THE ASCIDIANS OF SOUTH AUSTRALIA II.

NORTHERN SECTOR OF THE GREAT AUSTRALIAN BIGHT AND

ADDITIONAL RECORDS

by Parricia Kott*

Summary

Korr, Patricia (1975),—The Ascidians of South Australia TTT, Northern Sector of the Great

Australian Bight und Additional Records. Trams. R. Sov. JS. Aiest. 99(1), 1-20, 28 Feb-

ruury, 1975,

An account is given of 58 specics of ihe Ascidizcca from South Australia, of which

7 species are new, incliding two assigned to new genera in the sub families Euherdmantinae

and Botryllinge.

Records ef 22 species from the northern

part of the Great Australian Bight ure the first

from that area nnd suggest that the ascidian fauna there has a considerable endemic com-

ponent. Many of the species common in other parts of the Flindersian marine faunal Province

have not yer been recorded from this location.

Introduction

This account of the ascidian fauna of South

Australia is based on the following collections;

(1) from the northern part of Spencer Gulf

(made in connection with environmental

studies in that urea); (2) from the northern

part of the Great Australian Bight (made in

connection with an experimental Prawn Trawl

Survey, Explorer): (3) from Investigator Struit

(collected by J. Watson) and from West

Island; (4) additional collections from Elliston

Bay at the eastern end of the Great Australian

Bight (collected by S. Shepherd). The report is

supplementary to previous papers on the South

Australian ascidian fauna (Kott 1972a, 1972b).

Tt includes records of 57 species, including 7

that are new to science. Two of these new

spesies have been assigned to new genera in the

sub-families Kuhetdmaniinue and Botryllinae.

Four species are newly recorded from South

Ausiralia.

The occurrence of 6 new species in the

northern part of the Great Australian Bight

suggests an unusually high endemic component

for the ascidian fauna of that area, and its zoo-

geography is discussed.

Type material is deposited m Australian

museums as indicated by the abbreviations AM

(Australian Museum), NMV (National Mu-

seum of Victoria), QM (Queensland Museum),

and SAM (South Australian Muscum).

All available collection data for the speci-

mens discussed urc given in the Appendix.

Order ENTEROGONA

Suborder APLOUSOBRANCHIA

Family CLAVELINIDAE

Subfamily CLAVELININAR

Podoclavella cylindrica (Quoy & Gaimard).

Kott, 1972a: 5 (synonymy): J972b: 167,

New Records: Tipara Reef (Spencer Gult);

on reef NNW Douglas Bank (upper Spencer

Gulf).

Subfamily HOLOZOINAE

Distaplia australiensis Brewin, 1953: G1. Koit.

1957; 95.

New Records; upper Spencer Gulf (Stn DS).

Previous Records: Tas. (D’Entrecasieaux

Channel and southern Tasmania),

Description: Colonies consist of a rounded head

on a short cylindrical stalk, There is a single

terminal common cloacul aperture and the

zooids are arranged along cither side of the

cloacal canals that radiate from this uperture

and extend down the length of the head, There

are about 12 fine longitudinal thoracic muscles,

Ten stigmata ure present in each of the four

rows. and these are crossed by fine parastig-

matic Vessels, There are 8 rounded stomach

folds, The gonads extend, from the pole of the

* Queensland Muscum, Gregory Tce., Fortitude Valley, Qld, Australia 4006.

2 PATRICIA KOTT

but loop, milo a short posterior abdominal

extension separated fram the abdumen by a

shov. neck. Seven to 8 elongate testis lobes are

arranged in a circle with their long axes paral-

lel @ One another ta form a barrel shaped mass.

‘The vis deferens, extending trom the distal end

of the centre of this mass, passes around it

into the abdomen. There is also an ovary in

the posterior abdomen.

Remarks: The colonies are identical wilh those

previously assigned to this species. The zooids

differ from those described by Brewin in the

lesser number of stigmata in each row. Koti

(1937) has reported some variation in this

character and the differences an: not regarded

as significant, The presence of a purastiznalic

vessel crossing the rows of stigmata has not

previously been observed, but since thisas very

delicate it could have been averlouked.

Sycozoa pedunculata (Quoy & Gaimard). Kott,

IY72h: 170; 19720; 234 (synonymy),

New Record: upper Spencer Gulf (Sta B4).

Atapozoa tarshi (956; 31. Kott,

1972b: 168.

New Records: Investigator Strait (Stns YS,

ZA).

Description’ The specimens arc of the usual

form with a long evlindrical bead terminating

ima rounded point. The shorter fleshy stalk is

almost the same diameter as the head. The

colony from Sto Z6 is the largest yet recorded,

measuring 17 om of which the stalk is only 5

cm, The minute zooids are present jm the sur-

face layer of test with long posterior abdominal

stolons penetrating info the wentre of the lahe.

‘There ts the usual brown pigment patch over

the anterior end of the endostyle.

Family POLYCITORIDAE

Polycitor gigantcum (Herdman). Ko, 1972a: 9

(synonymy): 1972c; 244,

New Records: votthern Great Australian

Bight; West T. | Amphitheatre Rock}.

Family POLYCLINIDAR

Subfamily WUHERDMANIINAR

Euherdmunia ansiralis Kott, 1957: 103; 1972b:

172.

New Recards; Elliston Bay (outside bard;

Investigator Strait (Sin ¥5),

Deseription; The colonies are formed of the

usual long cluh-shaped lobes joined basully-

Rich lhe jis composed of 4 single zonid

covered hy as own separate sheath of sand-

Brewin,

stiffened test. ‘There pre 9-11 rows of 27-28

stigmalst, each row crossed hy a parastizmatic

vessel A pointed papilla is present in the

middle of each primary transverse and para-

stigmatic vessel on both sides of the body. The

mternal wall of the stomaeh is arranged in

longitudinal and transverse elimdular ridges

rather than folds.

Ritterella herdmania Koil; |972a: $1 (synu-

nymy); 1972b; 172; 1972: 246

New Record: Elliston Bay (oulside bury,

FIG. 1

Deseription: The present colonics sre smuller

than usual and sometimes each lobe contains

only a single zooid. Vhe lobes are the usual

spalulate, Jong, narrow-stemmed form, Each

zooid hus 5 rows of about 5 stigmata but there

ire 6 parastigmatie vessels, and a single

papilla is present in the middle of each trans-

verse’ Vessel. There are only single rows uf

testis follicles in the posteriar wbdomina. There

are 1--4+ embryos in the peribranchial cavity,

Larvee ate very. small, 0.3 mm lang. They have

3 median ampullac that alternive with the

papillae, and double rows of vesicles vhal

extend around the anterior aspect of the larvac

ou either side of the papillac and .ampullie sind

extend posteriorly along either side of the

dorsal mid fine. There is also a pated series of

vesicles that extends postero-ventrally (Fig. 1}.

Remarks: This species has been taken from

Elliston Bay (Kott 1972b) im May 1971. anu

the present colonies were collected in the pre-

Vious February, Only the latter ure sexually

mature and contain larvae, It is not cleur

whether the colonies taken in May were newly

settled forms, yet to reach reproductive matur-

ily, or whether they were older colonies thst

reproduced earlier in the year, However, the

species appears to reproduce sexually at the

end of summer, Collections from Port lacking,

N.S.W. (Kott 1972c) indicate that there,

although new lohes were being added to the

colonies at the end of August, the species dis-

appeared Curing the summer und did not return

until autumn. Recolonising stock must there

fore exist off Port Hacking, which reproduces

sexually at the endof summer or early autumn,

i.e. a similar seasonal cycle to that occurring at

Elliston Bay.

PATRIDIUM n. gen.

Zooids completely embedded with hoth

apertures opening separately to the exterior

and withoyt colonial systenis, Infernal Jong

ASCIDIANS OF SOUTH AUSTRALIA II

Figs 1. Ritterella herdmania., Larva,

Figs 2-3. Patridiuim pulvinatum. Fig. 2—Portion of branchial sac showing internal longitudinal

vessels, Fig. 3.—Thorax and abdomen -of adult zooid,

Figs 4-6. Aplidiium follorum. Fig. 4—Dorsal aspect of interior portion of thorax showing tripartite

atrial Jip, Fig. 5—Stomach. Fig. 6—Larva.

Fig. 7. Aplidium pronum, Thorax and abdomen of adult zooid.

Fig. 8- Aplidium digitarnim. Portion of colony.

Figs 9-11). = Leptotlinides volvus. Fig. ¥—Colony, Fig. 10.—Thorax,

tudinal vessels are present in the branchial sac.

The stomach is folded, Gonads are present in

the threaddike posterior abdomen, the testis

follicles arranged in a double row and. the

ovary present just anterior to the testes. The

heart is a U-shaped tube at the distal end of the

posterior abdomen.

Remarks: Only two genera of the subfamily

Euherdmaniinde are known in which either

longitudinal vessels or their vestiges are re-

tained in the branchial sac, These genera are

Tylobranchion Herdman, a monotypic endemic

antarctic genus (see Kott 1969), and Pretopo-

lyclinum Millar, in which 3 species are known,

4 PATRICIA KOTT

viz, P, pedunculatum Millar, 1960, trom New

Zealand; P. sabulosa (Millar, 1963), from Poi

Phillip, Victoria; and P. cleviforme Kott. 1963,

from Eden, N.S.W. Tylobranehion retains some

primilive churucters im the presence of the

heart half way down the posterior abdomen,

and a large ovary posleriur to the bunched

Testis follicles. In Prorepolyeliave: the stomach

does not have longitudinal folds and the testis

follicles are bunched in a short posterior ahde-

men ax in Pelvelinun spp, The present gequs

bears the same reli#tionship to A plidiner us

Protapolyctinum bears to Polvelinum. It differs

from both Protepolyclinum and Polyclinum in

the presence of stomach folds and in ity long

thread-like posterior abdomen in’ which the

testis. follicles are arranged in rows; it is these

characters that relate it to Aplicdium. It differs

from Ritterella, also in the subfamily Euberd-

maniinae, in the presence of the longitudinal

vessels, and the absence of parastigmatic Vessels

in the branchial sac, Ritreref/a is usually further

distinguished by the presence of 5 primary rows

of stigmata, although these are often subdivided

by parastigmatic vessels. ‘he restricied mum-

ber of primary rows of stigmuta suggests that

Ritterella may be more specialised than Purri-

dium, which demonstrates primitive affinities

it the presence of a large number of rows of

stigmata as Well as in the retention of the inner

Jonzitudinal vessels,

Patridium pulvinatum n- sp,

Type Loeytion) northern Great Australian

Bicht (32°24'S8, 133°30'R), 42 mm dvep,

S.v04T5, PL Syrnand. Heloivpes SAM,

E 1035.

FIGS 2, 3

Description The holotype only ty available. ty

is a circular cushion, & cm in diameter und 2

ci hegh, mure or less flat topped and with

rounded borders. It appears to have been ses-

sile and attached by & small area in the centre

of the bisul surfuce although there could have

been a short stalk in this position, The test is

very soft and semi-trunsparent, generally with-

out sand or other adherent foreign particles

except for u small sandy ures at one side of the

basal surface. The. zodidsy ure thread Jike, che

thorax und abdomen together are 1.5 im long

and the posterior abdomen about 4 mm. They

open all around the upper surface and the pos-

leclor abdomina project dewn into the centre

of the colony, The apertures are both 6lobed

Fine longitudinal muscle bands extend slong

both sides of the zooid for ils whole length.

‘There are about 25 rows of 16 short, oval, stig

mata; rather tall papillae are present on the

Iransverse yessels ahd these support longitu-

dihal yessels running the whole length of the

branchial sac. The longitudinal vessels are

crowded, being separated from one another by

wn Interval equivalent to the width of about

One anid a half stigmata. The oesophagus is

fairly long and there as a voluminous stomach

bout hulfway down the abdomen with 25 con-

spicuous longitudinal folds, The proximul) purt

of the posterior abdomen does not contain

gonads but (his region is often contracted, The

ovary 15 present just anterior to the double Tow

of testis follicles that occupy the greater part of

the posterior abdomen. ‘I'he heart is a wide U-

shaped utbe in the distal tip of the posterior

abdomen,

Pseudodistoma cereum = Michaelsen,

1972a: 12 (synonymy); 1972b: 173,

New Record: Margaret Brock Reef (Cupe

Jaffa).

Remarks: Specimens in the present collection

Measure up to 12 em of which the pomted or

rounded head represents. half of the total

length, The zooidy, wpening all around the

head, are small, the contracted therax and

abdomen together measuring only 2. mm. (Koti

19724, 2 cm sie).

Kou,

Subfamily Po. veuiNinar

Folyclionm = neptunium — Hartrever.

1972b; 175 (synonymy).

New Record: Investiggtor Strait (Sur ¥11,

Description: The present colonies are small,

with rounded heads only about 3 mm. in dige

meter on thin, branching stulks. Each head

contains about 6 zovidls surrounding a central

common cloacal opening, The test is very deli-

cate, There is no sand iniernally but externally

there is a heavy encrustiution.

The zooids are minute, The atrial lip is

typical of the genus (Mott 1963) and has 4

longitudinal muscle bands, It arises frany above

the upper tim of the opening and appears to

closc down over the apertyre which is prowuced

lo point directly sateriorly, UVhere are 7 musele

bands cacialing From the branchial aperture

hut these do not extend onty the posterior purt

of the thorax, There are 5-8 small oval stige

mati in cach of the 7 rews, and papillac on the

transverse vessels coincide with the stigmata,

The gonads ure not developed The stomach is

sTinott.

Kou.

ASCIDIANS OF SOUTH AUSTRALIA Il 5

Remarks Although the number of muscle

bands in the atrial lip, and the number of rows

of stigmata and the number in each row, are

very much less than that usually reported for

this species, the arrangement of the branchial

papillae is the same as that usually reported

(Kott 1972b), and it is passible that the colo-

nies are jirveniles.

Aplidiom Foliorum n. sp.

Type Location: northern Great Australian

Breht (32°24°S. 133°30'E), 42 m deep, P-

Sverond, 5.v,1973, Holoiype: SAM, E 1036.

FIGS +4

Deseripiion: The colony ts a circular cushion

6 ¢m in diameter and 2 em high, forming a low

dome, slightly concave basally where the zooid

hearimy surface layer of the test on one side has

grown atound énto the busul surface to form

a crescent shaped pockel invaginated towards

the border af the colony where the surface

zooid bearing layer of test has grown to over-

lap it. The test is soft, gelatinous atid semi-

transparent, There are about & common cloacal

apertures scattered over the surface of the

colony, about 1.5 to 2 em apart. Canals radiace

aut from the openings, lined on each side by

raws of zonids, These radiating canals sub-

divide many times und zooids lining them on

each side crowd the test, Zootds are at right

angles to the upper surface.

The branchial aperture is terminal with the

opening surrounded by a circular sphincter.

The atrial lip rising fram the upper border of

the aperture is very Variable and aay be simple

or tripartite. while the Jobes may be large and

foliate or small und pointed, There is a band

of muscles down the centre of each atrial lobe,

There are 14 raws of about 15 sligmata, The

stomach is large with 18 to 25 narrow longi-

tudinal folds,

The zoouls are long and threuul-like, the pos-

terior abdomen comprising the great part of

their length while the thorax und abdomen to-

gether are only 2 mm long, There are one or

twe embryos at very different stages of develop-

ment in a brood pouch that is formed by an

expansion of the distal end of the oviduct at

the postero-dorsal end of the thorax, Dense

testis follicles are present in two rows in the

posterior abdomen, The ovary is present antc-

nov to the testis (in the usual position for this

penis).

Larvens The larvae are 0.75 mm long with a

long lail that completely encircles the body.

There are the usual three anterior papillae

alternating with two median ampullae, Numer-

ous ampullary vesicles rise from the lateral

ridges extending anteriorly along both sides oF

the endostyle and to the post vential aspect of

the larval body.

Remarks: The species is distinguished trom

Anplidium pliciferunt by the larger size of the

eclony. the very distinct radinting double raws

af yooids which comprise the systems, and by

the characteristic foliaccous musculur atrial

johes that are present on many of the zooids,

The colony does resemble that of A. austra-

lienvixy which has similar systems and in which

the branchial sac is the same, In A astra

liensix, however, there is a lesser nUmber of

stomach folds and they are sometimes irregular

and oblique, while the zooids lack the distinc-

tive almal lobes of the present species. The

larvae of A. pliciferwm, A. australiensis and the

present species arc, however, identical. There

ure slight variations in size (¢g, larvae from

the holotype of A. ausiraliensiy are 0.9 mm

long) and in the length of the tail which

extends from half to the entire distance around

the body. However, the relationship of the

length of the tail to the larval body does not

appear to be constant for any single species.

The characteristic atrial lobes of the present

form nre similar to those described for A,

cletum Kott, 1972bh, which however dillers con-

siderably from the present specimen iv colony

form,

Aplidium flavolineatum (Sluiter) Kott, 1972h;

176 (synonymy).

New Record: morthem Great Australian

Bight.

Dexeriptiony The colony is mushroum shaped,

4 cm in diameter across the flat upper surface

and 2 om high. The fattenes! zoom-bearing

head narrows very suddenly to a short stalk

from the centre of the under surface, Sand is

present on the stalk and, to a lesser extent, on

the upper surface. The test is clear and glassy

but Soft, The zooids are crowded in the test

and it was mot possible 1o distinguish the form

of the systems, Zonids open only onto the

upper surface. They are 6 mm tong. of which

the thorax is 2 mm and abdomen only J mm.

The posterior abdomina cross one jnother in

the internal test although the thoraces are

parallel at the surface. ‘The atrial Jip is divided

into 3 very pointed lohes From the upper border

uf the opening, There are 12 fine longitudinal

muscles on the thorax, There ate 12 rows of

10 Jong rectangular stigmata. The stomach ts

6 PATRICIA KUTT

especially small with 17 distinct Jongitudinal

folds. ‘Vhere ire 2 rows of testis follicles in the

posterior abdomen.

Larvae: Up to 3 developing embryus are pre-

sent in the atrial cavity, They are 0.75 min

long, have 3 median papillae alternating with

single median umpullac and corresponding

laicral ampullac develop from the lateral ridge.

The median ampullac are narrow and in some

cases appear to be bifurcated. “There are also

clusters of ampullary vesicles both above the

endostvle und ventral jo the body of the larva.

The tail winds about three quarters of the way

uround the body of the larva,

Remarks: The colony and zooids are of similar

form to these deseribed previously far A, flave-

lindativa With the exception of the stomach in

which there ate only 17 folds. The sive and

shape of the stomuch and the course of the

longitudinal folds are similar to those described

previously for this species.

Aplidium coniferum Kott, 1963; 102.

New Records. Elliston Reef. Previous

Records: N.S,W, (near Twofoll Bay, 10-70

m deep; Montague North, 13 m decp)—Kou

1963,

Deseription. Sessile, rounded lobes ubout 4 em

in greatest diameter. Che lest is sindy inter-

nally, but the external layer of test is Pree of

sand and is smooth and gelatinous. Zooids ure

long and narrow and open all around the head.

There is # small pointed atrial lip from the

body wall anterior to 4 muscular siphon that is

preseiit whoul one third of the distance down

the dorsal surface. The thorax ts. long and

narrow with ubout 15 rows of 10 stigmata.

There are 5 stomach folds.

Remarks: Vhe specimen agrees with those pre-

viously described. ‘The clear external layer of

fest without sand and the form of the colony

are apparently charactenstic of the species.

Aplidium amorphatum Kott, 1963; 101.

New Record; Elliston Bay. Previany Record:

Vie. (38"51/S8, 144°55'E1—Kor 1963,

Deveriprion; The colony is soft. sessile and

deme shaped. ‘Vhe test is semi-transparent,

wilhaut sand. Zooids open all around dhe upper

suiface und no systems ure evident. The zonids

are very small and irregularly orented in the

lest So that they cross one another. There ure

[0 Jongitudinul thoracie muscles. The atrial

aperture is on a short sipham. ‘The upper rim

ef the aperture ws produced into a pointed lip.

There are 12 rows of about It stizmatn, Each

Tow is crossed by a pprastigmatic vessel, The

stomach is small with 5 folds.

Larvae: The atrial cavity is occupied by a single

lurge embryo. | mn long. [t hos the osual 3

mediun, stalked, papillae and numerous ampul-

lary vesicles are developed [rom the anterior

part of the bady-

Remarks: The specimen is identical with thit

previously described for the species. ‘The Jarva

is the same us thal of 4. paatherinum (see Kott

1963). The shape und consistency of the

colony differ, however, and resemble A. pri-

tecrans (Herdinan)); Kott 1443, fn the latter

species, however, the zooids ute larger, the

branchial sue larger, there are more thoracic

muscle bands and the parasusmalic vessels are

not present,

Aplidium pronumt cr, sp,

Type Location: Investigator Sirun (Sin X1),

Som deep, Marsan, Jan. 1971, Heloppe:

NMV. H 287.

FIG. 7

Deserinption: The colony consists ef small, Mhu-

topped lobes united basally. The lest is sofc anil

there is very little sand internally. There is

single common cloacal aperture in the centre

of each lobe, The zooids are more than 7 cm

long and thread-like. “The thorax and abdomen

ure of equal length. and about one third of the

total length of the zooid, The atrial lips is bifid

or trifd terminally and extends from the upper

border of the alrial opening which is on a shert

siphon, There are 1L rows of 12 stigmata and

8 very weak stomach folds, There are 20 longi-

tudinal thoracic muscles.

Remarks; The colonies resemble those of 34.

noveerelandine Brewin and 4, corrrelli (Brewin)

irom New Zealand, although the Jlat-topped

lobes. Brewin (1952, 1957 respectively) des-

¢ribed for both these species have several sys-

tems, oly S stomach folds, and the trial

siphon is not produced, The atrial siphon of A.

Mlaritiocen (Brewin, 19589) is, in Eaet. pro-

duced in the same way as in the present.

species. and the stomach has the same ill-

defined folds. However, the longitudinal tho-

Tacic muscles ate more pleotiful in the present

specimens, there are fewer stigmata in gach

tow, the pusterior pbdomen is longer and more

thread-like, and there ts only a single system in

each lobe. Burther, in Brewin’s species the

Jobes are separate and do not appear to. he eon-

ASCIDIANS OF SOUTH AUSTRALIA TIE 7

lijuous jm their basal half as m the present

species.

Aplidiun digitatum n. 5).

Type Cecativn: northern Greac Australiall

Bight (32°24'5, 133°30°R), 49 m deep,

5.v,1973. & Symond. Holotype: SAM, E

1030. Pararypes: QM, G 7508; AM, ¥

1982,

FIG, 8

Description; The colonies are long, branched,

cylindrical stalks 2-3 cm long, terminally

rounded and slightly expanded ta overlap the

stalk. The zooids open onto these terminal

expansions, Sand is absent only from the sur-

fuce fest where the vooids opett on the

expanded terminal portion of the lobes, ‘The

stalk is densely encrusted with sani. The test

is firm, especially in the stalk. und is impreg-

nated with sand throughout. Zooids are minute,

hut long and thread-like, crowded in the test

and exteading parallel to one another down the

slulk.. There are 15 rows of wbour § stigmata.

There is a long, pointed, atrial hp from the

upper border of the opening. The stomach has

12 longitudinal folds. There is a double row of

testis follicles in the long posterior abdomen.

Phere is a large common eluacul opening in the

ceuire af each lobe.

Remarks: ‘Tne comnivs resemble those of the

Antarctic species Aplidiwn recumbens; Kor,

1969, bul are distinguished by the large num-

her of distinct longitudinal folds in the sto-

mach. The zooids are especially delicate and

narrow,

Aplidium colelloides (Flerdman). Kott, 1972b:

176 (synonymy).

New Record: northern Great

Bight.

Australian

Family DIDEMNIDAE

Polysyacraton aspiculatum ‘Tokioka, 1949: 2.

Kott, 1962: 301.

Polvsyneruton magnilarvun Millar, 196: 13

incrgiert nudum); 1962: 165, Kott, 1972b;

178,

New Records: morthern Great Austvalian

Bight; ?investigaror Strait. Previous Records:

W. Aust (Rottnest T, I Peronj—Kou

1962, S. Aust. (Investigator Straiti—Kolt

1972b. Qld (Mackav}J—Kott 1962. 8. Africa

— Millar 1962. Mozambique—Millar 196),

Japan—Tykioka 194%,

Deserinvion. The colony from the Great Aus-

tralian Bight is a soft jellylike cushien, The

common cloucal system consists of narraw

canals at ocsophagea! level, rudiiting from

common cloucul apertures and Jined on either

side by zooids, There are no spicules, The

zooids. ure of moderate size. with 4 rows of

stigmata and a long bifureated atrial tongue

The oesophageal neck is long. Gonads are not

mature.

The colony from Investigator Strait (that is

doubtfully. assigned to this species) consists of

2 Jarge flattened lobes rising from a fleshy

evlindrical common basal stalk, The zoows are

embedded in the surlace layer of test. The com-

mon cloaca! canals extend between clumps of

zocids beneath an especially thin layer of sur-

face esl. There ure very extensive cloacal

spaces between the central, soft test that forms

the central core of cach lobe and the surfuce

that is only occasionally joined with the central

core hy solid test connectives. Secondary canals

extend between the zooids beneath ow very chin

layer of surface test, The test is colourless and

transparent. and the zooids show through it as

white duts, There are no spicules. Zooids have

a long oexophugeal neck, and the usual 4 rews

of stigmata. Gonads are not mature in this

colony.

Remarks. Specimens assigned to both P. aspi-

cularunt Tokioka, 1949 and Poo mageilarvene

Millar, 1962, are soft and rounded, stalked or

segsile, from 3 to 7 mm thick, with varighle

spicule disttibulion 10 an uspiculaur condition.

The. atrial lip 1 Jong, and often spread or biftr-

cale at its tip. There ate also. im both. a large

number of testis follicles (8-12) and large lar

vae (over 1 mm long) with up to 19 pairs of

lateral, finger-like ampullac and precocious

buds, Zooids af both species are alu charac-

terised by w long oesophageal neck. In Austra-

lian specimens the ventral surfaces are em-

bedded in the common test and they are

arranged along both sides of common cloucal

Cunals that demareate rounded zooid-free swell-

ings of the surface af the colony, In neither

species has the common eloaca been described

as posterior-abdominal, The shape of the pre-

sent colony from Investigator Strait is identical

with others. from this area that are assigned to

P. aspiculatunt (> P. omagnilarvurn; Kote

1972b). The posterior abdominal cavities tn

this colony, however, do not oceur in those pre-

viously described, Positive identification is nut

possible uwing to the lack of maiure gonads,

and the felationship of the extensive cloacal

system with the simple canals that have been

8 PATRICIA KOTT

Previously described is not known. The cloacal

system is the swme as that of Didemniunr fam-

bine and Pelvsyneraton chondritla but both

these species hive only a single aperture and

are nol known without spicules,

Leptoclinides yolvus 11. sp,

Type Lecagiion: northern Great Australian

Bight (32°24’S, 133°30°E), 42 m deep. P.

Symand, Holotype: SAM, E1034. Para-

wwresy SAM, F 1033; QM, GT75I1L

FIGS 9. 10

Description; ‘The specimeh designated as the

holotype ig a, flattened sphere 5.0 em in din-

meter and 3 em thick, wih @ thick but very

short stalk, constricted to 2 em in diameter

where it joing the body, The paratypes arc

enlively spherical, about 3 em in diameter, and

are without stalks although the base of ihe

colony is identified by the absence of zoids

and by foreign maller that is included in the

test material which has overgrown the ared.,

There ik a single, apical, sessile, and inconspi-

cuous common cloucal aperture, opposite the

base of the colony at the junction of several

common cloucal canals. Zooids are arranged

along both sides of parrow cloveal canals at the

abdominal level of the zooids and the surface

of the colony is depressed above these canals

These depressions demarcate rounded swellings

at the surfice of the test corresponding to

yodidelree areas The tose is very firm gela-

fines and translucent. There is 4 layer of blud-

der cells superficially, A sparse layer of spi-

Cilles is presept in the zonid Jayer of test and

these are most dense around the zonids, |hus

indicating their position through the test, The

spicules gre minute, 0,015 ta 0,02 mi in aia

meter, Ome stellate and others with needje-

like rays, There are minute, spherical. brown,

Pigment cells scattered throughout the test

Zooids are small, with about 8 stigmata per

row. The branchial siphon is of maderate

Jength, The atrial siphon js very short, casing

from the mid dorsiim, opposite the space

between the second and third rows of stigmata,

It ts surrounded by & circular sphineter muscle

and is. posteriorly or laterally directed. There

are 53 coils of the yas deferens and up to 10

testis follicles. Large ova are present in the test

at abdominal level but none of these appear to

be developing embryos.

Remarks: The spherical hody, constricted stalk

and single common clovesal Opening are unusual

in this genus, The lack of stalk in the two para.

types, together with the spherical! shape, suggest

that these colonies may be free living, although

the foreign particles that are embedded in the

basal region suggests that this part of the

colony was fixed to the substrate, hroke free

and wis overgrown by the surface test. The

constriction where the broadening stalk joins

the head in the holotype also supports the sug-

gestion thar the spherical head may break

away. Certainly the configuration of the sur-

face, with the projecting swellings of solid gelu-

tinous test, while the zooids, their openings, ynd

the common cloucal aperture are depressed into

the surface of the test, would all agcommadate

A free living habit in which the colony is able

to roll over (he sea floor as in some coral spe-

vies (see Glynn 1974, Pichon 1974),

The limited nature of the common cloacal

system Is unusual in this genus where extensive

postenor abdominal spoces are usually deve-

foped. It differs from species ia other genera,

in which the godids are urranged along both

sides of nantuw common cloacal canals that

extend around citcular zooid-free areas, in that

the canals gre at the abdominal rather than the

thoracic level (see Polysyncraron aspieularany,

ubove: and Didermrnim patulun; Kot 19724),

Stulked species are also unusual in this genus,

L. fungiformis Kott, 1972b being the other that

is known. It is distinguished! from the present

species by tts undivided testis folligle

Leptoclinides reticulatus (Sluiler). Kott, 1062:

285 (synonymy); 1972ac bw. 1Y7tb 180

New Reeord: northern Great Austrolian

Bight.

