal neck is long. Gonads are. not 1030. Potorypes: QM, G 7508; AM. Y mature. (982. The colony from Investigator Strait (that is tig. 8 doubtfully assigned to this species) consists nf long, branched, Description; The colonies are 2 large flattened lobes rising from a fleshy long, terminally cylindrical stalks 2- 3 cm cylindrical common basal stalk. The zooids are slightly to overlap the rounded and expanded embedded in the surface layer of test. The com- terminal stalk. The zooids open onto these mon cloacal canals extend between clumps of the sur- expansions. Sand is absent only from zooids beneath an especially thin layer of sur- zooids open on the face rest where the face lest. There arc vcrv extensive cloacal The expanded terminal portion of the lobe*. spaces between the central soft test that forms stalk is densely encrusted with sand. The test the central core of each lobe and the surface in the stalk, and is impreg- is firm, especially that is only occasionally joined with the central are minute, nated with sand throughout. Zootds core, hv solid test connect ives. Secondary canals hui long and thread-like, crowded in Lhe test extend between the zooids beneath a very thin another down lhe and extending parallel to one layer of surface test, The lest is colourless and stigmata. sialk. There arc J 5 rows of about 8 transpatent. and the zooids show through it as the There is a long, pointed, atrial lip from white dots. There are no spicules. Zooids have The stomach has upper Haider of the opening. a long oesophageal neck, and the usual 4 rows double of 12 longitudinal folds. There is a row of stigmata. Gonads are not mature in this testis follicles in the long posterior abdomen, colony. there is a large common cloacal opening in the assigned both P aspi- CCIlire of each lobe. Remarks Specimens to cnkvnm Tokioka, 1949 and /'. tnugmfarvum iistn/irks: 'the colonics resemble those of the Millar. 1962, are soft and rounded, stalked or Antarctic- Species Apluiinm rccurnhfttx; Kott. sessile, from 3 to 7 mm thick> with variable? 19n9. nut are distinguished by the large num- spicule distribution to an aspicufar condition. ber of di.%tinct longitudinal folds in the sto- The atrial lip is long, and often spread or bifur mach The zooids are especially delicate and caLe at its tip. There are also, til both, a large narrow. number of testis follicles (8-12) and large lar- ApUdfuiu colelloidc* (Merdrnan;. Kott, 1972b: vae (over 1 mm long) with up to 15 pairs of 176 (synonymy). lateral, finger-like ampullae and precocious AVh' Rcvoui northern Great Australian buds, Zooids of both species are also charac- Bight. terised by a long oesophageal neck. In Austra- lian specimens the ventral surfaces are em- Family DIDEMNIDAE bedded in the common test and they are Poh.sviicraton aspieubUum Tokioka, 1949: 2. arranged along both sides of common cloacal Kott 1962: 301. canats that demarcate rounded zooid-free swell- fvtvyynvrvtan ma^n'tlnrvum Millar, 1961: 13 ings of the surface of the colony. In neither (nomen nudum): 1962: 165. Kott, 1972b; species has the common cloaca been described 17*, as posterior-abdominal. The shape of the pre- Reronf.x: northern Great Australian Xt'w sent colony I torn InvcstigatCu Strait is identical Bight; ?!nvcsligutor Strait. Previous Recants: with others from this area that arc assigned to (Rnitncsi I l*i Peion) W. Alls'. , —Kou P. atpiculatitm (%P* nia^nilarvitm. Kott 1962. S. Aust. < Investigator Strait) —Kou 1972b). The posterior abdominal cavities in 1972b. Qld i.Mncfcay)— Kott 1902. S. Africa this colony, however, do nut occur in those pic- — MiMat 1962. Mozambique—Millar 1961. viously described. Positive identification is nut Japan—Tokioka 1949, possible Owing to the lack of mature gonads, Description. The colony from the Great Aus- and the relationship of the extensive cloacal tralian Bight is a soft jelly-like cushion. The system wilh the simple canals that have been 1 8 PAT RIC1A KOTT previously described is not known. Ihe cloacal types, together with the spherical shape, suggest system is the same as that of Dtdetunum fam- that these colonies may be free living, although ht/utti and Pofysymnttort cftondrWa but both the foreign particles that are embedded in the these species have only a single aperture and basal region suggests that this patt of the arc not known without spicules, colony was fixed to the substrate, hrokc tree and was overgrown by the surface test. The I.efKoclintdes voh'tts n. $p, constriction where the broadening stalk joins Type Locution: northern Great Australian the head in the holotype also supports the sug- Bight (32°24'S, 133 30' fc), 42 deep. P. m gestion that the spherical head may break Symond. Holotype; SAM. E 1034. PiVa- away. Certainly the configuration of the sur- types; SAM. F. 1033; QM, G 751 face, with the projecting swellings of solid gela- figs y. 10 tinous test, while the zooids, their openings, and the common cloacal aperture are depressed into Description: The specimen designated as the the surface of the test, would all accommodate holotype is a flattened sphere 5.0 em in dia- U free living habit in which the colony is able meter and 3 cm thick, with a thick but very to roll over the sea floor as in some coral spe- short stalk, constricted to 2 em in diameter cies (see Glynn 1974, Pichon J 974). where it joins the body. The paratypes arc The limited nature of the entirely spherical, about 3 cm in diameter, ami common cloacal system is unusual this genus ;irc without stalks although the base ol the m where extensive posterior abdominal spaces colony is identified by the absence of zooids are usually deve- loped. It differs front species in other $n Jengih, The atrial siphon is very short, rising rinsed with spicules. Spicules are dense at the Irom the mid dorsum, opposite the space zooid level but are absent basally. below ah- between the second and third rows of stigmata, dorainat level. Common cloacal cnnnls weie It is surrounded by a circular sphincter muscle not found and the thoraces oi the zooids and is posteriorly or laterally diieeicd. There appeared to be partially dislintegroted although arc 5j coils of the vas deferens and tip to 10 the abdumina were in good condition with 5| testis follicles. Large ova are presenl in the test coils of the vas delcrons around 5 ro *> testis at abdominal level but none of these appear to follicles. Small vegetative buds are present in be developing embryos. the oesophageal legion. fteuu/rLv: The spherical body, constricted stalk Reworks: The absence of common cloacal and single common cloaca! opening arc unusual canaW ;imi the condition of the zooids suggests in litis genus. The lack of stalk in the two para- that Ihe- colonies may present a ijinescenl winter ASUDJANS OF SOUTH AUSTRALIA 111 co.idiliun. The presence of vegetative buds gjk} the north, within the same biogcographical mature gonads suggests ihe onset rather than region, was arrived at after comparison of ihe end of ihis quiescent phase, Hastings type material (AM. GIJA49) vvrih other specimens Irom a wide range along the Lepttnliniiks rofiw (Sluiier). Kott, 1962: 286 southern and eastern Australian coast, includ- (synonymy); Hldredge, 1907: 221. ing the Great Barrier Reef and the Queensland Ltt>joefinidvs ttmts; Millar, 1963: 704. mainland. The type specimen of L. (liemetiewis Sew Records: Llliston Bay (outride bar. and has not been examined, although I okiolcu Millar 25 n\ deep). Previous Records: S. Aust. (1952), (1960). and kott (1962) have character that ) not been able to identify any i Port Noarl unga ) . Tas. (Maria I. . Vic. could distinguish the two species. Its synonymy I Port. S Shoieham )—Kott 1 962. N .S, W. Jackson—Knu 1962, Millar 1963. Qld (Bar- with L. ru/us is here maintained, The range of / rtiftts is, therefore, similar lo that of many jpra. Herun i.. Low Isles) —Hastings 1931; wide ranging species of this family in boih Kott J<>62. Indonesia U'alernosler 1., Ara- tropical waters Remarks; The synonymy of f . /furu Hastings, gether trout the basal half of the colony. The from Low Isles, with L. tufas originally des- common cloacal canals are deep, emending the cribed from Indonesia and the Arufura Sea to whole length of the zooids, bur they arc not . ID PATRICIA KOTT posterior abdominal. The spicules are 0.04- as the colony thickens and surface folds deve- 0.014 mm in diameter, They arc stellate with 12 lop. In the present specimens, both taken fuun conical rays in optical transverse section- The Sellick Beach, each species has spicules of dif- ionuls arc surrounded by black pigment par- ferent sizes although this siac difference W&4 ticles Ihat are often but nut always accumulated. not observed in specimens previously described. into the Usual pigment patch at the anterior end The number of spicule cays as previously of the endusryle. There is a distinct alrial reported, howevei. is greater for T. savignti siphon which I* laterally rather than posteriorly than for T. cerebrifortne. directed in these colonies. There is a distinct retractor muscle. The testis is not mature and Diploouma [ransUiculuin l.Hanmeyer) ihc vh< deferens was not distinguished. lwpio\'hnum I Leju'uclbumt ) trunsltKidum; Kou. 1962; 306 (synonymy | Remarks.' No further evidence is available from N,*w Record. Investigator Strait (Stn X1j. the examination of these specimens that could Previous Records: Indonesia- Sluiler 1909. clarify the relationship between this lndo North Western Australia—Hartmeyer 1919. Pacific subspecies and the Atlantic Ocean form W. Aust, (Oyster Harbour, Albany)—Kott T. vavignii subsp. a/roctnwm Van Name (sec Kott l%b). it should be noled that the Pacific Description: The colony K irregularly lobed, Ocean specimens (Eldrcdgc J 967) hnvc the investing weed or other ascidians.. Each lobe is 7—8 coils v\' ihc vas deferens that is associated flattened, about 2 wide, with (he Indo-Pacific form (Kott t966). cm 0.5 cm thick and up to 3 cm long. One colony completely enve- Tridiftemmim ccrebriforiue Hartmeyer. Kott. lopes a specimen of Pyura atisiralfs in which 1972c: 47 (synonymy) only the apertures are exposed. The surface K smooth, and the zooids Tcididemnutn stivt'tjnii; Tokioka, 1%7; 80; show through as while vt(r. j Remu/ks: The relationship of this species To '/ lamina is a double membra ne. ribbed on tl>e .? T. sovignii before its subsequent prolifcratiot ,, the flour nl the channel are encrusted with n ASUDJANS OF SOUTH AUSTRALIA III M layer of sand absent only from the siphons Amphirarpa Hiptyehu (Hartmeyer). Kott, 1972c which project upward from the middle pf the (synonymy). Upper surface. In this specimen, the right side New Record' northern Great Australian of lhe foody is narrower than the left and com- Bight prises the right side of the upper surface, while Ocutircuia auslralis Gray. Kot1, 1 972a: 29 the base, by which the animal is fixed, is the 1972b: 18-1- ventral surface and a large part of lhe left side Synonymy); Kolt, of the body. In specimens from G4, also on the New Record: ENiston Bay. floor of the channel, the siphons are especially Symplegnia viride Hcrdman. Kott, 1952: 252 long, both directed upwards, and the right side (synonymy); 1962; 129, Millar, 1966: 3ftS. of the body is similarly short, Plantc & Vasscur, 1966; 149. ToKiofca, The very long external siphons directed up- 1967? Itf? (synonymy). Viisseur, 1967: wards, and ttie sand encrustation, are unusual 111. in this species. Axcidia aclara is the only spe- Record: EDiston Reef cies of the genus in which a similar sand en- New crustation hardens the lest. The long, cylin- Subfamily kotryi unai. drical extension* of lhe test that, in A, aehtra* PARABOTRYLLUS n. gen. CWlte a canal or tube from the sessile *per. tutcs extending upwards from the animal, are Colonics are elongate branching stalks analogues rather than the homologues of the slightly expanded terminally. Otic to 3 circular long siphons in the. present forms. It is aJso of systems of zooids are present in each terminal interest that, in specimens from stations D> expansion, opening onto a more or less flat- and K4> where the siphons extend dorsally, tened surface of each free lobe. Each .system of their central position, from the middle of the zooids surrounds a central common cloaca! upper surface of the body, is achieved by rela- aperture. The terminal ampullae and conspi- tive narrowing (i.e. between the dorsal lamina cuous blood vascular system that arc present in and the endostyle) of the right side of the other genera of this subfamily are tthscnt The hotly; whereas in A. actaru it is the left side rim of the branchial aperture is smooth. The of the body that is narrower than the right. The in rial aperture hits a single anterior lip. There base of lhe present specimens is the ventral and arc only 2 internal longitudinal vessels in the two-thirds of the left surfaces of lhe body, branchial sac. Eggs are endogenous. while in A. achtra it is two-thirds of die light Remarks. The zooids are not conspicuous^ side, between the dorsal lamina and endostyle different from those of the genera Botrxllus and (Kott I972d). Botryijoides except jp the presence of only 2 The specimens appear to have adapted to internal longitudinal vessels in the branchial their free existence on a shifting sandy sea sac. The colony differs considerably, however. Hoar by these morphological variations in the both in its shape, and in the presence of only position an 12 PATRICIA KOI I FIGS 11-15 aperture, in the anterior third of the dorsal sur- Description; The colonics consist of narrow, face, has its upper lip produced into a single sandy lobes, 1-1.5 cm long, usually branched. pointed lip. The atrial aperture is very small I he superficial layer of test is encrusted with and is directed anteriorly so that the upper lip sand but is neither stiff nor brittle. Internally closes over it fas in Polyclhutfrt), There are sand Is absent and the test is very soft. There 10 rows of stigmata with about 10 stigmata in are circular systems of zooids opening onto the each row and two internal longitudinal vessels upper free end of the lobes surrounding the on each side of the body. The gut forms a central common cloaca! apertures that are single lighl loop on the left side of the bran- .slightly depressed into the surface. Generally chial sac. The stomach is pyrifonn wiih about there is only a single system in each terminal 8 distinct longitudinal folds and there is a short branch of the colony although occasionally curved caecum, of moderate length and ex- ihere are 1 or 2 smaller additional systems. The panded into a terminal bulb, from the pyloric blood vessels in the lest are short and relatively end of Lhe stomach. There is a connective ex- tew for rhis subfamily. They terminate in elon- tending from the pyloric region of the stomach gate rounded bulbs at the base of the zooids. to the intestine. There is a single, flat, testis The zooids arc about 2 mm long. The rim of follicle with lobed margins on each side or the branchial aperture is smooth and the atrial the branchial sac just anterior to the gut loop. Figs 11-15. ParabairyUm rwmorus. Fig. 11. —Portion of colony showing branching stulks. Fig. 12. Adult zooid. Fig. 13.—Hud (lateral aspect) showing connecting vessel. Fig. 14 —Bud (dorsal aspect) shoeing endogenous ova. Fig. 15. —Testis. Fit 16. Polycttrpa tincior. Aberrant individual with atrial siphon produced anlvTinrly Fig. 17. Pyura tendata. Section through the body wall, test and sandy coating. F;c. 18. Mtcmtosmite plaints. Gut and gonad on left side of the body. ASCIDIANS OF SOUTH AUSTRALIA Til 13 The vas deferens, arising from the middle of duced upwards so that it opened above the sea the mesial surface of ihe testis, is very priori. floor. In another specimen theic are long root- There arc three or four ova in (he botly wall like processes from the ventral border ot ihe anterior to the testis lobe. These are endo- body, which is otherwise typical of the specie* genous and project inlu the peribronchial (Fig 16), Jn the U-shaped specimen there is n cavity. Developing butls arc present in the test single row (17) of long polycarps around 1bc on either side of the posterior end of the adult ventral border of the body and only occasional zooids. These contain tour large ova on each pulycarps, representing a second row. scattered side of the body and a clump of large cells dorsal to these* dorsal to the ova. It is possible thai these may Polyesirpa perhinculata iHclIct't. Kott, 1972a; he precociously differentiating buds. The buds 35 (synonymy); 1972b; I8fi at this slaec of development are 0.2.5 -0.5 jnrn long- There is a blood vessel extending from the New Records: upper Spencer Gulf; northern posleriot end of each bud in the region of Ihe Great Australian Bight; Investigator SftraU. oesophagus. For Previous Records, Description, see Knit 1972a, 1972b. Remarks: The species is distinguished fmm Remarks: From the pari others jn the sublamilv by the large number 01 upper of Spencer oVvi and internal longitudinal branchial vessels. Gulf, arenaceous and naked specimens are lbs relatively limited blood vascular system in laken, sometimes growing side by side attached the test, and the form of the colony and the to the same shell or stone. There are a large limited development of colonial systems The number ol specimens and they are either internal tes! is also very soft in comparison stalked or sessile. Naked specimens vvilh a v\ith that ot other species in the subfamilv The leathery test were a bright yellow colour in life buds appear to undergo the major parr of their but pinkish in preservative. Living arenaceous* development in the test in connection with the specimens were a sandy colour with a reddish tinge colonial blood vascular .system and in the pre- There arc small, smooth, black indivi- sent colonies there wete no buds found directly duals in the preserved material and some that are larger associated with the parent zooids, The testis with rough leathery and rather thin lest. In of this species is reminiscent ol lhal in Sym- nature such specimens with a smooth piegma, while the multiplicity of ova resemble only fine sand encrusting it, and it appears that 1972c: 254: 1972d: 239 (synonymy). To- ihe animal had been half buried in the floor of kioka, 1967. Abbott & Johnson, 1972: 95. the channel and that the atrial siphon was pro- New Record: Pofl Rivet (St Vincent Gull). } N PAIRICIA. KOTT Remarks: The individuals arc small and the between folds. The branchial tentacles are of rounded swellings of the lest tire objured by varying sizes and twice pinnate. The dorsal epiphytic growth. However, the species is lamina is produced in a series, of pointed Jan- readily distinguished by Ihc short v$$$ elTeren- gucts. The gut forms a narrow Mraifiht loop ua, branched testis folliecs, undulating (some- and there is q mass of branched liver tubules iinies branched) ovarian tubes, lorn/ oeso- in the gastric area. Gonads, are divided into phagus, long recluni, deep secondary cut loop, separate paired polycarp-bkc sacs extending lomi voluminous stomach with internal folds along both sides ot the central common duct. and the small leaf-like endocarps that cover Remarks: This specimen agrees in most aspect* the body wall and the gut loop distal to the with those described from Investigator Snail, stomach. although the sandy coating is not as thick in The gut loop is reminiscent of thai of S. the present specimen. Nevertheless, the nature tvtmftcaw (Kott 1972d) although the rectum and orientation of the siphons are identical £S and oesophagus arc longer in the present spe- arc the internal organs* viz. the branchial sac cie^ and although the gonads resemble those the gut and the gonads, The atrial velum in the of S, partita (Stimpson), in the present spe- specimen from Investigator Strait was present cies the oesophagus is shorter and the vasa at the base of the atrial siphon rather than efferent ia -are shorter (Vasseur I9b7>. more terminally as in the present specimens, and this is possibly related to thickness of the Inoniiluuirpn «chcriduii (Hodman). Kott, sandy coating. The test beyond the velum is 1972a: 31 (synonymy); 1972c: 253, apparently produced to accommodate the thick- iVev Record: northern deal Austratum ness of the sand surrounding the animal, Bighl. Pyura pachydcrmattna (Herdmau) s. sp guV Family PYDRIDAK bosa (Heller). Kott, 1972b: 187. Cynthia gihhow (Heller). 1878: 27, Pyuratendata Kott, 1972b: 186. Pyura mbftow; Michaelssn & Wartmeypr, Reco?'d; south I. (off New of Goat Ceduna 1928: 410. Non P purhydrtrtwtitut vur. j|fA« FIG. 17 bosa: Kott, 1952; 265 (. pachvderrnaHita dras-vhii; Kott, l*J72b: IH7), Description: A single specimen only is avail- PyUrrt parhyilermatu Vara intvrmvditi; Kali, able. It is more or less a half circle in outline, 1952: 264 (synonymy) (part) Non P £lh* one cm in diameter. The external siphons ex- host* Mh'rtacdia Miehaelson, 1922.* 3<)l. ( and the atrial aperture is directed poslerodor- see Kutt 197 2b ( P pachydenmtlno villy. The external test is covered by a thick dtavchii) ; 1 9$2 ( P. pachydermatina inter- coating of sand held in place by hair-like exten- media ) sions from the test, There is a thin space Remark*: The present specimens have the between the sandy coating arid the surface of typical curved spines in the branchial siphon the test, traversed only by xhc base of the test, and the uiius is bordered with shallow rounded hairs. There is also n coating nf very fine sand I ones, on the surface of the test itself. The apertures P> lira spiiitfera (Quoy & CaMMrd), Kott, are lined with it very tough invagination of the 1972b: 186 (synonymy). test. The branchial siphon is especially mus- A/r Record: northern Great Austral tan cular and appears to be cversihlc. The bodv w Bight wall is also very muscular. The atrial siphon is muscular but not eversible and its aperture is Pyura aiistralis (Quoy &. Cat ward). Koit, protected by a well developed vehnn at the 1972b: 186 (synonymy). distal end of the siphon. Beyond this velum the New Rerord: reef, Douglas Bank (upper tvsi is produced into a cylindrical fibrous Spencer Gulf). extension for a short distance. There are seven hranchial folds on each side of the body with Pyura scorcsbicnsls Kott, 1972a: 36; 1972b: 14 strong internal longitudinal vessels very 1*7. closely placed on each fold. There are no in- New Record, upper Spencer Gulf rfStnv }'\, ternal longitudinal vessels in the interspace ASCIDIANS OF SOUTH AUSTRALIA ni 13 Pyura vtttaila (StimpsonJ. Kotl, 1972a: 37 Mieroeosmus planus n. sp. (synonymy); lV72d; 243. Type Lotulity: south of Goal L (olf Oduna), Net*. Record: upper Spencer Gulf (Stn Al). 31 m deep. I7-25.xii.1967, Hewlett. Hok). Remarks; The spines lining siphons are typically type; NMV. 4284. Paraivpes. SAM. £ H>32. long (0.1 and ncedlc-likc and ovcilappng. mm) QM, G 7510. "I he anal hordcr is smooth and two-lipped, FIG. 18 Pyura Irregularis (Herdman). Koit. 1972a: 38 (synonymy); 1972b: 187. Description; The individuals are circulat in out- New Record; upper Spencer Gulf (Stn B7). line and laterally flattened, with both apertures Remarks: The perttuhercular area has the usu- close together on the upper surface. The test is ally blister-like appearance tha? is characteitstic ihm and completely encrusted with sand, so of this species. The large dorsal tubercle, how- that the specimens resemble hardened discs of ever, is at (he top of that area rathei than, as sand. The sand is maintained around (he ani- has been previously described, at its base. The mal by hair-like extensions of the test, and lough leathery test and strong branchial &ac posteriorly these are longer, so that a flattened characteristic of this species are present. sandy keel is developed which interrupts the circular outline of the body, ft is apparent, Pyura slolnuifera (Heller) s. sp, praepijtialfe therefore, that these laterally flattened indi- Heller. Kott, 1952: 274 (synonymy); 1964: viduals are embedded upright in the sand 141, rather than lying on their side on the surface Pyura praeput talis; Millar. 1966. 372. of the sea floor. bfiip Record: Outer Harbour (St Vincent Gulf). Foi previous Records and Descrip- The apertures arc sessile. Longitudinal tion .see Kntt, 1952; Millar 1966. muscles from both siphons radiate over the crossing Remarks: This location apparently represents body, one another in the middle of each side as is usual in this genus. There arc the western extent of this species, which has ;i also circular continuous distribtilion from Queensland flown bands of muscles crossing the dor the eastern Australian coast. The specimens sal and ventral borders of the body. There is a conspicuous, elongate, dorsal ganglion here at the apparent end of its range axe be- smaller lhan have usually been recorded from tween the two apertures. The branchial sac has other locations, 7 folds on each side of the body with six internal No consta ni dilference has been detecttul longitudinal vessels on each fold. Parastigniatic vessels are present. There hciwecn ihe South African P. uolonifera s. sp. arc rectangular stigmata but no internal longi- itotorujera and ihe Australian P. xtotomfera s. sp. praepuriaUs and most characters demon- tudinal vessels between the folds. The dorsal strate a remarkable and overlapping ran.ee of lamina is a wide, plain-edged, membrane. The variation in the two populations. The rounded gut forms the usual long, narrow, curved loop with liver lamellae in the pyloric region. The fold of test enclosing ihe siphons, however* is never absent from Australian populations of branchial tenticles are twice pinnate although the secondary branches are very short and this species, although it also has been reported from South Africa; occasionally projections rounded The rectum extends anteriorly to the of test surround the apertures of South African base of the alnat aperture, forming a deep specimens but have never been observed in curve vvilh the gut loop. The anal aperture is Australian forms, (he different frequency with bi-labiatt!. The gonads are present in the sec- which these characters occur in each popula- ondary gut loop and consist of a long ovarian tube often forming close, curves, tion suggest that suhspectfic rank is appropriate. deep, Ihe ovarian tube has dense male follicles along its This matter is discussed more fully by KoH (in press). posterior border and on the lateral aspect nf the ovarian tube against the body wall. As the Halocyiithia hispida (Hcrdman). Kott« 1968: 7(i curves of the ovarian tube develop, the male (synonymy). 1972a: 41, 1972b; )S9, follicles appear to mingle with the ovaiy. rVw Record: upper Spencer Gulf (Stn G\. Owing to the curving of the ovarian tube, it Hcrdmonia mourns tSavigny). Kott, 19 72a: 4] often appears to project back into lobes (synonymy); 1972b: 1*9, bordered by male follicles. The gonad on the New Record; upper Spencer Gt)1l (Sin G); right side of the body is in a corresponding northern Great Australian BighL position to that on the left. I* PA UK I A KOI'I R*»iark\- The species is unusual in the semis circle. On The lateral aspect of Ihe ovary, it is ilt thai the gonads do nol cross mlo ihe apparent ihat the testis follicles embrace the primary gut loop. Id this respect only, it rc- end of the ovatian tube, hut on ihe mesial semhles Microco\tnus stotoniiera. the lorm ot aspect the U is completed to form S circle by the gonads in the present specimens is, how- the jerowth of the male fotlieles acmss Ihe sur- evcr, distinctive and the laterally Hnllened body face of the ovary. The male follicles are long, is also diagnostic of the species. and on their lateral aspe-cl lie along trie body wall dire:ted from the periphery of the circle VWcrocosmns squamigcf (MichacKcnl, (Cult, into Ihe eenlre. On the mesial surface of the 1972a: 43 (synonymy). gonads, the testis follicles can be seen to be New Retards upper Spencer Gulf Remarks: I lie specimens are small, more or with maturity. M. sabulosa is spherical with a less laterally flattened, spheres. The apertures sandy test ihat is hard and brittle, while M. are close together on the upper surface and a mollis, although encrusted with test, has fine ridge of slightly thickened lest extends Between hairs to which the sand adheres, the test itself them, 'there are line hairs on the lower part is more flucekJ, and the preserved specimens of the test which is completely encrusted with are laterally flattened. Ihe branchial aperture scjnd. Ihe rim of the aperture is lobed but is always protected by 6 pointed lobes from there arc no hollow test expansions surround- the sui rounding test a fiille dislance from the ing I hem as in M, xabidosa (sec below), The opening while the rim of the aperture itself is apertures are directed away from one another. produced into 6 smaller pomled lobes, thai an? there are 7 branchial folds on each side of the covered in the closed position by the rim of body, with up to 9 internal longitudinal vessels the larger lobes. 'Ihe atrial aperture is pro- nu each distributed over both sides of each tected by two flattened, wide tongues and theii fold, There are no internal longitudinal vessels border is separated into three rounded lobes between the folds, The testis follicles form a that arise from the test at the dorsal and complete* chele at the end of the ovary, wirh a ventral sides of the opening. The rirn of ihe ligumwl extending through the centre of this aperture itself is putduccd mto 4 small, ASC1D1ANS OF SOUTH AUSTRALIA Ul <• pointed, sandy lobes and these are covered by recorded here do nut therefore disagree with the larger lips in the closed position. All these previous information (Kott 1972b) Bftl Wlffc extensions from the test around the apertures ports Ihe existence ut* a marine fauna! Fioviuce are hollow and have prolongations of the body extending from Cockburn. Sound (or further wall extending into them They are characteris- to the north) un the -western Australian eoa.st tic of the specie* and are never present in M. to the vicinity of the eastern coast of South mollis. The gonad in the present species, >vhile Australia- superficially resembling that of M. tnollii,; has There is no evidence that would suggest lluil a very short vas deferens I hat opens at the the sample thai is available is not typical ttf proximal end of the ovary on its mesial surface. the fauna of the Great Australian Bight, This fauna, however, dom not, on the basis ol avail Uio&eO£r»pliy able data, appear to be Typical of the Flln- The 22 species rccoidcd from the northern dersian marine launal Province. Apart from part ol the Great Australian Bight (including the large endemic component, the species oc- Ccduna), a* far as 32"24'S, l-i3°30'E. can be curring there have n wide distribution around divided into the following groups: the Australian coast, especially along the east- ern seaboard. The ipectes that terminate then* 1. Possibly endemic to the Great Australian Bight. lange at the eastern end of the Flindeisian in collec- Patridium pnivbtatutn n. sp.; Aptidinm digi- Province have not been taken these tions from Ihe northern part or' I he Great Aus- uxtum n, sp.; Aplldium foiiontm n. sp.: Lep- although they have toclinides volvus n. sp,; Parabotryiltts ncmo- tralian Bight CKott 1972b), at easterly locations off rus n. sp.: Pyura tendata Kott; Mierocosmm been recorded more planus n. sp. South Austialia. Other species in this colled ion taken from 2. Southern temperate ( recorded aWo I rom other locations of! the South Australian coast South Africa), may be grouped according la ihe Soulh Aus- Aplhtium flavolineatum (Sluiler); Aplidtum tralian limits o\ their range in the following t'oIeUoides (Herdman). way: 3. Cirt tt tn -an sfrattan , J. Species ifmr do not extend eastwards into Pojycitor xixunteutn ( Herdman ; Lepto- ) the Muageati marine Provi/tce. clinides reriadutus (Sluitcr); Leptocliitides Podociavelta. cylindrica I Quoy & Gaimard); cuius (Sluitei ) ; Potysyncraton ospictdatum Pycrtoclavella dimittuta ( Kott ) ; Atapozoa Tnkiokn; Didemnum candidum Savigny: tnarvhit Brcwin; Diplosonm trausiuetdiuv Uidemnnm moselcyi ( Herdman); Atupht- Hai'tmcycr). Polyclinum neptunium (Hait ttirpa dipfycha (Hai'tmcycr): Polycarpa pe- meycr; Stafonhv carrwsa Millar; Afatguh dwtcuhita (Heller); Cncmidoctupa ciheridxii salntlusa Kott. (Herdman) (absent only from tropical Aus- 2. Species that do not extend westwards jnto tralia); tlerdmania momus (Savignyl the Flittdetsian murine Province. 4. Southern and eastern Australian species. Distaplia australicnsis Brcwin; Eidterdmania Ascidia aclara Kott; Sryeht pedata (Herd- australis Kott; Aplidium coni/eruoi Kott; man i Pyura pachydermatina ) (Herdman) ApUd'tutn amorphatum Kott; Ascidia thowp- nihhosa (Heller); Pynra spinileta I Quoy ami soni Kott; Polyandrocarpa iapidosa (Heid- Gaimard ; ) Microcosmus nicholhi Kott man): Pyuta irregularis (Herdman): Poly- (absent only from tropical Australia). carpa tinctor (Quoy & Gaimard); Polycarpa I he apparently endemic species comprise a papilhtta (Sluiter). considerable comporem (3l'# ) of the fauna 3. Species jor which the f1inderstart/ M andean in the northern part of ihc Great Australian boundary does not comprise a harrier, Bight. The ciTcum-Auslialian forms comprise Sycozoa cerehriformis {Quoy & Ciairnard): almost 5() f 7. of ihc species, while species with Sytozoa ptduHculuta (Quoy & Gaimard): a range to Port Jackson {Pyura p. #ihhoia* P. fiitterella herdmania Kott ; 1 rididemnutv Aptnifera) or Morvton Bav (Ascidia adam. stivignii f Herdman): 1 rididctnnutn cerebri- Styeitt pedidtt. Microcosmus nicholUi) also forme (Hartmeyer) ; Symple^ma virtde occur, the three latter records extend the Herdman; Pyura australis (Quoy & known range lo the west, although the first Gaimard); Microcosmus smiomjera KolL two species are already known to occur in 4. Ccdf faiota. southwestern Australia. The data thnt are PvtOa scoreshiensis Kotl. — IK PATRICIA KOTT References Aiibott, D. P & Johnson, J. V_ (1972).—The Koru, PAiKiCiA 196K).--A review of ihe genu-; Asvidians Styela harnharti. S. pircata, S. ctava, Hoiocv>ifhia Veridl. I«79, froc. I.ittu. Sec. and S. monterevensis in Californian Waters. A',\ |V, V3r ) 1 1, 7fV89. But! So, CrMjnhiitt Acad. 71 95-105. S$l {Z) y Ken, pAffticiA (JMA9).— AriUrclie Ascidiacea. A BRF.WIN, Bl.RVI I. U952).— Ascidians of New- monogiaphic uccnuni uf the known species Zealand, Pari VII, Ascidians from Olago based on upecinicns colleclcd under US coastal waters. Part Jl. Trans, R. Soc. N.Z, Governmcnl auspices 1947 to 1963. Antarcl. TlM^i). 452-458. ftef. Sfir. 13, i*XV, t-239. BnPWiN, ftp.RVf. T. Df the Mib-familv. Trans, H. Soc, N.Z, 8i( I ), atiJ Encounter Bav. IrvjU*. /?, 5'V. S. -las) 53-64. 9rt(l), 1-52 BreWuv, hFRVL I. ((95ft). Ampozoa nuirshl, a Kott. Patricia (1972b).—The ascidians of South compound uscidian from Wexlern Australia. Australia II. Eastern ncctor of ihe Great M'',.4. 31-32. J, R. Soc. 40(1), Australian Bieht and Investigator Stmir. Bur win, Bi-kyi \, (1957).— Ascidians of New 7Yew.v. R. Soc. S. Attsl. 96, 166 196. Zealand. Part X. Ascidians from North Kott, Patricia (1972c).—Notes on *omc sisci- 577- Auckland. Trans. R. Soc N.Z, 84(3), dians from Port Jackson. Botany Bay. and 580. Port Hacking, New South Wales. Proc. Lh*n. Bkiiwin, Bkkyl 1. (1958a).—Ascidiuns of New Soc, N.S.W. 97, 241-257. Zealand, Part XI. Ascidfans of the Stewart Koti. Patricia ( I972J) Some suhlatoral wsx-t- bland region. Trans. R, Soc. N.Z. 85(3), — 439-453. dians in Moreton Bay, Queensland. Mem. Qd Mas. 16(2). 233-260. BBEW1N. BtRYl. T. (1958b). -Aseidians ul New Zealand. Part XII, Ascidians of Ihc Hauraki Kott. Patricia (1972e).—-Paiina of the Gulf of Gulf Pt. III. Trans, R, SoC. N.Z. 85, 45^-45*. Carpentaria; 2. Asctdiacca (Chordala: Tuni- c-atal. W^fA. Notes t\\S. 2(3), 39-57- r.uwhoOK, L. O. (1967).—A taxonomic review of Indo-Pacifie didemnid ascidians and descrip- KQTT, Patricia (in piess).—The ascidian foutni tions of twenty-three central Pacific Specie*. of Wcslcrnport Bay. Victoria^ and a com- parison with that 61 Porl Phillip Bay. Mem. Mieronena I 1 61-261. Natn. Mas. Vict. Glynn. P. (1974).— Rolling sttsnes among ihe Scleractinia Prqc. Second lale*nationol Coral MroTuri.sTN, W. (1922). Ascidiae Ptycho- Reef Stkiposfom. Vol. II. 183-198. (Great Sranchiae und Diktyohranchiae von Neusee- Barrier Reef Commiliee: Brisbane). land und den Chatham-lnseln. (Papers from Dr. Tlr Morelensen's Pacific Fxnedilion Hart\u;yf.r. R. *IV19). -Ascidfevi. Results of Dr. 1914-lft. No, XI). Vtdfftk, McJd>, ttmsk K. Mjoberg's Swedish scientific expeditions to ttartirh, t'orfti, 73, 359-498. Australia 1910-13, K, wrnska Veim^k-Akatl- ifandl. 60(4), 1-150. Mu'.iiAr.LS.T.N. W, (1924). Ascidiac Kriko hranchiae von Nenseelond, den (_'hatham- hm\ Hastings, Anna B. { 1931 ).— "I nnicnta, Sclent* den Auckland-lnsclu. (Papers from Or. *l'h. Rep. Gt.. Barrier Reef Expcd. 4(3), 69-109. Mortemcns Pacific Expedition 1914-16. No, IIillCR, (I67K). Beitriige zur naheren C — XXII >. Viiicask. Mctfttr, tlaask. natuth. f'oren. Kenolnts der Tunicate i. S/nr. A Add. WLys. 77, 263-434. IVien. 77(1). 83-110, MltHAMSrN. W,, & HAHIMlVfcR. R. (I92K|.~ Kan, Patricia (1952)—The ascidians of Aus- Asdditic diktyobianchiae u.*d ptycho- iralia. I. Stolidobranchiatn and Phlebo- branchiae, Fauna SnJncsf-Audl. S, 251-4^0. hriuichiafii 1ltst. mar, Frcshh; Rv$. 3(3), A MiLiAR. K. H. f i960).—Asciduceu. Phcon'r* 206-333. Rep, 30. 1-60. Kott, Patricia (1957). —The ascidians of Aus- Millar. R. H. (1961).—^Avcidians from Mozam 1 nilf;i . H AplouNohranchiala I ahille; Clave- hi<|ue. Ann. M#x. Nat. Hht. t*3)4. 11-16 Imidac H'orbes and Hanly and Polyclinidac NfiLLAR. K. II. 11962) FuTther description* ' t. — ol Vorrill, Aust. J, mat, rfityrV1 Rc\, X( I 64-110. South African aockhan^ Ana, V, A jr. Mas. Kott, Patricia (1962)- -The ;e.adians of Au>- 46(7), 113-221. 1 ralin- |Tfi AploiiSiihriiDchiaia 1 ahille: Didcnv Miiiar, K H. (I9f»^),— Australian ascidians In hidac Oiard, Aast. J. mar. treshw. Res. Ihe British Museum (Naliual History I. Pew. 13(31, 265-334 Zttol Soc. Land. 141 1 4). 6X9-746- Koit, P.vnuuA. (1963).—The asvidiaus or Aob- Miliar, Is*. H. (1966).— Ascidlamt P<»rt Phillip iralia. IV, Aplousobranchiata LariiUe; Poly- Survey. Mem, natn. Mus, I'ict. 27, 357-375. clnudae Vern'll (continued ). Anst. J. ntar. Nott. J. T, (1892). —On the composire asciditins Freshw. Res. 14(1), 70-118. of the North Shore Reef. Trans. N.Z. fust. 24. Kott, PAiRtrtA (1964). — StoliJohianch and phle- 305-334. bobianeh ascidians of the Queensland coast. Prhon, M (1974). —Free living sclcractinian Pap, Dep. loot. Univ. Qd 2(7), 127-152. coral comnuinilies in the coial iccfs of Inlear Knrr, Patricia (1966) — Ascidians. of tiotlh Aus- ''Madagascar). Ptr.v. SvcotaJ JitTernniionai tralia. Pap. i)ep. loot. Univ. Qd 2fl5). Loral Herf Symposium. Vol It. 173-182 279-304., iGre.jt Barrier Reef Commilteo: Brisbane. 1 ) ASCIDIANS OF SOUTH AUSTRALIA III iy Pi.ante, R-. & VA&snm, f\ (1966),— Sur unc col- Tokioka, T. ( 1949) . — Contributions to the lection d'ascidies de la region de Tulcar (cote Japanese ascidian Fauna II. Notes on some sud-ouest de Madagascar). Ann. I'Univ. ascidtans collected chiefly along the coast Madagascar. Scric Sciences de la Nature et of Kii Peninsula. Pubis Seto mar. biol. Lab. Mathematiques, no. 4. (Tanarivc: Imprimcrie 1(2), 39-64. Nalionale. Tokiora, T. (1952 )— Aseidians collected by Qtjoy, J., & Gaimard, P, (1834).—Voyages de Messrs, Renzi Wada and Seizi Wade from decouvertcs de (Astrolabe 1826-29. MoL the pearl oyster beds in the Arafura Sea in 91-142. lusqucs. Zoologie 3, 559-626; 4 7 304-306. 1940. Pubis Seto mar. bwl. Lab. 2(2), Slliter. C. P. (1909).— Die Tunicaten der Siboga Tokioka, T. (1967), —Pacific Tunicata of the Expedition Pt. 2. Die Merosoraen Ascidten. United States National Museum. Bull. US* Siboga Exped. 56B, 1-112, nam. Mus. 251. 1-242. Sluiter. C. P. (1913). — Ascidien von den Am- VASSEim, P, ( 1967 ) . — Ascidies de Nouvelle- Inseln. Abh. sr.nckenh, nulurforsch. Ges. 35, Calcdonie. Edition de la Fondation Singer- 65-78. Polignae. 127-146, 2 pis. Appendix—Station List NORTHERN GREAT AUSTRALIAN BIGHT INVESTIGATOR STRAIT, January, 1971 (Coll. : (32°24'S. J33 '30'E). May 1973. Experimental J. E, Watson), Prown Trawl. Explorer (Coll. P. Symond). Station XI (depth 19 m); 42 m; Poly citor gigajiwum AplidiUm pronum n. sp. Patr'tdium puhinatum n. gen. n. *p Diplosoma translucidurt\ Aplidium cole I hides Pyura australis Aplidium foliorum n. sp. Station X3; Leptoclinides votvus n. sp. Pyura australis He rt Imania momus 49 m; Aplidium colelloides Station X7: A plidium flavoltneatum Hwdmaniti momus Aplidium digitutum n. sp. Stations X8, X9, XI 0: Lcptoclinidex reticulums Pxura scoresbiensis Polysynvraton nspiculuium Station Yl: Didemnum candidum ?Polyclinum neptunium Didemnum moseleyt Stolanicfi carnosn Amphlcarpa diptycha Polyundrocarpa lapidosa Polycarpa pedunculata Molgula moth's Styflu pedutti Station Y5: Cnemidocarpa etheridgii Atapozoa marsh't Pyitra pachydcrmntina gibbosa Euherdmania australi'i Pyura spinifera ?Pulysyncruton usphulatutn Tlerdmania mom Us Pyura australis Micrr/citsmus nichollyi Station 76: GOAT ISLAND, off Ccduna, Great Australian Atapozoa marshi Bight (Coll. P. Howlett). 32 m; Ascidia acjara UPPER SPENCER GULF, September. 1973. Pyura tendafa Transects and stations of S. A. Shepherd, Depart- Mkrocosmus planus m sp, ment of Fisheries, S. Aust. CLLISTON BAY. Feb. 1971 (Coll. S. A. Transects A-D: Shepherd). Polycarpa pedunculata (arenaceous and Outside bar: naked specimens, sometimes growing Fulierdmania australis side by side, attached to the same .shell Ritterelln berdmunia or stone, stalked or sessile). Aplidium ttmorphnlum Station A1 on Pinna, depth 0-1 m: Leptoclinides rttfus Pyura irregularis Oculinaria australis Pyura vittata Reef; Sytnplegrna viridc Station A5, depth 17 m: Aplidkurt caniJi'MM Ascidia tlwmpsont faces : Pyura pachydermatina Vertical (25 m) Polycarpa pedunculata Molgula sabulosa Station A7. depth 10m- WFST ISLAND; Amphitheatre RocV Polycarpa pedunculata (2 spec.) 7 m deep, 13.vii.1972; Station B4. depth 17 m: Pplycitor gtganteum Sycozoa pedunculata 17 m deep", 12,vh\1972: Sycozoa cerehriformis Station B7. depth 5 m: Polycarpa pedunculata (some naked, CAPE JAFFA; Margaret Brock Reef (3-4 m deep black; a few leathery) and in caves). 28.xi.1972. Polycarpa papillata (single specimen) Pseudodistoma cereum Pyura irregularis (one small aggregate) : 20 PATRICIA KOTT Station BH), depth 10 m, channel: Parabofrylins nemorus n. gen., ti. sp. Polycarpa iinctor Polycarpa peduuculata Molguhi mollis Pyura scoresbiensis Station C4, depth 12 m: fialocytithia htspidu Polyrarpa pedunculaia (naked and arena- Microcosmus squamiger ceous) Hcrdmania momus Station D3, depth 18 m: Reef 4 km NNW Douglas Bank: Ascidia thompsoul Podoclavclla cyl'mdrica Pyura aaMralis •Station D5, depth 15 m; Disfaplia austraiiensis SPENCER GULF. Station D9, depth 10 m; Tipara Reef, depth 11 m, 24jx.197J: Pyura scoreshiensis Pycnoclavella dimiuitta Station Eh depth 7 m: Pyura irregularis Polycarpa peduucalata Under stones; Microcosmus mchoflsi Hcrdmania momus Microcosmus stolon if era Depth 5 m, 2/V.1972: Station E3. depth 9 m: Podoclavclla cylirtdriva Ascidia tbompsoui Station E4. depth 5m: ST VINCENT GULF. Pyura irregularis Port River (near Ele.tricity Trust), depth 3 m T Station FL depth 19m; muddy bottom, 9.vi.l972: Polycarpa pedunculata (small stalked. .Slycia plicatii arenaceous) Outer Harbour: Station F3, depth 19 m: Pyui'n stolonifera Pyuta scoresbiepsis {without stalk) Microcosm us sqttam fgfiir Station F4. depth 16 m: Selliek Beach (S of Adelaide), Feb. 1972 (Coll. Ascidia thompsoni (with barnacles) K. Hammond) Station G, depth 9 in: Trididemnum savignii Ascidia thompsoni Indidemnum ccrebriUtrtnc Index to Genera and Species Amphicarpa diptycha II Parabofrylins ncmortis II Aplidtum amorphatun) 6 Patridium pulvinatum 4 - Aplidium colclloides 7 Podocbvella cylindrica I Aplidium eoniferum - 6 Polyandrocarpa lapidosa 19 AplkUum digitatum - 7 Polycarpa papillala - 13 Aplidium llavolincatum 5 Polycarpa pedunculate 13 Aplidium foliorum 5 Polycarpa tinctor 13 Aplidium pronum 6 Polycitor gigantetim - 2 Ascidia aclara - 11 Polyclinum neptunium 4 A.scidia thompsoni 10 Polysyncraton aspiculatum 7 Atapozoa marshi 7 Pscudodistoma ccrcum 4 Cnemidocarpa etheridgi 14 Pycnoclavella diminuta 20 Didemnum enndidum 9 Pyura australis 14 Didcmtium moseleyi - 9 Pyura irregularis 15 Diplosoma translucidum 10 Pyura pachydermatina 14 Distaplia ansfrnliensis 1 Pyura scoresbiensis T4 Euherdmania australis j Pyura spinifera 14 Halocynthta hispida - 15 Pyuta stolonifera 15 Herdmaiiia mom us IS Pyura tendata - 14 Leptoclinides reticulatus 8 Pyura vittata 15 Leptoclinides rufns - 9 Rittcrella herdmania Leptoclinides volvus - 8 Stolonica carnosa U Microcosmus nichollsi 16 Styela pedata I* Microcosmus planus - 15 Styela plicata 13 Microcosmus squamiger 16 Sycozoa cerebriformls 19 Microcosmus stolonifera 16 Syco/oa pedunculata Molgulu mollis 16 Symplegma virt'dc iT Molgula sabulosa 16 Trididemnum savignii Oculinaria australis - 11 Trididemnum cerebriforme to NOTES ON THE GENUS PSEUDOMALAXIS FISCHER (MOLLUSCA: GASTROPODA) AND ITS FOSSIL SPECIES IN AUSTRALIA byM. F. Buonaiuto* Summary BUONAIUTO, M. F. (1975). -Notes on the genus Pseudomalaxis Fischer (Mollusca: Gastropoda) and its fossil species in Australia. Trans. R. Soc. S. Aust 99(1), 21-29, 28 February, 1975. Two fossil species of Pseudomalaxis Fischer are discussed; P. asculpturatus Maxwell (Late Eocene) and P. praemeridionalis (Chapman) (Early Middle Miocene). The former is a new discovery in Australia and is one of a few forms common to both Australia and New Zealand; the latter is poorly known and is redescribed herein. The taxonomic position of Pseudomalaxis Fischer is reviewed and the genus is restored to the Architectonicidae. The possible synonymous or subgeneric relationship of Mangonuia Mestayer to Pseudomalaxis Fischer, of Calodisculus Rehder to Awarua Mestayer, and of Claraxis Iredale to Torinista Iredale, are considered. NOTES ON THE GENUS PSEUDOMALAXIS FISCHER (MOLLUSCA: GASTROPODA) AND ITS FOSSIL SPECIES IN AUSTRALIA by M. F. Buonaiuto* Summary Buonaiuto, M. F. (1975).—Notes on the genus Pseudomalaxis Fischer (Mollusca: Gastro- poda) and its fossil species in Australia. Trans. R. Soc. S. Aust. 99(1), 21-29, 28 Feb- ruary, 1975. Two fossil species of Pseudomalaxis Fischer are discussed: P. ascidpturatus Maxwell (Late Eocene) and P. praemeridionalis (Chapman) (Early Middle Miocene). The former is a new discovery in Australia and is one of a few forms common to both Australia and New Zealand; the latter is poorly known and is redescribed herein. The taxonomic position of Pseudomalaxis Fischer is reviewed and the genus is restored to the Architectonicidae. The possible synonymous or subgeneric relationship of Mangonuia Mestayer to Pseudo- malaxis Fischer, of Calodisculus Rehder to Awarua Mestayer, and of Claraxis Iredale to Torinista Tredale, are considered. Introduction nymy with Omalaxis, and separated the three The genus Pseudomalaxis Fischer is repre- remaining genera as distinct taxa. Also, he dis- sented in Australia only by three known spe- tinguished one of the two living forms pre- cies, two fossil and one living: the Late Eocene viously referred to P. zanclaea (Philippi) as a P. asculpturatus Maxwell, 1966; the Miocene different species, P. macandrewi Iredale, be- P. praemeridionalis (Chapman, 1912); and the cause of the latter's more evolute coiling, and living P. meridionalis (Hedley, 1903). Only the restricted Pseudomalaxis to this new living two fossil species will be discussed and des- form. cribed here. However, it is necessary to discuss Later, Monterosato (1913, pp. 362-3), from ( a ) the taxonomy of the genus Pseudomalaxis a different viewpoint, restored P. zanclaea as and the related genera Mangonuia and Awarua type-species of Pseudomalaxis and described Mestayer; (b) in which family the genus should the other living Mediterranean form, P. actoni be placed. Monterosato, previously mistaken by authors (a) Fischer (1885, p. 714) found in a living for P. zanclaea. Monterosato also instituted for form, then referred to the Neogene Bijrontia the above evolute form the subgenus Spirolaxis zanclaea (Philippi), a torinioid operculum and with P. (Spirolaxis) centrifuga Monterosato, therefore instituted for it Pseudomalaxis as a 1890 (syn P. macandrewi Iredale, 1911) as new subgenus of Torinia Gray. Sacco (1892, type-species (Monterosato 1913, fig. 2). Coss- p. 75) considered Pseudomalaxis Fischer a sub- mann (1915, pp. 122, 141) restricted Disco- genus of Discohelix Dunker. Dall (1892, p. helix Dunker to the Mesozoic forms and 331) recognized the intrageneric relationships Pseudomalaxis to the Cretaceous-Recent, re- between the Recent American Otnalaxis nobilis ferred both to Euomphalinac, and separated Verrill and P. zanclaea (Philippi), but insti- Omalaxis Deshayes in the Omalaxinae, a new tuted Discosolis for O. nobilis as a section or subfamily. possible subgenus of Discohelix, because of Rehder (1935. p. 129) recognized a very Verrill's description of the operculum of O. close affinity between Discosolis nobilis nobilis as trochoid. Iredale (1911, pp. 253-7) (Verrill) and Pseudomalaxis actoni Montero- tidied up the confusion existing in the use of sato and therefore placed Discosolis Dall in the names Discohelix Dunker. Omalaxis synonymy with Pseudomalaxis s. str. Rehder Deshayes, Bijrontia Deshayes, and Pseudo- (1973, pers. comm.) remarks: "Mangonuia malaxis Fischer; he placed Bijrontia in syno- Mestayer, 1930, is probably a junior synonym Department of Geology, University of Adelaide, S. Aust. 5000. 22 M. F. BUONAIUTO NOifcS ON eSUJtHhUAl 4X1S FISCHER IN AUSTRALIA 23 1 or [U leant a subgenus of PseitdoffteltixLs nobitts (Verxill) (Vcrrill IXR5, p. 421, pi 44. Fischer, 1885". Mcstaycr (1930, pp. l44-5> fig. (2). refers P. meridianalis (Hedley) to Manftpntda* Ircdalc (1936, p. 326) instituted a new In fact, the recent Mangontfin h**llon.*j Mefr family Mungonuiidae lor the Australasian layer, the type of this genus, displays the genera MctrtgonHiti, Awthna, Tnn'tmta and general pattern of spiral ornaments, the quad- Chwaxis, Apparently, Wenz (|939| considered rangular outer and the subcircular inner shape Mangonuiidac synonymous with the Architcc- o( body whorl characteristic of the Neogene- tonieidae. Because of a certain affinity heiween Reeent Pscudomalaxts a, str (l described below, the opercula of P. balesi Pilshry & McGinty and the .same type of pscudoplanispiral coiling and Parvitarbo zacallcx (Motycl;), Pilsbry & ILlf this group, McGinty (1945, pp. 9, 57. pi. 6. figs 2-2a. 5) Render further comments. "Calodhados placed Pseadomafaxis in the Cyclostrcmatidae. Render, 1935, is very close to Awanta Mes- Latei, Abhott (1954, p. 138) included Pxeudo- is its taycr, 1930, and probably synonym". rtialaxit in Vitrincllidae on the basis of <* vague Wenz f 1939. p, (567) considers Awanta a sub- and incorrect reference to a revision of the genus of Mangonuia. However, Calodhcuh** family by Pilsbry. Maxwell ( IV66, p. 4441 fol- and Awanta display the same kind of speciali- lowed Abbnii. zation in spiral ornaments, in particular a Rehdei (1974, pers. comm.) gave the marked development of heavily bended circum- opinion: '1 have examined young specimens of umhilical cords, Awanta ttmoena (Murdoch &. both Pscudo.vtu'aAtx and A rcliiieetoniea (Psl~ Suter) (Sitter 1913, p. 318; 1915, p] 15. jig, laxh) and I can find no essential difference in 21 ah) displays also a, torinioid operculum their protoeonehs: both show a heterostrophic- iMcstayei 1930. p. 146). anastrophic protoconch. 1 feel, therefore, that "Claraxix and Torinistti Tredale are very simi- Psfmdomalaxix should remain in the Archilec- lar, possibly synonymous, and might Ik sepa- tonicidac". There is support for this statement rated as a distinct genu?; whicb may be closer boih on the basis Of the torinioid conical oper- to Hchucus d'Orbiyny". Wenz (1939. pp. 666. culum and the protoconcli coiling of Pseudo- bhH) considers the former two as subcenem of waiaxis. Therefore the author* in agreemetii Mungonuia and the latter as a synonym of with Render, regards the restoration of Pxeudo*- Tnritt'ttt Gray. In this ca«e, Cbitttxis should fall malaxi* Fischer to the Architectonicidue to he into synonymy with lorttuxta because although justified. Iredalc (1A3& p. 327) published their initial dingrio;se-s on the same page, ihe former is des- Collections cribed after the latter. From the original des- P. cription and drawings of the type species, it is The specimens of astulpturutus Maxwell impossible to find substantial generic differ- here studied arc kept in the Pal aeon tologicai ences and Iredale did not specify any. Seciiun of the New Zcatarul Geological Survey the of of (h) The genus PseudotmdaXis Fischer is often (NZGSi, and in Department Mines placed in different families hut generally is South Australia (GSSA). The figured specimen referred to the Architcctonieidae. In introduc- of P. praemeridiomilis (Chapman) is kept in the Palaenntologicnl collection of the South ing the genus, Fischer referred it to the Sotor- — Australian Museum of Natural Hisiorv ridac f Architcctonieidae) on the basis of the conical torinoid operculum (Fischer 1885, p. fSAM). 714. see also Earncs 1952, p. 37), observed in Spccuncas M329S and M3299 were found the Recent form P. (Spirolaxis) centrifuge by the author and specimens M3300 M3303 Montcrosato fMonterosato 1890) and in P. were found subsequently by J. M. Lindsay. h»g. 1. Pwtuhrfmla.\fx (PsvudomuhtMS) asvalplanilus Maxwell. MeCullouah's Btiuee, New Zealand. NZGS. 9508 2; a, ad apical; b, abapical; c. axial views (all X 17.50). He. +JP. (PswHkmuitaxis) ascnlpiurattts Maxwell. McCullou^lt's Bridge. New Zealand. NZCS, 9508-1: iL, adapicul: b, abapical; c, axial view* (all X 17.3). Ffe 3. P {Pwutlontahais ) asadpturatus Maxwell. Adelaide Plains Sub-Basin, Adelaide Children* Hospital Bure No. 5; GSSA, M3299; a, ad apical <\ 161; n. abapical IX lfil; p, axial views IX 15) l'l£. 4. P. (Pjicudo/nataxis) pwmcridiojudis Chapman Clifton Bank. Muddy Cre^k; SAM. PtH^i: t t t adapicnl h tX 106); t abapical 1X10,6); < axial views (X 12.3) ) ) 24 M. F. BUONAIUTO Synopsis oj the history of tlie genus Pseudomalaxis Fischer on the basis oj the major authors, Author Family Subfamily Genus Subgenus Fischer (1885. p r Solariidae — Torinia Gray Pseudomahtxis 714) Fischer Sacco (1892, p. 75) Solariidae Discohelix Dunkcr Pseudomalaxis DaJl (1892, p, 331) Solariidae Discohelix Pseudomalaxis Discosolis'OaU Iredale (1911. Solariidae (?) Pseudomalaxis Pseudomalaxis pp. 253-7 Discohelix Discohelix Discosolis Monterosalo Solariidae — Pseudomalaxis Pseudomalaxis (1913, pp. 362-3) Spirolaxis Monterosato Cossmann (1915, Euomphalidae Euomphalinae Pseudomalaxis Pseudomalaxis pp. 122, 14L) Mestaver (1930, Arcbitcctonicidae Mangonuia p. 144) (syn. Solariidae) Mestayer Awarua Mestayer Rehder (1935, p. Architcclonicidae Pseitdomufasis Pseudomahixis 128) (?) (syn. Discosolis Dall) Paurodiscus Rehder Calodiscttltis Rehder Tredale (1936. Mangonuiidae Mangonuia p. 326) A wurtui Wen/ (1939. p. Arcbitectonicidae Manexmuia Mangonuia 668) (syns Solariidae, A wdrua Mangonuiidae) Pseudomalaxis Pseudomalaxis Spirolaxis Paurodiscus Calodisculus PiKbrv & IVfcGintv Cyelostrematidac Pseudomalaxis Pseudomalaxis (1945. p. 9.) Fames (1952, Architcctonicidae Pseudomalaxis Pseudomalaxis p. 37) Ahbolt ( 1954, Vitrinellidae Pseudomalaxis Pseudomalaxis pp. 138-9) Pchelintsev& Solariidae Pseudomalaxis Pseudomalaxis Korobkov (I960, pp. 137-8) Maxwell (1966. Viirinellidac Pseudomalaxis Pseudomalaxis p. 444) Gilbert (1973, Architcctonicidae Pieudomahnis Pseudotmdaxis p. 30) This paper Arcbitectonicidae Pwudtumduxis Pseudomalaxis (?«yn. Mangonuia) lAlangomtia Spirola.xis Paurodiscus A warua ( ?sy n . Calodisculus '^Calodisculus Systematic Descriptions flattened to subconcave; outer shape of the Phylum MOLLUSCA body-whorl rectangular to quadrangular, inner shape subcircular to similar to the outer; thin ClaSS GASTROPODA margins, two carinae at abaxial-adapical and Order Mp.sogastropoda abaxial-abapical ends. Straight growth lines Superfamily cerithiacea between the abaxial keels. Aperture rectangular Family ARCHITECTON1CIDAE to quadrangular; protoconch heterostrophic- Genus PSEUDOMALAXTS Fischer, 1SK5 anastrophic; operculum torinioid. thin, multi- Diagnosis; Shell discoidal, pscudoplanispiral; spiral, outside flat or conical (after Wenz 1939, nbapical side concave to suhconeave, adapical p. 668). 1 ; NOTES ON PShUDOMALAXlS FISCHER IN AUSTRALIA 25 Subgenus Pseudomalaxis *. itr. whorl is commonly quadrangular to subquad- is swbcircular. Diagnosis. Shell medium-large to very small; rangular, Ihe inner shape margins thin: pseudoplanispiral with tangent whorls: suture Hush to subimbricated; two ada- Pscudomalaxis (Pseudomalaxis) a*cul|>luralits pical and two abapical keels: the four keels MaNwelt. 1966 can be smooth or crenulatcd; growth lines a FIGS 1-3 little irregularly packed; operculum flat (after J9n6 Psendomtduxis metdpttiratus Maxwell, * Wcnz 1939). p. 444, figs 11-13, Material: well Observations: Pseudomulaxis s. str. appears lo 4 specimens very preserved with bo represented in the Tertiary by two main (he peristome slight! v damaged (GSSA, forms. The first predominant in the Eocene- M 3298-9; NZGS, 9508-1, 9508-2) and an- OHgocene, is commonly characterized by other 4 juvenile or damaged (GSSA. M3300-3). growth lines and the four carinae as the only Description: Shell very small \riL\ ihin, bicon- ornaments; a few species bear also spiral orna- cave, nearly planispiral; protoconch hctcrO- ments of fine riblets as in the Eocene P. dixoni strophic-unastrophic; whorls increasing far (Vasscur) and the Oligoeene P. italica (Saceo). more in diameter than in height and overlap- or of beaded cords and keels as in P. texana ping entirely in relation to the coiling axis and (Aldrieh); the outer shape of the body whorl very scarcely in relation to the normal to it- is rectangular to quadrangular, the inner is sub- Suture flush to subimbricated. Body-whorl circular to subquadrangular. the inner is sub- shape: outer subquadrate-subtrapezoidal; inner circular to subquadrangular or subrectangular. subcircular-ovoidal, wider than high. Whorl depending on the thickness of the margins ((he regions : adapical and abapical subconvex latter characteristic could be ontogenetic and abaxial flattened or subconcave. nearly vertical. related to age) Body-whorl regions connected by prominent The second form, predominant in the Neo- smooth abaxial carinae. Abapical side more gene-Recent, is characterized by the develop- concave than the adapical one. ment of a main spiral rib or cord on ihe abaxial Ornaments: Growth lines and rugae, prosocline region of the adapical margin, and often of in the adapical and abaxial regions, opistho- spiral costellae on the abaxial margin. The four cyrt in the abapical. Four carinae: at abaxial carinae and the adapical rib usually bear beads adapical, abaxial-nbapical, adaxial-adaptcal. or short spines. The outer shape of the body and adaxial-abapical ends. DIMENSIONS (in mm): (see Fig. 5) Specimen No, whorls Dw N Nis Hw Lis Hgc Dcg M329S 4 2.55 0.95 0.40 0.50 —0.10 0.60 0.65 M3299 4 2.50 0.92 0.39 0.55 —0,10 0.65 0.60 NZGS 3 L55 0J0 0.30 0.40 —0.05 0.45 0.50 950S-1 N2GS 3 2.15 0.90 0.35 0.5S 0.00 0.58 0.50 9508-2 RATIOS: (see Fig. 51 Specimen HwDw K - I is/Hw 9 Nis/N Hgc/Dgc M3298 0.1961 —0.2000 0.421 0.9231 M3299 0.2200 —0.1818 0.4239 1.0833 NZGS 9 SOS -1 0.25*1 —0.1250 0.42 Bfi 0.9000 NZGS 9508-2 0.2698 0.0000 0.3889 1.1600 Localities: Adelaide Plains Sub-Basin (St Vin- of Torrcns Lake, opposite Kintorc Avenue, at cent Basin, S. Aust); Adelaide Children's Hos- 20.10 m (M3300) and 24.50 m (M3301) pital, North Adelaide, hundred of Yalala, New Zealand: McCullOuglrs Bridge. "I own Acre 717, Bore 5, at 20.42-20.75 m Sfraiigraphic Distribution: Adelaide Plains Sub- IM329S) and 21.64-2 1..95 m (M3299); Bore Basin : Lindsay ( 1 966 ) gave a brief strati- 2, at 22.25-22,56 m (M3302, M3303); Ade- graphic summary of Adelaide Children's Hos- laide Metropolitan Subway, Bore 3, north bank pital Bores 2 and 5. "The interval 20.42-21.95 I lb M. F. BUONA1UTO rn in Bore 5 is low in Blanche Point Marls are certainly from near, and probably from ( Aldingan, Late Eocene). The Han/kt-nina slightly above, the //. primitiva zone. primitiva zone occurs in this bore at .19.20- Thus the vertical distribution of all the spe- 19.51 m, i.e. slightly above P. usculpturants. In cimens of P. asculptttratus found in the Ade- the type section at Maslin Bay, Hamkenina laide area spans a narrow interval from slightly ptim'ttiva is present only in a restricted band below to probably slightly above the H. pr'mti- 0,80-1.15 m above the base of Blanche Point tiva zone, in Blanche Point Transitional Marts, Transitional Marls. Late Eocene/' (J. M. Lindsay, Dept of Mines, 7 Other specimens from Adelaide Children s pers. comm., 1974.) Hospital, Bore 2, are within the Hantkenina New Zealand: Upper Waihao Greensand, Kaiatan. Late Eocene. primitiva zone. Although H . primitiva has not yet been found in Adelaide Metropolitan Sub- Ohsnvfitionv: The two Australian specimens way Bore 3, the specimens of P. asculptural its show an inversion in the coiling, variable in Hw Htr^. i ) f ) feq \V^ '-O.jCfl r : Li I — N'S — r . I N _— ^ Hw < Hgc JV „ Dsv 1 lis <0 Fig. 5. 'the paiumcteis measured in piaiiispirai (a) and in pseudoplanispiral (b) gastropod shell arc here defined an follows: No iv ; the total number of the whorls, aw the total number of revolutions Hgc. HW of the generating curve around the coil- ) !2& n ing axis. Hw; height of any whorl (in thus cas* the last one) as the projection of lh? T < foiling axis of the distance bcrween the - adapical point of the .generating curve Hw H 3 c r at the initial Us = Q position and ils abapieal point at the final position after a 2t revolution in the given interval In*" — 2 I he generating curve is here considered the projection on a plane, containing entirely the coil- ing axis, of the outline of the growing edge of a gastropod shell. The body whorl here repre- sents the part of shell cone generated by the growing edge in a given interval 2n* — 2(n H)**, T is the translation of the generating curve along the coiling axis per revolution. K and i arc respectively Ihe indexes of overlapping of any whorl; K, parallel to the coiling axis: r, uotmal to the coiling axis, and arc defined by the ratios K — Lis/Hw and v — Nis/N, (After Raup J966. Observations consistent with this theory will be presented in a subsequent publicalion.) NOTES ON PSEUOOMALAXiS FISCHER IN AUSTRALIA 27 amount, from a more hclicoidal protoconcb to (Lower Eocene) sample from North Canter- a more planisptral lelcoconch. Undescrihed in bury. The material is not well preserved but rhc Australia, this form was directly compared with shells closely resemble P. tistnlpturutus except two topotypos of tho only coeval species ul tor their much smaller size (Ihc larger speci Pxeiuhnuihtxis known in Australasia, P. asoulff- men is only 1 mm in diameter), i don't think (iirarux Ma\*ell. These two topotypes, more that the lower Eocene shells are [uve-nites ol juvenile than the Australian ones, display a P. Q$atl#Mtr#ilt£, however, as (hey also have somewhat higher spire, producing a decreasing much smaller proioconchs". concavity and consequent flattening of the He quotes also the occurrence ol P. ef, adnpicjil side, as a result o( the same inversion wulpmrntm Maxwell from Wharckuri, South of the coiling; fainter carinac. but the younger Canterbury ( Duntroonian, Ohgoccnc ) [ Max* of the two has them more marked. Protocoach, well 1969, p. 16!) but The sole specimen is growth lines, outer and inner shape of the body broken", however it *'i.s certainly close to the whorl are the same. Because toe .amount of Eocene species'*. inversion m the coiling of the New Zealand farms appears to differ only slightly from that of the Australian forms, and is also variable in !he Australian and New Zealand specimen*. It P*eudomaJaxb (Paeudomulaxb) is very difficult to distinguish fA the specific praenieridionalis (Chapman, 1912) level between these two tormx. Hence, they are heris considered couspecific and representing FIG. 4 probably different geographical morphs. 1912 Homahtxtx praemeritiinttulix Chapman,, hrom P. iaculptunuux Maxwell, the Anglo- p. 186, pi. 12, figs4-ri. Parisian Eocene P. dixoni (Vasseur) (Coss- Muieihil: 1 specimen Very well preserved manu 1D15, pp. 142-3), differs by its higher (SAM, PI 83421. spire and presence of spiral riblets; the Prnto- Atlrialic Priabonian P. hcyrlrhi (Oppcnhcim, Dexvription: Shell small, thin, planoconcave, 1896) by its much higher spire and an abaxial pseudoplanispiral, with the adapical side flat- margin anapically more convergent lo the axis: tened and the abapical concave; whurl.s increas- the Ljguria-Picmonte Middle Oligoccne P. ing far more in diameter than in height and hatirux Sacco. IS92, hy its lower spire and by overlapping entirely in relation lo the coiling if ic presence of spiral ribiets. The Indian axis and very scarcely to the normal tn it. Eocene P- ptmjohcnxix Eames, 1952, seems on Heterostrophic-anastTophic protoeonch with the coniiary >ery close, but the holotype being three smooth whorls, the inversion o\ the coil- very juvenile and the original illustration very ing already displayed in the third whorl. Suture poor, it is impossible to determine the actual grooved. Body-whorl shape, outer subquad differences and affinities between them, rate - trapezoidal: inner subcireular - ovoidal; slightly wider than high. Body whorl regions: The American Claiborne Kocene P. rotetla adapical convex in the adaxial part, concave in (li*a) (Palmer 1937. p. 176) differs by Us icct- .abaxial; abaxial flattened; ahapical convex; anj*utar aperture, higher spire, and lesser num- tho subquadrate peristome. Lip*. ber Of whorls; P. texana (Aldrich) {Palmer aperture with adapical elliptical with a gutter-shaped reflec- 1937, p. I7K) hy crenuhiled a basin I Veels and tion in the middle; parietal abaxial straight: SplcaJ cools ou ihc adapical margin; and P. and I .ip region plrtmwrr/jfi Palmer (Palmer 1937, p. 37R. by ahapical elliptical. and connections subclinical margins. more imbricated sutures, angular, marked by carinac. and the stdrfral part of the last whorl twisted to Orrutmvrttx; Four spiral beaded carinas, the the adapical. abnxiul ones with very short aVmlly elongated Otber Eocene species (Cownann 1915) are spines: adaxial-adapical. adaxial-abapicaL aba* xial-adapical. abaxial-abapical; a spiral cumu- * i- i Anglo-Parisian P. puieUatus (Sowerby), of and tbc Egyptian P. lyhicvs (Oppenbeim). lated cord on the abaxial part the adapical region and a smooth costella on the adapical Maxwell (Nov. 1974, pers. cotnm ) states: part of the abaxial region. Growth lines: proso- 'two specimens of a Punuhntahxis have re- cllne in the adapical region; prosocline in the cently been obtained from n Mangaorapan abaxial; orthocyrt in the abapical. 28 m. r buonau.to OlMHNSrONS I in mm)- (tope Fig- 5) Nil wfaftte Dw N Nis llw Ms Hoc Dgc 4 2 6 13 0.45 0.88 —0.02 0.90 0.9S RATIOS: Hw Dw K Lis.'HW v— Nis/N Hce Dgc 0.3381 —0.0227 0.3461 U.9KS4 Localities and Smitigntphic Distribution: dtotnthv iHedley) (Medley 1 903, p. 351) dis- Muddy Creek Maris, Clifton Bank. Muddy plays, on the other hand, i very close affinity Crock, 6,4 km west of Hamilton; "blue days". to P. ptaemeruiiOnalis ( Chapman ) , differing Newport Formation, AUnnu Bay Coal Shall. trom it only by two more abaxial riblels and Vic. the abapical-adaxial carina with heavier beads. Stnti (graphic Range: Balcombian, Early Middle The New Zealand Recent ?P, boftonsi (Mcs- Miocene (Ludbrook 1973). taycr. 1030. p. 144) differs by more whorls two beaded abaxial cords, broader abapicul- Observations: P. praenu-rklttJtjah's (Chapmnn) abaxial carina and slightly opisthocline growth was referred to Darragh (1970. p. 1 SS) to lines on the adapical region. Xfangotntia Mcstayer. The specimen here des- I'he American Recent P. nohilis (Verrili) cribed represents the second discovery of this iVcrrill 1885, p. 423, p]. 44. fig. 12) differs by species. From P. praenwndionaft's, the Palco- its shorter spire, a greater number of whorls. Mcditerranean Neogene /*. ranclaca (Philippi) slimmer body-whorl, adapical cord closer to iWenz 193?, p, 668. fig. 1906) differs by a the abaxial-ndapical carina, five spiral ribs on slimmer body whorl, higher spire, adapical the abaxial margin, and fine spiral ribs on the spiral costa closer to the adapical-abaxiai abapioil margin. carina, absence of spiral rihlets on the abaxial margin; from the Paleo-Mediterranean Pliocene Acknowledgment* P afdrovandi < Forest i) (Saccn 1 K92, n 75. pi. I dm very grateful to Dr P. A. Maxwell trf 2. rig. 65 a-d, 65 bis a-d) by smooth adapical- the Geological Survey of New Zealand for his abaxial carina, absence of spiral abaxial ri biers, assistance, particularly in lending topotypes of higher spire, and adapical costa Closer lo the Psrudontaiuxis asatlpturatus; to the Director of adapical-abaxial carina; from the Parathetys Mines. South Australia, for the loan of male- Miocene P. bocttgeri Cossmann (Cossmann rial; to Dr H. A. Render, National Museum of 1915, p. 143) by a larger body-whorl, lower Natural History. Smithsonian Institution, tor spire, lesser number of whorls, lack, ot spiral the information on taxonomy; to Dr H Silcock. ornaments except rarely, beaded abaxial Department of Pure Mathematics, University carinae, a semi-circular inner shape of the body of Adelaide, for verifying the definitions of whorl; from the Mediterranean Recent P. the pnramenlers; to Dr N. H. ludbrook for actoni Monterosato (1913, p. 362) by a greater encouragement and for reading the manuscript. number of whorls, slimmer body-whorL lower The work was carried out in the Department spire, more marked growth lines, more abaxial of Geology and Mineralogy, University of riblets, and an abaxial margin more parallel to Adelaide, during tenure of a University Re- the coiling axix- The Australian Recent P. meri- search Grant, References Aa&OTT, R. 1, iiyMl. "American Scashclls/ Darkagh, J. A. (19/0).—Catalogue ot Au&lialian (Van Nofctraud: New York.) Terliarv Mollusoa (except Chitons). Mem. Chapman, F. (1912).—New or lilllt; known Vic- Nalti. fats, Via. 31. 125-212, torian fossil* in the NutionaJ Museum. Pi IEamhs, F. E (3952).—A canti ibtuioti lo the study XVI. Some Tertiary Gasteropoda Prcc. H. of the Eocene in Western Pakistan and West- Soc. Vict, 25. 186-192. pis 12-13, ern India. C. The description of the Scapho- CossMTan, M. (1915).—"Essais de Paleoconcho- poda and Gastropoda from standard sections logic comparei?." Vol. 10. (Paris,) in the Kakhi Nala and Zinrta Pir areas of the Dall, W. H. (J 892). —Contribution to the Ter- Western Punjab and in ihc Kohat District. tiary £aun3 of .Florida with special reference Phil. Trans. R. Soc. London, B, 236(63 1), to the Miocene Silex Beds of Tampa and the 1-168, pis 1-6. Pliocene Beds of the Caloosahutchie River. FlSCUcft. P. (1880-87). —"Manuel de conchyologic Pt. 2. Strcptodont and other Gastropods ct de Pideontologie conchyologique on fnstoire Trans. Wagner free faith Sci. Phi fad, 3i'2l. nalurctle de?. mullusques vivunta el iWalts/' 201-473, pis 13-22. (Savy: Paris.) — NOTES ON PSEUDOMALAXIS FISCHER IN AUSTRALIA 29 Glirfrt, M. (1973).—Revision des Gastropoda Oppenheim, P. (1896).—Das Alttertiar tier Colli du Danien et tin Montien de la Belgique. Pt I. Rerici in Venetian, die slcllung dcr schiclcn Les Gastropoda du Calcaive de Mons. Mer&. von Priabona und die oligocane Transgression Inst. R. Sot\ Nat. Betg. 173, 1-116. 11 pis. in Alpinen Europa. Z. Dt. geol. Ges. 48. 27- Hedley, C. (1903).— Scientific results of the 152. pis 2-5. 1 trawling expedition of H.M.C.S. ''Thetis ', PALMbR van Winkle, K. (1937)-—The Clairbor- Mollusca. Pt II. Scaphopoda and Gastropoda. nian Scaphopoda, Gastropoda, and Dibran- 36-8. Mem. Austr. Mas, 4, 327-402, pis chiate- Cephalopoda of the Southern United T. Notes. Irt:dai.f. (1936). —Australian Molluscan States. Bull. Am. Paleont. 7(32), Pt I. 1-548; pis No. 2. Rec. Austr. Mus. 19(5), 267-353. Pt 2, Alias. 20-24. PCULLIKTSEV. V., & K.OROBKOV, I. A, ( 1 960). iREini e, T. (1911). —On some misapplied Mol- Osnovy Paleontologii. Spravochnik dlia pale- lusca generic names. Proc. Malac. Soc. T.ond. onlologov i geologov SSSR. Mulluski-Bruko- 9, 253-265. nogie. (Moscow.) Lindsay, J. M. (1969).—Cainozoic Koraminifcra and Stratigraphy of the Adelaide Plains Sub- Pilsbrv, H. A,. & McGinty, T. L. (1945).— Basin, South Australia. Bull. geol. Sun-. S. Cyclostrematidae and Vitrinellidae of Florida. Aost. 42. Pt. I & II. Nautilus 59, (1) 1-13, pis 1-2; (2), Li/obrook, N. H. (1973).—Distribution and strati- 52-59, pi. 6. graphic utility of Cenozoic Molluscan Faunas Raup. D. M. (1966).—Geometric analysts of shell in Southern Australia. I ohoku Univ., Set. coiline: general problems. /. Paleont. 41, Rep., 2nd Bejr: ( Geol ) , Special Volume. 6 43-65. 24- (Hatiti Memorial Volume), 241-261, pis Rrnor.R, H. A. (1935).—"New Caribbean Marine 28, Shells." Nautilus 48(4), 127-30, pi. 7. Maxwtll. P. A. (1969).—Middle Tertiary Mol- luscs from North Otago and South Canter- Sacco. F. (1892).—T Molluschi dei Terreni Ter- bury, New Zealand. Trews. H. Soc. N\Z„ ziari del Piemonte e delta Liguria, Part 12. GeoL 6(13), 155-185, pis t-3. (Clausen: Torino.) Maxwell, P. A. (1966). —Some Upper Eocene Sutlr, H. (1913-15),—"Manual of the New Zea- Mollusca from New Zealand. N.£. //. Geo!. land Mollusca." (MacKay: Wellington.) Geophys, 9. 439-57. Vkrrill. A. E. (1885),—Third catalogue of Mol- Mi:.stayer, M. K. (1930).—Notes on New Zea- lusca recently added to the fauna of the New land Mollusca. No. 5. Trans. N.7.. Inst, 61, England Coast and the adjacent parts of the 144-46, pi. 26. consisting mostly of deep-sea species. Monterosato, Marquis de (1913). —Note on the Atlantic, previously recorded. genus Pseudomalaxis Fischer and description with notes on others Trans. Connect. Acad. Arts Set, 6, 395-452. of a new species and subgenus. Proc. Mulai\ pig 42-44. Soc. Land. 10, 362-363. Moore. R. C. (I960).—"Treatise on invertebrate Wenz, W. (1939).