Deseriplion> A latee colony. investing a speci-

men of Herdmania momuy. There are streaks

of orange-brown, stellate, pigment cells scat-

tered amongst the spicules i) the surfyce test:

The spicules have 7 rays in optical irunsverse

section und are 0.03 to 0.05 mm in diameter,

There ix a superficial layer of bladder cells

mixed with spicules. Spieules ute dense at the

zooid level but are absent basally. below ab-

dominalt level. Common cloacal canals were

net feund and the thoraces of the zovids

appeared to be purtially distintegrated although

the abdomina were in good condition with 5+

coils of the vas deferens aruund 5 fo & testis

follicles. Small vegetative buds are present in

the oesophageal region.

Remarks; The absence of common cloacal

cxumuly aud the condition of the zodids suggests

that the calonies may preseat a yunescent winter

ASCIDIANS OF SOUTH AUSTRALIA Hl 9

coadition. The presence of venetative buds and

mature vonads suggests the onset rather than

the end of this quiescent phase,

Leptoclinides rufus (Sluiter). Kott, 1962: 286

(synonymy); Eldredge, 1967: 221.

Lepinelinides Nssus; Millar, 1963: 704,

New Records: Elliston Bay (outside bar. and

25 mi deep), Prepious Records: 5S. Aust

(Port Noarlunga). Tas. (Maris L). Vis,

¢(Shotvhain)—Kott 1962. NSW. (Port

Jackson—Kite 1962, Millar 1963. Old (Bar-

wara, Heron J,, Low ([sles)—Hhastings 1931;

Kott 1962. Indonesia (Paternoster 1, Ara-

fura Sev)—Stuiier 1909, 1973; Tokioka

1952, New Zeuland (North Island)—

Michaelsen 1924: Brewin 1958b; Millar

1960. Hawaii (Qahu)—Eldredge 1967.

Description. Living colonies are reddish brown

or grey with orange around the siphons. They

ure firm and investing, with common cloacul

“penings in the cetlre of evenly spaced

rounded swellings. “he common cloacal aper-

tures are about 1 om distant from one another.

There is a surface layer of bladder celle and

spicules ate especially dense below this layer.

They gradually becoine less dense and are

absent altogether from the basal half of the

colony, Although they wre present ina layer at

the base of the common clogeal cavity, they are

stellate, 0.02 1o 0,04 mm jn diameter with

pomted conical rays. The spicules arc cspeci-

ally densely accumulated around lobes of the

branchial apertures.

The common cloacal cavity is posterior

abdominal, extending imto circular chambers

heeath the common cloacal apertures.

The vooids gre about 2 mm long with up ta

12 Jongitudinal thoracic. muscle bands from

which fibres branch and wnastomose with adja-

cent bands. There is # minute cireular lateral

organ opposite the 4th row of sligmuta, The

atrial aperture is directed posteriorly as is usual

for the genus. There ure up to LO stigmata in

each row. The oesophagus is long. The gut

loop may be curved, although tL also occurs

ws a long straight loop. The gut is clearly dif-

ferentiated into duodenal and mid intestinal

regions and a posterior slomach swelling, The

stomach is smooth and rounded, Jonger than

its diameter. There are 9 to 11) testis follicics

and 74 coils of the vas deferens.

Remarks; The synanymy of ©, fisvus Hastings,

from Low tsles. with 2, ruéas originally des-

eribed from Indonesia and the Arafura Sea to

the orth. within the same bhioyeogriphical

region, wus arrived al after comparison of

Hustings type material (AM, G 13449) with

other specimens from a wide range along the

southern and eastern Australian coast. includ-

ing the Great Barrier Reef and the Queensland

mainland. The type specimen of L. diemenensis

has not been examined, although ‘Tokioks

(1952), Millar (1960). and Kott (1962) hive

not been able to identify any character that

could distinguish the two species, Its synonymy

with L. rejee is here maintained, The range of

t, rafus is, therefore, similar to that of many

wide iutging species of this family in both

tropical waters (Ketl 1974) and in Antaretic

waters ¢(Kott LY69a). The other related New

Zealand species, L. stutirert, L. auranticus and

L. nevae-selandiae, are distinguished only hy

the larger number of coils of vas deferens, the

relatively shorter oesophagus and the smaller

zooids, Further specimens are needed ta ude

quately determine the parameters of each of

these species. £. rvefay is characterised by its

relatively large zooids with distinct longitudinal

thorucic musculuture buy no reactor muscle;

by its long oesophagus and gul. Joop and the

clearly demurcaled gut regions; by its long and

muscular siphons; by the invasion of spicules

into the superficial bladder cell layer and by

the position of the small lateral organ in the

posterior part of the thorax. ‘The number of

testis lobes in the present specimens is greater

than the maximum of 7 previously recorded.

Didemnum candidum Savigny, Kott, 19724: 1%

(synonymy); 1972h: 179,

New Revord: northern Great

Bight.

Didemnum moseleyi (Herdman), Kore, 19720:

19 (synonymy), 19726: 179,

New Record: northern Great Australian

Bight,

Australian

Trididemmum savignli (Herdman) s. sp. savignii

Merdman, Kott, (966: 285) (synonymy);

Eldredge L967: 179.

New Record: Sellick Beach (south of Ade-

lutde), S, Aust.

Descripiien: Extensive investing colony with

round. smooth, margins, There is a spicule-lree

surface layer of bladder cells. Spicules are

sparse in the zooid layer und are ohsent alt

vether Prt the basal half of the colony. The

eommion cloacal canals are deep, exlending the

whale length of the zooids, but they are uot

in PATRICIA KOTT

posterior abdominal, The spicules ure 0.04—

0.08 mim in diwmeter, They are stellate with 12

conjeul rays in optical transverse section. The

zonuls are surrounded by black pigment par-

ticles that are often but not always accumulated

inte the usual pigment patch at the anterior end

of the endostyle. There is a distinct atrial

siphon which is laterally rather than posteriorly

directed in these colonies. There js a distinct

Tetractor muscle. The testis is not mature and

the vas deferens was nol distinguished,

Remurks: No further evidence is available from

the examination of these specimens that could

clarify the relationship between this Indo

Pacific subspecics and the Atlantic Ocean form

T. savignit subsp, afrocumunt Van Name (sec

Rott 1966). tt should be noted that the Pucilic

Ocean specimens (Eldredge 1967) have the

7-8 coils ul the yas deferens that 1% associated

with the Indo-Pacific form (Kou 1966).

Trididemnum cerebriforme Hartmeyer. Kott,

1972e: 47 (synonymy).

Trididemananm savignii; Tokioke,

vii, jolenye: B2.

New Record: Sellick Beach (south of Adce-

Iuide), S Aust,

Description: The colonies ure investing and of

vaniuble thickness. Conspicuous common

cloacal apertures with frilled lips are distri-

buted randomly over the surface of the colony,

Posterior abdominal cloacal canals radiate

from these apertures, Spicules are sparse in the

upper layer of test and apart from a layer lining

the test along the floor of the comman cloacal

cavity, they are entirely absent from the basal

layer of test. The spicules are stellate, 0.02 to

OAM mm in diameter, with about 6 rays in

optical transverse section.

The hoily wall of the thorax 1s covered with

bluck pigment purticles although these are

ubsent from the abdomen. The pigment par-

ticles ure uccunulated in 4 patch over the

anteriar end of the endastyle, The atrial siphon

is posteriorly difected. There are 64 coils of

the vas deferens around an undivided testis

follicle.

1967: Bt;

Remarks: The relattonship of this species to 7,

saved is perplexing since its three-dimensional

cummon cloacal system provides the principal

distinction, 7, eerebriforme acquires ereat cam-

pleaity in its common cloacal system with

growth. bur juvenile colonics must necessarily

display a cluacal system identical wirh thar of

T. savignii before its subsequen proliferation,

as the colony thickens and surtace folds deve-

lop. In Lhe present specimens, both taken From

Selick Beach, each species has spicules of dif-

fetent sizes although this size difference wis

not, observed in specimens previously described.

‘The number of spicule cays as previously

reported, however, iy greater for T. saviprii

than for T. cerebyiforine,

Diplosoma translacidum (Harimever)

Leploclinem — (Leytectinum)

Rou, 1962: 306 (synonymy),

New Record: Investigator Strait (Stn X14.

Previous Records: Indonesia—Sluiter 1909,

North Western Australia—-Hartmeyer 1919.

W. Aust, (Oyster Harhour, Albany)—Kott

1962.

Description: The culony is irregularly lobed,

investitig weed or other ascidians.. Bach lobe is

flattened, about 2 cm wide, 0.5 cm thick and

up to 3 cm Jong. One calony completely enve-

lopes u specimen of Pyura australis in whieh

only the apertures are exposed, The surface is

amooth. und the zooids show through as white

dots, They are small and crowded at the sur

Face of the colony in groups of ahout 8,

There is u Jarge Ovum present in most zooids

but the testis is not mature and no coils of the

vas deferens were detected,

Remarks: The tough, firey transparent test wil

the extensive cloacal system ts churacteristic of

this species.

translncidam:

Family ASCIDITDAE

Ascidia thompsoni Kott> 1972a: 27 (synony-

my); 1972b: 181,

New Records: upper Spencer Gulf,

Description: Specimens have a gelatinous test,

somnetimes thick and furrowed, Both apertiinss

are present on siphons, usuyily both directed

dorsally. or anteriorly and sometimes very long.

The animal is fixed ventrally and by most of

the lel! side. There is sometimes a coating of

sand cnerusting the boty, hut sand is never pre-

sent on the siphons. The body wall has the

usual meshwork ol muscles on the right side

of the body. The anterior part. of the dorsal

lamina is a double membrane. ribbed on the

outer sides but not tn the centre. ‘The ribs of

the dorsal lamina extend into pointed projec-

tions on the free edge of the membrane.

The neural gonglion is about one-third of

the body length from the dursal tubercle.

Remarks; The specimens from Station D3 on

the Nuor ol he channel are encrusted with a

ASCIDIANS OF SOUTH AUSTRALIA TIT ial

layer of sand ahsent only from the siphons

which project upward from the middle of the

upper surtace. In this specimen, the right side

nf the body ts natrower than the left and cam-

prises the tight side of the upper surface, while

the hase, by which the animul is. fixed, is the

ventral surface and a large part of the lefi side

of the body. In specimens from G4, also on the

floor of the channel. the siphons are especiully

long, both directed upwards, anil the right side

of the body is similarly short,

The very long external siphons’ directed up-

wards, und the sand encrustation, are unusual

in this species. Avcidia aclara is the only spe-

cies of the genus in which a similar sand en-

erustation hardens the test. The long, cylin«

drical extensions of ihe tost that, in 4. aelara,

ervate a candl or tube from the sessile apere

tures extending upwards from the animal, are

dmalogues rather than the homologues of the

long siphons in the present forms. It is also of

interest that, in specimens from stations D3

and K4, where the siphons extend dorsally,

their central position, from the middle of the

upper surface of the hody, is achieved hy rela-

live nurrowing (i.e. between the doy'sal lamina

and the endostyle) of the right side of the

hody; whereas in A. aclare it is the left side

of the body that is natrawer thin the right. The

base of [he present specimens is the ventral and

two-thirds of the left surfaces of the body,

while in 4. ¢elara it is two-thirds of the might

side. between the dorsal lamina and endostyle

(Kott [972d),

‘The specimens appeat to have adapied to

their free existence on a shifting sandy sea

floor by these morphological variations in the

position and length of the siphans.

Ascidia aclara Kott; (972d: 236 (synonymy).

New Reeords Goat 1 (off Ceduna).

Family STYELIDAE

Stolonica carnosa Millar, 1963; 734, Kott,

1Y72a; 28

New Record: Unwestigator Strait (Stn YT).

Remarks: The present specimens are small and

sandy individuals joined by stolons. There are

only 3 stigmata per mesh and the longitudinal

vessels in the branchial sac are slightly fewer

ihan previously reported, viz. DLO(5)2(4)1E,

It is probable that this is a young colony, which

is characterised by a pyriform stomach with

narrow folds and a Jong curved caecum that

extends into the gut loop rom the suture along

the lateral aspect of the stomach,

Amphicarpa diptyehu (Hartmeyer). Kott, 1972¢

(synonymy).

New Reeord>

Bight

Oculinaria «aistralis Gray, Kott,

(synonymy); Kolt, I972b: 184.

New Reevard: Elliston Bay.

Symplegma viride Herdman, Kott, 1952; 252

(synonymy); 1962; 129, Millar, 1946; 268.

Plante & Vasseur, 1966: L49. 'Tokioka,

nonhem Great Ausitraliin

|972a: 24

197: 162 (synonymy). Visseur, 1967;

ith.

New Record: Elliston Reef.

Subfumily BOTRYLLINAE

PARABOTRYLLUS no. gen.

Colonies sre elongate branching = stalks

slightly expanded terminally. One to 3 circular

systems of Zooids are present in each terminal

expansion, opening onto a more or less flat-

tened surface of each free lobe. Each system of

zooids surrounds a central common cloacal

aperture, The terminal umpullae and conspi-

culous blood vaseulur system that are present in

other genera of this subfamily are absent. The

tim of the branchial aperture is smooth. The

atrial aperture has a single anterior lip, There

ure only 2 internal longitudinal vessels, in the

branchial sac, Eggs are endogenous,

Remarks; The zooids are not conspicuously

different from thase of the genera Borryiius and

Borryiloides except in the presence of only 2

internal longitudinal vessels in the branchial

suc. The colony differs considerably. however,

both in its shape, and in the presence of only

one to three systems opening onto the flat ter-

minal surface of each Jobe, Buds are present in

the common test near the posterior region of

the adult zooids, to which they are joined by

narrow connectives from the oesophageal

region of the parent. The buds in this genus

therefore do nol, apparently, maintain a close

conection with the parent until a late stage

in their development as they do in other genera

of the sub-family. The zooids wre endogenous

and have three to four developing ova on each

side as in Borryllus, while Borrylloides produce

only single ova on each side of the body,

Parabotryllus nemorus n. sp.

Type Location: upper Spencer Gulf (Stn),

9 my deep. floor of channel, 5.ix,1973. Holo-

types SAM, E1031. Paratypes; AM, Y¥ 19815

OM, G 7507.

FIGS 11-15

Description; The calonies. consist of narrow,

sandy lobes, 1-1.5 em long, usually branched.

‘The superficial layer of test is encrusted with

sand but is neither stiff nor brittle. Internally

sund is absent und the test is very soft. There

are circular systems of zooids opening onto the

upper free end of the lobes surrounding the

central common cloacgl apertures that are

slightly depressed into the surface. Generally:

there is only a single system in cach terminal

branch of the colony although occasionally

ihere are 1 or 2 smaller additional systems. The

blood vessels in the lest are short and relutiyely

tew for this subfamily. They terminate in elon-

gate tounded bulbs at the base of the zooids,

The zovids arc about 2 mm long. The rim of

the branchial aperture is smooth and the atrial

z PATRICIA KOI

aperture. in the antenor third of the dorsal sur

face, has its upper lip produced into a single

pointed lip. The atrial aperture is very small

and is directed anteriorly so that the upper lip

Closes over it (#8 in Polyclinvin), There are

10 rows. of stigmata with about 10 stigmata in

each row und two interngl longitudinal vessels

on each side of the body. The gut forms a

single light loop on the left side of the bran-

chial sac, The stomach is pyritorm with about

8 distinct longitudinal folds and there is a short

curved caccum, of moderate length and ex-

panded into a terminal bulb, from the pyloric

end of the stomuch. There is » connective ex-

tending from the pyloric region of the stomach

to the intestine. There is a single, flat. testis

follicle with lobed margins on cach side oat

the branchial sac just anterior ta the gut loop.

Parabotryllus nemorus. Fig. 11—Portion of colony showing branching stalks. Fig, 12—

Adult zooid. Fig. 13—Bud (Jaterai aspect) showing connecting vessel. Fig. 14—Bud

Fies 11-15.

(dorsal aspect) showing endogeneous ova. Fig. 15.—Testis, ;

Fre, 16. Polycarpa tinclor, Aberrant individnal with atrial siphon produced anivrinrty,

Fig. 17. Pyura tendata. Section through the body wall, test and sandy cauting.

Fig. 18..

Microvesmus planns, Gut and gonad on leftoside of the bady.

ASCIDIANS OF SOUTH AUSTRALIA TT 13

The vas deferens, arising from the middle of

the mesial surface of the testw. is very xhort.

There are three or four ova in the body wall

anterior 10 the testis lobe, These are endo-

genous and project into the peribranchwl

cavity. Developing buds. ure present in the teyt

on either side of the posterior end of the adult

zooids. These contain Cour large Ovi ot cuch

side of the body and a clump of large ¢clls

dorsal to the ova. Jt is possible that these may

he precociously differentiating buds. The buds

at this slage of development are 0.25-0.5 mm

long. There is a blood vessel extending from the

posteriot end of each bud in the region of the

oesophagus.

Remarks: The species is distinguished fram

others in the sublanvily by the large number of

ova und internal longitudinal branchial vessels,

the reliutively limited blood vascular system in

the test. und the form of the colony and the

limited development of colonial systems, The

internal test iy also very soft in comparison

with thar of other species in the subfamily, The

buds appear tu undergo the major part af their

development in the test in connection with the

colonial blood vascular system and im the pre-

sent colonies there were to buds found clireetly

associated with the parent zooids, The testis

Of this Species: is reminiscent of that in Syin-

flegina, while the multiplicity of ova resemble

Heteyifus and ihe endogenous nature uf their

development and the small circular systems in

Ihe colony ate also reminiscent of the latter

genus.

Subfamily sirvetJnau

Polycarpa tinetor (Quoy & Gaimard). Katt,

1964: 134 (synonymy), 197Zh: 186)

[972e: 284; L972d: 242.

New Record; upper Spencer Gulf {Stn

Bi}.

FIG. |6

Remarks: One of the specimens is highly modi-

fied. It has the usual short branchial siphon

from the ynterior end of the body, Vhe atrial

siphon, however, extends anteriorly from the

posterior half of the dorsal surface parallel with

the anterior half of the body and opens at a

point more or less level with the branchial

aperture, so that the individual fxs U-shaped.

The lower half of the body ts encrusted with

large particles Of gamd but the upper half has

only fine sand encrusting it, and it appears that

the animal had been half buried tn the floor of

the channel and that the atrial siphon was pro-

duced upwards so that it opened above the sta

floor, In another spezimen there are long root-

like processes from the ventral border of the

body, which is otherwise typical of the species

(Fig. 16), In the U-shaped specimen there ts a

single row (17) of lony polycarps around the

veniral border of the body and only occasional

pul¥curps, tepresenting a second row. scattered

dorsal to these,

Polycurpa pedunculata (Helles). Kott, 1972u;

35 (synonymy); 1972b; 186,

New Recordy; upper Spencer Gulf; nerthern

Great Australian Bight; Investivator Strait,

For Previous Records, Description, see Kott

19724, 1972b,

Remarks: From the upper part of Spencer

Gulf, arenaccous and naked specimens are

luken, sOmelimes growing side by side attached

to the same shell or stone. ‘There are a lyrge

number of specimens and they are either

stalked or sessile, Naked specimens with a

leathery test were a bright yellow colour in life

but pinkish in preservative. Living arenaceois

specimens were a sandy colour with a reddish

linge. Vhere are small, smooth, black. indivi-

duals in the preserved material and some that

are larger with reugh Jeathery and rather thin

test. In nature such specimens with a smooth

fest are bright yellow (S. A. Shepherd, pers.

comm.)

Polycarpa paplilata (Stuiter}, Kutl, [972a> 24

(synonymy),

New Recerd> upper Spencer Gulf (Stn B7).

RenidAss 2 specimens only are available, One

is juvenile with a whitish coloured test und

without developed, polycarps. The oiher mature

individual has a row of eight long polycarps

avound the ventral aspect of the left bady wall.

with the ducts directed towards the atrial opet-

ing. There are five rounded anal lobes slightly

bifurcated and only two polycarps are present

in the middle of the tight body wall. The speci-

men otherwise conforms with previous des-

criptions of this spectes.

Styela pedata (Herdman), Kott. 1972b; 185

fsynonymy}.

New Records; northern Great Australian

Bight.

Styela plicata [Leseuer), Kott, 1972b; L&5-

1972e; 254; 1972d: 239 (synonymy). Te-

kigka, 1967. Abbott & Johnson, 1972; 95.

New Revord> Poyt River (St Vincent Gull).

4 PATRICIA. KOTT

Remarks: The individuals are small and the

rounded swellings of the test are obscured by

epiphytic growth. However, the species is

rendily distinguished by the short vasa elleren-

tla, branched testis follices, undulating (some-

times branched) ovarian tubes, long oesu-

phagus, long rectum, deep secondary gut loop,

long voluminous stomach with internal folds

and the small leof-like endocarps that cover

the body wall! and the gut loop distal to the

stomach,

The gut loop is remmiscernt of that of &.

camificara (Kott 1972d) although the rectum

and oesophagus are longer in the present spe-

cies; and although the gonads tesemble those

ofS, partita (Stimpson), in the present spe-

cies the ocxophagus is shorter and the vasa

eflerentia are shorter (Vasseur 1967),

Cnemiducarpa etheridgli (Hercdiman) Kutt,

L972: SL (synonymy); 1972c: 253,

New Record: northern Great Austratian

Bight.

Family PYURIDAE

Pyura tendata Kort, 1972h: 186,

New Record: south of Goat 1, (off Ceduni),

FIG. 17

Description: A single specimen only ig avail-

able, It is more or Iess a half circle in outline,

one cm in diameter, The external siphons ex-

tend from the anterior and posterior ends of

the more or Jess straight dorsal surface. The

branchial aperture is directed anterodorsally

and the atrial aperture is directed posterodor-

sally, he external test is covered by a thick

coating of sand held in place by hairtike exten-

sins from the test. There is a thin space

beeween the sandy coating and the surface of

the test, traversed only by the base of the test

heirs. There is also a coating of very fine sand

on the surface of the test itself. The apertures

ire lined with a very tough invagination of the

test. The branchial siphon is expecially mus-

cular and sppears to be eversihle. The body

Wall is ulso very muscular. The atrial siphon is

muscular but not eversible and its aperture is

protected by a well developed velum at the

distal end of the siphon. Beyond this velum the

test is produced into u cylindrical fibrous

extension for a short distance, There are seven

branchial folds on each side of the body with

l+ strong internal longitudinal vessels very

closely placed on each fold. There are no in-

temal longitudinal vessels. in the interspace

between folds. The branchial tentacles are of

varying, sizes and twice pinnate. The dorsal

lamina is produced in a series of pointed Jan-

guets. The gut forms a narrow straigh! loup

und there is a mass of branched liver tubules

in the gastric arcu. Gonads, ure divided mo

separate paired polycarp-like gacy extending

along both sides of the central common duct.

Remarks; This specimen agrees in most aspects

with thase described from Investigator Strait,

although the sandy coating is not as thick in

the present specimen, Nevertheless, the nature

and orientation of the siphons are identical as

are the internal organs, viz. the branchial sac,

the gut and the gonads, The atrial yelum in the

specimen from Investigator Strait was present

at the basc of the atrial siphon rather than

more terminally as in the present specimens,

and this is possibly related to thickness of the

sandy couting. The test beyond the velum is

appurently produced to accommodate the thick-

ness of the sand surrounding the animal.

Pyura pachydermatina (Hercdmin) 5. sp gil

bosa (Heller), Kott, 1972b: 187.

Cynthia gilbbosa (Heller), 187%: 27,

Pyura gibbosa; Michaelsem & Hartmeyer,

1928; 410. Non P pachydermating var. ihe

besa; Kott, 1952: 269 (<P. puchydermutina

draschii; Kott, 1972b: 187),

Pyura puchydermata var. intermediu; Katt,

1952; 264 (synonymy) (part) Non P, gin-

have seterimedia Michaelyen, (922: 391, (<P.

spinifera; Kott, 1972b: 187).

New Record: northern Great Australian

Bight. For Previous Reeords, Descripion,

see Ko 1972b (Pf. — pachydermaiina

aravchii), 1952 (P. pachyderinatinu inrer-

media)

Remarks: The present specimens huvy the

typical curved spines in the branchial siphon

and the unts is bordered with shallow rounded

lobes,

Pyura spinifera (Quoy & Gajrnard), Kott,

1972b: 186 (synonymy),

Now Record: northern

Bight.

Pyura, australis (Quoy & Gaimard), Kort,

1972b: 186 (synonymy).

New Revord: reel, Douglas Bank. (upper

Spencer Gulf).

Pyura scoresbionsis Kott, 1972a: 36, 1972hb:

187,

Mew Record; upper Spencer Gulf (Stns FA,

OV),

Geeat Australian

ASCIDTANS OF SOUTH AUSTRALIA TIT is

Pyura vittata (Stimpson). Kott, 19724: 37 Microcosraus planus a. sp.

(synonymy); 1¥72d; 243,

New Record: upper Spencer Gulf (Sta Al).

Remarks; The spines lintig siphons are typically

log (0.1 tom) and needle-like atid overlappne.

The anal border is smooth and two-lipped,

Pyurg irregularis (Herdman), Kott, 1972a: 38

(synonymy), 1972b: 187,

New Record; upper Spencer Gulf (Stn B7).

Remarks: The perttubercular arca has the usu-

ally blister-like appearance that is characteristic

of this species: The large dorsal tubercle, how-

ever, is.at the top of this area rather rhan, as

has been previously described, at its base, The

tough leathery test and strong branchial sac

characteristic of this species are present.

Pyura stolowifera (Heller) s, sp, praeputialis

Heller, Kott, 1952: 274 (synonymy); 1964:

141,

Pynra praepadalis; Millar, 1966: 372.

New Record) Outer Harbour (St Vincent

Gulf}. For Previous Records and Deserip-

tion see Katt, 1952; Millar. 1966.

Remarks: This location apparently represents

the western extent of this species, which has a

continous distrthution from Queensland down

the eastern Australian coast, The specimens

here at the apparent end of its range are

smaller than have usually been recorded from

other locations,

No constant difference hax heen detected

belween the South African P, stolowifera $, sp.

stalunifera and the Australian P. stolonifern 5s.

sp. praeputialis and most characters demon-

strate a remarkable and overlapping range of

vuriation in the two populations, The rounded

fold of test enclosing the siphons, however, is

never absent from Australian populations of

this species, although it also has been reported

From: South Africa; occasionally projections

of test surround the.apertures of South African

specimens but have never been observed in

Australian forms. The different frequency with

which these characters occur m cach popula-

lion suggesc that subspecific rank is appropnate.

This matter is discussed nore fully by Koti (in

press}.

Halocvnthia hispida (Herdman), Kott, 1968: 74

(synonymy). 1972a: 41, 1972b: 189,

New Record; upper Spencer Gulf (Stn Gy}.

Herdmania momus (Savieny). Kott, 1972a: 4)

(synonymy); 1972b: 189,

New Record; upper Spencer Guill (Sin G;

northern Great Australian Bight

Type Leeality: south of Goat L (olf Ceduna),

3t am deep, 17-25,x11.1967, Howler. Hols.

mpe: NMY. 4284, Paratypes: SAM, E 1032;

QM, G 7510.

FIG. 18

Description; The individuals are circular in out-

line and laterally flattened, with both apertures

close together on the upper surface. The test is

thin and completely encrustecl with sand, so

that the specimens resemble hardened discs of

sand, The sand is maintained around the ant-

mal by hair-like extensions of the test, and

postenorly these are longer, so that a flattened

sandy keel is developed which interrupts the

circular outline of the body. ft is upparent

therefore, that these Jaterally flattened indi-

viduals: are embedded uprivht in the sand

rather than lying on their side on the surface

of the sea floor,

The «apertures are sessile, Longitudinal

muscles from both siphons radiate over the

body, crossing one another in the middle of

each side as is usual in this. genus, There sre

also bands of circular muscles crossing the dor

sal and ventral borders of the body. There is

a conspicuous, elongate, dorsal ganglion be-

tween the two apertures, The branchial sav hits

7 folds on each side of the body with six

internal longitudinal vessels on each fold.

Parastigmatic vessels are present. There are

rectangular stigmata but no internal Jongi-

tudinal vessels between the folds, The dorsal

lamina is a wide, plain-edged, membrane. The

gut forms the usual long, narrow, curved Joop

with liver lamellae in the pyloric region. The

branchial tenticles are twice pinnate: although

the secondary branches are very short and

rounded, The rectum extends anteriorly to the

base of the alnal aperture, forming a deep

curve with the gut loop. The anal aperture is

bi-labiale. The gomuds are present in the sec-

ondary gul loop and consist of a Jong ovarian

tube often forming deep, close, curves, ‘The

ovarian tube has dense male follicles atnng irs

posterior border and on the lateral aspect of

the ovarian tube against the hody wall. As the

curves of the ovarian tube develop, the male

follicles appear to mingle with the ovary.

Owing ta the curving of ihe ovarian tube, il

often appears to project back into lobes

bordered by mile follicles, The gonad on the

right side of the body js in a corresponding

position to that on the left.

if PATRICIA KOT

Remarks: The species is unusual in the genus

i) that the gonads cdo nol cross into whe

pomary gut loop. In this respect only, ir re-

serililes Microcoyrnus stolonifera. Uhe lurm of

the gonads in the present specimens is, how-

ever, distincuve and the laterally futlened body

is also diagnostic of the species.

Microcosmps squamiger (Michaclsen}, Kate,

1972a: 43 (synonymy),

New Revords! upper Spencer Gulf (Stn G);

Outer arbour (St Vincent Gulf).

Remarks: The ptesent specimens hive a thick

test that is impregnated wih sand, and in some

tases numerous spesimens form avgresgutes,

The siphons are a rosy pinkish colour and

there are the characteristic flattened scales

lining the outer part of the siphons.

Microgosmus pichollsi Kott; 1972a: 42 (synony-

my): 1972d: 245

New Reeerds: northern Great Australian

Bight.

Microcosmus stolunifera Koll: 9724: 43

(synonymy), 197 2d: 243.

New Records: npper Spencer Gulf (Sta EL),

Remarks; The position of tre left wonad im the

secondary gut loop is characteristic of the

Species.

Fumily MOLGULIDAE

Molgula mollis (Hetdmun), Kott, 1952; 29%;

1964; 144; 149729; 45 (synonymy),

Molgula sabijlasay Kott, 972d: 248.

New Records! uppee Spencer Gulf (Sin

BIO); Investigator Strait (Ste Yiy. Foe

Previous Records, Deseriptian, see Kott

1972a (Molvula sabulasa).

Remarks: Vhe specimens ire small, more or

less laterally flattened, spheres. The apertures

are close tovether on the upper surface and a

ridge of slightly thickencd test extends between.

them, ‘Vhere are fing hairs on the lower part

of the test which is completely encrusted with

sand, ‘Phe rin of the aperture is Jobed ‘but

there arc no hollow test expansions sutround-

ing them as in M, xabulosa (eee below), The

apertures are directed away from one another.

Uhere are 7 branchial folds. on cach side of the

body, with up to 9 internal longitudinal. vessels

mm each ilistributed over both sides of each

fold, There ure no internal longitudimal vessels

between the folds, The testis follicles form a

complete citcle at the end of the ovary, with a

ligament extending through the centre of this

circle. On the lateral aspect of Ihe ovary, it ts

uwpparent Ubal the testis follicles embrace the

end of the ovarian tube, but on ihe mesial

aspect the U is completed to form a-citcle by

the growth of the (wale follicles across Ihe sur-

face of the ovary, The male follicles are long

and on their lateral aspect lie along tke bady

wall dirested from the periphery of the circle

into the centre. On the mesial surface of the

gonads, the testis follicles can be seen tu be

more or less fan-shaped and tightly packed

with their outer border divided into separate

lobes. ‘The vas deferens. extends from lhe

centre of the circle of testis follicles and,

viewed from the outside of the body, has twa

vasa eflerentia connecting ducts from each

individual testis follicle on each side. The vas

deferens then extends mesinlly and along the

surface of the ovarian tube aiid opens above

the opening of the oviduct, The proximal part

of the vas deferens is expanded into a seminal

vesicle.

Remarks The lengih of the oviduet and the

absence of hollow test extensigns pround the

apertures distinguish this species from the very

similar Af. yabutoya (see below) which hus

oflen been confused with it.

Molgula sabulosn (Quoy & Gainiard), 1834:

613. Michvelsen & Hurtmeyer, 1928: 449

(synonymy).Kott, 1952: 298 (part): 1972b:

190, Millar, 1966; 374.

Remarks: ‘Vbere ave no new records lor this

Species: however, a small specimen from Ells-

tou has made it possible to compare the species

characteristics with those of At. mollis with

which it has been confused, It is clear that the

differcnees in the lwo species are not associated

with maturity, Mf. sabulesa is spherical with a

sandy test that is hard and brittle, while Af.

mollis, although encrusted with test, has fine

hairs to Which the sand adheres, the test itself

1% more Macc, and the preserved specimens

are lutcrally Nattened. “The branchial aperiure

is always protected by 6 pointed lobes from

the surrounding test a liule distance from the

opening while the rim of the aperture itself js

produced into 6 smaller pomled lobes, that are

covered in the closed position by the rim of

the larger lobes. ‘The atrial aperlure is pro.

tected by two flattened, wide tongues and their

border is sepurated info three rounded lohes

that arise from the test at the dorsal and

ventral sides of the apening, The rim of phe

aperture itself is produced into + sittall,

ASCIDIANS OF SOUTH AUSTRALIA Il 7

pointed, sandy lobes. and these are covered by

the luge lips in the closed position. ALL these

extensions from the test around the upertures

are hollow and have prolongations ef the body

wall extending into them, They are characteris-

tie of the Species and are never present in A,

mollis, The gonad in the present species, while

superficially resembling that of M. nrelliy, has

a very short vas deferens. Ihal opens at the

proximal efd of the ovary on its mesial surface,

Biogeography

The 22 species recorded from the northerm

part of the Creat Australian Bight (including

Ceduna), as far as 32"24°S, 133°30’E. cun be

divided into the lollawing groups:

\. Possibly endemic w the Great Austratian

Bivht.

Patridium pulvinaiun a. sp. Aplidinm digi-

tatu no, sp; Aplidiiun foliorum o. sp.. Lep-

tuclinides volvus n, sp.; Parabortryllus nema-

rus. sp.v Pyare tendata Kott, Mieroceasiantis

planus n. sp,

2. Southern temperale

South Africa),

Aplidiumn favolineainm (Shutter); Aplidinn

colelloides (Herdmin),

3. Circum-australian,

Polyctior giganieum (Herdman); Lepto-

elinides reticulatis (Sluiter); Leptoclinides

yufuy (Sluiter); Palysyncraton aspiealatun

Tokioka; Pidenmum candidum Savigny:

Didermuin moselevi (Herdman); Ainphi-

enrpa diptyeha (Hartmeyer): Polyearpa pe-

chentenlata (Meller); Cnemidocarpa etheriedyit

(Herdman) (absent only front tropical Aus-

tralia); Merdmania momus (Savigny |

Southern tel easiern Ausiralian species:

Ascidia aclara Kott; Sryela pedata {Herd-

man). Pynra pachydermating (Herdman)

gibhosa (Heller); Pyara spintfera (Quey and

Gaimard); Mierocosmus nichalli Kott

(absent only from tropical Australia).

‘The apparently endemic spesies comprise a

vonsiderable component (31%) af the fauna

in the horthern part ot the Great Austratian

Bight, The circum-Australidn forms comprise

almost 50% of the species, while species with

a range to Port Jackson (Pyura p. pibbora, P,

spimiera) or Moreton Bay (Aaciedia avlara.

Sivela peduta. Microcasmus nickallsi) also

occut. he three latter records extend the

known range i the west, although the first

two species are already known to oceur in

southwestern Australia. The data that are

(recorded also from

recurded here do noe therefore disagree with

previous infgrmation (Kote 1972b) that sup-

ports the existence of 4 marine faunal Province

extending from Covkburn Sound (or further

lo the north) un the western Australian vost

to the vicinity of the castern const of South

Australna

There is no evidence that would sugeest thal

the sample that is available is out typical of

the fauna of the Great Australian Bight, This

Twuna, however, does nol, on the biayis of avail-

able data, appear to be iypical of the Plin-

dersian marine faunal Province. Apart froin

the large endemic component, the species oc-

curring there have a wide distribution around

the Australian coast, espectally wlong the cast-

ern seaboard, The species that terminate their

tange ul the eastern end of the Flindersian

Province have not been taken in these collec-

tions From the northern part of the Great Aus-

tralian Bight (Kott 1972b), although they have

been recorded ut more easterly locations off

South Australia.

Other species in this collection laken From

other locations off the South Australian coast

may be grouped according la the South Aus-

tralian limits of their range in the following

Way;

\_ Species thar do nat extend eastwards into

the Matizgeah marine Province,

Podoclavella cylindrica (Quay & Gaimard);

Pyenoclavella diminua (Katt): Adapozea

marshit Brewin; Diplosoma translicidian

Hartmeyer), Polyclinuim nepluniunt (Hart

meyer; Stolonica carnosa Millar; Molguta

sebuluse Kou.

2, Species that da not extend westwards intra

the Flindersian marine Province.

Distaplia australtiensis Brewin; Eteberderania

australis Kotl; Aplidium voui/erum Kate:

Aplidium amorphatum Kotts Ascidia themp-

sont Kott; Palyandrocarpa lapidosa (Herd-

man): Pyura irreeuluris (Herdman): Pely-

carpa tinetor (Quoy & Gaimard); Polvcarpa

papitata (Sluiter).

3. Species for which the Flindersian’ Mangean

houndary does not comprise a barrier,

Sycazea cerebriformix (Quoy & Gaimard):

Sieozadn pedutteuluta (Quoy & Gaimard);

Ritterelia herdmania Wott; Trididemnunr

savignii (Herdmain); Vrididemauam cerelri-

ferne (Hattmever); Syplegma viride

Herdman: Pyura ausiralix (Quoy &

Gaimard}; Microcosmus stelanifera Koll

4. Gall fannie.

Pyar scareshiensis Kol.

1% PATRICIA KOTT

References

Anporr, D, P.. & Jounson, JF, Vo (1972)—The

Ascidiaos Styela barnharti,.S. plicata, S. clave,

and S. montereyensis in Californian Waters.

Ball, $e, Catifornta Acad, Sei, 712), 95-105.

BREWIN, Beavt |. (1952).—Ascidians of New

Zealand, Part VIE, Ascidiuns from Otago

coastal warers, Part I. Trans. R. Soc NE.

TH 3-4). 452-458.

Barwin, Aervt. V, (1953)—Australbia aseidians

of the sub-family Hylozainac and a revicw

of the sub-famity, Srans, Ry Sav, NZ, BCL),

53-64.

Brewin, Breve 1. (1956)—Alepozoa praorvhl, a

compound ascidian from Western Australia,

J, R. Soe. WA. 40(1), 31-32.

Barwin, Berve |. €1957)—Ascidians of New

Zealand, Part XX. Ascidians from North

Auckland. Trans. R. See. N.Z, 84(4), 577-

380.

Baewin, Beryt I. (1958a).—Ascidiuans of New

Zealand, Part XL. Ascidians of the Stewart

Island region, J'rans, R, Sve. N.Z. 85(3),

439-453.

BREWIN, Beryr. J. (1958b).--Aseidiany ol New

Zealand. Part XT, Ascidions of the Hauraki

Gulf Pt Wb Trans, Ry Suc, NZ, 85, 455-458,

btoagEpGR, L. G. (1967).—A taxonontic review of

Indo-Pacific didemnid ascidiurs and deserip-

tions of twenty-threé central Pacific species.

Migriaeyie 2, 161-2461.

Gceynn. P. (1974).—Rolling Stones umong the

Scleractinta. frac. Second Interaarional Ceral

Reef Svuipoasium. Vol. IL 183-198. (Great

Barrier Reef Committee: Brishanc},

WaRTMEVER. R. (1919),—-Ascidien. Results of Dr.

FR. Mjiéberg’s Swedish: scientific expeditions to

Avistvatia 1970-13, Ky weensha Metensk-A kal.

Hanid!, 60(4), 1-150.

Hasrincs, AwNA B, (1931).—Tnnicota, Setens

Rep. Gt. Barrier Reef Baped. 4(3), 69-109.

Hletirer, ©. ((878),—Beitrhige zur niiheren

Keantnis der Tunicater. Sher. Akad. Wary.

Wien, 77(1), 83-110,

Kort, Pareto (1952)-—The asendians of Aus-

iralia. 1. Stolidobranchiata and Phleba-

branehlatu. fuse, J. mar. Freshiw. Res, 303),

206-333.

Korn, Faretei, (1957).—The ascidians of Aus-

aril, 1 Aplousobranchinta tahtlle, Clave-

linidae HKorbes 4nd Hanly and Malycliniduc

Verrill, tir, J) fier Fresh Res, BOL),

64LIO.

Kort, Patricia (1962).—The ascidians of Aus

tralia. IT, Aplousabranchinna tahitle: Didem-

nhiduc Giard, Anse. J. omar. tyveslow. Res.

(303), 265-334.

Kor, Aare, (1963)—The weidians of Aus-

tralia: TV. Aplousobranchiata Lahille; Poly-

clitidae Veerl (continued). cause. J. omar.

Preshw. Res. 14(1), 70-118.

Kort, Parricia (1964).—Stolidabranch and phle-

buobranch asvidians oF the Queensland coast.

Pap, Dep. Zool, Univ, Qd 27), 127-152.

Kort, Patric (1966) —Ascidians uf north Aus-

uolia. Pap, Dep. Zoal. Univ. Qa B15),

279-304,

Kor1,

Haloevithia Vert il.

NSM, 930}, 746-89,

Kori, Pataicia (1989).—Antarctic Axcidiacea, A

monographic account uf the know species

based on specimens collected under US

Government uwuspices 1947 to 1963. Adfured,

Ros, Ser, 13, xy, 1-239.

Kori, Paraicn (1972a)—The aseitians of Sotth

Australia L: Spencer Gulf St Vincent Gulf,

and Encounter Bay. Trans, R, Sec. 5. Ast,

9f(1), 1-42

Kort, Patricuy (1972b).—The uscidians of Sauth

Australia Il, Eastern sector of the Great

Australlan Bight und Investigator Strnir.

Trans. R. Soe. S&S. Aust. 96, 166-196.

Kott, Parnicia (1972¢)—Notes on same wsei-

dians fram Port Juckson, Botany Bay, and

Port, Hacking, New South Wales. Proc. Linn.

Soe, NSW. 97, 241-297,

Korr. Parkicia (1972d),—Some sublittoral asci-

dians in Moreton Bay, Queenshind, Mem. Qu

Mus. 16(2), 233-260.

Kort, Pararcrs (1972e).—Fauna of the Gut of

Curpentarias 2. Ascidiacea (Chordata: Tuni-

camil. Fish. Notes N.S. 2(3), 39-57.

Korr. Parrtcia (in press).—The ascidian fauna

of Weslernport Bay, Victoria; and a com-

parison with that of Port Phillip Buy. Mern.

Nain. Mus. Viet.

Mecnarosrn, W,. (1922). Ascidiae Plycha-

brunchiae und Diktyohranchine von Neusec-

land und den Chatham-[nseln. (Papers from

Dr. Th Pucific FRxneditian

Pargicia 1968)-—-A review of the genus

W879, Proc. Linu, Sec.

Morelensen's

1914-16, No, XT). Widensk, Meddr, dansk

natarh, Faren, 73, 359-498,

MICHARLSTN, W, (1924). Ascidiac = Krike

hranchiae von Nensecland, den Chatham und

deo Auckland-Inseln. (Papers from De. Th,

Mortensen’s Pacific Expedition 1914-16. No,

XNIM. EIReUS Ke Meddr. dansk. waturh. Foren,

TT, 263-434,

Micnatiskn, W., & Hanemever. R. C128 )—

Ascidine diktyobranchine urd — ptycho-

hranchiae, Fauna Sudwest-Anst. 5, 251440,

Mittar, BR. H. £1960) —Ascighiaces.

Rep, WO, 1-460,

Mra. R. H. £1961).—Aseidians from. Mezam-

bique. Ann. Mag. Nu FHivt. ¢63)4. 11-16.

Miniar, R: Wi. (1962),—Purther descriptions of

South Afreun ascidian Ang J. Afr. Mus.

46(7), 113-221.

Mingar, R. H. €1984),—

the British Museum (Natural Tistory 1

zinl Soe, Lend. 14114). A89-745.

Mitiak, R. H. (1966).—Ascidtaces. Port Phillip

Survey, Alem, natn, Mus, fet. 27, 357-375.

Nort. J, T. (1892).—On the composire ascidians

oi the North Shore Reef. Vyvans, NZ. Inst. 24,

35-334.

Picuon, M, (1974),-—-Free Iiving seleractinian

coral communities in the cofal reefs of Pilea

(Madovascar), Pree Sevond lyternatiowal

Coral Recf Symposiam. Vol, UW, 1735-182

(Great Barrier Reef Committee: Brisbane. |

Piscavery

Australian ascidians in

Mroe.

ASCIDLANS OF SOUTH AUSTRALIA Il 19

PLANTE, R,, & Vasseur, P. (1966),—Svr une col

lection d'ascidies de la region de Tulcar (cote

sud-ouest de Madagascar). Ann. UUniv.

Méadagasvar, Serie Sciences de la Nature et

Mathematiques, no. 4. (Tanarive: Imprimerie

Nationale, )

Quor, J. & GarmMarp, P, (1834),—VYoyages de

découvertes de lAstralabe 1826-29. Mol-

lusques. Zoologie 3, 559-626; 4, 304-308.

Sturrer. C. P. (1909).—Dic Tunicaten der Siboga

Expedition Pt. 2. Die Merosomen Ascidien.

Siboga Exped. 56B, 1-112,

Stuiter. C, P. (1913).—Ascidien von den An

Inseln. Ab. senckenh, naturforsch. Ges. 35,

65-78.

Toxioxa, T. (1949).—Contributions to the

Japanese ascidian Fauna IL Notes on some

ascidians collected chiefly along the coast

of Kii Peninsula. Puhls Seto mar. biel. Lab,

1(2), 39-64.

Toxtora, T. (1952)—Ascidians collected by

Messrs, Renzi Wada and Seizi Wade from

the pearl oyster beds in the Arafura Sea in

1940. Publy Seto mar. bial. Lab. 2(2), 91-142,

Toeroxsa, T. (1967).—Pacific Tunicata of the

United States National Museum. Bull. U.S.

nam, Mus, 251, 1-242.

Vasseur, P, (1967) —Ascidies de Nouvelle-

Calédonie, Edition de la Fondation Singer-

Polignac, 127-146, 2 pls,

Appendix—Station List

NORTHERN GREAT AUSTRALIAN BIGHT

(32°24'S, 133°30'B), May 1973, Experimental

Prown Trawl, Explorer (Coll. P. Symond).

42m; Polycitor giganteum

Patridium pulvinatum n. gen. 1. 8p.

Aplidium colelloides

Aplidium foliorum n. sp.

Léptoclinidey velvus n. sp.

Herdmania momus

Aplidium colelloides

Aplidium flavolinearum

Aplidium digitutnm n. sp.

Leptaclinides reticuletus

Palysyncraton aspiculatam

Didemnum candidum

Didemmim moseleyi

Amphicarpa diptycha

Polycarpe pedunculata

Styela peelute

Cnemidocarpa etheridgii

Pyura pachydermatina gib base

Pyura spinifera

Terdmania mamts

Micracosmus Hithallyi

GOAT ISLAND, off Ceduna, Great Australian

Bight (Coll. P. Howlett).

32m: Ascidie aclara

Pyura tendata

Micrecosmus planus 0, sp,

ELLISTON BAY, Feb. 1971

Shepherd),

Outside bar:

Fuherdmania auxtraliy

Ritterella herdmania

A plidiuen amorphatium

Leptoclinides rufus

Oculinaria australis

Synplegmu viride

Apliditon coniferian

Vertical faces (25 m): Pyura pachydermatina

Molgula sabulose

WEST ISLAND: Amphitheatre Rock

7 m deep, 13.vii,1972:;

Polyciter giganteum

17 m deep, 12,vii,1972:

Sycozoa cerebriformis

CAPR JAFFA: Margaret Brock Reef (3-4 m deep

and in caves), 28.xi,1972.

Pseudodistoama cereum

49m;

(Col, S. A,

Reef:

INVESTIGATOR STRATT, January, 1971 (Coll.

J.B, Watson),

Station XT (depth 19 m):

Aplidium pronum ua. sp.

Diplosoma translucidtim

Pyura ansiraliy

Station X3;

Pyura australis

‘Station X7:

Herdmuania momus

Stations X8. X9, X10:

Pyura scoreshiensis

Station Y1:

?Palyclinum nepiunium

Srolaniea carnasa

Palyandrocarpa lapidosa

Molgiula mollis

Station ¥5:

Atapozoa marshi

Euherdmania australis

?Polysynerauton aspiculanin

Pyura australis

Station 76:

Atapaezed marshi

UPPER SPENCER GULF, September, 1973.

Transects. and stations of S. A. Shepherd, Depart-

ment of Fisheries, S. Aust.

Transects A-D:

Polycarpa pedunciulata (arenaceous and

naked specimens, sometimes grawing

side by side, attached ta the same Shell

or stone, stalked or sessile).

Station Al on Pinna, depth 0-1 m:

Pyura trregularis

Pyura vittata

Station A5, depth 17 m:

Ascidia thompsonit

Polycarpa pedunculata

Station A7, depth 10 m:

Palycarpa pedunculata (2 spec.)

Station B4, depth 17 m:

Sycezea pedunculata

Station B7, depth 5 m:

Polycarpa pedunculata

black; a few leathery)

Polycarpa papillata (single specimen)

Pyura trregularis (one small aggregale}

(some naked.

PATRICIA KOTT

Station B10, depth 10 m, channel;

Polvearpa tinctor

Moleula meilis

Station C4, depth 12 m:

Polycarpa pedunculata (naked and arena-

ccous )

Station D3, depth 18 m:

Asctdia thompsent

Station DS, depth 15 m;

Distaplia australiensis

Station 19, depth 10 m:

Pyura scoresbiensis

Station E1, depth 7 im;

Palycarpa pedunculata

Microcosmus nichollst

Micracasmus stalonifera

Station E3, depth 9 m:

Ascidia thompsoni

Station E4, depth Sm:

Pyura irregularts

Station Fl, depth 19m:

Polycarpa pedunentata

arenazeaus)

Station F3, depth 19 m;

Pyura scoresbiensis (without stalk)

Station F4, depth 16 m:

Aseidid thompsani (with barnacles)

Station G, depth 9 m:

Aycidia thompsoni

(small stalked,

Amphicarpa diptycha

Aptidium amorphatum

Aplidium cole!loides -

Aplidium coniferum -

Aplidium digitatum -

Aplidium flavolineatum

Aplidium folicrum ==

Aplidium pronum ——-

Ascidia aclara - -

Ascidia thempsont = -

Atapozoa marshi -

Cnemidocarpa etheridgi

Didemnum candidum

Didemnum moseleyi -

Diplosoma translucidum

Distaplia australiensis

EBuherdmania australis

Halocynthia hispida -

Herdmania momus -

Leptoclinides reticulatus

Leptoclinidesrufus -

Leptoclinides volvus -

Micracosmus nichollsi

Microcosmus planus -

Microcasmus squamiger

Parabotryllus nemorus n. gen,, o. sp.

Polycarpa pedunculata

Pyura scoresbiensis

Halacynthia hispida

Microcasmus squamiger

Herdmania moamus

Reef, 4 km NNW Douglas Bank:

Podeclavella cylindri¢a

Pyura australis

SPENCER GULF.

Tipara Reef, depth 11 m, 24.ix.1971]+

Pyenaclavella diminuta

Pywura irregularis

Under stanes:

Herdmania momus

Depth 5 m, 2.v.1972:

Padaclavella cylindrica

ST VINCENT GULF.

Port River (near Electricity Trust). depth 3 m,

muddy bottom, 9.vi. 1972:

Styelq plicate

Outer Harbour:

FPywrq stoloaifera

Micrecasmus squamiger

Sellick Beach (S of Adelaide), Feb. 1972 (Coll.

R. Hammond):

Trididemnium savignii

Trididemnunt cerebrifarme

Index to Genera and Species

—

' 1

' '

= —e

' '

' 1

foot ac

Dwaartomeunwmwreooe py De-qnunAsanyntare

‘

Pr aw}

Microcosmus stolonifera - - 16

Molgula mollis - : - 16

Molgula sabulosa . - 16

Qculinaria australis - - - 11

+ Parabotryllus nemorus - - = 2

Patridium pulvinatum - = 3 =

Podoclavella cylindrica . - -

Polyandrocarpa Japidosa = - - - -

Polyearpa papillata = - - - - -

Polycarpa pedunculata - ~ - -

Polycarpa tinctor . - - - -

Polycitor giganteum = - - - x

Polyclinum neptunium - - - -

Polysyncraton aspiculutum = - 5

Pseudodistoma cercum - - = -

Pyenoclavella diminuta - - -

Pyura australis - - _ -

Pyura irregularis - - - - -

Pyura pachydermatina - - - -

Pyura scoresbiensis —- - - - +

Pyura spinifers . - - - -

Pyuta stolonifera - - - - -

Pyura tendats = - - - - - -

Pyura vittata = - - - - -

Ritterella herdmania - = = 2

Stolonica carnosa - - - -

Styela pedata =~

Styela plicata - - - - - ~

Sycozoa cerebriformis = - - -

Sycozoa pedunculata - - - 3

Symplegma viride - + - - -

Trididemnum savignii - - - 2

Trididemnum cerebriforme - - A

—

PURO DoO=p-Ppo

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: 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. Buonatuto*

Summary

Buonaluto, 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 Pseudomaiaxis 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 Pseudo-

malaxis Fischer, of Calodisculus Rehder to Awarua Mestayer, and of Claraxis Iredale to

Torinista Iredale, are considered.

Introduction

The genus Pseudomalaxis Fischer is repre-

sented in Australia only by three known spe-

cies, two fossil and one living: the Late Eocene

P. asculpturatus Maxwell, 1966; the Miocene

P. praemeridionalis (Chapman, 1912); and the

living P. meridionalis (Hedley, 1903). Only the

two fossil species will be discussed and des-

cribed here. However, it is necessary to discuss

(a) the taxonomy of the genus Pseudomalaxis

and the related genera Mangonuia and Awarua

Mestayer; (b) in which family the genus should

be placed.

(a) Fischer (1885, p. 714) found in a living

form, then referred to the Neogene Bifrontia

zanclaea (Philippi), a torinioid operculum and

therefore instituted for it Pseudomalaxis as a

new subgenus of Torinia Gray. Sacco (1892,

p. 75) considered Pseudomalaxis Fischer a sub-

genus of Discohelix Dunker. Dall (1892, p.

331) recognized the intrageneric relationships

between the Recent American Omalaxis nobilis

Verrill and P. zanclaea (Philippi), but insti-

tuted Discosolis for O. nobilis as a section or

possible subgenus of Discohelix, because of

Verrill’s description of the operculum of O.

nobilis as trochoid. Iredale (1911, pp. 253-7)

tidied up the confusion existing in the use of

the names Discohelix Dunker, Omalaxis

Deshayes, Bifrontia Deshayes, and Pseudo-

malaxis Fischer; he placed Bifrontia in syno-

nymy with Omalaxis, and separated the three

remaining genera as distinct taxa. Also, he dis-

tinguished one of the two living forms pre-

viously referred to P. zanclaea (Philippi) as a

different species, P. macandrewi Iredale, be-

cause of the latter’s more evolute coiling, and

restricted Pseudomalaxis to this new living

form.

Later, Monterosato (1913, pp. 362-3), from

a different viewpoint, restored P. zanclaea as

type-species of Pseudomalaxis and described

the other living Mediterranean form, P. actoni

Monterosato, previously mistaken by authors

for P. zanclaea. Monterosato also instituted for

the above evolute form the subgenus Spirolaxis

with P. (Spirolaxis) centrifuga Monterosato,

1890 (syn P. macandrewi Iredale, 1911) as

type-species (Monterosato 1913, fig. 2). Coss-

mann (1915, pp. 122, 141) restricted Disco-

helix Dunker to the Mesozoic forms and

Pseudomalaxis to the Cretaceous-Recent, re-

ferred both to Euomphalinae, and separated

Omalaxis Deshayes in the Omalaxinae, a new

subfamily.

Rehder (1935, p. 129) recognized a very

close affinity between Discosolis nobilis

(Verrill) and Pseudomalaxis actoni Montero-

sato and therefore placed Discosolis Dall in

synonymy with Pseudomalaxis s. str. Rehder

(1973, pers. comm.) remarks: “Mangonuia

Mestayer, 1930, is probably a junior synonym

* Department of Geology, University of Adelaide, S. Aust. 5000.

NOLES ON PYEVUDOWALANTIS PISCHER IN| AUSTRALIA 23

or at leas! a subgenus of Pseudometixts

Fischer, 1885". Mestayer (1930, pp. 144-5)

refers P. nreridianaliy (Hedley) to Mangoania.

In fact, the recent Mangoniia holland Mes-

tayer, the type of this genus, displays the

general pattern of spiral ornaments, the quasi-

rangular outer and the subcircular inner shape

of body whorl churacteristic of the Neogene-

Recent Psendomalaxis s, str,, described below,

and the same type of pscudoplanispiral coiling

of this group.

Rehder further comments, “Calodisculus

Rehder, 1935, is very close to Awarua Mes-

tayer, 1930, and is probably its synonym”.

Wenz (1939, p, 667) considers Awarne a sub-

venus of Mangonuid. However, Calediseuluys

and Awerna display the same kind of speciili-

zation in spiral Ornaments. in particular 2

marked development of heavily beaded cireum-

umbilical cords, Awarna amoena (Murdoch &

Suter) (Suter 1913, p. 318; 1915. pi LS, fig.

21 ub) displays also. a tortinioid operculum

{Mestayer 1930, p. 146).

“Claraxis and Torinista Tredale are very simi-

lor, possibly synonymous, and might be sepa-

rated as a distinct genus which may he closer

to Hehucuy d'Orbigny”. Wenz (1939. pp. 666,

648) considers the former two as subgenera of

Mangonuia and the Jatter as a synonym of

Torinia Gray. In this case, Cloraais should tall

inte synonymy with Toréiixta because although

iredale (1936, p. 327) published their initial

diggneses on the siime page, ihe former is des-

eribed after the latter, From the original des-

cription und drawings of the type species, it is

impossible to find substantial generic differ-

ences and Iredale did not specify any,

tb) The genus Pseudomualaxiy Fischer is often

Placed in different families but generally is

referred to the Architectonicidae. In introduc-

ing the genus, Fischer referred it to the Solur-

tidae (= Architectonicidae) on the basis of the

comcal torinoid operculum (Fischer 1885, p.

714. see also Bames 1952, p. 37), observed in

the Recent form P. (Spiralaxis) centrifuge

Monterosato (Monterosato 1890) and in P.

nabils (Verrill) (Verdi) 1885, p. 423, pl 44,

fiz. 12).

Jredale (1956, p. 326) instituted a new

fairly Mangonutidae for the Australasian

genera Mangonula, Awarua, Torinista and

Claraxis, Apparently, Wenz (1939) cunsidered

Mangunuiidac synonymous with ihe Architec-

tonicidae, Because of a certain aflinity between

the opercula of P. balesi Pilshry & MeGinty

and Parviturbo zacalles (Mazyck), Pilsbry &

McGinty (1945, pp. 9, 57, pl. 6, figs 2-2a, 4)

placed Psevdomafaxis in the Cyclostrematidue,

Later, Abhott (1954, p. 138) included Preude

malaxis in Vitrinellidae on the basis of a vague

and incorrect reference lo a tevision of the

family by Pilsbry. Maxwell (1966, p. 444) fol-

lowed Abboit,

Reider (1974, pers. comm.) gave the

opinion: "L have examined young specimens of

both Pseudomalaxsiy and Architectonica (PH-

faxis), and I can find no essential difference im

their protoconchs; both show a heterostrophic-

anastrophic protoconch, | feel, therefore, that

Pyeudomalaxixy should remain in the Architec-

tonicidac”’. There is suppon for this statement

both on the basis of the turintoid conical oper-

culum and the protoconch coiling of Psetde-

maluxis, Therefure the author, in agreement

with Rehder, regards the restoration of Pseude-

malaxis Fischer to the Architectonicidue to be

justified.