—Gastropoda- In O. H. Shinde- Palaeontology." Part I, Mollusca, 1. (Geol. wolf (lid.), ^Handbtich der Palaozoologic". Soc. Amer. & Univ. Kansas Press: New Vol. 6, Part 3(4). (Gebruder Borm'tracgcr; York.) Berlin.) THE CLINGING MECHANISM OF PSEUDOPHRYNE BIBRONI (ANURA: LEPTODACTYLIDAE) TO AN ALGA ON GLASS byN. Gradwell* Summary GRADWELL, N. (1975). -The clinging mechanism of Pseudophryne bibroni (Anura: Leptodactylidae) to an alga on glass. Trans. R. Soc. S. Aust 99(1), 31-34, 28 February, 1975. Despite the absence of an oral sucker, tadpoles of all stages from 26 to 40 (of Gosner 1960) were found to be capable of clinging by their jaws to an alga on vertical glass. When the glass was wiped clean of the alga, Phyllobium sp., tadpoles were no longer able to attach themselves. Therefore substratum algae are necessary for the clinging of the tadpoles to glass. As the nares appear to suffice as inhalent channels, the dental apparatus of tadpoles is adapted to maintain a firm grip on the alga. There is an absence of peripheral papillae adjacent to the most posterior of the tooth rows of the lower lip. Therefore this tooth row can bend farther forward and the security of its grip on the alga is probably increased. THE CLINGING MECHANISM OF PSEUDOPHRYNE BIBRON1 (ANURA: LEPTODACTYLIDAE) TO AN ALGA ON GLASS by N. Gradwell* Summary Gradwell, N. (1975).—The clinging mechanism of Pseudophryne bibroni (Anura: Leplo- dactylidae) to an alga on glass. Trans. R. Soc. S. Aust. 99(1), 31-34, 28 February, 1975. Despite the absence of an oral sucker, tadpoles of all stages from 26 to 40 (of Gosner I960) were found to be capable of clinging by their jaws to an alga on vertical glass. When the glass was wiped clean of the alga, Phyllobium sp., tadpoles were no longer able to attach themselves. Therefore substratum algae are necessary for the clinging of the tadpoles to glass. As the nares appear to suffice as inhalent channels, the dental apparatus of tadpoles is adapted to maintain a firm grip on the alga. There is an absence of peripheral papillae adjacent to the most posterior of the tooth rows of the lower lip. Therefore this tooth row can bend farther forward and the security of its grip on the alga is probably increased. Introduction has been compared with the suckers of five It is well known that anuran tadpoles are Australian species (Gradwell 1975). In addi- adapted to occupy a wide variety of ecological tion, Tyler ( 1 963 ) described the external niches (Noble 1931, Orton 1953). Of the mor- appearance of the sucker of Litoria arfakiana phological adaptations, an attachment mechan- (as Hyla angularis), which is much like that ism for clinging to substrata is a predictable of Ascaphus. association with a lotic habitat. It is thus pos- Another type of tadpole sucker is that which sible to avoid dislodgement by swift currents lies posterior to the mouth. The anatomy of even in torrential streams. In contrast, the ad- such a sucker has been described in Staurois ricketti hesive secretion produced by the ventral glands by Noble ( 1931 ) and in Staurois of most young embryos (at stages 18 to 24, of afghana (Bhaduri 1935). No data are available Gosner 1960) is too weak to withstand fast on the magnitude of its suction, except that if currents (Gradwell, unpublished). There is no a tadpole is manually lifted out of the water by published evidence that these glands can pro- its tail, its sucker can support a stone sixty duce subambient hydrostatic pressures. They times the weight of the tadpole (Hora 1922). atrophy after stage 24, and in the tadpole which The present paper reports a mechanism em- represents stages 25 to 40, either of two kinds ployed by lentic tadpoles of the Australian lep- of non-glandular sucker may develop. todactylid frog Pseudophryne bibroni which en- An oral sucker is the most usual adaptation ables them to cling to a vertical substratum, in lotic tadpoles and, accordingly, the degree even though they lack a sucker. I have ob- of development of this sucker would seem to be served such ability in nature for many other influenced by the velocity of the ambient water. suckerless tadpoles and have assumed that they In most lotic species the periphery of the upper use their teeth to secure a grip on rocks. and lower lips is continuous and forms a suc- torial disc, and the number of tooth rows in- Results and Discussion creases progressively with the velocity of the On 18 July 1974, 16 tadpoles of Pseudo- ambient water (Noble 1931; Gradwell. unpub- phryne bibroni (at stages 26 to 29) were col- lished). The structure and function of a tad- lected in a flooded ditch (ca 30 m long by 5 m pole's oral sucker has been described in wide) with sides sloping to a depth of 2.5 m. Ascaphus truei (Gradwell 1971, 1973) and it This habitat is about a kilometre from Boyd * Department of Environmental Biology, The N.S.W. Institute of Technology, P.O. Box 123. Broad way, N.S.W. 2007. 32 N. GRADWELL B w» % Fig. 1. A. The tadpole of Pseudophryne hibroni (stage 36) clinging to algae on the vertical glass side of an aquarium. B. Oral apparatus of a clinging tadpole as seen through a thin transparent layer of the alga Phy/lobium sp. growing on the aquarium glass. As the lower beak is enclosed within the" upper beak, only the latter is visible. The lower labial teeth are inclined forward to grip the algae. Adduction is accomplished by a slight lateral compression of the lips and teeth. C The oral apparatus dissected free from a tadpole to show the beaks in an open condition. Calibration bars = 1 mm. SUBSTRATUM CLfNOlNG IS rSFUDOVHRYSE u River Crossing, near Jenolan Caves. N.S.W. cial interest to find akinetcs m the manicotto suggests 1h«t The maximum depth of water was 0,5 Ffl and ("stomach"! of tadpoles, which the bottom consisted of mud and fallen Euca- thev hud been grazed "If the glass. II would lypws leaves and bait. At 3.00 p.m. the Wait* also seem that the bulbous akinetes on their temperature was 6 to ft C. Five hours later^ the strand-like thallus could easily be gripped by tadpoles were placed in an aerated aquarium the minute sharp teeth of tadpoles, thus provid- at approximately the same temperature. ing anchorage. It is possible that other filamen- For initial identification, the mouth pads of tous algae may also permit the clinging of the tadpoles were examined and compared with these tadpoles to substrata* the description of Martin (1965). Beeau.se the There are no papillae behind the most pos- absence of a sucker was noted, it was surprising terior row of lower labial teeth. Therefore 1 his to observe that all of the tadpoles clung to the row of teeth can bend farther forward than if vertical glass sides of the aquarium (Fig. 1A). papillae were present here and so the leeih can This attachment was insecure, for these tad- grip substratum plants more firmly. However. pole* were sometimes displaced from their some other suckcrlcss tadpoles (for example, clinging sites when other tadpoles collided with Lhoiki vemw(xi) can cling feebly to vertical them. Examination of the oral region of a substrata in similar fashion even though they clinging tadpole, by stereomictoscope through lack a posterior gap in, their peripheral papillae. the "lass, showed that the lower beak, was kept Four tadpole* were raised into Juvenile i rogs. closed within the edge of the upper beak (Fig. Until stage 40. the tadpoles < entire length, 16 .U IB) and that the upper and lower tooth tows mm; snout-to-vent length, 17.4 mm) were ahle were held adducied, though not touching one to cling by their teeth to the substratum- Alter another. The nates were the only inhalent chan- this stage the beaks and teeth were shed prioi nels for gill irrigation. in widening of the mouth. During preparations for photography it was It is proposed here that Psettdophryitr found that the. inside of the glass aquarium wan bihroni ladpoJes are able to cling to vertical covered with a thin layer of algae which pre- substrata by biting their teeth into algae and vented sharp focussing on the mouth and teeth. perhaps other vegetation. As the naruil inflows Therefore, half of one side of the aquarium appear to be Suffi&lcnt for respiration, the juw« was cleaned of its algae, and camera conven- need not open and close rhythmically to admit iently positioned to await the settling and cling- additional water, but can maintain their hue ing of a ladpole on the cleaned glass Although on plants. However* I have not had the oppor- tadpoles attempted to cling to the cleaned glass, tunity to compare these findings with the ability they were unsuccessful. However, they clung of P. bibroui in their natural habitat to cling readily to the uncleaned glass, demonstrating to .subslrata. The absence of an oral sucker the presence Of algae on the glass to be neces- in these tadpoles confers greater flcviHiUty on sary foi xhe clinging of these tadpoles. the faws and perhaps it broadens their food Apart from clinging to the glass, the tad- resources. On the other hand, an oral sucker poles also showed frequent feeding movements in lotic tadpoles facilitates grazing on algae over the elaas dunng which their jaws opened in swift currents (Gradwell 1971 ), where more and closed rhythmically in a scraping action. tenacious gripping is needed. Tigure IC shows the jaws its an open condition and both the upper and lower beaks ate visible Acknowledgements The predominant alga growing oil the glass was I am grateful to Miss P. McDonnell and Mr 1. Phvlfobium sp. t identified from Pntscolt 1970 B. Wilson Tor collecting the tadpoles, and KO which has a thallus of branched stiands. At the Miss Vf. Anstis for identifying them. Mr M. J. ends oi the strands are swellings called aki- Tyler kindly tend the manuscript and ofTefed nctes, which contain chloroplasts. It was of spe- helpful suggestions, Reference* (' ).- ttiiApLiu. J. L. (1935).—The unatomy of the Giunwri c, N- 197 1 Astaphux tadpole: experi- adhesive apparatus in the tadpole of Rana ments on the suetiou WW gill rnigniuui ajshatiu Gunthcr. with special reference to the mechanisms. Cun. I, ZooL 49, 307-332. adaptive modifications. Tranx. R. Sov. Edfft- tmrgh 58. 339-349 GR.vnweiL, N. < 1^73 ). —On the functional mor. GnsNErt. K- L. (I960).—A simplified table for phology of taction and gill miration in die staging aoucan embryos and fofV*c wiih noies tadpole of AsqaphUfo njvd miles Gradwfxl, N. (1975).—Experiments on oral suc- Orton, G. L. (1953).—The systcmatics of ver- tion and gill breathing in five species of Aus- tebrate larvae. Systematic ZooL 2, 63-75. tralian tadpole (Anura: Hylidae and Lepto- dactylidae). /. ZooL, in press. Prescott, G. W. (1970).—"How to know the Hora, S. L. (1922).—Animal life in torrential freshwater algae." (W. C. Brown Co., streams. /. Bombay Nat. Hist. Soc. 32, 111- Dubuque, Iowa.) 126. Martin, A. A. (1965).—Tadpoles of the Mel- Tyi.er, M. J. (1963).—A taxonomic study of bourne area. Vict. Nat. 82, 139-149. amphibians and reptiles of the central high- Noble, G. K. (1931).—"The Biology of the lands of New Guinea, with notes on their Amphibia." (McGraw-Hill: New York. Re- ecology and biology. 11. Anura: Ranidae and printed 1954, Dover, New York.) Hylidae. Trans. R. Soc. S. Aust. 86, 105-130. GENETIC EVIDENCE FOR THE EXISTENCE OF TWO SEPARATE POPULATIONS OF RATTUS FUSCIPES GREYII ON PEARSON ISLAND, SOUTH AUSTRALIA byL. H. Schmitt* Summary SCHMITT, L. H. (1975). -Genetic evidence for the existence of two separate populations of Rattus fuscipes greyii on Pearson Island, South Australia. Trans. R. Soc. S. Aust. 99(1), 35-38, 28 February 1975. A study of genetic variation of the enzyme glutamate oxaloacetate transaminase (GOT) reveals the presence of two distinct populations of the southern bush-rat (Rattus fuscipes greyii) on Pearson Island. Two allelic genes, Got-f and Got-l are present in the animals collected from the middle and southern sections of the island, while Got-f is absent in animals taken on the northern section. This is discussed in relation to the Pearson Island wallaby which was, until recently, restricted to the northern section of the island. GENETIC EVIDENCE FOR THE EXISTENCE OF TWO SEPARATE POPULATIONS OF RATTVS FUSCfPES GREYI! ON PEARSON ISLAND, SOUTH AUSTRALIA by L. H. Schmitt* Summary Schmitt, L. H. (1975).—Genetic evidence for the existence of two separate populations of Rattus fi/.scipps gXtyii on Pearson Island, South Australia. Traits. R. Sov. S. Aust. 99(1), 35-38, 28 February 1975. A study of genetic variation of the enzyme glutamate oxaloacetate transaminase (GOT) reveals the presence of two distinct populations of the southern bush-rat {Rattus fuseipes greyii) on Pearson Island. Two allelic genes, Got-l« and Got-l b are present in the animals f collected from the middle and southern sections of the island, while Gnr-f > is ahscnt in animals taken on the northern section. This is discussed in relation to the Pearson Island wallaby which was, until recently, restricted to the northern section of the island. Introduction Pearson Island, which lies 60 krn off South Australia's west coast, is divided into three dis- crete sections (Fig, 1 ) , The southern and middle sections are linked by a causeway, while the middle and northern sections are separated by a narrow sea channel. The total area of the island is approximately 325 hectares. The Pear- NORTH son J, wallaby, Petrogafe sp. (see Thomas & SECTION Dclroy 1971, for a discussion on its taxonomic status) and the native rat, Rattus fuseipes greyii, are the only terrestrial mammals which are known to inhabit the island. The native rat, which is found on all three sections of Pearson I., was first described by Thomas (1923) who named it Rati us murrayi. Tredalc & Troughton (1934) reclassified it as Rattus greyii murrayi recognizing its close rela- MIDDLE tionship to Rattus greyii greyii, the native SECTION bush-rat of mainland South Australia. It was later included in the species Rattus fuseipes by Ellenrmn (1949) as a distinct .sub-species, /c. fuseipes murrayi, along with the mainland form R.f. greyii. Recently, Taylor & Horner (1973a) have considered it to he subs peri fie ally indis- SOUTH tinguishable from RJ. greyii. SECTION Until I960, the Pearson I. wallaby was only found on the northern section of the island. No 250 evidence could be found to suggest the species ever inhabited the other two sections, despite Tig. 1 . Map of Pearson Island. Areas where ani- the suitable habitat available there. The channel mals were captured arc indicated by shad- appeared to act as a very effective barrier to ing. Department of Genetics, University of Adelaide, S. Aust. 5000. I. 36 K SCHMM I migration between the northern and middle TAuLII 1 sections. In I960, six wallabies, including either Family dura on GOT variation four or live females, were accidentally released N timber of Number and OOT phc on the middle seetiort and the species is- now »iF parentis ul oUs'Ting abundant on the middle and southern sections A AR (Thomas & Del toy iy69). Ax A 2 3 1) This paper describes a genetic difference in A at A* lf> 44 H the fl.f. xreyii population ol Pearson I., appa- AxB 3 9 A x AK 1 2 (1 rently caused by a restriction in migration HxAB 1 tl 4 (. across die channel separating the northern and middle sections*, * One parent was nut scored for GO'I phenotype. However, in each of the ten matings ii was Methods known tu have come from a population appa- Specimens of H.L grttyti were caught in rency monomorphic for the Uot-J" allele. "Shei-marT traps and transported alive to Ade- daia on the inheritance of the GOT variation laide. Tissue homo&euatcs were prepared and (Tabic 1) »s cosistenr with a 1 locus, 12 subjected to starch gel electrophoresis. The allele mode of inheritance. Llcclrophorelic buffers (pH SO) were essen- I he H.f* greyii population on the northern tially the same as those described by Scfander section is monomorphie tor the Got /'' allele, i-' (tl, >- glu- (1971 The methods used to detect h while hoih frV>r-/" and Got l are present in tamatc oxaloaeetate transaminase (GOT) acti- the population on the southern and middle vity and determine the subcellular locality of sections (Table 2). The genotypic frequencies the isozymes were modified from those given in the latter population 111 the Hardy-Wcinbere by De Lorenzo & Ruddle (l a 70). Heart ex- equilibrium frequencies (P>0.05). If the tracts were used predominantly, except in lhe Got-jf* allele is present in the population on latter procedure when liver extracts were used. Lhe northern section, then there is a V5% pro- Results and Discussion bability that its frequency is less than 3%. In Animals were caught from the areas shown any case, the Irequeney o! the Got~i h allele in ih Fig. 1. Seventy-five animals were caught in lhe northern section is significantly different ?iy ftpp nights, yielding a capture rate of from its value in (he muldle and southern **o- 34'% . This is coasiderably higher than the ttot* sent in gels, one migrating eathodaJly, the other Numhcr ftttfl GOT r>hcni*uyr>e h'tjUPtfnC] Alijft i:anj»Tn A AB B of anocially, The cathodal area of activity con- Oot-lb sisted of a single invariant band. This isozyme Nortlurn section I. DO Middle afTds&tnftiMn 0*3*5 predominated in mitochondrial extracts. The sve (ion anodally migrating; isozyme was found in the supernatant fraction and was variable. Three It seems unlikely rhat the marked difference distinct phenolypes, GOT-IA, GOT-1B and ii) allelic frequencies is maintained by selec- GOT-IAB were observed. This variation Is tion. All three sections of the island appear to consistent with the active enzyme heing a provide very similar habitats fur H.f. grt^ii. duneiic molecule and is similar to that found The observed absence of Gtyf-/* from the in man (Chen & (iihlett 1971 ) and Lire North northern section indicates a severe restriction in American old-field mouse. Fetomyncits pollo- gene flow between the areas sampled on the notuK (Selander ft af. 1971). Genotypes can middle and northern sections. The most he assigned to each phenotype. presuming that obvious point of demarcation is the channel the difference is under the control of an auto- separating the two sections. This surprisingly somal locus, with two co-dominant alleles. This low level of migration between the northern Incus has heen designated Got 1 and the alleles and middle sections is similar to rhat found in h Gat-J" and Got-i , the Pearson I. wallaby Laboratory matings of R,f4 greyii individuals It would appear improbable that the rats are from different, areas of South Australia, includ- not physically capable of crossing the channel sea is low, ing Pearson Li have been successful. Family When the calm and the tide ft is GFNETICS OF THE PEARSON ISLAND RAJ 37 erable time after the isolation of the island There is some evidence to suggest that since the isolation of Pearson Island from the main- land, the mean sea level on two or more occa- sions has been about 6 metres above ils presenl level (Twidalc. pcrs. comm.). During these limes of high mean sea level, the channel would have been considerably larger than at present. Thomas & Delroy ( 1971 I suggested that the wallahy did not cross the channel because it found the sea water distasteful. Another possi- bility is thai, because ot the action of the sea. there may be strong selective pressures against small animals which wander too close to the edge of the water. Under such an hypothesis. genotypes would be favoured which predis- posed animals to an aversion to moving close iltM to the water's edge. This would discourage the animal from crossing between the middle and northern sections. Various suggestions may he Offered to account for the present distribution of the Iwo A AB A B A AB alleles at the Got-I locus. All other popula- tions of R.f greyii studied {Greenly I.. Hop- kins I.. Kangaroo I., Eyre Peninsula and Mount Lofty Ranges) have been found lo be mono- 1 morphic for the Got-l * allele (Schmitl. unpub- lished). 1 he polymorphism on Pearson L may have been present before the channel was formed and at that time, or some time afier- wards, the Got-h 1 allele was lost from the northern section. Alternatively, one of the alleles could have arisen by mutation, since the 1 channel was formed. If the Got-l' allele is the Origin more recent mutant, then it has lo be postu- lated lhat it subsequently migrated across the channel. The channel separating the northern and middle sections has not only acted as an effec- Fig. 2. GOT variation found in /?./. greyii from tive barrier to migration lor the Pearson I. Pearson I. The pattern of variation is in wallaby, but from the genetic evidence pre- agreement With a Uimeric molecule being sented here, has the active enzyme. also had a similar effect on R ./. xreyii. easy for a man lo wade or step from rock to Thirteen other gene loci, for which about 70 rock between the two sections. However, the specimens of R.f. greyii from Pearson I. have eonfoimalion of the channel is probably in g been scored, are monomorphic on Pearson state of continual change. The relationship be- Island and show no differences across the chan- tween sea level changes and the occurrence of nel. However, all Pearson I. animals have one or more species of small macropod mar- haemoglobin and malate dehydrogenase pro- supials on small islands off the Western Austra- teins that arc electrophoretically distinct from lian coast all other has been discussed by Main ( 1961 ). populations studied (Schmitt, unpub- From Main's dala it appears that Pearson I. lished). has been isolated from the mainland for at least Further studies on the situation described 10.500 yearv The northern and middle sections here would be a useful part oi any future expe- would have remained connected for a consid- ditions to Pearson I. 38 L. H. SCHMITT Acknowledgments Island was generously arranged by Mr M. I wish to thank Dr D. L. Hayman, Dr R. M. Koch and provided by Mr R. Baker and Mr K. Hope and Professor J. H. Bennett for their White. The South Australian National Parks thoughtful discussion and help in the prepara- and Wildlife Service granted a permit to allow tion of the manuscript. Transport to Pearson me to collect specimens. References Chen, S. H.. & Giblett, E. R. < 1971).—Genetic Selander, R. K„ Smith. M. H.. Yang, S. Y., variation of soluble glutamic-oxaloacetic Johnson, W. E., & Gentry. J. B. (1971).— transaminase in man. Am. J. Hum. Genet. 23. Biochemical polymorphism and systematica in 419-424. the genus Peromyscus. 1. Variation in the old- field mouse {Peromyscus polionotus). Stud. De Lorenzo, R. J.. & Ruddle, F. H. (1970).— Genet. VI, 49-90. (University of Texas Pub- Glutamate oxalate transaminase (GOT) lication 7103.) genetics in Mus musculus: Linkage, poly- Taylor, J. M., & Horner, B. E. (1973a).—Results morphism, and phenotypes of the Got-2 and of the Archbold expeditions. No. 98. System- Got-1 loci. Biochem. Genet. 4, 259-273. atics of native Australian Rattus (Rodentia: — Muridae). Bull. Am. Mus. Nat. Hist. 150(1), R. ( 1949). "The families and Ellerman, J. 1-130. genera of living rodents." Vol. 3, pt 1. (Bri- — Taylor, J. M., & Horner, B. E. < 1973b). tish Museum (Natural History), London.) Reproductive characteristics of wild native Australian Rattus (Rodentia: Muridae). Aust. Iredale, T., & Troughton, E. LeG. (1934).—A J. Zool. 21, 437-475. check list of the mammals recorded from I. M., & Delroy, L. B. (1971 ).— Pearson Australia. Mem. Aust. Mus. 6, 1-122. Thomas, Island expedition. 1969. 4. The Pearson Island Main, A. R. (1961).—The occurrence of Macro- wallaby. Trans. R. Soc. S. Aust. 95, 143-145. podidae on islands and its climatic and eco- Thomas, O. (1923).—The native rat on Pearson's logical implications. /. R. Soc. W.A. 44, Islands, S. Australia. Ann. Mag. Nat. Hist. 84-89. Ser. 9, 11, 601-602. TWO NEW SPECIES OF THE GENUS CLOACINA (NEMATODA: STRONGYLIDA) FROM THE TAMMAR, MACROPUS EUGENII by Patricia M. Mawson* Summary MAWSON, P. M. (1974). -Two new species of the nematode genus Cloacina (Nematoda: Strongylida) from the Tammar, Macropus eugenii. Trans. R. Soc. S. Aust. 99(1), 39-41, 28 February, 1975. Cloacina smalesae and C. kartana are from the stomach of Macropus eugenii from Kangaroo I. C. smalesae is distinguished by the shape of the leaf crown elements, postoesophageal excretory pore, swollen cuticle at the anterior end of the body, and a long vagina. C. kartana is distinguished by the very small cephalic papillae and the presence of two oesophageal teeth situated shortly behind the nerve ring. TWO NEW SPECIES OF THE GENUS CLOACJNA (NEMATODA: STRONGYLIDA) FROM THE TAMMAR, MACROPVS ElIGENU by Patricia M, Mawsoic* Summary Mawson, P. M. (1974).—Two new species of the nematode genus Cloucina (Nematoda: Strongylida) from the Tammar, Macropus eugenii. Trans. R. Soc. 5. Aust. 99(1), 39-41, 28 February, 1975. Claacim smalesae and C. kartana are from tbe stomach of Macropus Fugenii from Kangaroo I. C. smalesae is distinguished by the shape of the leaf crown elements, postoeso- phayeal excretory pore, swollen cuticle at the anterior end of the body, and a long vagina. C. kartana is distinguished by the very small cephalic papillae and the presence of two oesophageal teeth siLuated shortly behind the nerve ring. Introduction The buccal ring is circular and stoutly built. The species which are described in this Its anterior and posterior walls are sligluty paper occur commonly and in considerable lobed (Figs 1, 2). The oesophagus is cylin- numbers in the stomach of the Tammar on drical for most of its length, ending in a small Kangaroo Island. They were taken in the swelling. There are no oesophageal teeth. The course of a quantitative analysis by Mrs thickened lining of the lumen is unusually dis- Lesley SmaJes of the nematodes present at dif- tinct, appearing in the whole mount as three ferent times of Ihe year in the stomach of this wavy longitudinal lines throughout the length host, undertaken as part of work for a Ph.D. of the oesophagus. In transverse section the degree in the Department of Zoology. lumen appears tri radiate with a thickening of Types of the new species will be deposited the lining In the midlength of each arm. The in the South Australian Museum; paratypes are nerve ring lies at about midlength of the oeso- in the Helmintbologicul Collection of the Zoo- phagus. The excretory pore is postoesophageal. logy Department, University of Adelaide. and the cervical papillae lie a little distance in Measurements of the two species arc given front of the nerve ring. :n Table 1. The bursal lobes are only slightly divided. The arrangement nf the rays (Fig. 6) is Qoacioa smalesae A. &p. typical for the genus. The genital cone is not 1-7 FIGS prominent; it bears a small ala on each side of Host and Locality; Mtxropus cugeni: ( Dcs- the cloaca, but no other appendages. The nurest). from Kangaroo T. f S. Aust. spicule is a little less than half the body length. This is a medium-sized species of Cloncinu, The tail of the female is conical, ending in with markedly swollen cuticle at the anterior a point. The vulva is slightly less than a tail end. The thickness of the cuticle increases length in front or tbe anus. The ovejectors unite from the level of the posterior end of the oeso- 6-7 tail lengths in front of the vulva, and the phagus to the base of the mouth collar, which vagina curves forwards before passing hack, it surrounds like a platform. The tnouth collar with one or two twists, lo Ihe vulva. is well developed and slightly lobed anteriorly; The rounded elements of the leaf crown it bears the four submedian cephalic papillae seen in this species have been described in only and two amphids. The distal segment of each one other species. C. tongifttbiata Johnston & cephalic papilla is longer and slightly thicker Mawson, 1939b (syn: C. minor J. & M., nee. than the proximal segment. Each clement of Davey & Wood, 1938). However, in C Jongi- the leaf crown is domed anteriorly and is with- labiata the vagina is shorter, Ihe cephalic papil- out the unguiform anterior projection usually lae are of a different shape and there is no present to species of this genu*. cuticular inflation at the anterior end. /Oology Department, University of Adelaide, S Aust. 5000. 40 PATRICIA M. MAWSON Figs 1-8 Claacina sttuttesae. Fig. 1.—Anterior end, lateral view. Fig. 2. —Anterior end, .showing leaf crown in everted position. Fig. 3. —Anterior end, en face. Fig. 4.—Oesophageal region. Pig, 5. —Transverse section of oesophagus, anterior to nerve ring, Fig. 6. —Posterior end of female. Fig. 7.—Bursa, flattened out and viewed from the inside. Fig. 8. —Lateral view of genital cone. Figs t, 2, 3, 5 and 8 to same scale, Figs y iA Cloacina keirtana. Figs and 10.—Sublateral and en face views of anterior end. FJg. It.— Oesophageal region. Fig. 12.—Transverse section of oesophagus, showing one oesophageal tooth Fig. 13.—Posterior end of female. Fig. 14.—Bursa. Fig. 15.— DorsaJ ray. Figs 10 and 12 lo same scale, Cloacina kartana n. sp. the proximal segment is longer and slightly FIGS 8-14 thicker than the distal one. The buccal ring is stoutly built, and is thicker posteriorly than Host and Locality; Macropm eugenii (Desmn- anteriorly. The six elements of the leaf crown rcst), from Kangaroo I., S. Aust. do not, in the resting position at least, project This is a medium-sized species of Cloac'ma above the collar. with a well developed mouth collar bearing the The oesophagus is cylindrical for most of four small submedkun cephalic papillae and its length, with a terminal bulb. There axe two the amphids, The cephalic papillae are small; small subvcntral oesophageal teeth, one a little — — TWO SPECIES Ob CLOACiNA 41 TABLE 1 Measurements of Cloacina sirutlesae and C. kartana; in mi unless otherwise stated. C. smalesae C. kartana Length (mm) 9.5- L 1.4 13.0-15.8 8.0-10.9 12.4-17.0 Oesophagus 690-720 710-790 680-790 810-900 Antr. end—nerve ring 360-400 360-490 300-350 340-400 —cerv. pap. 200-230 160-200 220-300 250-300 —excr, pore 790-S70 910-1000 600-740 700-800 Spicule length 4200-4700 1400-1600 Length/spic. 1. 2.2—2.5 5.7-7.1 Length/oesoph. lli-16.0 16.8 21.8 10.6-16.0 J 4.3-20.9 Tail 220-260 200-300 Postr.—vulva 350-490 380-500 Egg length x breadth 170-175x85-90 180-185x85-90 posterior to the other, at about midlength of a tail length in front of the anus; the vagina* the oesophagus, and shortly behind the nerve which is straight, extends about 3-4 tail lengths ring. The cervical papillae lie just in front of in front of the vulva. the nerve ring, and the excretory pore is at the The species most closely resembles C. obtusa level of the oesophageal bulb. Johnson & Mawson, 1939a, but differs in the The spicules are relatively short. The lobes spicule length. of the bursa are distinct but not deeply sep- arated. The rays are arranged as shown in Acknowledgments Figs 13 and 14. The genital cone is small and My thanks are due to Mrs. Smales for the no appendages were seen on it. collection of the specimens described. Part of The female tail is conical, ending in a nar- the work was done under a Grant from the rowed point. The vulva lies a little less than Rural Credits Development Fund. References Davxy, D. G., & Wood, W. A. (1938).—New JoHNsroN, T. H., & Mawson. P. M. (1939a). species of Trichoneminae (Nematoda) from Strongylate nematodes from marsupials in Australian kangaroos. ParasitoL 30, 258-266. New South Wales. Proc. Linn. Soc. N.S.W. 64, 513-516. Johnston, T, H„ & NfAWsON, P, M, (1938). Johnston, T. H.. & Mawson, P. M. ( 1939b). Strongyle nematodes from central Australian Sundry nematodes from eastern Australian kangaroos and wallabies. Trans. R. Soc. S. marsupials. Trans, R. Soc. S. Ausi. 63, Am. 62, 263-286. 204-209. THE GENERAL WATER CIRCULATION OF SPENCER GULF, SOUTH AUSTRALIA, IN THE PERIOD FEBRUARY TO MAY byD. A. Bullock* Summary BULLOCK, D. A. (1975). -The general water circulation of Spencer Gulf, South Australia, in the period February to May. Trans. R. Soc. S. Aust 99(1), 43-53, 28 February, 1975. Historical temperature and salinity data are presented together with the three-dimensional velocity structure of the general water circulation of Spencer Gulf, deduced from the data using a numerical model. In the southern area of the Gulf a cyclonic gyre is found which exchanges Gulf water with the ocean water outside the Gulf. On the western coast of the Gulf, a moderate boundary flow, which is called the "Port Lincoln Boundary Current", is evident at all depths. The water in the northern portion of the Gulf circulates both in the vertical and the horizontal and appears to form an almost closed system separate from the system in the southern region. THE GENERAL WATER CIRCULATION OF SPENCER GULF, SOUTH AUSTRALIA, IN THE PERIOD FEBRUARY TO MAY by D. A. But. lock* Summary BULLOCK. D, A. (1975) —The general water circulation of Spencer Gulf, South Australia, in the period February to May. Trans. R Soc. S. Aitst. 99(1), 43-53, 2S February,. 1975. Historical temperature and salinity data are presented together with the three-dimensional velocity structure of the general wa tcr circulation of Spencer Gulf, deduced from the data using a numerical model. In the southern area of the Gulf a cyclonic gyre is found which exchanges Gulf water with the ocean water outside the Gulf. On the western coast of the Gulf, a moderate boundary flow, which is called the "Port Lincoln Boundary Current", is evident at all depths. The water in the northern portion of the Gulf circulates both in the vertical and the horizontal and appears to form an almost closed system separate from the system in the southern region. Introduction Presentation and Interpretation of Cruise Data Ao investigation of the general water circu- Using measurements made by FR.V- Inves- lation in Spencer Gulf, and the region outside tigator (C.S.I.R.O. Aust., 1968c) on 7 May it. is made below from observation of property 1964, a vertical salinity section (Fig. 2a) was distributions and using a numerical model. The drawn for a line across the mouth of Spencer term "general water circulation" is used so that Gulf from Port Lincoln on the western side to tidal currents which basically involve a periodic Corny Point on Yorke Peninsula. The salinity north-south movement of the Gulf's waters are is approximately homogeneously distributed excluded. Tidal currents can be superimposed over depth for any particular watcr column. on the more sustained flows caused by thermo- although around the centre the situation Is haliue (and wind) forcing which arc particu- closer to that of a two-layer system since the larly important because of the part they play salinity is almost constant with depth until in the interchange of water in a substantially about 20 m, at which point a steep salinity enclosed body of water. Thermohaline currents gradient occurs. Below this, the salinity is are brought about by horizontal pressure gra- again constant with depth but greater than dients due to density variations in the Gulf. before. Also,, the salinity increases with dis- which are caused by temperature and salinity tance from west to east, indicating that water differences, produced in part by outside watcr enters the Gulf on the western side and leaves entering Ihe Gull', and in part by variation in via the eastern side after its salinity has in- chc effect of evaporation over the Gulf. creased due Id evaporation and mixing. The All available data for the region, obtained density section (Fig. 2b) has a similar form to On C.S.I.R.O. cruises, arc shown in Fig. 1, The the salinity section, indicating that salinity is cruises took place between 1961 and I9o6 dur- the dominant factor in determining thermo- ing the months indicated in the caption. No haline flow. direct current readings are available, although the most important feature of the hydrology all (Figs some results on surface drift do exist from a of Spencer Gulf r evident from data drift card experiment ( Marshallsay & Radok 10, 11), is lhat the watcr is approximately 1972). homogeneous from top to bottom, and hence The Flinders Institute for Atmospheric and Marine Sciences, Flinders University of South Australia, Bedford Pari:, S. Aust. 5042. Present address: Dept of Civil Engineering. University of Melbourne. Parkvillc, Vic. 3052. 44 D. A. BULLOCK A 8 s it y / B o /$ / ° :i ' "F / ^ /. o A j>~ w / * ^.r ^& .* b* / ' * ^ ^ / tj ^ Fig. 1 — WATER CIRCULATION Oh SPENCER GULF * The distributions shown in Fig. 3 are con- STAT I Oh 1(0. sistent with those in Fig. 2 and give additional '0? |W }Q\ tbt i? 3| information A How up the western shore is "ttt^ r r evident and it can be seen that the salinity in- 7/f creases eastward over the whole southern area of the Gulf indicating that the movement nf water is basically clockwise. It appears that the advective exchange with the water outside the Gulf is limited to the area below a latitude of about 33'45'S. Above this region where the u M Gulf narows. the water appears blocked-in . Along the eastern and central areas of the upper part of the Gulf, the high salinity water seems to flow down the Gulf until it meets the lower salinity water taking part in the advec- tive process described above. The data from the northern pari indicate thai the water moves as a mass in phase with the r tidal currents. At 34 00'S, 137'MO'E, two sfct- lions (76 and 89) were tKeupied, separated by a period of 29 hours during which the depth- average salinity changed from 38.40';,, to HijL 2. Vertical sections across Spencer Gulf near 38.14',,'-,.. At 33'57'S, I37°25'E (stations 77 the month, Marked — " — on Figure 1. and $5) t the aalinitites changed from 38.35^.;,. Jala taken from CSIRO Aust. 1968, samr> to 38.60'r, over a period of 21 hours. Using ling depths indicates by dots, the horizontal salinity gradients at this location. P«i£. ji Location of (Jeeanographic Stations in the South Australian Gulf System until W3. key to Swnhots a H.MA.S. Gascoyne (G2/61) Feb.-March 1961 (CSIRO Au 4(i D. A. BULLOCK KM SD lig. X Horizontal depth-average distributions in Spencer Gulf. Data taken from CSIRO Aust. t%8c. U (a) Salinity {#*}, (b) Temperature ( C), (c) <. ties. The process of evaporation requires the observed value (Fig. 3b). The neglect of advee- extraction of latent beat of vaporisation which tive exchange is justifiable in view of the ob- in turn lowers the temperature and nullifies the served small variation of temperature and effect of direct healing. The following rough closed nature of the circulation in the northern calculation shows that there is an approximate Gulf. Vertical mixing maintains the tempera- balance between the heat input from the atmos- ture difference between the atmosphere and the phere and the heat output due to evaporation. sea surface. From Haney (1972), the heat flux/ unit area Locations of the stations are shown by the is proportional to the difference between the dots in Fig. 3a-c; stations were not occupied apparent air temperature and the sea tempera- in some areas, notably above Hardwick Bay on ture (denoted by T\,* and TH respectively). the eastern side of the Gulf. Nevertheless, the Assuming this flux is all available for evapora- overall pattern of water properties can be in- tion, we obtain ferred reasonably accurately. E-MV-T.) The Numerical Model where h is the evaporative heat flux/unit area, In order to investigate whether the above cir- and approximately * 34 Wm--(°C)~l (Haney culation could be considered as primarily due * other 1972). Taking the values, Ta ~ 2TC (the to thermohaline forcing or whether some mean temperature for early April), and B = cause such as wind is dominant, and discover 2 100 Win , and substituting in equation (I) in greater detail the dynamics of Spencer Gulf, -^ U yields "C 1 C. which is approximately the a numerical model was used to derive theore- H WATER CIRCULATION OF SPFNCFR GULF 47 tically the field of flow from the density field horizontal pressure gradients induced by the 1 shown in Fig. 3c. The model entitled FLOW- density field . This is done by solving the finite DEN computes the currents generated in a sea diilcrcncc representation of the component of density independent of depth* by wind and transport equations written in a form applicable variations in atmospheric pressure, and by the to the terrestrial situation-. 3ct sx - k|ub|u (2) £ » *-*$*•-*¥ 3x b sy - k|ub|v (3) £-*»-*$-*•* 3y b where Ihc following notation is used; u udz, V vdz — H U eastward component of velocity (east = x direction) V northward component of velocity (north = y direction) x & y components of bottom velocity (uB ) z distance measured perpendicular to earth's surface, positive upwards 1 (* — 1 ) a l(K Pin kg!- f Coriolis parameter = Z^cos^ w — angular speed of the earth's rotation. *P = latitude H depth g* ICH *ar. ii.ng. "sr-DH =- specific volume of the standard ocean i gravitational acceleration V sea surface elevation K bottom drag coefficient density of a standard ocean — «i!nll,(j x & y components of surface wind stress The terms in equations (2) and (3) have the the thermohaline terms was 300 mmV-. This following physical meanings: the left-hand can he compared with the typical magnitude of sides arc the linearized accelerations, the first 100 mm-s 2 for the forcing term corresponding terms on the right-hand sides are the Coriolis to wind stress, so that the circulation in Spencer accelerations, the second terms are the hori- Gulf due to thermohaline causes can be ex- zontal pressure gradients due to elevation of pected to be quite significant. The bottom drag the surface, the third terms are the thermo- coefficient K = .0025. haline terms which take into account the hori- FLOWDHN was run with the precalculated zontal pressure gradients due to the inhomo- thermohaline term as the only forcing term and geneity of the horizontal salinity and tempera- with the thermohaline term together with a ture distributions. The final two terms are stress constant wind. After the equivalent of 17 hours terms describing wind and bottom friction res- an equilibrium current was reached (actually pectively. a small oscillation ahout the solution remained. The magnitudes of the thermohaline terms but this was ignored). The initial conditions were evaluated using another computer pro- were that the How velocity was zero everywhere gramme from the observed density distribution and the surface elevation was also zero every- (Fig. 3c) and the depths were interpolated where except at the boundary of the model from those given on a naval hydrographic across the south of the Gulf, which was chart. A typical value of the x component of assumed to have an initial slope similar to that 1 Bye, J. A. T. T 1975. Thallasso-seale nuraericaJ modelling. Computer Report. School of Earth Sciences. The Flinders University of South Australia. (Unpublished.) -Bullock, D, A. 1973. The general circulation uf St. Vincent and Spencer Gulf, Honours Thesis. The Flinders University of South Australia. (Unpublished.) 1 c 48 ft A. BULLOCK developed later itt the Adjacent pan of the To gain a detailed picture of the velocity Gulf. The boundary slope was found to have field in the Gulf, a routine was added to the Male effect on the current distribution and so program which printed out the surface anil bot- the assumption is presumed lo nol have serious tom currents. By involving the shallow water consequences. assumption, i.e. that the Ekman layer {the sur- The I'esuJtaul depth-average current, which face- layer influenced by wind) and the layer ii the transport divided by the depth, was ob- oli bottom frictionaJ influence ovedap fan tained al each point of n grid system formed assumption which is consistent with the waters' over the Gulf, in polar form so that yjfafc magni- vertical homogeneity), it can be shown (Bul- tude and direction of the current could be seen lock 1973—footnOLe 2) thai the curreni iom- immediately. The grid length was 6.4 km. ponents at nny level are given by. (z f'l) H f £y b v(z) = • n " f 3x c Appl> mg c4ua(ions (4) and (5) at z = *i - The magnitude of the velocity vectors shown is and at z " —H gives the surface and bottom typically about n.07 ms 1 (ranging from zero 1 currents. to I? mr ). ^ Since is independent of z it. can be seen r from Figs 6 and 7, which show the surlace Irom equaiions (4) and (5) that the vertical and bottom currents respectively, it can be seen homogeneity fit the Gulf implies that the cur- that at both the surface and bottom there is a vent is a linear function of depth z. Thus a very distinctive flow up the western side of the shear current, exists. The right-hand side of Gulf, which we shall call the "Port Lincoln equation (4) and (5) can be interpreted as fol- Boundary Curreni'* This current is broader in lows. The second terms are symmetric about a e-.tcnt at the surface than at the bottom wheie line Marking mid-depth (Fig. 4) and give, as a it tends to peel away to the east down the slope I unction ol depth, the current which is directly of the boiioin. The Port Lincoln Bounary Cur- a*latcd to Lhe local distribution of density. The rent is also somewhat stronger at tJie surface lirst terms on the right-hand side of the equa- than at the bottom due to the fractional tions, on the other hand, represent the depth- iniluencc of the latter. average current predicted by the model under The direction of the the constraints imposed hy the Boundaries of bottom current can he seen to swing eastward and to the south- the Gulf (the shores and bed). then east over a broad area before flowing south- The vertical components ol velocity were ward to exit down -shovel-shaped also found, using lhe bottom current and. the the topo- graphy through inc Gulf's mouth. The surface eradicate of the bottom topography, current displays a similar pattern; howcvei. since the Port Lincoln Boundary Current 3H 31 is w = U + V wider B ^ B 97 (61 at the surface, particularly near the mouth, the centre of the gyre of the surface Finally, the changes in the sea IcvcT which circulation is located further to the north-casr develop from the initial surface condition of than the centre of the bottom gyre. Evidence of slope obtained, zero are also the eastward movement of water from the Results of the Numerical Miukl boundary current is also provided hy Fig, 3a— where the contours bulge to the east in thi.s ( 1 ) Predicted Velocity Fields due to Thermo- halinc forcing only area. Fig. 5 shows the main features of the pre- The differences between the top and bottom dicted depth-average current. The figure is con- current gyres implies that the surface and bot- sistent with the water ma.ss analysis of the pre- tom currents on the eastern side of the Gulf vious section, i.e. the circulation in the wide, are, tn general, at an angle to each other, un- southern pari of Spencer Gulf consists ol a like the western boundary. This explains whv clockwise there is obvious salinity ff gyre, water entering via the western no or l tongue indi- side and leaving on the eastern side and the cating a return* How down the eastern side of circulation in the northern end appears closed. the Gulf in Fig. 3b and c. WA1ER CIRCULATION OF SPENCER GULF 49 gences of the currents exist. A large proportion of the water in the northern end appears to recirculate in a vertical gyre. Fig. 8, which shows the vertical components of velocity, lends support to this interpretation > by the downwclling at the top of the Gulf near Point / Lowly, which corresponds to an inflexion in the / salinity contours (Fig. 3a). Tn the southern part of the Gulf, a complex system of vertical cells of typical dimension 20 km is predicted. The following changes in surface elevation Fig. 4. Current as a function of depth, with refer- are predicted by the model: in the southern ence to mid-depth. part of the Gulf, a low develops, the sea level on the western side being between 10 and 50 Another important feature to be noted from mm higher than in the centre; the sea level also Figs 6 and 7 is that in the centre of the mouth increases to the north. We can conclude that the surface and bottom currents are in oppo- the circulation in Spencer Gulf which can be site directions so that a large shear is predicted attributed solely to thermohaline causes is of here. The closeness of the density and salinity significant magnitude and forms a major con- sections (Fig. 2b and 2a) To those of a two- tribution to the water exchange processes of the layer system is also explained by this pheno- Gull menon. It would be interesting to make direct current measurements in this area. (2) Velocity field due to Thermohaline and Further north, where the Gulf starts! to Wind Forcing narrow (at about the latitude of Wallaroo), the The numerical experiment was now extended flow pattern is more complicated and conver- to include the effect of wind stress in addition PORTAUGUSTA KM 50 "" P0RT\ ^ \ V .[NCOLN-yr*} ^ Fig. 5. Predicted depth-average thermohaline currents in Spencer Gulf. Fig. 6. Predicted surface thermohaline currents in Spencer Gulf. Fig. 7. Predicted bottom thermohaline currents in Spencer Gulf. 50 O. A. BULLOCK D DOWNWELUNG U UPVVCLUNG KM CI \ \ , IP fee Fig. 8. Predicted regions of up- and down-welling due to thcrmohalinc currents in Spencer Gulf. Fig. 9. Predicted currents in the presence of a 14 ms 1 SW wind- la) Surface, (b) Bottom. to the thermohaline forcing. The resulting sur- compared with the depression of 10 mm in the face and bottom currents predicted by the centre. model for the same density structure as before, combined with a 14 ms-1 south-west wind, arc ( 3 ) Possible reasons for the Persistence of the shown in Fig. 9a and 9b. Density Structure The pattern of circulation is basically the In the numerical model, the thermohalinc same but the velocity of the flow is accentuated, forcing terms remained constant throughout the 1 being typically about 0,16 mr , some velocity period from zero velocity to equilibrium; i,e, 1 vectors having a magnitude over 0.30 ms . wc assume that the observed density field per- Differences to the no-wind case do occur in the sists and is not an instantaneous feature which northern part of the Gulf where the surface subsequently runs down until the Gulf becomes current is predominantly northward in direc- a completely homogeneous body of water in tion, particularly on the eastern side where it which water motion of thcrmohalinc origin was directed southwards previously. Similarly would cease- the bottom current flows north adjacent to the It i.s surmised that the cause of the persist- shores, whereas in the no-wind case ihe bottom ence of the thermohaline circulation is external, current was slight here. It can also be seen that i.e. it i.s due to water from the Southern Ocean with the wind added, the surface and bottom moving up under influences that determine the everywhere. currents are almost the same pattern of flow in this region, perhaps the pre- The model predicts that the surface elevation vailing westerly winds. Fig. 1 2, which shows increases greatly (up to 0,37 rrO with distance the dynamic topography of the ocean south of up the Gulf, i.e. the water is piled up at the Australia (Bye 1971). indicates that the direc- northern end by the wind. The low in the tion of the geostrophic flow in the deep ocean southern half is also accentuated, the level is, in fact, basically perpendicular to the Aus- being raised by up to SO mm around the shores tralian coastline. WAlEK CIRCLXATION OF SPENCER GUEF 5! Fig. 10a and b show temperature and % sections drawn along Q line extending through Investigator Strait to past the end of the Con- tinental Slope, The data were obtained in Fcb,- March 1%1 (C.S.1.R.0 Aust. 1966) and March 1963 (C.S.I.R.O. Ausl. 1967a), One feature which is noticeable is the change in pat- tern from a vertically homogeneous situation in just inside the mouth of Investigator Strait to the more complicated horizontal stratification j in the deeper waters outside the Gulf. The most significant feature shown by these sections, however, is the gradient in salinity, temporal ure and density about Station 122. which was located nonh-east of Cape Borda on the north-west tip of Kangaroo Island. This density gradient implies the existence of tf thcrmohaline current, water to the west of Sta- tion 122 flowing approximately perpendicular to the plane of the section (into the page). This direction correlates very closely with the direc- tion of the thcrmohaline current entering Spencer Gulf which forms the Port Lincoln Boundary Current. Conversely there is also a southward directed flow between Stations 121 and 122 which could have its origins in the outflow from the eastern side of vSpencer Gulf. hi*. K), Vertical sections along Investigator Strau It is possible that upwelling is also occurring to the Continental Slope. Marked near the longitude of Station 122. The peak in on Figure 1, data taken from CSIRO the density lines for Aust. 1966 and iy67a. Sampling depths the surface water under indicated by dots, Station 41 is explained by the fact that this sta- (a) STft 1 1 D N r-l to. 5ft 6i 65 bl 6B 59 70 >918/ MH / <•«* <~-^fl*3 \ 2 = 6 2SO 23 / J 50 1-3£6 t'Sll : V// / i ait tr. 10 [I Fig. 12. Dynamic topography relative to 2000 dh., b south of Australia (taken from Bve 1971). Fig, II. Vertical sections along Spencer Gulf to the Continental Shelf. Marked — . on tributions of water properties in St Vincent Figure 1, data taken from CSIRO Aust. Gulf have been found (Bullock 1974) to be 1967a. Sampling depths indicated by similar to those in Spencer Gulf, it may be dots. ^ expected that the general circulation in St (a) Temperature CO, in the Gulf, it is obviously necessary to con- guidance and advice during this study. Part of duct a series of thorough cruises at various this material was presented at the Second South times throughout the year. In addition, direct Australian Regional Conference on Physical current measurements need to be made to Oceanography, University of Adelaide. 9 check the results from the model. Since the dis- Augusl lf>73. References Rowdhn, K. G. (1965).—Horizontal mixing in the CSIRO Aust. ( 1967ti ).—Oceanngraphicai obser- sea due to a shearing current. J. Fl. Mech. 21, vations in the Indian Ocean in 1963. 83-95. H.M.A.S. Gasvoyne, Cruise G 2/63. CSIRO Bullock. D. A. ( 1974).—Cruise Report 1. (Flin- Aust. Oceanoxr. Cruise Rep. 22. ders Institute for Atmospheric and Marine Sciences, The Flinders University of South CSIRO Aust. (1967b).—Oceanngraphicai obser- Australia.) vations in the Indian Ocean in 1965. Byi:. I. A. T. ( 1971 >.—^Variability south of Aus- H.M.A.S. Goscoyne, Cruise G 5/65. CSIRO tralia. Proc. hul Mat, Set. Sytnp. Institute of Aust. Oceanogr, Cruise Rep, 46, Marine Science. University oj fitgW South Wates, / 6- 17 Aug., 1971. CSIRO Aust. (1967c),—Oceanographical obser- CSIRO Aust. (1966).—Oceanographies! observa- vations in the Pacific and Indian Oceans in tions in the Indian and Pacific Oceans in 1965, H.M.A.S. Guseovne, Cruises G 2/62 1961. H.MA.S. Gascoyne, Cruise G2/6I. and G3/62. CSIRO Aim, Oceanogr Cruise CSIRO Aust. Oceanog. Cruise Rep. 10. Rep, 16. WATER CIRCULATION OF SPENCER GULF S3 CSIRO Aust. (1967d).—Oceanographical obser- CSIRO Aust. (1968f).—Investigations by F.V. vations in the Indian Ocean in 1964. H.M.A.S. EstelJe Star in South Australian and New Gascoyne, Cruise G 2/64. CSIRO At4st. South Wales waters in 1965. CSIRO Aust. Oceanogr. Cruise Rep. 34. Oceanogr. Stn List 71. Aust. (1968a).—Oceanographical obser- CSIRO CSIRO Aust. ( J 968g).—Investigations by F.V. vations in the Indian Ocean in 1965. H.M.A.S. Estelle Star in South and Western Australian Gascoyne, Cruise G 2/65. CSIRO Aust. waters in 1966. CSIRO Aust. Oceanogr. Stn Oceanogr. Cruise Rep. 43. List 76. CSIRO Aust. (1968b).—Investigations by F.R.V. CSIRO Aust. (1968h).—Investigations by F.R.V. investigator on the South Australian tuna Weerutta on the South Australian tuna grounds in 1963. CSIRO Aust. Oceanogr. Sin grounds in 1961, CSIRO Aust. Oceanogr. Stn List 64. List 55. CSIRO Aust. (1968c).—Investigations by F.R.V. CSIRO Aust. (1969).—Investigations by F.R.V. Investigator on the South Australian tuna Derwent Hunter on the eastern Australian grounds in 1964. CSIRO Aust. Oceanogr. Stn tuna grounds in 1961. CSIRO Aust. Oceanogr. List 67. Cruise Rep. 54. CSIRO Aust. (I968d).—Investigations by F.R.V. Investigator on the South Australian tuna Haney, R. L. (1972).—Surface thermal boundary grounds in 1962. CSIRO Aust. Oceanogr. Stn condition for ocean circulation models. /. List 60. Phys. Ocean. 1, 241-248. CSIRO Aust. (1968c).—Investigations by F.R.V. Marshallsay, P. G., & Radok, J. R. M. (1972).— Investigator on the South Australian tuna Drift cards in the Southern and adjacent grounds in 1965. CSIRO Aust. Oceanogr. Stn Oceans. Horace Lamb Centre Res. Report 52. List 70. (The Flinders University of South Australia.) VOL. 99, PART 2 31 MAY, 1975 TRANSACTIONS OF THE ROYAL SOCIETY OF SOUTH AUSTRALIA INCORPORATED CONTENTS Christophel, D. C.y and Blackburn, D. T. A New Procedure for Mounting Cleared Leaves Using Polyester Resin 55 Womersley, H. B. S., and Conway, Elsie Porphyra and Porphyropsis (Rhodo- phyta) in Southern Australia 59 Smith, Meredith J. The Vertebrae of Four Australian Elapid Snakes - - 71 lyier, M. J. The Ontogeny of the Vocal Sac of the Australian Leptodactylid Frog Limnodynastes tasmaniensis ------85 Dodson, J. R. The Pre-Settlement Vegetation of the Mt Gambier Area, South Australia 89 Tyler, M. J., and Martin, A. A. Australian Leptodactylid Frogs of the Cyclorana australis Complex 93 PUBLISHED AND SOLD AT THE SOCIETY'S ROOMS STATE LIBRARY BUILDING NORTH TERRACE, ADELAIDE, S.A. 5000 A NEW PROCEDURE FOR MOUNTING CLEARED LEAVES USING POLYESTER RESIN byD. C. Christophel* and D. T. Blackburn* Summary CHRISTOPHEL, D. C, & BLACKBURN, D. T. (1975).-A new procedure for mounting cleared leaves using polyester resin. Trans. R. Soc. S. Aust. 99(2), 55-58, 31 May, 1975. Palaeobotanical studies of leaf floras require detailed comparisons with extant leaves. Techniques for clearing and mounting extant leaves using either sodium or potassium hydroxide or a lacto-phenol solution as a clearing agent have long been known. This paper presents modification of this technique using an hydroxide dealing agent and mounting in polyester resin. In this procedure leaves are cleared in 15% KOH followed by saturated chloral hydrate. Leaves are then dehydrated, stained with safranin and mounted in Mulford EX-80 polyester resin. Use of this mountant shortens preparation time by several weeks as well as giving superior transparency to specimens. Photographic reproduction using direct printing of specimens placed in an enlarger is also discussed. A NEW PROCEDURE FOR MOUNTING CLEARED LEAVES USING POLYESTER RESIN by D. C. Christophel* and D. T. Blackburn* Summary Christophel, D. C, & Blackburn, D. T. (1975).—A new procedure for mounting cleared leaves using polyester resin. Trans. R. Soc. S. Aust. 99(2), 55-58, 31 May, 1975. Palaeobotanical studies of leaf floras require detailed comparisons with extant leaves. Techniques for clearing and mounting extant leaves using either sodium or potassium hydroxide or a lacto-phenol solution as a clearing agent have long been known. This paper presents modification of this technique using an hydroxide clearing agent and mounting in polyester resin. In this procedure leaves are cleared in 15% KOH followed by saturated chloral hydrate. Leaves are then dehydrated, stained with safranin and mounted in Mulford EX-80 polyester resin. Use of this mountant shortens preparation time by several weeks as well as giving superior transparency to specimens. Photographic reproduction using direct printing of speci- mens placed in an enlarger is also discussed. Introduction which are highly tomentose), and extreme The vast majority of plant macrofossils col- difficulty in conversion to a permanent mount. lected from Tertiary localities is in the form The second category of techniques, which of compressed or impressed leaves. Identifica- may be collectively termed the hydroxide cate- tion of these leaves is made difficult by the gory, was used initially for the study of cauline fact that most extant correlatives are described vascular tissue (Foster 1952). It usually com- and identified on the basis of floral parts. To bines either sodium or potassium hydroxide overcome this problem, some palaeobotanists treatment with further bathing in chloral are now studying extant plants in terms of hydrate or other bleaching agents. This method their "leaf architecture" (Hickey 1973). For is characterized by its relatively long clearing this type of study, detailed observation of the time (four to ten weeks) and frequent loss of higher order veins of angiospermous leaves is cellular and cuticular detail. It has the definite essential, and many workers (Chandrasek- all 1 advantages, however, of working on almost haram 1972 , Christophel 1973-, Hickey 1973) leaf types (given appropriate time) and of have found it advantageous to develop or being readily combined with many permanent modify techniques for clearing these leaves. mounting techniques. Of the many techniques developed to cope with this problem ( Lersten 1 967, Dilcher The technique presented in this paper is a 1974), most can be divided into two broad modification of the hydroxide method. The categories. The first may be termed the lacto- major drawback of the technique in the past phenol category, the most recent refinement of has been the large amount of time needed for which has been presented by Herr (1972). the preparations. For example, to prepare a This technique is characterized by rapid clear- large leaf of Cinnamomitm takes approxi- ing (1-5 days) and by preservation of cellu- mately four weeks in hydroxide, three days in lar and cuticular detail. Its disadvantages in- chloral hydrate, one day for staining, and an clude inapplicability to some leaf forms (e.g. additional three weeks for mounting and dry- those with extremely tough cuticles or those ing. This gives a total time of up to 54 days * Department of Botany, University of Adelaide, Adelaide, S. Aust. 5000. 1 Chandrasekharam, A. (1972).—Megafossil Flora of Genesee, Alberta. Ph.D. thesis, University of Alberta, Canada (unpublished). - Christophel, D. C. (1973).—Fossil floras of the Smoky Tower locality, Alberta. Ph.D. thesis, Uni- versity of Alberta, Canada (unpublished). 56 D. C. CHRISTOPHEL & D. T. BLACKBURN hydrate method. Photographed Fig. I. Strychnos lucida—leaf cleared by potassium hydroxide-chloral with a Leitz Aristophot 4x5 plate camera, x 2.5. enlargcr. 2.5. Fig. 2. Same specimen as in Fig. 1. Direct print using a Durst 1000 x * pro- from the date of collection to the date of drasekharam 1972, Hickey 1973), the here to facili- storage as a permanent mount. cedure is presented in its entirety its usage. The method is equally applicable O'Brien (1974) has pointed out that some tate fresh or dried leaves, with perhaps a slight time may be saved by auloclaving the material to advantage to the dried material, due to chloro- in hydroxide. Subsequent to this we have phyll breakdown. All times given are approxi- found that the drying time of nearly three mate, and will vary with the nature of the weeks for mounting media such as Canada material being cleared. Balsam, Euparal, Permount and Xam can be potassium or sodium cut to less than one day by using polyester 1. Soak leaves in 15% they are decolorised, resin. An added advantage of this procedure hydroxide solution until solution every two days or as is that the monomer of the resin is itself an changing the satisfactory vessels for this effective clearing agent. This not only shortens necessary. The most leaves placed the time needed in clearing, but also produces are glass petri dishes, with the plastic mesh screening a specimen with exceptional clarity and re- in them under discs of decolorisa- producibility for photographic work (Figs 1, to prevent flotation. The time for four days 2). tion varies with the specimen from to four weeks. Strongly resinous or tanniferous Methods leaves will usually take longest. While the basic technique described below 2. Wash the leaves under a gentle stream of hydrate was originally outlined by Foster (1952) and water and transfer to saturated chloral incorporates the modifications of others (Chan- until totally clear. This should take from two MOUNTING CLKARKD LEAVES USING POLYESTER RESIN 57 to seven days, with slight warming hastening 9. Drain the leaf of excess monomer and the procedure on slow material. mount, between two glass slides. Setting should 3. Wash leaves again (as in step two] and take about 2 hours at 3(TC. Higher tempera- transfer lo a dehydration series consisting of tures give shorter setting times but increase 10%, 50% and 90% aqueous ethanol. Finally the risk of the resin parting from the glass. transfer to absolute alcohol. This senes should 10. When the mount is dried, excess resin take less than one day. may be removed with a sharp razor blade or scalpel. Small particles or thin films may be 4. Stain in 1 % Satranin in 1:2 absolute alcohol/ toluol solution. Stain for three to five removed with xylene or acetone. minutes or until all vascular tissue has taken Up colour. Discussion 5. Destain the mesophyll in alcohol/ toluol Having obtained a permanent mount of a Liking care not w destain the ultimate vena- leaf with maximum transparency, it is then Iron. This step determines the final contrast necessary to reproduce the venation pattern between the mesophyll and th^ vascular tissue with as great a degree of contrast as possible and is consequently critical. Experience has for comparison with fossil material. For nor- shown thai mounted leaves having an absolute mal photographs, a Lettz "Artstophof 4 by absorbance oi' 1.2 to 2.5 at 525 am yield the 5 sheet film camera with a Macro-Dia attach- best density for transmitted light photography, ment for transmitted light gave excellent Absorbanees were measured on a Bcekman results (Fig. 1), With a large format camera Acta CIII Spectrophotometer with the leaves such as this it is possible to photograph the at the back of the sample chamber. By com- leaf at nearly life size. The negative can then parison, leaves having an absorhance of 1cm be contact printed and maximum detail js than 1.0 proved loo light for acceptable repro- obtained. duction while those with an absnrhance of Since maximum contrast between Ihe veins more than 2.5 proved too dense. As a com- and ihe mcsophyil tissue is desirable, however. parison, the ab.vorbance of 0.2% saframn the subtle shadings of greys provided by this measured over a 1.0 em light path is t.9, and method proved unnecessary. Galavazi {1963) a one cm cuvette of this solution may be used made brief reference to the direct use ot an as a control while detaining. enlarger for plant materia] cleared in methyl 6. Transfer correctly detained [eaves to benzoate. Ditcher < 1 974 j also successfully acetone for about ten minutes to remove re- used this technique on skeletonized and maining alcohol/ toluol. hydroxide cleared leaves. This technique was 7. Without allowing the leaf surface to dry, also admirably suited tor our cleared leaves. transfer the leaf to uncatalysed resin mono- The mounted leaves fit easily uitu the nega 11 live carrier of a 1000 mer . The monomer acts as a final clearing Durst enlarger and the agent and normally renders leaves quite trans- image can be directly printed on photographic parent in 30 minutes. Should the leaf surface paper. This technique produces a dark-light dry during transfer to the monomer, the result- reversed image (Fig. 2), but as the partem ing opacity may be cleared by leaving the leaf is the important aspect, this reversal b overnight m the monomer. This monomer bath immaterial. is reusable, and consequently this, step is best To achieve maximum contrast, a number carried out away from direct sunlight and of different stains were tried (Morley 1949). under fairly cool conditions to prevent poly- Bismarck Brown gave perhaps the greatest merisation of the resin. contrast, but stained very unevenly. Safranin 8. Prepare ihe final mounting medium by gave very nearly equal contrast, and had the adding about 5% of MEKP catalyst to ftesh advantage of staining much more uniformly. resin monomer. This is best accomplished by Grade four photographic paper was used to mixing in a small glass phiai and Tolling £ive the best contrast in printing. between ihe hands to combine the two com- While the above methods approximately ponents. Such gentle mixing prevents ihe for- halve the time of the total clearing and mount- mation of bubbles in the mountant. ing procedure, it can still take up to three :1 For mounting the specimens, Mutfoid EX-80 poUester resin was usetf, It is KVmlable from MulfotC Hustles Pty Lid.. 25 An*ac Highway. Keswick, S. Alist. 5035. 58 D, C. CHRISTOPHEL & D. T. BLACKBURN weeks for difficult leaves. Work is in progress this fashion. Even with this success, however, using a lacto-phenol clearing method and it is the hydroxide method, though longer, still believed that a procedure has now been found appears to give better results with a wider to permanently mount specimens cleared in spectrum of leaf types. References Dilcher, D, L. (1974).—Approaches to the iden- Hickey, L. J. (1973).-—Classification of the archi- tification of angiosperm leaf remains. Bot. tecture of dicotyledonous leaves. Amer. J. Review 40(1), 1-157. Bot, 60(1), 17-33. Foster, A. S. (1952).—Foliar venation in angio- sperms from an ontogenetic viewpoint. Amer. Lersten. N. R. (1967).—An annotated biblio- J. Bot. 752-766. 39, graphy of botanical clearing methods. Iowa Galavazi, G. (1965).—Clearing and staining State L ScL 41, 481-486. plant material in toto with phloroglucinol- HC1 in methyl benzoate for projection photo- Staining of plant graphy and subsequent serial sectioning. Morley, T. (1949).— materials cleared in Stain Tech. 231-235. Stain Tech. 40(1), 1-5. NaOH. 24(4), Herr, J. M. Jr. (1972).—Applications of a new clearing technique for the investigation of O'Brien, T. P, (1974).—Autoclaving as an aid in vascular plant morphology. /. Elisha Mitchell the clearing of plant specimens. Slain Tech. Sc. Soc. 88(3), 137. 49(3), 175-176. PORPHYRA AND PORPHYROPSIS (RHODOPHYTA) IN SOUTHERN AUSTRALIA byH. B. S. Womersley* and Elsie CoNWAYf Summary WOMERSLEY, H. B. S., & CONWAY, Elsie ( 1975) .-Porphyra and Porphyropsis (Rhodophyta) in southern Australia. Trans. R. Soc. S. Aust. 99(2), 59-70, 31 May, 1975. Two epilithic species of Porphyra, P. columbina and P. lucasii, occur on southern Australian coasts, mainly during winter; their seasonal morphology and distribution are described. One epiphytic species, the little-known P. woolhousiae, is described from additional collections, some of which are reproductive. Porphvrousis minuta sp. nov., epiphytic on Pterocladia capillacea and other cartilaginous red algae, is also described. — PORPHYRA AND PORPHYROPSIS (RHODOPHYTA) IN SOUTHERN AUSTRALIA by H. B. S. Womersley* and Elsie CoNWAYt Summary Womersley, H. B. S., & Conway, Elsie (1975). Porphyra and Porphyropsis (Rhodophyta) in southern Australia. Trans. R. Soc. S. Aust. 99(2), 59-70, 31 May, 1975. Two epilithic species of Porphyra, P. columbina and P. lucasii, occur on southern Aus- tralian coasts, mainly during winter; their seasonal morphology and distribution are described. One epiphytic species, the little-known P. woolhousiae, is described from additional collections, some of which are reproductive. Porphyropsis minuta sp. nov., epiphytic on Pterocladia capillacea and other cartilaginous red algae, is also described. Introduction Genus PORPHYRA C. Agardh Porphyra is common during winter on the Key to Southern Australian Species south-eastern coast of Australia, and a brief 1. Blades delicate, rose-pink, ovate to lanceo- account of P. columbina Montagne and P. late, epiphytic on certain brown (or red) lucasii Levring was given by Levring (1953). algae in the upper sublittoral Both these species occur on rock or firm sub- P. woolhousiae Harvey strates and are monostromatic with a single 1 . Blades lanceolate or ribbon-like, becoming rhodoplast per cell. Levring also described P. umbilicate, grey- red- dark-purple, denticulata Levring from Queensland, recorded to or growing on rock or hard substrates 2 P. naiadum Anderson (now Smithora naiadum 2. Thallus fairly tough, retaining its (Anderson ) Hollenberg ) from New South form Wales, and repeated the original record of when old, shrinking on drying and not P. woolhousiae Harvey from Tasmania. adhering strongly to paper; usually over Since it is largely a winter form, Porphyra 45 jum thick; carposporangialj groups has often been omitted from ecological prominent, scattered, with vegetative cells accounts. P. iimbilicalis (=P. columbina, from among the groups; spermatangia occur- Cribb 62.5 in ADU) was, however, recorded ring irregularly around the margin by Cribb (1954, p. 30) as forming an almost P. columbina Montagne pure association during winter and spring, on 2. Thallus usually delicate, disintegrating fairly rough-water coast at Port Arthur, Tas- when old, adhering closely to paper and mania, and P. columbina was recorded by not markedly shrinking on drying; usually GuiLer (1954, p. 64) from Blackman's Bay 20-30 ^m thick; carposporangial groups ( near Hobart ) , Tasmania. Womersley & usually not prominent; spermatangia Edmonds (1958, p. 247) recorded P. colum- occurring in (usually) narrow, elongate bina and P. iimbilicalis (=P. lucasii) as winter strips, extending inwards from the apical forms, mainly on the south-eastern coast of and side margins of the thallus South Australia, but sporadically further west. P. lucasii Levring This paper deals only with southern Aus- Porphyra woolhousiae Harvey 1863, pi. 265. tralian species of Porphyra and the related genus Porphyropsis. It is hoped that this J. Agardh 1883: 59. De Toni 1897: 15; presentation of the species will stimulate cul- 1924: 12. Guiler 1952: 84. Lucas 1909: tural studies to elucidate further their rela- 20; 1929: 15. tionships. FIGS L 2 * Department of Botany, University of Adelaide, Adelaide, S. Aust. 5000. t Department of Botany, University of British Columbia, Vancouver, Canada. i Although the true nature of the reproductive bodies in Porphyra is not fully understood, the classical terms carposporangia and spermatangia are used here to avoid confusion (Conway 1964). 60 H. B. S. WOMERSLEY & ELSIE CONWAY sio*. vitv 2 cm 200/jm Tig. 1. Porphyra woalhouxiae. A. Blackman's Bay, Tas. tADU. A44234). B, Thallus margin, from the type. C. Thallus with margin (A44234). PORPHYRA AND PORPHYROPSfS IN SOUTHERN AUSTRALIA 61 B ^ * "v " % J !ft. ft. '?£' f4 > y ^ . 50>um « -* D .1 A CD I : 1 * 200>um • * 5 g ' - • - ! : • * * " . f m . . . • <4lgf % »" " ^f ^ » ( lit ' * *^k i " g & * *^ * S ft / - it v£$ ft « > - - **•*.