Collections

The specimens of P. aseulpturatus Maxwell

here studied are kept in the Palucantological

Section of the New Zealand Geological Survey

(NZGS), and in the Department of Mines of

South Australia (GSSA), The figured specimen

of P. praemeridionalis (Chapman) is kept in

the Palaeontological collection of the South

Australian Museum of Nattiral History

(SAM)-

Specimens M3298 and M3299 were found

by the author and specimens M3300-M3303

were found subsequently by J. M. Linilsay-

big. 1. Pseudupnetexis (Pseudomalaxis) asculpluraius Maxwell. MeCullough’s Bridge, New Zeuland,

NZGS, 9508-2; a, adapiouk, b, abapical; c, axial views (all & 17,50).

Fig.

1 of (Psevidlomalaxis) asculptaratus Maxwell. McCullougit’s Bridge. New Zealand, NZGS, 5508-1:

ti, didupical: 4, abapical; ¢, axial views (ull X 17,3).

Pig 3, P

(Psevidemualaxis) asculptiratus Muxwell. Adelaide Plains Sub-Basin, Adelaide Chillrens

Tlespitul Bure No, 5; GSSA, M3299; a, adapical (X 16); A. abapical (X Wty ©, axial views

(X 15)

Vip. 2

#. (Psendomalaxis) pracmecidionalis Chapman, Cliflon Bank. Muddy Creek; SAM. P(83542:

#, Adapical (X 10.6); 6, abapical (X16); ¢,

axial views (XM 12,3),

M. F. BUONAIUTO

Synopsis of the history of the génus Pscudomalaxis Fischer on ile basis af the major authors,

Author Family Subfamily Genus Subgenus

Fischer (1885, p- Sulariidae — Torinia Gray Pseudomalaxis

714) Fischer

Sacco (1892, p. 75) Solariidue = Discohelix Dunker Pseudomalaxis

Dall (1892, p, 331) Solariidae — Discohelix Pseudomalaxis

Discosalis Dall

Iredale (1911. Solariidae (?) _ Pseudomaflaxis Pseudomalaxis

pp. 253-7) Discohelix Discohelix

Discosolix

Monterosato Solariidae —_— Psreudomalaxis Pseudomalaxis

(1913. pp. 362-3) Spiralaxis

Monterosato

Cossmann (1915, Euomphalidae Euomphalinae Pseudomalaxis Pseudomalaxis

pp: 122, 141)

Mestaver (1930, Architectonicidae _ Manganuia —

p. 144) (syn. Solariidae) Mestayer

Awarua —

Mestayer

Rehder (1935, p. Architectonicidae — Psendomalaxis Pseudomalaxis

128) ? (syn. Discosolis

Dall)

Panrodiseus Rehder

Caladisculus Rehder

Tredale (1936, Mangonuiidae Mangonuit _—

p. 326) Awarua

Wenz (1939, p. Archilectonicidae = Mangonuia Mungoniia

668) (syns Solariidae, Awarua

Mangonuiidie ) Psendomalaxis Pseudomualaxis

Spirelaxis

Paurodisceus

Calodiseulus

Pilsbry & McGinty Cyclostrematidae — Pseudomalanxis Pseudomalaxis

(1945, p. 9)

rene (1952, Architectonicidac = Pseudamalanis Pseudomatlaxis

p. 37)

Abhott (1954, Vitrinellidae ane Psendomalaxis Pseudomalaxis

pp. 138-9)

Pchelintsey & Solariidac — Pseudomalaxts Psendomalaxis

Korobkov (1960,

pp. 137-8)

Maxwell (1966, Vitrinellidac - Psetrdomalaxis Psreudomalaxis

p. 444)

Glibert (1973, Architcetonicidae — Pseundomalaxis Pyendomualaxis

p. 30)

This paper Architectonicidae — Pseudomalaxis Pseudamualaxis

(syn. Mangenuia)

IManganuia

Spiralaxis

Panrodiseus

Awarua

(syn. Calodiseulus)

Calodisculus

Systematic Descriptions

Phylum MOLLUSCA

Cluss GASTROPODA

Order MisoGAsTROPana

Superfamily CERITHIACEA

Family ARCHITECTONICIDAE

Genus PSEUDOMALAXIS Fischer, 1885

Diagnosis: Shell discoidal, pscudoplanispiral;

abapical side concave to subconcave, adapical

flattencd to subconcaye; outer shape of the

body-whorl rectangular 10 quadrangular, inner

shape subcircular to similar to the outer; thin

Margins, two carmae at abaxial-adapical and

abaxial-abapical ends. Straight growth Jines

between the abaxial keels, Aperture rectangular

to quadrangular; protoconch heterostrophic-

anastrophic; operculum torinioid, thin, multi-

spiral, outside flat or conical (after Wenz 1939,

p. 668).

NOTFS ON

Subgenus Pseudammalaxiy s. str,

Diagnosis: Shell medium-large to very small;

margins thin; pseudoplanispiral with lanvent

whorls; suture Hush to subimbricated; two ada-

pical and two ubapical keels: the four keels

can be smooth or crenylated; growth lines a

little irregularly packed; operculum flat (after

Wenz 1939),

Observations: Psendomalaxiy 8, str, appears. ta

be represented in the Tertiary by two main

forms, The first, predominant in the Eocene-

Oligocene, is commonly characterized hy

growth lines and the four carinue as the only

ornaments; « few species bear also. spiral orna-

ments of fine riblets as in the Eocene P. dixoni

(Vasseur) and che Oligocene P. italica (Sacco),

or of beaded cords and keels as in P. texana

(Aldrich); the outer shape of the body whorl

is rectangular to quadrangular, the inner is sub-

cireular to subquadrangular, the inner is sub-

circular to subguadrangular or subrectangular,

depending on the thickness of the margins (the

Jatter characteristic could be ontogenelic and

related lo age).

The second form, predominant in the Neo-

gene-Recent, is characterized by the develop-

ment of a main spiral rib or cord on the abaxial

region of the adapical margin, and often of

spiral costellae on the abaxial margin. The four

carinae and the adapical rib usually bear heads

or short spmes. The outer shape of the bady

DIMENSIONS (in mm): (see Fig. 5)

PSEUDOMALANIS FISCHER TN AUSTRALIA

whorl ig commonly quadrangular to subquad-

rangular, the inner shape is subcircular,

Pscudomalaxis (Pscudomalaxis) asculpturztns

Maxwell, 1966

FIGS 1-3

1966 Pseudomalaxts asculpnivatus Maxwell,

p. 444, figs 11=13,

Material: 4 specimens very well preserved wilh

the peristome slightly damaged (GSSA,

M3298-9; NZGS, 9508-1, 9508-2) and an-

other 4 juvenile or damaged (GSSA, M3300-3),

Description: Shell very small and thin, bicon

cave, nearly planispiral, protoconch hetero-

strophic-unusttophic; whorls increasing far

more in diameter than in height and overlap-

ping entirely in relation to the coiling axis and

very scareely in Telation te the normal to it.

Suture flush to subimbricated. Body-whorl

shape: outer subquadrate-subtrapezoidal; inner

subcircular-ovoidal, wider than high, Whorl

Tegions; adapical and abapical subconvex;

abaxial flattened or subconcave, nearly vertical.

Body-whorl tegions connected by prominent

smooth abaxial carinae. Abapical side more

concave than the adapical one,

Ornaments: Civowth lines and rugae, prosocline

in the adapical and abaxial regions, opistho-

cytt i the abapical. Four carinae: at ahaxinl:

adapical, abaxial-abapical, adaxial-adepical,

and adaxial-abapical ends.

Specimen No, whorls Dw N Nis Hw Lis Hee Deg

M3298 4 2.55 0.95 0.40 0.50 —0.10 0.60 9.65

M3299 4 2.350 0,92 0.39 0,55 —0,10 0.65 0.60

NZGS 3 1,55 0.70 0.30 0.40 —0,05 0.45 0.50

9508-1

NZGS 3 2.15 0.90 0,35 0,58 0,00 0.58 0.50

9508-2

RATIOS: (sce Fig. 5)

Specimen Hw/ Dw K — Lis/Hw = Nis/N Hgce/ Dye

M3298 0.1961 —0,2000 0.421] U.9231

M3299 0.2200 —0).1818 0.4239 10835

NZGS 9508-1 0.2581 —0,1250 0.4286 9000

NZGS 9508-2 0.2698 ,0000 0.3889 1.1600

Localities: Adelaide Plains Sub-Basin (St Vin-

cent Basin, S. Aust,); Adelaide Children’s Hus-

pital, North Adelaide, hundred of Yatala,

Town Acre 717, Bore 5, at 20.42-20.75 m

(M3298) and 21.64—-21.95 m (M3299}; Bore

2, at 22.25-22,56 m (M3302, M3303); Ade-

luke Metropolitan Subway, Bore 3, nerth bank

of Torrens Lake, opposite Kintore Avenue, at

20.10 m (M3300) and 24.50 m (M3301)

New Zealand: McCullough’s Bridge.

Stratigraphic Distrihurion: Adelaide Plains Subh-

Basin: Lindsay (1966) gave a brief sirati-

graphic summary of Adelaide Children’s Hos-

pital Bores 2 and 5, “The interval 20,42-21,95

26 M. F.. BUONAIUTO

m in Bore 3 is Jow in Blanche Point Maris

(Aldingan, Late Eocene). The Hantkenina

primitiva zone occurs in this bore ai $9.20—-

19.51 m, i.e. slightly above P. asculpturatus. Tn

the type section at Maslin Bay, Hanrkenina

prunitiva is presént only in w restricted band

0,80-1.15 m above the base of Blanche Point

Transitional Mars.

Other specimens from Adelaide Children’s

Hospital, Bore 2, are within the Hantkenina

primitiva zone. Although A. primitiva has not

yet heen found in Adelaide Metropolitan Sub-

way Bore 3, the specimens of P. .asculpturarus

are certainly from near, and probably from

slightly above, the 1, primitiva zone,

Thus the vertical distribution of all the spe-

cimens of P. asenulpturatns found in the Ade-

laide area spans a narrow interval from slightly

below to probably slightly above the A primi-

viva zone, in Blanche Point Transitional Marts,

Late Eocene.” (J. M. Lindsay, Dept of Mines,

pers. comm., 1974.)

New Zealand: Upper

Kaintan, Late Eocene.

Obserwerions: The two Australian specimens

show an inversion in the coiling, variable in

Waihag Greensand,

SS a

=a Cen

| L ) : Hgc, Hw Hyry

WL fg Z = i |

Dowd FOyael

le Nis —A reo

be N —H Hw<Has.,

——__—__——.-- Dw —- ---——"‘4H1 Lis <0

Fig, 3, The parameters measured in planispural

(a) and in pseudoplanispiral (b)

| gastfopod shell are here defined as

| Tollows:

Nea w the total number of the whorls,

——_«sSWS ————ernenr . 7? 1 we the total number of revolutions

lt +e ‘ ) WiC. bye Hw of the generating curve around the coil-

Na pe ee ee! = my ing axis.

__—— \ i s .

= ~~ => fiw: beight of any whorl (in this case

| the last one) as the projection of the

THU coiling axis of the distance between the

Hw = Hgeq adapical point of the generating curve

us = 0 ut the initial position and Wis abapical .

b point at the final position after a 27

revolution in the given intervul 2n7 -—

Zin+1)7,.

Lis; intersutural height of any whorl as the projection on the coiling axis of the dislunce

between the torial and final position of a given point placed on the adapical line of the hody

whorl after a 27 revolution, where the adapical line is the geometrical focus of any point

placed at the adapical end of the generating curye.

Hee: the height of the generating curve as projection on the coiling axis of the distance

between its adapical and ity abspical points.

Pw: the maximum diameter of any whorl as the projection on the normal to the coiling axis

of the pari of the planispiral or helicotd cone produced by the generating curve after a 7

revolution and included in the given interval (20-+-1)@ --- 2(n+1)7,

N: as the projection on the normal to the coiling axis of the distance between the adaxial

point of the generating curve al 2nw7 und ils abuxial point at 2(n |1)i positien,

Nis: iniersulural distance as the projection on the normal to the coiling axis of the distance

between the initial and the final position of a pomt placed on the adaxial line after a 27

revolution, where the adaxial line is the geometrical Jocus of any point placed at the adaxial

end of the generating curve-

Dee; diameter of the generating curve as the projection on the normal to the c. axis of the

distance between iis adaxial and iis abaxial point.

The generating curve is here considered the projection on a plane, containing entirely the coil-

ing axis, of the oulline 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 2nm -- 2(n |-1)™

T is the translation of the géeneraling curve along the coiling axis per revolution. K and 4 are

respectively the indexes of overlapping of any whorl; K, parallel to [he coiling axis: v, normal

to the coiling axis, and are defincd by the tatios K = TLis/Hw and v = Nis/N, (After Raup

1966. Observations consistent with this theory will be presented in a subsequent publication.)

NOTES ON PSEUDOMALAXIS FISCHER IN AUSTRALIA 27

amount, from a more helicoidal protoconch to

a more planisptral teleoconch, Undeserihed in

Australia, this form was directly compared with

two lopotypes of the only coeval species ol

Preudomalaxis known to Australasia, P. asculp-

trams Maxwell, These two topotypes, more

juvenile than the Australian ones, display a

somewhat higher spire, producing a decreasing

concuvity and consequent fattening of the

adapicnl side, as 4 result of the same inversion

of the coiling; fainter carinac, but the younger

of the two has therm more marked. Protoconch,

growth lines, outer and inner shape of the body

whorl are the same. Because the amount of

inverwon in the coiling of the New Zealand

furms appears to differ only slightly from that

of the Australian forms, and is also yattable in

the Australian and New Zealand specimens, it

is very difficult to distinguish at the specific

level between these Lwo forms. Hence, they are

here considered conspecific and representing

probably different geoxruphical mverphs.

From #. wycufpiuraies Maxwell, the Anglo-

Parisian Eocene P, dixeni (Vasseur) (Coss-

mann 1915, pp. 142-3), differs by its higher

spire and presence of spiral riblets; the Proto-

Adriatic Priabunian P. Aeyrichi (Oppenheim,

1896) by its much higher spire and an abaxial

margin abipically more convergent to the axis;

the Liguria-Picmonte Middle Oligocene P.

ttalicus Sacco. 1892, hy its lower spire and by

the presence of spiral riblets. The Indian

Rocene P. punjahensis Eames, 1952, seems on

the contrary yerv close, bul, the holotype being

very juvenile and the original illustration very

poor, it ix. impossible to determine the sctual

differences and affinities between them,

The American Claiborne Eocene P. retella

Chea) (Pulmer 1937. p. 176) differs bv its rect-

angular apertirc, higher spire, and lesser num-

ber of whors, Po texdnw (Aldrich) (Palmer

1997, pm T78) by crenulated abaxial keels and

spicwl! cords ou the adapical margin; and P.

pluntmerae Palmer (Palmer 1937, p. 178, hy

subelliptical margins, more Imbricated sutures,

and the adoral part of the last whorl twisted ta

the adupical

Other Eocene species (Cassmann 1915) are

the Angio-Purisian F patelfatus (Sowerby),

and ihe Egvptian P. /ybicus (Oppenheim),

Maxwell (Nov. 1974, pers. comm.) states:

“Two specimens of a Preudoimalaxis have re-

cently heen htuined From 9 Mangaoripan

(Lower Eocene) sample from North Cantes-

bury. The material is not well preserved but the

shells closely resemble P. asculptusatus except

tor their much smaller size (the large speci-

men is only | mm in diameter), I don’t think

that the lower Rocene shejls ure juveniles of

P. ascupiercius, however, as they also have

tnuch smaller protocenchs”,

He quotes also the occurrence af P. ef,

asoniptiumius Maxwell from Wharckurt, South

Cynterbury (Duntroonian, Oligocene) (Max-

well 1969, p, 161) but “The sole specinien is

broken", however ib “is certumly close to the

Eocene specics".

Psendomalaxiy (Pseadomalaxis)

praemeridionalis (Chapman, 1912)

FIG, 4

IG12 Hoemalaxis pracmeridionalis Chapman,

p. 186, pl. 12, figs 4-6.

Material’ 1 specimen very well preserved

(SAM, P18342),

Dexeription: Shell small, thin, planoconcare,

pseudoplanispiral, with the adapical side flat-

tened and the abupicul concave; whurls increas-

ing far more in diameter than in height and

overlapping entirely in relation lo the coiling

wais ugd yery scurecly to the normal to it.

Heterostraphic-anasirephic = protacanch with

three smooth whorls; the inversion of the coil-

ing already displayed in the third whorl, Suture

grooved. Body-whorl shape: outer subquad

rate- trapezoidal; inner subcircular - ovoidal;

slightly wider thaw high. Body whorl regions:

adapical conyex in the adaxial part, concave in

the ubuxial; abaxial flattened; abapical convex;

uperture with subquadrate peristome. Lips:

ndapical elliptical with a gutter-shaped reflec-

uion in the middle; parietal and abaxial straight:

abapical elliptical, Lip und region connections

ungular, marked by carmac.

Ornameniy: Four spiral beaded carinae, the

abaxial ones with very short asially elongated

spines: udaxial-adapical, adaxial-abapical, aba-

xial-adapical, abaxial-abapical, a spiral crenu-

fated cord on the abaxial part of the adapical

Tegion and a smooth costella on the udapical

part of the abaxial region. Growth lines: prosa-

cline in the adapical region; prosocline In the

abaxial; orthocyrt in the abupical,

2k M. F. BLONAIUTO

DIMENSIONS lin mm): (see Fig 99

No. whorls Dw N Nis Ly,

o 26 13 {145 0.88

RATIOS;

Hw/ Dw K Lis‘ HW vy — Nis/N

0.3384 —1).0227 0.3461

heoealities and Sreatigraphie Distribution:

Muddy Creek Marls, Clifton Bank, Muddy

Creck, 6.4 km west of Haniilton; “blue clays”,

Newport Formation, Altuna Bay Coal Shatt.

Vic.

Stratigraphic Range: Balcombian, Early Middle

Miocene (Ludbronk 1973),

Observations: P. praemvridionalix (Chapman)

was feferred to Darmgh (1970. p. 188) to

Manvonuia Mestayer. The specimen here des-

¢ribed represents the second discovery of this

species. From P. praemeridionalis, the Paleo-

Mediterranean Neogene P, zaneliea (Philippi)

{Wenz 1939, p. 668. fig, 1906) differs by a

slimmer hody whorl, higher spire, adupical

spiral costa. closer to the adapical-abaxial

carina, absence of spiral riblets on the abaxial

margin; from the Paleo-Mediterranean Pliocene

P_aldrovandi (Boresti) (Sacco 1892, p. 75, pl.

2. fig. 65 ul, 65 bis a-d) by smooth adapical-

abaxial carina, absence of spiral abaxial riblets,

higner spire, and acupical costa closer to the

adapical-abuxial carina; from the Parathetys

Miocene P. boettgert Cossmann (Cossmann

1915, p. 143) by a larger body-whorl, lower

spire, lesser number of whorls, lack of spiral

uraments extept rarely, beaded abaniul

carinae, a semi-circular inner shape of the body

whorl; from the Mediterranean Recent P.

action! Monterosato (1913. p. 362) by a greater

number of whorls, slimmer body-whorl, lower

spire, more murked growth lines, more abaxial

riblets, and an abaxial margin more parallel to

the coiling axis. The Australian Revent P. seri-

las Hee Der

—i,02 0.90 0.98

Hac! Dec

USTR4

dendliy (Hedley) (Hedley 1903, p, 351) dis-

plays, on the other hand, a very close ifinity

fo F. praemeridionaliy (Chapman), differing

trom it anly by two more abaxial ribleis and

the abapical-adaxial carina with heavier beads,

The New Zealand Recent ?P. bollonsi (Mes.

tayer, 1930, p. 144) differs by more whorls,

two beaded abaxial cords, braader ubspical-

abaxial carina and slightly opisthocline growth

lines on the adapical region,

the American Recent P. nobilis (Verrill)

(Vermill L885, p. 423, pl. 44. fiz. 12) differs by

its shotter spire, a greater number of whorls,

shimmer body-whorl, adapicul cord closer to

the jbaxial-adapical carina, five spiral ribs on

the abwxial margin, and fine spiral ribs on the

abapicul margin.

Acknowledgments

Lam very grateful to Dr P. A. Maxwell of

the Geological Survey of New Zealand for his

wssistance, particularly in lending tupotypes of

Prendomataxiy asculpluraius; to the Director of

Mines, South Australia, for the loan of male-

riql; to Dr H, A, Rehder, National Museums of

Natural History, Smithsonian Institution, for

the information on taxonomy; to Dr H- Silcock.

Department of Pure Mathematics, University

of Adelaide, for verifying the definitions of

the poramenters; to Dr N, H, Ludbrook for

encouragement and for reading the manuscript.

The work was catried out in the Department

of Geology and Mineralogy, University of

Adelaide, during tenure of a University Re-

search Grant.

References

Appoty, R, 1, 11954),—“American Scashells.”

(Van Nostrand: New York.) _

CHapMan, F. (1912).—New or little Known Vic-

torian fossils in the National Museum, Pi

XV). Some Tertiary Gasteropoda, Proc. #.

Soe. Viet, 25, 186-192, pls 12-13,

CossmMan, M. (1915)—"“Essais, de Paleoconcho-

logic compares,” Vol, 10. (Paris.)

Dat, W. H. (1892),—Contribution to the Ter-

tiary fauna of Florida with special reference

to: the Miocene Silex Beds of Tampa and. the

Pliocene Beds of the Caloosahutchie River.

Pt 2. Streptodant und other Gastropyds,

Trans. Wagner free favre Sci Philad., 32),

201-473, pls 13-22.

Dankacy, 1.4, (1970).—Catalogue of Australian

Tentian Mollusca (except Chitons). Mer.

Nair, Mus. Wier: 31. 125-212,

Eames, F. B. (1952).—A contribution to the study

of the Eocene nt Western Pakistan and West-

ern India. C. The description of the Scapho-

poda and Gastropoda from standiurd sections

in the Rakhi Nala and Zinda Pir areas of the

Westen Punjab and in the Kobar District.

Phil. Trans. R. Sac. London, B, 236(6311.

1-168, pls 1-6,

Fiscier, P. (1880-87 ).—"Manuel de conchyologie

et de Paleontologic conchyologigue ou histoire

naturelle des mollusques vivanty et fossiles”

(Savy: Paris.)

NOTES ON

GLutpert, M. (1973)—Revision des Gastropoda

du Danien et du Montien de Ja Belgique. Pt I.

Les Gastropoda du Calcaite de Mons, Meni.

Inst. R. Soe. Nat. Belz, 173, 1-116, 11 pls.

Hepiey, C. (1903).—Scientific results of the

trawling expedition of H.M.C.S. “Thetis”,

Mollusca. Pt I. Scaphopoda and Gastropoda.

Mem. Ausir. Mus, 4, 327-402, pls 36-8.

Trepan, T. (1936).—Australian Molluscan Notes.

No. 2. Ree. Ausir. Mus. 19(5), 267-353, pls

20-24,

Irepace, T. (1911).—On some misapplied Mol-

lusea generic names. Prac. Malac. Sac. Taare.

9, 253-263.

Lrenpsay, J. M, (1969)—Cainozoic Foraminifera

and Stratigraphy of the Adelaide Plains Sub-

Rasin, South Australia. Bull. geal Surv. S.

Alist, 42.

Lupproox, N. H. (1973).—Distribution and strati-

graphi¢ ulility of Cenozoic Molluscan Faunas

m Southern Australia. Tohoku Univ., Sci.

Rep., 2nd ser. (Geol), Special Volume, ©

(Hatai Memorial Volume), 241-261, pls 24-

28,

Maxwerie. P. A, (1969),—Middle Tertiary Mol-

luses from North Otago and South Canter-

bury, New Zealand. Trans. R, Soc. NZ,

Geol, 6(13), 155-185, pls 1-3.

MAXWELL, P. A. (1966).—Some Upper Eocene

Mollusca from New Zealand. N.Z. JL Geol,

Geophys, 9, 439-57.

Mrstayver, M. K. (1930).—Notes on New Zea-

land Mollusca, No. 5. Trans. N.Z. Inst. 61,

144-46, pl, 26.

MontEROsATO, Marquis de (1913).—Note on. the

genus Pseydontalaxiy Fischer and description

of a mew species and subgenas. Prat. Male.

Soe. Lond. 10, 362-363.

Moore, R. C. (1960)—"Treatise on Invertebrate

Palaeontology.” Part I, Mollusca, |, (Geol.

Soc. Amer. & Univ. Kansas Press: New

York.)

PSEUDOMALAXIS FISCHER IN AUSTRALIA 29

OPPENHEIM, P, (1896).—Das Alttertiiir der Colli

Rerici in Venetien, dic stellung der schicten

von Priabana und die oligocine Transgression

in Alpinen Europa, Z, Dt, geol. Ges. 48, 27-

152, pls 2-5.

PALMER VAN WINKLE, K. (1937).—The Clairbor-

nian Scaphopoda, Gastropoda, and Dibran-

chiate Cephalopoda of the Southern United

States, Bull, Am. Paleant, 7(32), Pt 1, 1-548;

Pt 2, Atlas.

PcueLIntTsey. V., & Koropkoy, lL. A, (1960).—

Osnovy Paleontologii. Spravochnik dlia pale-

onlologoy i geologoy SSSR. Molluski-Bruko-

nogie. (Moscow.)

Pirspry, H. A, & McGinty, T. L, (1945),—

Cyclostrematidae and Vitrinellidae of Florida.

Pl. 1 & UL. Nautilus 59, (1) 1-13, pls 1-25 (2),

52-59, pl. 6.

Rave, D. M. (1966) —Gcometric analysis of shell

voiling: general problems. J. Paleant. al,

43-65.

Reuprr, AH. A. (1935).—"New Caribbean Marine

Shells.” Nautilus 48(4), 127-30, pl 7.

Sacco, FL (1892)—T Molluschi dei Terreni Ter-

ziari del Piemonte é della Liguria, Part 12.

(Clausen: Torino.)

Suter, H, (1913-15),—"“Manual of the New Zea-

land Mollusca.” (MacKay: Wellington. }

VERRILL, A. E. (1885),—Third catalogue of Mol-

fusca recently added to the fauna of the New

England Coast and the adjacent parts of the

Ailantic, consisting mostly of deep-sea species,

with notes on others previously recorded.

Trans. Cannect. Acad. Arty Sei, 6, 395-452,

pls 42-44.

Wenz, W. (1939).—Gastropoda. Ji O. H. Shinde-

wolf (Ed.), “Handbuch der Palaiozoologie”.

Vol. 6, Part 3(4). (Gebruder Bornitracger;

Berlin.)

THE CLINGING MECHANISM OF PSEUDOPHRYNE BIBRONI

(ANURA: LEPTODACTYLIDAE) TO AN ALGA ON GLASS

BY N. 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 BIBRONI

(ANURA: LEPTODACTYLIDAE) TO AN ALGA ON GLASS

by N. GRADWELL*

Summary

GRADWELL, N. (1975).—The clinging mechanism of Pseudophryne bibroni (Anura: Lepto-

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

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.

Introduction

It is well known that anuran tadpoles are

adapted to occupy a wide variety of ecological

niches (Noble 1931, Orton 1953). Of the mor-

phological adaptations, an attachment mechan-

ism for clinging to substrata is a predictable

association with a lotic habitat. It is thus pos-

sible to avoid dislodgement by swift currents

even in torrential streams. In contrast, the ad-

hesive secretion produced by the ventral glands

of most young embryos (at stages 18 to 24, of

Gosner 1960) is too weak to withstand fast

currents (Gradwell, unpublished). There is no

published evidence that these glands can pro-

duce subambient hydrostatic pressures. They

atrophy after stage 24, and in the tadpole which

represents stages 25 to 40, either of two kinds

of non-glandular sucker may develop.

An oral sucker is the most usual adaptation

in lotic tadpoles and, accordingly, the degree

of development of this sucker would seem to be

influenced by the velocity of the ambient water.

In most lotic species the periphery of the upper

and lower lips is continuous and forms a suc-

torial disc, and the number of tooth rows in-

creases progressively with the velocity of the

ambient water (Noble 1931; Gradwell, unpub-

lished). The structure and function of a tad-

pole’s oral sucker has been described in

Ascaphus truei (Gradwell 1971, 1973) and it

has been compared with the suckers of five

Australian species (Gradwell 1975). In addi-

tion, Tyler (1963) described the external

appearance of the sucker of Litoria arfakiana

(as Hyla angularis), which is much like that

of Ascaphus.

Another type of tadpole sucker is that which

lies posterior to the mouth. The anatomy of

such a sucker has been described in Sraurois

ricketti by Noble (1931) and in Staurois

afghana (Bhaduri 1935). No data are available

on the magnitude of its suction, except that if

a tadpole is manually lifted out of the water by

its tail, its sucker can support a stone sixty

times the weight of the tadpole (Hora 1922).

The present paper reports a mechanism em-

ployed by lentic tadpoles of the Australian lep-

todactylid frog Pseudophryne bibroni which en-

ables them to cling to a vertical substratum,

even though they lack a sucker. I have ob-

served such ability in nature for many other

suckerless tadpoles and have assumed that they

use their teeth to secure a grip on rocks.

Results and Discussion

On 18 July 1974, 16 tadpoles of Pseudo-

phryne bibroni (at stages 26 to 29) were col-

lected in a flooded ditch (ca 30 m long by 5 m

wide) with sides sloping to a depth of 2.5 m.

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.

N. GRADWELL

wes

+ %

~~ :

Fig. 1. A. The tadpole of Pseudophryne bibroni (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 Phyllohium 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.

SLBSTRATUM CLINGING IN PSEUDUPNRYNE 33

River Crossing, near Jenolan Caves. N.S.W.