• «v* * ^^ »» „t \ . Fig. 2. Porphyra woolhousiae. A. Thallus with area of darkly-staining reproductive cells. (Blackmail's Bay, Tas.. ADU, A44234). B. Cell arrangement in older part (A44234). C. Thallus margin with liberation of reproductive bodies (St Kilda, S. Aust., ADU, A42722). D. Margin with spermatangia (A42722). ) €1 H. B. S. WOMERSI T-V & ELSIE CONWAY Thai I us (Pig. Ml epiphytic on brown algae the hitter appear to be juvenile All these, or rnrcly on red algae, to 17 cm high and 4 including the type, show similar cell structuie cm broad, from irregularly ovate or cuneate and presence of marginal growth on young elongate when young, to broadly expanded or parts and often also on mature thallL This elongate with curved margins winch are gently marginal meristem is not apparent on the convolute in older plants (e.g. the type), deli- other southern Australian species or on most cate, with one to a few blades arising from a other species placed in Porphvra. A fragment »hi/.oidal holdfast, rose-red to rose-purplish. of the type studied is not fertile, but the other specimens eharaetcristic Monostromatic, ( 1 2-) 16-24 /jn thick, show spennataogial ccUs isodiamcttic to slightly elongate in sec- groups which lend to deliquesce, and from non;»l vie.w, each with a single, laminale or marginal areas the contents of each cell appear stellate rhodoplast, with a pyrenoid. Growth to be liberated as a reproductive body. P t hy a marginal meristem (Figs 1B,C, 2A woolhousjae has n typical Potpfwta base with and by intercalary divisions: margins meris- one or a few fronds attached by rhi/oids from tematic, wilh anticlinal lows of cells; older the basal thallus cells, and in most features parts with cells often in rows but becoming agrees well with Porphvra. If, however, it re irregular, with cells often polygonal or angu- confirmed that ihc carposporangia aie formed lar (Figs IB. 2B), some rounded to ovoid, singly, then relationships wilh the genus Por- often with adjacent cells unequal in size. Mar phyrella Smith & Moilcnbcrg (1943, p. 215) ginal cells isodiametric to slightly elongate, must he considered, though Conway & Wylic < 1972) have shown that the angular, 6—10 ^m across; cells [ft older parts New Zealand ito-diamctric to twice as long as broad, 10-16 Porphyra sitbtuntetis does not form packet's of carpusporanjjia, i' -20) ft.m across. Reproductive bodies ( Fig. 1A ,C ) formed While it is desirable that mature plants on marginal areas, apparently singly, in spherical Matrocystix, corresponding to the type, be to slightly ovoid. 12-17 across, usually ^m studied in detail, and their reproduction fol- stellate rhodopla.st, with a lowed in cultuic, the other records arc Spermatangia (Fig. 2D) forming well sullicienlly similar to be placed under/*, woof- defined, irregular, patches near ihc thallus houxiac with some confidence. Most are epi- margins, the whole patch when matuie tend- phytic on brown algae, cither on marginal ing to deliquesce with numerous spermatangia spines or on spinous branchlets, and this hahi- irregularly arranged; spermatangia originally lal may be characteristic for the species. formed in packets of (16-) 32-64, in two P. w(fo(hoits{ae has been recorded from New tiers, individually 2-4 uTO across. Zealand (Lcvring 1955 T p. 412), followed by Type locality; Tasmania. D. J. Chapman fl962, p. 129), V J, Chapman Type: Herb. Harvey, TCD (presented by (1969, p. 20) and Adams (3972, p. 67). Miss Wool house of Sheffield). However Adams, following notes of E. Con- Distribution: As well as the type, this species way in CHR* expresses doubt as to whether is known from St Kilda, S. Aust., on Go;ar- the New Zealand records arc not P. colnoj- tina{'!) on Poxidftniu, I] m below Jow tide bina in an abnormal habitat. level (S. Lewis. 23.Viii.I972; ADU. A42722): The New Zealand specimens growing on Mallaeoota, Vic, on Seytothammts auxtralis Macrocyslis (and possibly Etklonia and Scyto- Pucker and King, 1 5.ii. 1970; ( MELU. thutnnus) need further comparison with the I>eal l. Bass Strait, Pfrithalia 20652); T on Australian plants referred to P. noolhottsUic. cmuhto (Kiw>, 2Xxi,1969; MELU. 21357); Ones from Hokio Beach, l.evin. N.Z., on Curtis I.. Ba&s Strait, on P. cutulata {King, Macrocystis i Moore, 1 7 .xi. 1 946; CHR. 8.ii,l97I; MELU, 21358); and from Black- 55566). determined by Levring j |y55, p. 412) (S. of Hobart), Tas., drift (Tyler. man's Bay as /'. wnofhnusiac (accompanied by notes of Oct (V) 1^73; ADU, A44234), E, Conway (1971) that they might be young The type of P. woolhousiae is a well plants of P. cohtmbitm), agree fairly well with developed specimen to 15 cm high, as in the P woolhousme as now known from the Aus- Blackmail's Bay, TasM specimen, whereas St tralian plants. They are similar in form, in Kilda specimens are 2-4 .-m huih, the Malla- thickness and cell arrangement, and in having coota specimens are only 1-2 cm high and a liierisleinatie margin, but their reproduction Ihose from Bass Strait less than I cm high. is inadequately known. PORPHYRA AND PORPHYROPS/S IN SOUTHERN AUSTRALIA 63 Further comparisons between Australian Distribution: From Elliston, S. Aust. to Syd- and New Zealand specimens epiphytic on Mac- ney, N.S.W. and around Tasmania; New Zea- rocystis are clearly needed. land, Auckland Islands and other sub-antarc- Skottsberg (1923, p. 4) recorded P. wool- tic islands. housiae from the Falkland Islands, also on P. denticuiata Levring from southern spines of Macrocystis, his and account shows Queensland is probably not distinct from P. similarities with the above description; details columbina and represents the range extreme of are not adequate for a full comparison. Hamel P. columbina. (1928, p. 55) recorded it from Kerguelen I., P. columbina is the commonest intcrtidal and Pujals (1963, p. 8) gives records from species of Porphyra in New Zealand and in South America. eastern southern Australia, where it occurs at a Porphyra columbina Montagne 1 842: 1 4; lower to mid (sometimes upper) eulittoral 1845: 33, pi. 9, rig. 2. J. Agardh 1883: level on rough-water coasts. In Australia it is 70, pi. II, figs 65-66. V. J. Chapman: essentially a winter form, persisting as late as 1969: 22. Dawson, Acleto and Foldvic December (rarely to February in Bass Strait) 1964: 32, pi. 615. Guiler 1952: 84. in cooler summers and reappearing in about Hamel 1928: 51. Kuetzing 1849: 693; May. 1869: 29, pi. 80e,f. Laing 1928: 39, figs 1-7. Levring 1953: 464, figs 2-4; 1955: Porphyra lucasii Levring 1953: 469, figs 6H-L, 410; 1960: 29. Pujals 1963: 8. 7. Wildemania columbina (Mont.) De Toni 1897: P. umbilicalis sensu Guiler 1952: 84. Womers- 22. ley 1950: 162. P. umbilicalis sensu Cribb 1954: 30. FIGS 5, 6 FIGS 3, 4 Thallus usually fairly delicate, varying from Thallus fairly tough, varying from ribbon- lanceolate or ribbon-like (Fig. 5^4) to broadly like (Fig. 3/4), often with undulate margins, ovate or cordate (Fig. 55), simple or irregu- to broader forms, sometimes furcate, and larly laciniate or basally branched, sometimes usually umbilicate (by loss of upper parts and becoming umbilicate from basal proliferation, basal proliferation) (Fig. 35) on rough-water usually adhering to paper and not shrinking coasts. Thallus often markedly shrinking on on drying; to 10 cm long and 15 cm broad, drying and usually not adhering closely to margin smooth to undulate. paper. Variable in size and width, reaching 40 Monostromatic, 20-30 cm long and 30 cm across. Colour varying ^m thick, cells with single from grey-red to red-purple. a axile, pyrenoid-bearing, rhodoplast. Cells 8-1 5^on across in surface view (Fig. Monostromatic, 35-50 uffl thick, cells iso- 6A , mostly irregularly arranged, separated diametric in section and with a single axile, ) by gelatinous matrix pyrenoid-bearing, rhodoplast. Cells 10-15 una i-1 times as wide as cells. across in surface view (Fig. AA), isodiametric Carposporangial groups (Fig. 6C) usually to slightly elongate, more or less in rows but not prominent, covering areas around the becoming irregularly arranged, separated by a spermatangial strips toward the margin of the gelatinous matrix |-1 times as wide as cells. thallus, without conspicuous intermingled vege- Carposporangial groups prominent (Fig. tative cells, about 8 carposporangia per group. 4C), of varying shades of red, forming irregu- Carposporangial groups often apparently lar marginal areas with vegetative cells inter- developing later than spermatangial strips, thus mingled, (8-) 32-64 carposporangia per giving an impression of dioecism. group, often giving an irregularly granular Spermatangial groups (Fig. 65, C) occur- ("spotty") red appearance to reproductive ring as well-marked elongate strips (Fig. 5B) areas. extending in from the apical regions and side Spermatangial groups (Fig. 4fi, C) scattered margins of the thallus; strips from a few mm among the carposporangial groups and in older to 2 cm long, 1-5 mm broad. Margin becom- plants occupying the marginal part of the ing "jfrisged" following shedding of sperma- thallus; not occurring in elongate strips as in tangial strips. P. lucasii. Type locality: Bunbury, W. Aust. Type locality: Auckland 1. (D'Urvilfe). Type: Herb. Levring, Goteburg. (Isotype in Type: PC (Herb. Montagne). ADU, A42700). 64 H. B. S. WOMERSLEY & ELSIE CONWAY \ ,t x '«> i C, Pi H.lt l.u*—*M~) ftnyVyffc. ,jU-,1, «* i>m Uy La-»*1, S ft«u n,* < J/Wrwl 7 ! t*?J toll u* - (,«!»•* "'"1 Fig. 3. Porphyra columbina. A. Ribbon-like forms, early winter. Little Dip, Robe, S. Aust. {W&mers- lev, 15. v. 1972; ADU, A42203). B. More umbilicate forms from spring. Cape Lannes, S. Aust. iiVomersley, 7.x. 1972; ADU, A42768). PORPHYRA AND PORPHYROPSIS IN SOUTHERN AUSTRALIA 65 lOOyum Fig. 4. Porphyra columbina. A. Vegetative cells. Nora Creina. S. Aust. KWomersley, 3.ix.l97I; ADU, A39559). B. Spermatangial marginal area (A39559). C. Area with carposporangial groups and young spermatangia (A39559). ' 66 H. B. S. WOMERSLEY & ELSIE CONWAY A J- / **%* K 1 23*9«7»9W * J -a B : y Robe, S.dwlr. 1 ' • ' : 1 . ' ' 1 1 1 1 m 1 1 1 1 m , h 1 1 1 1 1 1 , , ' ' ' : , 1 . , 1 1 1 1 , i , t < 1 , 1 1 1 . 1 1 1 1 1 1 1 1 : , m j | I | 1 1 1 1 r 1 V**j MCA 4«AtirwA t « "123456789 10 8 x mz 0»H "»-j>«a-. h. 8s.*d«?*.*-l*« Fig. 5. Porphyra lucasii. A. Ribbon-like forms. Port Stanvac, S. Aust. {Lewis, l.ix.1972; ADU, A42637). B. Broadly ovate forms showing spermatangial strips. Robe, S. Aust., in bay (Womersley, 8.X.1972; ADU, A42764). PORPHYRA AND PORPHYROPS/S IN SOUTHERN AUSTRALIA 67 100/jm Fig. 6. Porphyra lucasii. A. Vegetative cells (ADU, A42764). fl. Edge of a spermatangial strip with vegetative and carposporangial cells on the right (A42764). C. Part of a spermatangial strip releasing spermatia above and with carposporangial groups below (A42764). 68 II D. S. WOMERSLEY & ELSIE CONWAY Distribution; From Cotiesloe, W. Aust. to ^ra Jatis et aspectu fromali globosis vel Western Port Bay, Vic. and the north and east aliquantum elongatis, in sectione transversal! coasts of Tasmania, parum ovoideis. Rhodoplastus ut videtur, cellu- P. lucasii is the common Porphyta of calm Jam complet, pyrenoide absenti. to moderately rough waters, being replaced by Monosporangia subglobosa vel ovoidea. 5-7 P. eolutnbina where surf action is strong. It ^m lata, in cellulis prope marginem mem is essentially a winter plant of the mid to upper branarum formata. culittoral. with old plants being found in Octo- FIG 7 ber, P lucasii is found within calmer bays Type locality: Pearson I., S. Auil.. on Ptero- such as Port Phillip Bay and Western Port, cladia capillacea. upper sub-littoral (Spevht, Vic. whereas P. cotumbina occurs on rough- ? I7.U.1960). water coasts Type: ADU, A24525. Distribution; From Garden I., W. Aust. Genus PORPHYROPSiS Rosenvinge around southern Australia to Bateman's Bay PorphyropsLs minuta sp. nov. N.S.W., epiphytic on Pierocladia capillacea in Thallus epiphytic on certain Rhodophyta upper sublittoral (and pools) on rough-water with a firm surface, developing from a sub- coasts, and occasionally on Plocamium attxus- parenchymatous holdfast to form a hollow, tMftij P. tnertensii and Delisea pulchra in simi- sub-iphcncal to ovoid Madder (Fig. 1A ), later lar habitats. opening above to form irregular, monostro- P. minuta agrees well with P. coccinea in matic membranes often with convolute mar- the holdfast, form and development of the gins. Color greenish-brown to purplish, not bladder, and in reproducing apparently only jose-rcd. by monosporangia. The cctls of P. minuta are Bladders up to 1 (-2) mm across, torn similar in size to those in P, coccinea but art membranes to 5 (-8) mm high and across. arranged in distinct rows more or less at right Cell divisions intercalary. Holdfast 50-11)0 angles, in contrast to both P. coccinea from (-200) fjjn across, formed of irregularly Europe (Rosenvinge 1909, p. 69, fig. 9) and arranged cells without rhizoidal extensions, P. coccinea var. dawsonU Hollenberg & Ceils of bladder (Fig. IB, C) often paired or Abbott (196S, p. 1239, % 5a-c) from Cali- in fours following division, and lying in rows fornia, where the cells are more irregularly more or Jess at tight angles, sometimes becom- arranged but are grouped into elongate, some- ing irregularly arranged; membrane about 10 what lenticular patches. The life-history' 4tWj f.tTY\ thick, cells 3-5 ju.m across and rounded to relationships of the latter taxon have been somewhat elongate in surface view, slightly discussed by Murray, Dixon & Scott (1972), ovate in sectional view. Rhodoplast apparently and it is desirable that the Australian plant filling the cell, without a pyrenoid. should be studied in culture, Monosporangia (Fig- IB) formed from the A further difference is that in P. minuta the whole contents of cells near the margin of the whole contents of cells near the margins are membranes, subspherical to ovoid, 5-7 ^m liberated as monospores, whereas in P. across. coccinea the monospores are cut of! from the Thallus in Rhodophytis epiphyticus, solido parent cell by a curved wall, a residual cell superficie. ex subparenehvmato base ortus, remaining when the monospore is liberated. primum vesicam cavam, subglobosam vel Also, in P. coccinea some holdfast cells form ovoideam formans deinde supcrne membranas rhizoidal extensions whereas this has not been nrcgulares monostromaticasque saepe margine observed in P. minuta. The colour of P. minuta convoluto producens. Color brunneo-viridis is always a greenish-brown-purple, never rose- vel purpureas, sed nunquam carneus. red as in P. coccinea Vesicae ad 1 (-2) mm latae, membranae The only other southern hemisphere record laccratac ad 5 (-8) mm altae et latae. Divi- of Porphyropsu is by Adams (1972, p. 63) sura cellularum intercalans. Basis ad 50—100 who reported P. coccinea var. dawsonii from (-200) uffl lata. e\ cellulis sine projecturis New Zealand This taxon (e.g. CHR 248053 rhizoideorum irregulariter composite fonnata from Kaikoura, N.Z.; Parsons, 13.xLl973) has est. Cellulae vesicae saepe binae vel quaternae numerous ligulate fronds from a clumped base, post divisionem, seriatim plus minusve rec- each with descending rhrzoids. It is not a Por- tangulatae, tnterdum irregulariter compositae: phyropsis* but more closely related trj Porphyra merabrana circa 10 ^rn crassa. cellulis ad 3—5 woolhoustiu*. PORPHYRA AND PORPHYROPSIS IN SOUTHERN AUSTRALIA 69 r « &gm m 5 * * 5 S> TT-* a * • • • m J" • m m % m § 1 , w i 4p #. | m m % ^ 11 # " ^ 'I MM *ZT ^ & A 50yjiinn Fig. 7. Porphyropsis minuta. A. Small plants, on Pterocladia. B. Older membrane, with some cells re- leasing monosporangia. C. Cell arrangement. (All from the type, ADU, A42525.) — ) 70 H. B. S. WOMHRSLEY & ELSIE CONWAY Acknowledgments versity of Tasmania) who recollected P. wool- We are grateful to Mrs S. C. Ducker (Uni- housiue. The first author acknowledges pro- versity of Mclhoume) who rnade available vision of technical assistance by the Australian collections for study, and 10 Dr P. Tyler (Uni- Research Grants Committee. References Adams, Nancy M. (.1972). —The murine algae of Laino, R. M- (1928).—New Zealand Bangiales the Wellington area. A list of species. Rec. I Bang'ta, Porphyra, Erythrotricfua and ( ? Dom. Mus. 8(5), 43-98. Lrxtiirocladia). I rat is N^Z.. Inst 59, 33-59, Plates 1-15. Agardh, J. G. U883).—Till algerncs systcmaiik. (1953). Nya bidrag. Vt Ulvacene. Acta Univ. land LdVKiNG, T. — The marine algae of Aus- 19(2), 1-182, Plates 1-4. tralia. I. Rhodophyta Goniotrichales. Ban- giales and Nemalionales. Arkiv. for Botanik. Chapman, D. J (1962).— check list and key A Ser 2, 2(61, 457-530. to the Rhodophyceae of New Zealand Sec- Levrjng, T. ( 1955). —Contributions ft) the marine tion A: BangioiUeae. Trans, R. Soc. N.Z. algae of New Zealand. I. Rhodophyta: Bof, L 127-137. Goniotrichales, Bangiales, Nemalionales and Chapman, V. J. (1969). The marine algae of — Bonnemaisoniales. Arkiv lor Bot. Ser. 2. New Zealand. Part lilt Rhodophyceae. 1. 3(11 ), 407-432. Bangiophycidac and FlorideOphycidae (Nema- Lcvring, T. (I960). Contributions to the marine lionales, Bonnemaisoniales, Gelidiales). — algal flora of Chile. Lands Univ. Arssk. I Cramer: Germany.) N.F. Avd. 2. 56(10). 1-85. ( . onway, Elsie ( 1964).— Autecological studies of Lucas, A. H. S. t.J 909). —Revised list of the the eenus Porphvra: I. The species in Britain. Fucoidcac and Florideae of Australia. Proc. Brit. Phycol. Bull. 2(5), 342-348. Linn. Soc. N.SW, 34, 9-60. Conway, Elsie, & WYUBj Ann P. ( 1972).—Spore Lucas. A. H. S. (1929).—The marine algae of organisation and reproductive modes in two Tasmania. Pap. Proc. R. Soc. Tasm. 1928. species of Porphvra from New Zealand. Proc. 6-27. — VJt Int. Seaweed Symp., pp. TO5-107. (Univ. MoNVAUNk, C. (1842). "Prodromus Generum Tokyo Press; Tokyo.) Specierumque Phycearum Novarum In ltinere and Pol urn Antareiieum." (Paris.) Ckibu, A. B. (1954).—The algal vegetation of — Port Arthur, Tasmania. Pap. Proc. R. Soc. Montaone., C. (1845). "Voyage au Pole Sud les Corvettes )' Astrolabe Tasm. 88, 1-44, Plates 1-10 et dans 1'Oceanie sur et la Zelce." I. Botaniqu". (Paris.) Dawson, E. Y., Acitro. C, & Foi.dvic, N- Murray, S. N., Dixon, P. S„ & Scon. J. L. J. 1 1964 —The Seaweeds of Peru. Nova tled- J 972) .—The life history of Porphyrop\is wit>ia 13, J- i 11, Plates 1-81. ctHnnta var. dawsonii in culture. Br. phvcol. ( -"Sylloge Algarum Da Tom, G. B. 1897)— /. 7, 323-333. omnium hucusque CognitariunV\ Vol. 4. Pujals, Carmen (1963).—Catalogo de Rhodo- 1-388. Florideae. Sect. !, pp. (Padua.) phyta citadas para la Argentina. Revta Mus. Hi: Tom, G. B. (1924).—"Sylloge Algarum argent. Cienc. nat. Bernardino Rivadavia, Bot omnium hucusque Cognitarium". Vol. 6. 3(1). 1-139. Florideae, (Padua.) RoshNviNot, L. K. (1909).—The marine algae CituLER, E. R. 11952).—The marine algae of of Denmark. Part 1. Introduction. Rhodo- Tasmania. Check list with localities. Pap. phyceae. 1. (Bangiales and Nemalionales). Proc. R. Soc. Tasm. 86. 71-106. SW/« Danske Videnskah. Sclskab. Biol. Skr.. 7 Raekkf, AftL 7, M51, Plates 1, 2, 2 maps. (.hiikr, E. R. ( 1954).—The recolonization of SKorrsatRO. C. ( 1923). Botanische Ergebnisse rock surfaces and the problem of succession. — der schwedischen Expedition nach Patagonien Pap. Proc. R. Soc, Tasm. 88, 49-66. PJates und dem Feuerlande. 1907-1909. IX. Marine 1-6. algae. 2. Rhodophyceae. K. svenska Vetensk- Hamel. G. (19281.—Sur quelques Porphyra des Akud. Handi. 1-70. mers australes. Ann. Crvpt. E.xot. 1(1), 51- 63(8), Smith, G, M., & Holl£NREpg. G. J. (1943). — 57. On some Rhodophyceae from the Monterey HarvlV. W. H. (1863).—"Phycologia Auslralica". Peninsula. California. Amcr. /. Bot. 30(3). Vol. 5, Plates 24I-3(K), synop, 1-73. pp. 211-222. HoixtMBtun, G. Abbott, I. J.| & A. (1968). WoMBRSitv. IT B S (1950) —The marine algae New species of marine algae from Califor- of Kangaroo Island. III. List of Species J. nia. Can. J. Bot. 46, 1235-1251. — Trans. R. Soc. S. Aast. 73(2), 137-197. ). KutrziNU, F, T. j 1849 "Species Algarum". WoMKKStnv, H B. S, & Edmonds. S. J. (1958). ( Leipzig.) —A general account of the intertidal ecology Kuetzing, F. T. ( 1 869 ) .—"Tabulae Phyco- of South Australian coasts. Aust. J. mar. logicat". Vol, 19. (Nordhausen.) Fresh*. Res. 1(2). 217-260. Plates 1-12. THE VERTEBRAE OF FOUR AUSTRALIAN ELAPID SNAKES (SQUAMATA: ELAPIDAE) by Meredith J. Smith* Summary SMITH, MEREDITH J. (1975).-The vertebrae of four Australian elapid snakes (Squamata: Elapidae). Trans. R. Soc. & AusL 99(2), 71-84, 31 May, 1975. In vertebral morphology, the elapid species Pseudechis porphyriacus, Austrelaps superba y Notechis scutatus and Pseudonaja nuchalis conform with general descriptions and closely resemble each other. Features not previously noted are that epizygapophysial spines appeal* on the first 8 to 10 vertebrae, and that bilaterally on every precloacal vertebra a foramen opens through the accessory process near the anterolateral edge of the prezygapophysial facet. No morphometric feature was found to completely separate any two genera, i.e. there was some overlap in the values of all the characters (ratios) studied. However, in general P. porphyriacus vertebrae are distinguished by their relatively long accessory processes, A. superba by their short accessory processes and lesser width across postzygapophyses, and N. scutatus by their greater width across postzygapophyses and shorter neural spines. P. nuchalis vertebrae have strong subcentral ridges THE VERTEBRAE OF FOUR AUSTRALIAN ELAPID SNAKES (SQUAMATA: ELAPIDAE) by Meredith J. Smith* Summary Smith, Mkkedith I. ( I97i)»—Ifia vertebrae of four Australian elapid snakes (Squamata: Elapidae). Tarns, Soc. 1975. H. $ Aush 99(2) ? 71-84, 31 May, In vertebral morphology, fhfl efapid species Pseudediis porphyriacus, Austreiaps super ba, Soiethis scutams and P&cudonuja nuchuiis conform with general descriptions and closely resemble each other. Features not previously noted are that epizygapophysial spines appear on the first 8 to 10 vertebrae, and that bilaterally on every precloacal vertebra a foramen opens Through the accessory process near the anterolateral edge of the prezygapophysial facet. No morphoinclric feature was found to completely separate any two genera, i.e. there was some overlap in the values of all the characters (ratios) studied. However, in general P> par' phyruicuM vertebrae are distinguished by their relatively long accessory processes, A. superha by their short accessory processes and lesser width across postzygapophyses t and N, scutuius by their greater width across postzygapophyscs and shorter neural spines. P. nuchaUs vertebrae have strong subcemr&l ridges. Fossil vertebrae from a Pleistocene deposit differ from P. porphyiacus, A. superba and N. sc.utatus but resemble Pseudonaja in most features, The fossils differ from modern P. nuchalis chiefly in having a thicker zygosphene and relatively wider postzygapophyses. As these fire features which develop with increasing size of vertebrae, the fossil vertebrae are assigned to the genus Pseudonaja, Introduction the whip snakes by Worrell (1963)|. Hopio- Although snake vertebrae have been cephulus, and Oxyuranus been examined as recorded from Australian Pleistocene deposits iheir range is northern and eastern Australia (l.ydekkcr 1888, Merrilees 1968), the only (Worrell 1963). Specimens have been studied dustretaps, species that has been identified is the Carpet from the remaining four genera, Snake, Morelia spilotes variegata Gray (~- Notechis, Pseudechis and Pseudonaja, which Pyihon variegatux) from Marmor Quarry. arc each represented in southern Australia by Queensland (Longman 1925). No detailed one or more common species. studies have been published of Australian The exact column position of an isolated snake vertebrae and the diagnostic characters vertebra is impossible to determine (Johnson have not been established. need to deter- The 1 955 ) and in elaptds. with well-developed mine the reptile fauna in a Pleistocene cave hypapophyses on all pre-cloacal vertebrae, deposit stimulated the present study. division of the pre-cloacal column into regions The sixty-odd Australian species of clapids is virtually impossible. Nevertheless vertebral have been arranged in 29 genera (Worrell shape changes along the column, and to iden- 1963) or fewer (McDowell 1967, 1970), but tify single vertebrae it is essential to know the only seven genera contain species with a range of variation within individuals and recorded maximum length of over 0.9 ro. As species. Auffenbcrg (1963) based his descrip- centrum lengths of some of the Pleistocene tions and diagnoses on middle pre-caudal verte- vertebrae suggested that the specimens from brae (determined as such by the relative size which they were derived must have exceeded of the neural canal) and avoided considering 1.5 m, the 22 genera of smaller species (less intracolumnar variation. Although Johnson than 0.9 m maximum length) have not been (1955) measured 10 precloacal vertebrae at considered. Nor have Demansia [restricted to regular intervals along the column, he assumed Department of Zoology, University of Adelaide, Adelaide, S- Aust 5000. . 72 MEREDITH I. fiMJTH equnl variances i, between specimens) of ratios at least several individuals. Visual examination of these measurements and in his comparisons of the fossils revealed that 454 of then) were he used only the mean for each specimen, similar to each other in shape, anil dixlincdy Here, enure vertebral columns have been different from the rest, which were of seveial examined in an attempt to assess the morpho- kind.s, Ihe latter, heterogeneous group will be logical and murphornetrlc variation which discussed in a later paper. Of the larger group occurs within individuals and species, and to of 454 ihe HO most complete were examined find unique specific characters. As most elapids in detail and measured in the same way as the have over 200 vertebrae, the available samples modern vertebrae are large and the time required for detailed Because reputes grow ihroughuut their life. examination of one individual snake precludes absolute dimensions of the vertebrae are ol sampling many individuals. little use in comparing individuals or species. Because taxonomy of species of Fseudtmwt Ratios between dimensions have been calcu- and Austrclaps is confused (Rawlmson 1969, lated, the denominator being vertebra length Storr 1964) comparisons arc made ai the in most comparisons. Mean values of ratios I'encric level. rather than species have been prepared independent ly for each individual snake, and arc given as mean, T. Materials and Methods — standard error, followed in brackets by the A total of 2, 1 23 vertebnie from nine Recent number of vertebrae measured. The non-inde- specimens (Table I } and 556 Pleistocene ver- pendence of measurements of different verte- it:hr.:e were examined. Of the nine modern brne of the one individual precludes statistical specimens, four specimens of Pxeudcchis comparison of these samples (Siegel 1956) pnrphyrior\ti, one PseiuhnnU* nachulis Kroni a preliminary study of numerous (RI4064) and one Notechix $cntalus characters of snake vertebrae, characters were (R14059) were collected near Armidale. New selected that vary between species within the South Wales: the. one Austrclaps superba was tamily Elapidae, exhibit low intracolutnnai collected at Umidla. South Australia. The variation and arc well preserved in fossils. To localities of One P. vnchnfix (R 1 4065) and establish variation throughout the column, one jV. stututus (R 14058) arc unknown. every fifth vertebra was measured in two speci- The common brown snake. PscutJothij^ tex- mens of each of P. porphyriucits, P. nuchalis tile (Dumcril & Bibron), has beco described and N. sctttatttx, and one A. xttpertuj. as a separate species Jrorn the western brown Descriptive techniques and terminology snake. Psendonaja mtcholis Gunthcr. but It is follow Aullcnherg (1961). Measurements were not yet clear (Stnrr 1964) whether they are dis- made to 0.1 mm with dial Calipers, as in Fie. tinct or are merely races of a single species, P. I. tcxtilh. One clearcut diagnostic feature is ihe pr-po Ijcngth between zygapophyses-distance shape of the nasal bones, which are anteriorly between most anterior point of prczy- COnCHtVO on the lateral margins in P. ttuchulis gapohysis to most posterior point of but anteriorly convex in F. te.Kiilis (Worrell postzygapophysis. |9o_Vt. In R 1 4064 and R14065 the nasals arc ap-ap Width across accessory processes-dis- anteriorly concave and hence are attributable tance between outermost tips of acces- to P nuchaUs. sory processes, The cleaned skeletons were dried and com- po-po Width across postzygapophyscs-dis- pletely disarticulated for study. The specimens tance between outeimost points of the arc now lodged in the South Australian pnst/ygapophyxial facets, Museum, with register numbers as above. prl Length of prczygapophysis-the longest The Pleistocene vertebrae were excavated diameter of the pre/.ygapophysial facet from an extensive bone deposit in Victoria feven though this was almost perpen- Cave. Naracoorte. South Australia. The age dicular to the long axis of the cen- of this deposit is unknown, but the abundance trum). of extinct marsupials and the absence of pr Width of prezygapophysis-tbe maxi- remains of aboriginal man suggest that Ibc mum diameter at right angles to die deposit accumulated during the Pleistocene but length of the prezygapophysix (the was sealed during Recent time. As the verte- pre/ygapophysial width being almost brae were collected singly in many locations parallel to W*e lOrtg axis of the cen- in the bone deposit, they probably represent trum y VERTEBRAE OF FOUR E1.APID SNAKES 73 nsl Minimum length of neural spine-this usually occurred about halfway up the spine, as the dorsal edge overhangs posteriorly and/or anteriorly. zw Width of zygosphene-the maximum width of the tenon. cw Width of condyle-the maximum dia- meter in the transverse plane. hyp Length of hypapophysis-the vertical distance from the lower edge of the condyle to the tip of the hypapophysis. Results The vertebrae conform with the general des- criptions of AulTenberg (1963), HorTstetter & Gasc (1969) and Johnson (1956). po-po Pseudcchis porphyriacus (Shaw) The number of prccloacal vertebrae (Table 1) is consistent with the number of ventral scales (184, according to Worrell 1963). In every precloacal vertebra, the width across postzygapophyses exceeds the length between zygapophyses (Fig. 2). The hypapo- physis arises near the lip of the cotyle, extends as a low lamella for about two-thirds the length of the centrum and then deepens sharply before tapering to a sharp point which does not much exceed the posterior surface of the condyle in any but the most anterior vertebrae. The hypapophyscs of the anterior vertebrae are very long; they decrease in length fairly uniformly along the column (Fig. 2). The subcentral ridges are low and rounded. The dorsal articular facet of the paradiapopbysis projects as a little round dome; the lower facet is saddleshaped. The prominent parapophysial processes are rounded anteriorly. They do not extend closer to the midline than the most lateral lip of the cotyle, Interzygapophysial ridges are faintly distinguishable. The neural spine is a low, laterally-compressed blade, its dorsal edge parallel with the long axis of the vertebra, its anterior edge almost vertical and its posterior edge overhanging. The minimum length of the neural spine between the zygapo- Fik. J. Dorsal (D). ventral (V) and lateral (L) is about half the length views of an elapid vertebra showing where physes [x 0.53 ± .0056 (35); x 0.55 ± .0049 measurements were taken. (36)] (Fig. 3). The neural arch is slightly poi Length of postzygapophysis-the wider than high. The neural arches of most of longest diameter of the poslzygapo- the vertebrae do not extend backwards to form physial facet (even though this was epizygapophysial spines, but such spines are almost perpendicular to the long axis well developed on the first five vertebrae and of the centrum). are distinguishable on the sixth to tenth. The po Width of postzygapophysis-the maxi- cotyles and condyles are nearly round, but arc mum diameter at right angles to the slightly flattened dorsoventrally. The width of length. the condyle is about one third of the length 74 MEREDITH J. SMITH TABLE 1 Total length, snout-to-vent length and number of vertebrae of specimens of four species of elapids SAM Total Snout-vent Number o / vertebrae register length length Pre- Post- Species number Sex (mm) (mm) cloacal* Cloacal cloaca I Total* Pseudechis R14060 no data 1220 no data 181 8 >42 >231 porphyriacus R14061 no data 1375 no data 182 & 52 240 R14062 no data 1120 no data 182 5 >30 >217 R14063 no data 1070 890 177 5 48 231 Austrelaps R14066 2 515 433 143 4 42 isy superba Notechis R14058 c* 930 775 174 s S8 237 scuta tits R 14059 no data no data 800 178 5 52 235 Pseudonaja R14064 3 1555 1300 204 5 60 ?70 nuchalis R14065 no data 1100 910 206 5 62 273 * Atlas-axis complex not included in this count. •••••. 10 • • • • .' o°o°o * Cloacal region 8 °i ** °°°£ • ° a* l=K A AA * A AAAAA^A AA % . V AA fl " fcO A y **A • o i A o* a a o • o u I D DD o a a d DD D "", a D D D D n dDdddd J L J I L 50 100 150 200 250 Vertebra number Fig. 2. Variation throughout the vertebral column in (a) length between zygapophyses (A J, (b) width across * postzygapophyses (O • ) and (c) length of hypapophysis ot two specimens <) of Pseudechis porphyriacus. Hollow symbols R14060 solid symbols R 14061. VERTEBRAE OF FOUR ELAPID SNAKES 75 20 10- I RU0S1 S3 R^kJL. 20 10 5VojR14060 ELSjct RW063 ^^^ J^i 10 RU066 [Qfti D 20r g RK059 JL 2 or 10 J. R14Q64 30 20h T^i Fossil 10" L m to I I I I I I I i i 1 1 ' » ' ' ' ' ' i't i i 0.5 1.0 11 12 13 1.0 1.1 12 1.3 W 15 03 0A 06 Rel. width Ant. width: Post, width Rel. length spine Fig. 3. Distributions of values for the ratios (a) width across postzygapophyses to length between zygapophyses, (b) width across accessory processes lo width across post- zygapophyses, and (c) minimum length of neural spine to length between zyga- pophyses. Left-to-right downward hatching, Pseudechis porphyriacus; open columns, Austrelaps superba; right-to-left downward hatching, Notechis scutatits; solid columns. Pseudonaja michaiis; fine stipple, Victoria Cave, type A; coarse stipple, Victoria Cave, type B. 7fi MEREDITH I. SMITH 20 ris 10 b RU061 20p 10 |g RK060 Dfij, 20 10- RU066 a 20r 10- %% RK059 I m RUOSfi RU065 J R14Q64 40 r 30 w 20 Fossil 10 I I I'll I i ' I I L '''' a* OS 0.6 07 0.2 0.3 M 0.1 0.2 0.3 Zygosphene Condyle Postzygapophysis Fig, 4. Distributions of values for the ratios (a) width of zygosphene to length between zygapophyses, (b) width of condyle to length between zygapophyses, and (c) width of postzygapophysis to length between zygapophyses. Hatching as in Fig. 3. VERTEBRAE OF FOUR fcLAWl SNAKHS 7T TABLE 2 mens, each foramen of this pair opening accessory process near the antero- SiHtff c}}tTrmfm,\ii<* uf ttoC doc/cat /i\\iion ef four through the \)W.itfH'n.\ <>t pfi^rfitchis porphynactis lateral margin of the prezygapophysial facet. Vertebral structure fn the cloacal region Nu other Number varies widely between specimens, and the num- Joiai with Number iffjtfl ber of vertebrae with articulated forked ribs number of jrlicu- wilh nh* wiih cloacal toteU single deeply may be equal to or less than the number Specimen tosrwbtae ribs hypypophyvib forked n single hypapophysis (Tabic 21 The fork is rnosc T deep on most forked ribs but on the H 1 4060 ft 1 8 * > vertebra, the notch may be R 14061 1 3 anterior cloacal RM062 5 4 4 A* midway along the rib The lymphapophyses 2 3 51 ft 14063 6 project laterally and slightly ventrally, as do the pleurapophyses of the post-ctoaeal vertebrae * TIii' amcnor-nrasi pair of rite have shallow fark*. The haemapophyses of each post -cloacal between zveapophvses \x 0.33 ± 0047 (36); vertebra arise separately from the ventral sur- completely JIU? ± ".0067 (36)1 (Fig. 4). The zygo- face of the centrum, and remain anterior pusi-eloacal sphcrte. viewed from the front, is drill and separate, although on the haemapophyses con- straight or slightly convex; from abuve it is vertebrae the tips of the shape of the nearly straight or slightly concave with * faini verge slightly. In lateral view the median notch on some vertebrae. The width hacmapophysis is similar to thai of the hypapo- precloacal vertebrae The pleura- of the zygosphene is a little over half the physe.s of the anterolateral^* (viewed length between the zygapophyses [.? 0.53 ± pophyses are directed anterior to .0063 (36); F0.57 * .OU38 (35)] Tfic prczy- from above or below) and extend gapophysial facets are oblong [length/ width the cotyle. iiregularities occut in the skeletons. I.11M.58, xl.40 x .019 1 36): 1.01-1 .73. f Some facets of 1.49 ± 025 (36)|. The acute accessory pro- In R 14060 the right prezygapophysial vertebras are enlarged lo cesses extend laterally perpendicular lu both the J 30th 10 134th left-side facet. vertical and horizontal 3\es of the vertehra, almost double the si^e of the of these vcrte- I of K 14062, .v 1 physis occurs in Wo vertebrae [T I 39 ± ,014 |35); .38 ± .Q10 (35)] spongy bone has com- (Fig. 3>. The postzygapophysc< are large where an outgrowth ot [widen postzygapophysis/ length between pletely fused the prezygapophysis to the pro- accessory 2yjttpophy.se*: F0.2G ± .003 (3; "0.20 ^ ceeding postzygapophysis. and the prob- .002 |3b)| and have an anterior notch that process is reduced. These abnormalities but R 14063 shows gives their otherwise ovutc shape a kidney out- ably resulicd from injury, abnormalities, firstly on verte- line. The maximum diameter of the post- Slight congenital distinct epi/.ygapo- zygapophysis is almost perpendicular to Ihe brae 91. where a small, but the left side only, and long axis of the vertebra and exceeds the width physial spine appears on [length/width 1.03-1,51. T 1.27 ± .020 (3o); secondly in lour posl-cloacal vertebrae. Near beginning of the pusicloacal scries, the 1.02-1.46. x J 22 = .017 (36) J. the vertebrae com- The typical four pairs of foramina are centra of two consecutive are the left, the outline of the present on all precloacal vertebrae, and in pletely fused On many vertebrae they are unilaterally or posterior edge of the neural arch of the first seen, but the right the bilatcially doubled, For example, in ihe I S3 vertebra can be on is faintly distinguishable The precloacal vertebrae of R 1 4061 the lateral suture hnc only vertebrae lie close foramina are bilaterally double (H vertebrae), pleurapophyses ul both centrum ot double on the right only (II) or left only together, apparently fused to the of haem apophyses (13); the paracotylar foramina are bilaterally the firsl verlebia. Two paijs surface uf the double (I), double on Ihe right only (5) or arise separately on the ventral post-cloacal kit only 78 MEREDITH I. SMITH 8 •••••.. •••• A *4 • 00 00 °o 00 o a oA 6fe A * aa 4 a o ft* &6 9 °D . D ?°° D a I PD I , , I , 2_i 9D9 qODODOD , , ^— i i i 50 100 150 200 250 Vertebra number Fig. 5. Variation throughout the vertebral column in la) length between zygapophyses (A* ), (b) width across zygapophyses (O*), and (c) length of hypapophysis (W »" two species. Hollow symbols, Ausirdups superba (R14066), solid symbols. Notechis scutatia (R14058). similar abnormality has been observed in dor- from above. The ratio of zygosphene width to sal vertebrae of colubrid snakes (King 1959). length between zygapophyses (Fig. 4) has a mean of 0.50 ± .0038 (27). The prezygapo- Austrelaps supcrba (Giinther) physial facets are oblong, the postzygapo- The low number of precJoacal vertebrae in physial facets obovate. The acute accessory this specimen (Table ) is 1 confirmed as typical processes are relatively shorter than in P, por- of the species by the ventral scale number (151 phyriacus in most vertebrae (Fig. 3). according to Worrell 1963). The width across Of the four cloacal vertebrae, one has the postzygapophyes is less than the length articulated forked ribs and three have a between zygapophyses in the first ten and the hypapophysis. The hacmapophyses of the posl- last 35 precloacals (Fig. 3). cloacal vertebrae are long anteroposterior^, The hypapophysis (Fig. 5) is similar in form about half the length of the centrum, and are to that of P. porphyriacus and does not extend completely double. further posteriorly than the posterior surface of the condyle. Notechis scutatus (Peters) subcentral The ridges arc low and rounded, In all precloacal vertebrae, the width across as in P. porphyriacus, but the interzygapo- the postzygapophyses exceeds the length physial ridges of A. superba are stronger. Small between zygapophyses (Fig. 5). Each hypapo- epizygapophysial spines occur the first on six physis is extremely compressed laterally to a vertebrae, The condyle is smaller, relative to thin lamella that terminates posteriorly in a the length between zygapophyses, than in p. sharp point not extending posterior to the porphyriacus (Fig, 4). The zygosphene is thin posterior surface of the condyle in any but the and slightly convex from the front, convex first 15 vertebrae, where the hypapophysis is VERTEBRAE OF FOUR ELAPID SNAKES 79 1 cm Fie. 6. Line drawings, to exact scale, of the 80th vertebra of (a) Pseudechis porphyriacus (R14060), (b) Notechis scutatus (R14059), and (c) Pseudonaja nuchalis (R14064) in dorsal, lateral, ventral, anterior and posterior views. — so MLRHDrTH I. SMITH very long (Tig. 5.1. The suheenirnl ridges arc hypapophvsjs is well developed on all pre- low and rounded: the inter/ygapophysiu) i'(d;ees cloacal vertehrae. Thickenings on the rim of are weak hut distinct. The neural spine is the eotyle on either side of me ventral midline higher than in P. porphyriacm (Hip. 6) and jts give rise to a low ridge lhai narrows sharply horizontal dorsal edge overhangs boih in join the laterally-comptessed h>papophysis anteriorly ami posteriorly The minimum The hypapophvsis deepens from about the lenglh of the ncur-il spine is generally relatively middle of the vertebra and terminates in a shorter than in the specimens of P, porphf- rounded point thai extends posteriorly to the nacus [ratio of minimum length of neural condyle. Subccntral ridges are strong and inter- spine lo length between ?ygapoph\scs: 7 0.4S zygapophysial ridges are distinct, lite para- ± .0064 (35): x (X4fl =t .0049 (3n"l| (Fig. 3) diiipophyscN are well developed, with a pro- Hpizygapophysial spines are welt deveJopcd on truding. dors d i mi net neck fh'ig. 6). The ihin /ygosphenc is lateral border Of the eotyle. The neural spine straight or slightly convex from the front, is low and long (Tigs 3 and 6); its dorsal edge siraighl or slightly concave from .above. The overhangs slightly at the front, markcdlv prezvgapophysial facets are oblong (length/ bchmd Cpizygapophysiai spines are distinct on width I.OO-i.67, .7 1.47 =t .027 135); 1.09- the ftisi five vertehrae, family visible on the 1.58. x 1.35 ± .022 (35)]. The accessory pro- sixth to eighth and absent from all others, the cesses are obtuse and short, so that the ratio eotyles and condyles are almost round; in of width across accessory processes to width R 1 4065 some eotyles are depressed slighily, across posuygapnphyses is generally less than but »n R 1 4064 some are slightly compressed in P parphyitKus (Fig. 3). The posuygapo- laterally. The condyles are relatively much physcs arc obovate [length/ width 1.10-1.58, 7 larger m the smaller specimen than in the 1.35 i= .021 135); 1.00-1.47, J 1.2S t 022 larger one (Fig;. 4). The zygosphene is convex < 35)1* and their width is about one-fifth of from the front, concave from above, it is thin the lenglh between the zjgapophyses (Fig. 4), in vertebrae of the smaller specimen but is Prezvgapophysial foramina arc constantly thickened in those of the larger. The pre- present. zygapophysial facets are oblong [lenefh/ width Jn both specimens, all five cloacal vertebrae 1.09-1.72, J 1,39 - 022 (41); US-1.78. * have fused forked ribs. One has three, the 1.35 ± .018 (41)|, and the acute accessory other two cloacal vertebrae with a single processes extend laterally well beyoad the hypapophysis. articular facets (Rig, 3). Except fur a slight The baemapophyses are large, paddle-shaped notch posteriorly, the outline of the post- and completely paired. The plcurapophyses zygapophysial facet is almost round [length/ extend ventrally more than laterally. The width 1 OVI .35 5 1.17 ± .011 (41); 1.09- /.yganophysial facets extend anteriorly and 1.59. A 1.30 ± .019 (41)1. posteriorly, rather than laterally as in the pre- In the cloacal region the ribs arc all fused cloacal vertebrae, and acute nccessory pro- except for the anterior vertebra uf R 14064, cesses ore distinct on all the postcloacal verte- and a hypapophvsis occurs on one (R14065) brae. or three (R14064) cloacal vertebrae. Unilateral or bilateral doubling of the para- The haemupnphyses arc completely paired cotylar and/ or subeetUial foramina occurs in and extend anteriorly as two separate ridges very a lew prcctoacal vertebrae (no more than to the Jim of the eotyle The plcurapophyses four In cither specimen). are broad and flat and not pointed. In anterior view they project ventrolateral!}' but in ven- Pscudtuutju nucha lis Ciiinther tral view mainly laterally and only slightly Pwudonuiu nuchuJb has more vertebrae anlerinriy. than any of tire other species studied here In RN065, more than the one pair of para- fTnble IK and maximum length and width coiyh,r foramina appear on many vertebrae occur rn sequentially more posterior vertebrae 15 prccloacal vertebrae have an extra para- : than in the other species • *• • • • 10 • • A44A ## AA4 A A A A * AA A A*A* AA A • • A \Cloacal region 8 A G- 00oo A« O oo oo oo • A oo ooo °0° *; A fl A o° aAA^A^**^ %AAAAA A I O A^A *&*&.°0 4% - 8'° A "AA s * A • A A * A * A Aa o o »• • a o » o D o a D a o° - « on Q a° DO oa a u d d D D D D n D D DD n a D D J . I L 50 100 150 200 250 Vertebra number Fig. 7. Variation throughout the vertebral column in (a) length between zygapophyses ( A A ), (b) width across zygapophyses (0«), and (c) length of bypapophysis (D") "* Pseudotwja nuchalis. Solid symbols, R14064, hollow symbols, R14065, 8 precloacal vertebrae with additional para- convex in anterior view and slightly thickened; cotylar foramina. The left postzygapophysis of Type B: almost straight, with a median notch, the 114th vertebra is fused by an outgrowth of when viewed from above, almost straight in spongy bone to the prezygapophysis of the anterior view and extremely thickened. This succeeding vertebra, which lacks an accessory thickening is consistent with the robust process on the left side. The left rib of vertebra appearance of the vertebrae. The subcentral 114 shows a healed fracture near the articula- ridges are particularly strong (Fig. 8). tion with the vertebra. As well as the thickening of the zygosphene, its width relative to the length between zyga- Pleistocene fossil vertebrae from Victoria Cave pophyses differs significantly between the The precloacal vertebrae found in the Vic- types A and B [A, J 0.52 ± .0048 (40); B, x toria Cave deposit vary in length from about 0.54 ± .0057 (40); .002 < P < .01]. The dis- 2 mm to a maximum length between zygapo- tributions of the values for this ratio overlap physes of 11.1 mm. In their general conforma- widely not only between types A and B from tion they closely resemble those of Pseudonaja. Victoria Cave but also among the specimens However within the sample of vertebrae from studied (Fig. 4). The ratio of width across Victoria Cave, two types can be distinguished postzygapophyses to length between zygapo- on the characteristics of the zygosphene: Type physes tends to be greater in the fossil verte- A; slightly convex when viewed from above. brae than in A. super ba; the ratio of width 82 MEREDITH ). SMITH I cm 1 cm D Fig. 8. Precloacal vertebrae of Pseudonaja nuchalis; left, 80th precloacal of R 14064; right, Victoria Cave, Type B (P16126b) in dorsal (A, B), ventral (C, D) and anterior (E, F) views. VERTEHRAH Or FOtIR ELAPID SNAKHS 83 jygapophyses, the possibility exists that they 1*1 « are of a species different from Type A. How- ever, these characteristics are two which 31 - develop with age (Auffenberg 1963) and most of the Type B vertebrae are larger than the 6.0-8,1, 6.7; Type Type A (Type A t length x 2S B, 7.1—1 1 . K x 8.9) and also generally arc larger than the vertebrae of the modern speci- mens. When the correlation between length JfJ 5*# between zygapophyses and relative width of J* postzygapophysis of Type B vertebrae was tested by the Kendall Rank Correlation Test £ .,. - **£.% (Siegel J956) the correlation was found to be '|U a*V i i i i 1 1 1 1 1 The post-cioacaJ vertebrae recovered from L 5 6 7 B 9 >Q It (2 Cer I nurn I ETTjrth 'mm) Victoria Cave resemble those ot P. nuvhulh in the laterally-directed plcurapophyses. In the Kg. Double-logarithmic regression of width of other species studied the pleurapophyses pro- posslzygapophysfc on length between post- ject anterulatcraily. zvgapaphyses in Pseudonaja nuchaVts, K R 14054; + f R14064; Q, fossiL Type A: • . fossil. Type B. Discussion In all the vertebrae of the four species, the across accessory processes to width across hypapophysxs is well-developed (as in all posCzygapophyses is less than in most P. par- elapids) and hypapophysis length decreases phyr'iacus and the relative length of the neural slowly from anterior to posterior along the s»pwe is generally greater than in N. scitiafus column. There is no suggestion of two distinct (Fig. 3). In these ratios, the Victoria Cave regions as in Aehrochordus juvanieus where vertebrae closely resemble P. mtchalis. In the the anterior region ( to vertebra 96 ) has relative width of the condyle, the distribution hypapophyses long and fairly constant in of ratios for Victoria Cave vertebrae resembles length and the posterior region has hypapo- the larger P. rwchalis (R 14064)., though not physes short and of constant length (Hotl- Lhe smaller (RI4065). Finally, in the relative stetter & Gayrard 1964). width of the postzygapophyses the Victoria presence vertebral Cave vertebrae of Type A resemble the larger At family level, the of diagnostic value [e.g. the hut not the smaller P- nuchalis, and the relative foramina may have constant absence of lateral foramina in Achro- width of the postzygapophysis is generally (HorTstetter Gayrard 1964)1. but greater in Type B fossils than in any of the chordidac & the variability in the number of foramina at modern species studied I Fig. 4). subeentral, etc.) in This subjective and objective analysis of the each position (i,e< lateral, indicates the need for caution fossil vertebrae indicates that they most closely the one snake resemble Pseudonaja- The Type A vertebrae in the use of foramina in taxonomy. can be referred with confidence to this genus, Although middle prccaudal vertebrae may Because the Type 6 vertebrae differ in the be the most constant in their structure (within thick 2ygosphene and the relatively large ROSt- species) and hence best for identification — 84 MEREDITH !. SMITH (Auffenberg J 963), the difficulty of assigning fossil genera may be readily identified, but an isolated elapid vertebra to a particular further studies of congeneric species are region of the column precludes confidence in needed to determine whether specific identifi- selecting middle precaudals from a sample of cation is possible. fossils Because of the consistent variations along the column, together with some irregular variation (e.g. doubling of foramina), to iden- Acknowledgments tify isolated vertebrae it is necessary to con- Mr R. Shine, Mr G. Whitten and Dr R. T. sider not just the middle precaudal vertebrae Wells kindly donated the modern snakes. Dr of reference specimens, nor the mean of some Wells and members of the Cave Exploration value (even if it be given with standard error) Group ot South Australia helped in excavating but the range through which a given character the fossil vertebrae. I am grateful to Dr R. I varies. Also because of variations along the Wells, Mr I. M. Thomas. Dr T. F. Houston column, no unique specific character was and Mr N. Pledge for their criticisms of the found, and so it is necessary to consider several manuscript. Mr P. Kcmpsler prepared the characters in the identification of fossils. The photograDhs for Fig, 8. References Aufpenblrg, W. (1963).—The fossil snakes of Lydekxer, R, (1888).—"Catalogue of Fossil Rep- Florida TuUme Stud. Zoot. I0 f 131-216.- tiles and Amphibians in the Uritish Museum of Natural History." Part 1. (London.) Hoffstjctter, R., & Gasc\ J. P, ( 1 y Qld Mu.w 8, 1 11-112. (Angus and Robertson: Sydney.) THE ONTOGENY OF THE VOCAL SAC OF THE AUSTRALIAN LEPTODACTYLID FROG, LIMNODYNASTES TASMANIENSIS byM. 7. Tyler* Summary TYLER, M. J., (1975). -The ontogeny of the vocal sac of the Australian leptodactylid frog Limnodynastes tasmaniensis. Trans. R. Soc. S. Aust. 99(2), 85-87, 3 1 May, 1975. The ontogeny of the vocal sac of Limnodynastes tasmaniensis proceeds from initial bilateral evaginations of the mouth floor, through median fusion to a unilobular, submandibular structure. The acquisition of pigmentation by the submandibular skin is a concomitant process. It is suggested that the vocal sac evolved by a path now reflected by ontogeny, and involving progressive bilateral herniation. THE ONTOGENY OF THE VOCAL SAC OF THE AUSTRALIAN LEPTODACTYLID FROG, LIMNODYNASTES TASMANIENSIS by M. J. Tyler* Summary vocal of the Australian leplodaelylid frog Tyi.er. M. J , (1975).—The ontogeny of the sac Lirntiodynasfes tasmaniensis. Trans. R. Sac. S. Anst. 99(2). 85-87, 31 May, 1975. The ontogeny of the vocat sac of /.imnorfynanrs tasmaniensis proceeds from initial bilateral evayinations of the mouth Moot, through median fusion to a tmilohular, submandi- bular .structure. The acquisition of pigmentation by the submandibular skin is a concomitant process. It is suggested that the vocal sac evolved by a path now reflected by ontogeny, and involving progressive bilateral herniation. Introduction Wesl Beach near Adelaide on 1 September length Voca! sacs occur only in male anurans and. 1963 (SAM, R5290). The snout to vent 1 to in most species, comprise inflatable, epi- of the 8 males ranges from 30 34 mm. thelium-lined chambers located between the Dissections were performed with the aid of hyoid plate and the superficial mandibular a low-power binocular microscope, and musculature. Data on the ontogeny of vocal measurements made with a pair of dial cal- muscles stained with the re- sacs are limited to studies on only a few, lipers. The were mostly African, species (Jngcr 1956; Inger & versible iodine/ potassium iodide stain des- Circenberg 1956). The available information cribed by Bock ik Shear (1972), in order to differentiate the vocal sac from the surround- is so inadequate that the possible contribution vocal sae of ontogeny to an understanding of the evolu- ing striated muscles. Muscle and ( 1971 ). tion of vocal sacs has not been assessed. At terminology follow thai of Tyler present there is no published information on the ontogeny of the vocal sacs of any of the Observations 300 (approx.) species of frog* found in Aus- Secondary Sexual Characteristics tralia and Guinea New The secondary sexual characteristics of The Australian leptodactylid species Limno- IJrnnodynastes tasmaniensis are as follows: dynastes tasmaniensis represents an ideal males possess a unilobular, submandibular subject for studies of vocal sac onto- initial vocal sac, yellow pigmentation of the sub is superficial mandi- geny. This because the mandibular skin and glandular nuptial pads on bular musculature of adults has already been the first and second finger*. Females bear described in detail, and there are published broad lateral fringes to the first and second observations variation in the position occu- on fingers, and sometimes to the third. pied by the vocal sac when it is inflated (Tyler 1971). Vocal Sac Ontogeny The earliest step in the progress towards the Material and Methods development of vocal sacs involves the forma- Of 646 specimens of Limnodytiastex tas- lion of a shallow and elongate involution of maniensis in the South Australian Museum, the floor of the mouth on one side of the one series was found to exhibit ontogenetic tongue. There is a slight elliptical, ventral inclination (Fig. variation in the extent to which the vocal sac depression with a mediad lateral margin of the depression intrudes above the superficial mandibular mus- I A), and the border of the anterior culature. This series comprises 18 male speci- is level with the lateral mens from a group of 68 males collected at cornu of the hyoid. * South Australian Museum, North Terrace, Adelaide, S. Aust 5000. H M J tyler accurate external index of the presence, and stage of development, of the vocal sac struc- ture. In specimens lacking the initial evalua- tions in the mouth floor, the submandibular skin Was either entirely unpigmented 01 else bore a few scattered chromatophores at the periphery of the mandibles. Development of the evaginations was accompanied by an in- crease in the density of pigmentation and of its medial limit. The pigmentation, and the appearance of the hright yellowish background color of the submandibular skin, progressed in an identical sequence until, at completion of vocal sac development, the skin was entirely yellow. Discussion Beyond the sphere of its intrinsic interest, ontogeny can contribute to an understanding of the evolution of structure. In the present Fig. 1. Selected progressive stages in ontogeny of situation the progression of the vocal sac from vocal sac. A. Single, elliptic ventral' de- paired, lateral evaginations to a single, large pression, ti. Bilateral expansion mediad. C. Further medtad and initial caudad sac could readily be regarded as a recapitula- development. D. Media! unitv of separate tion of evolutionary history. However, the sacs. nature of the progression also indicates why the anuran vocal sac originates in the way that In four specimens, development was con- it does. lincd to such an evagjnation on the left side; in a fifth (here were bilateral evaginations. The floor of the mouth is supported by the From these slight folds, the vocal sac develops hyoid plate and its processes, and by muscles bilaterally, and as tar us could be determined communicating between the hyoid and the quite concomitantly, into roughly circular bags mandibles. These supporting structures provide intruding between the superficial, ventral, what can be visualised as a broad and complex mandibular musculature and the deep inter- sling in which the only gaps arc a narrow mandibular muscles situated above them (Fig. lateral zone on each side of the tongue, so IB). At this Stage of progress mediad develop- situated between the anterior cornua and the ment is more pronounced than anterior or pos- mandibles. In all anurans possessing vocal terior development. Simultaneously, that area sacs, apertures originate within these 'unsup- of the mouth floor between the anterior cornii ported' zones. and the mandible becomes depressed, render- The probable steps that lead to the evolution ing the aperture to each portion of the sac oi vocal sacs in this species, or its ancestral more conspicuous. stock be t can reconstructed quite readily As the two halves of the vocal sac approach Assuming an increase in the pressure of the one another, their posterior margins extend buccal cavity during vocal activity, the existing further caudad (Fig. 1C). This posterior en- sites ot I he vocal sac apertures are probably farsement is accompanied by comparable the areas of least resistance: in these regions enlargement of the interhyoiileus muscle into the superficial tissue is of a rather elastic- a slight lobe, extending beyond the post-articu- nature, and presumably subject to the greatest lar extremities of the mandibles. Ultimately distension. It follows that the first stage in the the vocal .sac occupies the entire muscular ontogeny of the vocal sac of Limnodynaxtes lobe, becoming united medially by loss of the tasnnmiensis is precisely that initial event, common medial wall (Fig. ID). Su& sequent stages could well have arisen from The presence and extent of submandibular* li'.lle more than progressive bilateral hernia- derma) pigmentation was found to provide an tion. ONTOGENY OF FROG VOCAL SAC 87 References Bock, W. J., & Shear, C. R. (1972).—A staining Inger, R. F., & Greenberg, B. (1956).—Mor- method for gross dissection of vertebrate phology and seasonal development of sex , , , , A i-ti i*v7 characters in two sympatnc African toads. muscles. Anat Anz. 130,n 122-227. ; Morphm 9% 549-574. Inger, R. F. (1956).—Morphology and develop- Tyler, M. J. (1971).—Voluntary control of the sh e US ~ ment of the vocal sac apparatus in the A trahanf ^ ^Zt'^vff^*}*™!}*leptodactylid frog Limnodynastes^ tas-, „ . Afncan frog Rana (Piychadena)t * porosissima maniensis. Trans. R. Soc, S. Aust. 95(1), Steindachner. /. Morph. 99, 57-72. 49-52. THE PRE-SETTLEMENT VEGETATION OF THE MT GAMBIER AREA, SOUTH AUSTRALIA BYR. 7. DODSON* Summary DODSON, J. R. (1975). -The Pre- Settlement Vegetation of the Mt Gambier area, South Australia. Trans. R. Soc. S.Aust 99(2), 89-92, 31 May, 1975. There are some conflicts as to the nature of the pre- settlement vegetation formations around Mt Gambier and Glencoe. European settlers long ago cleared the areas of their vegetation cover. Pollen analysis of Brownes Lake sediment reveals that the most likely formation around Mt Gambier consisted of open grassland with perhaps a sparse cover of woody taxa. THE PRE-SETTLEMENT VEGETATION OF THE MT GAMBIER AREA. SOUTH AUSTRALIA by J. R. Dodson* Summary Dooson, J. R. (1975).—The Pre-Settlement Vegetation of the Mt Gambler area. South Aus- tralia. Trans. R< Soc. S, Aust. 99(2), 89-92, 31 May. 1975. There are some conflicts as lo the nature of the pre-settlcment vegetation formation* around Mt Gambier and Gleneoe. European settlors long ago cleared Ihe areas of their vegetation cover. Pollen analysis of Brownes Lake sediment reveals that Ihc most Likely formation around Ml Gambier consisted of open grassland with perhaps a sparse cover of woody taxa. Introduction 1966). Hill (1972) recounts some of the early Crocker (1944) left gaps in his veget;uion records of water level changes of the Mt Gam- map in the areas around Mt Gambier and bier lakes and it appears that the most spec- Gleneoe, in south-eastern- South Australia, as tacular observed were in Brownes and Leg of Ihe original vegetation was no longer evident Mutton Lakes, These are the shallowest and thus changes may have been more obvious. at the time of his survey. Crocker ( 1944) and liver & Crocker (1951) hypothesized that the Henty's hut. the first building in the area, is areas were occupied by lightly wooded grass- said to have been erected in 1841 on the site land, and the trees cleared after settlement- of Brownes Lake, Brownes Lake and Valley filled Woods ( 1562) recorded the vegetation in the Lake and joined and ?be new lake Blue Lake crater at Mt Gambier as thickly reached its maximum depth in the 1890*s. wooded with several varieties of Melaleuca. This paper provides data on early vegetation The vegetation is sparse in the photograph in at Mt Gambier. Pollen analysis is an ideal Hill (1972, p. L08) taken in 1860 ancfyct Hill method for tackling this problem as the craier when describing an early record < 1861 ) of the walls are mostly steep-sided am! can support first road between Mt Gambier (then Gam- Jiirlc local vegetation. Therefore a significant biettown) and Port McDonnell on the coast proportion of the pollen rain is probably states (p. 109) regionally derived. Today the craters arc heavily exploited recreation purposes "the route to the 'Bay* was through dense for and most of the vegetation within is intro- bush counlry, mud and slush in winter, dust them duced. from native vegetation in summer, and tenanted by thousands of The change should in fossil kangaroos at all times." be recorded the pollen record. vSpecht (1972, p. 203) in his vegetation map Methods of the South East simply records the areas Before the Brownes Lake core site w«is around Mt Gambier and Gleneoe as cleared. selecied for investigation, areas of Brownes Brownes Lake occupies portion of one of Lake, Valley Lake and Leg of Mutton Lake the craters which formed in the Volcanic erup- were checked for undisturbed sediment. Access tions at Mt Gambier after 5000 years B.P, to sediment at Blue Lake was impossible with (Fergusson & Rafter 1957, Blackburn 1966). the equipment at hand. A 160 cm sample of It is 4-5 m deep, sits on collapsed volcanic peat was collected from Leg of Mutton Lake. debris and its water surface, like those of the The dry periods in Leg of Mutton Lake (Hill other three crater lakes, is an expression of the 1972. pp. 112-114) which are unfavourable regional water table (Bayly & Williams 1964, for pollen preservation rendered this core un- Department of Biogcography and Geomorphology. Australian National University, Canberra, ACT '2600. Present address: Geography Department. University of Canterbury, Christchurch, New Zealand r 90 J. R. DODSON *7^ Depth err; Restionaceae Poaceae htnuiypiui Type 1 ° m t Litaitypiux lype * it Cyperaceae l.iii'tiivpius iype4 ^j Myiaceae Type 1 8 Myrtaceae Type2 ^ Cuptessacaae P r ;_J1 L Unidentified pollen Cusmiiina (,>28^nVl S r J* L Ph'lhluuii n ( '(j\thJritui fe28pm) ° :c -\Unophyiiwn l.yuiftotfiitw Monolete spores H Acacnti ° Chenopodiaceae -n-n As;erac«ae (Tuoiliflorae) s g Asleraceae (uguhflnrafi) a t paendaceae Plan/two ititneohhi ^ Trigtochm ° r Pvfumngetim *n Brassicaceae J- :et— ftappw ° r i— Liliaceae u r Polyganaceae /.v/j/w ° r epitaenct Z, i I E Hygtrichospheres Haioragaceae Pimdia Gyrostemonaceae M^lva^eae Slylidiwn Restionaceaft Fig. 1. Pollen diagram for Brown es Lake. I \«LY MT GAMB1ER VEGETATION :> suitable for pollen analysis. No suitable sites region wi;h a heath fringe around the lake. The mainly Amhoeeros, we-.ret found &JL the accessible ureas in Vallev local vegetation was representing Lake nor in much or the Brownes lake area. Cypcraceae, and Afyriophylftwh a In February 1*574, a 45 cm core was shallow water environment at the core site collected with a D-vection sampler from the This pollen assemblage tends to confirm the core site on the eastern shore of lirownes Lake. Crocker hypothesis that trees were few. rather Hill, unless It consisted of 30 cm at black lake mud over- than the early report recounted by ling 15 cm of pale brown (straw coloured) the scrub described by the latter consisted of material which was largely clay-sized particles low pollen producers such as Banksia or of silica. This Is probably reworked volcanic Acactu. There is no evidence in the form of debris. The core was sampled on-site and remnant vegetation to support this, The organic pollen analysis was carried out in the labora- and pollen and spore content of ihe sediment tory using the standard hydrofluoric acid. was low (except for Atuhoceros spores which alkali and Erdtmnn's acetylosis methods as must have been derived locally), suggesting described by Faegii & iversen (1964). Resi- cither slow sedimentation during alternating dues containing pollen were dehydrated. wot and dry conditions (which could result in mounted in silicone oil. and counted until at the loss of pollen through oxidation) or fairly least 200 pollen grains of woody taxa had been rapid in-wash of inorganic material from the recorded. Relative peivrentages for terrestrial steep crater walls. On the evidence presented pollen and spore taxa were calculated against here it is not possible to favour either explana- r pollen sum of total land plant pollen exclud- tion. ing ihe recorded introduced taxa (Pimts and The post-settlement phase of the pollen dia- PlcwtagO hmceolata). Frequencies for aquatic gram is dominated hy Poaceae, herbs and vascular plant pollen and Fystrichosphere Phws ntdiata. and also shows increasing Euca- remnins were calculated aguinsl a pollen sum Ivp/ttx frequencies. Antlwceros and Cyperaeeac of total aquatic pollen. The results were decrease in importance and Myuophyllu/n plotted on a pollen diagram 'Fig. 1). Ecolo- dominates the aquatic spectra, indicating a gical information for taxa in the study area change to deeper water at the core site. Since and details of pollen identification have been the core site is near the edge ot' the lake, it given hy Dodson (1974). follow? that water would not be deep there until the level rose above its present position. Results' and Discussion Since the rise accompanies die increase in The short pollen diagram (Fig. 1) has not pollen of exotic plants, it seems likely that the been divided into zones, but the presence of rise is the one recorded for the latter part of introduced taxa divides the diagram into post- the i9th Century when Rrownes Lake and seUlemciu (0-10 cms) and pre-settlement Valley Lake were joined. The increase in phases (20-45 cms). Eucalyptus pollen is undoubtedly due to the The prc-scttlement phase was dominated by plantings established in the craters for recrea- Astcraccac (Tubulillorae). Poaceae and tion facilities and the wildlife Teserve and not Pteridium, with small numbers of herb pollen, to any change in the naiive vegetation Tree pollen was virtually absent, although small and increasing frequencies of scrub taxa Acknowledgments ( Ctisnwmct ( ^ 2$p$i ) -pi'obably C. pehtdosa. and Mvrtaccac Type i-Lepm$penvitr*i juni- It is a pleasure to lhank Gurdip Singh and perinum and l.eptQspermum myrsineides) were Joan Guppy who critically rend the manu- obtained. Assuming that regional pollen script, and the Australian National LTniversity dominates, then the assemblage most likely for supporting the work with finance and represents open (treeless) vegetation in the equipment. References Kwtv, 1. A. E., & Williams, W. D. (1964).-- TUylv. L A. C, & Williams. W. IX Blackburn, G. (1966).—Radiocarbon dates relat- Fergusson, G. J., & Rafter, T. A. (1957).—New 14 ing to soil development coastline changes, and Zealand C age measurements— 3. N.Z. J. volcanic ash deposition in south-east South Sci. & Tech. 38B, 732-749. Australia. Aust. /. Sci. 29, 50-52. Hill, L. R. (1972).—"Mount Gambier—the City . around a Cave." (Investigator Press: Crocker, R. L. (1 944 ) —Soil and vegetation Ade- relationships in the Lower South-East of laide.) South Australia. A study in ecology. Trans. Speciit, R. L. (1972).—"The Vegetation of South R. Soc. S. Aust. 68, 144-171. Australia." Edn 2. (Government Printer: Adelaide,) Dodson, J. R. (1974).—Vegetation history and Tiver,, N. S., & Crocker, R. L. (1951).—The water level fluctuations at Lake Leake, south- grasslands of south-east South Australia in eastern South Australia. I. 10,000 B.P. to relation to climate, soils and developmental Present. Aust. J. Bot. 22, 719-741. history. /. British Grassland Soc. 6, 29-80. Faegri, K., & Iversen, J. ( 1964).—"Textbook of Woods, J. E. T. (1862).— "Geological Observa- Pollen Analysis." Edn 2. (Munksgaard: tions in South Australia." (Longman: Lon- Copenhagen.) don.) AUSTRALIAN LEPTODACTYLID FROGS OF THE CYCLORANA AUSTRALIS COMPLEX byM. 7. Tyler* and A. A. Martinf Summary TYLER, M. J., & MARTIN, A. A. (1975). -Australian leptodactylid frogs of the Cyclorana australis complex. Trans. R. Soc. S. Aust. 99(2), 93-99, 31 May, 1975. Cyclorana australis as now defined is shown to comprise two closely related species: C. australis confined to northern Australia and C. novaehollandiae to eastern Australia. Notes are provided on the tadpole of C. australis, and the calls of both species are analysed. Call divergence is so limited that hybridization is considered possible in sympatry. ; \USTRALL\N LEPTODACTYLID FROGS OF THE CYCLORANA AUSTRALIS COMPLEX by M. {» Tyler* and A. A, Martin! Summary lYLtK, M. L & Martin, A. A. (J 975).—Australian Icptodactylid frogs of the Cyvtvnwa atistralis complex. Trans. #• Soc. S. Aust. 99(2), 93-99, 31 May, 1975. Cyctorana australis as now defined is shown to comprise two closely related species: C. australis confined to northern Australia and C. ttovaeholkmdiae to eastern Australia. Notes are provided on the tadpole of C. tiustruiis, and the calls of both species are analysed. Call divergence in so limited that hybridization is considered possible in sympatry. Introduction hvlhmdiae by Steindachner (1867), and as In recent years, examination of the biology Phractons ahttaceus by Peters (1567). Both and morphology of several geographically names were referred to the synonmy of aus- widespread "species^* of Australian frogs has tralis by Boulenger (1882), revealed that each comprises a complex of We have assembled and examined large col- species. For example, Crinia signifcm as recog- lect ions- of C australis (sensu lato) from nised by Parker (1940) is now known to be various sources. Here we report our findings a complex of seven species (Moore 1954; and propose the recognition of a complex of Lilflcjohn 1957; Main 1957; Straughan & two species. Main 1966; Tyler & Parker 1974): Mixophyes Methods fasriolaius is now four species (Straughan 1968) and Limnodynastes dorsalis is also lour The specimens reported arc deposited in the institutions: National Museum of | Martin 1972). following The most neglected leptodactylid genus Is Victoria (NMV): Naturhistoriska Riksmuseet. Cyctorana, of which the type species is A fates Stockholm (NR); Department of /oology. australis Gray (1842), described from material University of Melbourne (MUZD); Northern collected in the Northern Territory. This Territory Museum, Alice Springs ( NTM ) species, as currently defined, extends from Queensland Museum (QM); South Australian northern Western Australia to northern New Museum (SAM): and Western Australian South Wales: a geographic range of approxi- Museum (WAM) mately 3500 km. The conspeeificity of indi- Measurements of specimens (to 0.1 mm) viduals from the extremes of this extensive were obtained with a pair of Helios dial range is obviously suspect, and even the most callipers. Abbreviations employed in the text cursoiy comparison of specimens of C. aus- and tables are as follows; F — foot length (the tralis from the Northern Territory and distance between the proximal end of the tar- northern Western Australia with those from sus and the distal tip of the fourth toe); HI. Queensland reveals striking differences between — head length (the -distance between the an- them. The northern individuals lend io have terior extremity of the snout and the posterioi a rather elongated head* a distinct, dark rostral margin of the tympanic annulus); HW = head stripe and a narrow suhoeular ban Tn con- width ( the maximum width of the head, trast, most individuals from Queensland are usually taken at the posterior extremity of the particularly robust animals with a broad head, mandibles); TL — tibia length (obtained by and frequently obscure head markings: a placing the tibia between the callipers); S-V population described as Cyclorana novae- — snout to vent length (the distance between * South Australian Museum, North Terrace, Adelaide, S. Ausl. 5000. t Department of Zoology, University of Melbourne, Parkville. Vic. 3052. , 94 M. J. TYLER A A. A. MAfcTlN the anterior tip of the snout and the anterior of the maxillary, premaxillary, nasal, fronto- margin of the cloaca). parietal and squamosal bones. On the dorso- Ratios calculated and subjected 10 the Stu- lateral body surfaces there are continuous or dent (-test were TL/S-V, HL'HW. F/S-V, disrupted, longitudinally orientated skin folds F/TL and S-V/HW, Larval stage numbers commencing behind the skull, and terminating follow those employed by Gosner (I960). above the groin. Mating calls were recorded in the Meld using Nothing is known of the breeding biology a Uher 4000 Report portable tape recorder of the members of the complex, but they are and Beyer M69 dynamic microphone, af a tape probably opportunistic breeders. The eggs ate vpeed of 19 Calls era /sec. were analysed by small and pigmented (ovidiametors of ovi- use of a sound spectrograph (Kay Model ducnl eggs range from 1.1 to 1.3 mm), and the f»061-A with Sona-Graph) the overall res- tadpole (one species) is of the hylid type with curve in ponse maintained ihe FL-I position, Iwo upper and three lower rows of labial teeth, calls Three of each individual were analysed an acuminate tail tip and a median or slightly and mean values calculated. Kach call was. de^tral anus. analysed twice; a narrow-band 145 Hi band- The geographic range of the complex ex- pass) analysis at recording speed tn determine tends from northern Western Australia to duration and dominant frequency, and i wide- northern New vSouth Wales (Fig. I ), Of what band (300 Hz bandpass) analysis at half we demonstrate to be two component species, recording speed to resolve fundamental fre- C. australix (sanm strirta) occurs in northern quency. Western Australia, the Northern Territory and The Cjclorana anstralis complex northern Queensland lo the west of the Divid- ing C. is Frogs of the Cyctorana austratrs complex Range; ri R4306 7, Crystal Ck; WAM, R 1 558-5*), Dryttdaic River Mission; WAM. RZ1Z33, Kownl Down*; WAM, R1377. 