The muaimum depth of water was 0,5 m and

the botiom consisted of mud and fallen Euca-

Ivpttey leaves ancl bark. At 3.00 p.m, the walter

temperature was 6 to B°C. Five hours later, the

tudpoles were placed in an aerated aquanum

at approximately the same temperature,

For initial identification, the mouthparts of

the tadpoles were examined and compared with

the description of Martin (1965), Bycuuse the

ubsenee of a sucker was noted, it was surprising

to observe that all of the radpoles clung to the

vertical glass sides of the aquarium (Fig. LA).

This attachment was insecure, for these tad-

poles were sometimes displaced from their

clinging sites when other tadpoles collided with

them. Examination of the orl region of a

clinging tadpole, by stereomicroscope through

the glass, showed that the lower beak was kept

closed within the edge of the upper beak (Fig,

1B) and that the upper and lower tooth rows

were held adducted, thotigh not touching one

another. The nares were the only inhalent chan-

nets for gill irrigation.

During preparations for photography it was

found that the inside of the glass aquarium was

covered with a thin laver of algae which pre-

Vented sharp focussing on the mouth and teach.

Therefore, half of one side of the aquarium

was cleaned of its algae, and a camera conven-

iently positioved to await the settling and cling-

ing of # tadpole on the cleaned glass. Although

tadpoles ullempled to cling to the cleaned glass,

they were unsuccessful. However, they clung

readily to the tncleaned glass, demonstrating

Ihe presence Of ulgac on the glass to be neces-

sury for the clinging of these tadpoles.

Apurt From clinging to che glass. the tad-

poles also showed frequent feeding movemonts

over the slass dufing which their jaws opened

and closed rhythmically in a scraping action,

Pigure 1C shows the jaws 1 an oped condition

and both the upper and lower hewks ate visible.

The predominant alga growing on the glass was

Phyliokium sp. {identified fram Prescott 1970),

which has a thallus of branched strunus. At the

ends of the strands are swellings culled aki-

netes, which contain chloroplasts. It was of spe-

cial Toterest to find akineles in the maiticotto

(“stomach") of tadpoles, which suggests that

they had been grazed off the glass. $1 would

also seem that ihe bulbous akinetes on their

strund-like thallus could easily be gripped by

the minute sharp teeth of tadpoles, thus provid-

ing anchorage. It is possible that other filamen-

fous algae may also permit the clinging of

these tadpoles to substrata.

There ure no. papillae behind the most pos-

lerior row of lower labial teeth, Therefore this

row of teeth can bend farther forward than if

papillae were present here and so the teelh can

enip substratum plants more tirmly. However,

some other suckerless tadpoles (for example,

Livoria verreauexi) can cling. feebly to vertical

substrata in similar fashion even though they

lack a posterior gap in their peripheral papillae.

Four tadpoles were raised into juvenile frogs.

Until stage 40, the tadpoles (entire length, 36,0

min; snout-to-vent Jengih, 17.4 mm) weee able

ta cling by their teeth to the substratum. Alter

this stage the heaks und teeth were shed prios

to widening of the mouth.

It is proposed here that Psendoplyryste

Ajhron’ ludpoles are able to cling to vertical

subsiraia by biling their teeth inta algae anil

perhaps other vegetation. As the narial inflows

appear to be sufficient for respiration, the juws

need not open and close rhythmically to admit

additronal water, but can maintain their hite

on plants. However, I haye not hail the oppot-

Lunily lo compare these findings with the ability

of P. dibrent in their natural habilal fo cling

lo substrata, ‘The absence of an oral sucker

in these tadpoles confers greater flexibility on

the jaws and perhaps it broadens their food

resources. On the other hand, an oral sucker

in lotic tadpoles facilitates grazing on algac

in swift currents (Gradwell 1971), where more

tenacious gripping is needed.

Acknowledgements

T am grateful ta Miss P, MeDonnell and Mr

B, Wilson, for collecting the tadpoles, ad to

Miss M. Anstis for identifying them, Mr M, J.

Tyler kindly read the manuscript and offered

helpful suggestions,

References

Brapum J. L. (L935)—The anatomy of the

adhesive apparatus in the tadpole of Rana

afvhona Guather, with spectal teference to the

adaptive modifications. Trans, R. Soc. Edin-

horgh 58, 339-349

Goswea, K. L. £1960)—A sitmplificd cable for

stoging anuran embryas and Vrvac with notes

on identification, Herpetelogica 16, 183-190.

Granween, N, (1971).- Ascephus ladpole: experi-

ments on the suction and gill trrigation

mechanisms. Cua. ./, Zool. $9, 407,332.

GRADWELL, N. (1973).—On the functional mar-

phology of suction und gill irrigation in the

tadpole of Ascaphus, and noles on hiherm-

tion. Herpetologica 29, 84-9

34 N. GRADWELL

GRADWELL, N. (1975),—Experiments on oral suc-

tion and gill breathing in five species of Aus-

tralian tadpole (Anura: Hylidae and Lepto-

dactylidae). J. Zool., in press.

Hora, S. L. (1922).—Animal life in torrential

streams. J. Bombay Nat. Hist. Soc. 32, 111-

126.

Martin, A. A. (1965).—Tadpoles of the Mel-

bourne area. Vict. Nat. 82, 139-149.

Nosie, G. K. (1931}.—“The Biology of the

Amphibia.” (McGraw-Hill: New York. Re-

printed 1954, Dover, New York.)

Orton, G. L. (1953).—The systematics of ver-

tebrate larvae. Systematic Zool. 2, 63-75.

PrescoTr, G. W. (1970}.—“How to know the

freshwater algae.” (W. C. Brown Co.,

Dubuque, Iowa.)

Tyter, M. J. (1963).—A taxonomic study of

amphibians and reptiles of the central high-

lands of New Guinea, with notes on their

ecology and biology. I]. Anura: Ranidae and

Hylidae. Trans, R. Soc. S. Aust. 86, 105-130.

GENETIC EVIDENCE FOR THE EXISTENCE OF TWO SEPARATE

POPULATIONS OF RATTUS FUSCIPES GREYIT 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 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-(“ and Got-l’ are present in the animals collected from the middle and

southern sections of the island, while Got-/* 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 RATTUS FUSCIPES GREY! ON PEARSON ISLAND,

SOUTH AUSTRALIA

by L. H. ScHmitt*

Summary

Scumitt, L, H, (1975).—Genetic evidence for the existence of two separate populations of

Ratius fuscipes ureyii on Pearson Island, South Australia. Trans. R. Soc. §. 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 fuycipes

preyii) on Pearson Island, Two allelic genes, Gor-1¢ and Gai-1? are present in the animals

collected from the middle and southern sections of the island, while Gar-/* 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.

Introduction

Pearson Island, which lies 60 km off South

Australia’s west coast, is divided into three dis-

crete sections (Fig, 1), The southern and

middle sections are Jinked 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-

son J, wallaby, Petrogale sp, (see Thomas &

Delroy 1971, for a discussion on its taxonomic

status) and the native rat, Retruy fuscipes

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 Rattus murrayi,

Tredale & Troughton (1934) reclassified it as

Rattus greyli murrayi recognizing its close rela-

tionship to Rattus greyii greyii, the native

bush-rat of mainland South Australia. It was

later included in the species Rattus fuscipes by

Ellerman (1949) as a distinct sub-species, R.

fuscipes murrayi, along with the matoland form

R.j. greyit. Recently, Taylor & Horner (1973a)

have considered it to be subspecifically indis-

tinguishable from R.f. gréyii.

Until 1960, the Pearson I, wallaby was only

found on the northern section of the island. No

evidence could be found to suggest the species

ever inhabited the. other two sections, despite

the suitable habitat available there, The channel

appeared to act as a very effective barrier to

NORTH

SECTION

MIDDLE

SECTION

**SOUTH

SECTION

250 500

metres

Fig. 1. Map of Pearson Island. Areas where ani-

mals were captured are indicated by shad-

ing.

* Department of Genetics, University of Adelaide, S. Aust. 5000.

36 I.

migration between the northern und middle

sections. In 1960. six wallabies, including either

four of ive females, were accidentally released

on the middle section and the species is now

abundant on the middie and southern sections

(Thomas & Delray 1969).

This paper describes a genetic difference in

the Af. grevir population of Pearson 1, appa-

renily caused by a restriction in migration

across the channel separating the northern and

middle sections,

Methods

Specimens of A. greyli were caught in

“Sherman” traps and transported alive ta Ade-

faide. ‘issue homogenates were prepared and

subjected to starch gel electrophoresis. ‘lhe

vlectrophoretic bullers (pH 8.0) were essen-

tially the same as those described by Sclunder

eal (1971). Vhe methads used to detect glu-

tamate oxaloacetate transaminase (GOT) wet

vity and determine the subcellular locality of

the isozymes were modified from those given

hy De Lorenzo & Ruddle (1970), Heart ex-

tracts were used predominantly, except in the

latter procedure when liver extracts were used.

Results and Discussion

Animals were caught from the arcas shown

in Fig. 1. Seventy-five animals were caught in

218 trap nights, yielding a capture rate of

*4%%. ‘This is considerably higher than the

2.8% return obtained by Taylor & Homer

11973b) for Rf. greyvii on the mainland of

Australia ahd Kangaroo 1.

Two main tegions of GOT activity were pre-

sent in gels, one migrating cathodally, the other

anodally, The cathodal area of activity con-

sisted of » single Invariant band, This isozyme

predominated in mitochondrial extracts. The

anodally migrating isozyme was found in the

supernatant fraction and was variable. Three

distinct phenolypes, GOT-1A, GOT-1B and

GOT-LAB were observed. This variation is

consistent with the active enzyme heine a

dimeric molecule and is similar to that found

in man (Chen & Ciblett 1971) and the North

Ameticun old-field mouse, Peremprecns polio-

nots (Selander et al. 1971). Genotypes can

he ussigned to cach phenotype, presuming that

the difference is under the control of an auto-

somal locus, With wo co-dominane alleles. This

locus has been designated Gat-/ and the alleles

Go-l" aud Gar-l?,

Laboratory matings of &.f. greyi? individuals

from differen. ureas of South Australia. inehad-

ing Pearson J., have been successful, Family

H SCHMITT]

TABLE 1

Family dara en GOT variation

Sinber of

HOD slwnatyne Nomber and GOT phenytype

uF psrents manne wl gitsoring

AB B

AXA 2 3 0 ()

AxA* Lp 44 a 0)

AXxB a 0 9 0

Ax AK | 2 a ()

HxAAB | {) 4 o

* One patent was nol scored for GOP phenotype.

However, in cach of the ten matings it was

known to have come from a population appa-

tenily monomorphic for the Gated? allele.

dala on the inheritance of the GOV Variatinn

(Table 1) is cosistenr with a 1 locus, 2

allele mode of inheritance.

The &.f. greyii population on the northem

section is monomorphic for the Get /? allele,

while both Gai" and Got 1" are present in

the population om the southern and middle

seclions (Table 2), The genotypic frequencies

in the latter populavion fit the Hardy-Weinberg

equilibrium frequencies (P>0.05), HW the

Got-i" allele is present in the population on

the northern section, then there is a 95% pro-

bubilny that its Frequency is loss than 3%, Tn

any case. the frequency of the Got-/! allele in

the northern section is. significantly different

fram its value in the middle and southern ser-

tions (P< = 0,001).

TABLE 2

GOT phenotype distribution in animals

eanght on Pearson 7,

Number and GOT phenotype Frequency

4 AB B

Ajc@ tanebe of Goi-lb

Northern sztrion 0 ii} 49 0G

Midd'e and sotunerny wy 10 4 W375

schon

li seems unlikely thay the marked difference

in allelic frequencies is inuintained by selec-

lion, All three sections of the island appear to

provide very Similar habitats fur ALA previt.

The observed absente of Geri" from the

northern secuion Indicates a severe restrichion in

gene flow between the arcas sampled on the

middie und northern sections. ‘The mast

obvious point of demarcation is the channct

xeparaling the two sections. Phis surprisingly

low level of migration between the northern

und middle sections is similar ro rhac found in

the Pearson 1. wallaby.

Tr woulul appear improbable that the rats are

not physically capable of crossing the channel

When the sea is calm and the tide low, it is

GENETICS OF THE PEARSON ISLAND RAT 37

saat

A AB A B A AB

Origin

Fig. 2, GOV varation found in R.f. grevii from

Peurson 1. The pattern of variation is in

agreement With a dimeric molecule being

the active enzyme.

easy for a man to wade or step from rock to

rock between the two sections. However, the

conformation of the channel is probably in a

state Of continual change. The relationship be-

tween sea level changes and the occurrence of

one or more species of small macropod mar-

supiils on small islands off the Western Austra-

lian coast has been discussed by Main (1961).

From Main’s data it appears that Pearson [.

has been isolured from the mainland for at least

10,500 years, The northern and middle sections

would have remained connected for a consid-

erable time after the isolation of the island

There is some evidence to suggest that sinee

the isolation of Pearson Island from the main-

lund, the mean sea level on two or more occa-

sions has been about 6 metres above ils present

level (Twidale, pers. comm.). During these

times of high mean sea level, the channel would

have been considerably larger thin at present.

Thomas & Delroy (1971) suggested that the

wallaby did not cross the channel because it

found the sea water distasteful. Another possi-

bility 18 that, because of 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 un hypothesis.

genolypes would be favoured which predis-

posed animals to an aversion to moving close

to the water's edge. This would discourage the

animal from crossing between the middle and

northern sections.

Various suggestions may he offered to

uccount for the present distribution of the two

alleles at the Goi—/ locus. All other popula-

tions of R.f. greyii studied (Greenly 1, Hop-

kins L., Kangaroo L, Eyre Peninsula and Mount

Lofty Ranges) have been found to be mono-

morphic for the Gor] allele (Schmitt, unpub-

lished), The polymorphism on Pearson I. may

have been present before (he channel was

formed and at that time, or some time after-

wards, the Got-/* allele was lost from the

northern section. Alternatively, one of the

ulleles could have arisen by mutation, since the

channel was formed. If the Got—/” allele is the

more recent mutunt, then it has to be postu-

lated that it subsequently migrated across the

channel.

The channel separating the northern and

middle sections has not only aeted as an effee-

tive barrier to migration for the Pearson T.

Willaby. but from the genetic evidence pre-

sented here. hus also had au similar effect on

Ruf. greyii.

Thirteen other gene loci, for which about 70

specimens of R-f. grevif from Pearson Ll. have

been scored, are monomorphic on Pearson

Island and show no differences across the chan-

nel. However, all Pearson [. animals have

haemoglobin and malate dehydrogenase pro-

teins that are electrophoretically distinct from

all other populations studied (Schmitt, unpub-

lished).

Further studies on the situation described

here would be a useful part of uny future expe-

ditions to Pearson 1.

38 L.

Acknowledgments

I wish to thank Dr D. L. Hayman, Dr R. M.

Hope and Professor J. H. Bennett for their

thoughtful discussion and help in the prepara-

tion of the manuscript. Transport to Pearson

H. SCHMITT

Island was generously arranged by Mr M.

Koch and provided by Mr R. Baker and Mr K.

White. The South Australian National Parks

and Wildlife Service granted a permit to allow

me to collect specimens.

References

Cuen, S. H., & GipLetr, E. R. (1971).—Genetic

variation of soluble glutamic-oxaloacetic

transaminase in man. Am. J. Hum. Genet. 23,

419-424,

De Lorenzo, R. J., & RuppLe, F. H. (1970).—

Glutamate oxalate transaminase (GOT)

genetics in Mus musculus: Linkage, poly-

morphism, and phenotypes of the Got-2 and

Got-I loci. Biochem. Genet. 4, 259-273.

ELLERMAN, J. R. (1949).—‘The families and

genera of living rodents.” Vol. 3, pt 1. (Bri-

tish Museum (Natural History), London.)

IREDALE, T., & TROUGHTON, E. LEG. (1934).—A

check list of the mammals recorded from

Australia. Mem. Aust. Mus. 6, 1-122.

Main, A. R. (1961).—The occurrence of Macro-

podidae on islands and its climatic and eco-

logical implications. J. R. Soc. W.A. 44,

84-89.

SELANDER, R. K., SMITH, M. H., YANG, S. Y.,

JOHNSON, W. E., & GENTRY, J. B. (1971).—

Biochemical polymorphism and systematics in

the genus Peromyscus. |, Variation in the old-

field mouse (Peromyscus polionotus). Stud.

Genet. VI, 49-90. (University of Texas Pub-

lication 7103.)

Tay Lor, J. M., & Horner, B. E. (1973a).—Results

of the Archbold expeditions. No. 98. System-

atics of native Australian Rattus (Rodentia:

Muridae). Bull. Am. Mus. Nat. Hist. 150(1),

1-130.

TayLor, J. M., & Horner, B. E. (1973b).—

Reproductive characteristics of wild native

Australian Rattus (Rodentia: Muridae). Aust.

J. Zool. 24, 437-475.

Tuomas, I. M., & DELRoy, L. B. (1971).—Pearson

Island expedition, 1969. 4. The Pearson Island

wallaby. Trans. R. Soc. S. Aust. 95, 143-145.

Tuomas, O. (1923).—The native rat on Pearson’s

Islands, S. Australia. Ann. Mag. Nat. Hist.

Ser. 9, 11, 601-602.

TWO NEW SPECIES OF THE GENUS CLOACINA (NEMATODA:

STRONGYLIDA) FROM THE TAMMAR, MACROPUS EUGENIT

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 CLOACINA (NEMATODA:

STRONGYLIDA) FROM THE TAMMAR, MACROPUS EVIGENIT

by Patricia M, Mawson *

Summary

Mawson, P, M. (1974)—Two new species of the nematode genus Cloucina (Nematoda:

Strongylida) from the Tammar, Mucropus exgenti. Trans. R. Sac. §. Awst. 99(1), 39-41,

28 February, 1975.

Cloacina smalesae and C. kartana are from the stomach of Macropus eugenii from

Kangaroo I. C. snialesae is distinguished by the shape of the leaf crown elements, postoeso-

phageal excretory pore. swollen cuticle at the anterior end of the body, and a long vigina.

C. kartana is distinguished by the very small cephalic papillae and the presence of two

oesophageal teeth situated shortly behind the nerve ring.

Introduction

The species which are described in this

Paper Occur commonly and in considerable

numbers in the stomach of the Tammar on

Kangaroo Island. They were taken in the

course of a quantitative gnulysis by Mrs

Lesley Smales of the nematodes present ut dif-

ferent times of the year in the stomach of this

hest, undertaken as part of work for a» Ph.D.

degree in the Depariment of Zoology.

Types of the new species will be deposited

ia rhe South Australian Museum: paratypes are

in the Helminthological Collection of the Zaa-

logy Department, University of Adelaide.

Measurements of the two species are given

in Table 1,

Cloacina smalesae 1. sp.

FIGS 1-7

Hast and Locality; Macropes cugenui (Des-

marest), from Kangaroo T., 3. Aust.

This is a medium-sized spectes af Cloacirid,

with markedly swollen cuticle at the anterior

end. The thickness of the cuticle increases

from the level of the posterior end of the oeso-

phagus to the base of the mouth collar, which

it surrounds Itke a platform. The mouth collar

is well develaped and slightly lobed anteriorly;

it bears the four submedian cephalic papiilae

and two amphids, The distal segment of each

cephalic papilla is longer and slightly thicker

than the proximal segment, Each clement of

the Jeaf crown is domed anteriorly and is witb-

out the unguiform anterior projection usually

present in species of this genus.

The buccal ring is circular and stoutly built.

Its anterior and posterior walls are slightly

lobed (Figs 1, 2). The oesophagus is cylin-

drical for most of its length, ending in a small

swelling. There are no oesophageal teeth. The

thickened lining of the lumen is unusually clis-

tinct, appearing in the whole mount as three

wavy longitudinal lines throughout the length

of the oesophagus. In transverse section the

lumen appears triradiate with a thickening of

the lining in the midlength of each arm. The

nerve ring lies at about midlength of the ocso-

phagus. The excretory pore is postoesophageal,

and the cervical papillae lie a little distance in

front of the nerve ring.

The bursai lobes are onty slightly divided.

The arrangement of the rays (Fig. 6) 1s

typical for the genus, The genital cone is not

prominent; it bears a small ala on cach side of

the cloaca, but no other appendages. The

spicule is a little less than half the body length,

The tail of the female is conical, ending in

a point. The vulva is slightly less than a tail

length in front of the anus, The ovejectors unite

6-7 tail lengths in front of the yulva, and the

vagina curves forwards before passing back,

with one or two twists, to the vulva.

The rounded elements of the leaf crown

seen in this species have been described in only

one other species, C. /ongifabiata Johnston &

Mawson, 1939b (syn: C. minar J. & M., nec.

Davey & Wood, 1938), However, in C, /ongi-

labiaia the vagina is shoster, the cephalic papil-

lae are of a different shape and there is no

cuticular inflation ai the antenor end.

* Zoology Department, University of Adelaide, S Aust. 5000-

4) PATRICIA M. MAWSON

50 an

—

200 pm

Cloucina smidlesae. Fig. 1—Anterior end, lateral yiew. Fig. 2.—Anterior end, showing

leaf crown in everted position. Fig- 3—Anterior end, en face. Fig. 4—Oesophageal region,

Fig. 5.—Transverse section of vesophagus, anterior to nerve ring, Fig. 6.—Posterior end

of female. Fig. 7—Bursa, flattened out and viewed from the inside. Fig. 6—Lateral view

of genital cone. Figs 1, 2, 3, 5 and 8 to same scale,

Figs 9-15,

Cloacina kartana, Figs 9 and 10—Sublateral and en face views of anterior end, Fig. 11.—

Oesophageal. region, Fig. 12.—Transverse section of oesophagus, showing one oesophageal

tooth. Fig. 13.—Posterior end of female. Fig. 14.—Bursa. Fig. 15.—Dorsal ray. Migs 10

and 12 lo samé scale,

Cloacina kartana n. sp-

FIGS 8-I4

Hast and Locality; Macropus eugenii (Desma-

Test), from Kangaroo I., 8. Aust.

This is a medium-sized species of Cloacina

with a well developed mouth collar bearing the

four small submedium cephalic papillae and

the amphids, The cephalic papillae are small;

the proximal segment is longer and. slightly

thicker than the distal one, The bucciul ring is

stoutly built, and is thicker posteriorly than

anteriorly. The six elements of the leaf crown

do not, in the resting position at Icast. project

above the collar.

The oesophagus is cylindrical for most of

its length, with a terminal bulb. There are two

small subyentral oesophageal teeth, one a little

TWO SPECIES OF CLOACINA

Measurements of Cloacina smalesae and C. kartana; in vn unless otherwise stated.

C. smalesae

Length (mm) 9.5-11.4

Ocsophagus 690-720

Antr. end—nerve ring 360-400

—cerv. pap. 200-230

—exer, pore 790-870

Spicule length 4200-4700

Length/spic. 1, 2.2-2.5

ERB DESCDN 13.8-16.0

‘ai =

Postr,—vulva

TABLE 1

C. karfana

P 3 2

13.0-15.8 8.0-10.9 12.4-17.0

710-790 680-790 810-900

360-490 300-350 340-400

160-200 220-300 250-300

910-1000 600-740 700-800

— 1400-1600 —

— 5.7-7.1 _—,

16.8-21.8 10.6-16.0 14,3—20,9

220-260 — 200-300

350-490 — 380-500

Egg length x breadth —

posterior to the other, at about midlength of

the oesophagus, and shortly behind the nerve

ting. The cervical papillae lie just in front of

the nerve ring, and the excretory pore is at the

level of the oesophageal bulb.

The spicules are relatively short, The lobes

of the bursa are distinct but not deeply sep-

arated. The rays are arranged as shown in

Figs 13 and 14. The genital cone is small and

no appendages were seen on it.

The female tail is conical, ending in a nar-

rowed point. The vulva lies a little less than

170-175x85—90

180—185x85-90

a tail length in front of the anus; the vagina,

which is straight, extends about 3-4 tail lengths

in front of the vulva.

The species most closely resembles C. obtusa

Johnson & Mawson, 19394, but differs in the

spicule length.

Acknowledgments

My thanks are due to Mrs. Smales for the

collection of the specimens described. Part of

the work was done under a Grant from the

Rural Credits Development Fund.

References

Davey, D. G., & Woop, W. A. (1938).—New

species of Trichoneminae (Nematoda) from

Australian kangaroos. Parasitol. 30, 258-266.

Jounston, T, H., & Mawson, P, M, (1938).—

Strongyle nematodes from ceatral Australian

kangaroos and wallabies. Trans. R. Soc. S.

Aust. 62, 263-286.

Jounsron, T. H., & Mawson. P, M. (1939a).—

Strongylate nematodes from marsupials in

New South Wales. Proc, Linn. Soc, N.S.W.

64, 513-516.

Jounston, T, H.. & Mawson, P. M. (1939b).—

Sundry nematodes from eastern Australian

marsupials. Trans, R. Soc. S§. Aust. 63,

204-209.

THE GENERAL WATER CIRCULATION OF SPENCER GULF,

SOUTH AUSTRALIA,

IN THE PERIOD FEBRUARY TO MAY

BY D, 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. BuLLtock*

Summary

BuLLocK, BD, A, (1975)—The general water circulation of Spencer Gulf, South Australia, in

the period Februury 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 Curent”, 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

Ao investigation of the general water circu-

lation in Spencer Gulf, and the region outside

it, is mude below from observation of property

distributions and using a numerical model. The

term “general water circulation” is used so that

tidal currents which basically involve a periodic

north-soulh movement of the Gulf’s waters are

exeluded. Tidal currents can be superimposed

on the more sustained flows caused by thermo-

haline (and wind) forcing which are particu-

larly important because of the part they play

in the interchange of water in a substantially

enclosed body of water. Thermohaline currents

are brought about by horizontal pressure gra-

dients due to density variations in the Gulf.

which are caused by temperature and salinity

differences, produced in part by outside water

entering the Gaull, and in part by variation in

the effect of evaporation over the Gulf.

All available data for the region, obtained

on C,S\1.R.0. cruises, are shown in Fig. 1, The

crises took place between 1961 and 1946 cdur-

ing the months indicated in the caption. No

direct curren, readings. are available, although

some results on surface drift do exist from a

drift card experiment (Marshallsay & Radok

1972).

Presentation and Interpretation of Cruise Data

Using measurements made by F.R,V. Inves-

figator (C.S.I.R.O. Aust. 1968c) on 7 May

1964, a vertical salinity section (Fig. 2a) was

drawn for a line across the mouth of Spencer

Gulf from Port Lincoln on the western side to

Corny Point on Yorke Peninsula. The salinity

is approximately homogeneously distributed

over depth for any particular watec colunim,

although around the centre the situation is

closer to that of a two-layer system since the

salinity is almost constant with depth until

about 20 m, at which point a steep salinity

gradien( occurs. Below this, the salinity is

again constant with depth but greater than

before. Also, the salinity increases with dis-

tance from west to east, indicating that water

enters the Gulf on the western side and leaves

via the eastern side after its salinity has in-

creased due to evaporation and mixing. The

density section (Fig. 2b) has a similar form to

the salinity section, indicating that salinity ts

the dominant factor in determining thermo-

haline flow.

‘Lhe most important feature of the hydrology

of Spencer Gulf. evident from all data (Figs

10, 17), is that the water is approximately

homogencous fram tep to bottom. and hence

* The Flinders Institute for Atmospheric and Marine Sciences, Flinders University of South Australia,

Bedford Park, S. Aust, 5042. Present address: Dept- of Civil Engimeerig, University of Melbourne,

Parkville, Vic. 3052,

D. A. BULLOCK

ws, epi

TA een Ne ont”

”

a s

& K

as a

AA -

% pF

WATER CIRCULATION OF SPENCER GULF

STATION Hay

yo?

OEaTHiMl

DE=TH\ MI

Fig. 2. Vertical sections across Spencer Gulf near

the mouth, Marked — **' — on Figure 1,

data taken from CSIRO Aust. 1968, samp-

ling depths indicates by dots,

(a) Salinity (45), (b) wy.

itis reasonable that the quantities be repre-

sented by their depfh-average, Fig, 3a-c shows

horizontal depth-average distributions of salin-

uy, temperature and density drawn from data

obtained in the period 4-7 May 1964

{C.S.1,R.0. Aust, 1968c). Although other mea-

surements have been made in Spencer Gulf,

this cruise was by far the most comprehensive.

From experience it was found necessary to

restrict the data used to that obtained on one

cruise. whieh spans only a short duration, so

that seasonal and shorter term variations in

properties do nol cause misleading interpreta-

tions to be made.

Fig, 1.

key to Symbols

The distributions shown in Fig. 3 are con-

sistent with those in Fig, 2 and give additional

information. A flow up the western shore is

evident and it can be scen that the salinity im-

creases eastward over the whole southern area

of the Gulf indicating that the movement of

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°45’8, Above this region where the

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 part indicate that

the water moves as a mass in phase with the

tidal currents. At 34°00’S, 137°10°B, two sta-

tions (76 and 89) were occupied, separated by

i period of 29 hours during which the depth-

average salinity changed from 38.40", to

38.14%j,. At 33°57'S, 137°25'E (stations 77

und 85), the salinitites changed from 38.35%;,,

to 38.604, over a period of 21 hours. Using

the horizontal salinity gradients at this location,

Hig. 3 shows that this salinity variation corres-

ponds to a tidid excursion of a few kilometres

The mechanism responsible for the homo.

geneily of the water columns is based on ver-

ical mixing promoted by tidal currents, which

are amplified by the shallow water, running

over a rough bottom which enhances. vertical

diffusion by the shear effect (Bowden 1965).

The temperature variation over the Gulf is

small (Fig, 3b); a plausible explanation for this

observation is the following, Since, in the north

of the Gulf, the advective exchange of pro-

perlies is small and the water shallow, the

ellects of evaporation are more marked than

further south, os is reflected by the high sulini,

Locution of Oceanographic Stations in the South Australian Gulf System until 1973.