43282-86, 42399-42422, Knfum- buru; WAM, R22369-75, Kimberley Rcscaich Stn; SAM., R4769-70, R5O70. Kummurra; WAM, M 654-57, Landar Stn; WAM, R423X7, HO km S of La Grange; WAM, K4253G-40. 1238 1, 43478. 43491, Mitchell IMalcau; WAM, R42530-35, Main Ord River Dam Site (spillway): WAM, R42424, Mt Hait; WAM. R32099. Ml Anderson; WAM, R32291. Ml Barnell; NR, 1562, Mowh. Down; Fig. 1. Geographic distribution of ihe frogs of the WAM, KI3726, Oscar Ranges; NMV, D2J54-55, CyLloruita australis complex. Circles - Km George IV; WAM. R32I49, Si George Range. C. atosttiilis: triangles = C. tiovaehollan- WAM, Rl 1208, R 1 1894, RI2332, Woljulum: diae. Closed symbols indicate sites of The WAM, R26769-70. Point Springs. Webber Rang*; material examined, and the open circles WAM. R32351A, Wvndftam; WAM, R2*093. 40 one of ihe following literature references- km ST- of Wyndhom; WAM, R2O307. Yeda Cross- Braltsu-om 1 1 ) . Loveridee 1 93.S ing. 970 ( ) Northern Territory—SAM, R14.VU, AttMnide (!9filr Moore t Parser 1 1940). SloVXn River; NMV. DI2702. Harrow Ck. NMV. D8307, DXJI5. DS327. QM. J17K5. 2985, SAM. FROGS OF THE CYCLORANA AVSTRALIS COMPLEX 95 20mm 20mm Fig. 2. A. Cycloruna uustralis from Kununurra, W.A. B. Cyciorana tiovaeholliwdiue, 15 km N of Goondiwindi, Qld. R8968, Darwin; SAM, RI3453, Elsey; SAM. Description: The diagnostic characters of this R 13450, Howard Springs; WAM, R 1935-36. species are: size large, males 70.8-78.0 mm R21318, SAM. R14330-31. Katherine; SAM, and 71.0-81.0 in R4877, Mt Bundy Stn; SAM, RI3349 A-G, Smith females mm snout to vent Pt, Coburg Peninsula; WAM. R24007. Snake Ck; length; S-V/HW ratio high (mean 2.31); NMV, DI2704-08, SAM, R13275 A-L. Tennant head width only slightly greater than head Creek; NTM, 498, 525-26. 50 km of Tennant N length (mean HL/HW ratio 0.89); TL/S-V Creek. Queensland—SAM, R5010, R5070. Doom- ratio moderate 0.32 ; adgee Stn; NMV, D8437-38, SAM, R4934, Morn- (mean ) foot relatively ington I. long (F/S-V mean 0.40). 96 M. I. IYII.R & A. A. MARTIN Development and exostosis of the super- ficial skull bones are moderate in this species. The dorsal limit of the squamosal is such that I he re is a very hroad gap between the squamo- sal and the frontoparietal. On the fronto- parietal exostosis is confined to the lateral bor- 10mm ders of the bone. The sub-orbital portion of (he maxilla slopes steeply to the labial margin and is not expanded there into a lateral ridge. Cyclvtuna australis is usually pale olive or grey in preservative and bears a narrow and very sharply demarcated dark brown rostral hnr and a narrow sub-ocular bar which ter- minates far above the labial margin. The lateral body surfaces are commonly heavily .suffused with darker pigment. The backs of the thighs arc darker and densely variegated with light pigment. Geographic variation: The presence of darker irregular patches on the dorsum vanes through the range. Dorsal spots are absent from a series 1mm over 100 specimens from kununurra. Speci- of Fig. 3. Tadpole and tadpole mouthparis ui mens from Tcnnant Creek have light suffusions Cyctoruna ausiralh. of pigment, and those from the north-eastern portion of the range are heavily pigmented Cole in February. 1 963. The following notes with dark stippling. Immaculate and marked are based on four specimens from this series specimens occur on Mornington Tsland. at stages 36-3*5. All specimens are poorly pre- Variation in of the pertinent body pro- some served and badly distorted, so that their total portions is summarized in Table I. length range of 50-65 mm is only an approxi- bggs; A gravid female from Kununurra con- mation. tained approximately 1000 eggs varying from The spiracle is sinistral and the anus median 1.1 to 1.3 mm in diameter. The eges have or very slightly displaced dextrally. The over black animal poles. all appearance (Fig. 3) is similar to that of ('. fMrval morphology: A series at tadpoles was cttlrt'ipts and f . phvycL-phalus (Watson & Mar- obtained at Kununurra on the Ord River by K. tin 1973). TABLE I Geographic variation in prono) lions of Cyclorana species [Ranges are ^ivet) with means and standard deviations n> porenthews) Sf/et h's und Ii>iiii't1\ HL'UW TL/S-V S-V.1HW l/TL FS\ C. aiistniliv kiinumirru. W.A. 7 0.81 - 0.93 0.3K 0.45 2.13 — 2.33 0.9 1 1.00 0.38- 0.44 (0.86 _!_ 0.04) (0.42 ±0.03) (2.23 ± 0,09 J (0.97 ±0.03) (0.41 ±0.02) Smith Point, N.T. 7 0.K9 0.93 0.40 0.45 2.27 2A4 0.92 - 0.98 0.39 - 0,4} (0.91 ± 0.0 1) (0.43 ±0.02) <2 .36 ±0.06) (0.95 ±0.02) (0.41 ±001) lennant Greek. N T- 7 0.87 0.94 0.40 — 0.46 2.25 2.49 0.91 - 0.98 0,38 - 0.42 (0.90 ±0.02) (0.42±n.o:'> 12.36- ±0.08) (0,94 ±0.02) (0.39 ±0.01) Milctiell Pliileiui, W.A. 4 0.95 — 0.99 0.44 — 0..MJ 2.26 - 2.51 0.88 - 1.05 0.38 0.43 (0.97 ±0.01) (0.46 ±0.02) (2.41 ±0.U) (0.95 ± 0.07) (0.41 ±0.01) C\ nayavliidlundiuc - — Cooktown. QUI 7 0.8 1 0.8* 039 U.43 I.V3 2.08 0.87 0.96 0.36 0.41 (0.83 ±0.02) (0.41 sfcJXQl) (2.02 ±0.04) (0.91 ±0.03) (0.38 ±0.01) Calliope, Qld J 0.82 — 0.KK 0.39 — 0.40 USA - 2.17 0.R9 - 0-94 0.34 0.39 (0,86 ±0.03) (0.39 ±0.01) (2.12 ±0.01) (0.92 ±0.02) (0.36 ±0.01) Cunamutlu. Old 5 0.78 — 0.85 0.34 0.41 1,97 - 2.13 0.93 102 0.32 0.39 (0.82 ±0.03) (0.37 ±0.02) (2.05 ±0.06) (0.96 ±0.03) (0.36 ±0.03) I- RUGS OF IHh craMH4S>\ .tVSTRAUS COMPLEX 9? The mouth is subicmunal (Tit;. 3) with a TL.'S-V ratio rather tew (mean 0.40); foot large homy beak and papillae around the sides short Pftractojus ulutuvcHS PtWn l L#G7h goWeo and suffused with dark brown laterally ChirohpU's auairaits, Bou lenger i 1882) ipani. and inferiorty. The posterior surfaces of the Phravtops uuMnttis, Lovehdj;? (IM35J- Thighs were leaden jircy, whilst the ventral sur- CvcJoranu attxtruiis Paikcr (] Cecil Plains; D13049. 1 1525-24. NMV. SAM, R conclude initially that Iwo species were in- Opokbown; QM, !20*8V4I h km W &f rc>oUuv.ri. volved. Examination of larger scries, however, SAM, R%«J0. F.dwurd River Stn; QM. I2I84-H5. JI2M4. Eidsvold; OM, J 14383-84, tlilruth Plains has revealed the occurrence of Jotnw of inter- Cunnnnuilla; MUZD, S6-58/7Q. 75/70. 9 & 15 km mediate appearance, The variation may be Goondiwimli; E of QM. J56I1-I2. Mackav: SAM. summarized as follows. All specimens front R474?, Mupoun Mission Sin; SAM, R9734. Mtuy central and southern Queensland have high River; QM. J 14 159-67, Mitchell R. Miwion: SAM. KI04I9, Prcstwood, Gilbert River; SAM. JO 640, skulls with a gently sloping maxilla and a Rockhnrnpion; QM. 110482, SAM, R3686, Si densely pigmented dorsum. There is sinking George: .12186.89. Slannarv Hills: QM. SAM, variation in individuals from northern Queens- RW5, Stewart River; SAM. R9"S9l, Strathgordnn land, Some are densely pigmented whilst other* R.S . QM, 12222-7-2*. Surat. NMV. D7542, QM. 14644, Townsville: QM, .13480, Vieto, Cooncoota; are immaculate. The skull of the immaculate IJM. J 18063-65. Waratah Sin, Cunnamulla. individuals is either similar to that of the pig- Ot' n 1 1 ption' Snout to vent length ot mate* mented frogs, or is spatulate and distinctly rVM-SM mm. females 74.8-101,2 mm; head flattened. Unfortunately we have been unable noticeably hroadcr than long (HL/MW mean lo devise n mean* of objectively estimating 0.X3 1 : S -V.' HI. ratio low (mean 2.05 ). skull depth with any degree of accuracy We 9H M. 1. TV I ^K & A A. MAR11N hold the opinion that the high and the spatu- Discussion laic terms of the skull repiesenl different evo- The morphological complexity of C. novae- lutionary trends of development. hoUatidiue as defined here is unparalleled Whereas ihe extremes are dearly dilfeTen'., amongst Australian anurans. We believe that assessment of the significance of the observed a study of species isolating mechanisms, such variation rs complicated by the existence in as male mating call, in jjorth-etwtcnt Australia northern Queensland of a number of inter- could reveal ihe existence of two or possibly nediale fornix that cannnt he referred to either even throe species. Our action of resurrecting form. In addition there aie animals in which C. novtwhoUatuimt- from ihe synonymy of C- the terminal portion of the skull is more elon- ausftalis is therefore only the first step towards gated. This variant occurs only on the Cape an understanding nf Ihe C tmuratis complex. York Peninsula and at localities at the hase of The biological data, e.g. mating call structure. the Gulf of Carpentaria. Morphometric data of that are necessary for final resolution of the iwo small series are .summarised in Table I. problem may be extremely difficult to obtain. BggS: Oviducal eggs of three gravid females The northern Queensland populations arc ranged from 3 1 to 1,3 mrn in diameter, An apparently opportunistic breeders which may estimate ot the number in unc individual call at a locality on only one or two nights exceeded 1.000. The eggs have black animal each year (C. Tanner, pers, conim.). It seems poles. justifiable* therefore, to treat ihe complex ai Mating Call: The mating calls of two indi- this preliminary level, viduals fcJMj/dcd 5 Km S\V of Calliope. Queens- In terms of skull structure. C. attxtraJh rs land, 1 8.5.1 very on 970. are similar to those clearly the most simplified and primitive mem of ouxiralis. ho\n< laml C. The were calling on her of the complex, exhibiting limited develop- beside a rain-filled roadside ditch, with wet- a ment of skull bones and the least extensive bulb air temperature of 24.6*C. The spectral exostosis. All the variations in the foTm of the structure the is of calls of the two species skull and exostosis of the cranial bones such essentially identical, with novaahoHaiuiiitt' C. as the maxillary and squamosal m C fu>va?~ having also emphasized harmonic bands at hollantiiae can reasonably he derived from C\ about 600, its 800, and 1.000 Hz. However, australi.s. call duration is considerably longer (mean 249 of C. primi- m&ec: range 235—262 rmee). Judged on the The concept uusirahs being the tive mem her may be acceptable morphologi- hosts of the levels of difference in mating call cally, it is difficult structure nf sympatrie anuran spe.ies, this but more to conceive fcftog&fe- graphically in view of the absence of difference in duration does not represent diver- any mem- ber of the Cytloratm in gence of sulTjcicnl magnitude to achieve repro genus New Guinea. The geographic area occupied by ihe duciive isolalion. Hence if C. aust talis and members the C. aust/alh complex, includes hieh C. novaeholiatulioe occur hi sympalry (as they Of both rainfall and relatively arid areas: i.e. it may in the Gulf District) we would expect docs uot appear to be limited climatically. Thus if them ft) hybridi/.c. A .similar pal tern of the complex originated in northern or north- marked morphological differentiation, accom- western Australia it is surprising that it should panied by very little mating call divergence, (1V72) characterizes the Weslern Australian IJmnody- be absent from New Guinea, Jennings estimates that the most recent land communi- uastvs rfonaU.s and the eastern L. probable, iherefore, that the complex is uf rnosi readily from a rhomboid. That such a considerable antiquity. Two further circum- structure occurs in living Cyclorana is con- stances lend support to this suggestion. One is sistent With our hypothesis of its antiquity. tfac affinity of Cyclorana wilh the Hyhdae demonstrated by Tyler (1972) and Watson Acknowledgment* $ Murtin (J 973). The other stems from our observations on the pupiL shape of C. novae- For the donation of specimens or the loan hoflandiae. As stated above, the constricted of those in their care we are indebted to Mr K. Coventry, pupil of C. novae hoilandlae is almost rhom- Cole. Miss J. Covacevich. Mr J. Mr D. Howe, B. S. Low. Dr G. Storr, C. boid. In fact, the ventral margin is an obtuse Dr Mr )•*. angle and the upper a broad curve. This Tanner and Dr G. Ycstcrgren Mr G. Wat- son assisted in l he Meld. Photographs were pro- curvature is difficult to detect during extreme of constriction. Lynch (1971) considers the Ver- vided by Mr B N. Douetil and the tape the auxttatis tically orientated pupil to be the primitive and mating call of C. by Dr B Low and the horizontal pupil the derived state. Kow- Dr D. Gartside. cvcj. from the fact that Nvctimysres (a genus Our thanks are also due to Oi J. Ling tor that can only be derived from Litoriu. a hori- reading the manuscript, and to the Trustees of zontally-pupilicd stock) has a vertical pupil, it the Science and Industry Endowment Fund is clear thai, vertical orientation can be a for providing travel funds to M J.7. permitting derived State The trend to one or other orien- visits to various Australian museums ro tation of pupil shape could be accomplished examine collections. References — Boui enokr, Qj A, (, 1882). "Catalogue of the MooRr, i. A. (.1954). —Geographic and genetic Batraehia Salient iu 8. Ecandata in the collec- moThJiuu in Australian Amphibia. Anur. Nal. tion of the British Museum." 2nd Htin. »8> 65-74- (London > Parklr, U. W- 11940). -The Australasian frou-s U*ArrsruoM, B. H. ( L970). —Thermal acclima- of lbs tamily LeplodactyJidae. A'oWr Zool tion in Australian amphibians. Camp. Hio- 42, J-IOG. ehent, Physiol. 35, (S9-I03. Pfcrfcjts, VV. (1867).—Heip2loloi?isctie notizen. Krv. D. B. < 1914).- -On a collection of reptiles Afofiarsb. K. PreiiM- Akuci. Ifm. tier (in, and batrachians from Western Australia. /*»t. 1S67, 13-37 W. A ust. Mas. 1, 174-210. Sllvin, J. R. and Barrier: The natural and cultural his- of the genus Mixoplnts I Aouta: LeptOdacft- tory of Torres Strait"- Dept of Biogeotsranhy lidae). Proc Linn. Sac. N.5.W. SQlf), 52-5v. and Geomorphologv. Pubh I3C. 3, Australian Stkaoch^.1. K. & NHiN, A R. (I9fi6).— National University, Canberra. Specintion and polymorphism in the gcrtuS LintEjOHN, (1957). new spacies of M. L —A Criuia Tbchudi (Anura; Leptodactvlidac ) in frog the Cr'tnia. W. Aust. Nat. of genus 6, OueenslancL Prat: /?. Sot: QUI 78U.L H-3& 18-23. IvrrR. M i Superficial mandibular LovrjtiDGE, A. (1935) Austrian Amphihm jn U97$}<— — muscuJalurc. vocal «ac« and the phytogeny of the Museum of Comparative Zoology, Cam- Anstralo-Papuan l^plodactvlid nxies. Rev. S. bridge. Massachusetts. Hull, Mas, Camp, Aast. Mus, 16(9), 1-20. ZooL 78, 1-60. Tyllu, M. L, St Parkhr, K (1974). -New Hpecie^ I vnch, J. D. I 1971).—Kvolutionary relation ships, osteology, and zoogeography of lepto* ot hylid and leptodactylid frogs from southern dactylic! frogs. Misc. Pahl. Mas .Vu/. Wsi. New Cjuiucu. Trans. R. Sue- 5. Aast, V8(2). Vni'v K Maki'IN, A. A. 0972),—Studhs in Australian Watson, G. F.. Sl Martin. A. A, < 1973).—Life Amphihia III. Tte Ijmnmivnanu's dorsalis history, larval morphuloyy and relaliotwhips complex (Amua. Lcptodacivljdac) Aa\i. /, of Aitttraliftn Icptodfictvlid froas. 'thing. H 7,ool 20, 165 211. Sac.fr Aast. 97(1). 33-15. ) VOL. 99, PART 3 30 AUGUST, 1975 TRANSACTIONS OF THE ROYAL SOCIETY OF SOUTH AUSTRALIA INCORPORATED CONTENTS 1 Mahoney, J. A. The Identity and Status of Thomas "Lectotype" of Leporillus apicalis (Gould, 1853) [Rodentia: Muridae] . . . .101 Barker, S. Revision of the Genus Astraeus Laporte & Gory (Coleoptera: Buprestidae 105 Beck, R. G. Factors affecting the Distribution of the Leptodactylid Frog Geo- crinia laevis in the South-East of South Australia - - - 143 Edelstein, T. and H. B. S. Womersley The Thallus and Spore Development of Lobospira bicuspidata Areschoug (Dictyotales: Phaeophyta) - 149 PUBLISHED AND SOLD AT THE SOCIETY'S ROOMS STATE LIBRARY BUILDING NORTH TERRACE, ADELAIDE, S.A, 5000 THE IDENTITY AND STATUS OF THOMAS "LECTOTYPE" OF LEPORILLUS APICALIS (GOULD, 1853) [RODENTIA: MURIDAE] BY J. A. MAHONEY* Summary MAHONEY, J. A. (1975) .-The identity and status of Thomas' "lectotype" of Leporillus apicalis (Gould, 1853) [Rodentia: Muridae]. Trans. R. Soc. S. Aust. 99(3), 101-104, 30 August 1975. The specimen selected by Thomas as the lectotype of Leporillus apicalis (Gould, 1853) was misidentified by him and belongs to Leporillus conditor (Sturt, 1848). It does not belong to the type series of L. apicalis, therefore Thomas' lectotype selection for that species is invalid. The type material of L. apicalis and L. conditor is missing. Thomas' "lectotype" of L. apicalis and a second specimen of L. conditor in the British Museum (Natural History) could belong to the type series of L. conditor. Evidence for the occurrence of L. apicalis in Tasmania is lacking. THE IDENTITY AND STATUS OF THOMAS 1 "LECTOTYPE" OF LEPORfLLUS APICALIS (GOULD, 1853) (RODENTIA: MUR1DAE] by J. A. Mahoney* Summary MaHONEY, J. A. (1975).—The identity and status of Thomas' "lectotype" of Leporillus apicalis (Gould, 1853) [Rodentia: Muridael. Trans, R, Soc t S, Atut, 99(3*, 101-104, 30 August 1975. The specimen selected by Thomas as the; lectotype of Leporillus apicalis (Gould. 3 853) was misidentiftsd by him and belongs to Ltparilhts conditor (Sturt. J 848). ft does not belong to the lype series of L. apicalis, therefore Thomas' lectotype selection for that species is invalid. 1 The type material of f.. apicalis and L. conditor is missing, Thomas "lectotype" of L. apicalis and a second specimen of L. conditor in the British Museum (Natural History) could belong to the type series of L, conditor, Evidence for the occurrence of L. apicalis in Tasmania is lacking. introduction —the worst of the two, young, and with an The name Hapalotis apicalis was proposed imperfect tail. Thomas concluded with the by Gould (1853a) for a new species of rodent remark that probably from memory, and cer- from Australia. He did not stale in the original tainly wrongly, Gould stated that the species description if one or more specimens were had a white-tipped tail, but his overlooked heing described nor did he give any locality. second specimen [adult with nearly perfect Later, Gould (1853b) stated that he possessed skull- and quite perfect tail (BM, 53.10.22.14)1 has the latter organ uniformly blackish it single example procured by Mr Strange in or South Australia, and he illustrated the external brownish above and dull white below, and features, there is no indication of the white tail-tip Thomas (1906a) nominated Hapalotis api- found in so many Australasian Murtdae. calis as the type species of a new genus. A study of three Australian Museum speci- Leporillus, and subsequently (Thomas 1921a) mens of L. apicalis, and (he literature, enabled he selected British Museum (Natural History) Troughton (J 923) to confirm that Gould was 1 specimen 1 853.1 0.22.15 as the lectotype of correct in attributing a white tail-tip to the 1 LepoyUlus apicalis, describing it as a female species. Troughton stated also that Thomas hum S. Australia. Explaining his lectotype remark thai Gould seems to have done his selection, Thomas (1921c) stated that describing from only one of his two specimens although Gould had in his collection two speci- means that that specimen must be accepted as mens of that species, he seems to have done the holotypc. hi* describing from only one of them (BM, Tale (1951) treated specimens 1*53,10.27.14 53.10.22.14 (sici—lapsus for BM. 53.10.22.15) and 1853.10.22.15 as **cotypes" of L. apicalis '•' Department of Geology and Geophysics, University of Sydney, Sydney. N.S.W. 2006. 1 The first rwo digits of British Museum (Natural History) registration numbers or mammals are fre- quently omitted from publications. Thus Thomas uses 53.10.22.15 for 1853.10.22,15 "This skull is registered as 1854.10.21.1 and the Register entry mentions a stuffed specimen and relen lo 53.10.22.16. 1 have been unable to find a specimen numbered 1853-10.22.16 in tbe British Museum (N.H.) therefore 1 am following Thomas' conclusion that IS53.10.22.I4, 1*53.10.22.1 6 and 1854.10.21. J belong to the one individual; but it is possible that 185V 10,22.14, identified as Hapalotis apicalis in the Register, is jost and the skin novv numbered 1853.10.22. 14 is skin 1853.10.22.16 with an incorrect number. A note in Thomas' handwriting attached to skin IK53. 10.22.14 and stating that this specimen was considered to be the type seems \o refer to the Museum Register where "type"' has been written opposite the number 1853.10.22.16. A portion of the posterior half oi' the cranium, and the left mandibular ramus, are missing from the skull. 102 J. A. MAHONF.Y TAB Lb 1 1844-6 Expedition to Central Australia, the Skull measurements (in mm) of Thomas' "Iccto- original description was published in the t\>ve" of Leporillus ypieyli* (Cmuldl 8.M (N Wj, Narrative of the Expedition and Sturt did not 185 J. 1 0-22.1 5. say where his material was deposited. It seem*. Maximum width acio .s na sals 5.11 likely however thai at least one specimen. Minimum width across right zygo- illustrated by J. Gould and H. C. Richter in matte plate • t 4,4 a plate* accompanying Sturt's description of Length of tight M 1 -fl _ 9.7 the species, would have gone into Gould's col- Width of left Mi - 2.6 lection, perhaps from there into a t and Museum Width of left Ivt^ • 2.7 collection. A collection of mammals made by Width of lcftM» • • 24 the Expedition was presented to the British Length of left Mj- i - 9.1 Museum by Sturt in 1846. This collection is Widthof teftMj . . %& noted by Thomas (1906b) and docs not con- Width DricflM... . • - t 2.7 tain specimens of LepitrUlus. Width of left M A 2.4 Specimcus 1853. JO.22.I4 and 1853.10.22.15 were registered in the British The teeth measurements ate fur the crowns of the Museum on Octo- teeth ber 22nd, 1853, and were acquired from Gould. Labels attached to them refer to S. and referred to them as aduit and young Australia and F. Strange. The entries for them 1 females, from "South Australia ", collected by in the Register mention neither a locality nor F. Strange. He briefly described the skin ami Strange. S. Australia could be an abbreviation skull of each and. recorded measurements of of either South Australia or Southern Australia them. He does not refer, in hi.s account of L. and F. Strange presumably is Frederick ttpkatis, to Thomas' Jectotype selection or to Strange, a collector and dealer in natural his- Troughton's recognition of a hoiotype for the tory specimens who accompanied Sturt on >pccjes. some of his early surveys (but not the 1844-6 Expedition), and was an eaily settler in South Identity of the "leclotypc" Australia and later, in the 1840's, a resident British Museum (N.H.) specimens of New South Wales (Whittell 1947). Gould 1853.I0.22.J4 and 1853.10.22.15 are examples (1849) docs not mention Strange and Soulh of Lepohtlus conditor (Start, 1848) and not Australia in his account of L condiU*r. Subse- specimens of Leporttlus dpicalh (Gould, 1853) quently (Gould 1863) he gives only the as believed by Thomas, They do not agree with interior of New South Wales and Victoria as the original description of L. apicalh (Gould) localities for the species. It is possible that the and because of this 1 do not accept that either inscriptions on the labels are interpretation* of of them belong 10 the type series of that the origins of the two specimens based on species. Consequently, I regard as invalid Gould's account ot L. upica/tK. If they are riot Thomas' leelotype selection fur L. apicalis, interpretations, their significance is uncertain 1 since the citation of S. Australia is Thomas ( 1921b ) stated there is no specimen ambiguous of L. conditor in the British Museum, and and Strange might not be the collector of the tajther demonstrated his unfamiliarity with its specimen*. characters by suggesting that it possibly British Museum (N.H.) specimens belongs fo Notomys, a genus of Australian 1853.10.22.14 and 1853.10.22.15 could have hopping mice and rats. been collected on SiutVs JK44 ft Expedition Measurements ol the badly damaged skull of and might belong to the type series of L, condi- Thomas' "lectotype" arc given in Table L The tor. This is so even if they came from .South skull is illustrated in Figs 1-4. Australia. Sturt L. venditor is Jost. I he latter species was by Mr Browne and to one. a male, obtained described by Stun in IS4S as Mus conditor by himself from a native. However, Sturt (Vol from specimens observed and collected on hi* 2, Appendix, p 4) noted also that the last nest * Gould's name is printed on this plate and the Bpsciea name Mas conditor is attributed to him by Sturt, acveruVJess Sturt is the author of the name Mux conditor. THOMAS' "LECTOTYPE" OF LEPORILLUS APICALIS 103 iiimimiiiiiiimmiii Figs 1-4. Leporillus conditor (Sturt, 1848). British Museum (N.H.) 1853.10.22.15. Thomas' "lecto- type" of Leporillus apicalis (Gould, 1853). Fig. 1 —Ventral view of cranium (x3). Fig. 2 —Occlusal view of left upper molar row (x8). Fig. 3 —Occlusal view of left lower molar row (x8). Fig. 4—Right lateral view of cranium (x3). of L. conditor was found on the bank of the known, there is no uncertainty about which muddy lagoon to the north of the Pine Forest species of native rodents were named Hapalotis (N.S.W.), and the Expedition explored por- apicalis and Mus conditor by Gould and Sturt tion of South Australia before reaching the respectively, and neotype selections for them muddy lagoon. are unwarranted. Although the whereabouts of the type Gould (1853b) commented in his account material of L. apicalis and L. conditor is un- of L. apicalis that an animal in spirits in the 1 ) 104 J. A. MAHONfcY British Museum, presented by R. C, Gunn, Rattus lutreotux (Gray, 1841 ) ( 1845.5.2.3, from Van Diemcn's Land, accords very closely 1 852.1. 15. 16. 1852.1.15.17), Maswcomys with it in the colouring of the fur and in the fuscus Thomas, 1882 (1852.1.15.15) and rat-like form uf the tail. He added that it is of Pseudomys hifigmsi (Trouessart. 1897 much smaller size than L, apicatis and in all ( IX52J.15.IK). None of these are L. apicalis, probability will prove to be a new .species. and evidence for the occurrence of this species Gould's listing in 1 863 of Van Diemen's Land in Tasmania is lacking. as a possible locality for L. ttpicalis could be based on that material. Tasmantan rodent specimens in the British Museum (N.H.) and Acknowledgments attributable to Gunn are recorded m the Regis- This study was carried out in the British ter. The identities of these specimens and their Museum (N.H ) by arrangement with Or registration numbers are Rattus rttttus G. B. Corbel of the Mammal Section. The (Linnaeus, 1758) < 1837.6.10.56). Rattus nor- photographs were taken by Mr F. Greenaway, ve&cus (Berkenhout. 17b9) ( 1838.1.15.17), British Museum (N.H.) Photographic Unit. — References (Jouid. 3. ( 1849). "The mammals of Australia". foreign literature, .science, and the fine arts), Pi 2, pi. 8 and text (1863, vol. 3, pi. 6 and no. 1101. December 2nd, 1848. p 1207. eol. text). (J. Gould: london.) 2, and in a list of "Publications Received" in (iot'io, J. (1853a).—Remarks on the genus 1 he Spr* tutor, for the week ending Deeemher Hupalotis. Proc. zoot. Soe. Loud. 1X51, 126- 23rd, 1848. no. 1069. p. 1237. col. 1. Furthei- 127. [The approximate date of publication of more. p. 1243, col. 2, of The Spectator, for this work in 1853 is 29 April—see Prac. zoot. the wctk endine December 23rd. 1848. con- Sac. Lorut. 107, 81 (J 93 7). I tains a publisher's notice which states that Goui.o, J. (1853b).—"The mammals of Aus- Sturls work is "Now ready". tralia". Pt 5, pi. 12 and text (1863, vol. 3, pi. Tate, G. H. H. (1951).—Results of the Arch- 2 and text). (J. Gould: London.) [The date bold Kxpedilions. No. 65. The rodents of November 1st, 1853 is printed on the cover of Australia and New Guinea. Ball. Am. Mas. Pi 5 of Gould's work.l nat. Hist, 97, 183-430. Gould. J. (1863).—'The mammals of Australia". Thomas, O. (1906a).—On the generic arrange- Pt 13, Introduction, pp. xi-xl (1863, vol. 1, ment of the Australian rats hitherto referred Introduction, pp. xi-xl). (.1. Gould: London. ) to C'onilurus, with remarks on the structure (The date May 1st, 1863 is printed on the and evolution of their molar cusps. Ann. cover of Pt 13 of Gould's work. An uncor- Mas'- mn Hist. (7) 17, 81-85. rected proof-sheet version of Pt 13, pp. xi-xl Thomas, O. ( 1906b).—Mammals. Pp. 3-66. in of "The mammals of Australia'' is included "The history of the collections contained in in pp. 1-51 of a work entitled "An introduc- the Natural History Departments of the tion to the mammals of Australia" hy 3. British Museum". Vol. 2. Separate historical Gould and published in 1863. A copy of this accounts of the several collections included work in the Australian Museum library has in the Department of /oology. (British a date of presentation to Reverend John Museum: London.) Barlow L.R.S., May 6lh, 1863, inscribed on Thomas. O. (1921a).— Notes on Australasian the title page.1 rats, with a selection of lectotypes of Aus- Sti'itr, C. (1848).—"Narrative of an expedition tralasian Muridae. Ann Man. not. Hist. (9) into Central Australia, performed under the 8. 425-433. authority of Her Majesty's Government, dur- Ihomas, O. (1921b). —Notes on the species of ing the years I844 T 5. and 6. Together with Notomxs, the Australian jerboa-rats. Ann. a notice of the Province of South Australia, Mag. Hat. Hist. (9) S, 536-541. in 1847." (Boone. London.) Vol. 1, X. iv, Thomas, O. (1921c).—On three new Australian 5-416 pp. Vol. 2, vi, 1-308, 1-92 pp. [The rats. Ann, Afa&. not. fttst. (9) 8. 618-622. final 27 pages, i.e. pp. 66-92, of Vol. 2 are TbcjGhton, E. Le G. (1923). — A revision of the a botanical appendix by R. Brown.) [Although rats of the genus Lcporittus and the status ot this work has the date 1849 printed on the Htipalotis per.soitata Krefft. Rrc. Attst. Mus. title pages, Us date of publication is 1848 14, 23-4L Thus it is cited in a "List of New Books" in Whiiteil, H. M. (1947).—Frederick Strange. A The Atttennentn (Jnuttud of English and bioKiaphv. Attst. Zoot. II, ^6-114. REVISION OF THE GENUS ASTRAEUS LAPORTE & GORY (COLEOPTERA: BUPRESTIDAE) by S. Barker* Summary BARKER, S. (1975). -Revision of the genus Astraeus LaPorte & Gory (Coleoptera: Buprestidae). Trans. R. Soc. S. Aust. 99(3), 105-141, 30 August, 1975. A revision is presented of the Australian Buprestid genus Astraeus LaPorte & Gory. Following van de Poll (1889), but extending the status given by him to two separate groups within the genus, Iwo new sub-genera are proposed, Astraeus (Depollus) and Astraeus (sensu stricto). The genus is considered to contain 39 valid species. The species are keyed and a complete description or redescription is given of each, together with an illustration of all species not previously figured in a publication; an outline diagram is given of the male genitalia of 36 of the 39 species. Sixteen new species {aclamsi, bakeri, carnabyi, carteri, declariensis, fraseriensis, globosus, goerlingi, macmillanu minutus, obscurus, polli, robustus, smythi, tamminensisy and watsoni) are described. Two names, major Blackburn and mastersi MacLeay, previously regarded as synonyms are revalidated. Two names previously regarded as synonyms are confirmed as such. They are as follows: meyricki Blackburn (of badeni van de Poll) and tepperi Blackburn (of jansoni van de Poll). Three names are newly synonymised (the synonym first): splendens van de Poll = mastersi MacLeay; simplex Blackburn = mastersi MacLeay; and strandi Obenberger = dilutipes van de Poll. REVISION OF THE GENUS ASTRAEUS LAPORTE & C20RV (COLEOPTERA; BUPRESTIDAE) by S. Barker* Summary Barker, S. ( 1975).— Revision o£ the genus Axtraeux f.aPorle & Gory (Coteopterui Buprestidae). Jrans. R. Soc. $. Aust, 99(3 ), 105-141, 30 August, 1975. A revision is presented of the Australian Buprestid genus Axtraenx I a Porte & Gory. Following van de Poll (1889), but extending the status given by hinj to two separate groups within the genus, two new sub-genera are proposed, Axtraenx {Depoltux) and Axtraenx {sensu strtcto). The genus is considered to contain 39 valid species. The species are keyed and a complete description or redescription is given of each, together with an illustration of all species not previously figured in a publication; an outline diagram is given of the male genitalia of 36 of the 39 species. Sixteen new species (adamsi, bakeri, varnubyi, carter/, dedariensis, fraseriensis, glohosus, goerlingi, macmiUani, mlnuiux, ohseurux, palli, rahuxtux, stnytht, tammi- nensis t and watsom) are described. Two names, major Blackburn and masters! MacLcay, previously regarded as synonyms are revalidated. Two names previously regarded as synonyms are confirmed as such. They are as follows: meyricki Blackburn (of hiidrni van de Poll) and tcppt'ri Blackburn (of junsoni van de Poll), Three names are newly synonymmed (the synonym first); splendent van dc Poll • maxtcrxi MacLcay; simplex Blackburn = muxterst MacLeay; and xtrandi Obenberger = dihuipes van de Poll. Introduction same result is achieved in a different way. It There are 27 described species of the genus has no doubt evolved as an escape mechanism. Astmeus from Australia and one from New One other feature has apparently not been Caledonia. Originally established as Asthraeus commented on in the literature and that is the by La Porte k Gory (1837). the genus was fact that most of the species exhibit classical referred to by that name until Gcmminger & warning colouration of yellow markings on a de Harold (1869) used Astraeux, thus correct- dark background. Whether any of the species ting an incorrect transliteration from the are noxious to predators or whether they arc Greek. This has been followed by all subse- Batesian mimics has yet to be determined. quent authors. Barker (1964) requested the Since van de Poll's excellent monograph, International Commission for Zoological further species have been described and more Nomenclature to validate the emendation of discovered so that the genus is again in need Asthraeus to Axtraeus and this was eventually of revision. This poses some difficulty to .in carried out (I.C.Z.K 1966, Opinion 795). The Australian worker as most of the type speci- genus was last, revised by van de Poll (1889) mens are housed in overseas museums. Alv>. who gave excellent descriptions and illustra- through habitat destruction throughout Aus- tions of all species known al lhat time. Pre- tralia, some of the species are not abundant sumably van de Poll never saw a live speci- over their former range, and unfortunately the men of the genus, so would have been genus is very poorly represented in museum*- unaware of their unique spring mechanism and in most private collections in this country. involving the release of the elytra from the The adults of more than half of the species closed position. When the living beetle releases occur on Casuarina sp., but some are also the .spring, the elytra flick open with such force associated with species of Jaeksonia, BanhsUi, that the insect can be projected for up to Ximthorrhoea, CalUtris, Melaleuca, Acacia, •vevcral metres. The effect is reminiscent of the Daviesia, and possibly Dryandra and Hakeu, sternal spring of the Elatcridae except that the Adults of the earfy emerging species can he * Department of /xiology. University of Adelaide, Adelaide. S. Aust. 5000. I Oft S RARKFR collected in August and adults of some species men. Axtraeux is placed in the sub-tribe Bup- are present as late as March, More than haJf restes with nine other genera found in Aus- of the known species are found in the southern tralia. Axirtteux and Neohupresiis both have an hfltf of Western Australia- The oihcrs occur indistinct scutellum, but Neobuprextis has an over the rest of the continent excluding the exposed pygidium whereas in Astravtix it. is desert and northern-most tropical areas. One covered hy the elytra. All of the other Austra- species occurs on New Caledonia. The genus lian members of the sub-tribe have a distinct is unknown from Tasmania. scutellum. They are: Naxciohifs, Nose in, Neo- In this paper male genitalia have not been bubaxies, Notobuhasrex, Bubaxtex, Bttprextinti. used to diagnose the species, but have been BupreMocles n\\d huryspitux (Carter 1929"). mod as one character to place the species into related groups. Taxonomy This revision is based mainly on collections Genus ASTRAEUS LaPurte A Gory made by the author in South Wales'. New Avhractts LaPortc & Gory, 1837: I, pi. 1. South Australia and Australia between Western fiy. 1. (Invalidated, Opinion 795.) Imhoff. 1962-1975, now housed in (be South Austra- 1 856; 46. Laoordaire. 1857: 43. Saunders, lian Museum. The remaining specimens are IK6S: 10. pi. 1, Jig. 12. vCaltered throughout the collections listed Axtiwux: Gemniinger & de Harold, 1869, St below. The abbreviations used in the text for 1380. Saunders, (871: 43. Masters, 1871: museum and private collections arc as follows; 124. MaeLeay, 1873: 239-240. Kerremans of l\. EA Collection Mr K. Adams, 1885: 136, van de Poll. 1886: 176-178. Mas- Edungalbtu Queensland ters. 1836: 71. van de Poll, 1889: 79-110. pis AM "I he Australian Museum. Sydney J, 3, figs 1-19. Blackburn. 1890: 1256-1258, ANIC The Australian National Insect Col- 1891: 496. van de Poll. 1892: 67-6$. Kerre- lection. C.S.I. R.O.. Canberra. man*. 1892: 103-102. Blackhurn. 1892; 211- BPBV1 The Bermce I* Bishop Museum. 213: 1894: 101: 1895: 45-46. Kerremans. Honolulu. I9Q0: 295-296: 1902: 148-149. PmlveL BM The British Museum (Natural His- 1904: 116. Carter. 1925: 229, fig. 1 Oben- tory J. London. bcrgcr. 1928: 204-205. Carter. 1929: 265. Collection Mrs. KC of Mr and K. Car- 2S2, 302, pi. 33, fig. 43, Obenbcrgcr, 1930: naby, Wilga, Wcslern Australia .'65-367. Carter, 193.3 : 41 . ObenbergeT. )H Collection of Mrs J. Harslett. 1936: 133. Barker, 1964: 306-307. T.C.Z.N.. Amiens, Queensland. 1966: 269-270. MM The MaeLeay Museum. Sydney Ait>e?ia; Carter. 1929; 270. (types on permanent loan to ANIC) Typa species. A stratus jitrsoplrsux LuPorte MNHN Museum National d'Histoire Gory, 1837 monotypy). Nalurelle, Paris. & (by NM The National Museum of Victoria. Generic description Melbourne. Head. Medium size or large; the surface NMP National Museum. Prague. pined; with a median longitudinal impressed QM The Queensland Museum, Brisbane- line on the basal halt; clypeus short, the apical SAM The South Australian Museum. Ade- margin wilh a bow-shaped indentation; labium laide, incised: aniennal cavities small and rounded, WADA The Western Australian Department situated near the lower internal corner of the of Agriculture, South Perth- eyes. WAM The Western Australian Museum- Antennae, The first, segment longer than any Perth. of Ihe rest, rounder and thicker a! the apex The Australian Buprcstidae have been than at the base; 2nd segment short and divided into six sub-tamilies ( Britton 1970). obconic; 3rd segment longer than the 2nd bet Affrttffus belongs in the sub-family Bupres- not as long as the first, also obconic; 4th seg- tinae. the members of which have mouthparts ment slightly shorter than the 3rd^ dorsoven- not produced downwards to form a rostrum, trally flattened, the front edge projecting out- In the sub-tribe Buprestes the metathoracic wards to form a tooth, the remaining segment*.. epimera are completely exposed whereas in except the last, similar in structure to the 4th the Anihaxiae they arc totally or pailially ami each sligbly smaller iban the succeeding covered by the lateral extension oj the abdo- segment; the last segment flattened but not Rh VISION OF BUPRESHD GfcNUS ASTKAEUS LAPORTfc A OORY 107 toothed. In species belonging to Astrarus Key to the sub-genera of Astraeus tsensu stricto) the antennae are sexually dimor- I. Lateral lobes of pronotum projecting from rbc phic, being longer in male* than in females. elytra t edge 'ioc; elytra st date-punctate but never with longitudinal costac; numeral fold due to the segments after the 3rd being of rounded . . Astracus {Depol(us) approximately equal length, whereas in females I. Lateral lobes of pronotum confluent with the they become progressively shorter. In Asttaeus elytra I ed^e line; elytra sLriate-punclaLe with well iDepdlltts} the antennae do not show sexual or poorly developed longitudinal costac; humeral fold angled (Fig. I) . ,,,. dimorphism and are short in both sexes. Astracu.s (siwu .stricto) Pronotum. Surface pilled; wider than lung; Sub-genus Depollus sub-gen. nov. dorsaJly convex; rounded at the sides and Tvpe species Aitraeus abefrath van dc Poll, narrower a! the apex than median lobe. I have called this structure the lateral lobes with the posterior margin curved basal crypt (Fig. 24 >. upwards and outwards. Elytra striate-punctate. the edges of the striae never costate. The Scutellurn. Invisible. suture ends posteriorly in a short or very short Elytra. Convex dorsal ly, each side with a spine which may be blunt or pointed and cruvcenl-shaped lobe fitting over the notches curved outwards. With no marginal spine or in the base of the pronotum; the apex ol each in one species with a very short, blunt maiginjl clylron ends (with rare exceptions) in a sharp spine. The humeral fold A rounded and poorly point, the sutural spine, usually lonned by a developed. Only known to occur in Western crescent-shaped lateral emargination which Australia. lormx on the inner edge a smaller lateral to the speck* Astracus (Dcpollus) tooth, usually called the marginal spine; it' the Key of 1. Basal colour brown sp. flow, lateral enwginalion is absent, so is ihc mar- .,1. pwt I Basal colour black ... 2 ginal spine; below the shoulder the external 2. Elytra with 4 longitudinal red vittaa margin is dilated, sometimes the dilation is 5. iineatux van de Poll rounded posteriorly, sometimes siiongly pro- 2, BJytra without longitudinal red vitrnc -1 - jected backwards and angled forming a point. 3. Head, prunotum and uiidersiirfacc vciy hairy the whole being folded inwards and covering 4 1 Head, pronotum and undersurfa.ee not verv pan of the metastemal coxae—this structure 3 hairy 5 IS called the humeral fold I Fig. I); punctate- 4. Pronotum with clumps of hair emerging striate and in many cases also cosiate. there ft from depressions towards Ihe sides; no mar- a curved lip on the anterior undersurfnee of ginal spine 3. robustm sp. nov. each elytron which fits over an opposite curved 4. Pronotum with hair emerging singly, not projection on the mesothoras. together form- clumped; small marginal spine 4. akermwi van dc Pol) ing the catch of a spring escape mechanism. 5. Head and pronotum with deep punctures close 1} Undersurface. Variably pitted; anterior edge packed together 2. tammhtctts'u sp. nov, and with ol presternum straight or crescent-shaped and 5 Head pronotum shallow, separated punctures ,,.. .,....., ,6 a large prostcrnal process, mesostcrnum curving , 8. detlortettsis sp. ro>v- gressively shorter- : \sh.u-iis (Pepullus) polll sp. nov. Body outline. Laterally teardrop-shaped, the apex being constricted; dorsally convex; ven- FIGS 2, 22A trally straight at the anterioi end and curving TVfH'S Holotype. <£ Wungan Hills, W. Ausi.. On upwards ai (he end of the ahdnnien. rarely Casuurina campesirls, 9.il97l. -V- fair for, concave. SAM. T 20940. - I ox S. BARKPR ;. 1 W. Aust. T 12.U948, A, Douglas, NM; A Oedari, W. Aust, WADA; 2 d\ South Tam- min Mora Reserve, W. Aust. on C. catnpes- tris\ 6.i.l97l & 9.ii.l971, S. Barker, WAM ( Colour. Shiny. Head brown with the following yellow markings: twr o lunettes from the inner corner of the eye. concave towards the base of the antennae but not reaching the middle of the head; behind each of the former at the base is a single small spot; a small spot at the base behind each eye; an elongate band runs from the anterior underside of the eye almost to the base; there is a large spot on each side of the gular; antennae brown. Pronotum brown with bfue reflections and the following yellow markings: close to the anterior margin on each side of the centre, but not reaching it is a laterally elongate mark; in the same line, directly behind the eye, is. a smaller laterally elongate spot: towards the base, in the middle of each side, there is a longitudinally elongate spot, and between them a small median longi- tudinally elongate spot; laterally there h a longitudinally elongate spot commencing just after the middle, running to the base, which is flared outwards with a dark rim and the exposed inner surface is yellow. Elytra brown with blue reflections: the intervals between the striae are irregularly mottled with yellow, more so at the base than at the apex; the sutures, apical spines, and the lateral and antcnoi margins of the elytra are black hut elsewhere brown. Undersurface brown; the mesoslerxuim and metasternum each have lateral yellow «—1mm patches and each of the abdominal segments have lateral yellow spots; the coxal plates each have a yellow spot near the centre but not Pig. I. Outline diagram of the left humeral fold touching it; hairs silver. Legs: femur and tibia, of 4 species of A.straetia. The head end of each is to the top of the page and the left brown with blue reflections: tarsi dark brown side in uppermost. Note that the specimens with blue reflections. figured are all females and that (a) and Paratypes: 1 £ & 1 gj 2 km \V of Wongan Hills, the striae slightly convex and transversely W. Aust, on C. campvstris, 20.U973, 5. Bar- strigose with shallow punctures; parallel-sided /r?r, EA; 5 & 4 9. Wongan Hills, W. Altai. o* until after Ibe middle then rounded to the on C\ vumpestris. 9.1.1971. 5. Barker, ANIC base; with a small sutural spine projecting 1 1 rf arid I ?).BM(1 ^& I j-V MNHN laterally. I Undersurface (1 i/Al 2). SAM (2 £ & I S); ft Deduri. with shallow punctures, REVISION OF RUPRESTID GENUS ASTRAEUS LAPORTli k GORY 10:1 denser at the sides tluu in the middle; with a sided but abruptly rounded at the opex. Elytra lew short hairs. with the intervals between the striae trans- punc- Size. Males 10.5 ± 0.15 x 3.8 dr 0.07 mm versely wrinkled, and deeply nnd densely 118). Females 11,5 * 0.27 v 4,0 ^0.09 mm tured; parallel-sided from base to behind the MS). middle then uniformly rounded in the area where marginal spines occur in the other Oisthbutiim, Western Ausl ralia. sub-genus, then straight sided to 1hc apical General remarks. Named after the late J. R. H. spine which is minute. Undeisurfaoe closely Nccrvort van de Poll. and evenly punctured; wilh a lew short bails. Spt.t Unt'n.s e.ifiiniuitl. W Ami.: Types; 5 rf & 3 fee. Male 11.3 x 4.1 mm (1>. ?. J3 km N of Guomalliny. Dd Casuaritia campey- im. 9J.I97I, S. Barker, I ,? & I ?, Waddirtgton, Distribution. Western Australia. on C. campextri*. 9,1.1971, J. farker; \ J & 2 £ Central remarks. This species shows closest Deduii. on C. campffitris, 20.xii.l972. 5. Uar- attinity with Attraeus polti. gvrj 3 rf # 3 9, 2 km W Of Wongan 9%, on C campest ris, 20.U973, A'. Barker. Specimens examined. Type only. 2, Astra? us (Dcpnllus) tamniinejiMs sp. nov. 3. Astracus (Depotlus) robustus sp. nov. MCiS 3, 22H FIGS 21, 2ZC Types, Holotypc: J, South T&ramiu Flora Reserve. W Holutype: I7.ix.t970, .S'. Barker, laterally a small round spot between the eye BM; I -rf & I ? t 385 km along Pavnc's Find Rd, W. AusL on C. and the base of the head; ventially all yellow, diehitvia. i7.ix.l970. 5. Barker, MNHN; t almost continuous under with iirst the eye the 6*. Payne's hind. W. An si. on C". diefsictna, mentioned spot; antennae dark brown with I7.ix.1970, 5. Barker, WAM. blue rellections. Pronotum black, bottoms ol Colour. Shiny. Hcatl. pronotum and elytra most of the punctures yellow, with ihe follow- black wtih purple reflections. Antennae black ing yellow markings; a thin median longitudinal with blue and purple reflections, Elylra with Jim; from base to apex covering all bin the tip the following yellow markings: a longitudinal of the median lobe; two longitudinal stripes on spot in the 2nd outermost interstice near the each side of the first, curving inwards from the shoulder and another just before the middle; a base but ending before the apex, each capped similar but larger spot lies on Ihe 5lh Outer- with a circular spot near the apex; a (bicker most interstice after the middle but not reach- lateral stripe runs from the base 10 the apex: ing ihe apex; a large round spot before the underside with alternate irregular stripes ol' middle with the width of the 3rd-6th inter- bJack and yellow, the yellow predominating. stice from the suture; a variable number of Elytra with alternate black and yellow longi- small spots at the base of the elytra in the tudinal stripes broken irregularly, iheir breadth middle, the maximum numher is four per side conforming lo the intervals between the striae; but they are absent in some specimens. Under- predominantly black along the suture and the suilaee black with purple reflections; hairs outer margin, yellow in the middle. Under* silver. Legs brown: tfie joint between the femur surface mainly yellow with irregular black and tibia black nn the upper surface; 1st tarsal markings along the middle of each side across segment brown at the base and black at the the outer margins of the abdominal sclerites apex as are Ihe remainder. anil at the apex, the black markinjra all with Shape and .sculpture. Head with, dense shallow brassy metallic gleams; hairs silver. Legs dark punctures; with a glabrous longitudinal median with brown blue reflections; each femur with line between the eyes confluent with the basal elongate yellow marking?; on both surfaces, impressed line; bairy Pronotum sparsely and 5/m/k? und sculpture. Head with dense, uni- shatlowly punctured in the middle where the foitnly deep purcture*- Pronoium wilh uni- depressions are small Hut greatly increase in formly dense punctures; more or less parallel- sue laterally where the non-depressed areas 1(1 S. BARKER * * I ID g ^ m 1 H 1 K£ VISION OF BUPREST1D GENUS ASTRAEUS LAPORTE ft GORY 1 J arts glabrous; sides gradually rounded from middle to the small marginal spine; suiural base to apex: hairy, with hairs emerging singly spine small. Undersurfaee evenly and shallowly in Ihe middle ;tnd in clumps from the lateral punctured; hairy depressions. Elytra with intervals between the Size. Males 14.0 ± 0.2 x 4.9 ± 0.0* mm virkie slightly convex; more or Ises parallel- (25V Females 16.6 ft 0.43 x 5.3 * 0.17 mm sided from base to just after the middle, then (11). gradually ihe apc\; with sharp rounded to Diwtbutioth Western Australia. sumial spines; hairy. Undersurfaee covered General remarks. This species shows closest whh punctures, shallow in die middle, deeper affinity with Astravus mhusutx and denser at the sides; hairy, the hairs on the Sprritnctw exutitined. A list 1 r£, Carnomah. prosiernul process and presternum noticeably W. 30ax.lS65, E. Baker; 12 1n(i sp , l7.ix.l970, $, Par Sfctf Males 15.9 ± 11,31 x 5.6 = 0.16 mm (4). ker; 2 o. 106 km S of Payne's Find on CttMutnKU l conwulaia^ 18.ix.l )70, S Barker; 2 ft Wialki on Females I 7.0 ± 0.90 x 6.0 ± 0,25 mm (3>. cortticatnfa, V)A\.V)10 S. Rurker; I Wialkt C. t & Distnbuthm. Western Australia. on Casaurtna campestris, 21.ix.l970, 5. Barker: I Specimen,? examined. Types only. rfi South Taninun Flora Reserve on C, unnptstris, JO.ix.WO. S. Barker; 3JS*?, South Tamtnin 4. Astraens (Dcpollus) aberrant; van de Poll, Flora Reserve on C\ twmpesrns, fell .1**70, S, 1 Barker; 3 J & 1 % Quairadlms on C. eampesirit, 1S86: 1 76; IRSY: R4, 90, 91, pi. 2, fig. 7.xr*.l970, 5- Barker. 4, 4a. Kenemans. 1892; 101; 1902:149. Carter. 1929: 2K2 . Ohenberger, 1930. 365. 5. Astraeu* (Depollus) lineatus van de Poll. Asfraeas rt/irvtv>M.c vur pictkolUs van dt roll, 1*S9: 84, 87-R9, pi. 2, figs 2, 2a. Kerrc- IK8M;9l. Kenemans, 1S9M01. 1902:148. Car- mans, 1892: 102; 1902. 149. Carter. tel, 1929:282, Obenberser. 1930:365. 1929. 2S2 ObenbeTger, 1930: 366. FIG, 22D FIG. Type, MNHN (not .seen by author). 22E Colour. Shiny. Head pronotum and elytra Type- Holotype. MNHN (not seen by the author) hlack with purple reflections; antennae black Colour, Head and prunotum dull, elytra shiny. with blue and purple reflections. Elytra with Head black with bronze-green reflections u\ Iho following yellow markings: a small irregu- the middle m the base, the Test with purple lar numbec of elongate markings each being reflections, with or without ,» red spot under- restricted to the width of one interval between neath each eye; antennae black with purple striae and occurringly mainly in three areas: reflections. Pronotum black with bronze-green near the suture; in the middle of the elytra; reflections in the middle, the rest with purple jiexi to the anterior part of the outer margin, reflectrons: a thin lateral red stripe on each Undersurfaee metallic purple, the lateral mar- side: with or without a small red spot on each ba*e. gin tif the 1st, 2nd and 3rd abdominal seg- side of the median lobe near Lhe Elytra ments each with u yellow spot which is variable black with blue reflections and the following red m^rking.s: two broad vJtrae, the first near In she and absent from the 3rd segment in margin* but not it. running from some specimens; hairs silver I cgs dark with the touching blue and purple reflections. the shoulder to the preapieul area; the second in the middle commencing near the front edge Shape and satlpntre. Head deeply punctured, running parallel to the suture, bur not touch- in parts in Ihe middle less so than at Ihe sides. ing it. to the prxapical area. Each of the vittae the areas involved glabrous; with u longitudinal may be entire or broken into two or a number glabrous line between the eyes merging with of elongate marks. Undersurfaee metallic- the basal impressed line; very hairy. Prono- purple with lateral red marks on the pro- tum with large deep punclure* dispersed but sternum, 3rd coxal plate and 1st. 2nd and 3rd more so in the middle with the elevated parts abdominal segments, The presternal and 3rd glabrous; mostly parallef-vided but abruptly abdominal spots are absent from some speci- rounded and narrowed at the apex: anterior mens; hairs silver. Lejis: femur and tibia margin projecting forwards in the middle: very metallic purple; tarvi brown. hairy at the sides, the hairs emerging singly, not in clumps, Elytra with the intervals Shape and sculpture. Head evenly punctured between the striae convex and slight! v with a longitudinal gljhrous line between the wrinkled, each With two longitudinal rows of eyes; hairy. Pronotum evenly punctured, the small hairs; parallel-sided, rounded after the punctures deeper at the sides than in the . 112 S KAjttCER sides slightly rounded Size. t 1 middle; narrowed and Males 12.3 0.73 x 4.9 1 £ 0.3 mp 7 ( fumi the base to near the npex. where they nre Females 15.I ^ 0.95 x 6.0 ± 0.35 mm »6>. more noticeably rounded; the Croat edge pio- Dntributitm. Western Australia. jecling forwards in the middle; hairs longer Specimens examined. W, Aust t A Leonota. and more dense at the sides than in the tfr.ww!. H W. 2.0.1934, WAM, 34-511: I £ W middle- Elvira with the intervals between the km 8 of Coolyardte, ian. 1937, A. A*. P4Utlu& 37-722; 1 striae convex with shallow punctures and WAM, j & t fc Tmin ROtf, UJHOi A M. UoUiitwr. WAM, 40-32, 40-31. 2 t Wat- lainlly transversely slnguse, laterally slightly tf nint;, 20.ix.l950, R. r, MtMUhu % WAM, 73-3*4 t concave from the shoulders until after the 73-386; 2 j£ -K I % Waininis, 2b.xiU9S(i. ft. f. middle, then founded to (he smalt sutural McMillan, WAM. 73-385. 73-3S8, 1 £ & 2 ft spine; hairy. Undcrsurface: presternum wit!) WAM, 73-375/7: I % Red Gum Pass. Stirling Ranges On Castutrina sp- 2G,i. 1971. £', <£• A', Cor* large puncture*, the rest with small shallow nahy punctures. less dense in the middle: hairy. st Sizes. Males J 1.0 * 0.46 x 3-S 0.22 mm 7 Astraeus (Dcpullus) irregularis van de Poll, <3). Females 33.0 ± 0.29 >. 4.6 ± 0.U mm 1889; 84. $6, 87. pi, 2, fig, 1, la. Kerrc- <6). mans, IS92; 102; 1902: 149. Caller, Distribution. Western Australia. 1929: 232. Obenberger. 1930: 36h, Specimens examined. W. Auxr.: 1 d\ 3 km W of FIG. 22G Payne's Find on Cnsuaritnt sp„ 17. ix 1970, \, Bar- type, Hnlotypc, MNHN (not seen by author.) ker; 1 J, South Tainmiu Flora Reserve on Casuarina campestris< 30,ix.l97O, .S. Barker; 2 ?, Colon/, Head and thorax dull, elytra .shiny. Qutiirading on C. cumpe.stris, 7.\i, 1970. S. Barker: Male. Head black with blue reflections, with J £ & 2 ?, South Tammin Flora Reserve on l the following, yellow markings: continuous vamptslris. K.xi.1970. 5. Atflbr; 2 Si 13 km N of a Goomnlltm; on C. campestris, 9.i.t97I, 5. Rarler. hand under |he lower half of the eye; an elon- gate spot in the cenfre of the head which may 6 Wr;uus ( n«*|ji iHnM milHinotaius . '. r de he broken into several spots; two elongate spots Poll, 1889; 84. 89, 90. pi. 2, fig. 3, 3a. lateral and basal to the central spnr, each of Kerremanx, IR92: 102; 1 902 149. Carter which may be broken into two spots: a small 1929: 282. Obenberger, 1930. 366. spot on either side of the gular, Except for the FIG. 22F central spot, all other spots may be reduced oi absent. Antennae dark brown with blue t v/h-. Holotype- MNHN (not been by the author). reflections. Pronotum black with blue reflec- Colour. Head and pronotum dull, elytra >hiny. tions and the following yellow markings: a spot Upper surface and antennae black with blue on the margin of the apical edge on each side reflections The whole uppei surface cccept the of the middle; a ^pot on each side below the antennae speckled with small yellow spots. first, towards the base, two spots along the Undcrsurface black with yellow spots larger sides, a small one at the apex and an elongate than on the upper surface; hairs silver, Legs larger one at the base—the first may be absent dark brown; each femur vvilh a single elongate and the second broken into two spots. All of spur; the tarsi black, thc.se spots may be reduced and some or till of Shape and sculpture. Head wiLh shallow even them absent. Hlytra black with blue reflections, punctures and a small glabrous spot between each elytron with the following yellow mark- the eyes. Pronotum evenly punctured, the ings*, the basic pattern is of three spots in the punctures larger at the sides than in the middle but not touching the suture; Ihe lirsl middle; slightly inflated after the middle. is a small elongate spot at the base; the second rounding and narrowing quickly to the apex: a large spot before the middle; ihe third a virta ihe I root edge projects forwards in the middle. after the middle and extending to the pie- Clytra with intervals between the striae eon- apical area; Iben: are also two small donjratc vex, punctured and wrinkled: laterally slightly spots at the margin, one at the shoulder extend- concave, rounding well after the middle to the ing U> the legion of the humeral fold and a sutural spines which are turned out. Under- smaller one just behind it which is sometimes surface evenly punctured, but the punctures ahsent; an irregular number of small spots are larger at the sides than in the middle, the present along the basal edge between the vittii differences pronounced on the thorax. Apart and the sulure. Undersurfnce dark brown with Inim the legs, apical edges of the abdomen and blue reflections; with yellow spots at the base pronotum, the body is devoid of hair. of each leg and single lateral spots on the pro- REVISION OF Bl.'PRFSl ID GENUS ASTKAKVS T.APOKTE At GORY 115 sternum, thud coxal plate and abdominal seg- and pointing downwards but not touching the ments I—4-, 1—3 or 1—2. decreasing id size fTom mouth; a single circular spot In the middle. first to last; hairs silver. Legs dark brown with Antennae brown with hlue and purple reflec- blue reflections. tions except fur the first segment which is pre- Female. "Head, pronotum and elylra basically dominantly black. Pronotum black with purple similar to the male except thai the yellow and blue reflections and the following yel low- markings are larger and the irregular spols are markings: a small ovoid spot on each side at more numerous. Undersurface and legs as in the apex; a larger ovoid spot in the middle of the male except that lateral spots are present each side at the base, slightly angled outwards on all the abdominal segments;. on the apical side; laterally there is a line on each tide from the apex to the base where it is Shape (put sculpture. Head broad; with shallow extended by the flaring outwards of the lateral punctures larger at the sides than in the middle margin of each lobe exposing the yellow inner and a longitudinal glabrous line between (he surface Elytra black with purple and blue eyes. Fronoturu with shallow punctures larger reflections and the following yellow markings at the sides than in the middle; the lateral mar- which do not form a symmetrical pattern on gins rounded and narrowed gently at (he base, each side: a large basal spot; behind and in more abruptly at the apex. Elytra with the line a large elongate spot, commencing befote intervals between the striae slightly con- the middle and ending al the. middle, close to vex, with a shallow row of punctures ami the suture but not touching it; an elongate spot taiully transversely wrinkled; parallel -sided commencing alter the middle and running to until after the middle, then rounded to the the apex, close to the suture but not touching sutural spine which is well developed ai>d it; an elongate spof at. the shoulder, not touch- iurntd out. Undersurface more closely punc- ing the outer margin hut close to it, behind the tured at the pdfe than in (he middle, with a last a much smaller spol not touch- few short hairs. ing die margin: after ihe middle there are Size Males 1 1 .2 ± 0.24 x 4,1 - fi.OK mm several elongate %poK intending almost to the (II >. Females 13.5 ± 0.22 x 4.7 ± OM mm apex but not touching the outer margin; several (ID). small eccentric oval spots. Undersurface black Distribution. Western Australia. with purple TcrtcetLons; the first three Specimens examined. W. AuM-: 3 & & 5 ?. Cion- abdominal segments have lateral yellow spots nclk on Casuarirui up., 24."V»k19fi7, 1 S Barker; which progressively diminish m size from in tJ & 2 9, Gosnclls on Casuavina sp . 27.xJi, 1967. fronl backwards; hairs silver. Legs; raeso- V. 'Darker; 2 S A 3 ?. Red Hill Rd near Midland Junction on Casuurina sp.. -4.i.l9rJS, S. Barker: 2 sternal and mctastcrnal coxae with yellow spots ?. 77 km along main York Rd on Casuatina «p. t on the outer margins; the rt$t brown with 5.i I06R> S. Barber, 3 4 ik 4 ?, South Tammin purple reflections; the upper surfaces of the Mora Reserve on CasuaruHt hne^cliana, C1,1971. tarsi are black. S. Barker; I & & 2 8; 142 km E of Norseman on C, fuu'Reliarut, 19.xii.1972. S, Barker; 1 ft 58 km Shape and sculpture. Head with shallow punc- of Tammin on York -Tammin Rd on C. W tures and without hair. Pronotum with shallow WeWfVv/w, 23\x\U97t, S. Barker; 1 •& 1 5. 3 rf punctures; a short median km E of Gosnclls on C. lutegciiana, 27.xii 1972. impressed line pro- .V. frmm jects forwards from the basal crypt; parallel sided at the base, abruptly rounded at the K A stratus iOej#ollu^> dednrieiiMs sp. nov. apex; with short hair on the anterior edge only. Elytra with the intervals between the striae FTGS 4, 22M slightly convex and transversely strigose; Typ?!- parallel-sided until after the middle then Holotype* tj? DeuVil, W. Aust , fi t W. Brvwa. WAVf, 71-1779. rounding off to the apex; each ending in a Allotype: ?, Dcctari. W. Aust, H, W Browrn Lhiek short spine projecting backwards; no WAM, 71-1778. marginal spine but a small mound is preseni in Pamtypes: 7 ?, Dcdari, W. Aast., WAM, the region where they occur in other species. 40-1207, 40-1208; 1 f, Borden, W Aust., SAM: I ?, Deduri. Vv\ Ausu, AN1C; 1 ?. Undereurface with shallow punctures; with a Dedari, W. Aust. t on Camanrta Lwmeulatih lew short hairs, 21X1936, fj W, fa own, MNHN. Size, Males J 1.2 .\ 42 111 Fem.Ves 11.0 Colour, Shiny. Head black with purple reflec- mm ± 0.15 :< tions and the following yellow markings; on ftp m 0,1 mm (6). ovo>ii spol touching the underside of each eye Distribution. Western Australia. 114 S. BARKER General remarks. This species shows closest long. Pronotum deeply notched on both sides affinity with Astrueus irrcyttlctris. at the base, with the median lobe and the two Specimens examined, Types only, lateral lobes pointed, Elytra striatc-punctatc and/ or costute, the outer edge of the intervals SflgpgftfH$ AstratMi* (seiisu stricto) sub-gen. between the striae costatc, either down the nov# entire length or towards the apex only. The Type species Astweus flovopiclus LaPorte & Sl j flirc cn Key to the species of AstraeuS (sensu slrklu) I. Head with a median keel ..,,_.. 2 l.Head without a median keel ... ,., ...... 15 2. Hairs silver . , ... _ 2. Hairs yellow ., ,., , ,.,. ., )* x Part or all of the anterior nndersurface red-brown 4 .3. None of the anterior undcrsurfaec red-brown 6 4. Guiar, prosternum. meso- and metastcrnum, 2nd and 3rd coxae and 1st abdominal segment .. 4. Prosternum and coxae red-brown ,..,.,,..,. - 10. minutu.\ sp. nov, 4. A red-brown area on either side of the proslermd process 5 5. Usually shorter than 7.5 mm; humeral fold well developed and acutely angled , II. frasniensis sp. nov. 5. Usually longer than 7.5 mm: humeral fold moderately developed and angled 15. ohsaiYux sp. nov. . 6. Most of the leg testaceous . .... 7 6. Tips of the tibia and 1st tarsal segment testaceous ...... 8 6, "None of the leg testaceous .. .. -.,.- .._ .. 10 7. 1st and 2nd legs testaceous except for the outer margin of the femur, 3rd leg testaceous except '. for the femur . „ . . _ ...... 19. dlhttipt's van do Poll 7. Tibia, 1st and 2nd tarsal segments testaceous , , 13. ,s?nythi sp. nov. 8. Usually shorter than 7 mm 12. pypnaeus van dc Poll 8. Usually longer than 7 mm - 9 9. Humeral fold well developed, acutely angled 17. nmstersi Mad env 9. Humeral fold well developed and angled !8. samoueUi Saunders 10. Humeral fold moderately developed and angled II 10. Humeral fold poorly developed, slightly angled .,., ,. 13 I I. Head and pronotum green and coppery-purple 21. intrkaius Carter IT. Head and pronotum black ...... „ .. 12 J2. Broad and rounded species 16. gloko.\u.s sp. nov. 12- Elongate species ...... ,_ _ ,,. 35. watxoni sp. nov. 13. Black species; elytral markings a number of yellow spots 22- crasxits van de Pull 13- Blue species; elytral markings two yellow fascia 25. fraternrius van de Poll 14. Elvira! markings three yellow fascia and red ureas 23. major Rlaekbuin 14. Elytral markings iwo yellow fascia without Ted areas .. 24. navarchis (Thomson! t£. Body elongate and cylindrical . I IS l5.Body not elongate and cylindrical , . , 17 16. Pronotum conically elevated in the middle 26". prothoma'ats van de Poll 16. Pronotum not conically elevated in the middle 27- slnntwttts van dc Poll 17. Su rural spine with a rounded internal edge . IR |/i Suturaf spine with a straight internal edge 26 J8. Legs a red-brown colour IV 18. Legs not a red-brown colour 20 19. Elytral markings spots and fascia 30. mmrnHhmi sp. nov. 19. Elytral markings two vittae on each elytron ,., 28. vttuittts van de Poll 20. Hsad, pronotum and legs metallic brown or hron/.e , , 29. ffavopictus LaPorte & Gory 20. Head, pronotum and legs not metallic brown or bronze 21 21. Humeral fold well developed and angled . 22 21. Humeral told poorly developed, slightly angled ...- ,-. 23 22. H-ad black or coppery-purple; undersurface cc-pp^y-purple • 20. adamsi sp. nov. 22. Head blue or green; undersurface blue-green . ... 14. Abnuhi/i/t van de Foil REVISION Oh BUPRESTID GENUS AStttAEVS LAPORTE k GORY 115 1} Basal spot expanded along the trout edee of ihe elyuou ... •• *..- • .. 31. carnuh\i tip. nov. 23. Basal spot not expanded along Ihe front margin of the elytron 24 24 l £Jytral mark/rigs eitlici two spots and two fascia oi tbt first fascia may be broken making lour spots and a fascia on each etvtron ... 32. bndenl van de Poll 24 Flytrnl markings either $rx spots and a fascia or the fascia may be broken iuaJcnie eight spots on each elytron 33. jansonl van de Poll 24. Elytra! markings 7 spots on each eJyiron ... 25 25. Kronotum parallel-sided from bass? to the middle then strongly rounded and narrowed to the apex; dorsally convex in lateral profile ... - 34. edwrthuri van lateral profile , . . .. _, ..... _ _ s6, carter! sp, nnv. .slightly excavated .. 26. Head between the eyes , ... 37. goertingi sp. nuv. 26. Head deeply excavated beiwccn the eye* 27 27. Head excavated mainly at the base; propotOiU laterally inflated with an oval patch of hexag- onal cells in the middle ...... 38. e.yaneus Kerremnns 27. Head excavated mainly ai. the apex; pronotum not laterally inflated and without hexagonal cells 39. catedoniciis Fauvel 9. Astraeus (Astraeus) bakeri sp. nov. lobe without a basal crypt; sides rounded and narrowed from base to apes, hairy, Elytra FIGS 5, 221 costalc. the intervals between fiat with a pro- minent row of punctures; at the sides gently Holotypc: i Dryandra, W. Aust. on rounded from the base to just before the C'ostfarifiii middle hucgeliana, 8,xii,l£70 ( S, Barker. then rounded and narrowed to the SAM, 1 20945. strong marginal spine: sittural spine rohust; Allotype: 9, DryandrB. W. Aiixt. on C*. /»«<#•*• humeral fold we'l developed .ind angled. H«hu> 8 xii.1970, 5. Barker, SAM, I 20946. Undcrsurfacc sparsely punctate in the middle, Furalypes: 1 Casuariaa sp., 2. i. 1968. S. Barker, SAM; 4 She. Males 7.5 ± 0.07 x 3.0 as 0,03 mm (44). 3 & 4 ?, Drvandra, W t Aust. on C, hong* Females 7,9 i 0.12 x 3.2 ± 0.06 mm (23). liana, 8.xu\1970. S, Barker. ANIC (1 liarta, 1 broken into spots in some specimens: a liirge 9\ii.lV74 T S. Barker; ^. 115 km S of Bnlladonin on Casnarttut humitis. 9.xiiM974. spot below the middle, Undersurlutcc; gulaf, S~ Barker; I 6*, SW valley of Ml Rafieed,' on C pro- meso- and meiasternum, second and third huepcliumi. I0.xii.l974, S. Barker; » 9, 3 km S ot coxae and median anterior part of the first Mt Ragged on Israelite Bay toad, on C. lutmiffx, abdominal segment ail reddish -brown with KUii.1974, S. Barker. light blue rellections on the lateral part of the above; remaining abdominal segments 10. Asfraeiis (Astracus) minufns sp. nov. black with metallic blue reflections; hairs sil- FIGS 6, 22J ver. Legs brown with metallic blue rellections Types. on the upper surface, of the end the first tarsal Holotypc: d\ South Tammin Flora Reserve. W. segment and all of the remaining tarsal seg- Aust. on Casuartna hue^eltana, 6.i. 1971, &> ments metallic blue, Barker, VSAM, T 20459. Allotype; ?, Souih Tammin Mora Reserve, W. Shape and tfcttrpwre. Head evenly punctured; Aust. oa C. huexetiana, 23.xii.l972, .S Bar- small median keel; h.iiry. Pronotum deeply ker, SAM, I 204/V0. Panjfypcs: 1 t Soalh Tammin Flora Reserve. and evenly punctured, with a median longi- .J W. Aust. on C. huexttinna, 23.xii.1972> Si tudinal impressed line at Ihe apex; median Barker, ANIC (1), MNHN (1); 1 «? South 116 S. BARKER K 1 I ft 5 ^ 1 . REVISION OF BUPKfcSUD GtNUS ASTRAEUS MPOftTB & GORY 11*7 Tammin Flora Reserve, W. Aust, on C I9.dii.1972. S. Barker, ANTC f I cf & I 9K 1 1 1 2}-: 1 Aust., huv-elkma. 6.LI971. 5. £a/A?r. SAM: jjf, BM (Id * A J-ate Grace. W. South Tammin Tloni Reserve;. W. And. on 9.L1972. E. A K. Caraabv, KC: 3 & Fyre C. tlUdf&IUMA 9.i.l9?4, 5. MrWi SM; 2 .il974> A, ^tifte/i WAM. SAM; 2 6 &. I 9, South Tammin Flora Colour. Shiny. Head black with purple reflec- Reserve. W. Aust. un C hueselitma. 6.U971. V. Barker, tions; antennae black with blue reflections WAM. except for the tips of the first and second seg- Colour, Shiny. Head, antennae and pronotum ments which are brown. Pronotum black with black with violet and blue reflections. Elytra purple reflections in the mdldlc, red-brown at black with purple reflections, each with the (he sides. Flytra black with purple reflections following yellow markings: a large basal spoi each with the following yellow markings: a in ihe middle; a fascia covering the humeral large basal spot; a fascia before the middle fold, running towards rtie suture but not touch- covering the humeral fold bui not reaching Ihe ing it, and slightly concave towards the base: a suture; a large median spot after the middle, large spot after the middle not touching the Undersurfacc: presternum and coxal plates suture or the margin; sometimes there is a red-brown, the rest black with metallic purple preapical spot. Undersurfacc metallic purple reflections; hairs silver. Legs: upper femur red- e She Males 5.7 0.22 x. 0.10 ± 2J * mm (7). SSte Males 07 ^ 0.05 x 2.6 ± O.02 mm (72). Females 2.7 (I). 64 x mm Females 7.0 ± O.ritS \ 2.S =r 0.U4 mm (2S) fiistrihutUm. Wcslern Australia. Disitibnrton. Western Australia. Gaieraf remark*. The smallest species known General remarks. The three speciei Atfraeus at this time. bakvri, Asrraeits mfrwrus and Astnwu* fra&et* Specimens extfflv'Wii Types only. tenxis show a close affinity to each other. Specimens' examined. W. Aust.; Types: I V. 58 km 1 1 . Astraeus (Astrueus) fraserfrnHs sp. nov. VV of Tammin on York-Tanuniii Rd, un FIGS 7. 22K. 24. 25 Camarina huea? litt nu, 23.xii.1972, S. Darter; I o- Types, 58 km W of Tammin nn York-Tnmmin Rd, on C, Holotypc: £ Vraser Range, W. Aim on hu & I 5). MNHN (I cf & I 2): 2 $ & 2 ?, 5 S* Ju/anda jockhole 106 km S of Balladouiu, Fraser Ran*e, W. Aust. on C Imey.eliamt on C huecetiana. 9.xii.l974, ffi Barker; 1 & 19 Ilfi S. BARKER km W of Balladonia. on C. tuimifis. 23.ii.l97S, punctures and transversely wrinkled; parallel- V. Hotkey, sided until after the middle then rounded and 12 \fitraeus (Aslraeus) pygmaeus van de Poll, tapered to Ihe marginal spine: both spines well developed; fold 1 8Kb I7K-IH0; ISS9: 85. 104. 105, pi. 3, humeral well developed and figs In*, 16a, loc, Blackburn, IR91: 496. angfeo. Undersurface: prosier nal punctures van de Pod, 1892; 67, 68. Obenoetgor, larger at the sides than in the middle, punc- 19*0; 366. Carter, 1931: 107; 1*3,3: *!. tures uniform on the abdomen: hairy. Astt tti tts pvginui'ii\ var. suhjawiutus van dc Size. Males 6,3 '- 0.05 x 2.5 ± 0.02 mm (66). l»olK 1886' 178-180; 1889: 104. 105, pi 3. Females 6.6 at Shape and St ulpturv. Head evenly and closely Female, Head; apex turquoise, base green; punctured; with a median keel; hairy. Prono- antennae black with blue and green reflections. tum closely punctured, the punctures larger at Pronotum green in the middle, turquoise at the the sides than in the middle; parallel-sided sides. Elytra black with blue and purple rcflcc- from the ha.se until just before the middle then tioas. yellow markings as described in the male tounded and tapered to the apex; hairy. Elytra but less prominent. UrKtersutface and legs as custutc. the intervals tint, cadi with a row of in the male. . _ REVISION OF BUPRESTID OFNl-S A5TRAEUS LA PORTE & GORY IH> Shape and sadputn. Head evenly punctured; Shape and sculpture. Head with close uniform a median keel; hairy. Pronotum with punctures punctures; no median keel; hairy. Pronotuin latget at the sides than in the middle; a faint evenly punctured; basal third parallel-sided longitudinal impressed line commences at the then slightly rounded and narrowed to the middle and runs to the apex? parallel-sided apes; anterior margin project big forward in from the base to just before the middle then the middle, that general area glabrous: hairy. rounded and tapered lo the apex. Elytra F.l\lra eostale. the intervals between flat but costate, the intervals between flat each with a slightly wrinkled; parollcl-sided from the base. row of punctures and faintly transversely rounded before the middle then tapering to the wrinkled; parallel-sided until before the marginal spine; both spines well developed: middle, then rounded and tapered to the mar- humeral fold well developed and angled. ginal spine; both spines well developed; Undersurface evenly punctured; hairy, developed and angled. humeral fold well 5a. Males 6.7 ± 0.10 x 2.5 ± 0.05 mm ( II >. Undersurface. prostcrnal punctures shallow in Females 7.0 - 0.24 x 2.7 ± 0,09 mm (6)- deeper at the sides; punctures on the centre, Dhtriluition, Queensland. the abdominal stemites sparser and shallower: General remark* Of the 17 specimens I have hsiry. examined, 7 c? and 2 ? have preapical spots Size. Males 6,5 ± 0,08 H 2.6 ±0.15 1 2) mm on the elytra, the other eight specimens have Female* 6.5 ± 0.18 x 2.6 £ 0.01 14) mm no preapical ivpots. Although male genitalia pi\?rihnti'->n- : Queensland. ( Fig. 22N ) is similar in shape to the previous Genera! remarks, A WWhi shows cfnse two groups of species, the external morphology affinity with A, pygmaean. I place these species of A, simulator is dissimilar to those five alongside *he previous group of three species species. Because of this 1 do not grnup it with Named after my late colleague, Dr M. Smyth. any of the above mentioned species. SpecitHwt* vxatmnefL Qld : Types; I 2, One Tree Specimens examined. Qld: 'lyre: 1 c?. 3..xii.l 07 1 Hill, Brisbane. 1920. i\ Mwr, nPBM. E. E. Adams. KC, 4 (t I !?»: 1 2, Fdunjulba on 14, Astraeus (Asiraeuil si initial or van dc Poll. SAM rf & 9>, EA C. equisrtifoth, 15 xiU*>73. t E Adam*. EA. 2 188°: 85. 102. 104. pi *. rigs 15. |5& rf, Edungalba on Caxuar'ma sp.. I on each o( 15l>. Keiremans 1892; 102. 1902; 148. 23JU.1974 aUtl 8jrf|4$H E, E, <* 5. dtfeWi HA; Carter. 1929" 282. Obcnbcrgcr, 1930; 366. 1 2 c?» Edtinjealba on Ca&uarirta sp., 15.xii.l974. J y/if* Hn|otypc 9, Peal Downs, MNHN (seen by E. E. tfi S. Adams, EA. :i u tlioi I Colour. Malt'. Shiny. Head green at the apes 15. AsflnAn (Astraens) onscurus sp. nov. with golden-green rcllecions, black at the base with blue reflection*: antennae black with FIGS 9. 220 J ypes. golden-green and blue reflections. Frenulum Hototype: 3*, Eraser Range, W. Aust.. un hlack with blue and purple reflections- Hlytra Casuariua htuKvu'ana, ]9.AJi. 1972. &. BarUvv. blciek with turquoise reflection*, each with the SAM. I 20957. <"*- following markings: a large nasal spot", a lasCta Allotype: ?, Eraser Range, W. Ausl.. on I9.xir.1972, ,y, timier, ) covering the humeral fold, running transversely Imefiehww, SAM, 20958. Towards the suture but not touching it. concave Paratypes: 2 J &. 2 £ Eyre Highway 142 km E towards (he base and clubbed ut ihc end; u of Norseman. W. Aust. on C . Iiue^eh'iwu. fascia after the middle running transversely I9.xii.1972, S. Barker, EA U 6* & 1 $K ( 6* i Range. from the margin but not touching the suture. ANIC I A ?»; 2 o* & 2 ?» rrftttr W. Aust. on C. huik'eliaua. I9.xii. 1972. 5. Undersurface metallic blue: hairs silver. Legs Barker, BM (1 o & I 2). MNHN ( 1 rf &. 1 metallic blue; tips of libia and first tarsal seg- ¥>; 1 ri\ 24 km NE of Lake Grace. W Auhi.. ment brown: second, third and fourth tarsal on C. ht/wtiana. 24.U973. 2f, Barker, KC; segments black with blue reflections. 1 ^, Tammin, W. Aust.. //. W. firowih WADA; I 9. 2 km N of Nccdilup, W. Auu.. Female. black with blue reflections: Head 23.xiiJ972, K, Kewhy. WADA; I 9, 30 *m .antennae black with blue-green reflections Pro- W uf No. 8 pumping Hlattim ( Kalgoorlie vvalcr m;iin) on Casuarhta >'ampesirts notum black with blue reflections. Elytra, black 24.I.IM5X, WAMs 71-1775; I ?, WAM, 7t* with blue and purple reflections. The rest as 1777; 1 S; South Tammin Flora Reserve on In male. the C fmmttmi* 23.xii.l972. .?. BarUt WAM 1 SO S. BARKER i i O I 3' ; 1 ^ rf I * 1 £ in I REVISION Of BUPRRST1D GH-KUS 4&T&4&V& LAW1RTE & GORY .2. Colour. Shiny. Head purple or black with In Astrae-ns (Astraciis) gltitaosus sp. ttov. metallic purp/e reflections at the apex, metallic FIGS 10, 2211 blue reflections at the sides and underneath; antennae black With blue and purple reflec- Holotype: d, 77 km alone main York Rd. w. tions, tips of first, second and third segments Aasl. on Cusuunna kHetoeJhlNA 5, i. 1968, .V. dark brown. Pronotum purple M black with Bar^r SAM, T 2U953 metallic blue reflections at the sides. Elytra Allotype: $ 77 km along main York Rd, W. Aust. on 1 black with blue and purple reflections C hbfg$l(0Jl8, 4 196* , V. Horker* SAM, J 20954. each with the following yellow markings: a Paraiypes: 2 A & I ?. Dryandra. W. AusT- co large round basal spot; a broad fascia before C. fute&liana I2.JJ973, S. Barter* EA; 3 3 the middle concave towards the base commenc- & 2 $, Dryaiidra. W. Aust. on C. hne# i. small preapical spot in the form of a lunette 23. 1072, E A A' T Carnaby, KC; I 8, 77 sometimes missing. Undei surface black with km along main York Rd". W. Aiwt. on Casuarina sp., J. i. 1968, a\ Barker, SAM; 1 <£. purple reflections, except, lor a red-brown luuaning Reserve. W. Aust. on Casuarina patch either side of the prosternal process on to., 111968. S. Barker, SAM; I g, Spencer's extending lo the laleral margin; hairs silver. Brook. W. Anst, 26..\ii.l94.S. R, p, McMillan. 71-1763. Legs: femur, basal half of the tibia and WAM. Colour. second, third and fourth tarsal segments dark Shiny. Head and pronotum and elytra hrown with blue and purple reflections, apical black with coppery reflections; antennae black with coppery blue reflections. hall' ul (he libia and first tarsal segment brown. and Elytra; each elytron with the following yellow markings; a Shape tmd sculptta?. Head evenly punctured: large spot at the- bn.se; a broad fascia before a large median keel; hairy. Pronotum deeply (he middle, commencing at the shoulder and punctured; a short impressed line projects for- ending near the suture but not touching it, ward from the basal crypt, becoming a groove concave towards the bn**e; a second fiucia after and extending to the apex; short with rounded the middle, commencing ,it the margin but sides gradually narrowing from base to apex; not reaching the suture; an oval prcapicai spot; hajry Hlytra costatc, the intervals between flat a single very small spot between the two fascia each with a row of shallow punctures; more and close to the suture. Undersurface metallic or less parallel-sided until before the middle coppery; hairs silver. Legs: femur metallic then rounded and narrowed to the marginal coppery, tihia and tursi brown with varying spine: both spines well developed; humeral fold amounts of metallic blue on Ihc femoru and moderately developed and ungled. Undersur- tibia and on the upper surfcicc of the tarsi. face evenly and shallowly punaured; hairy. Shape and .w.'fiputre, He;u1 closely punctured; with a median keel; hairy. Size. Males 9.1 £ 0.05 x 3.4 ± 0.02 mm (105). Pronotum deeply punctured but Females 9.2 ± 0.08 x 3.4 ± 0.03 mm (64). more closely at Ihc side* than in the middle; short with rounded sides Distribution. Western Australia. narrowed towards the apex; H shon imposed line projects forwards from the basal crypt, Specimens examined. W. Aust.: Types; 2 S & 2 ?. hauy. Elytra coslale, the intervuls ilitt each Dedari, If. W Bntw/>, WAM, 7M77J/4; 1 J, Spencer's Brook. 24.xii.l94A. K. P. McMillan. with a row of shallow punctures: rounded. 71-1764; WAM. t S, Bcjoording, I6i,liwy. diverging slightly from the base to a maximum WAM, lirliWX 3 6*& I '?. BefooixSIOfc, 29.xi.J950. width before the middle, thereafter rounding 3 K. P. McMillan, WAM.. 71-1765 /& V3. 1 km E off of No. 8 pumping, station (KalgOOritfe water mam) to the marginal \pine; buth sprnes well oil Casuarina eampextrit, 2(J.i. 1958. WAM, developed: humeral fold modetafely developed 71-1776; 33 A & 2J 9. Coragina rockhole 66 tan and angled. Undetsurface: finely and sparsely S Of BalladOllla on Casuarina hutaeluma, punctured in the middle ot the sternal seg- 5xii,l974. 5. Bar/car: 21 J & 13 9. Ponic? tock- ments, closely hole 70 km S of Balladonia on C. htiegrtiatia. more coarsely and punctured at S-xii 1974. S. Barker; 14 cf & 3 ?, Tummf* rock- the sides: abdominal sternircs finely and closely hole 106 km S of Balladonia un C hut'Kelitwa. punctured; hairy 9X11197*1. S. Barker; i if & 2 Distribution, Western Australia. suture: a small spot neat the base, ai the General remarks. This species shows closest shoulder, may or may not be continuous with the middle a tavcia affinity with A obsenrn:,- the base of lhe fascia: after 1 running transversely from the margin towards Alls!.: 'types; 1 Saeeitnens examined W. J , 77 km along main VorV Rd on Casuaritm sjv, the suture but not touching iu a preapical 5.U968. Si Barker; Z t& 77 km atone main York spot. Undersurface blue with metallic, gleams; 7.xii.l970, Rcf i>n Cnsuuritm huexefiana, 5- Barker; hairs silver. Legs metallic blue; lips of tibia J »;*, Dryandra on C- hta^Hana, 7.xi.lW), tS\ and first tarsal segment testaceous. Barker; 9 J & \ 9- Dryandra on C". hut^elutna, X \M.IS>7(I. .V. Barker. Shave and sculpture, Head shallowly, closely but uniformly punctured; with a median keel; 17. Astracus (Agtraetts) masters! MaxJ coy, hairy. Pronotum shallowly punctured with 1873; 239. Kerremans, 1S85: 136. punctures larger at the sides than in the Masters. 1886: 71. Kcrremans. 1892; 102. middle; a median longitudinal excavated line Astraeus stanoaetli; van dc Holl» 1886: 176, from base to ape*; straight sided from the IS89: 80, 107-109, pi. 3, fit*. 18b. Kcrrcnmns, base to the middle then rounded and narrowed 1 i0? 1 48. Gctfftfi 1 : 2S2. Obenherger, 089 to the apc.\; hairy. Elytra costate, the inter- 1930; 366. vals flat but slighlly wrinkled each with a A.tnaru.s spletnh'HS van dc Poll, 188*); gg, 108, row of punctures; more or less parallel-sided 109, pi. 3, figs 19. 19a. Kctrcmans, IS92: 102: 1902: 148. Carter, 1*29: 2S2, tig. to before the middle, rounded at the middle fjj. 3, 45. Obeiiber»er. 1930; 367 (new synonym). then tapering to the marginal spine; both spines Astraeas simplex Blackburn, 1802: 211, 212 well developed; humeral fold well developed kern-man-,. J 902: |4!>. 1929: Carter, 2S2. and acutely angled. Undersurface with shallow Obenberger, 1930: 3f>h inew synonym"). uniform punctures except in the middle anil HGS U, 22Q on the outer edges of the abdominal stcuijtes. both areas being glabrous; hairy. r.v/u. Holotype; 2, Gaynclab. AM. X32099 (seco by author). Steej Males 7.9 * 0.16 x 2.7 * 0.22 mm (17). Colour. Mute. Shiny. Mead ami antennae Females 8.4 - 0.26 x 3.2 ± 0,10 mm ( 10). golden-coppery. Pronotum copper coloured in Dtxtnbuiiw. Queensland and northern New the middle at the apex, wiih a dark blue heart- South Wales. shaped area in the middle, outlined by a Is only species of copper coloured margin; at the sides, blue at General temaeka. This the greatly (see the base arecn at the apex. Elytra black with Astraeus that has been confused reason for blue or purple reflection*;., each elytron vvitb van tie Poll 1889. p. 19). The basic A. nutstersi and A. the following yellow markings; a large spot at this h that the females of in although Lhe base; a broad fascia running from the Mimouetli are very similar pattern shoulder covering the humeral rold and run- the males of (hesc species are quite distinct. ning Hansvers^Iy towards the suture but not Unih .-species were described from female types hold collecting experience was the teaching a. after the middle n broad fascia The lack of from the margin running towards the suture most likely reason for van dc Poll redesc.ribing the male of A. mast erst as A. spteiittens, as he hvi not reaching it; a pt'eapical spot —in some would noL have had access to an associated specimens the preapicM spot \$ absent and females ol lhe two species. usually in these the first fascia is broken in the scries of males and inrddle forming two spots, Undersurface blue; Syeehm-ris examined. N.SAV-' Type; I 6 paratvpe. Ash 1.. ANtC: I A & 1 S paratync, CiaynriAh. hairs silver. Legs: blue: the eud of the tibia and J ANTO. Qld; Type; £ A. splendent van de l oli, first tarsal segment leslaceous. Kockhumpton, 3 ?, Da [by, Mrs, F. H. Jlokler. I Sloiuhorpo. Cemmetl; 14 dT ft 5 female. Shiny. Head blue-gicen at the apex. SAM; J, A 9, Edongalba on CwtfitUia rqnisetifolia. 24.x. 1971. dark at the base and aides with purple tcIIcc- F. E. Adam*. Type $, A. aimulex Blackburn KM lions; antennae black With blue or purple re- (locality of collection uncertain). flections. Pronotum dark blue in the middle. metallic blue at ibe sides, Elytra black with 18. Astraeus (Astraeus) samoueHi Saunders. fig. purple reflections, each with the following 1868: 10. pi. 1. 12, 14S7I : 43. yellow markings: a large spot at the base; a Masters. 1 871: 124. KcTremans. IRK5: lateral fascia commencing M the margin and 136. van de Poll, 1886; 176. 178, 1889: covering the humeral told, running towards lhe «6, 107, pi, 3. fi^s i8 t 18a. WacVburn. suture but Interrupted to form a spot near the IN91: 496. Kerremans, 1S02. i02. van dc REVISION OF BUPRCST1D GENUS ASJXAtUSS l.APORTF. A GORY 123 Pol), 1892; 67. Blackburn, IS92; 2J2, Females 9.2 =: 0.14 x 3.3 ± 0.06 mm (40) Kerremans, i902. US. Cariet, 1929; 282. G+nrrai remarks. The type of A, .splcadens var. Axtrueas samouelki: Cernmirigcr & tie Harold, emhtikieUus Obenbergei is a V specimen of A IH69: 1380. Masters, 1**6; 7J. Blackburn. samouetli. This species IK90: 1256. Obcnberger 1930: 366. shows closest ailinitv Astraetts sufHOticlli: Ker remans, 1900: 295. With A. masters!. Astmcus .iplenden,\ var roihrikieflas Oben- Distribution. New South "Wales. oer$ei\ 1936: 133. Specimens examittrt/. N.$,W., Type: 2 6* & 3 9, FIGS lb, 22R Bredbu on Ctisttarina striata, 8.1.1963, S. Barker; 12 o* & 20 % Majors Creek on CvSUttriuti fit- Type. Hofotype: ? BM fseen by author). 1 Totals 2fr.i,1963, 6'. Barker; 8 d & 2 2, Khanyunis Colour. Male. Shiny. Head golden-green, Station. Mimima Range on C. Uttatalls, 9,ii.19fiV golden or coppery; antennae coppery or tur- S. Darker, 4 d & 2 ?. Gipttiins Hat to Braidwood quoise. Pronotum uniformly gtildcn-grcvn or Rd. on C UitoraHs. 9.ii.l96.V S. Barker; Type 9 A. splendent Vac, enthr<\teltti> Obenberger, golden or enppcry in the middle then golden- Comoro, NMP, 219S9 A.C.T.; V, Mt A indie on green blending into turquoise ai the sides. C strhiu, 26.xii.l962, S. Barkrr; 7 & & 12 ?, Tng- Elvira hlack with blue reflections, each with jeeranong on C ihfota, I xii 1962. S. Barker; 3 rf & 8 *?, Tufigetauong on xtricw, ,S\ (he following yellow markings: a large gpol ut C. 2.xii 1962, Barker: 4 A St 2 5, Tugsernnotte on C. f/ri*tJff, the base; a fascia just before (he middle cover- l.i.Wtf. V tiarker. ing the humeral told, extending backwards then transversely towards the suture but not 19 \ st rums (Astraciis) diluUpe* van de Poll, touching it: a spot at ihe shoulder which ma> ISS9: 86, ir>5. 106. pi, 3. fig, 17. 17a. be discrete or continuous with the base of the Illackburn. 1892; 2l2. Kerremans, 18922 lin>t fascia; a fascia after the middle, commenc- 101: J902: 148. Carter. 1929- 282. Obeu- ing at the margin and running towards the berger. 1930: 365. suture but not touching it; variably, a small Asrraeits samunelli var. ditmtpes vun de PnlL 1886: elongate spot between the two fascia closer to 180. Astrarat MrumH OberuSerger, 1028: 205;'Carter. the first and near ihe sulure hul not touching 1929: 302. Ofccnbcrgcr. 1930: 367 (new syno- it; a preapical spot. Undcrsurfaec bluc-gteen; nym). hairs silver. Legs blue-green; tips of the tibia FIG. 22S and most of the first tarsal segment testaceous. J>0« Hofotyps, MNHN Uiot seen by author). Female: Shiny- Head gieen ;il the ^pcx with Colour. Mate, Shiny. Head green at the apc.\ golden reflections, dark blue with purple reflec- and golden green w\ ihe base or jroldcn-green tions in the middle of the base, turquoise at at the apex and coppery at ihe base; antennae the sides; antennae ^reen. Pionoium dark blue black with blue reflections. Pronotum divided with purple reflections in the middle* turquoise in u transverse direction by an M-shmped line. iii the sides. Elytra black with blue reflections, the small apical part being preen or golden- with markings as in the except male that the .green, the larger basal part bein? dark blue tit first fascia is broken in The middle to form a black and the line of demarcation, purple or fascia spot marginal anil a near the suture. coppery. Elytra black with hluc reflections Underxurface turquoise: hairs silver, ljcgs tur- each wiih the following yellow markings: in quoise: tips of the tibia and mosi of the basal the middle but not touching ihe sulure there part of first tarsal the segment testaceous. is; a large basal spot; a spot before the middle; Shapt- artel sculpture. Head shallowly. closely a preapical spot. On ihe margin- a large spot but uniformly punctured; with a median keel: covenn^ ihe humeral fold; after the middle a hairy. Pronotum shallowly closely hut uni- transverse fascia not touchine the suture. formly punctured; with a median longitudinal Undersurface blue or green; hairs silver. Legs: excavated line; uniformly rounded and first and second femora hrown on the outer nanowed From base to apex; vciy hairy. Elytra margins, third femur totally or partially solid oostatc. the intervals heiween flat; slightly colour similar io the rest of the iindersurfHce; lapered from the base to tile middle, then lemamder of leg and tarsi testaceous. rounded and tapered to ihe marginal spine; Female. Head green at the apex, purple in both spines well developed. Undersurfiiee the middle, dark blue at the base with purple shallowly punctured, more closely sides. at the reflections: antennae black with blue rcflec- less so in the midsiirface middle; glabrous; lions Pronotum black in ihe centre with hairy. purple and blue reflections, bine at the sides. Size, Males 8.6 ± 0.07 x 3 ± 0.03 mm (52) Elytra as in the male. Undersurtace dark hlue; A 124 S. BARKfR hairs silver. Logs: outer margin of femora middle nor touching the suture and just behind brown, tibia and tarsi testaceous. the second spot these last two in the form of fascia; Sfta/H- and sculpture. Head very densely and 8 broken J4JJt behind the break, near evenly covered with deep punctures: with a the margin but not touching it, there is a small thin medial) keel; covered with long hair Pro- spot (not present in a third of the specimens noium evenly punctured, the puncture* at the examined): after the middle a fascia concave but not sales larger than those in the middle; slightly backwards commencing at lhe margin rounded bui narrowed from base to apf.v, reaching the suture: a small preapical spot. median excavated line from base to apex; Undcrsurfncc coppery-purple; legs pale brown,, hairy. Ulytra cost ate* the intervals Hut with a the basal ends ol the tibia darker: baits silver. row ut deep punctures and faintly transversely Shape and sculpture, Head closely and evenly strigose; narrowed from base until after the punctured; nu median keel, lightly haired. Pro- middle then rounded and again narrowed to notum closely and evenly punctured except lor the marginal spine; both spines moderately a short glabrous line in the middle ai the ape.v. develuped; humeral told well developed and ut the sides gradually rounded and narrowed angled. Undcrstirfaec evenly and shallowly from base to apex. Elytra costate. each inter- punctured; wiih long hairs. val with a row of deep punctures and faintly transversely wrinkled: parallel-sided lu jusi Skt. Males &&--£ 0s2i K~3J ' 0.0R mm (ll). before the middle rounded and tapered to Fenuites 9.4 ± 0.13 v 3.4 s= ti.til mm (7), then the sharp marginal spine, sutural spine well Diunhtiwon. New* South Wales and Queens- developed: humeral fold well developed and land. angled. Undersurfaee closely nml evenly pune Ginttal remarks. The type of A. stnmdi Obcn- lured, the punctures shallower on the prostcr- hergcr is a small V specimen uf A. dtitu'tpex. nal process than elsewhere; lightly haired. The externa! morphology of this species is like Size. Miiles 7.1 ± 0.14 X 2.9 ± 0.05 mm (6). that of A. Muitet'si and A. sanwuctll but male Females 7.7 ± 0.10 x 3.2 J* 04 (9). genitalia (Fig. 22S) is different. 1 do not group Distribution. Queensland. it with the other two species. Specimens e\ SAM, I. 201*44. FIGS 1c, 12, 21 Allotype; V. tidutt^alba, yjd On C. equisciijolia. lypr, Ilolotype: $, Montiro. MM (seen hy 15.xiUP73, fi, H, Adorns, SAM, I 209V). K author). l aratypes: 3 £ & 3 £, HduueaJba, QUI on C. rtfinsrtifolia, l5.Tn.J97*, E. iC S. Adams. EA Colotdr. Shiny. Head metallic green ai the apex (1 <3 & 1 S), ANIC (T 6' * 1 2), BM (1 2 & and sides, purple nl lhe base: antennae black I c?); I d & i ?. rdtmgulba, QM un C. with coppery -purple reflections. Pronotum etfaiseltfoliu, ?.?.Ni.l!>74 & S.xii 1974. £ «* £ O S <* I2f> S. BARKER ker; the humoral fold and running along the ouier 23 S & 9 V. Orunmeir on C nana, 26. i. 1^63, margin to the middle then extending obliquely S. Barker; 3 eft lf> km S of Countceativ SUiion on C nana, 3.H.1963, Barker; 20 rf'A towards the suture, reaching slightly below but £ 15 ?. Klutnyunis Station on C, nana 9ii.l963. S. Bar not touching (he middle of the three median ker. spots; after the middle a fascia commencing ai the outei margin ;tnd running transversely 12, Astraoiis (As^raeos) crassus van tie Poll. towards the suture but not touching it; an elon- 1889: 85, 95-97. p|. 2, %. 9, 9«. Kci re- gate preaptciil Close suture spot, to the but not mans. 1892; 101; 1SI021 148. Carter, touching it. Undersurface deep coppery-purple; 1929: 282. Obetiberger, 1930: 365v halts silver. Legs deep coppery-purple, ends AstwvuA fUnopictus van de Poll* 3S86: ISO. of tibia, first and second tarsal segment* the PE 3B biown, third ;ind fourth tarsal segments d;*rk hrown. Type. Holotype, MNHN (not seen by author 1. Colour. Shape and sculpture. Mead closely punctured; Shiny. Rend and pronotum black with purple or blue purple with a median keel; hairy. Pronotum more and reflections; antennae black with blue and purple reflections. closely puneiured fit the sides than in the Elytra black with blue middle, with a median longitudinal impressed and purple reflections each wilh the following yellow line visible nt the base and apex only; parallel- markings: the basic pattern consists stded at the bu*c, gently rounded until after of irregular spots in two rows; lour the middle, strongly rounded and narrowed to spots in the middle near the suture, but not touching it, the apex, projecting forward lightly in the one at the bu%c, one before the middle, middle uf the apical edge; hairy- Elytra eoslate. one after the middle and one in the preapical area, three spots the intervals between flat, each with a row along the margin, ol shallow punctures; parallel-sided from the one ft] the shoulder and coveting ihe humeral fold, base, rounded off well after the middle and one ot the middle and one after the narrowed strongly to Ihe murginal spine; both middle; there may also be a few irregular spnts spines well developed, humeral fold moder- and in two specimens there are spots at the of the ately developed and nngled. Undersurface; base sutur.it spine. Undersurface and sternal segments finely and sparsely punctate lee.s black with blue and purple reflection*; silver. in the middle, deeper and larger punctures at hairs the sides: abdominal Ntciniies finely and closely Shape and sculpture. Head with a median keel; punctured; hairy. evenly punctured; hairy. Pronotum with small and sparse punctures in the middle leaving Ske. Males 9.6 ± 0.10 x 3.5 ± ffcgjl mm £50*. glabrous areas, larger and more dense at the Females 9.8 £ 0.15 x 3.6 ±0.07 mm (35). sides with a short impressed line projecting Distribution. New South Wales. forwards from the basal crypt; gently rounded General remarks'. The species was described hy and narrowed at the aides from the base |o Carter from a unique specimen In the MacLeay two-thirds way to ihe apex, then tapered to Museum* Only two of the S5 specimen* I the apex with the apical edge projecting have collected conform to the colour pattern slightly in the middle. Elytra punctate-striatc of the type. I he pattern on the elylr;. is with the intervals flat towards the base and variable, ranging from the basic pattern as slightly concave towards ihe apex, each with described herein (nine specimens) to patterns a row of shallow punctures; parallel-sided until in which all of the yellow markings are con- after the middle, then gently rounded and fluent, giving the appearance of yellow elytra narrowed to the small marginal spine; suttiTai with black, borders. Between the extreme*, in- spine well developed; humeral fold poorlv termediary patterns occur, including the intri- developed but slightly angled. Undersurface cate pattern of Carter's type. There is no dif- shallowly punctured, more closely at the sides ference between the sexes in si/.e or colour than in the middle; hairy. pattern. All of the specimens I collected were Size. Mates 14.4 rt: 0.25 x 5.3 ' 0.06 mm (5) taken on low Casiwrina nana heath in the FcmaJcs J6.2 ± 0.40 x 6.3 * 0.15 mm (7). Minuma and Kybcao Ranges. Monaro District, Distribution, Queensland and New South N.S.W. I do not group A, intricatus with any Wales. other species. Gcnentf remarks. This species cannot be Specimens examined. N.S.W.: Type; 5 <$ & Ml ?, Ornnmeir on Cnsuarti^a inula, 9.xii 1962. S. Bar- grouped with an} other REVISION OF BUl'KESTID GENUS ASTR4EUS t.APORTC & GORV 127 Spotirnvns exmwrtcd. Qid: I xi-, £i SuUoh. NSW is found in low rainfall areas of three states, I Barker. I d. Khanyunis Station, tvlinumn on C. species. tittorotisr 26.1.1963. 5. Barker. Spvcintcttv CMittiiWii, M-*. Ann.: I o*. WAM 73 2^. Astraetrs (Astracus) mnjor Blackburn, 62> I cm hitm, SAM; I ol- 1S90; 1257, 125S; 1891:496. Kerremans, Rurtlicfuh. JS.ix.1957. £ Barker. WAM. 73-60*. I 1892: (02; t902: 149. V, Cnllvim, fi.ix.l9tfl. /?. r MMUWi\ WAM, 73-61. Sf. A urt.: I Monartu. /. G. O. Ifpfivn A-itiwti* ntworthts var. major- Carter, 1929; % SAM. I'it:.: 1 B, Sea Lake, Gomiic, NM. 2S2. Obenbcrgci. J930; 366. FIGS I7,23C 24 Aslraciis (AsUaeus) navarchis I Phonicon) Type. Holoiype, HM inor s:en by author). Corto$rtatha rtavunhix Thomson, 1856: Colour. Shiny. Head coppery with metallic 115, W6> pi. 6, fig. 2. Masters, (886; 79 t reflections; .antennae coppery-green or blue. Astttieus navi/nhif: Saunders, 1S7J; 4J. Prouotum dark blue with purple reflections. van de Poll. 1886: 176. Masters, 1886; tlytr.i dark blue with blue and purple reflec- 71. van de Poll, 1S89: 84, 91. 92, pi. 2. tions, each elytron with the following yellow fig 5, 5a. Blackburn. 1890: 1257. Kerre- markings a Fascia running from the shoulder mans, 1892; 102- Blackburn, 1892. 211. iransversely across the base towards the \uture Kerremans, 1902; US. Carter. 1929; but not touching it; before the middle a thick 282. Obenbcrger, 1930: 366. fascia funning frv>m the margin transversely FTG. 23D to the suture; after the middle there in a fascia Type. Holotype. MNHN l not seen by author). commencing at the margin, running towards Colour. Shiny- Head coppery at the apex, dark the suture but uot reaching it. There is an oval hluc and purple at the base; antennae dark red spot in Ihe region of thy marginal spine. brown, segments one and iwo with golden- The outer margins of the fascia and humeral green reflections, the rest wilh blue and purple Told arc red. Undorsurface dark blue with icflections. Pronotum coppery al the apex and metallic purple reflections; hairs yellow. Legs sides, dark blue and violet at the base, Elytra rcd-hrown; tarsi blue. black with violet reflections: each elytron with Shape ami xculpture He^d with sparse shallow two yellow fascia, the first commencing at the punctures at the base, wrinkled towards the margin and covering the humeral fold, slightly apex; with a thin median keel: hairy. Prnno- concave towards the base and touching the tum with .small shallow punctures in the suture, the second after the middle, commenc- middle, larger deeper ones at the sides with an ing, at the margin and running transversely irregular glabrous longitudinal area in the towards the suture but not touching it. Under- middle; there is a short median longitudinal surface: prothoracic steiniles metallic golden- impressed line piojecting forwards from the green, abdomen metallic violet; hairs yellow h.tsal crypt; short with the sides gradually Legs red-brown with violet reflections. rounded from base to apex. Elytra punctatc- Shape and sculpture, Head closely punctured striatc from the base for most of the length, the at the base, grooved and wrinkled at the apex; intervals between the striae convex, costate at slightly excavated between the eyes towards the apex only; parallel-sided unhl just after the base, btit wilh a thin glabrous median 1 he middle then giadually rounded to the n'lar- keel commenciny anterior to this; hairy, Pro- g*na! spine; both spines well developed: notum with small punctures in Ihe middle, bnl humeral fold poorly developed and slightly mainly lacking in the centre, forming an in- angled. Undersurface densely punctured at the definite median longitudinal glubrous line, the .\idev sparsely ami shallowly puuciuied in the punctures larger and dccpci at the sides; a middle; hairy. median longitudinal impressed line projects 5.."? Sizr. Males 1 ± 0.70 x 5.9 ± 0-22 mm «4)- forwards from the basal crypt to ihe middle, Females 15.5 ± 0.98 x 6.3 ± 0.91 mm 1 2). short, rounded at the sides and narrowed from Distribution Western Australia. South Aus- base to apex. Elytra punctaic-stnate. the in- tralia and Victoria. tervals convex at the base, flat at the apex, Gewntl r**mitks. I have no evidence of over- those in the centre with a shallow row of punc- lap in the distribution of this species, which tures, those at the sides with deep punctures; . F m S. BARKtK parallel-sided until after the middle, then and A. ttavarchis, however male genitalia is rounded and narrowed to i.hc rnargm.il spine, different (Fig. 23E). Therefore I do not group boQj spines well developed; humeral fold the three species together but place A. jraler- pooily developed and slightly angled. Under- chIus next to A. major and A. navarcbis, surface: anteriorly the- punctures arc small in Spcihthtt\ rwntineti, W. Ausl.; 3 rf & I ?. Borden the middle and larger at I he sides, on the on ffakea trif areata, November 1939. H. W. Hrown, VV abdomen uniformly r-matl; hairy. A M, 73-378/38 1 : I ?, Bnshmead, 18.xi.1939, /?. P. McMillan, W'AM 73-383. 2 - Site. Males 14.4 ±0.41 x SJ = 0.13 mm (7). & I 2. Hopetoim, 18x1946, Mrs. Morris. WAM '- Females 16.2 0-34 x (g & - 0.08 mm (5). $6-191071912-. 1 cf, Wcmblv on Davie.Mu divori- cuta, 5.ix.l970. S, Bather; 1 & Borden, Distribution. Victoria 4 WAM. 73-382, General remarks. This species shows close •ifliniiy wilh A. major 26 Astnmi* (Astraeus) ptotttoracicus van de '2 Poll. 1889: R5, 98, pi. 1 1. Specimens rvambh'd Vic: rf, SAM; 3 ?, Jan 99, 3. fig. 11a. lUC^ NM; S df &4S. NM. Kcrrcmuns, 1892: 102: JV02; 149. Carter. 1929: 282. Obenbergcr, 1930; 366. 25. A^tiaeiis (Astrueus) fralerculus van de Poll, FIG. 23 1889: 84, 92. 93, pi. 2, %s 6. 6a. Konc- mans, 1892: 102: 1902: 149. Carter. Type. Holoiypc. MNHN (not seen by author). 1929: 282. Obenhcrger. 1 930. 365. Colour. Shiny. Head, antennae and pronotum bronze, hlytra dark brown with bronze FIG. 23E rclloc- tiom. each wilh Ibc following yellow mark- type. Rolotyne seen by author). MNHN (HOC in cs: along the margin there is a vitta. com- Colour. Shiny. Head antennae and prnnoi.um mencing at ihe shoulder and ending in the hiue-black wilh blue reflections. Elytra blue- preapical area which may be broken into Mack, each elytron wi(h two yellow fascia; the several elongate spots; in the middle buL not first commencing: at the shoulder and cover- touching the Suture there is a vftta commenc- ing the humeral fold, then running trans- ing on the anterior edge and ending in the versely towards the suture but not touching it. preapical area, this is usually broken into * concave lowards ihe base; the second alter the basal spot, a spot before the middle and an middle, commencing at the margin and run- elongate spot after the middle. Undersurface ning transversely towards the suture bill not bronze; hairs silver. Ixgs red -brown. touching it. Undersurface deep metallic blue; Shape and sculpture Head Closely and evenly hairs silver. Legs deep metallic blue, punctured; no median keel; hairy. Pronotum Shape and sculpture. Heat! shallow ly and with puncture* wrinkled at the front and sides, sparsely punctured; with a small median keel; projecting conically in the middle where the hairy. Pronotum sparsely hut evenly punc- puncture coalesce and become siriynve; tured, the punctures at the sides larger and gently rounded at the sides from the base to deepei than those rn ihe middle; a short the middle, then tapered and narrowed to the medinn longitudinal impressed line projects apex, the apical edge brood ly convex in the lorw.uds from Ihe basul crypt; short the sides middle; Sairy, Hlytra costate, the intervals flat tapering acutely from base to apex, slightly with a deep row of punctures; parallel-si tied rounded in ihe middle. Elytra cost ate, the in- until alter the middle then very gently rounded tervals ftai v\ilh irregular shallow punctures; to the marginal spine; both spines poorly parallel-sided until after the middle, then developed, blunt and close together; humeral rounded and nai rowed to I he marginal spine; fold poorly developed and slightly angled. httih spines well developed; humeral fold Undersurface: punctures close* at ihe sid«s poorly developed and slightly angled. Under- than in the middle: legs and whole under- surface with small shallow punctures in the surface densely covered with hair. middle, larj/er and deeper punctures ai the Size. Males 10.4 l 0-33 x 3,7 ± 0.13 mm \%\. sides; hairy. Females 10.5 ± 0.25 x 3.9 * 0.17 mm (6). Sir.c. Males 9.3 t 0.30 x 4.2 ± 0,10 mm (7). Distribution. Western Australia and Queens- Females 8.8 = 0.30 x 4.0 ± 0.1 1 mm (3). land Disirihmion. Western Australia. (tenerr.il remarks, van dc Poll ( 1 889) listed the General remarks. I he external morphology uf two specimens he had as coming from this species is very similar to that of A. major Clarence River and Champion Bay, and Cjif- REVISION OF BUPRF.STID GENUS ASTRAEVS LAPORTE & GORY I2!> ter (1929) gave the range as Clarence River. Females li.8 ± 0.28 x 4.2 ^ 0.J I mm H2>. N.S.W. It is now known that in Western Aus- Bistrihwion. Western Australia. specie* is associated with Banksia tralia the Genet'ril remark?. A Size. Male 9.9 x 3.7 < t )- late-striate at the base costate towards the mm apex, the intervals flat and slightly trans- Distribution. Western Australia. versely wrinkled; more or less parallel-sided, Specimens exatniftcd. W. Ausl.: I H*. W&lnllftgi rounded after the middle to the marginal spine: 10.xiL19M), R~ P. McMitlnn, WA.VT, 71-1761. hoth spines well developed; humeral fold 29- Astraeits (Astraeus) flavopjetus LaPoTte A slightly angled. poorly developed, Undersur- Gory. 1837 2. pi. 1. fig 1 Imhnff. 1856. closely face punctured; hairy. 46. Lacordaire, 1 857: 43. Geinminget A Size. Males 10,5 =t 0.4 x 3,7 * 0.13 mm <$* de Harold. IK69; 1380 Masters. 1K7I: 130 S- RAKKbR 124. Saunders, 1871 ; 43. Kcrremans. 1885: 136. van de Poll, ISS6: 176. 177. Koiotypc: tf, 77 km along main York Rd. W. Aiisl- on Ca\uaihm hut'fHlfwia, 21.x. 1970. S. Musters, 1886; 71. van de Poll, 1889* fittt&rt, SAM. 1 20955. 85, 97 98. pf. S, fig. 10, 10a. kerremans, Allotype; =?, 77 km along fgAto "York Kcl, W. 1892; 101; 1902" J 48. Carter. 1929: 282. Aiwl. on C liiivnetwtKi, 5 i. 19f>8, S. Batk,r, Obcnbctger. 1930: liS5. Barker, 1964: SAM. t 20956. Paraiypes: I t?), SAM (4 rf & 2 ?>, WAM (1 rf); I * lions, with the following yellow markings in tf 2 ? II km S of Walebing, W, Aust, two rows, one in the middle but not touching 6.x. 197 1, K. T. Rivhntth, WAIM: 1 & 77 the suture, the other along the margin; in the fcm alone main York Rd. W. Aust. on t xiuddle; an elongate basal spot; a transverse iHn'Mthwu, 5.1.1968, ,V. Barker. WAM. bar before the middle: an elongate spot aiier Colour. Shiny. Head black with purple reflec- ihe middle; a long thin preapical mark. Along tions: antennae black With metallic bine the margin: an elongate spot from the shoulder reflections, each ol segments 1-4 with dark covering (he humeral fold; a spot in the brown apex. Pronotum black with blue-green middle: a spot alter the middle. The three- reflections in the middle, purple reflections spots along the margin may coalesce forming a in the front and at the sides. Elytra Hack with single vitta or be separated into two or three purple reflections, each with the following yellow spnis, Understirfacc and legs bronze; hairs sil- markings: a large spot at the base; a ver. fascia commencing at Ihe shoulder, covering the humeral fold and running Shape and sculpture. Head densely punctured; transversely towards the suture but not touching h, con- no median keel; hairy. Pronotum closely and cave towards the base (sometimes broken into evenly punctured: basal half pnralleUsided, a marginal and H medial spot): a small spot thereafter rounded to the apex; apical edge after the middle on the outer margin, projecting forward in the middle; a short im- an oval preapical spot; between the last two spois pressed line projects forwards from the basal a shon fascia, running From the margin, half crypt: hairy. Elytra cosiate. the iniervals flat way to the suture. Undeisurface metallic but deeply punctured and slightly wrinkled; purple and bronze; hairs silver. Legs rcd- parallel-sided from base until after Ihe middle, brown. then narrowed to the small marginal spine: sutural spine short; humeral fold moderately Shape and sculpture. Head closely punctured; developed and angled. Under.surface closely no medinn keel; hairy. Pronotum deeply and sides, and evenly punctured: densely h Harkrr; I \ c? & 13 9, 64 km along main York Rd, at the sides; abdominal segments finely and un Jacksonia sp., ti.xi. (970. 5. Barker, closely punctured; hairy. Size. Males 10.5 ± 0.10 x 3.9 ± 0.04 mm 30. Astraeus (Astraeu*) iijueruillvmi sp, new, (47). Female* 11.7 = 15 >. 4.3 ± 0.06 mm FIGS 13, Til 1 KbVTStON OF ttUPROSTID GENUS ASTRABVS LAPORTE & GORY i Pt-itrihufson, Western Australia. longitudinal channels on apical part; uo General rvmurhx. In same specie* trie marginal median keel; hairy, Pronotum evenly punc spot is reduced ox lost, in others (here is an cured, with n median glabrous line at the base additional spot close to the suture and between and apex: sides rounded gradually from base the two fascia. Named lifter Mr R. P to ape*; harry. Elytra eo-state, the intervals McMillan. between flat, each with a row of shallow punc- until Specimens examined. W. AuU.: Types; 5 of & 4 ?, tures; parallcl-stded just after the middle, 77 km alone mam York Rd on (.'osuarina sp., then rounded off to the strong marginal spine; 5.i.l968, .9. Darker; 11 j. Diyandra oq Cusuarirta well developed sutural spine; humeral [old sp, 21 x. 1970. 5. Barker; 6 cf & 6 9, Dryandra on 1 poorly developed but slightly -angled. Under- 1 Catuarhta httt^lionci t 1 xi .1970, 5 fi4/&4 2 -d * punctured, hairy. 2 ?, 6 km E of North Bannister on C fytteuc liana, surface evenly and shallowly 5". 4 77 along 19jaJ970, JfVA<7, 6 c* & 9. km Size. Male* 9.X * 0.17 x 3.7 ± 0.06 mm (8). main York Rd on C. hucxoliutw, 2J.xi 1^70. S. Females 10.6 ± 23 x 4.0 -i 0.1 1 mm (7). /toAe/. I rf & 3 9* 77 km along main York Rd on C. httrgetiorta, 7.*ii. 1970, .$ Barker, $ J & Dixtrshution. Western Australia. 1 ?, Dryandra on C huexeliami, 8.xii.t970. 5. /?ur- (leneral rentarta. Named after Mr K. Carnaby. At'r; L rfi Dryandra on C, huepxliam, 12t,l973. Specimens examined, Types only. ;M. AUraeus tAstraeus) camabyi sp. nov. 32. Astraens (Astroeus) budenj van dc Poll, FIGS Id. 14, 23K 1889: 84, 93. 94, pi. 2. fig. 7, 7a. Black- Types. burn. 1891: 496. Kerremans, 1892. 101. Ilolotype: r?, 16 km NE of Lake Grace, W. van dc Poll, 1892: 67. Blackburn, 1895: Aust., on Ca\uurhtu ImegeUoiia, 9.1.1972, Carter, /:. MNHN (1 «?) WAM fl?); 1 9, 24 km S blue reflections, tips of first and second seg- of Lake Kjn&, W Aust. on C. knvxelumtt, ments brown. Elytra: each elytron with the 25J.I973. 5. Knftir* MNHN; 4 touching jt: a small prcapical spot. Under- middle; sit the sides gradually rounded and surl'aee metallic purple; hairs silver. legs dark, nan owed fiom base to apex; hairy. Elytra upper sides with metallic blue reflections, costate, the intervals between flat each with a undersides metallic purple. row of shallow punctures; parallel-sided until Shape and sculpture. Head with medium sized after the middle then rounded olT lo the mar- punctures on basal half merging into irregular ginal spiuc. strong sutural spine; humeral fold 132 S. BARKER •a .5 9 LO S g ©0 K a g REVISION OF BUPREST1D GENUS AS'i'KAtVS LAPORTH it GORY 133 pooxly developed but alightly angled Under- middle, a prcaptcal spot Undersurface and Ices .surface, punctmev sparser In the middle than metallic bronze-green; hairs silver. at the aides' hairy. Straps and sculpture. Head evenly punctured, $&*. Males S.8 i 0.14 x 3.4 ± 0.07 mm (16). stichtly excavated in the tnt-ddte between the Females S.^x0,|6x 3.5 ± 0.07 mm (17). eyes; no median kech hairy. Pronotum evenly punctured: a short longitudinal median Distribution. All of mainland Australia except impressed line projects forwards from the basal the Northern Tctntorv. crypt; at the sides rounded basal ly, then General rework*, This species was described tapered and narrowed to the apex; hairy, Elytra almost simultaneously by van de Poll as A. costatc, the intervals between llat. each with htnfeni and by Blackburn as A meyricki, a deep row of punctures and faintly trans- Blackburn (1891) recognised A. titcyrieki a* versely wrinkled: parallel-sided until after the h*tcleni but later changed his u synonym of A. t middle, (hen rounded and narrowed to the mind and cnlled A. meyr'tcki a good species marginal spine: both spines well developed: (Blackburn 1S95) Distance between the two humeral fold poorly developed and slightly populations was the main argument used by angled. Undersurfacc with small shallow punc- Blackburn (1895) in favour of calling the tures in the middle, larger and deeper at the eastern and western representatives two dis- sides; hairy. tinct species. I have been unable to separate Size. Male*. 8.1 ± 0.08 x 3.2 ± 0.04 mm (37>. specimens collected in Western Australia from Females 8.6 ± 0.15x3.4 ±0.06 mm (12). those collected in South Australia. Distribution. Mainland eastern Australia from Specimens examined. 6". Aust.; Type; 1 c? & X ?. South Australia to Queensland. Morgan, A, M. Lea, SAM: 1 ?, Murray Bridge, Ctnetui remarks. yellow pattern Is Oct. 191 1, SAM; 8 rf 4 6 ?, on Melaleuca sp., The Dernu Pass (probably the same as Puttapa Gap). variable; there aie either six spots and a fascia 21 km S of Copley, 25.x.) 969, ,V WT^fe/ttfi on each elytron or the fascia may be broken SAM; 7 3 & 9 ?, Puttapa dap. Flinders Rangev in the middle giving a total of eight spots on Melaleuca t/iomerata, 21.X.197I, S. ftarkct, W. Ausi.t Patalype of A. tnevrickt Blackburn. Specimens' examined. S. Aust,: 2 ?, McDonald SAM; 1 ?, T8 km SW of Three Springs on Dry- Femes N.P. on CaVlttis prei.wii, Z.xi. 1967, A, Bar* andra cirsioides, $.xi.l968, N. McFxvland, SAM: krf 1 o* & I ?, 2 km E of Hiii (ley on C. un'tswi, 3 S & 3 H, luianda rockholc. 106 km S of Balla- 15.xi.1969, S, Barker; 7 6* & 7 ?, McDonald donia on CaltitrFx preissii. 9xii.l974, 5. Barker. Ferries N.P. on C. prvissii, 15.xi.19f>9, S* Barker; -? K.S.fy,: 1 2, W. du Boutay, WAM, 73-5-1. QUI 2 & 2 9, on road N of Pa era Wina N.P. on C. <* 1 Type £ A. kaleni var disiunetus ObenbergeT. pr'fissii, lO.xii.196V, S. Rnrket: 7 & 1 5, Tothill NMP. 21 990. Ranges near Brady Creek on C. preissii, IZjuLT969« S. Barker; I 9. Onknparinpu Gorge near Hack fi am on Caltitri'i rhomhofdeus, VV Astraews fAstraeus) unison i van de Poll, 27.Xii.1V6V, 8. Barker; II suture bui not touching it; in the middle near FIG. 23N the suture bu; not touching it, a spot at the Type. Ho]ot>pc black with purple ami blue neJU'-ciiuns, Elytra J5 Astmeim WriiUv. \ 1 5. ^ t 77 km along mam xOci Rd. on FIGS 230 C. fuwxelianu, 7,\iU970. 5. Barker; 3 S & 2 ?. Type* xii. Wnnnnma! oh € htteveUatm, 10, 1H7", S. Bar- Holutype: rf, 3S3 km ahmjj: Payne'* Find Rd* ker; 2 $, 135 km along Albany Highway on (7. W. Aust, on Ca.wiatina otehiana, 17 iX. 1970. hiwxeliana, KU.I973, 5. Barker. S. Barker. SAM, 1 201M9. REVISION OF BUPRES'UD GENUS AStRAEOS LAPORTO & OORV 135 Allotype: & Lake King. W. Aust.. 18.xii.197i)> (13). Females J 2.0 ± 0.92 X 4.3 ±ft4rtmm £. <$ A. Cartmhy. SAM. T 20950. O). Patau-pes: 3 J, 383 km along Pavne's hind Rd, Distribution Western Australia. Aust on fftetitanth n.ix.WO, S. Bar- W , C General remarks, A. carters show* closest ker, ANIC (1 i), BM (1 ?), MNHN t L rfjj 3 but not touching it: three spot* close to the Colour. Shiny, Head, antennae and pronotum suture but not touching it, the first Ihc largest black with bronze, blue or purple reflections, above the middle, the next smaller below the or a combination of these colours. Elytra black, middle and a longitudinally elongate prcapical with purple reflections, each with the following spot (the first of these sometimes coalesces yellow markings: a basal spot; a fascia com- with the spot covering the humeral fold to mencing at the shoulder and covering the farm a fascia, concave towards the base). humeral fold, running towards the suture but t;ndersurfacc and legs dark metallic purple; not touching it, concave towards the base; a hairs silver. spot at the middle touching the margin; a pre- apical spot; a spot midway between the two Shape and .sculpture. Head closely punctured; previously mentioned marks, elongate and keel; hairy. Pronotum evenly punc- no median touching the margin; a spot near the suture, tured; with a short median longitudinal but not touching it; midway hetween the fascia project forwards from the impressed line ing and the pneapical spot Undersurface and legs basal crypt, running forwards from the purple, coppery and bronze; hairs silver. impressed line is a glabrous line formed by Shape and sculpture. Head closely and evenly lack of perforations, better defined in females punctured; slightly excavated in the middle where it reaches the anterior margin than in between the bases of the eyes; no median keel; mattes where it runs only ro the middle, sides hairy, Pronotum with punctures larger at the rounded and narrowed to the strong marginal sides than in the middle, median longitudinal apex; dorsally flattened in lateral profile; hairy glabrous line from base to apex formed by the at the sides but less so in the middle. Elytra absence of punctures; gently rounded at the costalc, the. intervals between flat with a row sides from base to apex; hairy. Elytra costate of shallow punctures; sides at first diverging at the apex, punctate-striafe at the base, the slightly outwards from the base then parallel- inteivals between flat at the apex and convex sided to before the middle* then gradually at the base, each with a row of punctures: rounded and narrowed to the strong marginal parallel-sideU to the middle then rounded and Spines; well developed sutural spines: humeral narrowed to the marginal spine, which is well fold poorly developed but slightly angled. developed: sutural spine sharp but shortened Undcrsurfacc finely and sparsely punctured in by the sutural margin being straight and super- the middle more closely at the sides: densely ficially appearing to be broken; humeral fold hairy pooTly developed and slightly angled. UnJer* Site. Males 1 1.2 ± 0.16 x 4.0 ± 0.0S mm surface evenly punctured, the punctures S. BARKKR 5 m 5m m Fig. (7. Asiraeus major Blackburn Fig. 18. Astwus watsoni sp, nov, deeper at the sides than in the middle; hairy. Colour. Shiny, Head and pronotum bluc-grccn: In the male the lasi ahdominal slernire has p antennae black with blue reflections. Elytra marginal indentation in the centre. black with blue-green reflections, each with the Size. Males 10.8 £ 0.12 x 3.8 ± 0.05 mm following yellow markings; a large elongate (26). Females 11.7 =t 0.19 » 4.2 ±; 0.07 mm basal spot not reaching the anterior margin or <22>. suture; a fascia at the middle, expanded towards the apex near the lateral margin bui Pitfribution, Western Australia. not touching it or the suture; a huge spot after General remarks. A. fioerlitigl shows features the. middle, not touching the margin or the in common with the preceding group of species Miture; a. spot covering the humeral fold; a (Fig. 23P) and also with xhc following group small elongate spot in the form of a lunette of two species, and because of this I place it hy near the preapical margin and ending at the itself between the two groups. Named after the marginal spine (present in the illustrated speci- late Mr A. Goerling. men-, ahseni in ihe holoiype) Undersurface Sp/urlmetis cxaminrd, Types; Aust.: : 14 W. 20 i & blue-green; hairs silver. Legs metallic blue. i\ Marloo Stn, Wurarga. 1951-1941. A. UoartiriK. ANIC. Shape find sculpture, Head evenly punctured: deeply excavated eyes, mainly IK, Astraeus (Astraeus) eyaneus Kerremans, between the at the base; no median keel; 1900: 295; 1902: 148. Carter. 1929: 282. sparsely covered with Obenberger, 1930: 365. long, fine hair. Antennae strongly serrate. Pronotum FIG. 19 deeply punctured at the sides, towards the middle shallow punctures with type. HoIOlype: & Standing, N.vS.VV.. DM liseen a by author). central ovoid area consisting of hexagonal RFVISION OF BLIPRFSTID GENUS ASTRAEVS LAPORTE & GORY 137 5 mm ^^^— B rti m f Fig. 19, Astraeus tiffinm Kerremans Fig. 20. Astraeus caledonivus hativel depressions, each with a small central punc- Specimens examined. N.S.W.: Type. Qld: I ?, Acacia Creek via Killarnay, Jan. ture; inflated at the sides just before the 1948, Mrs J. Harslet!, JH. middle, then straight sided and strongly tapered to the apex; the lateral lobes with their 39. Astraeus (Astraeus) caledonicus Fauvel, apices turned downwards, convex at the apex, 1904: 116. Obenberger, 1930: 365. flattened at the base; covered with fine hair. FIG. 20 Flytra flattened; punctatc-striatc anteriorly, Type. Holotype: 2j Baie du Sud* N. Caledonie, costate posteriorly, the intervals between con- Delauney, MNHN (seen by author). Colour. vex towards the base and flat at the apex, each Upper ytirface glabrous. Head and pro- with a single longitudinal row of shallow punc- notum black with green reflections; antennae tures and slightly transversely wrinkled; paral- purple, Elytra black with yellow reflections, lel-sided to the middle, then rounded and each elytron with the following yellow mark- narrowed to the small marginal spine: sutural ings; a large basal spot; a small spot after the spine shortened by the sutural margin being middle near (he suture but not touching it: a straight and turned slightly upwards; humeral spot at the margin covering the humeral fold: fold poorly developed but slightly angled. a spot after the middle ai the margin but not Undersurface evenly but shallowly punctured touching it; and slightly behind the last a spot the it. Undersur- in the middle; lateral presternum and abdomi- near suture but not touching nal sternitcs longitudinally grooved: sparsely face black with green and purple reflections; haired. hairs silver. Legs red-brown with purple reflec- tions; tarsi dark-brown with blue reflections. Size. Males 11.6 x 3.9 mm (1) Females 13.9 Shape and sculpture. Head shallowly bul x 4.8 mm CI). evenly punctured; excavated between the eyes Distribution. New South Wales and Queens- mainly at the apex; no median keel; without land. hairs. Pronotum shallowly but sparsely punc- 138 S. BARKhR 5 mm Fig. 21. Astraeus robmtus sp. nov REVISION OF BUPRESTID GENUS ASTRAEUS LAPORTE & GORY 139 ^ A /!\ A B c D E F rw /l\ i\\ /T\ /1\ rt\ /I\ M N (II m r ^\ /\\ /l\ rf\ i_ lmm_i Fig. 22. Outline diagrams of the parameres of male Astraeus (Depollus) species (A-H) i_1m m-i and Astraeus (Astraeus) species (I-T), dorsal surface uppermost. A-polli; B- Fig. tamminensis; C-robustus; D~aberrans; 23. Outline diagrams of the parameres of male Astraeus E-lineatus; F-multinotatus; G-irregu- (Astraeus) species, dorsal surface laris; H-dedariensis; \-bakeri; J-minu- uppermost. A-intricatus; B- tus; K-fraseriensis; L-pygmaeus; M- crassus; C-major; D-navarchis; E- smythi; N-simulator; O-obscurus; P- fraterculus; V-prothoracicus; G-elonga- globosus; Q-mastersi; R-samouelli; S- tus; H-vittatus; \-flavopictus; l-macmil- dilutipes; T-adamsi. lani; K-carnabyi; L-badeni; M-jansoni; N-oberthuri; O-carteri; V-goerlingi. tured; a short but deeply impressed median spine with a straight internal longitudinal line projects forwards from the edge; humeral fold rounded and barely obvious. basal crypt; parallel-sided at the base, before Undersurface shallowly and sparsely punctured; lightly the middle rounded and obliquely narrowed to haired. the apex; median lobe short and blunt, apices of lateral lobes sharp and turned downwards. Size, Females 11.5 x 4.1 mm (1). Elytra punctate-striate, the intervals between Distribution. New Caledonia. convex with a few faint transverse wrinkles at the base, without hairs or punctures; parallel- General remarks. A. caledonicus shows close sided until after the middle, then gently affinity with A. cyaneus. rounded to the strong marginal spine; apical Specimens examined. Type only. 3 6 1 40 S BARKER. micrograph of Fig. 24. Scanning electron Fig. 25. Scanning electron micrograph of the the median lobe of the pronotum head of A. frosen'ertsis showing the showing the of A. jraseriensis median longitudinal keel. basal crypt. Index To THE Vaud St'KIF.s and Synonyms op ASTRAEUS With the Numbering System Adopted In the Text Valid Species Synonyms Acknowledgments uh^mms van de Poll 4 1 would like to thank the following people (uiarnsi sp. nov. 20 lor their assistance; Mr G. Gross, Dr E. httdvni van de Poll 32 -— nwvrhki Blackburn Mathews and Mrs B. K. Head. South Austra- buktri sp. nov. 9 lian Museum; Dr E. B. Britton, C.S.J.R.O., caledomcus Fativel 39 rarnabvi sp. nov. 3 I Division of Entomology; Mr K. Dahms, curttri sp. nov. 36 Queensland Museum; Mr A. Neboiss, National cntssus van dc Poll 22 Museum of Victoria; Dr C. N. Smilhers and cyanvus Kerremans 38 Dr D. K. McAlpine, Australian Museum; Mr dedariens'ts sp. nov. 8 ddutipes van de Poll 19 - strandi Obenberger L. Koch, Western Australian Museum: Mr B. tlvngaUtii van dc Poll 27 Levey and Miss C. M. von Hayek, British Gory 29 (lovopictus LaPorte & Museum, Monsieur A. Descarpentries, Paris fraxerit'tisis sp. nov. 1 A. Samuelson, Bernice P. jraterculus van de Poll 25 Museum; Dr G. gfobosus sp. nov. 1 Bishop Museum; Dr J. Jelinck. National ftoerlhigi sp. nov. 37 Museum of Prague; Mr K. T. Richards. iniritattis Carter 21 Western Australian Department of Agriculture: irregularis van de Poll 7 George, Western Australian Herbarium; jansoni van de Poll 33 teppvri Blackburn Mr A. lineatus van de Poll 5 Mr EL Slater. Canberra: Mr E. E. Adams, tntwrnillatii sp. nov. 30 Edungalba; Mrs J Harslett, Amiens; Dr F. H. major Blackburn 23 Uther Baker, Applecross; Mr R. P. McMillan, masters! MacKeay 17 - splendent van de Poll - simplex Blackburn Cottcsloe; Mr and Mrs K. Carnaby, Wilga. mimttus sp. nov. 10 Dr K. Bartusek, Dr S. J. Edmonds, Dr E. mnltinotatus Vfcn dc Poll 6 Wollaston. Miss R. Altmann, Miss B. Jones navorcliis (Thomson) 24 and Mr P. G. Kempster all of the University of oberthun' van de Poll 34 National Parks Board of tihxt unts sp. nov. 15 Adelaide. The polli sp. nov. i Western Australia for permission to collect in prothonicicus van de Poll 26 Flora Reserves; the Director, National Parks p\\i>tnacus van de Poll 12 and Wildlife Service of South Australia for robustus sp nov. 3 samoueilt Saunders 18 permission to collect in National Parks. The simitlutor van de Poll 14 Royal Society of South Australia provided a smyilti sp. nov. 1 grant to cover the cost of one illustration. The tammuu'Hsis sp. nov. 2 Australian Biological Resources Committee vtttarns van de Poll 28 watsoni sp. nov. 35 provided a grant-in-aid of research. — — REVISION OF BUPKESTID GENUS ASTR,1EUS LAPORTE & GORY 141 References Barker, S. (1964). Asthraeus LaPorte and Gory, K.erk£M.vns, C. (1892).—Catalogue synonymique 1837 {Insecta; Coleoptera): Proposed emen- des Buprestides decrits de 1758 & 1890. dation to Astraeus. Butt. zooi. Namend. 21, Menu Soc. enl. Bela. 1. 1-304. 306-307, Kfrrrmans. C. (1900).—Buprestides nouvcaux ct Blackburn, T. ( 1890) . —Notes on Australian remarques svnonymiques. Ann. Soc. ent. Belg. Coleoptera, with description!; ol" new species, 44, 282-351! pan v. Proc. Litw. Soc. N.SW, 4, 1247-1276. Kkrrkmans, C. (1902). —Genera lnsectorum 12. Blackburn, T. (1891).— Notes on Australian Coleoptera; Senicornia Fam. Buprestidae. Coleoptera, with descriptions of new species. Lacordaire, T. (1857). —"Histoire naturelle des Proc. Linn. Soc. N.S.W. 5, 479-550. Insectes." Genera des Coleopleres, Vol. 4. Biackbukn, T. < J892) . Further notes on Aus- — (LibraJrie Encyclopedique de Roret: Paris.) tralian Coleoptera with descriptions of new F, Gory, H. ( 1837).— 'Histoire genera and species. Trans. H. Soc. S. Attst. L^PORTr, L, & 15, 207-261. Nalurcllc ct tomographic des lnseclcs Blackburn, T. (1894).—Notes on Australian Colcoptercs." Tome I. Monographic des spi- Coleoptera with descriptions of new species. locera. euiydera, nyeieis, eunosuis. Bupies- lides (Chrysoehroiles et AgriUtes). Pmt: Linn. Soc. NSW. 8, 85-108. tP. Buackbcrn, T. (1895).— Further notes on Aus- Dumenil* Pari*.) tralian Coleoptera, with descriptions of new MacI.raY, W. (1873).—Notes on a collection of genera and species. Trans, tf. Soc, S, Aust. insects from Gayndah. Trans, ent. Soc. 19. 27-60. N.S.W. 2, 239-318. Brttton, JS R. (1970).—Coleoptera- in ''Insects Mastkks, G, (1871).—''Catalogue of the des- of Australia". Ch. 30. (Melbourne University cribed Coleoptera of Australia." (Sydney.) Press.) Masters, G. (1886).—-Catalogue of the des Carter* H. J. (1925). Revision of the Austra- — eiibed Coleoptera of Australia- Part HE. lian species of Chrysobothrls (Fam. Bupres- Proc. Linn. Soc. N.S.W. II, 1-106. tidae), together with notes, and descriptions of new species of Coleoptera. Proc. Linn. OnrNfirRriRR, J. (1928).—Opuscula B;ipresto- Soc. N.S.W. 50. 225-244. logica I. Arch. Naturxexch. 92, 113-224. Carter, H. J-, (1929).—A cheek list of the Aus- Ouj-nihw(.;h,k, (. (1930). —In **Catalogus Coleop- tralian Buprestidae. Aust. Zooi 5, 265 304. terus." Vol. 12, Buprestidae 2, pp. 365-367. Carter, H. 1. list of 4 (1931).—ChecV the Aus- Oblnblroer, I. (1936).— "Festschrift zum 60 tralian Buprestidae, Corrigenda. Aust, Zooi. Gebudst Embrik Strand, Rica." Vol. I, pp. 6, 107-108. 97-145. Carter, H. J. (1933).—"Gulliver in the hush." van de Poll, J. R. H. N. (1886).—Description of (Angus & Robertson: Sydney.) three new species and a synopsis of the Bup- Fauvfl, A. (1904).—Fatme analylique des restid genus Astraeus, C. & G. Notts Leviien la Coleopleres de Nouvelle-Caledonie. Rev. Mtts. 8, 176-180. d'Ent.JX 113-208. van nE Poli > J, R. H. N. (1889).—Monographi- CJemminger, Dr. & de Harold, B. (1869). cal essay on the Australian Buprestid genus "Catalogus Coleoptorum hucusque descrip- Astraeus C. et G- Tjschr, v, Ent, 32, 79-1 10. torum synonymicus ct systcmaticus." Vol. 5. (E. H. Gumnti, Monachii.) van de Volu J. R, H. N. (1892).—Note sur quel- 1.CZ.N. (1966).—ASTHRAEUS La Porte & ques especies d' Astraeus, Tjschr. m. Em. Jfi, Gory, 1837 (Insecta, Coleoptera): Validation 67-68. of emendation to ASTRAEUS. Ball. Zooi. Saunders, E. (1868). A revision of tbe Aus Nonwnc. 23, 269-270. tralian Buprestidae described by The Rev. F. Trans, cut. 1S68. Imhoff, T.. (1856).—"Dersuch einer Eiufuhrung W. Hope. Soc. Land. in das studium der Koleoptcrn." (Basel.) 1-68. — Kr-KUEMANS, C. (1885).—Enumeration dc Bup- Saunders, E. (1871). "Catalogus Buprcstidarum restides decrits posteneuremcni au Catalogue synonymicus et systematieus.'' de M. M. Gemminger & de Harold IS70- Thomson. J. (1856).—Description dc dix Colcop- 1883. Ann, Soc. ent Bel}>, 29, 11W57. teres. Rev. Mas. Zooi. 2, 112-118. FACTORS AFFECTING THE DISTRIBUTION OF THE LEPTODACTYLID FROG GEOCRINIA LAEVIS IN THE SOUTH-EAST OF SOUTH AUSTRALIA byR. G. Beck* Summary BECK, R. G. (1975). -Factors affecting the distribution of the leptodactylid frog Geocrinia laevis in the south-east of South Australia. Trans. R. Soc. S. Aust. 99(3), 143-147, 30 August, 1975. Geocrinia laevis (Giinther), a species of leptodactylid frog previously recorded from Victoria and Tasmania, was first reported in South Australia in 1966. A survey has revealed that this new population is an extension of that known to exist in south-west Victoria. The frog is, extremely rare in South Australia, occupying only a fraction of the potential habitat for which it shows preference. Some thoughts on this are tendered as a basis for further studies. FACTORS AFFECTING THE DISTRIBUTION OF THE LEPTODACTYLID FROG GEOCR1NIA LAEV1S IN THE SOUTH-EAST OF SOUTH AUSTRALIA by R. G. Beck* Summary Bkck, R. G. (1975). —Factors affecting the distribution of the leptodactylid frog Geocrinia luevis in the south-east of South Australia, Trans- 7?. Soc. S. Artst, 99(3), 143-147, 30 August, [975. Geitcruihi laevis (G anther), a species of leptodactylid frog previously recorded from Victoria and Tasmania, was first reported in South Australia in 1966. A survey has revealed that thi.s new population is an extension of that known to exist in smith-west Victoria, The frog i3 extremely rare in South Australia, occupying only a fraction of the potential habitat for which it shows preference. Some thoughts on this are tendered as a basis for further studies. Introduction survey, field work was concentrated around The leptodactylid frog GeotrJnta laevis Marsh's Swamp (Site 3, Fig. J. Grid reference (Ciunther) has been described in detail by 353362, Australian Army Survey Map, Penola, I jttlejohn & Martin ( 1 964 ) . The specimens 1:250,(100 Sheet SJ/54-6). From here the sur- found in the south-east of South Australia vey extended to the north-west, following the have been up to 25 mm long, with dorsal general /one of influence oi the Reedy Creek skin dark grey and slightly warty, ventral skin drainage system, and to the south-east along smooth and paler grey, All had distinctive the Dismal Swamp complex to the Glenelg pink markings on the groin and thighs and River. Most field work was carried out during some specimens showed this colour under the the daylight, but night road surveys were con- forearms as well. ducted in likely areas, yielding one specimen Prior to 1966, Geocrinia laevts was known only. to exist in four disjunct populations in Aus- When six specimens had been found, tralia: Tasmania, King Island, the Grampians, detailed botanical surveys were made of the and the area in south-west Victoria from surrounding areas, particularly of the nearest Dartmoor to Port Campbell (Fig. 1). With the probable breeding site. Following the establish- discovery (Woodruff & Tyler 1968) of a speci- ment of definite ecological patterns, soil sur- men from Marsh's Swamp near Me Burr, some veys of these areas were undertaken. SO km west of Dartmoor, it was desirable to establish whether this was a fifth isolate or an Typical habitats extension of the Victorian population, Geocrinia taevis normally lays eggs in areas which later become flooded, Breeding sites Methods may be I he edges of permanent swamps, or The study area comprises the following non-permanent swampy areas, often situated Hundreds in County Grey: Mt Muirhead, to the east of sandy Tises. ranging from a few Mayurra, Riddoch, Hindmarsh, Grey, Young, to 200 m away. The soils of the rises are Nangwarry, Mingboof, Blanche and Gambier, podsolised sands, usually Mt Burr sands as and the adjoining area east of the state border described by Stephens *i ah (1941). The to (lie Gleuelg River. swamps occur in Wandillo sands, and there The survey was undertaken from 1968 to may be several intermediate soil types between 1974, and most areas were visited in both the rises and the swamps. For this reason, the summer and winter. In the early stages of the natural dry -sclerophyll forest may vary m lype Lynwood Park, Mil Lei, S. Aust. 5291. 144 R. G. BECK Fig. I, Map of the lower south-east of South Australia and adjacent south-west Victoria, showing main collecting sites of Gcocriftia laevis in relation to the Reedy Creek-Dismal Swamp Corridor. The inset shows known Australian distribution prior to 1966. (After Littlejohn & Martin 1964, 1965.) and consist of any of Eucalyptus baxferi, E. Details of the findings are given in Table 1. obliqua, E. hitheriana or E. ovata according to Representative specimens have been lodged the soil type. However, the understorey in the with the S\A. Museum. viciulty of the swamps is remarkably con- Of particular interest is the specimen col- stant, and four species have been found in all lected at Grid rcf. 376346 (site 7, Fig. 1 ), in areas, viz. Acacia nielunoxylon, Leptospermum a deep pit dug to observe pine tree root growth jurriperimmi, Melaleuca squarrosa and Heli- at the Forest Research Station north of Mt chrysitm detultoiclea. Gambier. This site is two km from the nearest soil of Stephens er al. plus The map (1941 ) potential breeding area, giving an indication of the plant indicator species greatly facilitated the actual mobility of the species, which is later survey potential areas work and most in regarded as sluggish compared with other- south-east the have now been examined, local species. Results The distribution is shown in Fig. 1, and A total of 20 specimens of G. laevis have with one exception corresponds with normal now heen found in South Australia, arid a dispersals from the Reedy Creek and Dismal further 10 in Victoria between the State bor- Swamp complexes. It is suggested that the der and the Glenelg River, which, prior to Canunda specimen (site 1, Fig, 1 ) is from a 1966, represented the known western limit of community established from eggs or larvae distribution. The species is common at Dart- washed down one of the many man-made moor, where on one occasion, 12 were found drains which cross the area between the Milli- under a log in the bed of the Glenelg River. cenl Hills and the coast, i GEOCRINIA LAEVIS IN THE SOUTH-EAST OF SOUTH AUSTRALIA 145 TABLE i Recorded distr'thnt'ton of Geocrinia laevis in south-east South Australia and nearby Victoria Refer A.A.S, Map, Penola, 1:250,000, Sheet SJ/54-6 Grid Site No. No. of Reference (Fig. 1) Specimens Collector Remarks 353362 3 I H. Minchan & First S. Aust. specimen, SAM, C. Taylor R8118 353362 3 1 D. Woodruff Melbourne University Zoology Dept. 220/67 353362 3 6 R. Beck All within 2 km of original site at Marsh's Swamp 353362 3 3 D. Klcm Marsh's Swamp. One specimen, SAM, R10583 354364 3 1 F, Aslin 2 km NE of original site at Marsh's Swamp 38(35! 6 1 C. Taylu; Earl*K, 16 km N of Ml Gambier 381353 6 1 R. Beck Hem's scrub, 17 km N of Mt Gambier 378351 6 1 F. Aslin Telford's scrub 378349 6 1 D. Klem 2 km S of Telford's scrub 376346 7 1 D. Klem Forest Research Station, Soil Pit, SAM, R 13974 368355 5 1 D.KJem Hogarth's scrub 1 J, Aslin 326360 | Canunda Reserve, SAM R 13975 342366 2 1 F. Aslin Night Road Survey, 5 km NE of MilliceiU 361361 4 ] A. Rowley Lake Leake, SAM, R 141 99 Refer A.A.S. Map, Hamilton, 1 :250.000, Sheet SJ/54-7 414325 8 6 H. Roach 16 km E of State Border on High- way No. 1, SAM, R 10780 414325 8 1 R. Beck Same locality 427324 9 Many R. Beck Common in Glenelg River 2 km up and downstream from Dart- moor Refer AA.S. Map, Portland, 1:250,000, Sheet SJ/54-11 4283 1 10 F. Aslin East of Glenelg River near Jones' Lookout 409313 12 F. Aslin Lower Glenelg River 424310 II J . Aslin Lower Glenelg River The major geological and physiographic into the Glenelg River just north and south of features of the region have been described by Dartmoor by way of the Scott and Ardoo Sprigg (1952). The Reedy Creek and Dismal Creeks. Swamp complexes are separated geologically In the study area, G. laevis is restricted to by the Gambier Upwarp, and flow in wet years areas receiving an average annual rainfall of to the north-west and south-east respectively. 