H.M.AS. Gascoyne (G2/61) Feb.-March 1961 (CSIRO Aust 1966)

® ALM.A.S. Gascoyne (G5/65) March-April 1965 (CSIRO 1967b}

* HM.A.S, Gascoyne (G2/62) June-Aug. 1962 (CSIRO Aust. [967¢}

* HM_AS. Gascoyne (G2/65) Feb, 1965 (CSIRO Aust. 1968a)

+ HM.AS, Diamantina (Dm2/66) April 1966 (CSIRO Aust. L969)

2 F.R,YV. Investigator 1963 (CSIRO Aust. (968b)

« F.R,V. Investigator 1964 (CSIRO Aust. 1968c)

*« F.RV_ Investigator Dec. 1962 (CSIRO Aust. 1968d)

& TR.V, Investigator Feb. 1965 (CSIRO Aust, 1968e)

~ EV, Estelle Star April-May 1965 (CSIRO Aust. 1968f)

* FY, Estelle Siar April 1966 (CSIRO Aust. (968g)

a FRY. Weerutta fan-March 1961 (CSIRO Aue 1968h)

4h D, A. BULLOCK

Hig. 3, Horizontal depth-average distributions in Spencer Gulf. Data taken from CSIRO Aust. 1968c.

(a) Salinity (%-), (b) Temperature (°C), (ct) oe.

ties. The process of evaporation requires: the

extraction of latent heat of vaporisation which

in turn lowers the temperature and nullifies the

elfect of direct heating. The following rough

culculation shows that there is an approximate

balanve between the heat input from the aimos-

phere and the heat output due to evaporation.

From Haney (1972), the heat flux/unit area

is proportional to the difference between the

upparent air temperature and the sea tempera-

ture (denoled by T,* und T,, respectively).

Assuming this flux is all available for evapora-

tion, we obtain

E=A(T,* — Ty)

Where E is the evaporative heat flux/unit area,

and approximately & = 34 Wm-=(°C)-! (Haney

1972). Taking the values, 'T,,* = 21°C (the

mean temperature for carly April), and E =

100 Wnri?, and substituting in equation (1)

yields T ~= 18°C, which is approximately the

observed yalue (Fig. 3b). The neglect of advec

tive exchange is justifiable in view of the ob-

served small variation of temperature and

closed nalure of the circulation in the northern

Gulf, Vertical mixing muinlains the tempera-

iure difference between the atmosphere and the

sea surface.

Logations of the stations are shown by the

dots in Fig. 3a-c; stations were not occupied

in some areas, notably above Hardwick Bay on

the eastern side of the Gulf. Nevertheless, the

overall pattern of water properties can be in-

ferred reasonably accurutely,

The Numerical Model

In order to investigate whether the above cir-

culation could be considered as primarily due

to thermohaline forcing or whether some other

cause such as wind is dominant, and discover

in greater detail the dynamics of Spencer Gulf,

a numerical model was used to derive theore-

WATER CIRCULATION OF SPENCER GULF

tically the field of flow from the density field

shown in Fig, 3c. The model entitled FLOW-

DEN computes the currents generated in a sea

of density independent of depth, by wind and

variations in atmospheric pressure, and by the

horizontal pressure gradients induced by the

density field’, This is done by solving the finite

difference fepresentation of the component

transport equations written in a form applicable

to the terrestrial situation?.

au on. gx H? dot , tsx | mn

a 7 MBH eax tp, 7 Kluelee (2)

av on, H* Boe , tsy

ge AU - a Gy Bt ay to lua vg tal

where the following notation is used;

7 7

U= | udz, VW= | vdz.

—H —H

u ~ eastward component of velocity (east = x direction)

Vv = northward component of velocity {north = y direction)

Uy. ¥_ = xX & y components of bottam yelogity (ug)

z — distance measured perpendicular to earth's surface, positive upwards

a, = (P—I) a 104. ein kel

f = Coriolis parameter = 2~cosi, » = angular speed of the earth’s rotation. # = latitude

H > depth

ge = 10% egnog, 83500 = specific volume of the standard ocean

& — gravitational acceleration

n -- sea sutfacc elevation

K = bottom dray coefficient

ae l

density of a standard acean = -

5.0.0

The ‘terms in equations (2) and (3) have the

following physical meanings: the left-hand

sides arc the linearized accelerations, the first

terms on the right-hand sides are the Coriolis

accelerations, the sccond terms are the hori-

zontal pressure gradients due tg elevation af

the surface, the third terms are the thermo-

haline terms which take into account the hori-

zontal pressure gradients due to the inhomo-

geneity of the horizontal salinity and tempera-

ture distributions. The final two terms are stress

terms describing wind and bottom friction res-

pectively-

The magnitudes of the thermohaline terms

were evaluated using anather computer pro-

gramme ftom the observed density distribution

(Fig, 3c) and the depths were interpolated

from those given on a naval hydrographic

chirt. A typical value of the x component of

> x & y components of surface wind stress

the thermohaline terms was 300 mm*s-*. This

can be compared with the typical magnitude of

100 mms for the forcing term corresponding

io wind stress, so thal the circulation in Spencer

Gulf due to thermohaline causes can be cx-

pected to be quite significant. The bottem drag

coeflicient K = .0025.

FLOWDEN was run with the precalculated

thermohaline term as the only forcing term and

with the thermohaliné term together with a

constant wind. After the equivalent of 17 hours

an equilibrium current was reached (actually

a small oscillation about the solution remained,

but this was ignored). The initial conditions

were that the How yelocity was zero everywhere

and the surface elevation was. also zero every-

where except at the boundary of the model

actoss the south of the Gulf. which was

assumed to have an initial slope similar to that

1 Bye, J. A. T., 1975. Thallasso-scale numerical modclling. Computer Report. School of Earth Sciences.

The Flinders University of South Australia. (Unpublished. )

2 Bullock, D; A. 1973, The general circulation of St. Vincent and Spencer Gulf. Honours Thesis, The

Flinders University of South Australia. (Unpublished.)

48 Db, A, BULLOCK

developed tater if the adjacent part of the

Gulf. The boundary slope was found to have

litle effect on the current distribution and so

the assumption is presumed to not have serious

cHMsequences,

‘The resultant depth-average current, which

is the transport divided by the depth, was ob-

lumed at cach point of a grid system formed

over the Gulf, in polur form so that the magni-

tude: and direction of the current could be seen

immediately. The grid length was 6.4 km,

rey = Uy BP Bor

ulz) = F*F

. - V_ gt 804

vite) = Ho £ 3x

Applying equatwns (4) and (3) az—7= 0

and at z ~- —H gives the surface and bottom

CUPrents,

Since », is independent of -z it can be seen

Irom equations (4) and (5) that the vertical

homageneity of the Gulf implies that the cur

rent is a linear function of depth z. Thus a

shear current exists. The right-hand side of

equation (4) and (5) can be interpreted as fol-

lows. The second terms are symmetric ybout a

ling marking mid-depth (Fig. 4) und give, as a

function of depth, the current which is directly

reluted to the foeal distribution of density. The

first terms on the right-hand side of the equa-

lions, on the other hand, represent the depth-

uverage current predicted by the model under

the constraints imposed hy the boundaries of

the Gulf (the shores and bed).

The vertical components of velocity were

also found, using the bottony current anc the

gradients of the bottom topography,

oH aH

“a dx * YB By (6)

Finally, the changes. in the sea level which

develop from the initial surface condition of

zero slope ate also obtained,

Results of the Numerical Mudel

(1) Predicted Velocity Fields due ta Thermno-

haline foreing only

Fig. 5 shows the main features of the pre-

dicted depth-average current, The figure is con-

sistent with the water mass analysis of the pre-

vious section, ie. the circulation in the wide,

southern park of Spencer Gulf consists of a

clockwise gyre, water entering via the western

side and legving on the eastern side and the

circulation in the nerthern end appears closed.

Ww =

To gain a detailed picture of the velocity

field in the Gulf. a routine was added to the

program which printed out the surface and bot-

tom curreats. By involving the shallow water

assumption, ie. that the Ekman layer (the str-

face layer influenced by wind) and the layer

ol bollom frictional influence overlap (an

ussuniption whieh is consistent with the waters’

vertical homogeneity), it can be shown (Bul-

lock L973—footnole 2) thal the curren: @om-

ponents at any level are given by,

4th (4)

cr) er

The magnitude of the velocity vecturs shawn is

typically about 0.07 ms? (ranging from vere

to OVS ms),

Prom Figs 6 and 7, which show the surlace

and bottom currents respectively, it can be scen

thar at both the surface und bottum there is a

very distinctive flow up the western side of the

Gulf, which we shall call the “Port Lincoln

Boundary Current”. This current is brouwder in

extent ut the 4uufuce than at the bottom where

it tends to. peel away to the east down the slope

of the bottom. The Port Lincoln Bounary Cur-

rent is also somewhat stronger at ihe surfuce

than at the hettum due to the frictional

influence of the Jatter.

The direction of the bottom current cun he

seen. to sWing eastward and then toa the south-

east over a broad urea before Mowing south-

ward lo exit down the shovel-shaped topo-

graphy through the Gulf’s mouth. The surface

current. displays a similar pattern; however,

since the Port Lincoln Boundary Current is

wider al the surface, particularly near the

mouth, the centre of the gyre of the surface

circulation is located further to the north-east

than the centre of the’hottom gyre, Uvidence of

the gastward movement of water frou the

houndary current is also provided by Fig, 3u—c

where the contours bulge to the east in this

aren.

The differences between the top and botlum

current gyres implies that the surface and bot-

tom currents on the eastern side of the Gulf

are, in general, at an angle to cach other, un-

like the western boundary. This explains why

there is no obvious salinity or % tongue indi-

eating a return flow down the eastern side of

the Gulf in Fig. 3b and c,

WALER CIRCULATION OF SPENCER GULF 49

o |

ra i

——:

Fig, 4. Current as a function of depth, with refer-

ence to mid-depth.

Another important feature to be noted from

Figs 6 and 7 is that in the centre of the mouth

the surface and bottom currents are in oppo-

site directions so that a large shear is predicted

here. The closeness of the density and salinity

sections (Fig. 2b and 2a) to those of a two-

layer system is also explained by this pheno-

menon. It would be interesting to make direct

current measurements in this area.

Further north, where the Gulf starts to

narrow (at about the latitude of Wallaroo). the

flow pattern is more complicated and conver-

PORTAUGUSTA

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 io this interpretation by the

downwelling at the top of the Gulf near Point

Lowly, which corresponds to an inflexion in the

salinity contours (Fig. 3a). In 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

are predicted by the model: in the southern

part of the Gulf, a low develops, the sea level

on the western side being between 10 and 50

mm higher than in the centre; the sea level also

increases to the north. We can conclude that

the circulation in Spencer Gulf which can be

attributed solely to thermohaline causes is of

significant magnitude and forms a major con-

tribution to the water exchange processes of the

Gulf,

(2) Velocity field due te Thermohaline and

Wind Forcing

The numerical experiment was now extended

to inchide the effect of wind stress in addition

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.

D DOWNWELLING

U UPWELLING

WALLAROO

D. A. BULLOCK

Fig. 8. Predicted regions of up- and down-welling due to thermohaline currents in Spencer Gulf.

Fig. 9. Predicted currents in the presence of a 14 ms! SW wind.

(a) Surface, (b) Bottom.

to the thermohaline forcing, The resulting sur-

fuce and bottom currents predicted by the

model for the same density structure as before,

combined with a 14 ms south-west wind. are

shown in Fig. 94 and 9b.

The patiern of circulation is basically the

same but the velocity of the flow is accentuated,

being typically about 0,16 ms", some velocity

vectors having a magnitude over 0.30 ms?.

Differences to the no-wind case do occur in the

northern part of the Gulf where the surface

current is predominantly northward in direc-

tion, particularly on the eastern side where i

was directed southwards previously, Similarly

the hottom current flaws north adjacent io the

shores, Whereas in the no-wind case the bottom

current was slight here. {t can also be seen that

with the wind added, the surface and bottom

currents are almost the same everywhere.

The model predicts that the surface elevation

increases greatly (up to 0,37 m) with distance

up the Gulf, ie. the water is piled up at the

northern end by the wind, The low in the

southern half is also accentuated, the Jevel

being raised by up to 80 mm around the shores

compared with the depression of 10 mm in the

centre.

(3) Poysible reasons for the Persistence of the

Densily Structure

In the numerical model, the thermohaline

forcing terms remained constant throughout the

period fram zero velocity to equilibrium; i,¢,

we assume that the observed densily ficld per-

sIsls and is not an instantiuneous feature which

subsequently runs down until the Gulf becomes

a completely homogeneous body of water in

which water motion of thermohaline origin

would cease,

It is surmised that the cause of the persist-

ence of the thermohaline circulation is external,

i.e. it is due to water from the Suuthern Ocean

moving up under influences that determine the

pattern of flow in this region, perhaps the pre-

vailing westerly winds. Fig. 12, which shows

the dynamic topography of the ocean south of

Australia (Bye 1971). indicates that the direc-

tion of the geostrophic flow in the deep ocean

is, in fact, hasteally perpendicular to the Aus-

tralian coastline.

WATER CIRCULATION OF SPENCER GULF $i

S{ay ion nue

#8 «(0

a7 fen

mW?

Fig. (0, Vertical sections along Investigator Strait

to the Continenlal Slope. Marked —--

on Fivure 1, data taken from CSIRO

Aust. [966 and 1967a. Sampling depths

indicated by dols.

(a) Temperature (°C), (b) of.

The other factor which exerts a continuous

effect on Spencer Gulf is evaporation, which

has the effect of maintaining a high salinity and

henee a high density at the northern end. It is

warth noting that in Fig. 34 the salinity tongue

extending downward from the Gulf's northern

end occurs over a favourabie bottom gradient,

whereas this is not so for the tongue which

Corresponds to the Por Lincoln Boundary Cur-

rent.

(4) The Region Outside Spencer Gulf

The locations of measurements made in the

area outside Spencer Gulf are shown in Fig. 1.

Data from this region were obtained primarily

to aid the fishing industry and suffer from the

drawback that there aré no synoptic surveys of

the whole area, which would enable horizontal

distributions of properties to be drawn. Only

vertical sectiens and profiles can be satisfac-

torily constructed. The paths. along which sec-

tions fave been drawn are shown by the

various lines in Fig. 1.

Fig. 10a and b show temperature and +,

sections drawn along a line extending through

investigator Strait to past the end of the Con-

tinental Slope, The data were obtained in Feb,-

March 1961 (C.S,1.R,O. Aust. 1966) und

March 1963 (C.S.1.R.0. Aust. 19674), One

feature which is naticcable is the change in pitt-

tern from a vertically homogeneous situation

just inside the mouth of Investigator Strait to

the more complicated hurizental stratificution

in the deeper waters outside the Gulf,

The most significant feature shown by these

seclions, however, is the gradient in salinity,

temperature and density about Station 122,

which was located north-east of Cape Borda on

the north-west tip of Kangaroo Island. This

density gradient implies the existence of a

thermohaline current, water to the west of Stas

tien 122 flowing approximately perpendicular

to the plane of the section (into the page). This

direction correlates very closely with the direc-

tion of the thermohaline current entering

Spencer Gulf which forms the Port Lincoln

Boundary Current. Conversely there is also a

southward directed flaw between Stations 121

and 122 which could have its origins in the

outflow from the eastern side of Spencer Gull.

It is possible that upwelling is also occurring

near the longitude of Station 122. The peak in

the density lines for the surface water under

Station 41 is explained by the fact that this sta-

tion was occupicd in a different year to the

adjacent stations (although at the same time

of the year). so that the surface water tompers-

tures ure not consistent.

The scetion shown in Fig. tla and b runs

mughly in a north-east direction from the Con-

finental Shelf up Spencer Gull (C.S.1.R.0.

Aust. 1967a}. The gradient of the lines adja-

cent te the rapidly deepening region outside

Spencer Gulf under Station 64 again indicates

a current inwards across the plane of the sec-

tion, probably angled ap slope so that the water

inundates the more level sea bottom of Spencer

Gulf, ‘This is again in agreement with the pre-

viously deduced direction of flow into Spencer

Gulf which is NNW and thus crosses the Jing

of the section,

Fig, 12 shows that the direction of the circt-

lation in the Southern Ocean is consistent with

the flow in the region outside the Gulf as de-

duced trom Figs 10 and 11,

Conclusion

The various data presented were mainly col-

lected in the months of February to May,

ion

™

station no.

JEPTRIM |

' 1-946 4/922 +542

1 Seed | 4

b-2Sp

z sof i F a / {

5 ah me é' A\

re ae ; / |

iat - | | = ; b

Fig, 11. Vertical sections along Spencer Gulf to

the Continental Shelf. Marked —— on

Figure 1, data taken from. CSIRO Aust,

1967a. Sampling depths indicated by

dots.

(a) Temperature (*C), (b) ¢;.

although during different years, so that it would

appear that the general circulation deduced

occtirs at least seasonally. However, toa gain a

complete and detailed picture of the circulation

in the Gulf, it is obviously necessary to con-

duct a series of thorough cruises at various

times throughout the year. In addition, direct

current measurements nced to be made ta

check the results from the model. Since the dis-

D. A. BULLOCK

2 DB

ten ey

130°E

cre) vase

1 s®

4 pose | i i 4 =i

Fig. 12. Dynamic topography relative to 2000 db.,

oat) of Australia (taken from Bye

1971).

tributions of water properties in St Vincent

Gulf have been found (Bullock 1974) to be

similar to those in Spencer Gulf, it may be

expected that the general circulation in St

Vineent Gulf would also display a pattern

hasically similar to that in Spencer Gulf.

Acknowledgement:

Tam most grateful to Dr J. A. T. Bye for his

guidance and advice during this study. Part of

this material was presented at the Second South

Australian Regional Conference on Physical

Oceanography, University of Adelaide. 9

Avgust 1973,

References

Rawonen, K.G. (1965) —Horizontal mixing to the

sca duc to a shearing current. J. Fl. Mech. 21,

83-95.

Buttock, D. A, (1974).—Cruise Report 1. (Flin-

ders Institute for Almospheric ane Marine

Sciences, The Flinders University of Sautl

Ausiraliit.)

Byrn, §. A. T. (1971).—Yariability south of Aus-

tralia. Proc. Intl Mar. Sei. Symp. Institute of

Marine Science, University of New Sonzh

Wales, 16-17 Auv., I97L. ;

CSIRO Aust. (1966).—Oceanographical observa-

tions in the Indian and Pacific Oceans in

1961. H.M.A.S, Gascoyne, Cruise G 2/61.

CSIRO Aust. Ocvanog. Cruise Rep. 10,

CSIRO Ausr. (1967a).—Oceanographical obser-

vations in the Indian Ocean in 1963.

H.M,.A.S. Gascoyne, Cruise G 2/63. CSIRO

Aust. Oceanvysr. Cruise Rep, 22,

CSIRO Aust. (1967h).—Oceanographical obser-

vations in the Indian Ocean in 1965.

H.M,A.S. Gascayne, Cruise G 5/65. CSTRO

Aust. Oceanogr. Cruise Rep. 46,

CSIRO Aust, (1967c)—Oceanographical obser-

vations in the Pacific and Indian Oceans in

1965, H.M.A.S. Gaseryne, Cruises G 2/62

and G3/62. CSIRO Aust, Oceanogr. Cruise

Rep. 16.

WATER CIRCULATION OF SPENCER GULF 53

CSIRO Ausr. (1967d).—Oceanographical obser-

vations in the Indian Ocean in 1964. H.M.A.S.

Gascoyne, Cruise G2/64. CSIRO Aust,

Oceanogr. Cruise Rep. 34.

CSIRO Aust. (1968a).—Oceanographical obser-

vations in the Indian Ocean in 1965. H.M.A,S.

Guscoyné, Cruise G2/65. CSIRO Aust,

Oceanogr. Cruise Rep. 43.

CSIRO Aust. (1968b).—Investigations by F.R.V.

Investigator on the South Australian tuna

grounds in 1963. CSIRO Aust. Oceanogr. Sin

List 64.

CSIRO Aust. (1968c).—Investigations by F.R.Y.

Investigator on the South Australian tuna

grounds in 1964. CSIRO Aust. Oceanogr. Sin

List 67.

CSIRO Aust. (1968d).—Investigations by F.R.V.

Investigator on the South Australian tuna

grounds in 1962. CSIRO Aust. Oceanogr. Stn

List 60.

CSIRO Aust. (1968e).—Investigations by F.R.V.

Investigator on the South Australian tuna

grounds in 1965. CSIRO Aust. Oceanogr. Stn

List 70.

CSIRO Austr. (1968f).—Investigations by F.V.

Estelle Star in South Australian and New

South Wales waters in 1965, CSIRO Aust.

Oceanogr. Stn List 71.

CSIRO Aust. (1968¢).—Investigations by F.Y.

Estelle Star in South and Western Australian

waters in 1966. CSiRO Aust. Oceanogr. Sin

List 76.

CSIRO Aust. (1968h).—Investigations by F.R.V.

Weerutta on the South Australian tuna

grounds in 1961. CSIRO Aust. Oceanogr. Stn

List 55.

CSIRO Aust. (1969).—Investigations by F.R.V.

Derwent Hunter on the eastern Australian

tuna grounds in 1961. CSIRO Aust. Oceanogr.

Cruise Rep. 54.

Haney, R. L, (1972).—Surface thermal boundary

condition for ocean circulation models. J.

Phys. Ocean, 1, 241-248.

MarsHacisay, P. G., & Rapox, J. R. M. (1972).—

Drift cards in the Southern and adjacent

Oceans. Horace Lamb Centre Res, Report 52.

(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., and Blackburn, D. T. A New Procedure By Mowising

Cleared Leaves Using Polyester Resin - - 55

Womersley, H. B. S., and Conway, Elsie Porphyra and La sees:

phyta) in Southern Australia - - - 59

Smith, Meredith J. The Vertebrae of Four Australian Elapid Snakes - - 71

Tyler, M. J. The Ontogeny of the Vocal Sac of the Australian epee te

Frog Limnodynastes tasmaniensis - - - - - 85

Dodson, J.R. The Pre-Settlement Negeation of the Mt Gambier Area, South

Australia - - - - - = c . - 89

Tyler, M. J., and Martin, A.A. Australian LPR OaneT eS Ero of the

Cyclorana australis Complex - - - 93

PUBLISHED AND SOLD AT THE SOCIETY’S ROOMS

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NORTH TERRACE, ADELAIDE, S.A. 5000

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 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

The vast majority of plant macrofossils col-

lected from Tertiary localities is in the form

of compressed or impressed leaves. Identifica-

tion of these leaves is made difficult by the

fact that most extant correlatives are described

and identified on the basis of floral parts. To

overcome this problem, some palaeobotanists

are now studying extant plants in terms of

their “leaf architecture” (Hickey 1973). For

this type of study, detailed observation of the

higher order veins of angiospermous leaves is

essential, and many workers (Chandrasek-

haram 19721, Christophel 19732, Hickey 1973)

have found it advantageous to develop or

modify techniques for clearing these leaves.

Of the many techniques developed to cope

with this problem (Lersten 1967, Dilcher

1974), most can be divided into two broad

categories. The first may be termed the lacto-

phenol category, the most recent refinement of

which has been presented by Herr (1972).

This technique is characterized by rapid clear-

ing (1-S days) and by preservation of cellu-

lar and cuticular detail. Its disadvantages in-

clude inapplicability to some leaf forms (e.g.

those with extremely tough cuticles or those

which are highly tomentose), and extreme

difficulty in conversion to a permanent mount.

The second category of techniques, which

may be collectively termed the hydroxide cate-

gory, was used initially for the study of cauline

vascular tissue (Foster 1952). It usually com-

bines either sodium or potassium hydroxide

treatment with further bathing in chloral

hydrate or other bleaching agents. This method

is characterized by its relatively long clearing

time (four to ten weeks) and frequent loss of

cellular and cuticular detail. It has the definite

advantages, however, of working on almost all

leaf types (given appropriate time) and of

being readily combined with many permanent

mounting techniques.

The technique presented in this paper is a

modification of the hydroxide method. The

major drawback of the technique in the past

has been the large amount of time needed for

the preparations. For example, to prepare a

large leaf of Cinnamomum takes approxi-

mately four weeks in hydroxide, three days in

chloral hydrate, one day for staining, and an

additional three weeks for mounting and dry-

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).

2 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

]

= is

sgh a .

oe,

ove

narnia

& D. T. BLACKBURN

Fig. 1. Strychnos lucida—leaf cleared by potassium hydroxide-chloral hydrate method. Photographed

with a Leitz Aristophot 4 x 5 plate camera. x 2.5,

Fig. 2. Same specimen as in Fig. 1. Direct print using a Durst 1000 enlarger. x 2.5.

from the date of collection to the date of

storage as a permanent mount,

O'Brien (1974) has pointed out that some

time may be saved by autoclaving the material

in hydroxide. Subsequent to this we have

found that the drying time of nearly three

weeks for mounting media such as Canada

Balsam, Euparal, Permount and Xam can be

cut to less than one day by using polyester

resin. An added advantage of this procedure

is that the monomer of the resin is itself an

effective clearing agent. This not only shortens

the time needed in clearing, but also produces

a specimen with exceptional clarity and re-

producibility for photographic work (Figs 1,

2).

Methods

While the basic technique described below

was originally outlined by Foster (1952) and

incorporates the modifications of others (Chan-

drasekharam 1972,1 Hickey 1973), the pro-

cedure is presented in its entirety here to facili-

tate its usage. The method is equally applicable

to fresh or dried leaves, with perhaps a slight

advantage to the dried material, due to chloro-

phyll breakdown, All times given are approxi-

mate, and will vary with the nature of the

material being cleared.

1. Soak leaves in 15% potassium or sodium

hydroxide solution until they are decolorised,

changing the solution every two days or as

necessary. The most satisfactory vessels for this

are glass petri dishes, with the leaves placed

in them under discs of plastic mesh screening

to prevent flotation. The time for decolorisa-

tion varies with the specimen from four days

to four weeks. Strongly resinous or tanniferous

leaves will usually take longest.

2. Wash the leaves under a gentle stream of

water and transfer to saturated chloral hydrate

until totally clear. This should take from two

MOUNTING CLEARED LEAVES USING POLYESTER RESIN 3

to seven days, with slight warming hastening

the procedure on slow material.

3, Wash leaves again (as in step two) and

transfer lo a dehydration series consisting of

10%, 50% and 90% aqueous ethanol. Finally

transfer 10 absolute alcohol. This series should

take less than one day.

4. Stim m 1% Satranin in 1:2 «absoltite

alcohol/tolaol solution. Stain for three to five

minutes or until all vascular tesue has taken

up colour,

S. Destain the mesophyll in alcohol/toluol

taking cure not to destain the ultimate vena-

ton. This step determines the final contrast

between the mesophyll and the vascular tissue

and is consequently critical. Experience has

shown that mounted leaves having an absolute

absothance of 1.2 to 2.5 at 425 nm yield the

best density for transmitted light photography,

Absorbances were measuted on a Beckman

Acta CIIT Spectrophotometer with the leaves

at the back of the sample chamber. By com-

parison, leaves having an absorbance of less

than 1.0 proved too hght for acceptable repro-

duction, while those with an absorbance of

more than 2.5 proved too dense. As 2 com-

panson, the absorbance of 0.2% safranin

measured over a 1.0 em light path is 1.9, and

a one cm cuvette of this solution may be used

a4 a contro) while destaining.

6, Transfer correctly destained leaves to

acetone for about ten minutes to remove re-

maining aleghol/tohioal.

7. Without allowing the leaf surface to dry,

transfer the leaf to wheatalysed resin mono-

mer’, The monomer acts as a final clearing

agent and normally renders leaves quite trans-

parent in 30 minutes. Should the leaf surface

dry during transfer to the monomer, the result-

ing opacity may be cleared by leaving the leaf

overnignt in the monomer, This monomer bath

is reusable, and consequently this step is best

carnicd out away from direct sunlight and

under fairly cool conditions to prevent poly-

mensation of the resin.

8 Prepare the final mounting medium by

adding abont 5% of MEKP cutalyst to fresh

resin Monomer. This is best accomplished by

mixing in a small glass phiul and rolling

between the hands to combine the two com-

ponents, Such gentle mixing prevents the for-

mation of bubbles in the mountant.

9. Drain the leaf of excess monomer and

mount between Iwo glass slides. Setling should

take about 2 hours at 30°C. Higher tempera-

tures give shorter setting times but increase

the risk of the resin parting from the glass.

10. When the muunt is dricd, excess resin

may be removed with a sharp razor blade or

sealpel, Small particles or thin films may be

removed with xylene or acetone.

Discussion

Having obtaincd « perminent mount of a

leaf wiih maximum transparency, it is then

necessary to reproduce the venation pittern

with as great a degree of contrast as possible

for comparison with fossil material. For nor-

mal photographs, a Leitz “Aristophot” 4 by

5 sheet film camera with a Macro-Dia attach-

ment for transmitted light gave cxcellent

results (Fig. 1), With a largé format camera

such as this it is possible to photograph the

leaf at nearly life size. The negative can then

be contact printed and maximum detail is

oblained.

Since maximum contrast between the veins

and the mesophyll tissue is desirable, however,

the subtle shadings of greys provided by this

methad proved unnecessary. Galavazi (1945)

made brief ceferente to the direct use of an

enlarger for plant material cleared in methyt

benzoate. Dilchee (1974) also successfully

used this technique on skeletonized and

hydroxide cleared Jeaves, This technique was

also admirably suited for our cleared leaves:

The mounted leaves fit easily into the nega-

tive carrier of a Durst 1000 enlarger and the

image can be directly printed on photographic

paper. This technique produces a dark-light

reversed image (Fig. 2), but as the pattern

is the importam aspect, this reversal is

immaterial.

To achieve maximum contrast, a number

of different stains were tried {Morley 1949).

Bismarck Brown gave perhaps the greatest

canirast, but stained very unevenly, Safranin

wave Very nearly eqiial contrast, and had the

advantage of staining much more onifurmly.

Grade four photographic paper was used ta

give the best contrast in printing.