700 mm or more, whereas in Tasmania and However, the watershed gradient is so gradual, Victoria it is found where rainfall is greater being only a few cm per km, that in extremely than 500 mm (Martin 1967). If the species wet years such as 1S96 and 1946 there was an in South Australia followed the Victorian rain- almost continuously wet corridor from the fall pattern, it would be reasonable to expect Kingston district to the Glenelg River. Even in the distribution to extend laterally about 100 years of normal rainfall, swamps are close km. Likewise, if it occupied all sites considered enough to provide ready access for frogs to the suitable on the basis of soil and vegetation lower south-cast the Glenelg River. from patterns, an extended distribution pattern The last occasion on which the Dismal could be expected to the extent of about 50 Swamp actually flowed was in 1946, emptying km. : |4G R, O BECK 1 Discussion continent at the same time.' This is supported, Three tacts emerge from ihe survey: on a one year basis at least, by an examina- tion of the Commonwealth Bureau of Mtrieoro J While it is certain that more specimen* will logy rainfall figures from major Australian be discovered within and beyond the present mainland centres for 1967, Perth. Broume. known range of distribution, (7. lovvis is Darwin, Cairns. Brisbane and extremely rare in south-east South Australia. Sydney had greater than average rainfall, Geraldton Even at Marsh's Swamp, where most speci- and Alice Springs were Only slightly lower than mens have been found. I have not positively average, whereas Adelaide and Melbourne tdentitled its calK during the Aprit-May- received approximately half their average Junc breeding season, when calls are com- amount. The coastal strip from the head of monly heard at Dartmoor, Victoria. How- the Great Australian Bight to c«ist of Mel- ever, Woodruff & Tyler (!96«) have re- bourne obviously was the worst affected area. ported the recording of a mating call al Marsh's Swamp. Cluirchilf (I968-) has shown that, in 2. According to prevent known records, the Western Australia, during the past 5000 years species occupies only a fraction of the poten- there have been several fluctuations in climate of sufficient magnitude to cause the replace- tial habitat for which it shows preference wilh respect to soils, vegetal ion um\ rain- ment of jarrah forests with karri and vice fall ier\a. Similarly, recent wofk by Dodson (1974) in the study area indicates variations i. G. Icwvi.s was always found under the shelter in climate, wilh relatively drv periods between of logs, litter, or stones during the day. As 5000 and 2000 R.P., and -.gain since 1300 a result, the species has not been found in BP. areas cleared tor agriculture or pasture pro- duction Gentilli (1^72) also supports the concept of changing climate. "It must be stressed thai Some suggestions for the reason tor ih'S climate, being rhe result of numerous variables restricted distribution are tendered as a banis variously combined in space and time, can for further work by someone wilb more time vary, fluctuate, oscillate, or just change." It and resources than the present author. Rain- follows that changes such as these must pro- i.iU and associated weather patterns are duce equally dramatic changes in the local probably the major factors influencing the fauna. spread of any frog species. In 1967, an extreme draught was experienced in the lower south- With pluvial conditions prevailing in east of South Australia. The aveiage annual southern Australia during the last glacial period, Bassian species extended their range rainfall at Mr, Gambier is 776 mm, but in that ( I.itllejohn ]'>67) and it is reasonable to year only 402 mm fell, and unofficial figures expect that from the Diurnal Swamp area were as low as G. fumh occupied much of the lower south-east of South Australia- 2SU and 331,1 mm respectively. In this single With the return dry year, many of the local swamps previously of present-day weather conditions, il.s considered permanent, dried up completely, range would have decreased but not to the limits today. is suggested and most of the non-permanent swamps stayed found U therefore that in the recent past, possibly dry throughout the winter. As a result, none of much more iccently than thai postulated the Spring breeders bred, and only a few of by Crocker ik the autumn arid winter breeders actually Wood 1 I947K there has been a period suffi- spawned ciently arid to cause the withdrawal of G. Ifwvis and perhaps some other species Since Crocker & Wood (1947) first presen- anuran to the favoured of Vic- ted evidence for a recent at id period, many more puTtS south-weM toria or workers have commented, and they arc about even the Grampians. It must be emphasised that in this context the "'arid" equally divided in their acceptance or rejec- term tion of the concept (Mulvancy & Golson 1971 is relative rather than absolute. Littfejohn 1967). Inoccupation of the south-east of South Gentilli i 1961 ) had already offered an Australia by G. /«em would have occurred by explanation for this diversity of opinion; ^Aus- way of the Cilenelg River and the Dismal tralia is a targe land, spanning several maior Swamp when wetter conditions returned. This climatic belts, and may have experienced dif- may have taken place even as recently js with ferent climatic change*, in various parts of the in historic times. — GEOCHINIA LAEVIS IN THE SOUTH-EAST Ob SOUTH AUSTRALIA Since the settlement of the isouth-eaKl of distribution has been prevented by a combina- South Australia by white man, the aTea tion of clearing the natural habitat, and the reached its physically wettest stale in 1S96. drying out of the district by drainage, pine This Information was obtained from S.E. plantations and the establishment of better Drainage Board records and also, some years pastures. It is obvious that much more study ago. from old residents who remembered the is required to explain satisfactorily this limited district during the nineties. They claimed "you distribution. could tow a boat across country from King- Adin owlcelgm en Is ston to the Glenelg River" While this is no The author gratefully acknowledges the doubt a slight exaggeration, it is surely signifi- col lection of specimens by the following mem- cant to the distribution of frogs. bens of the South East Field Naturalists' Colville & Holmes (1972) attribute the in- Societies: Mr and Mrs F. Aslin, Messrs D. crease of wetness during the second half of Klcm, P Roach, C. Taylor, and Miss A. Row- to the clearing of the nineteenth century ley. Also thanks are due to Mr M. ). Tyler and natural scrub. Subsequent widespread plant- Dr M. Littlejohn for advice and encourage- of drainage ing of pines and establishment ment, and to Mr D. Lewis from the S.A. Dcp1* schemes have greatly reduced surface waters of Agriculture for his help with soil identifica- Very recent reoccupation of the south cast tion. The Royal Society of South Australia v account Irtc, kindly granted $50 towards petrol by O . /tft'Ws as outlined above would ? for the limited spread of thz species. Further expenses. References J., Martin, (1964). CM rcuill, T). M. ( 19681.—The distribution Littlejohn, M. & A. A. and prehistory of Eucalyptus diversicolor F. The Crima laevis complex (Anura: Lepto- Muell., E. marxinula Donn. ex JSm,» and E, dactvlidae) in south eastern Australia. Autt. J. Zool. 12. 70-83. calophylhi R.Hr f in relation to rainfall. Ausi. I. Bot. 16, 125-51. Littlejohn, M. J.. A Makiin. A. A. (1965). The vertebrate htuna of the Bass Strait Colville, J. S« & Holmes, .1. W (1972) Water islands: 1. The Amphibia of Flinders and table fluctuations under forest and pasture King Islands. Hoc, H, Sol. Vict, 79, 7.47-256 in a karstic region of Southern Australia. Martin, A. A. (1967). —Australian Aoitran lite Hydrology 17, 61-80. Histories: Some Evolutionary and Ecological Crocker, R. L., & Wood, I. C. (1947).—Some Aspects, In A. H. Weuthcrfey (Ed.), "Aus- historical influences on the development of tralian Inland Waters and their Fauna, Eleven the South Australian vegelational communi- Studies." (Australian National University: ties and their bearing on concepts and classi- Canberra.) fication in ecology, Trans, H. Sac. S, Anst. 71, Mulvanev, D J., & Golson, .1. (1967).— 91-136. "Aboriginal Man and Environment in Aus- tralia/' (Australian National University: Dodson, J. R. ( 1974).-- Vegetation histury and Canberra.) water level fluctuations at Lake I. cake, south- Sprjgg, R. C. (1952).—The Geology of the eastern South Australia. I. 10,000 B.JV to Soulh-Easl Province. South Australia, with Present. Attst. '. Bot. 22. 719-741. Special Reference to Quaternary Coast-line Gentilll J. U96I),—Quaternary climates of the Migrations and Modern Beach Developments. Australian region. Ann. N.Y. Acad. Set. 95, South Australia Department of Mines, 465-WI. Bulletin 29. Stephens, C. G, Crocker, R, L.. Butler, B.. £ CiLNiit.LL J. (IV72).—"Australian Climatic Pat- Soil - Smith, R. (1941)—A and Land U BYTlKVAHEDELSTEIN*ANDH. B. S. WOMERSLEYf Summary EDELSTEIN, TIKVAH &, WOMERSLEY, H. B. S. (1975).-The thallus and spore development of Lobospira bicuspidata Areschoug (Dictyotales: Phaeophyta). Trans. R. Soc. S. Aust. 99(3), 149-156, 30 August, 1975. The apical growth of Lobospira bicuspidata, release of tetraspores, and growth of the spores in culture to plants up to 1 cm across, are described. Both development of the axes and growth of the sporelings is from a marginal row of apical cells, and the thallus is monopodially developed. Lobospira is therefore placed in the Zonarieae group of the Dictyotales. THE THALLUS AND SPORE DEVELOPMENT OF LOBOSPIRA BICVSPJDATA ARESCHOL'G (DICTYOTALES: PHAEOPHYTA) by TiKVah Edelstein* and H. B. S. Womersleyt Summary fcDELSTFiN, Tikvar, & Womersley. H, B, S, (1975).—The lhallus and spore development of Lubospira hicuspidata Areschouu ( Dtctyotatcs: Phaeophvta). Trans. H, Sac. S. Aitsi. 99(3), 149-156, 30 August, 1975. The apical growth of Lobospha hicuspidata, release of tetrasporcs, and growth of the spores in culture to plants up to I cm across, are described. Both development of the axes and growth of the sporelings is From a marginal row of apical cells, and the thallus is niono- podially developed. Lohcfspira is therefore placed in the Zonai ieae group of ihe Diclyolales. Introduction of the thallus, made in 1972 while the firM Lohospifa bicuspichsta Areschoug is a dis- author was on leave at the University of tinctive brown alga referred to the Dictyotales. Adelaide. It occurs from Nickol Bay, Western Australia, Methods around southern Australia to Eden, N.S.W.. Plants (ADU, A42264) were collected in and around Tasmania (Womersley 1967. p. drift at Aldinga reef. South Australia, on 27 2! 5) and is frequently abundant in regions of May, 1972, and transferred to the laboratoiy moderate To strong water movement, from in sea water. The specimens (Fig. IB) bore just below low tide level to 35 m deep. mature sporangia, many of which released The alga (Fig. 2A) is easily recognized by tetrads of spores. Fertile branches were placed its spirally twisted axes, with a phyllotaxis of in a glass jar with Provasoli ES medium about 1/3, bearing laterals with bicuspid, de- (Starr 1971, p, 359) in a 15X culture room, terminate ramuli (Harvey 1858, pi. 34). and and spores allowed to settle on slides durinv with lower branches bearing recurved attach- the next two days. On day 3 the slides with ment tendrils. Kjellman (1897, pp. 295, 297) attached sporelings were transferred to pe:n and Oltmanns (1922, p. 185) considered thai dishes (5 cm in diameter), and germanium ihe thallus develops from an apical cell, with dioxide at a concentration of 5 p. p.m. added sympodial branching, and Lobospira has thus to the medium. Single sporelings from the been considered as a member of the Dictyo- slides were detached to be grown in free cul- tcae. The sporangia, about I00/*m in diameter, ture; they were washed several times in a well- occur scattered over the thalitis (Fig. IB), they slide and inoculated into a new set of dishes are developed from cortical cells and sunken each with 15-18 sporelings. Afler 4 weeks in the thallus (Fig. IB). Neither division of the cultures were maintained in SWM 3 medium sporangia nor release, of spores has been pre- (Chen. Edektein & McLachlau 1969). viously reported, and release was only obtained As the sporelings developed, considerable by the present authors on the one occasion. difficulty occurred with bacterial (and at one 1 Sexual reproductive cells alio have never been stage fungal ) contamination. Addition of peni- observed. While Lohospira has usually been cillin to the Provasoli medium bad little effect, placed in the DictyotaJes, and the Dtctyoteae but streptomycin (100-150 mg streptomycin I Womersley 1967, p. 215), its relationships sulphate/ 1 of seawaier) eliminated most of Ihe have not been established. bacteria, and a commercial fungicide, Myco- This paper reports observations on apical statin-Dusting Powder ( 1 ,000.000 unils I development, spore release and early growth (E. R. Squibb & Sons Ltd, Melbourne) proved 1 Atlantic Regional Laboratory. National Research Council of Canada, Halifax, N.Sw, Canada. I?\RCC Mo, 14511.) -" Department of Botany, University of Adelaide. Adelaide, S. Aust. 5000. 50 TIKVAH EDfcLSTHIN & H. B. S. WOMERSLEY Fig- f. A. Apical development, with two -young latctals present, the older one (right) becoming bicus- pid. Trie axis meristem (a.m.) continues growth of the branch, and the lateral meristcms (I.m.) may or may noi develop further into laterals (ADU, A42264). B Cross section of ramulus bearing sporangia (ADU. A42264). u C. Various sporclings I week old, with a rhizoid 3-5 cells long and early .stages of the ceU- mass". O. A sporeling 3 weeks old, with a well developed cell-mass. to be effective when used as a single treatment tinued to develop to plants consisting of clus- of 200 mg of "Mycostutin" per 100 ml of sea ters of branches up to 1 cm across, but from water for 20 hours. Repeated treatments with which it was impossible to clean the epiphytes. 3 streptomycin and Myeoslalin were used and In general, cultures were maintained at 15 C frequent cleaning of the sporclings with glass under a light intensity of 600-800 lux provided needles was carried out. While this damaged by 40 watt white fluorescent lights, and a some sporclings, the majority survived and con- regime of 14 hrs light/ 10 hrs dark. The me- THALLUS AND SPORE DEVELOPMENT OF LOBOSPIRA 151 B m ? 9 jPJ 1 1 c ' 1 1 * 'H HP? Fig. 2. 152 TIKVAH EDELSTEIN & H. B. S. WOMERSLEY B *-* '»2ffin - **% kj «... *...*? I \^>i3P *" ». \ 4 " ' ! %* 100/jm ^1 50/UlTi 4 M 200/jm 200/jm Fig. 3. THALLUS AND SPORE DEVELOPMENT OF LOBOSPIRA 153 200/jm Fig. 4. 154 TIKVAH EDELSTEIN & H. B. S. WOMERSLEY Fig. 5, 1KALLUS AND SPORE DEVfcLOPMFNT OF LOHQSMRA 155 Fijj, 7- A Thallus of Lobospira, showing basal entangled branches with tendrfJ-beaflng laterals, and erect branches with ultimate bicuspid ramuli. Stanley Bench, Kangaroo I„ S. Atwl. {f( h'h(tin & Krujl 126, 25.iv.197Z). B. Surface view of a branch showing tctrasporanam (undivided) and released tcttaspurcs (arrow). (ADU, A42264.) Fi£. 3. A. A multicellular sporeiing, with rhizoid. about 2 weeks old. B. A plant about 3 weeks old, showing development, of a flat thatlus wkh a memtemalic margin. C Planl t? weeks old, with two lobes. O Plant 6 weeks old, showing development of lobes from the margin and especially from the rhizuidal region. Fig. 4. A, Plant Ik weeks old, wilh 3 main lobes. H. Planl 71 weeks old. becoming convolute. C Plant Ml weeks old. with several irregular lobes. D. Plain 174 weeks old, consisting of many irregular branches, each with marginal lobes of various sizes. Fig. 5 4, Plant 24* weeks old: general view of merislemalic apex, with degenerating cells in CCftt/c of each lobe. H, As in A. with two darkly-staining cells (arrows) separating in centre of margin of lobe, C. As in A, with degeneration of cells between two darkly-staining cells. References — Chen. L. C-M- Edei.sthin, T.. & McLachlan, J. Oltmanns, F. (1922). "Morphologic una" Bio- (I9fi9).—Bonncmuisoniii hantifcra JIariot in logic dcr Algen/* Vol. 2. (Jena.) nature and in culture. /. Phyeot. 5, 21 1-220. Stark, R. C. (1971).—The culture collections of 1 algae at Indiana University—additions to the 11 (1858), Phycologia Australica Harvey. W. collection July 1966-Jiily I97K /. Phycol. 7, Vol. I, Hates 1-6(1. 350-362. Kjfu man, r\ R. (1897).— Fhoeophyceae. In A. WoMERS^EV, H. B. S. (1967).—A critical survey Hnglcr and K. Prantl, "Die Naturlichcn Pflan- of the marine algae of southern Australia. 11 Attst. 189-270. zenfamklicn". Th. 1, Abt. 2, pp. 176-297. Phaeophyta. A Dot. 15, VOL. 99, PART 4 30 NOVEMBER, 1975 TRANSACTIONS OF THE ROYAL SOCIETY OF SOUTH AUSTRALIA INCORPORATED CONTENTS Watson, Jeanette E. Hydroids of Bruny Island, Southern Tasmania - - 157 Maconochie, J. R. Shoot and Foliage Production of Five Shrub Species of Acacia and Hakea in a Dry Sclerophyll Forest - - 177 Dolhunty, J. A. Shoreline Shingle Terraces and Prehistoric Fillings of Lake Eyre 183 Late, P. K. Notes on the Relict Palm Livistona mariae F. Muell. in Central Australia 189 Twidale, C. R., and Bourne, Jennifer A. Geomorphological Evolution of Part of the Eastern Mount Lofty Ranges, South Australia - - 197 Haslett, P. G. Woodendinna Dolomite and Wirrapowie Limestone—Two New Lower Cambrian Formations, Flinders Ranges, South Australia 211 Womersley, H. B. S., and Cartledge, Sally A. The Southern Australian Species of Spyridia (Ceramiaceae: Rhodophyta) - 221 Obituary—Graham Frederic Whitten, M.Sc. 235 Annual Report of Council, 1974-75 237 Award of the Sir Joseph Verco Medal -238 Balance Sheet 239 PUBLISHED AND SOLD AT THE SOCIETY'S ROOMS STATE LIBRARY BUILDING NORTH TERRACE, ADELAIDE, SA. 5000 HYDROIDS OF BRUNY ISLAND, SOUTHERN TASMANIA byJeanetteE. Watson* Summary WATSON, Jeanette, E. (1975).-Hydroids of Bruny Island, southern Tasmania. Trans. R. Sot: £ Aust. 99(4), 157-176, 30 November, 1975. A systematic collection of the sublittoral hydroids of Bruny Island, southern Tasmania, using SCUBA, yielded 34 species, including three newly described, three new records for Australian waters and 11 new records for Tasmania. Most Haleciidae, including two new species, are epizoic, occupying sheltered microhabitats. Few species of Sertulariidae are recorded, and Amphisbetia operculata is now rare in a former habitat. The Plumulariidae is represented mainly by small epiphytic forms, and Plumularia angusta, P. crateriformis and P. wilsoni are recorded for the first time from one locality. Two species newly described, Halecium bruniensis and H. luteum, are each closely related to endemic New Zealand species. The occurrence of these, and the first record of Salacia farquhari outside New Zealand waters (where it also occurs south of 43's) suggests active progress of speciation and dispersal across the Tasman Sea. HYDK01DS OF BRUNY ISLAND, SOUTHERN TASMANIA by Jeanette E. Watson* Summary Watson, Jeanette, E. (1975).—Hydroids of Bruny Island, southern Tasmania. Ttan.v. R. Soc. R Atmt. 99(4), 157-176, 30 November, 1975. A systematic collection of the sublittoral hydroids of Bruny Island, southern Tasmania, using SCUBA, yielded 34 species, including three newly described, three new records for Australian waters and 1 1 new records for Tasmania. Most Haleciidae, including two new species, are epizoic, occupying sbellcred micro- habitats. Few species of Sertulariidae are recorded, and Amphisbetia operculata is now rare in a former habitat. The WumuJariidae is represented mainly by small epiphytic forms, and first Ptutmdaria an$u$ta, P r ermeriformis and P. wilso/ti are recorded for the time from one locality. Two species newly described, Halecium bnmienxh and H. luteum, are each closely related to endemic New Zealand species. The occurrence of these, and the first record of C Salacia farquhari outside New Zealand waters (where it also occurs south of 43 S) suggests active progress of speciation and dispersal across the Tasman Sea. Introduction topes would make any significant contribution e to the hydroid fauna of Bruny Island. Bruny Island (43°25'S, l40 20'E) is situated In his revision of the hydroid fauna of Tas- oft the cast coast of Tasmania, 25 km south o( mania, Hodgson (1950) listed 64 species known Hobart. The island is approximately 50 km from the deep and shallow waters of the Tas- long and is separated from Ihe Tasmanian manian coast and Bass Strait. His list includes mainland by the narrow waters of the D'Entre- 22 species from the D'Entrecasteaux Channel castcaux Channel, The indented coastline of and the adjacent Dcrwcnt Estuary, and one Bruny Island provides a range of environ- species from Adventure Bay, mental conditions varying from the sheltered The present survey yielded 34 species (in- bu* swift flowing tidal waters of the D'Entre- cluding two identifiable only to genus) and ensteaux Channel to the rough-water eastern 2 varieties of one species. There are 1 1 new coastline of Adventure Bay and Penguin records for Tasmania, including 3 new records Island, open to the Tasman Sea. for Australia, and 3 species are newly described. Systematic collecting of the sublittoral Only 19 species of Hodgson's list appear in the hydroid fauna was undertaken during two present collection. weeks in February. 1972 t Sampling, using Holotypc and parutye mieroslides of new SCUBA equipment, was carried out. over the species, and other microslides and material, are entire depth range (0-20 m) presented by the lodged in the National Museum of Victoria, rocky sublittoral along the coastline at Satellite Melbourne (referred to as NMV). Island, Great TayJor Bay, Simpsons Bay, the No athecate hydroids were recorded from adjacent DTIntxecasteaux Channel, and at Bruny Island. The Campamilanidae is repre- Fluted Cape, Penguin Island and Adventure sented by 5 species, Lafoeidae by L Haleciidae Bay on the eastern coastline of Bruny Island. 6, Syntheciidae J, Sertulariidae 9, and the Plumulariidae by 12 species, including 2 varie- No collection was made of hydroids from ties of one species. the littoral zone, as these localities comprise either steep rocky cliffs facing the Tasman Sea LIST OF SPECIES or the sandy beaches of the D'Entrecasteaux * Denotes a new record for Tasmania Channel. It is unlikely that cither of these bio- t Denotes a new record for Australia National Museum of Victoria, RussclJ Stteet, Melbourne, Vic 3000, , I5K /EANRTTF Jr WATSON Til ECATA species occupied a microhabital in sheltered Family CAMPANULARHDAK crevices at a depth below turbulence due K> *i Campanularia ambiplica Mulder A: Trcbifcock- surge. "* Campttnuluria pulerath ecu Mulder & Trcbdcocfc. This is the first record of 0. ambiplica from Clytiu -\p. Tasmania. Other localities: Victoria; Champion Orthopyxix caliculata Hincks. Bay, W, Aust. Silieitlaria mwv? Meyen. hamiiy LAFOEIDAf (Jainnumiluria pulcratheca Mulder & Trebil flfbt-lhi ?funu Millard. cock. 1914a: 11, pi. 2, figs 1, 2. Blackburn. Family HALECUDAK 1942; 105. Paracalix pulcratheca (Mulder & Trebilcoek, HoU'cium dt'Jiartulum Coughtrey. 1.91 4ft). Stechow, 1923a: 3 flatccium sp. Satellite I. - Halecium heanu (JohuNlun), Record: (no depth recorded), on HuU'riuM brutriensis n.sp, red alga Defixea. Halecium htteum n.sp, Material: Colonics infertile. Hydrorhiza tubu- Phylactothcca armata Stechow. lar. Stems 0.H3-1.33 mm long. 0.06-0,09 mm Family SYNTHRCJIDAF diam.. perisarc thick, a spherule between stem Synihi'ciutn patalum Busk. and hydrotheca Hydrothecae long and tuhular. Family SERTULAiUIDAE perisarc thickening distally, a distinct dia- E Sutu Halopleris campantda var. campanula (Husk). FIG. 1 Ptumularia filicaulis Kirchenpauer Record: Adventure Bay, 10 m deep on *l«m * Plumttlurta hyalina Bale. of Thccocarptis divaricatus var. typt'ea. * Phttmduria attRusta Stechow. Material: A few infertile stems. Stems of vari- * Plumuhuia crater iformix Stechow. length, 0.70-1,90 irregularly undu- * Plunmlaria wilsoni Bale. able mm, AgUtophvma plumosa Bale. lated, in some places smooth. Hydrothecae Thccocurpt4st xUvaricafus var. typiea (Busk). campanulate, expanding from base Eo margin, * Thccocarpux dharicatus var. hrt&gsi Bale. perisarc fairly thick, and a well defined dia- Haiieornaria tanxiroxtrix (Kirchenpauer) phragm with a thickening of the thecal wall Systematic Section below. Depth to diaphragm 0.30-0.40 mm. A small spherule between hydrolhecae and pedi- Family CAMPANULARIIDAK cel. Margin 0.20-0.32 mm diam., with 12 ambiplica Mulder HYDROIDS O* BRUNY ISLAND, SOUTHERN TASMANIA J5« 1924: 232. Hodgson. 1950: 7, ligs 14-16. panulate, pcris3rc delicate, smooth. Hydrolheca Shepherd & Walson, 1970: 140. asymmetrical, one side convex, the other Ctitnpanularin adiculata Htntfc* 1853: 178, pi. straight or slightly concave, this side always *. fig. 5. inclined to the hydroid host. Margin entire, with tucopclUi calk-utata (Hincfcs) HirohitO. 1969; a thin slightly everted rim. Pedicel of variable tig. ft length, perisarc thick, strongly undulated to Record: Penguin l.„ 15-20 m deep, on a red smooth, widening distolly to pass into base alga and on sponge in a crevice. of hydrotheca below diaphragm. Immature Material: A lew fertile colonies, tlydrothecae hydrothecac truncated with a thin cap-like very shallow ;ind expanding; hydrothecal pedi- operculum, cels spirally annotated, a few wilh smooth Dimensions (mm.); South regions. Hydratuh wkh 24 tentacles. Gwoiheca Hruny 1. Africa smooth with thick pcrisaic, containing mature Hydrolhecit eonophorcs. depth to diaphragm 0,80-0.92 0.74-0.S4 citato, at margin 0.50-0.f.8 0.55-06^ Remarks: The specimens from sponge possess dtam. at diaphragm 0.1 4-0. 18 0,22-0.2* longer pedicels and have a thinner perisarc Pedicel than those epiphytic on algae. length from diaplnagm 20-0.29 0.52-0.72 O. colic alata was not abundant at Bruny minimum diam. 0,1)5- 0.08 0.08-0. 10 Island, occurring in sheltered only a crevice on Remarks: Comparison of the Bruny I. material the rough-water side of the island at a depth wilh paxatype microslides of flebefla fura\ below major turbulence. This accords wilh Millard from False Bay, South Africa (pro- previous findings on habitat preferences of this vided by Dr N. A. H. Millard) shows that species (Shepherd &. Watson 1970; Watson although similar in shape to the South African 1973). specimens, the hydrothecae of the Tasniunian material are generally deeper and Silfciilaria rosea Meycn, 1834: 204, p]. ?5. figs narrower at the 1-11. Millard. 1968: 259. diaphragm. lack the distinct thickening of the lower thecal wall, and have a more pro- Silicuhria bilahiam (Coughirey, 1875 i. kaiph, 1957; 842. nounced eversion of the margin than the South African specimens. bacvpella campatwUlrtu voa Lendenreld. 1883, However, in the absence Dale, tfigg; 751. pi 13, figs 9-J5. of gonophorcs it is difficult to determine the SiltcuUtria campamduriu (von Lcndenfehl, specific status uf hydroids of simple morpho- IWO). Hodgson. figs 1950: 6, 12. !3. logy, hence the presenl specimens arc pro- Record; Fluted Cape (no depth recorded) on visionally assigned to Ih furax, the brown algae Seirococcus axillaris ,,nd Sryto- This is a new record for Australia. thalia dorycarpa. Family HALECIIDAE Material: Luxuriant fertile mate and female Halecfium Uelicatufum Coughtrcv, colonies on the algae. Developing gonophnres 1#7fia; 299; IS76b: presenl. 26. pi. 3, figs 4, 5. Ralph, 1958: 334, %s 1 1, 12 (synonymy >. Remarks: The present material conforms to Hateciitm flexile Allman. 1883: U, p|. 5, fit. 2- descriptions given by Ralph (1956. 1957) for Hodfiwta, 1950: 16. figs 25-27. Silicularia bilabiata forma subtropica (demon- Records: Adventure Bay, Penguin I., Satellite strated by Millard (1968) to be a synonym of I., on the kelp Macrocystix pynfera. 2-7 in S. rosea Meyen), a form occuring only north deep; Fluted Cape, 10 m deep, on Tbecorarpus of the Subtropical Convergence, and typical of divarictUiis var. typica (Busk). the southern Australian coastline, including Tasmania. Material: Luxuriant fertile colonies. Stems to I cm long, simple und branched. Many hydro, Family LAFOEIDAE phores with marginal replications; secondary HcbcUa furax ? Millard. 1957: 200, % 8; hydropborcs arising trom the pedicel ot 1964; 10, fi& 2B-D. Millard & Bouillon, primary hydrophorcs. Colonies dioecious with 1973: 59. mature gonophores ansing from proximal parb. FIG. 2 of stem and on hydmrhiza. Distal parts of Record: Penguin I., on stem of ThectKarpus blastostyie cap-shaped |p both sexes. Colour, divarieatuit var. typita. depth 20 m. irophosomc yellow, gonophore bright orange. Material: One small colony. Hydrorh'tza a Remarks: Hodgson (1950) described ami Uroscly winding tube. Hydrothecae large, cam- figured H. flexile Allman (= H. deficatulum) 160 TEANETTE B. WATSON Fig, 1. Chtia sp, Hydrolheca. Tig 2, Hebella ?jurax Millard. Hydrothcca epizoic on J hecocarpus elivaricaius. Figs 3.4. Halecium sp. Fig. 3.—Whole colony. Fig. 4—Distal part of colony enlarged. Figs 5,6. Halecium beanii (Johnston). Fig. 5.—Whole colony. Fig. 6.—Part of stem showing hydro- phores. from Eaglehawk Neck, but did not record the thick, with one proximal annulation and a distal substrate. apophysis giving rise to the succeeding inlcr- The colonics of H, delicttmlum in the pre- node. Hydrophore fairly deep, expanding to wall sent collection, while very abundant on Macro- margin, slightly asymmetrical, adcaulinc wall. Depth to cystis and other brown algae, were, however, more expanding than abcauline 0,06-0.07 to base of strictly epizoic, always growing on the surface diaphragm. mm, depth 0.12-0.14 of the crustose bryozoan Membrhwpora mem- hydranth, 0.05-0.06 mm. Margin rolled rim. brinacea, a common epiphyte on the aging mm diam., with a distinct outwardly fronds of Macrocystis. Diaphragm concave, approximately 0.01 mm below line of attachment of hydranth, usually Halecium sp. a strong thickening of the inner wall immedi- FIGS 3, 4 ately below diaphragm, best seen in older Record: Penguin I., on a red alga in sheltered hydrophorcs, often absent in younger terminal water, no depth recorded. hydrophores. Punctae not visible; if present, hydrophorcs Material; One infertile stem. Stem 3 mm high, obscured by hydranths. Pedicels of pedicels usually lightly fascicled at the base, branching irregu- of variable length; shorter pedicels smooth. Hydro- larly sympodial. Stem internodes of variable annulatcd, longer 1-3 regenerated pedi- length, 0.30-0,40 mm, narrow, perisarc fairly phores regenerated times, 6 5 HVDROIDS OF BRUNY ISLAND, SOUTHERN TASMANIA 161 eels often with a deep proximal constriction Millard (1957) and Ralph (195S), the speci- Hydramh short and stubby, wfrih approximately mens, although infertile, are assigned to ibis 14 tentacles. species. Retmrkx; The single specimen from Hnmy 1. An unusual microhabilat is the epizore resembles H. tewllum Hinks, 1861, and Hale- growth on crustose sponge on the underside of dum sp. recorded from Pearson 1. (Watson the thick, plaie-like thallus of Sonderophycus. 1973. p. 167). However, H. tenellum is muno- H. heanii is a cosmopolitan species, not siphonic tuid the dimensions given by Millard previously recorded from Australian waters. It (1957, p. 193) and Ralph (1958, p.~34(l) for is rare at Bruny L H. tenelhtm arc creator than iho.se of the pre- Hakiium bruniemis a. sp. sent material. The specimen from Pearson J-, FJGS 7-1 while similar >n habit, is monosiphonk. the margin of the hydrophore is more everted, and Type material and records: Holotype, NMV. the overall dimensions are smaller, Without C2494-microslidc; G2495-prcserve"d material, adequate fertile remainder of holotype cofony; material it is not possible to Penguin f. t 20 m make a decision on the specific status or rela- deep, on sponge and bryozoa in crevice. tionships of the specimen from Bruny I. Description from hohtype; Etea stem 3 cm high, growth habit arborescent, in one plane, Ifahciiim bcatiifi (Johnston, 1 838). Millard, stems sparingly fascicled at base, woody, the 1957' IKK; 1966; 464; ]96S; 256, fa, polysiphonic tubes running up stem, forming 9A-F. Ralph. 1958: 3.12, fig. 10a, b, c-k. the branches Branches monosiphonic distally,, FIGS 5, 6 with markedly sympodia! growth. Stem inter- nodes Records: Adventure Bay (no depth recorded) of ultimate branches of fairly constant length, 0.48-0.64 on encrusting sponge on underside of the red mm, narrower proximally, widening distatly 0-0.] alga Sotiilerofthycux auitrafiv: Penguin 1„ 15 m to 0.1 2 mm at node. Nodes deep, on sponge in crevice. oblique, well defined, sloping alternately left and right, often 1 or 2 annulalions above Material: Busby infertile colonics to 15 mm node. Pedicef of hydrophore given off distally long. Proximal parts of the colonies fascicled, from a well defined apophysis 0.04-0,14 mm becoming monosiphonic distally,. usually where long, at the same level as the node. Pedicel regrowth has occurred from broken poly- tubular. 0.13-0.20 mm long (node to dia- siphonic tubes. Branching irregular, occasion- phragm), perisarc of younger parts smooth, in ally tendrils given off distal ends of branches older regenerated parts heavily internally through the orifice of the terminul hydrophore. ridged, a deep fold just ahnvc apophysis giving Stem internodes of variable length, 0.28-0.52 pedicel an offset appearance. Hydrophore fairly mm, narrowest at node, 008-0.12 mm, widen- deep, 0.06-O.Ofc mm margin to diaphragm, ing distally to 0. 13-0.23 mm to accommodate slightly expanding with an everted margin, hydrophore. Nodes distinct, with a slightly 0.15-0 18 mm in diam,, and distinct rim. Dia- oblique slope alternately right and left, occa- phragm well defined, thin, with a ring of sionally straight, Hspirophores alternate, shal- punctac just above and a pseudodiaphragm low, saucer-shaped, adnale to internode, diani. below, frequently only marked by a thickening at margin 0.12-0.14 mm. Diaphragm distinct, of the adenine wall. Secondary and tertiary 0.02-004 below margin, tilted towards node, regenerations of the pedicel from the orifice of marked by a thickening of perisarc, p ring the preceding hydrophore common, each suc- of punctac (frequently not well seen) above! ceeding pedicel uutally shorter than the last; in Secondary hyttrophores given off on a short some instances regeneration is reduced to a pedicel from diaphragm of primary hydro- mere replication of the hydrophore. Secondary phore. adcauJine wall convex, abcauline wall branching of pedicels rare. Hydranth elongated, straight or very slightly bulged. Body of slender, with 10-16 long filiform tentacles. hydranth delicate, wilh approximately 14—1 Female gonorheca very large, flattened, lenti- long filiform tentacles borne on a long cular, but somewhat variable in shape, usually peduncle, a deep constriction between slightly longer than wide: greatest width G,9&- peduncle and hypostome, 1.33 mm; length (excluding pedicel) 1.20-1,43 Rrmarkx; Since the Bruny I. specimens con- mm, tapering proximally into a short pedicel form lo descriptions, and fall well wilhtn the arising at base of hydrophore, usually ol junc- range of dimensions given for U, beonii by tion with main stem. Perisarc very delicate, 162 JEANETTE E. WATSON Figs 7-15. — HYDROTDS OF BRUNY ISLAND. SOUTHERN TASMANIA m distal extremity puckered, a small circular Branching more or less in one plane, the ulti- Otificc sealed by an operculum situated in the mate branches in any one part of the colony distal third of Ihc abcauline wall Male gono- all directed anteriorly: in other parls they may theca small, sausage-shaped, 0,25 mm long, face in other directions. Stein tnrentotlex nf 0.10 mm wide, arising from a short pedicel at variable length, 0.40-0.65 mm, expanding base of hydrophore. Buth sexes arising only dismally, with 1, occasionally 2, extra nodes. on younger, monosiphonic pans nf colony, the Nodes distinct, oblique, parallel in each inter- males more distal than females Colour, $tems uode. sloping alternately left and right in ad- straw coloured, female gonophores pink. jaccat uitcrnodcs; perisarc indistinctly internally Remarks: Hatecium bninienM\- is closely allied ridged. Width at proximal node, measured to //. lentiatlure Trcbilcock, 1928. in sym- parallel to node, 0.12-0,14 mm. Hydrophore podial habit, shape of female gonotheca and seated on distal third of distal segment of inter- hydrorheca However, //. knilculure as known node, well below node, Hydrophore sessile, at present (Ralph 1958, p. 331) is n mono- very tlat and shallow, wails thin and .UTOngly siphonic species with smaller stems I less than constricted just above diaphragm, abcauline I cm high ) , a considerably smaller female wall more concave than adcauline. Margin or. gonotheca, and a male gonotheca of somewhat cular, 0.14—0.16 mm in dianv. with outrolled different shape. rim. Depth from margin to puncjae, 0,015- The type material of H. hnmiensix shows 0.02 mm; depth from margin to diaphragm. growth stages of the female gonophorc from 0.025-O.O35 mm. Diaphragm veiy strong, with earliest development to maturity (Figs 12-15). a distinct riug of punctae above, and a wedge- Development of J he uonophorc begins with shaped thickening oi the perisarc ol the inter- formation of a hook-shaped hlasiostyltr sur- node below. Below the wedge the wall of the rounding a central body (Fig. 12). Further internode thins into a large circular fenestra- growth and diilcrcntation into 6-41 large bodies, tion from which the apophysis of a branch posstbly larvae (but material insufficiently well may arise, Hydnatth large with an annular preserved for positive identification), then oc- hyposlomc and 25-28 tentacles. Colour, bright curs, filling gonotheca (Figs 13. 14), A small yellow, Gonotheca. absent. circular aperture with slightly thickened rim Remark*: then develops in the distal third of the ah- Ntdectum htieum is superficially similar caul'mc wall through which the reproductive to //. corrugatissimum Trebilcock. 1V2S. (rom Zealand, il products escape (Fig. 15). The orifice of ihe New as has strongly ridged intcrnodes and now empty gonotheca then becomes rcsealed shallow hydrophore* character- istic of this hv a very Ihrn operculum. species. In the latter species, bow- ever, the ridges Only one group of colonies of H. bnudensh of the stem are merely strung wat lound These were cpuoic on sponge and annular constrictions, not definite oblique nodes as in fi. encrusting bryozoa in a sheltered crevice. furcunh Furehe/mure, Ihe hydm- phores, while shallow, axe somewhat deeper I lutecium hiteum sp. n. than those of the new species. Also, H. eorru- FIGS 16-lS s}athsimum is a monosiphonic spectes, while type material mid Records: Holotype. NMV. H. tutemn has a strongly fascicled hahit. G24%-microslide Figs 7-15. Hatecium brtwiensis n.sp. He. 7.—Holotype colony, natural size. Fig *.—Distal part of colony, Shwtofc fasciculation oi stem and empty female fiOOOthevae. Figs 9, 10. Hydro- phurc-v enlarged, showing offset pedicel and teplications of the hydrophore Fig. II.—Male gonotbec*. Figs 12 15.— Development ot the female gonopbore. Fig. 12.—Parly stage development of showing booX-shaped bbstosrvie with developing central mass Fig. 13. tatei stage ot development. Fig. 14.— Mature gonophore. Fig. 15.—Gonotheca after dis- charge of reproductive products. Note residual mass remaining at site of the circular orifice- through which contents have been discharged. 164 JEANETTE E. WATSON 1 cm Kgs 16-18. Halevium luteum n.sp. Fig. 16,—Paralype colony, twice natural size. Fig. 17.—Distal part of branch. Fig. 18.-—Stem internodes with hydrophores enlarged. Fig. 19. Phylactotheca armata Stechow. Gonotheca with developing male gonophorc. bulky hydranth. The extreme thinning of ihc holdfasts of Phyihsporn comoca, 10-22 m wall of the internode in the fenestration below deep. the hydrophore must produce a serious struc- Material: Luxuriant colonies, some fertile. tural weakness of the hydrocaulus. This is. St^ms of variable length, 4-15 mm, usually however, offset by the support given to the branched. Colonies dioe- hydrophore by the very strong wedge of peri- simple, occasionally thickly on hydrorhiza sarc extending across the base of the hydro- cious, gonophores borne gonothecae large, flatly ovate, phore from the stem. at base of stems, shape, widest at The group of colonies were both epizoic and both sexes of same size and or perisarc thick, slightly undulat- epilithic, growing down from the roof of a middle lop, ing, on very short pedicel, length 1.14— cavern in sheltered conditions. borne a 1.56 mm (excluding pedicel), maximum width Phylactotheca armata Stechow, 1924:59; 1925: 0.90-1.17 mm Gonophores mature, of creamy 204, fig. C. Blackburn, 1942: 106. Hodg- white colour, almost filling gonothecal cavity, son, 1950: 17, fig. 31, Watson, 1973: 166. male surrounded by a thin blastastyle, female Ophiodixw fragHfr Blackburn, 1937: 365, fig. I. packed with mature ova. Hydnmths with a FIG 19 single row of large lenticular nematocysts Records: Penguin 1. and Adventure Bay: epi- (probably stenoteles) alternate with the ten- zoic on solitary ascidians, bryozoa and sponge; tacles surrounding the hypostome. These also epiphytic on crustose coralline algae and on occur in the nematocyst batteries in the capi- HVDROTDS OF BRLNV ISLAND, SOUTHERN TASMANIA 165 tulum of the retracted daciytozooids, and are to distinguish between Synthecium etegans f. scattered throughout the hydrocaulus in some suhventrieosum and S. patulum on character) stems No discharged neniatocysts were seen. of the hydrothecae alone. The present material is Remarks; Blackburn ( 1937) described the thus provisionally assigned to S. pawtum. gonophore of "Ophiodissa fragilis" (*= P ar- Further work may eventually prove that the mata) as being "subspherical, arising at the two are conspecific. junction of stem and peduncles, as well as stem No morphological differences could be and hydrorhiza". Blackburn's type mieroslide detected between stems growing in the environ- of "O. fragilLv" t'NMV collection) shows three mental extremes of exposure to current (Simp- extremely delicate structures which appear to sons Bay), sheltered situations under ledges (Satellite I.) be either directly attached to, or enveloping or exposure to surge I Penguin J.>. the hydrocaulus. Although two of these contain Family SERTLLARflDAE a central mass which could possibly be a developing gonophore. they resemble neither Salaefo farquhari (Bale, 1924). Ralph. 196!a: in shape or structure 1he immature and mature 760. fig. 7, gonophores of P. armaia as seen in the present FIGS 20-22 material. They dp, in fact, closely resemble Thuiartu farquhari Bale. 192*1: 2^4, fig, 10 &gg capsules of certain minute gastropods. TreWlcock, 1928: 19, pj, 8, fig. 4. Records: I., P, artnata shows a wide range of substrate. Penguin 15 m deep; Satellite I., The epiphytic colonies, particularly those from 3 m deep, on sponge under ledge. the rough-water sites among Phylfaspora hold- Materia}: Two infertile colonies. Stems mono- fasts, were usually short and robust, with siphonic, to 18 mm long, simple and branched, heavily ridged cauline perisarc. Eptzoic the simple stems shortest. Stem Jnternodes colonies, particularly those from more shel- tapering proximally and distally, with one pair tered situations under ledges in deeper water. of opposite hydrothecae adnate in front, widely separated were lax, branched, with a more delicate peri- behind. Branching regularly alternate sarc and fewer intranodal ridges. All stems, from aii apophysis of the inter node 0.25-0.4 however, show a tendency towards thickening mm long, usually 2 pairs of hydrothecae oE the perisarc and increase in cauline ridges between branches. Branches given off at an with age. upward angle with a distinct proximal genieu- lation This is the most abundant occurrence of P. at the apophvsis and a V-shaped distal armaia so far recorded In Australian waters, joint. Some secondary branching, but where developed, and demonstrates a greater variability in stem these branches carry few hydro- characteristics than formerly known. thecae. Family SYNTHECHDAE Remarks: The specimens compare well with microslides of Thuiarfo farquhari Bale (NMV Synlhectum patulum (Busk, 1X52). Hodgson, collection) and with the redescription of S, 1950: IS, figs 32, 32. farquhari given by Ralph f 1961a) The present Sertutasia paiula Busk, 1852.* 390. material does, however, exhibit certain dif- Records: Satellite I., 14 m deep, tinder ledge; ferences from the species as described from Simpsons Bay, 2 1 m deep, on scallop Equl- New Zealand. These are the greater length and chlamys hifrvHSi Penguin l M 20 m deep, on the proximal geniculation of first sponge. the branch rnternode. as well as the tendency towards Materiaf: A few infertile colonies. Stems to thickening and loss of the stem lnternodes 25 mm long, some stems immature Proximal in older parts of the colonies, features recog- internodes of sterns short, with I pair of oppo- nized by Ralph as being more characteristic of site hydrothecae, followed by an Joternode Solatia bicalycuhi (Coughtrey, 1R76a) than 5. with 2 pairs of opposite hydrothecae and farquhari. Since the present material agrees in ilislal hydrociadia; distal lnternodes with either most respects with the latter species, particu- I pair Or 2 pairs of hydrothecae. Hydrothecae larly in the size of the colonics., it is assigned on older stems with thick perisarc and replica- to S, farquhari. tions, of the margin. Colour, purple and white. This is the first record of S, farquhari for Remarks: The present material conforms to Australia, and the first record of the species descriptions of Synthec'mm patulum given by outside New Zealand watery where it occurs Bale (1914a, p. 5) and Hodgson (1950) How- only south of 43°S, the same latitude z% ever, it is very difficult I Watsou 1973. p. 167) Brnny I. : i«6 jFANrTTT T WATSON SirrcoUuM-ii elongata (Lamouroux, 18it>). zoically on dead shell and other material in Ralph, 196 la: 7*2. fig. 4. Waison, 1973: the lyEntrecasteaux Channel in situations of 170. .good current How, those of intermediate stem Set tula flu clotizuta Lamouroux, 1816: 189. pf length being from cryptic habitats beneath 5. Bale 1884: 75, pi, "short-stemmed" ocean form frequently asso- internode" form of the rtruny I materia! cor- ciated with red algae. responds very closely with Sertalarella simplex (Hutton, 1873) described from Pearson I- Scritilarolla robusta Coughtrcy, IS76h: 300, fig. (Watson 1973). A specific distinction based on undula- 2. Hodgson, 1950: 33, fig. 58. Ralph. thickness of perisarc and faintness of 1961a: 824, fig. 22a-d. Watson, 1973: tions of the thecal wall as defined by Ralph arti- 171, fig. 21. ( 1961a, p. 820) seems to be somewhat ficial, Thus, if further material with smooth F[GS 23, 24 hydrothecae and thin perisarc is found, 3. Records-: Satellite I.. 3 m deep, on sponge- wbitsta must be referred lo the synonymy of under ledges, and 6 m deep on the red alga S. simplex. Sonderophyctts aastralis; D'Enlrecasteaux Channel, 1 1 m deep on dead seawhip Prim- Synipkctoscyphus pyjjmneus (Bale, 18JU). noeita australasiae, and on old scallop shells. Walson 1973: 176. Equichlumys hijrons; Adventure Bay, 5 m deep Sertuhrella pv.unuwa Bale. IS81: 25. pi. 12, fitt- on sponge under ledges: Fluted Oupe (no depth Sfj 1KS4: H>8, pi, 3, fig. 8. pi. 19, fig. 19. 64. recorded) on stem of a brown alga. Hodgson. 1950: 36. figs 63. in Material: Stems 3-4- mm long, the longer stems Eectjrd; Penguin I.. 16 m deep on bryo/oa llexuoiJs. with long inlernodes, occasionally sheltered situations. with short branched Shorter stems robust, A'fiirrnV//* Sparse infertile colonies. Stems to occurring iniefnodex. each stem type in 7 mm long, simple, or with one branch. A "long separate colonies. Hydrothccae of the delicate 3-flapped operculum visible in most iniernodc" form large with smooth, very faintly hydrothecae. undulated walls* hyilrothecac of the "short Remarks: The line of small dots rri the thecal internode" form distinctly smaller (see dimen- wall diagnostic of 5. pyxmueas is obscured by sions), with thinner perisarc. heavily ridged the hydranths. However, Ihe material corres- wirh 3-4 annulations and a Strong submarginal ponds closely with specimens of S, pygmnens constriction of the thecal neck on the abcauline in the Bale collection (NMV) and for this side, All material infertile except for one stem reason is Assigned to this species. of the '"short intermule" form. Dimensions (mm) tons short Sertularia acuta (Stechow, 1921). Millard, 195*: internode internode 192, fig. 8. Shepherd A Watson. 1970: 140. form form Tridentata acuta Stcchow, 1921: 231. Length, abcauline wall 0.55 fl.fiO 0.33-0.45 Sertularia favulo.w Bale, 1884: 91. pi. 4. fiji figs 1915: 272. Length, free ndcaulinc wait 0.35-0 4? 0,28-0.38 5, 6: 1913: 121 pi. 12. 7, 8; Hodgson. 1950: 25. figs 43, 44. Remarks: Specimens of S. robusta from Bruny Record: Adventure Ray, 10-22 m deep, on red I. show a complete range of variability between algae. the extremes of the long and short internode forms, and large and small hydrothecac. Some Material: Luxuriant fertile colonics. Sietm to correlation appears to exist between stem type, 7 mm long. Cottothecae with 4—6 strong annu- environmental conditions HYDK01DS OF BKUNY ISLAND. SOUTHERN TASMANIA H,T Remarks: Hodgson',*; infertile specimens came of "pumiloides" type, and a distal rcgrowth of from storm-drifted Microcystis. S. acuta was thiee hydrothecae of unmistakably "intermedia" r\o\ associated with this kelp at Bruny 1 type following stem breakage. The basal pair of "intermedia" hydrothecae bave well Serfularin macrocarpa Baie. 1&S4: ftO. pi. 5, developed tubular nematothecae. Dimensions fig. 2. pi. 19. fi&. 11: !9I4a: 14; 1915; of the proximal and distal groups of hydro- 277. Mulder & Trebilcock, 191 4ht 42. thecae on this stem are given for comparison: Hodgson, 1950; 27, rig. 47. Shepherd & Dimensions (mm) j Watson, 1970: 140. Watson, 1973; 177. ptter* pumiioides media Record: Penguin J., sheltered side, 10-22 rn type type deep, among algal holdfasts. Internode lenpih 0.33-0. *5 30-0.34 S'iawriai Rare colonies, comprising few a widrh at node 0.06 0.04 stems each Stems to R cm long, infertile- HydroJheca length 0,20 0.21 Remarks: The internal suhmarginaJ loolh is not width across ba^e 0.24 0.25 0.18-0.20 as well developed in these specimens as in (hose widrh across margin 0.43-0-52 0.32 0.37 ffom the Australian mainland. The finding of iwo varieties of A. minima on This dark brown species wjlh distinctive one stem lends support to evidence (Watson white-tipped hydrocladia has previously been 1973) that these varieties are in reality eco« recorded among the hold I si fauna red algae a of tnorphs, the development of which may be at Pearson I. (Watson 19731. dependent upon the type of substrate available, This is Ihe first record of S- macrocarpa from or environmental conditions prevailing during Tasmanian waters. growth. Hodgson (19501 did not differentiate be- Amphisbctia minima var. intermedia Bale. tween the varieties of A. minima in his collec- 1915. Watson, 1973; 179, fig 29 tions; however, according lo his measurements, FIG. 25 and the ahsence of nematothecae. as well as Sertutaria min Thompson 1 : 1 pi ima T 879 04, the material having come from Maerocystis 17. fig. 3. Bale, 1881: 21, 45, pi. 12 fig, 2; and "drift" (brown?) algae, the material may 1884, 89, pi. 4. V pi. fits 12, fifiS t 10, ft, 13; 1 be ascribed to the "var. '. Mulder & Trebilcock, 1 9 14b: 39, Hodgsnn, pumiioides 1950; 23, figs 41, 42. Amphisbctta operculafa I Linnaeus. T 75K). Records: Adventure Bay, 4-6 m deep, on iod Ralph, 1961a: 775. fig. 8. alga Rhodymenia and on sponge under ledges; Scrtidaria opercuhta Linnaeus, 1758: 808. Bale, Penguin 1., 16 deep, on red algae in m and 1884: 67. pi. *. fig. l, pi. 19. tm. 3. Hodgson, in crevices rough water; Satellite 1 , 1 m deep> 1950; 22, figs 36. 37. on Laurencia. Record: Simpsons Bay. 11 m deep on stem of Material: Luxuriant fertile colonies, mainly on dead seawhtp Primimella australiastac. red algae, some on sponge. Stems to 5 mm Material: One small infertile colony of a few long, internode length 0.30-0,38 mm; diam. at short stems. node 0.03-0.06 mnu HydrOlheca 0.19-0.28 mm Remarks. Although Hodgson remarks that long. Tubular nematothecae present in the base "this species is very abundant in the D'EoUe- of proximal intrathecal chamber. casteaux Channel, being constantly taken in Ihe Remarks: Although the hydrothecae are larger form of large tangled masses in scallop v than those recognized as var. intermedia (Wal- dredges . only one colony was found in the son 1973, p, 181), the present material is present survey, This apparent rarity may be referable to this variety on the ba\ts of the due to seasonal growth (Hodgson's material shape of the hydrothecae and the presence of was collected in August) or to permanent the characteristic tubular nematothecae. How- changes in the environmental equilibrium cvei, several of the stems have rather robust caused by scallop dredging. The substrate of hydrothecae and the wedge of perisarc hetween old shells preferred by A, operculata (pers. hydrocaulus and hydntfheca considered typical observ.) ts no longer available as the seafloei of (he var. pumiloides. Furthermore, one stern of the estuary has now heen invaded by enor- (Fig. 26) from sponge., shows distinct transi- mous numbers of the New Zealand gastropod tinnal features between the pumiloidcK and Maoricoiptts roseus. Since no colonies of /I, intermedia types. The stem of this specimen operculata were found associated with M hits four hroad* robust, proximal hydrothecae roseus> it may be concluded that the smooth 168 JEANETTE E. WATSON Figs 20-27. HVDROTDS OF BRUNY ISLAND, SOUTHERN TASMAMIA 169 shell of this jtaunopod is unsuitable for settle- Width across margin between teeth 0.07-0,09 ment of larvae of A. opvrculata, mm; width across paired hvdrothecae (outer- most teeth) 0.52-0.69 mm. A Plicate internal Amphisbetia aria o. sp. sheath often present within margin Gotto* FIGS 26, 27 tftetwe large, obovate, 1.15-1.33 mm long (in- Type Material and Records: Holotype, NMV, cluding pedicel) expanding from base to sum- G2499-micros!ide; G2500-preserved material, mil; max width 0.75-0.83 mm. tapering evenly remainder of holotype colony: Adventure Bay: into a narrow pedicel arising from the inf*a- paratypes. G2501, G2502-rjrncrosltdes; Satellite ihecal chamber of hydrotheca on lower stem. I., all colonies on the brown alga Carpoglossum Apenurc circular, 0.35 mm in diam., centrally conflttens, 3 deep, m situated at distal end, with a slightly raised Description from holotype and paftttypex; collar, a ring of minute denticles within, and a RydrorhhM tubular, 0.09 mm diam., reticulate, Oat operculum. Gonorhecae identical in both very loosely wound on algal surface. Stems saxes, only one borne on each stem: male and arising at slolonic junctions, simple, un- female gonophnre on same colony. Female brartched, to long, 5 mm beginning with I or 2 gonophorc narrowly elliptical, not filling gono- twists, then a V-shaped proximal pint, followed thecal cavity, with 16-20 eggs; male gonopho:e by first thecate intemode, PcnsaTC of stem and of same shape and size as female, spermato- hydrotbecae Ihick. and very brittle. Stem inter- geaic mass surrounded by a thin blastostyle. nodcx O.3&-0.46 mm long, with one pair of Remarks: Amphisbetia avia is closely related hydrotbecae, nodes slender. 0.06^0.03 mm lo Ihe Amphisbetia minima group in size, wide, distal node collaT-shapcd. proximal node colour, habit, and preference for algal sub- V-shapeJ and sockettcd into (he collar of pre- strate, and is not easily distinguished from the ceding infernodc; if node absent, it is replaced varieties of A, minima in the field. However. by a nanowirtjr of the intcrnodc. Hydro/necac the deep retroflexion of rhc abcauline wall, the opposite, on distal half of intcrnodc, tubular. horizontally directed distal part of the hydro- narrowing to margin, actuate for one third of theca, and the Jong marginal teeth immediately length, proximal adcauline wall more or less distinguish A. avia from A. minima. parallel to axis of intemode, in contact or A. avia was associated only with one species slightly separated, base of hydrotheca hori- of alga. Curpogfossum confluens, growing in zontal; a very deep notch, and occasionally a shallow water. short oblique intrathecal fold about one third distance up abcauline wall from ba^e of hydro- FamiJy PLUMULARMIME theca, and a corresponding, hut not so deep Halicornopsis elegant tLamarck, 1816). Bale, inflexion (sometimes missing altogether) of the 1914a: 56: 7915; 303; Briggs. 1914: 296; adcauline wall, opposite, but more distally 1915: 309. Blackburn. 1942: 107. Hodg- situated, just behind margin. Width of inicr- son, 1950: 48, fig, 79. Watson, 1973. 195 node just below hydrotheca, 0.24-0.29 mm; Pltwwtaria at^ans Lamarck, f816: 129. length of fixed adcauline thecal wall (measured Record Adventure Bay. 15 m deep, epiliibic diagonally) 0. 15-0.1 y mm, length of free un vertical face. adcauline wall (to end of tooth) 0.12-0. J 8 mm; Material: One fertile colony. SterftX short. :o lenoih of abcauline wall (measured diagonally 7 cm. Gonothecae borne prolifically on main from to end bade of tooth) 0.26-0.31 mm. stems and occasionally at base of branches on Margin horizontal, facing upwards, with two the apophysis of the hydrocfadium. Gona* long, shaip, laterally placed teeth with a deep shecae mature, irregularly ovate, flattened dis- hortontal cmbayment between, connected lo tally or slightly flattened on one side, aperture the internode by a thick wedge of perisatc. closed by a thin membrane. Only female gono- Figs 20-22. iWwm farauhari (Bale), Fig. 20.—Part of stem. Fip. 21.—Stem mternodes, enlarged. Fig. 22.—Single hydrotheca. anterior view. Figs 23.24. St'rwfan-Ua robusta Coughlrcy. Fip. 23.—"Long internoUc" farm with large hvdroThecae and -smooth thecal walls. Fig. 24—"Short intemode" form, showine smaller hydrolhecae with undulated wajls. Fig- Amphisbetia "25. minima var. intermedia Bale. Aberrant stem with both ^pumiio'nhs" (proxt- 5«J P*rt of ylem) and "inter/nvtifa' (dista?) hvdrothecac. Hp 26.27. Amphisbetia avia n.sp. Holotype Fig. 26.—Stem with femnle gonophore. Fig. 27 —Stem mternodes, enlarged 170 JEANKITE L. WATSON phorcs present, spherical, half filling gnno- shows that the Bruny I. specimens correspond theeal cavity, packed with niaiure ova sur- closely to a micros! ide of P. mirabilis from rounded by a ihin granular blastostyle. Buss Strait. Gonotbccac of P. producta col- Remarks: Only one small colony of H. clematis lected at Port Jackson, in 1886, arc smaller WAS found growing in a relatively exposed Pvcnotheca mirabilis (Allrnan, 1883) var, mira- Plumuluria hlicaulK Kirchenpaner. 1876: 28. bilfe Stechow, 1925; 241. Ralph, 1961b: p\. 5, fig. 6. Bale, I8S4: 134, pi. II. figs 50. tig. 7a, b. 6, 7. pi. 19, figs 41, 42. Icloup, 1934; 41 1958: FIG, 28 Hodgson. 1950; 42. fig. 72. Millard. 209,^ fig. 13D E. Shepherd & Watson. Pvenotlteca mirabilis (Allrnan, 1883). Hodgson, f 1950; 50, fiss 81. 82. 1970: 140 Diplocluilas mintbilis Allrnan, 1883: 49, pt. 8, Record: Adventure Bay, 10-22 m deep, on red fig* 4_7. algae. Kirr}vni>t m*tla mirabilis (All man. 1883). Bale, 1894: 109; pi. 6. flgfi 4-7. Briggs, 1915; 308, Material: Luvuriam infertile colonies. Usdro- Blackburn. 1942: 106. rhiza pegged, forming a reticular network on Records: Satellite I., 9 m deep; Adventure Bay, the algal frond, Stems short, simple and pin* 10-22 m deep, on red algae naLe on the one colony. Simple stems to 2 mm, Material: Fertile stems to 5 cm high. Stems pinnate stems io 4 mm high. monosiphonic, arising singly from bydrorhiza. Remarks: P. filicaulh is a common epiphyte on lower stems devoid of hydrocladia. Gonothecae several species of red algae 1 Shepherd & large, adnate to lower 9tem. hydrorhiza, of Watson 1970). jlgac. Perisarc very thick, strongly ridged with Plimtularia hyalina Bale. 1SK1: 41. pi. 15, fig, 9: up to 9 ridges, more prominent on ahcauline 1884: 14T.pl. 12. figs 4, 5 Ralph, 1961 h: side. Colour of gonolheca, deep red brown, 41 lionophore.s, female. FIG. 29 Remarks: It is difficult to distinguish between Record: Fluted Cnpe, 16 m deep, on a red alga infertile stems of P. mirabilis and P. products fertile colonies. Bale. 1KR1. Hxamrnation of fertile material of Material: Abundant, sparingly hydrocladia! both species in the Bale collection (NMV). Seems io 3 mm long, stems and HYDROIDS OF BRUNY ISLAND. SOUTHERN I ASMANIA 171 Fig_ Pycrwthecu mirahUts vjir, 28. mirabilis (Allman). Gonoiheca with heavily ridieed perisarc and female gonophore. Fig. 29, Plumularia hyalina Bale Stem with ripe female gonophore. Figs 30,31. Plumularia angu.sta Stechow. Fig. 30,—Part of hydrociadium. Fig. 31.—Gonoiheca Fig. 32. Piumuhma crateriformis Siechow Part of hydrociadium. Figs 33, 34. Ptumularia wilsoni Bale. Fig. 33.^Hart of hydrociadium. Fig. 34.—Gonoiheca showing downwardly directed growth habit. internodcs strongly ridged, hydrocladia with Some stems of the material from Bruny I. one terminal hydrotheca. Gonothecae large, have a well developed "stolonic plate" identical top-shaped, greatest widlh distally, 0.7 mm with that described for Plurmdaria epihtacteo- wide. 0.9 long; gonophores mm mature, female hsa Watson, from Pearson I. only, completely filling gonothecal cavity. This is a new record for Tasmania. Other Colour, stems white to yellow; gonophores localities: Vic.; S. Aust.: New Zealand. bright yellow. Plumularia angusta Stechow, 1923b: 226. Remarks: This is the first record of mature Blackburn, 1942: 108. P. hyalina in Australian waters. Ralph (1961b) FIGS 31 described much longer (1.3-1/7 mm) and 30, Plumularia setaceoides vars slightly a, b, d, Mulder & narrow (0.5-0.65 mm) gonothecae for Trebilcoek, 1910; 117, pi. 2* fig. 9, pi. 3, figs her mature specimens from New Zealand. 3, 6. Judging by this considerable difference in size Records: Penguin I., 15 m deep, on Macro- of gonothecae, it seems that an as yet undocu- cystis holdfast and sponge; Adventure Bay, on mented range of geographic variants of P, Macrocystis holdfasts and the brown alga hyalina may exist. \4yriodesma quercijolium. IT JP.ANETTE U. WATSON Malarial: Abundant fertile stems to I cm long. very short pedicel from proximal part of stem, Hydrothecae pitcher-shaped, but variable, some distal end directed downwards towards s»b- with a pronounced concavity of the adcauline smiie- Petisare delicate, very faintly undulated. wall and a constriction behind margin, others no operculum. Gonophorcs—mule, with a thickening of the abcaulinc wall. Colour, Remarks: The U'Ophosome of the Bruny I. ^traw-coloured, gonotbecae bright orange. specimens corresponds very closely with Bale's Remark*. The hydrothecae arc very variable material of Pi wihoni from Griffiths Point. in shape even on the one hydrocladium. the Victoria. The submarginal constriction behind thecal walls ranging from pitcher-shaped to the hydrothecae of the present material is almost straight, often closely approaching the rather variable, being more pronounced in some shape ui the hydrotheca of P. sefaceoides. hydrothecae tiuin In othets even on the same hydrocladium. The hydrothecae with a shallow Pluinularia eratcrifnrmis Siechow, 1923b: 227. constriction closely approach those forms of FIG 32 P. setaceoides with more recumbent hydro- PhtmulnHa sctaceoutes var. craterijormis thecae. Tiebltcoek, 1 18, pi. 3. figs 8, Mulder & 1V10: The gonothceac of the Bruny I, specimens Ra; 1VI5: 51. pi. 7. figs 3, 3a. arc identical with those of Mulder & Trebil- Record: Adventure Buy, 4-6 T on the brown m cock's (1910) microsHde from which they alga Xlphophora gladiata. described the gonorheca of P. wihorii. Their Material: Infertile stems to 1 cm long. figure is. however, misleading, as the walls of Remarks on P. angusta and P, crateriformis: the figured specimens, like those of the present Sicchow (1923b, pp, 226, 227) raised Mulder material, are only faintly undulated, not heavily k Trebilcock's varieties of P. setaceoides (the ridged, as may be inferred from their figure. unnamed vars a, b. d. and var. t:ritterijormh\ A new record for Tasmania. Other localities: to specific rank, rm distinction between Hie Victoria; New Zealand. two species resting largely on the shape of the Agfaopbenia plumosa Bale, 1881' 25, pi- 14, hydroiheca, a character of considerable varia- fig. 6. 1884; 153, pi. 14. tig. 5. pi. 17. bility in this group and thus of doubtful diag- fig. 12; 1924: 257. Blackburn. 1942; 110. nostic value- The material from Bruny U Hodgson. 1950; ?& fig 87. Shepherd & although showing some intergnidaiion. can. Watson, 1970: 140. however, be fairly readily assigned to one or Record. Fluted Cape, 20 m deep, on sponge. other of these two species. This, together with the (act that the two Material; Rare infertile colonics. Stems to species occur together in the one locality, 1 cm long. although at different depths and on different Remark*. This is the robust form of A. pfo- olgal substrates (and one. P, angusia, was fer- PWW vvith short hydrocladial and stem inter* tile, while P. crateriformix was not) indicates nodes. Hodgson notes that his material col- thai these are valid, although closely related lected from Macroeywtis and drift had shorter species radiating From ihe central P. setueeoid?$ stems (L.c. the short internode foi'iVU than those stock. steins collected from the scagrass /.o.uera, Neither P, tmgnsta and P. crateriformis have which growls in sheltered waters. previously been recorded from Tasmania. They Thccocarpu* divaricatub var. typica iBusk, may have been confused with P. setaceotdts 1852). Shepherd & Watson, 1970; 140. P. an gusto is. known from Victoria and S. Aust.; Aftlaophenta divuricutu Busk, 1852; 398. P. crateriformis from Victoria. Records: Adventure Bay, 1-10 m deep, on PlumutariA wilswii Bale. 1926: 21. Ralph, horizontal faces among holdfasts of brown alga 1961b: 31. fig* 2, 3, Phytlositora eomosa; 10-22 in deep, cptlithic FIGS 33, *4 in sheltered situations. to 10 in height, Ptumularia drlfrotufa Bale. ISftl: 28. pi 15. Material: Stents infertile, cm in fig. 2; 1884: 137, pi 11, fig. !*, Midder & branched. Cawline nematothecac similar Tiebitcoek, IV 10: 1 15. pi. 2. fig. 2 shape to those of 7\ divaricatus var. cystifera, Records; Penguin I.; Adventure Bay: Satellite but much smaller, ilydrocladia close-set, hydro- I,, 3 m deep, on sponge. thecae crowded on hydrocladium, no oblique Mit/eritU; Sparse fertile stems to 1 cm long. intrunikU' sepia on ihecatc hydrocladial inter- (ionothecae u>p-^hapcd, 1 or 2 arising on a lude. Colour, dark brown. JIYDROIDS OF BRUNY ISLAND. SOUTHERN TASMANIA m lltecocarpns divancaru* (Busk) var. briggtJ cpizoic colonies on T, divarkaius var. briggsi Bale, 1926: 22. fig. 5. Watson. 1973; 194. from both Penguin I. and Satellite J. H.e, from Record- Satellite I,, on the red alga Thamno- protected and relatively rough-water situations) ilnm\tw dichototnum. depth recorded No had somewhat shorter, lax stems, givea off Material; Straggling, irregularly branched, singly from a stolon creeping on the stem of stems to 15 cm long. Colonies infertile, Colour, the hydroid host. The cauline internodes of hlacfc. these latter stems are long, with distant hydro- This is the first record of var. briggsi from clatUa, the hydrocladia themselves having long Tasmanian waters. Other localities, Port interludes, and the. mesial ncmatoiheea extends Jackson, N.S.W. (Bale); Pearson L S. Aust well over the mouth of ihe hydrotheca. Rare (Watson). epiphytic colonies on red algae at Adventure Bay had Remarks on the varieties oj T. divaricafus at the shorted stems. There were no discernible Bruny Island The two varieties of J\ divari- differences in microstruetures be- tween these cates recorded at Bruny I. are easily distinguish- three ecomorphs. Briggs (1915) mentions able by the presence of an intranodal ridge in that his specimens from Storm Bay were "4*6 in height the hydrocladium of var. btiggsi, as well as in mm (and) were found asso- differences in habit and substrate preferences ciated with Agiaophetu'u divaricate", Hodgson of each. (1950) did not record the substrate of his The var, brfa&si was found only in sheltered material from Black man's Bay in the Derwenl Estuary. rvefs near I Satellite T in the D'Entrccasteaux Channel, where it was a common epiphyte on Discussion Thamuoctonium, The more robust var rypka Ecology occurred on rock faces and as a common epi- Campanulariidae were recorded from all phyte on PhyUoxpiua holdfasts in situations of depths, being epiphytic on red algae growing moderate exposure to surge. mostly in sheltered places. One exception, Sili- Although showing affinities with van cysri- ctdaria rosea, was associated with the brown fera, the latter form docs not display the dis- algae Scytothaita dorycarpa and Seirococcu, tinctrve planar growth habit recorded for this axillaris in situations of moderate water move- variety (Watson 1973). nor the enlarged ment. cauJinc nematothecac. In microstrudures it The only representative of the Lafoeidac. most closely resembles a fragment of Busk's Flebella furax, provisionally recorded for the type of A g'.aophenia divaricate from Bass Strait first time in Australian waters, is a small form (NMV collection) and U thus recognized here cpizoic on the stems of Thecacarpat divarica- as var typica. tes var. hrigpxi Bale. The majority of species T, divaricates is a common and variable of the Lafoeidac known from Tasmaman and species of the southern Australian coast. mainland Australian waters arc larger forms Further study is necessary to elucidate the sys- from the deeper continental shelf, hence are tematic status and ecological relationships of unlikely to be found in a shallow water collec- the varieties of this tion species. such as the present one from Bruny I. With the exception of Phylactorhectt Hatfcornaria longlrostris (Kirchenpauer, I872>. armata and Halccium ..II Bale. I8S4: sp., haleciid species in the |SJ. p (. 13, fig. 7, pj. !6. collection are epizoic forms. Hulecium delicatu- fig. 3, pi, 19, tie. 30. Hodgson. 1950: 51 lam Cotightrey [H. fig, 83. Watson, 1973: 197. flexile of Hodgson (1950)) is one of i he most abundant Records; l. bydroids at Bruny Penguin f 10-20 m deep, on Iheco- the luxuriant orange-yellow carptts divaricarus: Adventure Bay, 10-20 m J„ colonies growing on the crusiose bryozoan Mcmbrinopora deep, on red algae and epilithtc on vertical mem- brinacca epiphytic on old stipes of the large faces: Satellite I. 12m deep on T. divaricates r kelp var. wtggth Macmcystn pyrifem. The two species newly described, Hafeciutn hrunieruis and Material: Luxuriant colonies, stems 3-8 cm H. luteuni, as well as H beanii. are of cryptic long, unhranched, infertile habit, the two new species growing on sponge. Remarks: The material from Bruny I. displays bryozoa, and rock in crevices, white H. heanii the same range in substrate as already noted for occurs on the underside of the large plate-like ff. hagirostris at Pearson I. (Watson 1973), alga Sondvrophycus australii, While fertile Epilithic colonics from faces* exposed to surge colonics of Phyiactotheca armata were recorded have the longest jud most robust stems, while abundantly at all rough-water sites, rhey in fact 174 JEANETTE ft W A' I SON also occupied a relatively sheltered niierohahitat Tecords for Tasmania, including 3 new records among the holdfasts of die brown alga rhyilo* for Australian waters. Seven of these specie* xpora comosu. are already known Irom ihe Vieto/ian coast- Remarkably lew species of Seuulariidae were line of lias* Strait, so would be expected to recorded- Of greatest interest is ihe first record occur among the Tusmanian fauna. Theeocar- from of Sixluctu farquhm (Rule) outside New Zea- pux ttfpatitatus var. &ff{gpjf4 now known land waters (sefe further discussion beluw). S. N.S.W. (Bale 1926). Tasmania and -South Aus- iarquitari was. however, rate at Bruny I. Sertn- tralia (Watson 1973) has not yet been recorded lart'a acuta Stechow. while one ol the most from Victoria. abundant epiphytes in the collection, was asso- Ol" Ihe 3 new records for Australia, flcde- ciated only with red algae, and not with chtm beanii is cosmopolitan, Hebella juntx is Mucracysiis as were Hodgson's specimens known only from South Africa, white Salutki Hodgson (1950) also recorded very abundant fa/Qithtnl is a New Zealand .species recorded colonies of AmpfvsbcJia operculum on old only from the South Island, where it does no:. scallop shells from the DT.ntrecasleaux Chan- however, occur north of 43 *S. The absence of nel. Careful search in Ihe area of the DT.ntre- S. farquhari from mainland Australian waters casteatix Channel covered in this survey pro- may thus be attributable to this southern distri- duced only one attenuated colony. Although bution, as Bruny I. also lies close to 43 S. display strong sea- A. npercuiata is known to The profound influence ol the East Australia the rootstock and pat is sonal growth, some of Current And the West Wind Drift in the dis- persist from one season Of rhc colonics usually persal of species and the biological and zoo- virtual absence ro another (per*, obscrv.l. The geographic relationships of the trans-Tnsman habitat may be of this species from a former hydroid fauoa have been discussed at length changes in the eco- explained by permanent by Ralph (1961c). In this repaid, three species about by invasion of the gastro- system brought recorded from Bruny I. are of considerable following the collapse pod Maoriculpux rosetts, interest. These are Ihe two halceiids newly Probably the of the scallop dredging industry. described, and the occurrence of S. jurquhari. shell of M. roxeux does not offer an attractive Hidccium bnmwndK bears a strong resemblance larvae of A. operadata* substrate to the to H. knlk-idun\ a species known from the occurrence in Of particular interest is the South Island of New Zealand and Cook Sir air. Plumtdaria an- unc locality of Ihree species, while H. htleum displays close aflmiiies with wilxoru, all of gUSlfo P cmterijormis and P H. vors'tiMtussUmm* a rare species recorded HVDROIDS OF BRtTNY ISLAND, SOUTHERN TASMANIA 175 References Allman, G. J. (1872).—A Monogiaph or the Bvuk. G. 11852).—An account of the Polyzoa. Cymnobtasric or Tubiilarian Hydrnids, R(JJ, and Scriularian zoophytes. In J. MacGillivrav, Part I. 11 Site Lond. 1871; Purl f !S72> "The Narrative of the Voyage of the H,M.$ l Allnun. G. J. ilg&a i.—Report on Ihe Hydroida. ^Rattlesnake' \ Appendix 1V. 385-402. Part J. Plumulariidae. Rpi. Set. Res. HM.S. (Bonne- London) Challenger. Zoology 7(2), 1-55, Plates 1-20. Coughtrev, M (1875).—Nous on the New Zea- land Hydroideae. AUMAhr, G. J. (1888),— Kepoit on ihe Hydroida Tratif. Ptot: A'.Z. Inst. 7, 281-293, PI. dredged by H.M.S. "Challenger" during the 20 Couohirey, M. ( Critical years 1873-1876. Part 1J. Ihe Tubularmae, 1876a)— Holes on Ihe New Zealand Corymorphinae, Campanulannjje, Scrtu- Hydroida. front. Proc. NZ 298-302. larinae and Thalamorpha. Rpt. Vow Challen- iml S, Co-lghtrev. ger 1873-1876, 23(70). 1-90. M. i l£T76b>.—Critical notes on the New Zealand Hydroida. Ann. Mug. MoL ftteL Bale, \V. M, (1881).—On the HyOroida or south- scr, 4, 17. 22-32. PI. 3. eastern Australia, witb descriptions of sup- Hincks, T. 1 1853) —Huther notes on Bnush posed new species, and notes, on the genus Zoophytes, with description* oi' new species. Axtaophvnia. J. ntfer, Soc. Vkx. 2(1), 15^47, Ann. Max. Nnt, Mist, ser 2, 2. 178-185, Pis Pis XH-XV. 5, 6. Bale, W. M. ( 1884 1.—"Catalogue of the Aus- Hiroi inr>, £mt>efor of h.pan ( 1 y69 ) —Some tralian Hydroid Zoophytes." (Auit. Museum; hydroidN of the Amakusa Islands. Sydney.) Mai. Lob. Imperial Household, Tokxo 9, l->2. Bale, W. M. 11888).—Un some new and rare Hodgson, M. revision hydroid.s in the Australian Museum collec- (1950),—A of the Tav mnnian Hydroida. Pap. Proc. tion. Proc Linn. S'oc N.S.W. 3, 145-199. R. Soc. 7asm Pis XH-XXt. 1949. 1 65. HciniN. F. W. (1873).—On Ihe New Zealand Bale, W, M. (1894).— Furfhcr notes on Aus- seilufarianv. Trans. Proc. vV.Z. fn.u tralian Hydroida 5, 25(*- whfi descriptions oi some 259. new ssecfe& Proc. R. Soc Met. (n.s.) 6, 93- 1ohnston> G. (1R3«).—-A historv of Ihe British 117. i>ls Ul-VJ. Zoophyl-s." 1st ed. (W. H. Uzars: Edin- Balk, W, M, I JV|3).—Further notes on Austra- burgh. I lian hydroids. Jl. Proc. H. Sot'. I'fct. \t\.s/\ KtRCHtNPAUEa, G. H. ( 1 872 ) .—C'ber 26, 114-147, Ph Xll XIII. dk* Hydroidcnfamille Plumulandae. cinzelnr Kaie, W. \f. (19l4 t -KcpOil On the O.— Hvdroida Gruppen derselben [inc\ ihre Fnichtehtiltcr. 1. collected In the Great Australian Bight and Aglaaplwruu, Abh. naiurw- Vet, Hambun* 5. vt other localities, Biol. Rrs. FJ,S fc.'ilde^vour*' 1-52, Pis IVIU. 2(1), 1-62, Pis I-Vri. — Lamahck, J. B. 1 1816). "Hisioire naturalk dss Bale. W. 1 M. ( 19 Ub).— Further notes on Aus- Animaux sans Vertehres, II, Polypes." tralian hydroids. Ill, Proc: R. Soc. Vict. Lamourolix, M J. V. F. <; 1816).— His|aire dea (n.s.) 27, 72-93, Pis XI-XH1. Polypiers Coralligenes f*c\ib]e<, vulgniremeni Baif, W\ Mi (1915).—Report on the Hydroida nommes Zoophytes" (Caen) collected in the Great Auslialinn Bight and LnooK. F. (1934).—Tmis hydmpolvpes de \u other localities. Part Til. Biol. Res F.t.S. bale de la Tabic. Ainqne, Anstrale. Bull "Endeavour" 3(5), 241-336. PH XLVI- Mhs, Hist. nat.Bvts. 10(19). 1-7. XLVII. LcNDENPRi.o, R. von. Oher Baie. W. IVf. (1924).—Renort on some hydroid*. fl883).— CoeJen teratcn del Sud^ec. IV. from the New Zealand coast with noles on Miuheduna Eueopelhi cumpanukiria nov New Zealand Hydroida generally, supple- gen. Zvhsvhr. f.iHss Zoot. LtipzwWA), 497-582. Pis 27-32. menting Parquhat's list. Trans. N Z. Inst. 55. 22<-268. TjNNACUfi, C. (1758).— 'Svstema Namrae." lOlh ed. Hale. W. M ( 1 926).—Further notes on Austra- (Lipsjac.1 lian hydroida. V. Proc. R Sot: Via. (n.s.) MtVEN, K, J. F. (1834),— Beitrage ztir Zoolouie 13-23. **> V'esammelt >ad* einvr Reise urn die Erde.^V M. BucKbUkn. (1937).—Report of the expedi^ Ucbei das l.eueh1en de^ Mtere« und Bcschrci- rion oi the McCoy Society for held investiga- bun^ einigci Polypen und anderer niederer tion and research II. Cuelenterata, Proc. it. Thicrc. Nova Acta AteiL Caes. Leopold tine. Vki (rtAJ 49. 364-371 Caret, siippl. 16, 125-216. Blackburn, M. (1942).— A systematic list of The MniAfltv N. A. H. (1957) —The Hvdro^oa of Hydroida of South Australia with a summary False Bay, South Africa. Ann. S. Aft: Xfus. of their distribution in other seas. Trans H 4% 173-243. Soc. 5. Aust. 66(1 ( 104-118. Mrlartj, N. A H (1958). -Hydrozoa from Ihe Bhiuos, E A. (1914),— Hydrozoa from one co39t5 of Natal and Portuguese E-A*f Africa. hundred lathoma, seven miles enst of Cane Part r. Cahptotdtntea. Ann. S. 4h\ Mux. Pillar. Tasmania. Rec. Aust Mut. 10(10), 44, 165-226. 285-30!. Pis 25. 26, Millard. N. A. H. (1964) .^The Hydrozoa of BUM IL A. (I?|J), Notes on Tasmania* ihe 176 JEANETTE E. WATSON Millard, n, A. H. {1966).—The Hydrozoa of Ralph, Patricia M . ( 1961a).—New Zealand the south and west coasts of South Africa. Lhecate hydroids. Part III. Family Serin- Part III. The Gyinnoblastca and small lariidae. Trans. R. Soc. N.Z 88(4), 749- families of the Calvptoblastea, Ann. S, Afr. 838. Mas. 48, 427-487. Ralph, Patricia M. (1961b).—New Zealand Millard, N. A. H. (1968.).—South African thecate hydroids, Part IV. Plurnularitdae. hydroids from Dr. Th. Moi'tensen's Java- Trans. R. Sac. N.Z. (n,s.) 1(3), 19-74, South Africa expedition. 1929-1930. Vidensk. Ralph, Patricia M. ( 1961c). -New Zealand Meddr. dansk nuttnh. t'oren. 131, 251-288. thecate hydroids. Part V. The distribution of the Zealand thecate hydroids. Trans. R. Millard, N. A. H., & BoutLLON, J. (1973).— New Soc. N.Z. Zool. 1(7)> 103-1 1 L Hydroids from the Seychelles (Coelentcrata). f Shepherd, S. A., & Watson, Jeanette E. (1970).— Artnls Mm. r. Aft. cent. ser. 8, 206, 1-106. The sublittoral ecology of West Island, South Mulder, J. F. & Trebilcock. R, E- (1909).— f Australia. 2. The association between on Victorian hydroida with descrip- Notes hydroids and algal substrate. Trans. R. Soc. tions of new species. Geclong Nat. 4, 2nd S. Aust. 94, 139 146. ser., 29-35, PL I. Stkchow, E. (1921).—Uebcr Hydroiden der Mulder, J. F., & Trebilcock, R. E. (1910).— Dcutschen Tiefsee—Expedition nebst Bemcr- Notes on Victorian hydroida with descrip- kungen liber cinigc andere Formen. Zool. 4 2nd tions of new species. Gedona Nat. ( Anz. 53, 223-236. ser., 115-124, Pis 11, III. Siechow, E. (1923a).—Neue Hydroiden der Mulder, J. F.. & Trebilcock, R. E. (19 14a). Deutschen Tiefsee— Expedition nebst Bcmer- Victorian hydroida with descriptions of new kuttficn liber einige andere Formen. Zool, species. Part III. Geetonx Nat. 6(1), 6-15, Anz, 56, 1-20. Pis MIL Stechow, E. (1923) —Zur Kenntnis der Mulder. J. F., & Trebilcock, R. E. (1914b).— Hydroidenfauna des M ittelmeeres. Amerikas Victorian hvdroida with descriptions of new und anderer Gebiete. Zool. Jb. (Svst.) 47, species Part IV. Gccionx Nat. 6(2), 38-47, 29-270. Pis 1V-VL Stechow, E. (1924).—Diagnosen neuer Anz, 59, Mulder. J F n &. Trebilcock, R. E. (1915). Hvdroiden aus Auslralien. ZooL Victorian hydroida with description of new 57-69, species. Part V. Geeiong Nat. 6(31, 51-59, Stechow, E. (1925).—Hydroiden von West- und pis vn-ix. Sudwestaustrulien nach den Sammlungen von Ralph, Patricia M. (1956).—Variation in OMm Prof. Or Michaelsen und Prof. Dr Hart- Syst. 191-269. geniculate (Linnaeus, 1758) and Sifkularia meyer. ZooL Jahrb. Abt. f. 50, hilabiata (Ooughtrcy, 1875) (Hydroida, F. Thompson, D'A. W. (1879).—On some new and Campanulariidae). Trans. R. Sac. N.Z. rare Hydroid Zoophytes (Sertulariidae and 84(2), 279-296. Thuiariidae) from Australia and New Zealand. Nat, Hist. (s*r. 5)» 3, Ralph. Patricia M. ( 1957).—New Zealand Ann. Mag. 16-19. thecate hydroids. Part I. Campanulariidae and 97-114, Pis Carnpanulinidae. Trans. R. Sot. N.Z, 84(4). Trebllcock. R. E. (1928).—Notes on New Zea- 811-854. land Hydroida. Proc. R. Soc. Vict, (n.s.) 1-31, Pis 1-7. Ralph, Patricia M. ( 1958).—New Zealand 41(1), (1973). Pearson Island thecate hydroids. Part 11. Families Lafccidae, Watson, Jeanette E. — Lineolariidac, Haleciidae and Syntheciidae. Expedition 1969.—9 Hydroids Trans. R. Soc. Trans. R. Soc, N.Z. 85(2), 301-356. S. Ausl 97(3), 153-200. SHOOT AND FOLIAGE PRODUCTION OF FIVE SHRUB SPECIES OF ACACIA AND HAKEA IN A DRY SCLEROPHYLL FOREST BY J. R. MACONOCHIE* Summary MACONOCHIE, J. R. (1975) .-Shoot and foliage production of five shrub species of Acacia and Hakea in a dry sclerophyll forest. Trans. R. Soc. S. Aust. 99(4), 177-181, 30 November, 1975. An examination has shown that shoot growth of Acacia niyrtifolia, A. pycnantha, Hakea rostrata, H. rugosa and H. ulicina is distinctly seasonal. Growth commenced in spring, finished by mid- summer and was preceded by flowering. The maximum rates of loss of foliage occurred towards the cessation of active shoot growth and both mature and juvenile foliage was lost. Measurements of size of shoots of the three species of Hakea over a series of years suggested that the available soil moisture during the growth period was the controlling factor for shoot size. It is further suggested that the growth habit of these three species is reflected in the pattern and size of new shoots along a parent shoot. H. ulicina, the species which showed a tendency toward apical dominance is an erect, several stemmed shrub, whereas H. ro strata and H. rugosa are rounded spreading shrubs. ) SHOOT AND FOLIAGE PRODUCTION OF FIVE SHRUB SPECIES OF ACACIA AND HAKEA IN A DRY SCLEROFHYLL FOREST by J. R. Maconochie* Summary Maoonochh:-, J. R, (1975).—Shoot and foliage production of five shrub species of Acacia and Hakea in a dry sclcrophvll forest. Trans. R. Sac. 8, Attst, 99(4), 177-181, 30 November, J 975. An examination has shown thai shoot growth of Acacia myrtifolia, A. pyenantha, Hakea rostratu, H. rugosa and //. ulicina is distinctly seasonal, Growth commenced in spring, finished by mid-suromer and was preceded by flowering. The maximum rates of Joss of foliage occurred towards the cessation of active shoot growth and both mature and juvenile foliage was lost Measurements of size of shoots of the three species of. Hakea over a series of years suggested that the available soil moisture during the growth period was the controlling factor for shoot size, It is further suggested that the growth habit of these three species is reflected in the pattern and size of new shoots along a parent shoot. /?. ulicina, the species which showed a tendency toward apical dominance is tin erect, several stemmed shrub, whereas //. rostwta and H. rugosa are rounded spreading shrubs. Introduction These data for axillary and terminal shoots This study on shoot production of the five were separated and the annual mean shoot temperate shrubs. A cacia myrtifolia (Sm. sizes compared by analysis of variance. Willd., A. pyenantha Beruh., Hakea rostraia F. Climatic data were supplied by the Austra- Muell, ex Meisn., H. rugosa R. Br. and H. lian Bureau of Meteorology, ulicina R. Br,, was part of a project in which Results shoot growth was measured on other shrubs from the arid and semi-arid areas of South Qualitative Aspects The percentage cumulative gains and losses Australia ( Maconochie & Lange 1 970, of foliage for Maconochie 1973), to obtain basic phenologi- A. pyenantha and A. myrtifolia are cal data in a range of habitats. presented in Fig. 1, .4. myrtifolia did not produce any new shoots in the first year of the Study Site and Methods study but the burst of growth during the This study was carried out on the boundary second year was in phase with that of A- of the Para Wirra Reserve in the Ml Lofty pyauintha. The "stepped" effect displayed by Ranges, South Australia from April 1965 until A. pycnaiuha was also produced by H. January (967. The occurrence and size of new rostratu, H. rugosu and //. ulicina. New shoot shoots produced on about 120 tagged shoots growth was distinctly seasonal- of each species of Hakea and A, pyenantha^ Figure 2 presents the relative rates of gain and of 25 tagged shoots on two bushes of A. and loss for the five species. Climatic data are myrtifolia, were recorded monthly. The tech- presented in Fig- 3; soil moisture values were nique has been described previously taken from Martin & Specht (1962). (Maconochie & Lange 1970). Shoot growth reached a peak during Octo- From 1965 until 1971, samples of shoots ber when daily temperatures were increasing were collected at the end of the growing and soil moisture was available (Figs 2 and season from the three Hakea species and data 3). In both 1965 and 1966, flowering, which recorded on size and position of new shoots, commenced in early August and rinshed by the * Animal Industry and AgricuHure Branch, Arid Zone Research Institute. Dept. of the Northern Terri- tory, Alice Springs, N.T. 5750. 178 L R. MACONOCHIE loss of vycnantha total new positions; A Fig. 1. The percentage cumulative leaf gain and A. (, rtifolia new positions; A, leaves; •, scars). leaves; Q, scars), and A T my (, branches up to 5 end of September, preceded shoot growth. in some cases clipping off Immature flower buds on the Acacia species mm thick. nticina and H, roslrata were observed to develop as early as January. Some losses of H, 1966 were a result Both axillary and terminal flower and shoot during January-February plants. These bushes buds on the Hakea species were enclosed in of the death of several examined for cause of death, but bracts and these buds developed as the new were not was 223 below leaves matured. This observation suggested since rainfall in 1965 mm probable that localised drought that the size of new shoots for the next season average it is most likely cause of may be predetermined by conditions at the during summer was the time of bud development. death. once foliage had matured, Although the A. myrtifolia bushes did not Generally new loss the Hakea species produce new shoots in 1965, flowering did the rate of leaf on zero. The needle-like, rigid, take place. declined to almost leaves of these species arc The peaks in rates of foliage loss coincided sclerophyllous physical damage with or immediately followed the peaks of obviously more resistant to the mesomorphic production, with loss comprising both mature than the leaves of more senescing and soft immature foliage, Some species of the community. further loss was caused during a severe wind Quantitative Aspects and hail storm in December 1966 in which number of leaves per shoot for trees and branches were felled, and also by The mean for the years 1965 to 1971 bird pruning during July-August 1966. Parrots each species and presented in Table h Analysis were observed pruning all the trees and tall inclusive are that for some years shrubs, apparently at random, in the area and of these data showed 1 SHOOT AND FOLIAGE PRODUCTION IN ACACIA AND HAKEA U? //. rostrata—terminal and axillary shoots r=-f-0.52 P>0.10 y = 5.1 + 0.008* H. rtigosa—terminal and axillary shoots r= + 0.83 .05>P>.G2 y « 4.1 + 0.029x H, rugosa was the only species showing an acceptably significant correlation between mean shoot size and rainfall during the grow- ing season. Calculation of the correlation coefficient between the mean shoot size and rainfall period September-November of the preceding year gave the following: H. ulicirtu—0.65, H, rostram—0.07, and H. rugosa—0.55. 4 Analysis of the data for 1966 on the rela- tive positions of shoots showed that there were no significant differences in shoot sizes between pairs of positions distal to the apex (Table 2) except for H. ulcina. In this species there was a significant difference (P < 0.05 between the mean of the terminal and that of the first axillary position. Further analysis showed significant differences (P < 0.05) between terminal and axillary positions two and three. It would appear, therefore, that H. ML idkina is the only one of these three specie* which has a tendency towards apical dominance. Discussion Cambagc (1918, 1927) measured the height growth of A. pyenantha at Sydney Botanic Gardens and his study showed i * 9 C the normal pattern of rapid growth during the juvenile Fig. The relative rates of gain (block) and loss stage followed by a decreasing rate as the of foliage of (a) A. myrtijal'to, (b) A. plant matured. pyrnarttha, (c) i/. ulicina. (d) //. rostrata. The plants studied at Para and (e) //. rugasa, for the years 1965 and Wirra were mature and the rates of growth as 1966. retlected in both Figs 1 and 2 were of a "steady state" nature, as occurs at tbc plateau there was a significant difference between the of the sigmoid growth curve. sizes of shoots produced in successive years. Martin & Specht (1962) measured the In other years, the variation in shoot size was moisture relationships in a dry sclerophyll so large that no significance could be attached forest, of which these species arc shrub to the differences. com- ponents. Their studies showed that the more A set of correlation coefficients (r) and mesie community of this vegetation type had regression equations were computed between a higher index of evapotranspi ration and could total rainfall (x) for the period September to November be subjected to a drought period during the and the mean shoot sizes (y) for mid-summer. The three species of Hakea and successive years during which there were sig- two Acacia species all distinct nificant show a season- differences ( P<05 ) between the ality of shoot growth with a cessation occur- means. This rainfall period was selected ring in mid-summer probably during the because it was the time of active growth. drought period or when soil moisture is only H. M//Wna—axillary shoots sufficient to maintain a dormant growth phase. r^ + 0.74 0.10>P>.05 The negative correlation between mean y - = 5.0 4- 0.014x shoot size and soil moisture (as reflected by 180 J. R. MACONOCHIE soil moisture ex Martin Speehl (1962), other data Fig 3 Climatic data of Parra Wirra study site; & poten- temperature; 9 rainfall; Q, means for 1965-1970. , soil moisture; O, mean duration of active growth. tial evapotranspi ration (P/H 0.75); , florescence and TABLE 1 Mean number of leaves per shoot for seasons 1965 to 1971 Shoot Year 1970 1971 Species Posn. 1965 1966 1967 1968 1969 9.2 9.4 9,8 Hakea ulicina Total 7.4* 8.1* 8.2 8.7 Terminal 9.6 10.2 9.4* 9.4 10,9 10.6 11.2 Axillary 5.9t 7.1' 4.9 % 8.1* 6.7t 8.9 8.4 5.4$ 6,5 6,7$ 9.0 Hakea rostraia Total Mf 5.8t 5,2 Terminal 6.1 6.1 5.7 5.4$ 7.0 6.5$ 10.6 Axillary 6.5$ 5.7 5.2 5.4$ 6.4 6.8* 7.5 8.4r Hakea ru^osa Total 1.1% 9.1$ 4.6$ 8.7 9.2$ 13.7 Terminal 7.3* 9.4 8.1 8.8 8.8 9.5$ 15.9 Axillary 6.9$ 9.0^ 4.4 % 8.8 8.3* 8.9$ 12.7 indicated. Significant differences between mean and mean of succeeding year are * P < 0.05; fP< 0.01; t P < -001. TABLE 2 tip year 1966 Mean number of leaves per shoot for terminal and axillary positions distal from shoot for Relative AX< AX* AX r, ; AX T Position Term. AX AX. AX;! AX4 7.6 5.6 H. ulicina 109 7.8 7.5 ?.: 7.6 7.4 6.6 H. rosrraia 6.2 6.0 5.3 5,6 5.3 5.3 H. rugosa 9.3 8,6 9.6 10.2 10. 8.2 ::; Indicates significant difference (.05 > P > .01) between mean and that of succeeding position, h SHOOT AND FOLIAGE PRODUCTION (N ACACIA AND HAKEA 181 rainfall) during the growing period of the contrast, actively grew only during the spring previous year suggests ibat shoot size in the period at the study site, and although shoot following year is not necessarily determined growth on A. myrtifolia was only recorded at the bud formation stage. Rather these during the second year, it appears that from results suggest that shoot size is more likely this slender evidence that the shoot growl to be determined by the soil moisture during follows a seasonal pattern also. the period of active growth. Both H. routram and H. ru^osa have a more The time of shoot growth for these species spreading bushy habit than H. ulkhia which contrasts with that of heath studies of Specht has a tendency to be more erect. The habit (1957), Specht & Rayson (1957) and Groves of these plants is reflected in the sizes of new f J { 1 65 ) . who recorded that the shoot growth shoots on a parent shoot. Both H. roxtmra and commenced in December when soil moisture H. rugosa did not produce significantly smaller was decreasing Specht (1957) showed that 3xiilary shoots in comparison to the. terminal, drought conditions occurred in both December but the terminal shoots of //. ulicina were sig- and January. Groves (1965) noted that shoot- nificantly larger than shoots on the three suc- growth continued throughout the summer of ceeding positions below the apex. This suggests his study period; however a recharge of soil a tendency towards apical dominance and moisture from a mid-summer rain was thus explains the more erect habit of this recorded- species. Maconochie & Lange (1970) and Macon- ochie (1973) have reported the seasonality of A cknowledgments shoot growth on A> sowdeniu /i. Uxukifu and The author is indebted to Dr R. T. Lange, A, tnunayarWt and possihle non-seasonal and Botany Department, University of Adelaide, seasonal shoot growth responses on A. aneura for his encouragement in the first part of the and A. kempeana, Weiherell (1966) recorded project. Dr R. W. Rogers, now of the Univer- Mushes of growth on A. harpophylla m Queens- sity of Queensland, supplied samples during land during spring and summer, and during the years 1967 to 1970 and his assistance is one period of early winter. A. pycrtanthtt, by gratefully acknowledged. References Camdace, R- H. vertical (1918).—The growth of Martin, Helerte A. ? & Specht, R. L. (1962).— Irees. I. /. Proc R. Soc N.S.W. 52, 377-384. Are mesic communities less drought-resistant? study on moisture relationships in dry Cambagh. R. H. (1927).—The vertical growth of A sclerophyll forest at Inglewood, South Aus- trees. IJ. Proc, /?. Soc. N.SW. 61. 279-2K4. tralia. Aim. J. Bot. 10(2). 106-118. Grovhs, R. H. < 1965).—Growth of Heath Vege- Spechi. U. L.. & Rayson, Patricia. (1957).—DaTk tation, II. The seasonal growth of a heath on Island Heath (Ninety-Mile Plain, South Aus- ground-water podzol at Wilson's Promontory, tralia). T. Definition of the ecosystem. Auss Victoria. Aust. A Bot. XX 281-289. J. Bot. 5, 52-85, Swht, R. L. (1957).—Dark Island Henlh MicONOCHfE, J. R., & Lange, R. T. (1970), — (Ninety-Mite Plain, South Australia"). V. The Canopy dynamics of trees and shrubs with watei relationships in heath vegetation and particular reference to the arid- zone topfecd pastures on the Makin Sand. Aitst. /. Bot. species. Trans. R- Soc. S. Ai4st 94, 243-244. 5, 151-17^. Maconochiii, J,. R. (1973).—Leaf and shoot Wr.TirEHAU., A. J. ( 1966).—Leaf growth of Briga- growth on Ac arid kempcana T\ Muell. and low (Acacia harpophylla) suckers in relation selected other arid-zone species, Trnpiuil to seasonal conditions. QUI J, (fttrip, /iw/w. Grasslands 7(1), 49-55. Set, 23. 453-456. SHORELINE SHINGLE TERRACES AND PREHISTORIC FILLINGS OF LAKE EYRE BY J. A. DULHUNTY* Summary DULHUNTY, J., A. (1975). -Shoreline shingle terraces and prehistoric filling of Lake Eyre. Trans. R. Soc. S.Aust 99(4), 183-188, 30 November, 1975. Investigations were carried out of (i) development of contemporary wave-built shingle terraces during the 1974 record historic filling of Lake Eyre, and (ii) old shoreline shingle terraces built by prehistoric fillings of the Recent salina to levels above that of 1974. Evidence indicates at least three prehistoric fillings to levels of 280, 160 and 70 cm above that of the 1974 filling. No precise evidence of the ages of the prehistoric fillings was obtained, but estimates inferred from field studies suggested dates of the order of 3,000, 1,500 and 500 years before present. The "3,000" year old filling was probably the deepest during the Recent salina episode of the history of Lake Eyre. The possibility of more permanent, deeper and widespread pre- salina filling of the lake, and late Recent subsidence of its southwestern areas, was also inferred from field observations. SHORELINE SHINGLE TERRACES AND PREHISTORIC FILLINGS OF LAKE EYRE by J A. Dulhunty* Summary Dclhuntv, J. A. (1975). —Shoreline shingle terraces and prehistoric filling of Lake Eyre. Trans. R. Soc S, Ausl, 99(4'),, 183-188, 30 November, 1975 Investigations were carried out of (i) development of contemporary wave-built shingle terraces during the 1974 record historic filling of L<>kc. Byre, and (ii) old shoreline shingle terraces built by prehistoric fillings of the Recent salina to levels above that of 1974. Evidence indicates at least three prehistoric fillings to levels of 2S0, 160 and 7t> cm above that of the 1974 filling. "No precise evidence of the ages of the prehistoric iillings was obtained, but estimates inferred from field studies suggested dates of the order of 3,000, 1,500 and 500 years before present. The "3,000"' year old filling was probably the deepest during the Recent salina episode of the history of Lake Eyre. The possibility of more permanent, deeper and widespread pre-salina filling of the lake, and late Recent subsidence of its south- western areas, was also inferred from field observations. Introduction tudinal sand ridges rising steeply to as high j.s 10 m above the lake bed. Lake Eyre consists of a large northern area —Lake Eyre North, connected by a narrow Around Babbage and Hunt Peninsulas and channel to a small southern area —Lake Eyre in Belt Bay ( Fig. 1 A) , shores are beinij South (see 1:250,000 Topographical Sheets eroded in hard dolomite, silcrete and ferru- giniscd siltstonc 1 Noolyeana, Lake Eyre, Curdimurka) . It is a (Williams 1973 ) Wave salina forming the "sump" of a large internal action, during periods of filling, has produced drainage system (Bonython 1955; Mason quantities of pebbles or shingle consisting Q of slightly flattened 1 55; Wopfncr & Twidale 1967). Aridity and largely and rounded high rate of evaporation, over the greater part pebbles from 2-10 cm in diameter. Single of the drainage area, normally prevent river strands or beaches have developed, and break- water from reaching the lake. On infrequent ing waves have washed up across the beaches occasions when rivers flow into it, part or long parallel deposits of shingle. These whole of the lake bed is covered with water deposits, formed during the present salina which cannot escape and must evaporate. The episode of the lake's geological history, as dis- bed of Lake Eyre North slopes gently from cussed later, are Recent to contemporary in north to south, and evaporation of brines in age. They are wave-built, strand, shingle ter- the southern bays has produced salt crusts races forming ridges, terraces, ramparts and (Oulhunty 1974; Bonytlum 1956). banks, closely associated with the present shoreline of the salina. For the purpose of the When the lake contains water, its shores arc present are referred as "the actively eroded by wave action, producing paper they to well-defined shorelines with shelving strands shoreline shingle terraces of Lake Eyre salina". or low clitfs rising abruptly from its almost Gravel deposits were observed and recorded 1 flat bed. At some places, cliffs up to 10 m or by Williams (1973 , 1975) on Hunt and more in height have been cut in soft, partly Babbage Peninsulas and on the western side consolidated sediments. At other places shore- of Belt Bay. He described them as "strand lines lie along, or around the ends of, longi- gravels" and "old shoreline deposits". They • Department of Geology and Geophysics. The University of Sydney, N.S.W. 2006. 1 Williams, A. F. (1973).—Explanatory Notes for the LAKE EYRE 1:250,000 Sheet S. Aust. Dept. Mines. R.B 72/93 (unpubl.j. 1X4 J. A. DULHUNTY include the Recent shoreline shingle terraces water level at any point along the .shore* desert bed in this paper, a nd other gra vel measured after a calm period of not less than deposits, possibly at higher Levels, and more 6 hours, could be regarded as mean lake level remote from the present shoreline, which may plus or minus 10 cm. be older and related to prc-Recent history of Contemporary the Lake. Williams fpcrs. coram- J 975) also shingle terraces noted shoreline shingle terraces along the Studies of shingle terrace development southern shores of Lake Eyre South, which associated with the 1974 filling of Lake Eyre he regards as probably similar in age and were carried out in August of that year Mean origin to those of Lake Eyre North described lake level reached its maximum during May in this paper. He also found and examined It rose by only about 20 cm during ApTil. more remote terrat.es up to 20 km from %he remaining steady during May and fell by present lake shore about 20 cm during June and July Therefore, rhe The tilling &l Lake Eyre in 1974 to the shingle terraces studied in August had been Greatest depth known since European settle- built over a period of four months during ment, provided a unique opportunity for the which mean lake level rose and fell through study of contemporary shingle terrace develop- only about W cm. ment in relation to lake levels and wave action. Well-defined shingle terraces were built This was undertaken to gam a better under- along all shelving shores with shingle beaches standing of the nature and origin of the old around Hunt and Babbage Peninsulas, and in shoreline Jerraces and enable interpretation of Belt Bay They were built during periods of their significance regarding the maximum strong wind, by waves breaking and washing deplh to which the lake had been filled during shingle up across the beaches to form lei races its role as the salina of the present internal above mean lake level. It was evident that shingle moved slowly drainage sysfcm* and the time since it was Up the beaches as the filled tu. or above, the level of the 1074 filling- lake filled, then formed into fully-developed Such investigations and results, recorded in terraces during the maximum mean lake level this paper, are confined to the significance only period, and finally remained stranded when of terraces along present shorelines, arid do water fell, providing evidence of the highest not represent a cornprehensvic study of all level reached, shingle deposits and their regional distribution The 1974 contemporary shingle terraces around l-ikc Eyre, varied in width up to 500 cm. and in depth or thickness up to 151) cm above pre-exislmg beach material. The height of their lops above Investigation of shingle terraces maximum mean lake level ;ii determining levels of shoreline shingle ter- group, although the height of their lops above races tn relation to each other and to mean mean lake level varied from place to place lake level. Short term variations in wafer level depending on degree of exposure to wave were caused by wind tides, currents and action. It was found that the general level of possibly other factors. Wind tides were most the tops of ihe highest terraces, on exposed pronounced on windward shores where water shores, was a uniform feature of a group, and level rose and fell, within a maximum range of the most significant feature in determination, about 45 cm, very quickly with rise and fall after the water level had fallen, of the mean of wind velocity and wave height. When wind lake levd at which they had formed, There- dropped, wave action usually ceased and water fore, it was concluded that the maximum mean evel fell lo "nnrmar wllhfn 5 to 6 hours. lake level of any past filling would be very "Normal" watci* level or the level free of wind clove to 90 cm below the general level of the effects during periods of calm, varied as a original tops of the highest shingle terraces result of other factors within a range of about produced dining that filling. 20 cm. Tn view of this n was assumed, for Greater depths of water during prehistoric the purpose of Ihe present investigation, that fillings are not likely to hnve produced waves t SHINGLE TERRACES AND FILLINGS OF LAKH EYRE UCALfTV BUtft SOUTH.V^- -^lOM prehistorc iAflAce »-i'.-E:. I LAKE dWUB MIUCLE t TERGAL COO' Tfi fcF "I'lrii"" i'fi * f-Ou" rff &p '5W&Y 5M1WGLE LQMITE - 1 ''fr _| J f _"_"_.'" OflY.LAKE GEO BHffiEUNE 7hVtL-'-~- " |^>£j - !--—! 1 I — D* SECTION F- ri£.E?CftrjS10NI ACWOSH SHINQts TEKKACES Fig. L Wave-built shoreline shingle terraces in Lake Eyre North. of greater amplitude than in 1974, or terraces at levels above that of the 1974 terraces, fol- with tops more than 90 cm above mean lake lowing contours across sloping terrain behind level, as the 1974 water depth was more than and above the 1974 beaches. Excellent one half the wave length of average strong examples occur along the eastern shoreline of wind waves. This was sufficient "to allow Hunt Peninsula between Willow Head and development of waves as high and long as the Campbell Point, immediately north of the limited fetch across the lake could produce. Elliott Price National Park fence, at numerous Increases in depth of water, within the limits places on Babbage Peninsula and Bonython of beach heights, would not appreciably in- Head in Belt Bay, and on Silcrcte Island off crease distances of water across the lake as the Bonython Head (Fig. 1A). Although eroded, shores are relatively steep. If average strong the old prehistoric shmgle terraces all bear winds during prehistoric fillings were similar close resemblance in general form and nature to those of today, heights and lengths of waves to the 1974 terraces, and follow contours con- must have been similar to those which built forming to the present shoreline. They were the present day terraces. formed in a manner similar to the 1974 terraces, by previous fillings of the lake to Old shoreline shingle terraces depths greater than that of 1974 which was At many places along the shingle-strewn the deepest known filling in white man's his- shores, old partly eroded shingle terraces occur tory. Wave action which formed the prehis- 1*6 S. A. DULHUNXY shingle terraces of the lOtic terraces would seem to have been similar Mostly the old greatly eroded or to that of the [VTA filling, as already discussed. highest group have heen Cower rates of evaporation in prehistoric mid completely removed. However, in protected erovion has occurred, their 10 laic Rec«nl rime could have resulted in places, where less which greater duration of peak lake- level periods and tops reach uniform maximum levels Singer limes of terrace format iom than in vary in different areas from 280 cm to 350 1974- However, this would not necessarily cm above the 1974 maximum mean lake level. have produced terraces higher than the equilib- ' They reach levels of 1 541 cm along both Peninsula. rium height, at a given mean lake level, for a eastern and western sides of Hunt of ftabbage Peninsula, given wave height. From observations, 320 cm on both sides equilibrium terrace height appeared to have and 280 cm on Silerete Island. Comparison 1974 bee* reached after 4 months in 1974. So the with the shape and general nature of the of the terrace top height of 90 em above mean lake terraces 1 to from the H973 ) at distances up 8 km I'i-l ! liiU no Fillings ouiside the field of present b;ke shore iyinji Aa discussed above, the terrace-lop level of contain Coxiella which Is this investigation, the highest group of old shore-line shingle recent shoreline terraces. They absent in the terraces on Hunt Peninsula wus 370 em above disturbed by IcetODism, fcnrt may have been the peak mean lake level of the 1974 idling, older, possibly Pleistocene, could teprescnt and the contemporary 1974 shingle icnacc-top of an earlier phase of lake history water levels level was 90 cm above the mean lake level at as discussed later. which the terraces were formed (Fig ID). Therefore, the peak mean lake level of the It was concluded that the highest, group of Jilting which built the highest group old shingle terraces on the present shnrelme prehistoric terraces must have been very slope was built bv the deepest lilHng of Lake of old shingle that of the 1974 tilling- Pyre during the Recent geological history of close to ISO'cm above This that the peak mean lake level of the snlina as we know it to-day. Terraces Of means highest prehistoric filling of Lake Byre this group were studied up greatest detail to the Recent lime, would have reached a ascertain as precisely as possible the height salina, in above the lake bed and age of the deepest prehistoric filling of depth of about 640 cm shores of Madman Gulf. the presen! valina. along, the southern SHINGLE TFRKAChS AND KILLINGS OF LAKE EYRE m about 850 cm in the deepest pari* or" the gulf, Shell material was absent from both con- and a level of approximately 280 cm above the temporary and old terraces, suggesting that the 1974 water level in Lake Byre South and along kilter were built hy intermittent fillings he I Centra? Australian Railway line. between which aridity and dryness of the fevefs i terrace tops in The of he the middle salina prevented establishment of shell fish and lowest or the three groups of terraces on The more remote and higher gravel deposits Hunt Peninsula measured 230 cm and 140 cm, described by Williams all contain small gastro- respectively, the above 1974 peak mean lake pod shells (Williams 1973', and per?;, comm. level. Alluwing for 20 cm of erosion, and 90 1973) supporting the suggestion that they were for cm (he height of the terrace top levels deposited during an earlier pre-salma stage above the lake mean levels at which they when the lake was permanently filled wuh were built, the two groups would represent water carrying a shell fish community, Studies fillings the ot lake tn depths of 160 cm and by King fl°5r\ I960) suggest that gypseous 70 cm above the 1974 peak mean take level lacustrine clays with Coxiella occurring along iPig. ID). the southern shores of Lake Eyre are nf The three groups of old shingle terraces Pleistocene ago. and that younger shoreline described above, were built at widely spaced deposits snd sand ridges originated in associa- intervals of time, during which many other tion with recent delation of the lake and major fillings must have buik terraces, a: inter- development of the present salina. mediate and lower levels, which were probably Fluvial erosion of the old shingle terraces destroyed or modified by redistribution of bears evidence of their antiquity The highest shingle during subsequent deeper fillings. Any and oldest group has been extensively eroded, terraces built by a Tilling to a level intermediate and completely removed in many places, whilst between the depths represented hy the lowest the middle and lower groups have each suf- and middle groups, during the Interval fered successively less erosion. Rainfall is very between the building of the middle and low, bin this is offset by high runoff from arid highest groups, were partly or wholly surfaces. FJuvial erosion under these condi- destroyed by the filling which built the middle tions could be comparable with that of higher group. Similarly terraces built by fillings to rainfall area*, where runoff Is icdured by depths less than thai of the 1974 filling, includ- vegetation. Wind erosion is not effective as the ing terraces of the 1950 filling, have been shingle is too large te be removed. modified or removed by the 1974 filling. Where shingle terraces of the highest group Antitftiifv of fitting have not been completely removed, they have The ages of the three prehistoric fillings of been breached in places by flowing! water dur- Lake Eyre, to depths greater than that of the ing rare occasions of heavy rain Terraces J 974 ve-Ty filling, arc difficult to ascertain pre- built across pre-existing or antecedent creeks, cisely. The only available evidence is very trend upstream in L'-bends as would contours. vatfuc and indefinite, and the only conclusions Where hreached, their eroded ends still turn possible are of th^ order ot* magnitude of age upstream (Fig. 1B). Terraces built across rather than exact periods of time. gently doping surfaces with no pre-existing Extensive searches were made for creeks, have been hreached in places by water carbonaceous materia) within the old shoreline held back on the sloping surface, leaving the shingle terraces, suits hie for carbon dating. No eroded ends of the lerraee "in-line" and not shell material or animal bones were found. No turned upstream (Ftg. 1B, C). Such breaches plant matcrrat such as driftwood was found, are frequently about 8 m wide, and the newly although sticks and fragmentary pieces of formed subsequent creek channels running wood were plentiful in the contemporary 1974 through the breaches have eroded down into terraces, Plant roots Were found in varying underlying weatheiing dolomite to depths of degrees of decay, but rt evident was that they 130 cm below the bases of the terraces. Tribu- belonged ro plants which grew on the terraces tary consequent creeks carrying water along after rhev were built. The environment of the top sides of the terraces towards the burial in the old shrngie terraces was evidently breaches have eroded channels in dolomite to one of maximum aeration, oxidation and pene- 50 cm deep (Fig. 1C). Whilst erosion oF (his tration of rain water, lending to complete des- kind is very difficult to refale |p time, it is truction ^nd loss of all plant material evident thai the terraces concerned are of con originally Included in the terraces. siderahle antiquity. — !** J. A. DUL1IUNTY The close proximity and relations of the old After making extensive field studies, taking shingle terraces to> the contemporary terraces all factors into account, and balancing the indicate very little modification of the shore- xnmewhai conflicting significance of" different line in general since their formation, suggest aspects of evidence, it was concluded that the ing limited age. oldest and highest group of shingle terraces Exudation of highly mineralised ground could date from more than 1,000 years befoie water, due to aridity, has produced much sur- present, but. less than 5,000 years, Further face and shallow sub-surface cementation of narrowing of these, limits must involve infetred Pleistocene and early to mid Kecent sediments. speculation, but in the author's opinion the Cjypcrete, silcrete, ealcrcte and ferruginite have age may be of the order of 3,000 years, which 1 estimate of the age of the deepest been formed (Williams 1973 ; Wopfncr & Twi- provides an ddle 1967). and the processes still appear to prehistoric filling of the present salina. The he in operation. No cementation has been middle and lowest groups of old shingle found ih the old shingle terraces or the more terraces arc certainly younger, and could well sandy and earthy material forming their basal he of the order of 1,000 and 500 years, respec- layer*. Suggesting limited antiquity. tively. The age of 500 years for the lowest and The evidence outlined above is not at all youngest group of old terraces, occurring just precise or satisfactory in attempting to estab- above the contemporary 1974 terraces, pro- lake lish ages in years, but it is all that is, as yet, vides an estimate of the time since the available For purposes of geological history, had previously been filled to the level of the the evidence is more significant, indicating that J374 iUling. the old shoreline shingle terraces are almost Acknowledgments certamly of mid to late Recent, and most probably late Recent, age. For the purposes it is wished to- acknowledge () valuable ol physical geography and hydrology of Lake assistance and co-operation of Muloorinp Sta- hyre, estimates are required for (i) the age tion which made field investigations possible, A. F. in years of the deepest prehistoric fining of the (ii) valuable discussion with Mr salina under existing environmental conditions, Williams of the Geological Survey of South provided and (ii) the number of years since the lake- Australia, and (iii) research facilities had been previously filled to the level of the by Ihe Department of Geology and Geo- 1974 filling. physics, University of Sydney. References Bonytmon, C. W. (1955).— in uU#e F.yre, South ICiNC, D. (I960)-—The sand ridge desert* of land- Australia. The Great Flooding Ot !94SM95(ft South Australia and related Aeolian Irons, The report of ihe Lake Eyre Committee, K. forms of the Quaternary arid cvclc. H- eeogr. Soc Aust. (S. Aust. Branch), pp. 7-9, Soc. S. Aust. 83, 93-108. 27-56. 63-70. (Griffin Press: Adelaide.) Mason. B. (1 955). In "Lake Byre, South Aus- tralia. The Great Flooding of 1949-1950* W. (1956). The Salt of Lake BoNYtftON, C The Report of the Lake Eyre Committee, R fcvrc »t.s occurrence in Madigan Gulf and — grogr Soc. Aust. (S. Aunt. Brunch), pp. II- Its possible origin. Trans. H. Snc. S. Aust. 2&. I'CitttTin Press: Adelaide.) 66-92. 79, Wuuams, A. V. (1975).—LAKH FYRF map distribution Dirt monty, .1. A. (IV7*1)<— Salt crnst sheet. Geological Atlas of South Australia areas of and lake bed conditions in southern 1:250.000 scries. Geoi $W\ Si AtiSJL Take F.vre North. Trans. K- Soc. S. Aust, Wupfner, H., & Twumi.k. C. R. (1967),—Geo i»8(3), 125-I33. morphological history of the Lake Eyre stratigraphie Basin, in J. N. Jennings, & J, A, Mahhutt Kin<7, D. ( 1956),—The Quaternary studies from Australia and record at Take Eyre North and the. evolution Fd.s.. "Landform of existing topographic forms. Trans. R. Sov. New Guinea", pp. 118-143. (A.N.U. Press, S. Aust. 79. 93-103. Canberra) NOTES ON THE RELICT PALM LIVISTONA MARIAE F. MUELL. IN CENTRAL AUSTRALIA BY P. K. Latz* Summary LATZ, P. K. (1975). -Notes on the relict palm Livistona mariae F. Muell. in central Australia. Trans. R. Soc. S. Aust. 99(4), 189-195, 30 November, 1975. Comparison of photographs taken after intervals of up to 56 years indicate that the relict palm Livistona mariae may reach an age of up to 300 years. Aspects of the ecology of the palm are presented. NOTES ON THE RFXICTPALM UVISTONA MARIAE F. MUELL. IN CE3STRAL AUSTRALIA by P. K. Latz* Summary Lata P. K. < 1975).—Notes on the relict palm Lhistona mariae F; Muell. in central Australia. Trans, R.Sac, S,Au&t. 99(4,), 189-195. 30 November, 1975. Comparison of photographs taken after intervals of up lo 56 years indicate thai the relict palm Uvistotia mariae may reach an age of up to 300 years. Aspects of the ecology of the palm are presented. Introduction distribution in central Australia. The majority Palm Valley, situated in the Finke Gorge of these rarer species are restricted to areas Ln National Park in the MacDonneli Range sys- the valley fed by permanent water seepage. a large field been dtv tem, Northern Territory (Fig. 1) is of con- Recently gas has siderable interest to tourists and scientists alike covered in the area adjacent to and north of largely because of the presence of a stand of the Park, Two gas bearing wells are situated the relict cabbage palm Livistona mariae F. only a few kilometres north of Palm Valley and wells for Muell. This species of Livistona is restricted tapping of these commercial produc- tion is being considered. W "«n area of about 60 km- on the Finke River and its tributaries. It is a relict species separated The Habitat by about 1000 km from the nearest Livistona The palm has a shallow fibrous root system to the north. The closest relative to L. mariae and is situated in an arid area. Therefore a occurs on the Fortescue River in the Hamcrsley suitable habitat for its continued survival must Ranges, W. Atist., a stand in many ways simi- have a permanent shallow water supply over lar lo that at Palm Valley. The relict nature of an area large enough to support a viable breed- the palms is discussed by Keast (1959), Bur- ing population The area must also be pro- bidgc (1960) and Chippendale (1963). tected from severe erosion forces or have a Although the explorer Ernest Giles was the stable soil environment, as the palm will not first to discover the palms in the Finke Gorge survive scouring of surrounding soil. in 1872, he almost certainly bypassed Palm PaJm Valley appears to be one of the very VaMey itself, and its discovery is attributed to few areas in the central ranges which meets ail a Lutheran missionary from Hermannsburg of these requirements. The permanent water Mission some time later. The valley was in- supply in the valley appears to be associated vestigated by members of the Horn expedition with the peculiar stratification of this area. The during 1S94 (Tate 1896). After a visit to the gently sloping conglomerate and sandstone Valley, Lothian (1959) discussed aspects of strata percolate water from a large catchment the palm's reproductive behaviour. area, down lo where the valley dissects these Chippendale (1959) listed 200 plants found strata. The position of this dissection ensUTes to occur in Palm Valley. Since this time a total a continuous permanent seepage area along a of 333 plant species have been recorded from considerable distance of the valley floor and the valley (about a quarter of the total number lower slopes. Although other springs and of species recorded for the whole of central seepage areas occur in the Ranges, few, if any, Australia!). About 10 percent of these species extend over such a continuous area. The Finke can be considered to be of rare or restricted River bed below Palm Valley has a permanent * Arid Zone Research Institute, Animal industry and Agricultural Branch, Department of Northern Aus- tralia, Alice Springs, N.T. 5750. 190 P. K. IATZ Fig. 1. The distribution and numbers of mature individuals of the cabbage palm Livistorm marwr in the Finke- Gorge National Park, Northern Territory. shallow water table for much of its length, but which apparently were absent in the earlier It is subject to severe water wash and the few photographs, taken in 1918, are now up to 12 palms found there are restricted to areas m high, a minimum mean annual growth rate which escape the full force of the Hoodwaters. of 22 cm. Palms grown from seed in Miami, Florida, Palm Growth Rales and Ages on the bank of a permanent stream, grew to a The author was fortunate in obtaining a height of 9 to 18 m in 27 years—an excep- number of accurately dated photographs of tional growth rate of up to 60 cm per year certain areas of the valley taken as early as (Lothian 1959). <\ palm in the cooler climate 1917. Exact relocation of many of the source of Melbourne Botanic Gardens is reputed to points was made possible by the cliff back- have grown to only 3.75 m in 30 yearsr a grounds of most of the photographs and the growth rate of 12.5 cm per year (Australasian fact that the narrow valley is restricted in the Post, 31 May, 1973). number of pholographie vantage points, In its natural habitat, the rate of growth of Of the 13 photograph sites relocated, three the palm appears to be mainly dependent on are presented in Figs 2-4. 'I he palms in Fig. the water supply. The palms showing the fastest 2 show the least change of any of the palms growth rate (Fig. 3) are at Palm Bend on The investigated and almost all of the individuals of the Finke River about 6 km east of present in 1917 (when the photograph was bank the Valley. This area has a greater depth of taken) are now still present. The present fertile alluvial soil and a better water supply heights of the living palms were estimated with than most areas in Palm Valley. Palms in Fig a Suunio clinometer. By using the cliff back- 4 arc situated on one of the best spring areas ground ns a scale, palm heights at the time of in the valley, whereas those in Fig. 2 occupy the earlier photograph were estimated. Distor- one of the drier areas of the valley, as is tion due to angle of viewing and to lens aberra- reflected by their slower growth rates. tion was estimated to be less than 10 percent. A mean annual growth rate of 10 cm was esti- The tallest palms in the valley have reached mated for this group of palms. a height of 25 m. Making allowances for dif- Palms in Fig. 3, how r ever, have grown ferent growth rates at various ages, 100 to 300 approximately Hi m in 38 years, a mean years seems a reasonable estimate of the age annual crowth rate of 30 cm. Palms in Fig. 4, of" the oldest individuals of Livistona mariae* THE RELICT PALM UVISTONA MARIAE 191 Hg, 2. The upper photograph was taken in 19 17 and the lower in 1973, Effect of Fire deep) were found at the bases of the majority The areas of high palm densities are sus- of the palms. ceptible to fire as large amounts of plant debris Ninety-six established palms were affected by accumulate around the base of the palms. A the fire, thirty of which subsequently died. To small area of the valley was subjected to a ascertain whether certain height classes of the natural fire in the early part of 1973 and the palms had higher survival rates, the heights area was visited by the author approximately of all affected palms were measured with a 2 months after the event. The fire appeared to Suunto clinometer. Results indicated that the have been intense; a palm 28 m high was burnt palms in the height range of 5-8 m had greater at the crown and thick ashes (up to 40 cm mortalities than those in the 1-3 m height 192 P. K. LATZ Fig. 3. The upper photograph was taken in 1935 and the lower in 1973. The white bed of the Finke River can be seen in the foreground. range. However, statistical tests showed no sig- through the stand in Little Palm Creek early nificant difference between the two height in 1959, but little evidence of this fire is now populations. apparent. A localised fire was reported in the same It does appear that many of the palms can area in the 1940's and this fire was severe survive a fire and subsequent regeneration is enough for the smoke to be seen from Her- quite rapid. However, an excessive accumula- mannsburg Mission, 12 km distant and on tion of debris over a long time period could the other side of the Ranges. After several sub- provide enough fuel for a more severe and sequent rains, regeneration of palms was extensive fire than those previously observed, reported to be abundant. (A. Latz. pers. and could effect a higher palm mortality. If this seed produc- comm.) A fire is also known to have occurred fire followed a few years of poor THE RELICT PALM UVISTONA MARIAE 193 Fig. 4. The upper photograph was taken in 1918 and the lower in 1973. This area was usl 7S.the large rock right of centre in the earlier photograph marker.mi?E tw as a This rock is hidden by palms in the later photograph tion, or if a severe flood subsequently removed individuals at all stages of growth were present the majority of seeds, palm densities could be in the valley. severely reduced. An exact count of all individuals in the Palm Numbers valley is impractical because of the high den- Tate (1896) stated, "... Livistona mariae sities of the palms in some areas and the many is known by only one colony estimated to com- younger palms at all stages of growth. How- prise not more than a hundred mature indi- ever, individuals of reproductive age (ca. 3 m viduals". A few were also observed by him or taller) were found to have at least some of along the Finke River as far as Boggy Hole. their trunk clear of leaf debris and are rela- Keast (1959) estimated a total of at least 300 tively easy to count. A count made early in I'M i». k i \\y washed away 1973 indicated that about 750 palms of repro- the rock fissures before they are is present in the ductive age occur in Palm Valley, or die of desiccation, there wide range of all growth stages. This is a much greater number than those populations a usually slow to ger- previously given. Several recent visitors to the Seeds of other palms are lose their viability unless valley were asked for their estimates of palm minate and quickly Leon 19581, but numbers and these ranged from 30 to 800 with kept in humid conditions (dc can germinate within an average of 150. These observations tend to seeds of the cabbage palm viable for two or more show thai rough estimations give a lower months and remain dry storage. number than is actually present and may years under account for the previous low estimates. How- Discussion ever, the early photographs, including those The cabbage palm is restricted to a small not presented in this article, do show lower area and its numbers are small. However, it palm densities than those at the present time. has a relatively fast growth rate, high pou-niial that the white basal ability Stirling i 1896) reported for reproduction, and a relatively rapid parts of the inner leaves of the young palms to recover after fire, all factors that indicate Thi& were eaten by aborigines in the area. the palm is in no real danger of extinction, pro- would have caused the death of at least some vided the present ecosystem is not substantially of the palms. This practice has now ceased In altered. In fact, numbers appear to have in- the past, aborigines frequently lit fires to hunt creased somewhat in the last 50 years. game, stimulate rcgrowth ol some edible plants The greatest danger to the continued health of and signal their presence to other members of the stand would arise ftotu the lowcrinr til (he tribe. Only two tires of restricted extent the water table or general disruption or pollu- are known to have occurred in Palm Valley tion of the seepage area. Fortunately the struc- during this century. Before this time the ture of the sediments probably ensures that this fre- aborigines may have burnt the area mote possibility is remote. The relict nature of this quently to stimulate growth of young palms unique palm warrants close attention to the which could then be used tor food. general health of the stand and prevention of Contrary to Tate's observations, there are any disturbances to the area. The planting of as many palms outside Palm Valley as in the palms m other areas of Australia to ensure largest valley itself fRg. D. The second that replacements arc available in case of iuiliuiv is in little Palm Creek where 550 natural disaster is recommended. counted. Except mature individuals have been Conclusion for about 6 palms further downstream in the Palm Valley is situated on the edge of a Finite River, the location of all individuals of targe gits iield. The valley is unique botanically. Lhhtotw mariav known to occur naturally is not only because of the presence of the palms, indicated in Fig. I. All ot the palms of repro- but also because of the high number of other ductive age have been counted and total 1 500 rare plants and ihe overall diversity ol the flora, Counts of all of the palms in small representa- A recluse area such as this, is by nature fragile, areas (including established seedlings) tive and therefore any development in this area show that the palms of reproductive age con- will require great care and foresight. stitute less than half of the population There- Acknowledgments fore the total number or established individuals the help in interpreting and of Livhrona mariae (the total naturally occur- I acknowledge photographs lent to me by my father ring world population) is estimated to be up- dating the indebted to G. Griffin, the wards of 3000. Arthur Latz. 1 am Valley, for his able assistance Alter a good season, each mature palm pro- ranger at Palm facts for this report, and to his duces an abundance VA seeds with a high in gathering Northern Territory Reserves natural rate of geimination. Although few of employers, the these seedlings arc able to get their roots into Board. References G. ( 1963 —The rehe nature of BuRHRirxih, N. T. (19601.—The r>hytoieo?.raphy Chippendale M J 7?- Centra) Australian plants Trans. R. Sc*\ ol the Australian region. Au.\t. J. Bat. 8. some 31-34 211. \. Aust. 86. o( Palm CHii'PtNDAi.t, O. M ( 1 9?V>—Plants 1958).—Viability of Palm Seedi Valley. Central Australia. Kto Nor, 75, 192- r>E LnoK *N. J. ( l >N 19V. /Vi>fc//^v2(3>.9h- — ) THE RELICT PALM LIVISTONA MARIAE 195 — Giles, E. (1875). "Geographical travels in Cen- Tate, R. (1896).—Botany. In W. B. Spencer tral Australia, from 1872-74." (McCarron (Ed.) "Report of the Horn Scientific Expedi- Bird and Co.: Melbourne.) tion to Central Australia Pt. Ill—Geology Keast, A. (1959).—Relic animals and plants of and Botany." (Melville, Mullen and Slade: the Macdonnell Ranges. A ust. Mus. Mae. Melbourne.) 13(3), 81-86. Stirling, E. C. (1896).—Anthropology. In W. B. Lothian, T. R. N. (1959).—Further notes con- Spencer (Ed.) "Report of the Horn Scientific cerning the Central Australian Cabbage Expedition to Central Australia Pt. IV—An- Palm Livistona mariae. Principes 3(2), 53- thropology." (Melville, Mullen and Slade: 63. Melbourne. GEOMORPHOLOGICAL EVOLUTION OF PART OF THE EASTERN MOUNT LOFTY RANGES, SOUTH AUSTRALIA byC. R. Twidale* and Jennifer A. Bourne* Summary TWIDALE, C R., & BOURNE, Jennifer A. (1975).-Geomorphological evolution of part of the eastern Mount Lofty Ranges, South Australia. Trans. R. Soc. S. Aust. 197-209, * 99(4), 30 November, 1975. Scattered relicts of a lateritic erosion surface of probable early Mesozoic age occur as the Whalley Surface high in the relief in the eastern Mount Lofty Ranges. However the greater part of the prominent summit surface of the uplands, the Tungkillo Surface, is an etch plain due to the stripping of the weathered bedrock and exposure of the irregular weathering front. The surface of the adjacent Murray Plains is of Pliocene age and was graded to a shallow estuary which then occupied the present lower Murray valley. It is cut across Miocene strata, though there are relicts of embayments eroded in Cambrian rocks where major rivers debouch from the uplands. It carries a veneer of ferruginous grit, Quaternary calcrete, dunes and alluvia. CEOMORPHOLOGICAL EVOLUTION OF PART OF THE EASTERN MOUNT LOFTY RANGES, SOUTH AUSTRALIA by C. R. Twidale* and Jennifer A. Bourne* Summary Twidalh, C. R., & Bourne:, Jennifer A. (1975).—Geomorphcilogical evolution of part of the eastern Mount Lofty Ranges, South Australia. Trans, R. Soc. S. AusL 99(4), 197-209, 30 November, 1975. Scattered relicts of a I a ten tic erosion surface of probable early Mesozoic age occur as (be Wnalley Surface high in t\\t relief in the eastern Mount Lofty Ranges. However the greater pari of the prominent summit surface of the uplands, the Tungkillo Surface, is an etch plain due to the stripping of the weathered bedrock and exposure of the irregular weathering front The surface of the adjacent Murray Plains is of Pliocene age and was graded to a shallow estuary which then occupied the present lower Murray valley. It is cut across Miocene strata, though there arc relicts of embayments eroded in Cambrian rocks where major rivers debouch from the uplands. It carries a veneer of ferruginous grit, Quaternary calcretc, dunes and alluvia. The scarps associated wilh the Milendella and Palmer faults are primarily of late Mcso- zoic-carJy Tertiary Age. It has been claimed that there has been 60-90 m of dislocation on the Milendella Fault since the Miocene, but this is questioned partly because the evidence is equivocal, and partly because in some areas the lower part of the scarp is erosional, being due to the exploitation by streams of intensely weathered rocks in the fault and scarpfoot zones in Quaternary times. as the riverine features of the Finnis valley (de Introduction Mooy 1959), the granite forms of the Palmer The Mount Lofty Ranges have received and Caloote areas (Twidale 1968, p. 95 et seq.; considerable attention from geomorphologists, 1971. pp. 5-44), and the Permian glacial but even in studies which are ostensibly con- features of the Strathalbyn area (Maud cerned with the uplands as a whole it is the 1972) have been investigated- Although there western areas that have been investigated in has been significant geological mapping of the detail (sec Benson 1911: Fcnncr 1930, 1931, region (Kleemun & White 1956; White & 5 1938; Sprigg 1946), There has been passing Thatcher 1957: Mills 1965 : Thomson 1969) reference to particular landforms which occur there has been no consideration of the evolu- in the eastern Mount Lofty Ranges (e.g. Fen- tion of the eastern Mount Lofty Ranges as a ner 1931. p, 23); there have been reconnais- whole, no attempt to correlate its development sanec studies of small parts of the area (Saies with that of the western Murray Basin, and nc 1 s 1968 ; Breucr I96S-'; Forrest 1969 ; Frahn investigation of the interplay of tectonism and 4 1971 ); and specific features or problems such land form development in the area. * Department of Geography, University of Adelaide, Adelaide. S. Aust. 5000. 1 Saies, Janet U968).- ''Geomorphology of the Marmum Falls Area". (Unpubl, B.A, Hons thesis, Univ. Adelaide.) - Breuer, Margaret (1968).—"The Geomorphotogy of Harrisons Creck"\ (Unpubl. B.A. Hons thesis, UnivT Adelaide.) * 4 Forrest, G. L (1969). —"Geomorphoiogical Evolution of the Bremer Valley" (Unpubl. BA. Hon* thesis, Univ. Adelaide.)— ' Frahn. D. (1971). "Geomorphology of the Milendella Area of South Australia*. (Unpubl. B.A. Honi thesis. Univ. Adelaide.) : — - Mills, K. J. (1965). "The Structural Petrology of an Atcr Hast of Springton, South Australia. (Un- publ, Ph.D. thesis, Univ. Adelaide.) w C. K TWTOALE &: )£NN1?£K A BOUKNF. 1--"-* jy- 5*be.?, . map. Fig. Physiographic reeioilS- of part of lhe eastern Mount Lofty Ranges, and (inset)- location The area nrouncf Rueljcns Hill and depicted in Fig. 5b is indicated. A-R ind CD are lhe tides of seciion shown in Fig- ^a- and X-Y lhe sec-lion represented in Tig. 7. Genera! Description rocks, perched rocks or loganstoncs (Fig. 3a) The area under consideration lies between together with lhe many clusters of boulders lhe River Marne in the north and Gorge Creek iorm a distinctive landform assemblage, The gneisses has been exploited by in the souih. Il falls naturally into three physio- foliation of the j^uphic regions; the high plain, lhe Murray weathering agents to produce pen item ( Acker- tombstone rocks paiivozolQ socinTientt Fig. 2. Generalised geological map of part of the Mount Lofty Ranges (after S. Aust. Cieo]. Surv, mop sheet SI/54-9, Adelaide). Gorge creeks. Their valleys are however sur- the Millcndella Bench, is greatly dissected by prisingly narrow in proportion to their depths, the present Milendella Creek and its tributaries, and in many places the streams run through but there are nevertheless broad flats at eleva- gorges or defiles. tions between 260 and 280 m, which are inter- The Escarpment preted as remnants of a once continuous flood plain. They stand some 80 m higher than the The eastern escarpment of the Mount Lofty plains to the east, and deseend by way of a Ranges is, like its western counterpart (Fig. steep slope to the Murray Plains (see Tepko 2). a complex feature. However in broad view 1 .50.000 topographic series (6728-111) M comprises two north-south trending scarps bftween grid lines 320 and 347 ? and 440 and which arc so gently arcuate that they can be 493; see also Fig 5a and b). The bluff leading regarded as linear. The two are offset with up to the summit surface from the perched respect to one another by a distance of some 3 bench is precipitous and arcuate in plan. km, the northern component extending as far Several features suggest that it is an abandoned south as Milendella. the other from the vicinity meander bluff: its shape, the presence of a of Raerjens Hill and south through Palmer central hill interpreted as a meander core with- (Fig.2). in the horseshoe-shaped valley which forms the The scarps are intensely dissected. The northerly extension of the bench, and the Milendella Scarp is greatly eroded, the zone presence in the valley floor of several metres of dissection being some 4-6 km wide. How- ot alluvium. It was associated with an in- ever, dissection behind the Palmer Scarp ex- trenched meander simiiai to those found on the tends for only some 2 km. The latter is also present Milendella Creek but of roughly 5—6 lower, being 100-145 m high compared to the times greater radius. The meander loop and Milendella which rises 250 m above the scarp core developed when the precursor of Milen- foot. della Creek flowed at a level some 80 m Various minor breaks of slope of probable higher than at present. structural origin can be discerned on the scarp. A second bench occurs where Harrisons In some places the upland scarp displays a basal Creek leaves the uplands to llow across the steepening and scarpfoot depression, as do Murray Plains, where it is known as Reedy some of the ridges within the upland (Fig. 4a). Creek (see Tepko 1 : 50.000 topographic In some few sites, prominent flats occur at series (67284II) around MR3I3374; see Fig. levels intermediate between the Tungkillo Sur- 4b). Less extensive than the Milcndelld Bench, face above and the Murray Plains below. One, it stands at an elevation of 150 m above sea 200 C. R. TWIDALE & JENNIFER A. BOURNE Fig. 3. a. Boulder cluster and loganstone or perched rock on the granite outcrop near Palmer. (C. R. Twidale.) b. Penitent rocks or monkstones developed on granitic gneiss near Tungkillo. (C. R. Twidale.) c. Low blocky outcrops of granitic gneiss east of Mount Pleasant. The even skyline is part of the Tungkillo Surface. (C. R. Twidale.) GEOMORPHOLOGICAL EVOLUTION OF THE EASTERN MT LOFTY RANGES 201 Fig. 4. a. ^Ijjcarpfoot depression developed at the base of a gneiss ridge at Raetjens Gap. (C. R. b. The Palmer Scarp west of Tepko, showing the Tungkillo Surface, the low scarp (a) the scarpfoot equivalent of the Palmer Bench (b). the break of slope on the hogback scarp (c) 0Ot Val ey (d) " P and ,he lower stce P« "»rp face (e). (C. R. Twidale) ft?e \v£ n e ! 