While the above methods approximately

halve the time of the total clearing and mount-

ing procedure, it can still take up te three

For mounting the specimens, Milford EX-80 polyester resin was used, Il is available Trom Mulford

Phistics Pty Lid., 25 Anzac Eighway, Keswick, S. Aust. 5035,

58 D. C. CHRISTOPHEL & D. T. BLACKBURN

weeks for difficult leaves. Work is in progress

using a lacto-phenol clearing method and it is

believed that a procedure has now been found

to permanently mount specimens cleared in

this fashion. Even with this success, however,

the hydroxide method, though longer, still

appears to give better results with a wider

spectrum of leaf types.

References

Ditcuer, D. L. (1974).—Approaches to the iden-

tification of angiosperm leaf remains. Bot.

Review 40(1), 1-157.

Foster, A. 8. (1952).—Foliar venation in angio-

sperms from an ontogenetic viewpoint. Amer.

J. Bot, 39, 752-766.

Gatavazi, G. (1965).—Clearing and staining

plant material in foto with phloroglucinol-

HCl in methyl benzoate for projection photo-

graphy and subsequent serial sectioning.

Stain Tech. 40(1), 1-5.

Herr, J. M. Jr. (1972).—Applications of a new

clearing technique for the investigation of

vascular plant morphology. J. Elisha Mitchell

Sc. Soc. 88(3), 137.

Hickey, L. J. (1973),—Classification. of the archi-

tecture of dicotyledonous leaves. Amer. J.

Bot. 60(1), 17-33.

LERSTEN, N. R. (1967).—An annotated biblio-

graphy of botanical clearing methods. Jowa

State J. Sci. 41, 481-486.

Mor _ey, JT. (1949) ,—Staining of plant materials

cleared in NaOH. Stain Tech. 24(4), 231-235.

O'Brien, T. P. (1974).—Autoclaving as an aid in

the clearing of plant specimens. Stain Tech.

49(3), 175-176.

PORPHYRA AND PORPHYROPSIS (RHODOPHYTA) IN

SOUTHERN AUSTRALIA

BY H. B. S. WOMERSLEY* AND ELSIE CONWAY}

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

Porphyra is common during winter on the

south-eastern coast of Australia, and a brief

account of P. columbina Montagne and P.

lucasii Levring was given by Levring (1953).

Both these species occur on rock or firm sub-

strates and are monostromatic with a single

rhodoplast per cell. Levring also described P.

denticulata Levring from Queensland, recorded

P. naiadum Anderson (now Smithora naiadum

(Anderson) Hollenberg) from New South

Wales, and repeated the original record of

P. woolhousiae Harvey from Tasmania.

Since it is largely a winter form, Porphyra

has often been omitted from ecological

accounts. P. umbilicalis (=P. columbina, from

Cribb 62.5 in ADU) was, however, recorded

by Cribb (1954, p. 30) as forming an almost

pure association during winter and spring, on

fairly rough-water coast at Port Arthur, Tas-

mania, and P. columbina was recorded by

Guiler (1954, p. 64) from Blackman’s Bay

(near Hobart), Tasmania. Womersley &

Edmonds (1958, p. 247) recorded P. colum-

bina and P. umbilicalis (=P. lucasii) as winter

forms, mainly on the south-eastern coast of

South Australia, but sporadically further west.

This paper deals only with southern Aus-

tralian species of Porphyra and the related

genus Porphyropsis. It is hoped that this

presentation of the species will stimulate cul-

tural studies to elucidate further their rela-

tionships.

Genus PORPHYRA C. Agardh

Key to Southern Australian Species

1. Blades delicate, rose-pink, ovate to lanceo-

late, epiphytic on certain brown (or red)

algae in the upper sublittoral .....0000.00.0.0..

P. woolhousiae Harvey

1. Blades lanceolate or ribbon-like, becoming

umbilicate, grey- to red- or dark-purple,

growing on rock or hard substrates .......... 2

2. Thallus fairly tough, retaining its form

when old, shrinking on drying and not

adhering strongly to paper; usually over

45 wm thick; carposporangialt groups

prominent, scattered, with vegetative cells

among the groups; spermatangia occur-

ring irregularly around the margin ..........

P. columbina Montagne

2. Thallus usually delicate, disintegrating

when old, adhering closely to paper and

not markedly shrinking on drying; usually

20-30 ym thick; carposporangial groups

usually not prominent; spermatangia

occurring in (usually) narrow, elongate

strips, extending inwards from the apical

and side margins of the thallus ................

P. lucasii Levring

Porphyra woolhousiae Harvey 1863, pl. 265.

J. Agardh 1883: 59. De Toni 1897: 15;

1924: 12. Guiler 1952: 84. Lucas 1909:

20; 1929: 15.

FIGS 1, 2

* Department of Botany, University of Adelaide, Adelaide, S. Aust. 5000.

+ Department of Botany, University of British Columbia, Vancouver, Canada.

¢ 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. 8. WOMERSLEY & ELSIE CONWAY

Shida Wty

Povphyra woolhousiae Hare,

Blackwans Bey (S. of Hobart), Tas.

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Fig. 1. Porpliyra woelheusiae. A. Blackman’s Bay, Tas. (ADU, A44234). B, Thallus margin, from the

type. C. Thallus with margin (A44234),

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42 H. B.S. WOMERSLEY & ELSIE CONWAY

Thallus (Fig. 14) epiphytic on brown algae

er rarely on red algac, to 17 em high and 4

cm broad, from irregularly ovate or cuneate-

elongate when young, to broadly expanded or

elongate with curved margins which are gently

convolute in older plants (e.g. the type), deli-

cate, with ove to a few blades arising from a

rhizoidal holdfast, rose-red to rose-purplish.

Monostromatic, (12-) 16-24 ym_ thick,

cells isodiametric to slightly elongate in sec-

vional view, each with a single, laminate or

stellate shodoplast. with a pyrenoid, Growth

hy w marginal meristem (Figs 18,C, 2A)

and by ‘ntercalary divisions: margins meris-

tematic, with anticlinal rows of cells; older

parts with cells often in rows but becoming

irregular, with cells often polygonal or angu-

Jar (Figs 18. 28), some rounded to ovoid.

often with adjacent cells unequal in size. Mar

ginal cells isodiametric to slightly elongate,

ungular, 6-10 pm across; cells in older parts

iso-diametric to twice as long as broad, [0-16

{-20) ym across.

Reproductive bodies (Fig. 24,€) formed

in marginal areas, apparenily singly, spherical

te slightly ovoid, 12-!7 ym across, usually

with a stellate rhodoplast.

Spermatangia (Fig. 2D) forming well

defined, irregular, patches near the thallus

margins, the whole patch when mature tend-

ing to deliquesce with numerous spermatangia

irregularly arranged; spermatangia originally

formed in packets of (16-) 32-64, in two

tiers, individually 2-4 4m across.

Type locality; Tasmania.

Type: Herb. Harvey, TCD (presented by

Miss Woolhouse of Sheffield).

Distribution; As well as the type, this species

is known from St Kilda, S. Aust., on Gigur-

tina(’?) on Pastdonie, 14 m below Jow tide

level (8. Lewis, 23.viii 1972; ADU, A42722):

Mallacoota, Vic., on Scyfothamnus australis

(Ducker and King, 151.1970; MEBLU,

20652); Deal L., Bass Strait, on Perithalia

caudata (King, 23.xi1,1969; MELU, 21357);

Curtis I., Bass Strait, on P. caudata (King,

87,1971; MELU, 21358): and from Black-

man's Bay (S. of Hobart), Tas., drift ('yler,

Oct(7) 1974; ADU, A44234),

The type of P. woolhousiae is a well-

developed specimen to 15 on: high, us in the

Blackman’s Bay, Tas., specimett, whereas St

Kilda specimens are 2<4 ¢m high, the Malla-

coota specimens are only 1-2 em high and

those from Bass Strait less than | cm high;

the latter appear to be juvenile. All these,

iicluding the type, show similar cell structure

and presence of marginal growth on young

parts and often also on mature thalli. This

marginal meristem is mot apparent on the

other southern Australian species or on most

other species placed in Porphvra. A Tragment

of the type studied ix not fertile, but the other

specimens show characteristic spermatarigial

groups which tend to deliquesee, and from

marginal arcas the contents of each cell appear

to be liberated as a reproductive body, P.

woolhousiae has a typical Porphyra base with

one or a few fronds attached by rhizoids from

the basal thallus cells, and in most features

agrees well with Porphyra. Uf, however, it is

confirmed that the carposporangia are formed

singly, then relationships with the genus Per-

phyrella Smith & Hollenberg (1943, p. 215)

must he considered, though Conway & Wylie

(1972) have shown that the New Zealand

Porphyra subtumens does not form packets

of curposporangia,

While it is desirable that mature plants on

Mecrocystis, corresponding to the type, be

studied in detail, and their reproduction fol-

lowed in culture, the other records are

sulliciently similar to be placed under P. wew!-

housiae with some confidence. Most are epi-

phytic on brown algic, either on marginal

spines or on spinous branchlets, and this habi-

tat may be characteristic for the species.

P. woulhonsiae has been recarded trom New

Zealand (Levring 1955, p. 412), followed by

D. J. Chapman (1962, p. 129), V. J, Chapman

(1969, p. 20) and Adams (1972, p. 67).

However Adams, following notes of EB, Con-

way in CHR, expresses doubt as to whether

the New Zealand records are not P. coelym-

hina in an abnormal habitat.

The New Zealand specimens growing on

Macroeystis (and possibly Evklonta and Seyto-

thantnas) oeved Turther comparison with the

Australian plants referred to P. woolhorstae.

Ones from Hokio Beach, Levin, N,Z, on

Macrocystis (Moore, 17.%i1.1946: CHR.

55566), determined by Levring (19455, p. 412)

as P. woalhousiae (accompanied by notes of

E, Conway (1971) that they might be young

plants of P, columbina), agree fairly well with

P. woalhoustae as now known from the Aus:

tralian plants. They are similar in form, in

thickness and cell arrangement, and in having

a meristematic margin, but their reproduction

is inadequately known.

PORPHYRA AND PORPHYROPSIS IN SOUTHERN AUSTRALIA 63

Further comparisons between Australian

and New Zealand specimens epiphytic on Mac-

rocystis are clearly needed.

Skottsberg (1923, p. 4) recorded P. wool-

housiae from the Falkland Islands, also on

spines of Macrocystis, and his account shows

similarities with the above description; details

are not adequate for a full comparison. Hamel

(1928, p. 55) recorded it from Kerguelen I.,

and Pujals (1963, p. 8) gives records from

South America.

Porphyra columbina Montagne 1842: 14;

1845: 33, pl. 9, fig. 2. J. Agardh 1883:

70, pl. II, figs 65-66. V. J. Chapman:

1969: 22. Dawson, Acleto and Foldvic

1964: 32, pl. 61B. Guiler 1952: 84.

Hamel 1928: 51. Kuetzing 1849: 693;

1869: 29, pl. 80e,f. Laing 1928: 39, figs

1-7. Levring 1953: 464, figs 2-4; 1955:

410; 1960: 29. Pujals 1963: 8.

Wilhemanig columbina (Mont.) De Toni 1897:

P. umbilicalis sensu Cribb 1954: 30.

FIGS 3, 4

Thallus fairly tough, varying from ribbon-

like (Fig. 34), often with undulate margins,

to broader forms, sometimes furcate, and

usually umbilicate (by loss of upper parts and

basal proliferation) (Fig. 3B) on rough-water

coasts. Thallus often markedly shrinking on

drying and usually not adhering closely to

paper. Variable in size and width, reaching 40

cm long and 30 cm across. Colour varying

from grey-red to red-purple.

Monostromatic, 35-50 ym thick, cells iso-

diametric in section and with a single axile,

pyrenoid-bearing, rhodoplast. Cells 10-15 ym

across in surface view (Fig. 4A), isodiametric

to slightly elongate, more or less in rows but

becoming irregularly arranged, separated by a

gelatinous matrix 4—1 times as wide as cells.

Carposporangial groups prominent (Fig.

4C), of varying shades of red, forming irregu-

lar marginal areas with vegetative cells inter-

mingled, (8—) 32-64 carposporangia per

group, often giving an irregularly granular

(“spotty”) red appearance to reproductive

areas.

Spermatangial groups (Fig. 4B, C) scattered

among the carposporangial groups and in older

plants occupying the marginal part of the

thallus; not occurring in elongate strips as in

P. lucasii.

Type locality: Auckland |. (D'Urville).

Type: PC (Herb. Montagne).

Distribution: From Elliston, S. Aust. to Syd-

ney, N.S.W. and around Tasmania; New Zea-

land, Auckland Islands and other sub-antarc-

tic islands.

P. denticulata Levring from southern

Queensland is probably not distinct from P.

columbina and represents the range extreme of

P. columbina.

P. columbina is the commonest intertidal

species of Porphyra in New Zealand and in

eastern southern Australia, where it occurs at a

lower to mid (sometimes upper) eulittoral

level on rough-water coasts. In Australia it is

essentially a winter form, persisting as late as

December (rarely to February in Bass Strait)

in cooler summers and reappearing in about

May.

Porphyra lucasii Levring 1953: 469, figs 6H—L,

P. umbilicalis sensu Guiler 1952: 84. Womers-

ley 1950: 162.

FIGS 5, 6

Thallus usually fairly delicate, varying from

lanceolate or ribbon-like (Fig. 5A) to broadly

ovate or cordate (Fig. 5B), simple or irregu-

larly laciniate or basally branched, sometimes

becoming umbilicate from basal proliferation,

usually adhering to paper and not shrinking

on drying; to 10 cm long and 15 cm broad,

margin smooth to undulate.

Monostromatic, 20-30 ,m thick, cells with

a single axile, pyrenoid-bearing, rhodoplast.

Cells 8-15u4m across in surface view (Fig.

6A), mostly irregularly arranged, separated

by gelatinous matrix 4—1 times as wide as cells.

Carposporangial groups (Fig. 6C) usually

not prominent, covering areas around the

spermatangial strips toward the margin of the

thallus, without conspicuous intermingled vege-

tative cells, about 8 carposporangia per group.

Carposporangial groups often apparently

developing later than spermatangial strips, thus

giving an impression of dioecism.

Spermatangial groups (Fig. 6B, C) occur-

ring as well-marked elongate strips (Fig. 5B)

extending in from the apical regions and side

margins of the thallus; strips from a few mm

to 2 cm long, 1-5 mm broad. Margin becom-

ing “fringed” following shedding of sperma-

tangial strips.

Type locality: Bunbury, W. Aust.

Type: Herb. Levring, Gotebiirg. (Isotype in

ADU, A42700).

64 H. B. S. WOMERSLEY & ELSIE CONWAY

Par cat

Forphyra

Little Dep, 9f, Rabe), S. Qatt

Pd ah Mea

ving pairing iain aii adie aia

7 2 eos wee aD

ev 0

CoM Wt uhemertey

Fosphyve tolumbing Mont,

Calpe Lamnes, 5 Aas

PA culitterad

PUA WAY MOT WK KO

‘pene EB

y 2 39 £4 £8 F SC @ a

Loh. Dee Bf, omanwling

Fig. 3. Porphyra columbina. A. Ribbon-like forms, early winter. Little Dip, Robe, S. Aust. (Womers-

ley, 15,v.1972; ADU, A42203). B. More umbilicate forms from spring. Cape Lannes, S. Aust.

(Womiersley, 7.x.1972; ADU, A42768).

PORPHYRA AND PORPHYROPSIS IN SOUTHERN AUSTRALIA 65

it. he

TA pas

a> eer

ae"n

sinents

i eeertr

tf etre

a heal

Fig. 4. Porphyra columbina. A. Vegetative cells. Nora Creina, S. Aust. (Womersley, 3.ix.1971; ADU,

A39559). B. Spermatangial marginal area (A39559). C. Area with carposporangial groups and

young spermatangia (A39559).

66 H. B. S. WOMERSLEY & ELSIE CONWAY

Au2,637

For physa Vucasa Leven

Post Stanvac, $ Aust

MA talitrevad wn prac

ve RAD

vn pn ypeeygevapenyjving snag canoe cnn nnqey

OU a ]

, 2 2 * 5 6 * 8 @ Cot RR Lewes

PA BS Wememky

Poxphy ya lucas Levuing

be, 9. Aust.

TPH KD Robe,

vane

TA audittoval, 19 bay nea

“ 2 3 4 5 6 7 8 9 10 x

Bw. ID

Col. + Det. W.B.S. Womenley

Fig. 5. Porphyra lucasii. A. Ribbon-like forms. Port Stanvac, S. Aust. (Lewis, 1.ix.1972; ADU,

A42637). B. Broadly ovate forms showing spermatangial strips. Robe, S. Aust., in bay

(Womersley, 8.x.1972; ADU, A42764).

PORPHYRA AND PORPHYROPSIS IN SOUTHERN AUSTRALIA 67

20%

KY y %5°> ~

<0 <8 8 5 2,2 = *

Basse tseus nes eae

MNT ESE oT

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449 § Dat Ges on Oa here et

et ate tn GO as Tete

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sre

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8 tas ge 03 5S ER Page 08a tet et o%

SESS ING Ara ces baateras abenaee atone

e +

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oot oy

YY)

Fig. 6. Porphyra lucasii. A. Vegetative cells (ADU, A42764). B. 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 Ht, B.S. WOMERSLEY & ELSIE CONWAY

Distribution; From Cottesloe, W. Aust. to

Western Port Bay, Vic. and the north and east

coasts of Tasmania,

P. Ineasii is the common Porphyra of calm

to moderately rough waters, being replaced by

P. colymbina where surf action is strong, It

is essentially a winter plant of the mid to upper

culittoral, with old plants being found in Octo-

ber, P. lveasii is found within calmer bays

such as Port Phillip Bay and Western Port.

Vic,, whereas P. columbina occurs on rough-

water coasts.

Genus PORPHYROPSIS Rosenyinge

Porphyropsis minuta sp. nov.

Thallus epiphytic on certain Rhodophyta

with a firm surface, developing from a sub-

parenchymatous holdfast to form a hellow,

sub-spherieal to ovoid bladder (Fig. 74), later

opening above to form irregular, monostro-

matic membranes often with conyolute mar-

gins, Color greensh-brown to purplish, not

Jose-red.

Bladders up to 1 (-2) mm across, tarn

membranes to 5 (-8) mm high and across.

Cell divisions intercalary, Holdfast 50-100

(-200) ym across, formed of irregularly

arranged cells without rhizoidal extensions,

Cells of bladder (Piz. 7B, C) often paired or

in fours following division, and lying in rows

more or less at right angles, sometitnes becom-

ing irregularly arranged; membrane about LO

pm thick, cells 3-5 wm across and rounded to

somewhat elongate in surface view, slightly

ovate io sectional view. Rhodoplast apparently

filling the cell, without a pyrencid.

Monosporangia (Fig. 78) formed from the

whole cantents of cells near the margin of the

membranes, subspherical to ovoid, 5-7 ym

across,

Thallus in Rhodophytis epiphyticus, solido

superficie, ex subparenchymato base ortus,

primum vesicam cavam, subglobosam vel

ovoideam formans deinde superne membranas

ittegulares monostramaticasque saepe rnargine

conyoluto producens. Color brunneo-viridis

vel purpureus, sed nunquam carneus.

Vesicae ad 1 (—-2) mm lutae, membranae

laceratae ad 5 (-8) mm altae et latae. Diyi-

stra cellularum intercalaris, Basis ad 50-100

(—200) jm lata, ex cellulis sine projecturis

rhizoideorum irregulariter compoasitis formata

est. Cellulae vesicae saepe binae vel quaternae

post divisionem, seriatim plos muinusye rec-

langulatae, interdum irregulariter compositae:

membrana circa 10 ym crassa, cellulis ad 3-5

pm Jatis et aspectu frontali globosis vel

aliquantum elongatis, in sectione transversall

parum ovoideis. Rhodoplastus ut videtur, cellu-

lam complet, pyrenoide. absenti.

Monosporangia subglobosa vel ovoidea, 5-7

pm lata, in cellulis prope marginem mem-

branarum formata.

FIG, 7

Type locality: Pearson I, S. Aust. on Plero-

eladia capillacea, upper sub-littoral (Speeln,

17.11.1960).

Type: ADU, A24525,

Distribution; From Garden J., W. Aust.

around southern Australia to Bateman’s Bay,

N.S.W., epiphytic on Prerocladia cupillacea in

Upper sublittoral (and pools) on rough-water

coasts, and occasionally on Plocaynium atigus-

tani, P. mertensii and Delisea pulchra in simi-

lar habitats.

P. minuta agrees well with P. coccinea in

the boldfast, form and development of the

bladder, and in reproducing apparently only

by monosporangia. The cells of P. mina are

similar in site to those in P, coccinea but are

arranged in disinct roWs more or less at right

angles, in contrast to both P, ceccinea from

Europe |Rosenvinge 1909, p. 69, fig. 9) and

P. coccinea var, dawsonli Hollenberg &

Abbott (1968, p. 1239, fig. Sa-c) from Cali-

fornia, where the cells are more irregularly

arranged but are grouped into elongate, same-

what lenticular patches. The life-history and

relationships of the latter taxon have been

discussed by Murray, Dixon & Scott (1972),

and it is desirable that the Australian plant

should be studied in culture,

A further difference is that in P. minuta the

whole contents of cells mear the margins are

liberated as monospores, Whereas in P-

coccinea the monospores are cut off from the

parent cell by a curvéd wall, a residual cell

remaining when the monospore is liberated,

Also, in P. coccinea some holdfast cells form

rhizoidal extensions whereas this has not. beets

observed in P. mitfauta, The colour of P. minuta

is always @ greenish-brown-purple, never rose-

red as in P. coccinea.

The orly other southern hemisphere record

of Porphyropsis is by Adams (1972, p. 63)

who reported P. caccinea var. dawsonit from

New Zealand. This taxon (e.g. CHR 248053

from Kaikoura, N.Z.; Parsons, 13.x1.1973) has

numerous ligulate fronds from a clumped base,

each with descending rhizoids, It is not a Por-

pityropsis, but more closely related ta Porphyre

woolhoausiee’.

PORPHYRA AND PORPHYROPSIS IN SOUTHERN AUSTRALIA

7 ]&

on +*,.

der membrane, with some cells re-

phyropsis minuta. A. Small plants, on Pterocladia. B. Ol

leasing monosporangia. C. Cell arrangement. (All from the type, ADU, A42525.)

Fig. 7. Por,

70 H. B. 5. WOMERSLEY & ELSIE CONWAY

Acknowledgments

We are grateful to Mrs S. C. Ducker (Uni-

versity of Melbourne) who made available

collections for sttidy, and to Dr P. Tyler (Uni-

versity of Tasmania) who recollected P. waol-

hausiae. The first author acknowledges pro-

vision of technical assistance by the Australian

Research Grants Committee,

References

Abams, Nancy M. (1972).—The marine algae of

the Wellington area. A list of species. Rec.

Dom. Mas, 8(5), 43-98,

Acarpr, J. G. (1883).—Till algernes systematik,

Nya bidrag. Vi Ulvaceae. Acta Univ. lund

19(2), 1-182, Plates [-4.

CHAPMAN, D. J. (1962).—A check fist and key

to (he Rhodophyceae of New Zealand, Sec-

tion A: Bangioideae, Trans, R. Soe. NZ,

Bor. 1. 127-137. d

Culapman, V. J. (1969).—The marine algae of

New Zealand. Part LIT: Rhodophyceae. 1.

Bangiophycidac und Florideophycidae (Nema-

lionales, Bonnemuisoniales, Gelidiales).

(Cramer: Germany.)

Conway, Elsie (1964).—Autecological studies of

the genus Porphyra: 1. The species in Britain.

Brit. Phycal. Bull. 2(5), 342-348.

Conway, Elsie, & Wyte, Ann P. (1972).—Spore

organisation and reproductive modes in two

species of Purplyra from New Zealand. Prov.

FU Int. Seaweed Symp., pp. 105-107. (Univ.

Tokyo Press: Tokyo.)

Curse, A. B. (1954).—The algal vegetation of

Port Arthur, Tasmania. Pup. Proc. R. Soc.

Tasm. 88, 1-44, Plates 1-10,

Dawson, RB. Y., Acieto, C., & Foipvie, N.

(1964).—The Seaweeds of Peru. Nova Hed-

wigia 13, 1-111, Plates 1-81.

De Toss, G, B. (1897).—Sylloge Algarum

omnium hacusque Cognitarium”. Vol. 4.

Florideae: Sect. 1, pp. 1-388, (Puduu,)

Re Tow, G. B. (1924)—"“Sylloge Algarum

omnium hucusque Cognitarium”. Vol 6.

Florideae, (Padua.)

Ginter, E. R. (1952).—The marine algae of

Tasmania. Cheek fist with localities, Pap,

Proce. R. Sac. Tasm. 86, 71-106.

Gumer, FE, R. (1954).—The recolonization of

rock surfaces and the problem of succession.

Pap, Proc. R. Soc, Tasm, 88, 49-66, Plates

1-6.

Hamev. G. (1928)—Sur quelques Porphyra des

mers australes. dan. Crypt. Exor. 1(1), 51-

57.

Harvey, W. H. (1863) —"Phycologia Australica™.

Vol. 5, Plates 241-300, synop, pp. 1-73.

Ho.Lenpera, G. J., & Apporr, 1. A. (1968).—

New species of marine algac from Califor-

nia. Cun. J. Bot, 46, 1235-1251.

Kuerzing, F, T, (1849)—"Species Algarum”.

(Leip2ig..)

Kuerzinc, F. TT. (1869).—'Tabulse

Phyco-

logicae’. Vol, 19, (Nordhausen. }

Lame, R. M. (1928)—New Zealand Bangiales

(Bangia, Porphyra, Erythratrichia and (?)

Erythractadia). Trans, NZ. Inst, 59, 33-59,

Plates 1-15.

Levainc, T, (1953).—The marine ulgae of Aus-

tralia. I. Rhodophyta: Goniotrichales, Ban-

giules and Nemalionwes. Arkiv. for Botanik,

Ser, 2, 2(6), 457-530.

Levring, T. (1955).—Contributions to the marime

algae of Wew Zealand. J. Rhodophyta:

Goniotrichales, Bangiales, Nemalionales and

Bonnemaisoniales. Arkiv fdr Bot. Ser. 2.

3(11), 407-432,

Levrinc, T. (1960).—Contributions to the murine

algal flora of Chile. Lunds Univ, Arssk,

N.F. Avd. 2, 56(10), 1-85.

Lucas, A. H. S. (1909).—Revised list of the

Fucoideae and Florideae of Australia. Proc,

Linn. Soc, N.S,W, 34, 9-60.

Lucas. A. H. 3, (1929).—The marine algac of

Tasmania. Pap. Proc, R. Soc. Tasnt. 1928,

6-27.

Montaane, C. (1842).—"Prodromus Generum

Specierumque Phyceurum Noyarum In

Itinere and Polum Antarcticum.” (Paris. )

MontaGne, C. (1845)—“Voyage au Péle Sud

et dans /Océanie sur les Corvettes ]'Astrolahe

et la Zélée.” 1, Botanique. ( Paris.)

Murray, S&S. N., Dixon, BP. S&S, & Socom, J. L.

1972).—The life history of Porphyropsis

cuecinea var. dawsonii in culture. Ar. phyceal.

J. 7, 323-333.

Puyats, Carmen (1963),—Catulogo de Rhodo-

phyta citadas para la Argentina. Revia Mus.

argent. Cienc. nat. Bernardina Rivadayia, Bot

3(1), 1-139.

Rosenvincre, L. K. (1909),—The marine algac

of Denmark. Pari 1. Introduction, Rhode-

phyceae. 1. (Bangiales and Nemalionales),

Kel. Danske Widenskab. Selskab. Biol. Skr.,

7 Raekke, Afd. 7, 1-151, Plates 1, 2, 2 maps.

Sxorrsperco. C. (1923) —Botanische Ergebnisse

der schwedischen Expedition nach Patagonien

und dem Feuerlande, 1907-1909. [X. Marine

algue. 2. Rhodophycene. K. svenska Vetensk-

Akad. Harndl. @3(8), 1-70,

SmirH, G, M., & HoLLENeerc. G. J. (1943),—

On sume Rhodophycese from the Monterey

Peninsula, California, Amer. J. Bot. W(3).

211-222.

Womersiey, H. B.S, (1950)—The marine algae

of Kangaroo Island. UWI. List of Species 1.

Trans. R. Soe. 8. Aust. 7312), 137-197.

Womersiry, H. B. S, & Epmonops, 8. J. (1958).

—A general account of the intertidal ecology

of South Australian coasts. Aust, J. mar.

Freshy. Res, 9(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. S. Aust. 99(2), 71-84, 31 May, 1975.

In vertebral morphology, the elapid species Pseudechis porphyriacus, Austrelaps superba, Notechis

scutatus and Pseudonaja nuchalis 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 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 VERTEBRAF. OF FOUR AUSTRALIAN ELAPID SNAKES

{SQUAMATA;: ELAPIDAE)

by MEREDITH J. SMITH*

Summary

SmMiTH, Merrepity J. {1975).—The vertebrae of four Australian elapid snakes (Squamata:

Elapidae). Traits, R, Sec, §, Aust, 99(2), 71-84, 31 May, 1975.

In vertebral morphology, the elapid species Psewdechis porphyriacus, Austrelaps superba,

Norechis seytatus and Psendonaja nucheliy conform with general descripuions and closely

resemble each other. Features not previously noted are that epizygapophysial spines appear

an the first 8 to 10 vertebrae, and that bilaterally on every precloucal vertebra a foramen

opens through the accessory process near the anterolateral edge of ihe prezygapophysial facet.

No morphometric ferture was found to completely separate any two yenera, ie. there was

some overlap in the values of all the characters (ratios) studied, However, in general P. por-

piiyriacus vertebrae are distinguished by their relatively long accessory processes, 4- superba

by their short accessory processes and lesser width across postzygapophyses, and N, scutatus

by their greater width across postzyeapophyses and shorter neural spines. P, nuchalis vertebrae

have strong subcentra) ridges.

Fossil vertebrae from a Pleistocene deposit differ from P. parphyriacus, A. superba and

N. seutatus but resemble Pserdonaja in most features, The fossils differ from modern P.

nuchalis chiefly in having a thicker zygosphehe and relatively wider postzygapophyses. As these

fire features which develop with increasing size of vertebrae, the fossil vertebrae are assigned

to the venus Preudenaja.