6 (a> and Tl n i110 S,,rface (b) west of Mount - Pleasant. Outcrops of LgneissP ,T^countrynL f ! ^ the rock are seen in the foreground (c). (C. R. Twidale.) i 202 C R- TWIDALF & JENNIFER A. BOURNE » 3 .'JM run i. i"'.m'! KILOMETRES study area showing the Whalley and Tungkillo Fig. 5. a. Profiles (A-B and C-D on Fig. 1) across the benches. Drawn from S.A. surfaces, the east-facing scarps, and the Milendella and Palmer Lands Dept. 1:50,000 topographic sheet 6728-111, Tepko. Okm contour interval 20 metres i showing the old meander loop and the Fie 5 b Generalised topographic map of the Milendella Bench g ' iSSSoM?S^hm 339). Drawn from S.A. Lands Dept. 1:50,000 topographic sheet 6728-1 UTcpko. GfcOMORPHOLOGIOAI. EVOLUTION OF THE EASTERN MT LOFTY RANGES 2£l% Ifrfit, 50 m above the adjacent plains and 130 type, argued ihat several stages could be m below the Tungkillo Surface and in detail detected in its development: Includes several levels which represent stages 1. Initiation of The fault in the Palaeozoic with in downcutting. This Calmer Bench (so called the east side upthrown by many hundreds of because the site of the original Palmer town- metres. ship was only a short distance away) carries a 2. Early Tertiary erosion and peneplanation mantle of quartzkic blocks, some of which are with no faulting. iron-stained It has been dissected by Reedy 3. Renewal of faulting during the early Tertiary Creek which in this sector displays wclE- to Miocene with the east side subsiding by developed intrenched meanders. about 2fi0 m. Other, similar, perched benches have been 4. Miocene marine incursion with strata over- noted near the mouth of Pine Valley, south oF lapping the eroded fault-scarp. Tepko, and particularly west of Strathalhyn 5. Recurrent movement on the fault wince the where there is an extensive dissected bench, Miocene, the east side being lowered by 60- but as these arc outside the study area they are 90 in. not considered further. Sevetal aspects of this chronology can be challenged, and in particular the date and ex- Murray PUthts tent of faulting arc open to question. The proh- The western Murray Plains comprise a roll- Icm involves a consideration of the age and ing lowland which stands 200-150 m high character of erosion surfaces identified in Che near the scarp foot, to 70-50 m at the cliff top Mount Lofty Ranges and on the adjacent overlooking the River Murray. In several sec- Murray Plains. tors there is a scarpfoot depression up to 50 rtl deep fronting the Mount Lofty Ranges fFig. Age and Nature of mW Summit Surface I). The summit surface is a complex feature. Tin; These western Murray Plains are underlain iatcritic remnants nf the Whallcy Surface arc by Caino/oic strata 80 m or mare thick. In the crucial to the interpretation of the whole up- west. Quaternary fnnglomeTates predominate land. They arc sufficiently scattered to suggest hul these thin to the east where Miocene that they arc relicts of a once-contiguous marine slrata occur in bores, in Uibutaiy weathered surface of low relief, over which valleys, and in tfie high precipitous bluffs bor- lluwed streams carrying, a gravelly and dering the Murray trench (Fig. 2). Borelog bouldcry quatuose bcdload, Remnant deposits data indicates thai the Miocene strata rest on of these boulder beds occur in places, and a an irregular surface cut in acid and basic crys- discontinuous mantle of angular quartz frag- talline rnckv Basic crystalline rocks ale ments forms a veneer on Lhe Inw divides. The exposed at Black Hill, where Ihc so-called weaihering extended only some 10 m beneath 'norile' ha* been extensively quarried; similar the surface but it is notable that the base of rocks have been located in bores near Sedan significant weathering—the 'weathering front* and Walkers Flat. Granitic rocks and schists of Mabbuft (1961) —is everywhere coincides arc exposed extensively in the valley of Reedy with the local lever of the tungkillo Surface. Creek, but also in other many minor outcrops. underlain by intrinsically fresh rock. Thus it is reasonable to suggest that the Tungkillo Stu- face is an etch plain (Wayland 1934), formed Hmv did this landscape evolve? Of what age as a result of the stripping of the Interitic deep origin ,? and are the ntains What is the nature weathering profile and the consequent exposure nf ihc scarp which separates the high from the of the essentially fresh rock beneath (Figs 4c low plains? and 6). The penitent rocks, small castle koppies The lineai ity of these scarps and their and boulders typical of various areas of the association with faults exposed in Ihc gorges of Tungkillo Surface (Fig. 3) were- developed by River the Myrnc and Saunders Creek or in- differential weathering at the weathering frntil ferred from the displacement of strata, suggests and exposed as a result of the stripping of that the topographic forms are fault generated weathered material. and probably fault scarps, i.e. due directly and The age of the WhaMey Surface and of the wholly to tectonic displacement This is ccr- deep weathering is Meso/oic and probablv 1 laittJy the view of Mills who with resneci early Mesozoic (Triassic). The latente rem- lo the Milendclla Fault, which fe of reverse nants can be traced to ihc southern Mount F 204 C rt TWTDALE & JENNIFER A BOURN in minot outcrops throughout the western Mutray Basin, and they are commonly en- countered at shallow depths west of tbe River Murray. But there is one feature missing, namely buried laterile. though weathered Fig. jS. Diagrammatic section Through the eastern t.kaoltnised) Cambrian bedrock is recorded in Mount I ofty Range** showing the rela- tionship between the laleritiscd Whaliey bores beneaih the fresh crystallines and tbe Surface muferjalll by weathered bedrock overlying Cainozoic strata. There are also and the etch plain, coincident with the superficial ferruginous encrustations on high weathering front , uf the TungkLllo Sur- plain remnants and at higber elevations than face, the Miocene strata and therefore of lalei Cainozoic age, in the Murray Plains, for example just south of the Mannum Falls. Lofty Ranges where the laterite surface stands Further reference is made to Ihese occurrences high above valleys pnitially iilled by Miocene below, but no laterite profiles have been found marine sediments, and where the fault angle cither in outcrop or in bores. depressions resulting from the fault-dislocation There are a number ol' possible explanations, of the summit surface contain basal marine For some reason or reasons at present un- sedimeni of early Tertiary age. and with in- known, laterite may not bavc developed on clusions of laterite blocks I'Glacssner 1953; Ihose rocks of what was- to become the down- Glacssner & Wade 1958; Campana 1958). thrown block; this is rejected as being Thus the summit .surface and the laterite it irrational. Second, The laterite could have been carries must be of earliest Tertiary or late eroded before burial by Miocene and later However, evidence from Mesozoic age Kan- sediments, but then some remnants should garoo Island, just, across Backstairs Passage surely have been preserved. Third, the laicrtie Fleurieu Peninsula, suggests a much older from may indeed be preserved beneath the Cainozoic age. the laterite plateau formed on Cam- There cover, but has not yet been discovered. This is dissected brian and Penman strata was partially unlikely because many bores have been sunk before the extrusion of Middle Jurassic basalts anil though there is some reference to just west of Lhe present site of Kings- near and weathered crystallines, there is none lo laterite. suggest the eoie. Pataa »climatic consideration Fourth, tbe weathering ot the Cambrian rocks most likely period fur laterite Tiiassic as the could have taken place beneath the Cainozoic development (Daily, Twidale & Millies 1974). cover, but it is unlikely thai the oxidation i\^- Thus by short-distance correlation the rcsetired by the weatheted bedrock could have Whaliey Surface and the associated deep developed in such conditions. Fifth, the latenre weathering are considered to be of early Ivfeso- could have "been buried and iron oxide duiicrust aoic age. The development of the Tungkillc deslroyed by groundwater attack in the sub- Surface followed the disruption of the upland, surface, Such dissolution of iron oxide by water probably by faulting. Streams were rejuvenated charged with organic acids bas been shown to and they exploited the contrast in cohesiveness be possible (Bluomrield 1957*. Coulson, Davjs and resistance between Ibe rcgolilh and the & Lewis I9b0; Hingston 1963), fresh Tocks beneath, wirh the result that an This last explanation *eems to be the tnosi surface of low relief, the Tungkillo Sur etch likely. All other indications arc that the It i* still extending. face, was initialed. Whafley Surface wav dislocated by faulting, with the cast side subsiding by a matter of 260 Disruption nf (he Summit Surface m if] the north. The concentration of ground- scarp foot zone (see Ruxtun In view of the character of the Milendella waters in the 1958; Twidale 1962, 1%7) would account for and Palmer scarps it is obvious that the reason the ferruginous for the disruption of the summit surface could the subsurface elimination of have been a renewal of movement on tbe two crust, recognised faults, If this were so, rocks similar Thus if it is accepted that faulting occurred 1o those of the nearby upland should be found after the development of the lateritised sumnm beneath the Cainozoic strata of the western surface but prior to the Miocene marine trans- Murray Basin GTamtes and gneisses arc gD&slrJt], a numbei of geomorphological exposed widely in the valley of Reedy Creek, changes eau bo related to this period; GrOMORHHOLOOlCAL EVOLUTION 01- THE EASTFRN MT LOFTY RANGTS MS 1 Pie rejuvenated streams began the stripping Pleistocene age, being related to low glacial of the lafeWte regolith and the development stands of the sea. of the Tungkillu Surface. However, in Pliocene times the regional baie 2. The rejuvenated streams incised into the nn- level on the western Murray Plains stood 50- weaihered bedrock creating quite deep, nar 70 m higher than it does now, and it is to this row, and in places meandering gorges, higher ba.selevel that the streams responsible for the erosion 3. As they emerged from (he uplands, lhe of the plains were graded. These streams transported streams eroded quite broad ewhayments in to the then Murray estuary the crystalline basement rocks of U>e uplands the detritus which in part comprises The Norwest contiguous widi the then surface of the Bend Formation. Thus the Murray Surface Js .Murray Plains. The Milendella Bench stands in the study area of Pliocene age. There is considerably higher than either the Palmer some corroboration from the thin ferruginous Bench or bench remnants at the mouths ol duricrusts found on some parts of the surface, as for instance the Saunders and Marne gttt^cji. This may just south and south-west reflect its development at a stage before of Mannum Fatls. for surfaces of comparable faulting hud ceased, and its grading to the age on northern Yorke Peninsula (Hbrwftz Daily Miocene shore, or if may indicate toca* late & 19581 and Eyre Peninsula Cainuzoic faulting, (Twidale, Bourne & Smith 1976) also carry & ferruginous encrustation, as indeed do posl- When did faulting begin? There is. no direct Miocene valley floors in the southern evidence, but the stratigraphy of the Murray Mount Lofty Ranges (Horwiu 196CM, and Bnwn vO'Driscoll I960; l.udbrook 1969) sug- the Karoonda Surface of the central gests that part of (he region became a jeMrictcd Murray Plain* (Firman 1973, p. 23). marine emhayrnent during the early Crcla- lhe occurrence ol iron staining, eeuus. The next proven marine transgression as well as their elevation, suggest took place during the Late Eocene ami marine that lhe several perched benches found at the mouths major influence continued through to the Middle Mio- of upland gorge* were also formed at this time, even cene. However, though strata of Late Creta- though they are now separated from ceous age have not been located, they may the plains by scarpfoot valleys up to 50 deep. have been deposited and .subsequently been m This is corroborated by profiles relating the benches to removed by erosion (l.udbrook 1969. p. 159). the main plains surface (Fig. 5a). The fail It dislocation was probably gradual and Although little is known of the geometric possibly occupied the whole of this late Mesu- rclationship zoic-middle Tertiary period between river discharge, channel width md the radius of intrenched meanders Age of tlie Plains Surface tcf. Bales 1939 and Dury 1954, 1964; with respect to flood plain meanders, The broadly /oiling plains surface of the also Gey I 196$) tf>e abandoned meander loop preserved western Murray Basin is bfkhg dissected by the o*i the Milendella Bench few through-flowing streams such as the Marnc suggests rivers of much greater I? occasional t discharge Milendella and the Harrisons-Reedy systems, than those active at present. which emerge Horn the Mount Lofty Ranges Thus, during the Pliocene, |hc scarp foul and persist to rcaef> the Murray. The River stood 80 m higher than present at the Murray trench is a Quaternary feature, though mouth of Milendella Creek and 50 above there wa-s an earlier shallow estuary coincident m present Ivucleve! where Harrisons Creek with the present Murray valley as tar north as debouches oi: to the plain. Is the post Pliocene scarp tectonic Morgan. In this long narrow and shallow estu- or erosional, or are there ary, marine sediments, the Norwesl Bend For- element oi" both origins present? mation, of Late Pliocene age were deposited (l.udbrook 19931}, Remnants can be seen The Question of Post I ate Cainozoie Faulting perched high on the modern cliff-tops in many The lower part of the Milendella, and by places, occupying a shallow valley cut in the implication the Palmer, fault scarps are seen Miocene strata. As the trench now occupied by by Mills (1965, pp £36-442)* as due to post the river has been incised below these Pliocene Miocene faulting. He discovered several out stiata, it must be younger and there is much liers of Miocene sediments perched on the general .i-gumem as well 473 metres perched Miocet<« Inoostone 162 _ QuarernaTyalluviiim upper Miocene sand Miocene* limestone 76 . Eocene lipniticsands 40 Sea levei Ckm \ Creek, I After Mills 1965) fig. 7 Section through the Milcndella Scarp near Suundm (pers. comm.) and Milcndella areas, and took the eievational tic, Mr. J. M. Lindsay reported Ci recrystailised quartzosc difference between the two occurrences (i.e. in 1975: A sample of from a perched oulliet 60 and 90 m) as a measure of post Miocene fine-grained limestone, north of Sander- dislocation on the faults (Fig. 7). He also noted of mid-Tertiary rocks 3.5 km small benthonic forammifcra that boulders in ihe Pleistocene faoglomcrates ston> contained lithological abutting the fault plane display orientation sug- bur no diagnostic forms and no distinguish between Mor- gestive of recent movement along the plane. features which would This suggestion of recent and continuing dis- gan Limestone and Mannum Formation." As stratigraphic position of the Miocene en- location is borne out by the seismic record the also not been deter- (Sutton & White 1968) which shows that the countered in bores has it is not possible to identify eastern border of the Mount Lofty Ranges is mined to this date, outliers or the strata intersected in subject to earth tremors. The distinctness of either the precision. Thus cufre- the escarpments both here and to the south bores with certainty and It can be argued that the suggests geological youthi'ulness (see e.g. .leune lation is unwarranted. topographically higher outliers are Morgan Si Chittleborough J974). The higher level of left behind as remnants of cireum- ihe Milcndella Bench (see earlier) can also be rormatioo after the removal by solution and interpreted as indicating local late Cainozoie denudation the bulk the Formation dur- dislocation. fluvial action of of and Quaternary, that the Mio- On the other hand, the scarps have been ing the Pliocene the bores is Mannum For- deeply and intricately dissected, the Milendetla cene encountered in in situ, and thus that no post~Mit»ccne scarp in particular being greatly cut about yet mation of any significance is implied by the retaining its overall linear morphology. More dislocation 8). This crosional hypothesis significantly, the evidence cited by Mills is evidence (Fig, support from the character of the equivocal and is equally well accounted for in receives scarps. the scarps other terms. Milcndella and Palmer If then the scarps The Miocene of the western Murray Basin, were wholly of tectonic origin, elevation or comprises two major formations: the strati- ought everywhere to be of similar systematically and gradually graphically lower or older Mannum Formation at least to vary differences the throw of the and the higher and younger Morgan Formation according to in fact (Ludbrook 1969). The two are very difficult to fault dislocations. But the scarps in according to their location with distinguish in the field and can only be dilTeren vary in height streams. Where there are riated palaeontologieally. The faunas of the respect to scarpfoot the scarp foot perched outliers are unfortunately not diagnos- streams either heading back to UEOMORPHOLOGICAL EVOLUTION OF THE EASTERN MT LOFTY RANGES 207 tion is implied^ This is not to suggest that there has been no recent movement on the fault: on a fatting the contrary, the seismic evidence and the dis- impMSd turbance of pebbles in the fanglomerates which abut the fault plane point to recent and con- tinuing dislocation, What is argued here is that the lower, steeper part of the fault scarp is Morgan principally erosionai and that tectonic disloca- b no posl Miocene faulting implied tion has been a relatively minor factor in its evolution. There remains the problem of the contrasted Fig. 8. Two possible modes of evolution for the eastern scarp of the Mount Lofty Ranges. state of dissection displayed by the Milendclla and Palmer scarps. It is not a matter of con- trast in or debouching from the hills on to the plains, rock type or volume of run-off, for Hie lithological units of the the scarp is higher, there is a faceted scarp with eastern Ranges run the lower unit steeper than the higher, and north-south and are essentially similar behind both scarps. It cannot there is a distinct scarpfoot valley running be a question of climatic differences. It may be due to the Milendella parallel to the escarpment (see Figs. 1, 4b and and 5a), frequently with intensely weathered rock Marne draining a higher scarp, and eroding regressivcly exposed in the valley floors. In between these more quickly to capture the head- waters sectors arc others where (here are no scarpfoot of the Torrens drainage. More likely however, the dissection streams, where the escarpment is of lesser of the southern part ot the study area amplitude and where the hill-plain junction is has been retarded by the base- level control higher and coincident in elevation with the exerted on the Harrisons-Reedy break of slope on the higher scarps nearby. creek system by the large mass of crystallines This suggests that die upper part of the scarp exposed west of Caloote. The lower elevation of the is of fault or tectonic origin, though of con- Palmer as compared to the Milendella scarp siderable antiquity (late Mesozoic or early Ter- may be a contributary factor. tiary), but that the lower part, where present, is due to erosion by streams eroding the Conclusion weathered strata of the scarpfoot zone (Twi- The development of the eastern Mount Lofty i.e. is dalc 1967): that the lower scarp of fault- Ranges and the parts of the Murray Plains is line type and that no significant recent disloca- summarised below. Cambrian Deposition in Adelaide Geosyncline Early Palaeozoic Orogenesis, granite emplacement and mctamor- phism l-ale Palaeozoic Permian glacialion (not evidenced in study area) Early Mesozoic Planation and deep weathering. Marine deposi- Whalley Surface tion in lower part of Murray Basin in early Cretaceous (Thornton 1974) Late Mesozoic- Faulting, eastern block down. Marine sedimen- Tungkillo Surface. Middle Tertiary tation Eocene- Miocene, Stream rejuvenation Scarp formed. Gorges and stripping of regolith developed Pliocene Essential withdrawal of sea, only estuarine sedi- Perched benches and mentation. Planation of Miocene sediments and Murray Surface of emhayments in crystalling rocks on face of scarp Quaternary Lowering of sea level. Murray Trench formed, Scarp foot valleys, tributaries rejuvenated, valleys and scarpfoot abandonment of scarp zone excavated benches . 20K C. R. IW1DALH fr JHNNIFKK A BOWKNk Acknowledgment* Kiugsley Mills, Department of Geology and rhe writers wish id thank Dr Brian Daily, Geophysics, University of Sydney, for helpful licology Department, University or Adelaide, discussion and continuing interest in ihe evolu- lor a critical reading or" the paper, and Dr tion of the area under investigation. References Ackhrmann, E. (1962).— Biisscrsteinc-Zjcugen var- Horwitz. R. C. ( 1960).—Geologic de la region zeitlicher Grundwassetscb.Wank\irjgen. '/.. Geo- de Mt Compass (feuille M'huig), Anslrulte morph. (N.S.) St. Vincent Basin in M. F. GlaessneT & 1 . W. TwtDAifc. C. R, (1962).—Steepened margins of Parkin teds). "The Geology of South Aus- inselbergs from northwestern F.yie Peninsula tralia", pp 115-126. (Meib.U.P.) Soudi Australia Z. Gcomorptr (N.S.) 6. 51- Hinumun, F. J. (1963),—Activity of polypheno- 69. ls constituents of leaves of Etu:alvptus and 'IwiDALt, C K. (1967).—Origin of the piedmont other spevtes in completing ;md dissolving angle as evidenced in South Australia. J- iron oxide. Auxt. /. Soil. Rr.w HI), 63-73 OVaf, 75,393-411. — GEOMORPHOLOGICAL EVOLUTION OF THE EASTERN MT LOFTY RANGES 209 Twidale, C. R. (1968). Geomorphology with von der Borch, C. C. (1968).—Southern Aus- special reference to Australia. (Nelson: Mel- tralian submarine canyons: their distribution bourne.) and ages. Mar. Geol. 6, 267-279. Twidale, C. R. (1971).—Structural Landforms. Wayland, E. J. (1934).—Peneplains and some (A.N.U. Press: Canberra.) erosional platforms. Geol. Surv. Uganda. Ann Twidale, C. R., Bourne, J. A., & Smith, D. M. Rept. Bull, 77-79. (1976).—Age and origin of palaeosurfaces on White, A. J. R., & Thatcher, D. (1957).—Man- Eyre Peninsula and the southern Gawler num Geological Atlas 1 mile series, No. 811 Ranges. Z. Geomorph. (in press). Zone 6. (S. Aust. geol. Surv.: Adelaide.) THE WOODENDINNA DOLOMITE AND WIRRAPOWIE LIMESTONE - TWO NEW LOWER CAMBRIAN FORMATIONS, FLINDERS RANGES, SOUTH AUSTRALIA BY P. G. HASLETT* Summary HASLETT, P. G. (1975) .-The Woodendinna Dolomite and Wirrapowie Limestone-two new Lower Cambrian formations, Flinders Ranges, South Australia. Trans. R. Soc. S. Aust 99(4), 211-219, 30 November, 1975. Two new formations, the Woodendinna Dolomite and the Wirrapowie Limestone, are proposed for two distinctive units of the Lower Cambrian carbonate sequences of the Flinders Ranges, which have not, hitherto, been differentiated from the presently defined formations. Both formations are widespread in their occurrence, but are best developed in the eastern Flinders Ranges, north of the line of the Blinman, Wirrealpa and Frome Diapirs. The Woodendinna Dolomite and Wirrapowie Limestone record a significant period of supratidal and shallow, sheltered, intertidal deposition respectively within the Cambrian of the Flinders Ranges. THE WOODENDINNA DOLOMITE AND WIRRAPOW1E LIMESTONE — TWO NEW LOWER CAMBRIAN FORMATIONS* FLINDERS RANGES, SOUTH AUSTRALIA by P. G. Haslett* Summary HytSLfixr, P- G. (1975).—The Woodendinna Dolomite and Witrapowie Limestooe—two new Lower Cumbrian formations. Flinders Ranges, South Australia. Trans. /?. $ Introduction the WUkawiliina Limestone, the Parara Lime- Formations within the Hawker Group (Dal- stone and the Ajax Limestone (Table 1), several garno 1964) of the Lower Cambrian of the distinctive lithofacies exist. Not only do the Adelaide Geosyncline characteristically show present formations contain separate and InJi- frequent changes in both thickness and facies, vidually mappable Ethologies of significantly This results in complex intertonguing of litho- different depositions! origin but in the past the logical units. The Ten Mile Creek type section formation names have in some instances been (Daily 1956), measured within a syndeposi- loosely applied (Dalgarno 1964, pp. 134-1 35"L tional graben structure northeast of the Ora- table i parinna Diaplr, represents one of the thickest F.xixrinz jotmatinn names ami their sources for the Lower carbonate sequences Flinders Rang*;* ««.j «*»*•*. .,~~,.,i .*~ *. A„ «.««„.. «£ *u~ xj ...1,^ Cambrian of the and most complete sequences or the Hawker : : — in the Flinders Ranges. CE NI> Group Mapping by the western ea1stern S.A. Department of Mines (Dalgarno & lake torrens flinders flinders i,i~ laii, r-w* i tanri\ „a u. t» AREA RANGES RANGES Johnson 1966; Coats et al. 1973) and by B. Daily and a number of his students (Gehling1 ; limesi'ONE 1 :i Haslett-: Hatcher : Mi>unl ; Hull"; Daily andamooka. ajax (Duit> iSSfi) . -,- Irt-fc-, , ... It. c LTMESTONE. LIMESTONE |y/2a) has resulted m the recognition of (Johns 196S) (Daily 1956) wilkawillina several significant litbological sequences not ^e^JwTSgF present within the Ten Mile Creek type section. The thick basal Cambrian carbonates in par- Whilst it is extremely impractical and un- lietilar exhibit complex and varied facies inter- desirable to introduce an excessive amount of relations and it has become obvious that within formal stratigraphic terminology for the Lower 4 Department of Geology and Mineralogy, University of Adelaide, Adelaide. S. Aust. 5000. 1 defiling, J G. (1971). —The Geology of the Reaphook Hill Area. Honours Thesis, Univ. Adelaide (unpublished). " Haslett, P. G. (1969).—The Cambrian Geology North of the Wirrealpa Dinpir, Flinders Ranges, South Australia. Honours Thesis, Univ. Adelaide ("unpublished), s Mount. T. J. (1970).—Geology of the Mt Chambers Gorge Region. Honours Thesis, Univ. Adelaide (unpublished). 5 Hatcher, M. I. (1970*),—The Geology of the Mt Chambers Mine Region, Northern Flinders Ranees. S. V Honours Thesis. Univ. Adelaide (unpublished). ^Htdl, K. G. (1973).—The Lower Cambrian, Puttapa Synclinc, FlfudetK Ranges. S.A. Honours Thesis. Univ. Adelaide (unpublished). 212 ?. O. HASTKTT sufficient nomenclature be available Ic map and above the Pound Quan/ite W\ beneath the red describe, on a regional scale, the major ruck elastics of the Billy Creek Formation in the relationships preseni within the depositions Mt Scort Range area. Thrs comparatively thin basin, Careful recognition of mappable units carbonate sequence incorporates a number of lithological types. Daily (1972b) has fil uniform or repciitive lithulogical nature different should facilitate rather than cucumber further subsequently distinguished a 1 ower and Upper work, Such accurate lithologieal subdivision, Member, and In view of the thickness of this coupled with thorough palaeontologieal unil, any further subdivision of the Aja.x Lime- evaluation, will result in a more lucid inter- stone may prove unpractical on k regional pretation of the deposiiional history within the scale. It is very imponanc to realize, however, lithotypes present in the ba&in. Jt is with this intention, that the new that a large number ol formation names Woodendinna Dolomite and Ajax Limestone are present within the Cam- Wirrapowie LimeMone are proposed. brian carbonate formations mapped elsewhere in the Flinders Ranges (Daily, pers. eomm.) Present Nomenclature of the Lower Cambrian and that the change in stratigraphic ter- Carbonate Sequence in the Flinders Ranges minology in the Mt Scott Range area is in large Wilkawillina Limestone part historical rather than geological. The Wilkawillina limestone, in its Ten Mile Proposed New Formations Creek type section, discotiformabiy overlies the Bonncv Samisione Member of the Pound A significant propoition of the basal Cam- those hi Quartzitc, and ii extensively dolomitizcd ai the brian carbonate sequences, particularly 1>3SC In general the unit is massive, pure and the north-east Flinders Ranges, are not readily pale cream to pink coloured. II consists pre- assignable to any of the above Mrali graphic dominantly of ooid and skeletal grainstnnes. units. Although mapped c>n the COPLEY which may he patchily silicified and lecrystal- 1:250.000 and parts of the PARACHILNA Wilkawillina Jurd. An abundant fauna js characteristic of 1 :250,tl00 geological maps as the upper parts, some beds consisting almost Limestone, the carbonates show only scant as defined entirely ot broken skeletal remains of arcbeo- similarity to Wilkawillina Limestone eyathids. and associated braeh'topods. hyo- in the type section. In part, these carhnnates the Parara liihids. and trilobites. Algal Temains arc show greater similarity with Lime- common throughout and although some stone of the type section, hut crucial differences cryplalgalaminatcs and oncolitc horizons are in Ikhology and depositionai environment are present at the very base, stromatolites arc vir- apparent on careful examination. It is for these basal Cambrian tually ahsem. Suhaerial desiccation features arc distinctive and widespread Dolo- not represented in the type section Daily units that the new names WtwlenJittna proposed. (1956, 1972o) considers the Wilkawillina Lime- mite and Wirrapowie Limestone are stone 1o have hecu deposited on a stable shelf Type Section conditions. under shallow marine The type section is located in Witrapowic t'antra Limestone Creek and one of its tributaries, about 14 km Flaggy to tubbly dark to black impure lime- due east of Point Well Station homestead in Hie stones with intcrbedded dark calcareous shales Central Flinders Ranges (Fig. 1). Unfortunately form the lower Member of the Parara Lime- the section presents some difficulties with Upper of stone m i he Wilkawillina graben. The respect to access, which involves \S km Member consists of uaggy and thin bedded travel by four-wheel drive vehicle from a point dark jrrey to black aphanitic limestones and on the Point Well-Narrina road to Wooden (bin calcareous .shales. Although scantily din mi Bore, and a further 2 km on foot alon.a Ussilifcrous throughout, the Parana Limestone a creek to the section. 'Ibis disadvantage is tar contains trilobites, hyolufiids, brachiopods, outweighed by the completeness of the sponge spicules, evnehostracans and rare sequence and the good exposure of both the uichcocyathids (Daily 1 956). Stromatolites, Upper and lower boundaries of both forma- ITarpebblc conglomerates and desiccation tions, features are all completely absent. The combined Woodcndinna Dolomite find AjdX Limestone Wirrapowie Limestone in the type section con- carbonate rocks which over- Daily (J 95b) proposed that the term Ajax sist of 392 m of Ltmcstont- should be used 'or all carbonates lie, with apparent amformtty, Diplorratcrion LOWER CAMBRIAN FORMATIONS, FLINDERS RANGES 213 WOODENDINNA DOLOMITE MAP A WIRRAPOWlE LIMESTONE TYPE SECTION LOCATION 31 00 — PARARA LIMLSTONE WIRRAPOWIE LIMESTONk WOODENDINNA DOLOMITE HARACHILNA FORMATION POUND OUARTZlTL All facks shown are passable to fouf-wheel drive vehicles onlj SOUTH AUSTRALIA MAP B \ 1 A — 31 00 £ MAP B J'h\ ^ { ) \ i i i —I ! \ PGM 75 Fig. 1. Location of type section. . 214 P. G. HASLETT TYPE SECTION woodendinna dolomite and wirrapowie limestone Very r«fti,fiiiy indued, .5tratiq"ly EiuttrappHg finC grtlifcd ir-tei-beJs . PARARA "ill iter tori as ^Ufi 1 itjnler coloured inalcy l 'jparsaly fossil i fet ous and lacking dcsT^catiOi foafUfM i tldt-gelljlt: cciitjloniei aiOE,. LIMESTONE I and 392in -r-fWv-w^ ^^Xk '*^VVyyWv> Tina.- ijraincd riffij t« tii*c1v ?iity 1 iivestones .-ind 300m LdTrfi^fftaua siit'jiu.iifMi, w.vrti very vtstjit far 3-'ij wr ii't'tdi"^ riapk fiiifiy lamim*tPi} a]qnl Uruno.Uj.l Tie ! di' LUtinloiuei'titfs .ire £ed$ (5) 4ty I PP1T1 ru tld.-btfbble cairj-ior,. I'li-io'r- cross-aedded utk ^nnstone beds (.3) PBJ.W* l.iii-oughnul. Sui iwn-i; ai'teofititti -letri tus is WIRRAPOWIE j w—uriil," dbsi-nL m the type section. Tire upperiros-t LIMESTONE m jmiii-v -^ this ufi~ is spJicwfeS tjradati'crtal in*& Eli* r iiftt3 - at Uw tawicst: al' i*i i-'ARMA LIMESTONE Btyecd : stftuM-JftHM bed. 200rn Tellcw-We£^icrinc|, flatly to n,i>.sivn 41q1ut,;-qs cress- 4fi; loTonitie l iiaestotrES'i Wf? ujiiniiurw tloiSni bedded qaafM SriivisMirti-. j;,), in!."j1 ut nntional conglomerate and ile^c^uLiail iflJilrtWfcS-. Auundan: 1'imafituncs •jtrouia*Ql1te beds .( = ), 0C]it.k (VI o:uir sda< ttw Up. 100m MdSbive outuiiflpiHT] dull (pey dulaml L'7t."d 1 fl>itfii±EH*&. U^ally cry^UHine! prm..vy n«r>o-si ticrial ;eKti*T5 hw*«j WOOOEN0INNA difficult ta toUmnr.e nr tiic liieid. jHj4ra1 I1n«it«tifv cm'.:.-.- bedded air.d uralruunes wure u'^ticmtr>-av>t I _y DOLOMITE juiitiittn-mi mattered qujrcz s^rd ^r^nii (foSiiVf to! cuwui'- I a. Litr'ei "ly*'l inf.ed stroma T t&s are Idi-IIMi.: HlSSiVtt LU fl-l'liv yd, low-WEMtH(>rn".g JulOI'ilitflS 3-nd rffi limr^trn as. Tit WS^ h4 Ww 'aIKMITWIE UKC STONE tS v, Lire itnrievtying pARAClltLflA rtp-jd^nily confonr^hla i Lh HHMAilUN and is plat«»t: it tht* v-cla !.i vclj' dbnpt Kf&nttWtfiiiB Fig. 2. Type section. LOWER CAMBRIAN FORMATIONS, FLINDERS RANGES 215 ROCK RELATION DIAGRAM Woodendinna Dolomite and Wirrapowie Limestone (Not to scale ) Wirrealpa WEST CAMBRIAN Mt Scott ehe Range Nepabunna Sittstone €hd Midwerta Shafe Ghr Parara Limestone Ghw Wilkawillina Limestone Ch, Ajax Limestone Ghp Parachilna Formation Gu Uratanna Formation WOODENDINNA DOLOMITE PROTEROZOIC Ewp Pound Quartzite Disconformity Fig. 3. Rock relation diagram. sandstones and shales of the thinly developed sandstones. Intraformational conglomerates, Parachilna Formation. The entire sequence often with dolomite clasts in a quartz sandstone dips 25° to the SW, on the western limb of a or ooid grainstone matrix, are also common large, slightly southward plunging anticline. In (Fig. 5). the type section, the Woodendinna Dolomite Between 69 m and 119 from the base of conformably underlies the characteristically m the Woodendinna Dolomite in the type flaggy grey limestones of the Wirrapowie Lime- section, all carbonates have been extensively stone, the formations being 176 m and 216 m dolo- mitized. This results in the development of a thick respectively (Fig. 2). massive dull grey unit which appears, super- Lithological Characteristics ficially at least, to be lithologically homogen- Woodendina Dolomite eous, but which has pre-diagenetic lithologies comparable to The Woodendinna Dolomite in the type sec- those directly below and above, but with tion is apparently conformable upon the Para- a greater number of ooid grainstone beds. Some of the chilna Formation, and its base is selected at the grainstones are rich in dis- abrupt appearance of carbonate rocks above persed quartz sand. Thick, dolomitized algal stromatolite the shales and Diplocraterion sandstones of the beds and intraformational con- Parachilna Formation. The basal carbonates glomerates are also common. are partly or completely dolomitized oolitic and Above this dolomitized zone to the top of pisolitic limestones, interbedded with very the Woodendinna Dolomite the sequence con- minor green shales. Massive to flaggy yellow sists of interbedded flaggy yellow dolomites, weathering dolomites with pronounced desicca- ooid grainstones, stromatolites, flat pebble con- tion features are abundant just above the base, glomerates and quartz sandstones, all with in association with low, broadly domed stro- beautifully preserved primary sedimentary matolites (Fig. 4) and, beginning about 36 m structures. Desiccation mudcracks, some up to from the base, thin cross-bedded, pure quartz 8 cm deep, are common in the dolomitic mud- 216 P. G. HASLETT .- :: 7"! 1 ft.; *^Miiln iiint^Plf** '^*^to Figs 4-11, LOWER CAMBRMN FORMATIONS. FLINDERS RANGES 2i: Stones (Fig, 6), Ripple maTks and chevron-type rnudstones of the Woodeodmna Dolomite, and cross bedding are pieseni in (he ooid grain- minor trails and apparent btotuibarjcm of bed- stones and quartz sandstones (Figs $ and 9), At ding which may be found in impure limestones 135 rn from the base, a thick cross-bedded of the Wirrapowie Limestone. quartz sandstone With abundant creamy yeJIow Dwribution dolornitic mudsione cla^ts is developed. Toward The Woodendinna the top of the Woodendinna Doiomite> ooid Dolomite and the Wirra- grainstone beds commonly show a primary powie Limestone form the basal Cambrian car- bonate succession wbbly or cobbled upper surface, reminiscent of over much of the Flinder.v Ranges. that found in modern environments on area< They are best developed in the eastern part the ranges, of stihaemdly exposed, partially lithificd car- of north of the line of outcrop bonate sands (Fig, 7). of the Blinman, Wirrcolpa and Chambers db- pirx (sec Fig. 3). In this region their combined Winapowie Limestone thickness ranges from 300 m to 500 m and they The hase of die Wirrapowie Limestone coin- appear to thin gradually to the wesi, where cides with the relatively abrupt disappearance comparable Jithologics are thinly developed in of dolomite in the sequence Flaggy fine- iht AJax Limestone at Mt Scott fDaily, pers, grained, dark grey, limestone and green grey comm.) and the Andamooka Limestone in the calcareous ailtstones dominate the sequence. vicinity of Yarrawurta Cliff (Wopfner 1 969; Desiccation features arc less ahundant but stro- Daily 1972a). To the south, along the above- matolites, intraformational conglomerates and mentioned line of diapirs. the Wirrapowie minor cross-bedded ooid grainstones arc Limestone thins very abruptly and is overlain pieseni throughout (Fig. 11). Quartz sandstones by, and intcrlingers with, Wilkawillina Lime- are virtually absent, As in the Woodendinna stone. Syn- and post-deposttional tectonism Dolomite, stromatolites arc finely laminated. along this line of diapirs. and the consequent low, laterally linked domes which, on bedding development of complex local facies variations, plane exposures, show little if any preferred has complicated the regional facies relation- orientation fFig. 10). ships along ihi.s line. Lateral intcrfingering of The Wirrapowie Limestone in the type sec- the Wilkawillina Limestone and the Wirrapowie tion is overlain by Parara-type limestones. The Limestone- h strikingly demonstrated, however, topmost limit of the Wirrapowie Limestone is in outcrops just north of the Wineatpa Diapir selected at the top of the uppermost algal (Haslctt, in press). Abrupt lateral change south- stromatolite bed. Although a superficial simi- ward from Wirrapowie Limestone may also be larity with the Wirrapowie Limestone remains seen in tbe vicinity of Fregunda Creek. Iu addi- above this limit, the sequence lacks ooid grain- lion, Hatcher* indicates a similar southward stones, flat-pebble conglomerates and evidence thinning of a wedge of Wirrapowie-type lime- of subacrial desiccation. The dark limestone* stones within the northern part of an area ol are far more strongly outcropping, purer and thickly developed Wilkawillina Limestone near crystalline more above the contact. They also M t Frome. tend to he sparsely fossil if with erous. archeo- In tbe north-eastern Flinders Ranges, the cyathids, hyolithids and various hraehiopods Wirrapowie Limestone is in all cases overlain present. This fa in contrast to the underlying by Pararn-type limestones. To tbe best, of the Wirrapowie Limestone and Woodendinna author's knowledge, Wilkawillina Limestone is Dolomite which are notably depleted in fossils. absent from the north-eastern Flinders. Those Apart from alga! stromatolites, tbe only evi- rocks which are mapped as Wilkawillina Lime- dence organic of activity found to date are very stone in this area on tbe COPLEY 1:250.000 minor irregular hurrows in some dolomitic Geological Map are assignable to the Wood- Fig, 4. Bedding plane exposure of stromatolites, Woodendinna Dolomite. ' oI n ,,e c,ast £lg *' m inlraformational conglomerale. Woodendinna Dolomite, 7 i? ? i ? hig. Well2 6. developed rmidcracks on the underside of a bedding plane, Woodendinna Dolomite. Ng. 7, Cobbled upper bedding plane surface of ooid erarnstone, Woodendinna Dolomite, hig. 8. Chevron-type cross beds in quartz sandstones, Woodendinna Dolomite. Fig. 9, Ripple marks cm bedding surface of a thin quartz sandstone, Woodendinna Dolomite. Mg. 0. Squat stromatolite bioherms on bedding plane, Wirrapowie Limestone. frig. It Impure flaggy-bedded limestones of the Wirrapowie Limestone. Note the lenticular agereealcs of uitra-formatKMial conglomerate. 2IK P. O. HASIJtTT cndinna Dolomite and Wirrapowie Limestone of Brachina Creek to south of Mt Aleck, and in Ibi! lower pari and obviously hitherto undif- Dalgarno (1964) records similar rock types in ferentiated, and in the upper part can be much the vicinity of the Chase Range- More recent more justifiably related to Parara Limestone work ha* confirmed the presence of both Itthologies than those of the Wilkawillina LiuKV Woodendinna- and Wirrapowie-type carbonate stone. sequences in these southern and south-western Flinders (Daiiy, pers. Thin but well developed Woodendinna Dolo- parrs of the Ranges mite does oceiir south of the line of the Klin- comm.). man-Wirrcalpa-Chamhers diapirs along the Depnsitinnal F.nvironraenl southern limb o( the westward plunging ami- From the above descriptions, it is apparent dine near Mt Lyall (see PARACHILNA that although the Woodendinna 1>olomiie and 1:250.000 Geological Map). Here it is overlain Wirrapowie Limestone are distinctively dif- with apparent conformity by the Wilkawillina ferent in gross mineralogy and in field appear- Limestone. Further south, toward the Wilka- ance, they share a number of lithological simi- willina Gorge area, in the Central Flinders larities. They are intimately associated with one Ranges, the Woodendinna Dolomite rapidly anuthcr both temporally and spacially and thins in response to erosion, or nondeposition clearly have formed in closely related deposi- or both. At the Ten Mile Creek type section tional environments. itself, Wilkawillina Limestone disconformably The Woodendinna Dolomite xhowi- overlies the Bonncy Snndstone Member of the numerous features characteristic of deposition 1 Pound Quart2ile (Dalgarno 1964; Pierce' ). on emergent high intertidal and suprMidal basal Wilka- I he very highly dolomitired mudflats. Prevalent desiccation muilcracks however, willina Limestone in this section, record repeated periods of suhaeriaJ exposure, parts of the Wonden- superficially resembles ami the highly restricted nature of the environ- tfinns* Dolomite, but due to the degree of dia- ment is suggested by the paucity of fauna and these basal car- genesiv, the original nature of dominance of primary dolomhic mudstones or convenience bonates has no! been resolved. For pene-eontemporanoously dolomitized car- represent they are considered at this siage to bonate mudstones, The only organisms capable Limestone litho- dolomitized Wilkawillina of tolerating the extremes of restriction and facics. desiccation of the environment were algae, as Tn the east-central Flinders Ranges Gcnling* evidenced by the ahundance of stromatolites describes approximately 25 m of carbonates, in the sequence. which are tithologieally similar to Ihuse of the Cross-bedded ootd grainstones and quartz Woodendinna Dolomite, from the base of the sandstones record brief episodes of high energy Cambrian carbonate sequence at Reap hook conditions penetrating the mudflats These con- Hill. ditions, which probably occurred during Along the entire western outcrops of the periods of storm activity or times of unusually Cambrian sequence* from I'arachilna Gorge high tides, resulted in the spreading of .sheets south, a thin sequence of flaggy grey lime- of coarser elastics from more open marine stones, dolomitie mudstones, stromatolites, nat areas, over the exposed mudflats. Quart/ sand- pebble conglomerates, and imcrbedded sand- siones Sn the Woodendinna Dolomite show con- stones, occur beneath typically fossiliferous siderable variation in iheir lateral development, Wilkawillina Limestones'. These beets are reflecting the local availability of w qnariz sand assignable to the Woodendinna Dolomite and source, which in many cases probably cor- unlithif*ed the Wirrapowie Limestone and roughly cor- responded to areas of exposed, respond to what Dalgarno called the Lower Pound Quartzitc. Member of the Wilkawillina Limestone (DaJ- The Wirrapowie Limestone has developed in garno 1964). although as staled above, these a sheltered but less restricted upper intertidal dolo- lilhologics are not present within the defined to lagoonal environment. The absence of type Wilkawillina Limestone, Dalgarno & mite and the rarity of desiccation mudcracks Johnson (1963) report significant thicknesses suggest that only limited exposure and restric- of Wirrapowie-type limestones from jus! north tion prevailed. The sequence lacks coarse detri- — Flinders Ranges South B Pierce. P. R. 11*691. "the Cumbrian ecology south of thr Wirreatpa Piapir. Aitttralin. Honours Thesis, Univ. Adelaide ^unpublished). LOWER CAMBRIAN FORMATIONS, FLINDERS RANGES 219 tus, current bedding and any preferred elonga- Acknowledgments tion of stromatolite bioherms indicating that The writer wishes to acknowledge the use of energy conditions were in general still very low. the facilities of the Geology Department, Uni- As described above for the Woodendinna Dolo- versity of Adelaide, and to thank both Dr Daily mite, however, thin grainstone sheets attest to for his generous assistance over a considerable the periodic penetration, probably during period of time, and Mr C. R. Dalgarno who storms, of high energy conditions into the shel- read a draft of this paper and offered valuable tered tidal mudflat environment. criticisms. References Coats. R. P., Callen, R. A., & Williams, A. F. Dalgarno, C. R. ? & Johnson, J. E. (1962).— (1973).- COPLEY map sheet, Geological Cambrian sequence of the Western Flinders Atlas of South Australia, 1:250,000 Series. Ranges. Quart, geol. Notes, geoL Surv. S. (Geol. Surv. S. Aust: Adelaide.) Aust, 4. Daily, B. ( 1956) —The Cambrian of South Aus- Dalgarno, C. R., & Johnson, J. E. (1963). tralia. In HI Sistema Cambrico, su Paleoeeo- Lower Cambrian of the Eastern Flank of the grafia y el Problem de su Base 2: 91-147. Flinders Ranges. Quart, ^eol. Notes, geol. (xx Congresso GeoL Tnternacional, Mexico.) Surv- S. AusKl. Dai.garno, Daily, B. (1972a).—Aspects of carbonate sedi- C. Rm & Johnson. J. E. (1966).— mentation in the Cambrian of South Aus- PARACHTLNA map sheet, Geological Atlas tralia. Abstracts, Joint Specialists Groups of South Australia, 1:250,000 series. (Geol. Meetings, Canberra, GeoL Soc. Aust. cl0-cl4. Surv. S. Aust.: Adelaide.) Haslett, P. G. (1975, in press).—Lower Daily. B. (1972b).—The base of the Cambrian Cam- brian stromatolites from open and sheltered and the first Cambrian faunas. Centre for Pre- intertidal environments, Wirrealpa, cambrian Research, Univ. Adelaide, South Aus- Spec. tralia. Hap. L 13-41- In M. R. Walter (ed.) "Stromatolites". (Elsevier: Amsterdam.) Daily. B. (1973).—Discovery and significance of Johns, R. K. (1968).—Geology and basal Cambrian TJratanna Formation, Mt mineral resources of the Andamooka-Lake Torrens Scott Range, Flinders Ranges, South Austra- area. Hull. geol. Surv. S, lia. Search 4(6), 202-205. AusL 41, 1-103. Wopfner, H. (1970).—Early Cambrian Paleogeo- Dalgarno. C. R. ( 1964).— Lower Cambrian graphy, Fromc Embayment. South Australia. stratigraphy of the Flinders Ranges. Trans. R. Bull. Am. Ass. Petrol. GeoL 54(12). 2,39*- Soc. S, Aust. 88, 129-144. 2.409. THE SOUTHERN AUSTRALIAN SPECIES OF SPYRIDIA (CERAMIACEAE: RHODOPHYTA) byH. B. S. Womersley* & Sally A. Cartledge* Summary WOMERSLEY, H. B. S., & CARTLEDGE, Sally A. (1975).- The southern Australian species of Spyridia (Ceramiaceae: Rhodophyta), Trans. R. SOC. S. Aust. 99(4), 221-233, 30 November, 1975. Four species of Spyridia are recognised on southern Australian coasts. S. filamentosa (Wulfen) Harvey (including S. biannulata J. Agardh, S. breviarticulata J. Agardh, and S. spinella Sonder) is common in sheltered to moderately rough water. S dasyoides Sonder (including S. opposita Harvey and S. prolifera Harvey) is fairly common on rough-water reefs and in deeper water. S. squalida J. Agardh (including S. wilsonis J. Agardh) is a less common, usually deep-water, species. S. tasmanica (Kuetzing) J. Agardh occurs in relatively calm localities but where there is often a strong current. The Australian species differ in vegetative aspects such as arrangement and diameter of the ramelli, but agree well in the development of the thallus and reproductive structures, and emphasize the generic uniformity of the species ascribed to Spyridia. THE SOUTHERN AUSTRALIAN SPECIES OF SPYRIDIA (CERAMIACEAE: RHODOPHYTA) by H B. S. Womersley* & Sally A. Cartledge* Summary Womersley, H. B. S. t & CumtooK, Sally A. (1975).—The southern Australian species of Spyridia (Ceramiacejie: Rhodophvtn). Trans. R. Sox:. S, Attst. 99(4), 221-233, 30 Novem- ber, 1975. Four species of Spyridia are- recognised on southern Australian coasts. $, fiiantwtosa tWuifen) Harvey (including Si bianmdata J. Agardh, S. breviarticulata J. Agardh, and S. spinefla Sender) is common in sheltered to moderately rough water. S dasyoides Sonder (including S. opposita Harvey and S. proVtfera Harvey) is fairly common on rough-water reefs and in deeper water. S. squalida J. Agardh (including S. wilsonis L Agardh) is a less common, usually deep-water, species. S. tastnanica (Kuetzing) J. Agardh occurs in relatively calm loca- lities but where there is often a strong current. The Australian species differ in vegetative aspects such as arrangement and diameter of the rarnelli, but agree well in the development Of the thallus and reproductive structures,, and emphasize the generic uniformity of the species ascribed to Spyridia. Introduction and internodal cells is found in all species (the While Spyridia is a distinctive and easily cell shape varies considerably m S, fdamen- recognised genus of the Ceramiaceae, the taxo- tofin), and all species have radially disposed nomy of the southern Australian species has rarnelli (opposite in S. dasyoides) which (ex. 5. squalida) been confused. Some 1 species have been cept commonly have mucronate credited to the region, but the only recent end cells. account species of the by Lucas & Perrin Growth of the uniaxial thallus is from an (1947) and a key by May (1965) offer little apical cell which cuts off a row of short axial help in recognising or separating the species. cells which develop into a branch of unlimited The type species of Spyridia, S, filamentosa growth, from each cell of which one or more (Wulfen) Harvey, has recently been described rarnelli develop laterally. The ramelli are Of in detail by Hommersand (1963, p. 177), who limited growth, developing rapidly to between reviewed earlier studies and clarified its vege- 10 and 30 cells long by divisions of their apical tative and reproductive features. Indian mate- cell, and following cessation of cell division rial of this species has been studied by Krish- they expand by cell elongation to their mature namurthy (1968). Other species referred to length of generally 1-3 mm. The axial cells Spyridia agree well with the type in general also cut off in alternating sequence a ring of morphology, presence of nodal and internodat periaxial* cells, which form a band around the cell bands, branches of limited growth (ramclli node between two axial cells. Each of these * H or brachyblasts") and unlimited growth, and periaxial (nodal) cells cuts off two cells from in reproductive features. However, the features its lower end. and these elongate and become which Hommersand (1963, p. 177) used to attached by pit-connections to th$ nodal cells distinguish 5. fdamentosa from other species of of the next lower segment. Thus the thaUun Spyridia do not apply satisfactorily to the Aus- shows bands of shorter and broader nodal cells tralian species. Cortication by tiers of nodal alternating with the bands of iutemodal cells. * Department of Botany, University of Adelaide, Adelaide, S, Aust. 5000 t This term is used for cells cut off from, but of different form to, an axial cell, Cells of similar form to the axial cell, as found in the pofysiphonous families of the Ceramiales. are still referred to a* per [central cells. ) 1 222 H- R. S. WOMERSLEY & SALLY A. CARTLEDGE which arc longer narrower, and approximately Tetraspornngia occur on the lower ceils of mostly on the twice as many as the nodal cells, Purine! cor- the nunelli. being sessile and ucation occurs some distance from the branch upper (adaxtal) side. Hommersand (196$, p. arise directly as pro- apices, from descending rhizoidal cells deve- 196) considers that they Krishn;,- loped from the nodal cells, anil this ohscurcs trusions from the axial cell, while i 47) considers thai they arc the regular pattern of nodal and internodal cell munhy IU6S, p. hands, especially in certain species (e.g. S formed from periaxial cells- •qaallda). Some 10 species of Spyndia have been study The cells uf (he ramelli each cut off a ring of recorded from southern Australia. This S. 6-8 celts from their upper end. and these deve- recognises only four species, including fikt- lop into a nodal band 1-3 cells broad in S. mentoxa. The thallus development ajid mor- reproductive (ihmwntosa. Spyridhi is readily recognised by phology, and the structure of the alternating nodal and internodul bands, and organs, are very similar to those of S. filtt- ramelli with corticated nodes. mvntosd* and the Australian species differ mainly in vegetative features. 'I he descriptions Ueproductively Spyhdia is also distinctive, below are therefore confined largely to recog- especially in (but the carposporophyte becomes nition of the species, with brief notes only on surrounded, by periearpic filaments developed reproductive details from the segments above and helow the one bearing the procaip These filaments can eive the appearance of a eystocarp with a well deve- Key to southern Austrian species loped pericarp wall heint; held together by a ]. Ramelli robust, opposite and decussate, usually pit-con- mucilaginous sheath and some lateral 100-150 /mi thick with i»H(Ji;»metTic cells and nections, but ihey disintegrate fairly readily in nodal bands 3 5 cells broad tkuyoidt'.s (p, -31 preserved material. & V Ramelli slender, -single or whorled but not oppo- small cells Procarps (V-6) are produced on site, less than 70aiu thick, usually with 1-3 oclls lateral branchlets of rcslricied growth, Accord- longer than broad, imd nodal band* • ,. ( broad , 2 ing lo Hommersatul (1%3. p 191 .), three peri- bmnchleis stout (t-l mm thick), central (periaxial) cells are normally formed 2. Ultimate markedly basatly constricted, hi?avily corti- segment, one of which (the sup- in each fertile cated tn their apices, bearing slender, irregu- earpugonial branch, and porting cell) bears the larly branched ramelli . S. squub'da '• aim each pericentral forms an auxiliary cell Hom- 2. Ultimate branch let* slender < under not or only sliehily hastily constricted, mcrsand reported that the carpogoniurn fuses thick), corticatinn only on older branches, with third cell of the curpogonial branch with the ramelli eirher one per segment or veuualluic which then connects with the auxiliary cells by cells. Thus two (or rarely 3."5-t>5 /urt thick, nodal means of connecting J; Ramelli one per segment, 2-t S. 222) three) gonimoblasts are initiated and the ma- hands cells broad glshieitt&m 4. Pt Sranvae. S. Atist. (Uwh. MaMfc^ FiEB I &9ttfamm**te&. * Kraft, l4:o..W1: ADO. Mil flW.« MuJc cystoearp . Denison. W. A*tf. . plant with vo.ing ft °- 30-" 1 ADU plant with "sperLtangial ramelli f&lfai, S. Ausl. CWlMft W2; - **»»! Tetrasporangia! plant (A41KI5V THE SOUTHERN AUSTRALIAN SPECIES OF SPYRWIA 223 B-D , # 300/um * ft f Fig. 1 224 H. B. S. WOMERSLEY & SALLY A. CARTLEDGE THE SOUTHERN AUSTRALIAN SPECKS Oh SPYRIfUA 225 1 832: J 8. Tisdall 1898: 505. Wilson cells I Fig. 3 A) which taper abruptly to a 1692; iSl Wornersley 1958: 157, mucronate cell, f 35-) 4f>-55<-65 > pn\ thick Ftwuf fxUovctitaws Wulfcn 1803: 64. with cells (I J-) N-241-3) times as long as Jf, fftvmtMtim VflPi urhtixcuta Sender 1855; 518. broad; mucronatc end cell often Inst from btamHtfote J. £ Agardb Jg76: 267; |gfe: 13. older ramelli. ramelli with about 9 nodal cells, De Toni 1903: 1426. Guiler 1932; 98. Lucas each usually cutting off 1 (-2> cells anleriortv, 190$: 52: 1929a: 25; 1929b: 53 Lucas & Perrin giving a nodal band 2-3 1947: 363. May 1965: 369. Oltamura 1932: cell* broad. 130. Retnhotd 1897: 60; IS99; 50. Sender I8S0; Cys/ocarps (Fig. \B) short-stalked, usually 16. Tadall 189ft stXV Wiison 1892: I8L bilobed, lobes globular, 300-700 pip across. Wornersley J950: 180. Spermutangia covering the .V. hrrviarUttttaUi J. Agardh 1876: 268; 1897 lower (except 13. Dc Toni 1903: 1427. Guile* 1952: 98. basal) several segments of ramelli (Fig. IC). Lucas. I9U9: 52; 1929a: 25; I9&&! 53. Lucax & forming maie organs 75^120 jam in diameter. Pen in 1947; 363 May 1965: 369. Osamura letrasporanfjia 1932: 130. Reinbold 1897: 60; 1890: 50. Sonder (Fig. ID) sessile, 1-3 per 1**0: 16. cell on lower cells of ramelli, mostly on the i. stnneUa Sonde r 1845: 53: 1846 168; ISS0- upper (adaxial) side, spherical, 50-75 ,»m in J6. J. Agardb 1852- 342; 1876: 269; IH97 13 diameter, fctrahedrally divided, He Tooi 1903; 1430. Harvey |B63, SVflOB.! 42. Type locality; Adriatic Sea. KuctzJng 1*49; 668; 1862: 16, pi. Sled. Lucas 1909: 52. May 1965: 369. Mzzzu 1925. no. 824. Type: 1 Okamum 1932: 130. Distribution; All around the Australian coast FIGS I, M.B (including. Tasmania) in conditions of mod«r- Thatlui (Fig. usually M) 7-18 cm high, atc to slight water movement. epilithic or epiphytic on various larger algae Spyndia fifonwitosa is recognised as a and scagrasses. lax and soft, irregularly much widely distributed species, having been branched on all sides with longer and shorter recorded from most seas, and manv authors branches intermixed, with one to several axes (e.g. Harvey 1846, pf. 46; Felduiiinn-Mazoyer (often poorly defined) from an originally dis- 1°40, p. 348) refer to h as a very variable coid holdfast, soon becoming fibrous or stoloni- species J. Agardb (1876, p, 268) in segregat- tcrous and entangled, commonly grey to grey- ing two Australian species [S. hUmwtota and red, sometimes red-hrown, in colour. Axes and S. hreviar/jculata) from 5. filamentosa, referred larger branches corticated, terete; axes to their i-1 similarity in habit with Si fitamentoxa t—If) mm thick, tapering to branches 300-500 and the large number of forms classed as this /ntO thick and hranchlets 100-300 ,/m thick; species. J. Agardb apparently regarded his two laterals arising from periaxial cells or advents segregate species with some doubt, and a de- liously from cortical Cells. Segment* usually tailed study of extensive collections of Ausira- clearly defined on branchlcts (Fig. J/?), vari- lian material does not provide any satisfactory able in length and proportions but usually way of segregating S. binnnukita and 5". brevi- (i—H-l times a.< long as broad, with bands of art'wulQia from S. filamentosa. shorter nod^l cells and longer internodal cells The type of 5. bimmulata is from Tasmani;. alternating; nodes with 11-14 periaxial cells, (Georgetown, Tav. Gtwn. LD, 51300, selected each corresponding to two internodal cells ex- as lecto(ype) and was distinguished by J, cept for the (.usually) larger periaxial eel) bear- Agardh in having more conspicuous bands of ing the famellus. Cortication usually commenc- nodal and internodal cells and being ing less cor- few mm from the apices but very ticated above. Si breviarticttlaiu is based on variable, consisting of rhi/oidal cells lying specimens from Whitsunday T., Queensland between the internodal cells and gradually (lectotype in LD, 51311). forming and was distin- a continuous cortex l(-2) cells thrck\ guished by having the nodal and internodal Raoidh (Figs }B 3A, B) t single per segment, bands shou and of ~ almost equal length. The irregularly spirally arranged, j-j* mfn | ong variation in S, filammtosa with encompasses the 12-201-27) cells, linear or gently taper- above features of both S. buttmutata jng apart and S from the terminal 2-3 very short hwviaeticHfaUt. Fl * % A ForuuHngtott, SESfettSP** Vic. (Woilastrm, I7.vlli.1956; AM), A20S67) B Femak- ".melll (Taplcy Shoal. A„s, , 15 rtwot —n.ivnv, ajju, % m SEKh%^f^A^^ri^^Aj>>38). D. Tetrasporangial plant %W (A4fQ$gt, 226 H. B. S. WOMHRSLEY & SATXY A. CARTLEDGE in face view mid region oC ramellus, with periaxial cells Pig 3 4, B. C Wumentout. Apex and (A41815). , fli,n^ C\ O. 5. wsmanica. Ditto (A41066). c.br.-carpofipnial branch; p.a.-pcnax.al cell; 8 .c.-suppw t- jSjf£' 7885 ADUT A35287>. face view region of lamellus, with periaxial ceils in flf f\ snualida. Apex and mid (A26375). /, K. S, tla^oides. Ditto (A20I70) THE SOUTHERN AUSTRALIAN SPECIES OF SPYRID1A 227 S, filamentosa var. arhuscula Sonder (1855, periaxial cells are usually transformed into p. 518) from Wilsons Promontory, Vic, May tetrasporangia. The number of periaxial cells 1853 (type in MEL, 45181) is typical of the in a ramellus does vary slightly, so it is not species and not a distinct variety. possible to state as Hommersand does that S. spinella Sonder (type in MEL, 502090) tetrasporangia must arise directly from the was based on Preiss material from Western axial cell because the number of periaxial cells Australia. The type has somewhat broader and is the same in sterile or fertile segments. stouter ramelli than most specimens of S. Spyridia tasmanica filamentosa, with cells about as long as broad (Kuetzing) J. Agardh 1852: 342. and the nodal bands of the ramelli 2-3 cells Gordon 1972: 39. Harvey 1859: 329. broad. Collections from Cottesloe, W. Aust., Kuetzing 1862: 14, pi. 42 c, d. reef pools {Parsons, 14.xi.1968; ADU, Sonder 1853: 680. S. filamentosa A34072) and from Elliston, S. Aust. (Womers- var. tasmanica Kuetzing 1849: 666. ley, 15.1.1951; ADU, A 15142) agree well with S. filamentosa var. verticillata Harvey 1 844: the type in having densely aggregated ramelli 449. 400-800 yxm long, composed of 15-18 cells, Wrangelia setigera Harvey 1859: 309. pi. I9M; 50-65(-90) fim thick and 1-H times as long 1863, synop.: 27. J. Agardh 1876: 622; 1879: as wide. pi. 32, fig. 3. De Tom" 1897: 133; 1924: 149 This is within the extremes of S. fila- Gordon 1972: 39. Guiler 1952: 99. Lucas 1909: mentosa and is probably typical of plants 23; 1929a: 16. May 1965: 365. Mazza 1919. no. occurring in rock pools subject to moderate to 678. Okamura 1932: 133. Sonder 1880: 29. Tis- considerable water movement. These plants dall 1898: 511. Wilson 1892: 170. are much closer to typical S. filamentosa than FIGS 2, 3C-G, AA-C the Brest specimen discussed below, and are Thallus (Fig. 2A) usually 8-25 cm high, provisionally placed under S. filamentosa, but irregularly much branched, epilithic or on further studies on this "spinella" form and its Amphibolis, with a small, discoid holdfast, variation are desirable. grey-red to red-brown in colour. Branching S. filamentosa has been studied by several irregularly alternate, branches terete, with one authors, most recently by Feldmann-Mazoyer to a few main axes and prominent lateral (1940, p. 348), Hommersand (1963, p. 183) branches, bearing lesser branches and branch- and Krishnamurthy ( 1968, p. 42). The Austra- lets on all sides, with whorled ramelli. Axes lian material agrees well with these descrip- l-li(-2) mm thick, branches about £ mm tions, and also with material from Leghorn, thick, lesser branchlets 200-400 ^m thick. Seg- Italy (Sartoni, 18.viii.1973; ADU, A43938). ments (i-)J-H times as long as broad (Fig. However, material from Brest, France 2D), with usually 12 periaxial cells, each pro- (Cabioch, Dec. 1972; ADU, A43050) has dis- ducing two internodal cells; cortication by rhi- tinctly more robust ramelli (about 100 ^m zoidal cells from the nodal cells, commencing thick, cells scarcely longer than wide) which within 1-2 cm of apices and becoming heavy have 2-3 very small basal cells with the parent on axes and main branches. Ramelli (Figs 2D, periaxial cell not enlarged, in contrast to the 3C-F) l(-3) per node near apices, becoming full-sized basal cells and enlarged periaxial cell whorled (3-6(-8) per whorl with the addition of typical S. filamentosa. These two collections of adventitous ramelli), arising from enlarged indicate that there may be greater variability periaxial cells, £-2 (-3) mm long with 20-30 in European S. filamentosa than in the Aus- (-35) cells of greatest diameter (15-) 20-35 tralian material. (-40) ^m and 3-5 times as long as broad, the Hommersand (1963) and Krishnamurthy end cell mucronate; robust form with ramelli ( 1968) differ in some details in their accounts ( 1 2-) 1 5-20 cells long, greatest diameter of reproduction in the material they studied, as 30-40(~45) ^m and l£-2i times as long as mentioned above in the introduction. Austra- broad. Ramelli with a single row of 8-14 nodal lian material (e.g. Aldinga, S. Aust. Cartledge, cells (Fig. 3D. F). 30.iii.l972; ADU, A4188I) shows three peri- Cystocarps (Figs 2B, 4A) terminal on a axial cells in fertile segments, but clarification short branchlet bearing ramelli, i-£ mm in of immediate post fertilisation stages has not diameter. been possible. No clear stages have been ob- Spermatangia cover 2-6 cells (Figs 2C, AB) served of a tetrasporangium arising directly within 1-2 cells of base of ramellus, forming a from the axial cell of a ramellus (Hommersand cylindrical male organ 70-130 ^m in diameter 1963, p. 194), but it appears more likely that and 1-6 times as long as broad. 22X H. B. S. WOMERSLEY & SALLY A. CARTLEDGE Fig. 4. THE SOUTHERN AUSTRALIAN SPECIES OF SPYRIDIA 229 Tetruxporanxia (Figs 2D. 4C) on 1-4 cells more characteristic of plants found under near the base of the ramelli, borne mostly on rougher-water conditions. Flutter studies on the upper (adaxial) side, 60-100(-120) Mm in this form are. however, desirable. diameter when mature, tetrahedraliy divided. Spyridia squalida ), A*»ardh J 876: 270; 1897; Type hiatity; Tasmania (probably Cmnth ex 16. De Toni 1903; 1436. Lucas 1909; 52; Hooker I 1929b: 53. Lucas & Perrin 1^47; 364 Type: l (941, 3ti...37n. May 1965: 369. Okamura 1932: 130. Distribution; Elfiston. From S. Aust. lo Reinbold 1897; 60. Sotukr IKKO; 16. Western Port, Vic. and aiound Tasmania. Gen- Tate 1882: 18. erally in relatively calm localities, often wilb $ wiJsortisJ. Agardh 18^7: 16, De Toni (90S: cuiiNtdcrablc current. 2-35 deep, occa- M35. Lucas 190V; _S2. m lue;is ft Perrin 1941 sionally in partly sheltered habitats and shaded 3c»4. May 1965: 396". Okamuia 1932: 130. •V. KttlkUi Sontlrr 18X0* tuck pools on tough-water coasts. 16 (uomen nudum). Development of reproductive structures in FIGS 3//. /, 4 A /. S. itisniattica appears to be very similar to that ThaUus (Fig. 40) usually 10-30 cm high, in 5. ftlamentaw. The female axes (Fig. 3G) robust, erect, irregularly and proliferouvly arise as short laterals and bear to 5 up pro- branched, epilithic with one to several axes carps, separated by one or two sterile segments from a small discoid holdfast, usually with each with eight periaxial cells, one of which long, much branched, laterals on all sides, hears a ramellus. Each procarp consists of 3 grey- red to red-brown in colour and when (rarely 4) periaxial cells, one of which is the dried Often appearing somewhat farinaceous, supporting cell bearing the 4-celled carpogonial All branches terete and corticated to thcij branch. Immediate post-fertilisation stages have apices, axes and main branches linear, branch- not been clearly followed, but usually two lets (Fig. 4E) basally constricted and bearing groups of gonimohlast cells develop, probably densely arranged ramelli, especially on their from fusion cells originating from auxiliary upper parts, sometimes denuded below. Axes cells cut off from the supporting cell and one li-2i mm thick, denuded below or with short, of the other ferule periaxial celts. Sterile peri- proliferous branchlcts, tapering slightly in carp filaments arise from the periaxial cells of branches 1-1 f mm thick and lesser branchlcts segments above and below the proearp-bearing 4-1 mm thick. Segment* largely obscured by segment, forming a cysfocarp similar to that in eorticatiou, \b**& as long as broad, with 16 S fiiamettfoxft. periaxial cells and about twice as many inter- Development of spermatanvia and tetraspo- nodal cells; eortication commencing within a rangia is similar to that in S. fifamentoxa. few axial cells of apices, pseudo-parenchyma- tous, 2-3 S, fasmanica is a distinctive species with its cells thick on branchlcts, several cells thick v* hoi led ramclli and single row of nodal cells. on axes. Ramelli (Figs 3//, /, 4E) one The ramelli are per segment close to apices typically slender (20-35 /Am and derived from thick), with matuie cells 3-5 times as long as periaxial cells, but scattered adventitious hroad. However, some plants from rougher- ramelli (usually) densely cover the hnmehlcts. water localities |e.g. Elliston, S. Aust., 7 m sometimes persisting onto larger branches; deep (Shepherd, 20.x. 1970; ADU, A37622) ramelli *-1(-li) mm long with (IO-)I4-20 and Cape Lannes, S. Aust., in shaded pool (-24) cells of greatest diameter (20-) 30 40 I 12.ii. 1972; (-45) pzh and (1-j I Krttft, ADU, A41S09 ) | have J-2(-2J ) limes as tone more robust ramelli, 30-40f-45) yjix thick as broad. Ramelli with a single row of small 3£. F, nodal cells (Fig. CFifiS 45, C) and mature cells 1 t-2i 31) derived from 5-6 peri- times as long as broad. Otherwise the latter axial cells each of which cuts off 2-3 outer specimens are similar to the majority of plants, cells in the same transverse plane. and the type, of tasnumha, & and the more Cystocarps short-stalked, globular-bilobed, robust* shorter-celled rametli are regarded as \-i mm in diameter 4. Spyrun tasmanca (robust 1% form . A. Female plant with quashed mature cvstocarp (EUislon S. Aust., J deep in m bay. Shepherd, 20.X.I970; ADU, A?7ft2?). B. Male plant langud wiUi spermP ramelli (A37622). C. Telrasporangbl plant (A3TO2) Spyndta sxjualidti D. Victor iLirhor, S. Aust. iWotnerslev, iK.iii iVfcft; ADU UOtmi K Branchlcts and ramelli (Pt Elltol. S. AnM- Dodd, J2.iii.I463; ADUi A26575J 230 H. B. S. WOMERSLEY & SALLY A. CARTLEDGE . 1 THE SOUTHERN AUSTRALIAN SPECIES OP SPYJUDlA 23 Sperrnatattgia cover the lower 2-6 cells Spyridia dasyoldes Sonder 1853: 680; 1880: (except basal cell) of ramelli, forming a male 16. J. Agardh 1876; 272. De Toni 1903: organ 50-1*0 plti in diameter, 1437. Harvey 1863, synop.: 42. Lucas Tetraspvrangia borne on the iower several 1904: 52. Lucas & Pcrrin 1947; 364. Mav ( ceffs of tbe ramellK largely on the apper 1965; 369 Okamura l >32: 130. Tate (adaxial) side. 1 (—2) per cell, sessile, spherical I8S2: l«. Tisdal! 1S98: 505. to slightly ovoid, 40-60 pm in diameter when S, oppoxha Harvey 1855b. 256; I860: pi, 15*. mature., tetrahedralfy to sub-cruciately divided. Adi.ms. 1972: S3. J. Agardh 1876; 270; 1897: Type locality: "Nov. Holland, australem". H De Toni 1903; 1431; 1924: 502. Guile* 1952: 9fc. Lucas 1909: 52; 1929a: 25; 1929b- Type: Herb. Agardh. LD, 51533. 53 Lucas * JVrrin 1947* 363. fig, 1*2. Mav Distribution; From Geographe Bay, W, 1965: 369 Mama 1912, no. 421. Okamura 1932: 130. Reinboid 1897: 60. Shepherd & AusL to Waoatafi Bay, S. Gippsland, Vic., Womersley 1970: 135. Sonder ISSOr 16. Taie- usually in deep water i 2-24 m deep) IS82: 18. Tiwlall 1X98 505. Wilson 1892: 1*1. Female axes of 5. squckluto develop as short, WnnwNley 1950: 180; 1966; »51. adventitious branchlets which are more heavily 5. pralifera Hnrvcy 1863: (A. 274. J. Agardh 1*76: 1597: 14. Toni -a r f ;:.it. 269; De 1903: 1431. , d than in other species but less so titan Lucas 1909- 52. Lucas & Perrin 1947: 362. fie. in vegetative branchlets of this species, Alter- Ifil. Mav 1965. 369. Mazza 1912. no. 422. nating segments each bear a procarp, with the Sonder 1880: 16. sterile segments be^Ting ramellr. Usually three FIGS 3i, A', 5 periaxial cells occur in fertile segments, one line supporting cell) producing a 4-oeUed car- Thallus (Fig. SA-C) usually 10-20 cm high. pogontal branch. Two, or probably ofien 3. eroc(, eprlithic or epiphytic, much branched auxiliary cells are formed, leading to >i carpo- with one to several axes from an originally div sporophyte with two or three lobes. The pen- coid holdfast which soon becomes fibrous and develops carp similarly to that in other species stoionrferous, dark red to red-brown in colour. S. squaUda is a distinctive and robust species Axe* and larger branches heavily corticated, of Spyridia, having cortication to the apices terete to angular and becoming four-Mdcd with and thus forming swollen, basally-constrided, thickened cortical flanges in line with the 4 branchfets, bearing ramelli often densely scat- ranlcs of ramelli, densely branched. Axes 1-2 tered hut usually soon denuded. The farina- (—2i) mm thick, often denuded but sometimes ceous appearance is also a common feature of with numerous, short, proliferous branchlets, older, dried plants tapering to branches }-£ mm thick and lesser S. wiltot\ii J. Agardh is typical S. sqttaJida, branchlets k-\ mm thick; branching usually The type of the former is from Pt Phillip Hds, subdistichous (Fig. $C) with laterals arising Vic {J. B. Wilson, 1857; LD 51532), and tbe from nodal cells. Segments largely obscured by thallus is not compressed ns stated by J. cortication, Kl times as long as broad, with $ Agardh ( 1-897) and May (1965V perineal cells producing 16 intemodal cells 5. vul'ulu Sonder (1880: 16) is a nomen and the 8 cells soon with interposed rhteoidal nudum, based on a specimen in MEL (45195) cells giving both nodal and mternndal rings of from Geographe Bay. W. Aust. (Bunbury. 16 cells (Fig. S/)i; cortication commencing IH75), accompanied by Sonder* drawings. It within a few segments of apices, of clongalc is typical S. sauaiida. cells later appearing pscudo-parenchymatoui, a «Y. xtjuulUh was recorded from Port Alfred few cells thick on branchlets, many (especially (Kowie), Souih Africa by Barton (16%, p. on flanges) ccih thick on axes. Ramelli (Figs ?96>, and the record repeated bv De Toni 3/, K* 5£>, E) arising from an enlarged peri- (1903, p. 1436) and Lucas & Porria (1947, axial cell, in opposite and more or less decus- p. 364), This record almost certainly applies sate pairs (Fig. SD) on successive segments to some other species, probably to S. pfamosa (often displaced to two rows on each side in Schmitz ex J. Agardh (G. F. Papenfuss. pers. the plane of branching), n-}|*-2(-2*) mm comm.1. long with (16-HX-22 cells, relatively uniform Fig. 5 Spyridic desyoides. A Holotype (MEL, 45128). B. Robe. S. Aim. 232 H. B. S. WOMERST.EY & SALLY A. CARTLLDOE in diameter and tapering fairly abruptly to a by rhizoidal cells, thus forming both noda) and point, (70-) 100-1 50 ^m thick, cells about intcmodul rings of usually lf» cells, the bands l( H) limes as long as broad. Ramclb with being about equal in length. 16-20 nodal cells, each cutting off 1-2 cells The type of S dosynidts (Fig. 5A) in MEL (which otren divide again 1 on both skies (an- is typical of this f-peeies, previously known teriorly first), producing a nodal hand (2-) 3-5 mainly as S. opposita, Sender's hand-wriucn (-6) cells broad (Fig. 3A K). label with the holotype gives the locality as Cystocarps short-stalked, irregularly globular "Adelaide'', inland from Holdfast Bay. The to bilobed, 400-600 ^.tn in diameter. type of S. opposha (in TCD) is from Preserva- Spermutansict cover the lower (except ha.sal) tion Harbour, on the souih-west coast of the several segments of young ramclli of adventi- south island of New Zealand (LyalL Jan nous hranchlets lying between older ramelli. 1851), and agrees very well with the southern forming male organs 120-200 /*ni in diameter. Australian plants. Adams f 1 972, p. 83 Terrasf'Oruttgia (Fig. 5E) sessile, 1-3 per records S. vppoxtta from several localises near cell, mostly on the upper (adaxiul) side of the Wellington, New Zealand, and although it is ramelli, Mibspherieal, 50-90 /j,m in diameter, apparently not widely known in New Zealand, tcUahcdrally divided the specimens (e.g. CHR, 55775, from Patu- rau, Nelson) agree well with Australian mate- Type totality: Holdfast Bay, S. Ausl. (F.v. Harvey, in TCD, Mueller). rial. The type of S. protifera is from Fremantle. Western Australia (Clifton), ry^.MHL, 45128- and is a plant with denuded branches hearing Mxfribittion: From Port Dcnison, W. Ausl. proliferous hranchlets. S. prolijtta represent *> u> L, Vic. S. dasyoides usually occurs on Gabo older plants of S. dasyoides. where Ihe ihick coasts from low tide level to rough-water axes and branches arc probably remnants of growing plants are depths of 33 ra. Deeper the previous yeats' growth, from which num- than those growing in usually more delicate erous short branchlets have arisen prolifcr- turbulent conditions. nusly. The structure of the ramelli. is idenlicnl reproductive cells develop very sunilarly The with that of 5. da$y*>Ui?s. Plants referred to 5. in or S, 7asmank- pericarp is relatively lirrn at its periphery. vided by Mrs Hnid Robertson and Miss Cheryl S. tUisyoides h characterised hy its robust, Andcisoo- Dr G. Sartoni. instrluto Botanico, opposite and more or less decussate ramelli. Fircnze, Iialy. and Or J. Cabioch, Station Biu- with cells ahout as long as broad and nodal logtquc de Rosccff. France, kindly made avail- bands 3-5 cells broad., logcther with the largely able material of .Y. fflamentoM from I'm/ope distichous branching. Eight periaxial cells are The Director of the National Herbarium, Mel- formed in branches *nd soon become separated bourne, is thanked for the loan of specimens. References Adams, Nancy M. (1972).—The marine Algae of Bauton. Hihel S. (181*6), —Tape Afepe J. Hvl the Wellington area. A list of species. Hec- M, 193-1^8. Dom. Mus. 8(5), 43-*>8 BOBRQtsaN, F THE SOUTHERN AUSTRALIAN SPfcCifS OF SPYRWIA 23» hFLDMANN-M^OYEft, G, (1940).—"Reche by P. B. Webb Summary 2>5 OBITUARY GRAHAM FREDERIC WRITTEN, M.Sc. 16TV.I924-16.I1U975 Graham Whitten, former Deputy Director of his premature retirement due to serious illness, Mines in South Australia died on 16 March, in November 1973, 1975, after a long illness, With his passing the Whitten made a significant contribution State lost an outstanding geologist towards the understanding of the iron forma- Whitten graduated as B.Sc. from the Univer- tions of South Australia. In relation to this sity of Queensland in 1946 and subsequently work he visited Europe and North America m jnined the Electrolytic Zinc Company where he 1958 and 1967, and was awarded an M.Sc worked as geologist and later Senior Geologist degree from the University of Adelaide. at the Rosehery Mines in Tasmania, After He was elected Fellow of several years at Rosehery. he became respon- the Society in 1962 and was Secretary from 1970-72. played sible for exploration for the company in He a prominent part in Tasmania, New South Wales and Queensland, the affairs of the Geological Society of Australia and was also In 1954 he accepted an appointment as an active member of the Australasian Senior Geologist in the Geological Survey Institute of Mining and Metallurgy. Division of the South Australian Department He was also a member of the Society of Economic Geologists of Mines. He was promoted m 1965 to Super- and of the Society of Exploration Geophysicists vising GeoJogtai and during the next five years (U.S.A.). was responsible for all activities of the Explora- His energy and drive will be remembered by tion Services Division, and in 1970 was all who weie privileged to know and to work appointed Chief Geologist, with him. His many official reports and With the untimely death of the then Acting published papers are a fitting record of his Director of Mines, Dr K. R. Miles, ui March career. 1972, Whitten was left in administrative con- To his wife Beuy and to his three daughters*, trol of the entire Department for a period of we offer our deepest sympathy. He wiD be sadly eight months. He was subsequently appointed missed by his professional associates and bv Deputy Director, The post which he held until his many personal friends, B. P. WEBB Bibliography I9tf Investigation of Aeromagnetic Anomalies iyf>7 Investigation of Carrow and Neill aero- hundreds of Carina, Chandada and Ripnn— magnetic anomalies. Min. Rev , Western Adelaide, Eyre Peninsula. Kept, fnxesi,, %rol. 121, 40-46. Sitrv, S, Ausi, 23. 1968 Type Section of Iron Formations, Tarcoola Ironstone deposits. Radium Hill district District. Quart, Geo!. Notes, Min. Rev. Adelaide, 117, 80-87. geof. Sarv, S Aust. 26. |yM Metallurgical testing of Pceralilla H||I latcn're. Min. Rev Adelaide, 117, 89-90. 1968 Atlas of Technical Data. Quart, Gent J Notes, 965 Investigation of Kopi acromagnelic ano- geot. Surv. S. Ami. 26. malies. Min. Rev, Adelaide, 118, 116-123. 1 5*8 With Kisefy. B. G. A ground magnetic and 1965 The Uhc of AeronrtHgndii Maps ill Geo- gravity survey of the Wanamboo Aeromag- logical Regional Mapping. Quart. Geo!. netic Anomaly, Central Ryre Peninsula. Notts. $eo!. Sttrv. S. A MM. 15. Repl. Invest, gcal. Sur\'. S. Aust. 32, 1965 Iron Ore deposits in South Australia out- 3 969 Regional .gcochemical investigations in the side the Middleback Kanges. Ptor, Eighth Ctare one-mile sheet Comm, Min. Metull. aTca. Mineral ftesour Congress. Aust. AVir Rev.,S. Zealand, Aust. 127,34-45. J 965. 1 , 309-3 11. 19711 The 1065 Leached Outcrops, Prov. Eighth Ccmm. Investigation and Exploration of ;he Rtuorbacfc Min, MetalL Conpress. Aust, Nwot Zealand Ridge Iron Deposit Rrp. Invest., 1965. % 193-204. f?eol, Surv. S. Aust, X$, 1966 The Geology of some South Australian Iron 1971 The Investigation of the Almanda mine Deposits. M.Sc. Thesis, University of Ade- area. Mineral Resour. Re\\, S. Aust, 131 laide (unpublished). 25-37. 1*66 Suggested Correlation of Iron Deports 1972 Annual Report of the Director of Mines within South Australia. Quart. Geo}. Notev and Government Geologist—for year ended zeal. Snrv S A us/. 18. 30th June, 1972. South Australia. ROYAL SOCIETY OF SOUTH AUSTRALIA INCORPORATED Patron: HIS EXCELLENCY THE GOVERNOR OF SOUTH AUSTRALIA SIR MARK OLIPHANT, K.B.E. OFFICERS FOR 1975-76 President: C. R. TWIDALE, Ph.D. Vice-Presidents: B. R WEBB, M.Sc. N. H. LUDBROOK, M.A., PLD. D.I.C., F.G.S. Secretary: Treasurer: C. J. M. GLOVER, M.Sc. S. A. SHEPHERD, B.A., LL.B. Editor: Assistant Editor: H. B. S. WOMERSLEY, PLD., D.Sc. W. K. HARRIS, M.Sc. Librarian: Program Secretary: I. M. THOMAS, M.Sc., MX, Biol. W. V. PREISS, Ph.D. Minute Secretary: R. H. FISHER, A.U.A. Members of Council; P. J. M. GREENSLADE, M.A., Ph.D., R. W. R. RUTLAND, PhJX Did J. H. J. SZENT-IVANY, Ph.D. f J. K. KING, Ph.D. F.R.E.S. Auditors: Messrs MILNE, STEVENS, SEARCY k CO. Registered for posting as a publication Category 'C* Printed by Graphic Services Pty Ltd, 86a PIrle Street, Adelaide 6000