Intraduction

Although snake vertebrae haye been

recorded fram Australian Pleistocene ‘deposits

(Lydekker 1888, Merrilees 1968), the only

species that has been identified is the Carpet

Snake, Morelia spilotes yvariegata Gray (=

Pyrhen variegatus) from Marmor Quarry,

Queensland (Longman 1925). No detailed

studies have been published of Australian

snuke vertebrae and the diagnostic characters

have not been established. The need to deter-

mine the reptile fauna in a Pleistocene cave

ueposit stimulated the present study.

The sixty-odd Australian species of clapids

have been arranged in 29 genera (Worrell

1963) or fewer (McDowell 1967, 19703, but

only seven geners contuin species with a

recorded maximum Jengih of over 0.9 m. As

centrum lengths of some of the Pleistocene

vertebrae suggested tbat the specimens from

which they were derived must have exceeded

1.5 m, the 22 genera of smaller species (less

than 0.9 m maximum length) have not been

considered. Nor have Demansia [restricted io

the whip snakes by Worrell (1963)|. Hopic-

cephalus, and Oxyuranus been examined as

their yange jis. northern and casiern Australia

(Worrell 1963). Specimens have been studied

from the remaining four genera, Austrelaps,

Notechis, Pseudechis and Pseudenaja, which

are cach represented in southern Australia by

one Of morte common species.

The exact column position of an isolated

vertebra is impossible to determine (Johnson

1955} and in elapids, with well-developed

hypupophyses on all precloacal vertebrae,

division of the pre-cloacal column inta regions

is virtually impossible. Nevertheless Vertebral

shape changes along the column, and to iden-

lify single vertebrae it is essential to know the

tange of variation within individuals and

species.. Auffenberg (1963) based his descrip-

tions and diagnoses on middle pre-caudal verte-

brae (determined as such by the relative size

of the neural canal} and avoided considering

imtracolumnar variation. Although Johnson

(1955) measured 10 precloacal vertebrae at

Tegular intervals along the column, he assumed

* Department of Zoology, University of Adelaide, Adelaide, S$. Aust, SODG.

72 MEREDITH J. SMITH

equi variynées (between specimens) of ratios

of these nleasurements and in his comparisons

he usted enly the mean for each specimen,

Here. entire vertebral columns have been

examined in an plempt to assess the morpho-

logieu)! and) morphometric variation which

occurs within individuals and species, and to

find unique specific chatacters. As most elapils

have over 200 veri¢brae, the available samples

are large and the time tequyred for detailed

examination of one individual snake precludes

sampling many individuals,

Because taxonomy of species of Preurdonayce

and Austrelaps is confused (Rawlinson 1969,

Storr 1964) comparisons are made at the

gener tather than species level,

Materials and Methods

A total of 2,123 vertebrae from nine Recent

specimens (Table 1) and 556 Pleistocene ver-

lehrie were examined. Of the nine modern

specimens. four specimens of Pyreuedectiy

porphyeiacys, one Pseudonala — nuchalis

(R14064) and one Norechix — scutatus

(R14059) were collected near Armidale. New

South Wales: the one Aastrelaps superba was

collected at Uruidla. South Australia, The

localities of one P, nuchalis (RV4065) and

one N, seuratuy (R14058) are unknown.

The common hrown snake, Poendunafa tex-

filiy (Duméril & Bibron}, has been described

as # separate species Jrom the western brawn

snake. Psendenaja nuchalis Gunther, but it is

not yet clear (Storr 1964) whether they are dis-

tibet or are merely races of a single species. P.

featilis, One clearcut diagnostic feature is the

shape of the nasal bones. which are anteriorly

eoncuve on the lateral margins in P. nauelialis

but anteriorly convex in P. /extilis (Worrell

1963), In RI4064 and R14065 the nasals are

anteriorly coneave and hence are attributable

to FL onrichatis,

The cleaned skeletons were dried and com-

pletely disarticulated for study. The specimens

ure now lodged in the South Australian

Museum, with register nambers as above,

The Pleistocene vertebrae were excavated

from an extensive hone deposit in Victoria

Cave, Naracoorte. South Australia. The ave

of this deposit is unknown, but the jihundance

of extinct marsuptals and the absence of

remains of aboriginal man suggest thal ihe

deposit uccumuluted during the Pleistocene but

was sealed during Recent time. As the verte-

lirae were collecied singly im many locations

in the bone depasit, they probably represent

al feast several individuals. Visual eximinution

of the fossils revealed that 454 of them were

similar to each other if shape, and distinctly

different from the resi. which were of several

kinds, The latter. hererogencoux group will be

discussed in a later paper. OF the larger group

of 454. the S0 most complete were examined

in detail and measured in the same way as the

modern vertebrae

Because reptiles grow throughout their life.

absolute dimensions of the vertebrae are al

little use im comparing individuals or species.

Ratios berween dimensions have been calcu-

lated, the denominator being vertebra length

im most comparisons, Mean values of ratios

have been prepared independently for cach

individual snake, and are given 4s mean, T,

+ standard error, followed in brackets by the

number of vertebrae meusured, The non-inde-

pendence of measurements of different verte-

hrae of the one individual precludes statistical

comparison of these samples (Siegel 1956) _

From 4 preliminary study of numerous

characters of snake vertebrae, characters were

selected that vary between species within the

tamily Elapidat, exhibit low intracolummar

variation and are well preserved in [fossils To

eslablish vanation throughout the column,

every fifth vertebra was measured in. two speci-

mens of each of P. porphyriacns, Po nuchaliy

and N. scutetus, and one A. superba,

Descriptive techniques and terminology

fallow Auflenberg (1963), Measurements were

made to 0.1 tam with dil calipers. as in Fig,

pr-po Length between zygapophyses—distanoe

between most amecior point of prezy-

gapohysis lo most posterior point of

postzygapophysis.

ap-dp Width across accessory procusses—dis-

tance between outermost tips of acecs-

Ory Processes,

po-po Width across postzygapophyses—cis-

lance between outermost points of the

postzygapophysial facets,

pri Length of prezygapophysis—the longest

diameter of the prezygapophysial facet

feven though this was almost perpen-

dicular to the long axis of the cen-

trum).

pr Width of prezygapophysis-the maai-

mum diameter at tight angles co the

length of the prezygapophysis {the

prevygapophysial width being almost

parallel tu the toi axis of the ¢en-

trum),

Vig. |.

VERTEBRAE OF FOUR BLAPID SNAKES 7

Dorsal (D), ventral (V) and Jateral (L)

views of an elapid vertebra showing where

measurements were taken.

Length of postzygzapophysis—the

longest diameter of the postzygapo-

physial facet (éven though this was

almost perpendicular to the long axis

of the centrum).

Width of postzygapophysis-the maxi-

mum diameter at right angles to the

length.

Minimum Jength of neural spine-this

usually occurred about halfway up the

spine, as the dorsal edge overhangs

posteriorly and/or anteriorly.

Width of zygosphenc-the maximum

width of the tenon.

Width of condyle-the maximum dia-

meter in the transverse plane,

Length of hypapophysis-the vertical

distance from the lower edge of the

condyle ta the tip of the hypapophysis.

Results

The vertebrae conform with the general des-

criplions of Auffenberg (1963), Hoffstetter &

Gase (1969) and Johnson (1956),

Pseudechis porphyriacus. (Shaw)

The number of precloacal vertebrae (Table

1) is consistent with the number af ventral

scales (184, according to Worrell 1963).

In every precloacal vertebra, the width

across postzygupophyses exceeds. the length

between zygapophyses (Fig, 2). The hypapo-

physis arises near the lip of the cotyle, extends

as a Jow 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 hypapophyses of the anterior vertebrae

are very Jong; 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 anteriarly. 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 minitnum length of the neural spine

is about half the length between the zygano-

physes [¥ 0.53 = .0056 (35); ¥ 0.55 = .0049

(36)] (Fig. 3). The neural atch is slightly

wider than high, The neural arches of most of

the vertebrae do not extend backwards to form

epizygapophysial spines, but such spines are

well developed on the first five vertehrae and

aré distinguishable on the sixth to tenth. The

cotvles and condyles are nearly round, but are

slightly flattened dorsoventrally. The wadth of

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

Oo, ee ha eee

SAM Total Snout-vent Number of vertebrae

Apecies number Sex (mm) (mm) cloacal* Cloacal cloacal Total®

Pseudechis R14060 no data 1220 no data 181 8 >42 >231

porphyriacus R14061 no data 1375 no data 182 6 §2 240

R14062 no data 1120 no data 182 5 >30 217

R14063 no data 1070 890 177 $ 48 231

Austrelaps R14066 ge $15 433 143 4 42 184

superba

Notechis R14058 g 930 775 174 5 58 37

Scutatus R14059 no data no data 800 178 5 52 235

Pseudonaja R14064 é 1555 1300 204 6 60 270

nuchalis R14065 no data 1100 510 206 5 62 273

* Atlas-axis complex not included in this count.

50 100 150 200 250

Vertebra number

Fig. 2. Variation throughout the vertebral column in (a) length between zygapophyses

(A 4), (b) width across postzygapophyses (QO @ ) and (c) length of hypapophysis

(C1m) of two specimens of Pseudechis porphyriacus, Hollow symbols R14060, solid

symbols R14061.

VERTEBRAE OF FOUR ELAPID SNAKES

+ Ws N R14061

: X\ we a R14063

‘TCH al Fi ac

“ i) yp i

al Dy. DL Lo pes

730

320

Lie

10 i R14065

; a _ ries

20 Fassil

bitiririts LiLLitrisiirs Litiviry

10% 2 B 10 12 13 1% 15 03 04 OS O6

Rel. width Ant. width: Post.width Rel. length spine

Fig. 3, Distributions of values for the ratios (a) width across posizygapophyses to Jength

between zygapophyses, (b) width across accessory processes to 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 scitatus; solid columns,

Pseudonaja nuchalis; fine stipple, Victoria Cave, type A; coarse stipple, Victoria Cave,

type B.

MEREDITH J. SMITH

10 R14061

10 R14060

10 R14066

10 R14059

10 R14058

>30

1=]

$20

iC 10 R1G065

10 R14064

20 Fossil

Loot) iy bout tty Liga s

0.4 05 0.6 07 02 03 O4 OL O2 03 :

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 ELAPIN SNAKES 7

TABLE 2?

See charactorstics of Mie eloucal region of four

specunens of Poeudechis porphyriacus

Nuober Nuiviber

Total with Number with

mumber of articu- with ribs

cloxcal lated sifigle deeply

Specimen verielsrme riks hypypuphysis Forked

Hidn6u 5 1 2 8

RAANAL = 1 3 $

RLAg62 5 4 4 ar

KA4N63 6 2 3 sv

Seen

* The anreriarsmost pair of rihs Rave shallow farks.

between zyeapophyses [x 11.33 = 0047 (36);

¥ 0.47 + .0067 (36)] (Fig. 4). The zvgo-

sphene, viewed Irom the front. is thin und

straight or slightly convex: from abuve it is

nearly straight or slightly concave with a faint

meiian notch on some vertebrae. The width

of the zygosphene is a little over half the

length belween the zvgapophyses [x 0.53 =

.0063 (36); 0.57 + ,OU38 (35)]. The prezy-

gapophysial facets are oblong [length/ width

1181.58, F140 = O19 (36): 101-173, %

1.49 + 025 (36)|. The acute accessory pro.

cesses extend laterally perpendicular tu ‘both

vertical and horizontal Axes of the vertebra,

and project well beyond the articular facets.

Hence the ratio of width across accessory pro-

cesses tu Widih across postzygapophyses is high

{r 1.399 = Ol4 (35); F 1.38 + O10 (35)]

(Fig, 3), The postzygapophyses are Jarge

[widch postzygapophysis/length between

zygapophyses: * 0.20 + .003 (36); * 0.20 =

U2 (36)| and have an amterior notch that

gives their otherwise ovate shape a kidney oul-

Ime. ‘fhe maximum diameter of the post-

zyzapophysis is almost perpendicular to the

lung axis of the vertebra and exceeds the width

[length/ width |.03-1,51, ¥ 1.27 + .020 (36);

1.02-1.46, ¥ 1.22 = O17 (36}].

The typical four pairs of foramina are

present on all precloneal vertebrae. and in

many vertebrae they are unilyterully or

bilaterally doubled, Mor exumple, in the 183

preclaacal vertebrae of RI4061 the lateral

foramina are bilaterally double (8 vertebrae),

double on the right only (11) of left only

(13); the paracotylar foramina are bilaterally

double (1). double on the right only (5) or

left only (8); the subcentral foramen 1s: double

on the aght (2) or left (3) and in each of

three vertebrae three pairs of subcentral fora-

wuts appear. A fifth pau of forming 1

present in the precloacul vertebrae of all speci

‘

mens, each foramen of this pair opening

through the accessory process near the: antero-

laterul margin of the prezygapophysial facet.

Vertebral structure in the cloacal region

varies widely between specimens, and the nurm-

her of veriebrae with urticulated torked ribs

may be equal to. or less than the number with

a single hypapophysis (Table 2). The fork is

deep on most forked ribs but on the most

anterior cloacal veriebra. the notch may be

midway along the rib, The lymphapophyses

project luterally and slightly ventrally, as do the

pleurapophyses of the post-cloacal vertebrae

The thaemapophyses of cach post-cloacal

vertebra arise separately from the ventral sur-

face of the centrum, and rémain completely

separate, although on the anterior pust-cloacal

vertebrae the tips of the haemapophyses con-

verge slightly, In lateral view the shape of the

haemapophysis is similar to that of the hypapo-

physes of the precloacal vertebrae. The pleura-

pophyses are directed anterolaterally {viewed

from above or below) and extend anterior to

the cotyle.

Some irregularities occur in the skeletuns.

In R1-4060 the right prezygapophysial facets of

the 130th 10 134th vertebrae ure enlarged to

almost double the size of the Icft-side facet,

and the right postzygapophyses of these verte-

brae are slightly enlurged. A similar, bul even

greuler, enlargement of the night prezygapu-

physis occurs in two vertebrae of RI4062,

where an outgrowth of spangy bone has com-

pletely Eused the prezygupophysis to the pre-

ceeding postzygapophysis. wand the sccessory

process is reduced. These abnormalitics prob-

ably resulted from injury, but R14063 shows

slight cangenital abnormahties, firstly on verte-

brae 91, where a small, but distinct epi4ygapo-

physial spine appears on the left side only, and

secondly in four post-cloacal vertebrae. Near

the beginning of the posicloscal serics, the

centra of Wo canstcutive vertebrae are com-

pletely fused, On the left. the outline of the

postering edge of the neural arch of the first

vertebra can be seen, but on the right the

suture line is only faintly distinguishable The

pleurapophyses of both vertcbrac lie close

together, apparently fused to the centrum of

the first vertebra. Two puirs of haemapophyses

anise separately on the ventral surface uf the

combined centrum. Later in the post-cloacal

series two vertebrae ore more fully fused and

share a commen neural spine. Two pairs of

haemapophyses are fused ul their hase near the

condyle of the second vertebra, Ali apparently

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MEREDITH J. SMITH

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150

Vertebra number

200 250

Fig. 5. Variation throughout the yertebral column in (a) length between zygapophyses

(A 4), (b) width across zygapophyses (O@).

Hollow symbols,

(Cle) in two. species,

Notechis scutatus (R14058).

similar abnormality has been observed in dor-

‘sal vertebrae of colubrid snakes (King 1959),

Austrelaps superba (Giinther)

The low number of precloacal vertebrae in

this specimen (Table 1) is confirmed. as typical

of the species by the ventral scale number (151

according to Worrell 1963). The width across

the postzygupophyes: is less than the length

between zygapophyses in the first ten and the

last 35 precloacals (Fig. 3).

The hypapophysis (Fig. 5) is similar in form

to that of P. porphyriqeus and does not extend

further posteriorly than the posterior surface

of the condyle.

The subcentral ridges are low and rounded,

as in P. porphyriacus, but the interzygapo-

physial ridges of A. superba are stronger. Small

epizygapophysial spines occur on the first six

vertebrae, The condyle is smaller, relative to

the length between zygapophyses, than in P.

porphyriacus (Fig, 4). The zygosphene is thin

and slightly convex from the front, convex

and (c) Jength of hypapophysis

Austrelaps superba (R14066), solid symbols,

from aboye, The ratio of zygosphene width to

length between zygapophyses (Fig. 4) has a

mean of 0.50 + .0038 (27). The prezygapo-

physial faccts are oblong, the postzygapo-

physial facets obovate. The acute accessory

processes are relatively shorter than in P, por-

phyriacus in most vertebrae (Fig. 3),

Of the four cloacal vertebrae, one has

articulated forked ribs and three have a

hypapophysis. The haemapophyses of the post-

cloacal vertebrae are long anteroposteriorly,

about half the length of the centrum, and are

completely double.

Notechis scutatus (Peters)

Tn all precloacal vertebrae, the width across

the postzygapophyses exceeds the length

between zypapophyses (Fig. 5). Each hypapo-

physis is extremely compressed laterally to a

thin lamella that terminates posteriorly in a

sharp point not extending posterior to the

posterior surface of the condyle in any but the

first 15 vertebrae, where the hypapophysis is

VERTEBRAE OF FOUR ELAPID SNAKES 79

Fig. 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. *

sb MEREDITH J. SMITH

very lone (Pig. SJ. The sulycentral ridges ure

low. and rounded: the interzygapophysial ridges

are weuk hut distinct. The neural spine is

higher than in P. porphvriacus (Fig. G) and its

herizental dorsal edge overhangs both

anteriorty aml posteriorly. The mininium

lengih of the neural spine is generally relatively

shorter than in the specimens of FP. porphy-

vineus [ratio of minimum length of neural

spine to length between zygapophyses: ¥ 0.48

+ .0064 (35): ¥ 0.48 + 049 (361) (Fig. 3).

Epizyzapophysial spines are well developed on

the first four vertebrae. faintly clistinguishable

on the filth to eighth. but absent from all

others.

The round condyle is ser on a short but

uistine, neck (Fig. 6). ‘The thin zyzosphene is

straight or slightly convex from the front,

straight or stightly concave fram above, The

prezygapophysial facets arc oblong flength/

width 1-00-1.67, © 147 % .027 (35); Loo-

1.58, ¥ 1.35 4 022 (35)], The accessory pro-

cesses are obtuse and short, so that the ratio

of width across accessory processes 10 width

ucross posteygapophyses is generally less than

in P. porphyriacus (Fig. 3). The postzyyapo-

physes ure obovate [length/ width 1.10-1.58, ¥

1.35 + 071 135); 1.00-1.47, ¥ 1.28 + 022

(35)), and their width is about one-fifth of

the length between the zyzapophyses (Fig, 4),

Prezygapophysial foramina are constantly

present,

In both specimens, all five cloacal vertebrae

have fused torked mbs. One has three, the

other two cloacal vertebrae with a single

hy papophysis.

The haemapophyses are large. paddle-shaped

and completely paired, The pleurapophyses

extend ventrally more than laterally. The

vygapophysial facets extend anteriorly and

posteriorly, rather than laterally as in the pre-

cloacal vertebrae, and acute necessary pro-

cesses are distinct on all the postcloacal verte-

brae.

Unilateral or bilateral doubling of the para-

cotylar and/or subcentral foramina occurs in

a very few prechoacal vertebrae (no more than

four in cither specimen).

Pseudonaja nuchalis Giinther

Pseudonaja nuchalis has more vertebrae

than any of the other species studied here

(Table 1), and maximum length and wielth

OCCUE i sequentially more posterior vertebrae

than in the other species (Fig. 7), All pre-

cloacal vertebrae wie wade thin Jone The

hypapophysis is well developed on ull pre-

eloucal verlehcie, Thickenings on the rim of

the cotvle on either side of the ventral midline

ive rise to jy low ridge that narrows sharply

In juin the laterally-compressed hypapophysis

The hypupophysis deepens from abour the

middie of the vertebra and lerminates in a

rounded point that extends posteriorly ta the

condyle, Subcentral ridges are strony and inter-

aygapophysial ridges are distinerz. The para-

diapophyses are well developed, with a pro-

ftruding, dorsoventrally-clongated upper facet

und saddjeshaped jower facet, The parapo-

Physial processes appear in ventral view us a

flut surface neatly as broud as long; the ventral

projection of these processes is medial to the

lateral border of the catyle. The neural spine

is low and long (Tigs 3 and 6}; its dorsal edge

overhangs slightly at the front. markedly

behind, Epizvgapophysial spines are distinet on

the first five vertebrae, faintly visible on the

sixth to eighth and absent from all others. The

coryles und condyles are almost round: in

R14665 some cotyles are depressed slightly,

but in RId064 some are slightly compressed

laterally. The condyles are relatively much

Farver in the smaller specimen than in the

larger one (Fig. 4). The zygosphene is convex

from the frent. concave from above; it is thin

in vertebrae of the smaller specimen but is

thickened in those of the larger. The pre-

zygapophysial facets are oblong [Jength/ width

1,09-1.72, ¥ 1.39 = 022 (41); L_L8-1.78, %

1.35 + 018 (41)], and the acute accessory

processes extend laterally well beyond the

articular facets (Fig. 3). Except fur a slight

natch posteriorly. the outline of the post-

zygapophysial facet is almost round [length/

width LOG-133 F007 + 011 (41): 10g

1.59, ¥ 1.30 + 19 (4151.

In the cloacal regian the ribs are all fused

except for the anterior vertebra uf R1d064,

und a hypapuphysis occurs on one (R14063)

or three (RL4064) cloacal vertebrue-

‘The huemapophyses are completely paired

and extend anteriorly as two separate rmuges

to the sim Of the cotyle, The plewrapophyses

ire broad and flat and not pointed. In antenor

view they project ventroluterally but in ven-

tral view mainly laterally and only slightly

anteriorly,

In R14065, more than the one pair of pura-

colvlyr foramina appear on many verlebrac—

15 prectoacal vertebrue have an extra puru-

coWlar foramen on the right side, 18 wn the

left and 34 on both sides. RIGN64 has only

VERTEBRAE OF FOUR ELAPID SNAKES

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50 100

200

150 250

Vertebra number

Fig. 7. Variation throughout the vertebral column in (a) length between zygapophyses (A 4 ),

(b) width across zygapophyses (C@), and (c) length of hypapophysis (jm) in

Pseudonaja nuchalis. Solid symbols, R14064, hollow symbols, R14065,

8 precloacal vertebrae with additional para-

cotylar foramina. The left postzygapophysis of

the 114th vertebra is fused by an outgrowth of

spongy bone to the prezygapophysis of the

succeeding vertebra, which lacks an accessory

process on the left side. The left rib of vertebra

114 shows a healed fracture near the articula-

tion with the vertebra.

Pleistocene fossil vertebrae from Victoria Cave

The precloacal vertebrae found in the Vic-

toria Cave deposit vary in length from about

2 mm to a maximum length between zygapo-

physes of 11.1 mm. In their general conforma-

tion they closely resemble those of Pseudonaja.

However within the sample of vertebrae from

Victoria Cave, two types can be distinguished

on the characteristics of the zygosphene: Type

A: slightly convex when viewed from above,

convex in anterior view and slightly thickened;

Type B: almost straight, with a median notch,

when viewed from above, almost straight in

anterior view and extremely thickened. This

thickening is consistent with the robust

appearance of the vertebrae. The subcentral

ridges are particularly strong (Fig. 8).

As well as the thickening of the zygosphene,

its width relative to the length between zyga-

pophyses differs significantly between the

types A and B [A, ¥ 0,52 = ,0048 (40); B. ¥

0.54 = .0057 (40); .002 < P < .0]]. The dis-

tributions of the values for this ratio overlap

widely not only between types A and B from

Victoria Cave but also among the specimens

studied (Fig. 4), The ratio of width across

postzygapophyses to length between zygapo-

physes tends to be greater in the fossil verte-

brae than in A. superba; the ratio of width

MEREDITH J. SMITH

: =

a ten

right,

dorsal (A, B), ventral (C, D) and '

Pseudonaja nuchalis; left, 80th precloacal of R14064

ave, Type B (P16126b)

vertebrae of

(E, F) views.

Precloacal

Victoria C

ig. 8.

VERTEXRAE OF POUR ELAPID SNAKES $3

¢ *

. a

“he te

E\s ee

— oo by -

£ a+

2 rd

: é

= gy

ed thee

x eat

‘

t 5 6 7? & & WN Te

Centrum length tmmt

Fig. 9 Douhble-logarithmic regression of width of

posizygapophysis on length between post-

zygopophyses in Pseudonaja nuchalis, 4,

R14054: +, R14064: ©, fossil, Type At

@. fossil, Type B.

across accessory processes to width across

postzygupophyses is less. than in most P. per-

phyriacus and the relative length of the neural

spine is generally greater than in N.. scutafus

(Fig. 3). In these ratios, the Victoria Cave

vertebrae closely resemble P. nuchalis. In the

relative Width of the condyle, the distribution

of ratios for Victoria Cave vertebrae resembles

the larger P. nuchaliy (R14064), though not

the smaller (R14065). Finally, in the relative

width of the postzygapophyses the Victoria

Cave vertebrac of Type A resemble the larger

but not the smaller P_ nuchalis, and the relative

width of the posteygapophysis is generally

greater in Type B fossils than in any of the

modern species studied (Fig. 4).

This subjective and objective analysis of the

fossil vertebrae indicates that they most closely

resemble Pseudonaja. The Type A vertebrae

can be referred with confidence to this genus.

Because the Type B vertebrae differ in the

thick zygosphene and the relatively large post-

zygapophyses, the possibility exists that they

are of a species different from Type A. How-

ever, these charucteristics are two which

develop with age (Auftenbetg 1963) and most

ef the Type B vertebrae are larger than the

Type A (Type A, length 6,0-8,1, ¥ 6,7; Type

B, 7-1-11.1, ¥ 8.9) und also generally are

larger than the vertebrac of the modern speci-

mens, When the correlalion between length

between zygapophyses and relative width of

postzygapophysis of Type HB vertebrae was

tested by the Kendall Rank Correlation Test

{Siegel 1956) the correlation was found to be

highly significant (" = 0.41, z = 3,765, P <

001), On a double-logarithmic plot of post-

zygapophysis width against length between

zygapophyses, the distribution of values for the

fossi] vertebrae (of both types, A and B) falls

hear the same straight line as those of R14064

and R14065 (Fig. 9). The similarities between

Victoria Cave preloacal vertebrae and those of

modern P. nuchalis so far outweigh the slight

differences: that recognition of the fossils as a

scparale species seems unnecessary at least

until the limits of variations of modern species

ure better known.

The post-cloacal vertebrac recovered from

Victoria Cave resemble those of P, avchelts

in the hiterally-directed pleurapophyses. In the

other species studied the pleurapophyses pre-

jech anterolalerally.

Discussion

{In all the vertebrae of the four species, the

hypapophysis, is well-developed (as in all

elapids) and hypapophysis Jength decreases

slowly from anterior to posterior along the

column. There is no suggestion of two distinct

regions as in Achrochordus javanicus where

the anterior tegion (to vertebra 96) has

hypapophyses long and fairly constant in

léngth and the posterior region has hypapo-

physes short and of constant length (Hoff-

stetter & Gayrard 1964).

At family level, the presence of vertcbral

foramina may have diagnostic value [c.g. the

constant absence of lateral foramina in Achro-

chordidae (Hoffstetter & Gayrard 1964)], but

the variability in the number of foramina at

each position (i.e, lateral, subceritral, etc.) in

the one snake indicates the need for caution

in the use of foramina jn taxonomy,

Although middle precaudal vertebrae may

be the most tonstamt in their structure (within

species) and hence best for identification

&4 MEREDITH J. SMITH

(Auffenberg 1963). the difficulty of assigning

an isolated elapid vertebra to a particular

region of the column preciudes confidence in

selecting middle precaudals from a sample of

fossils, Because of the consistent variations

along the column, together with some irregular

variation (e.g. doubling of foramina), to iden-

tify isolated vertebrae it is necessary to con-

sider not just the middle precaudal vertebrae

of reference specimens, nor the mean of some

value (even if it be given with standard error)

but the range through which a given character

varics. Also because of variations along the

column, no unique specific character was

found, and so it is necessary to consider several

characters in the identification of fossils. The

fossil genera may be readily identificd, but

further studies of congeneric species are

needed to determine whether specific identifi-

cation is possible.

Acknowledgments

Mr R, Shine, Mr G. Whilten and Dr R. T.

Welly kindly donated the modern snakes. Dr

Wells and members of the Cave Exploration

Group of South Australia helped in excavating

the fossil vertebrae, | am grateful to Dr R. T-

Wells, Mr Lt. M. Thomas. Dr T, F. Houston

and Mr N, Pledge for their criticisms of the

manuscript. Mr P, Kempster prepared the

photographs for Fig, 8.

References

AUEFENBERG, W. (1963)—The fossil snakes of

Florida. Tulane Stud. Zool, 10, 131-216,-

Horrstetrer, R., & Gasc, J.. P, (1949).—Verte-

brae and ribs of modern reptiles, Jn C, Gans

(Ed.) “Biology of the Reptilia”. Vol. 1, pp.

201-310.

HOFFSTETTER, R., & GAYRARD, Y. (1964) —Ohser-

vations sur l’osteologie et Ia. classification des

Achrochordidae (Serpentes), Bull, Mus. natn.

Hist. nat., Paris (2). 36, 677-696.

Jounson, R. G, (1955).—The adaptive and phylo-

genetic significance of the vertebral form in

snakes. Evolution 9, 367-388.

Jounson, R. G. (1956),—The origin and evolu-

tion of the venomous snakes. Evalution 10,

56-65,

Kina, W, (1959).—Vertebra duplication, an osteo-

logical anomaly widespread in snakes. Her-

petologica 15, 87-88.

Loncman, H. A. (1925).—Ophidian Vertebrae

from cave deposils at Marmor Quarry. Mer,

Qld Mus, 8 111-112.

LYDEKKER, R, (1888):—"“Catalogue of Fossil Rep-

tiles and Amphibians in the British Muscum

of Natural History’ Part 1. (London.)

McDowéi, S. B. with a letter by H, Cogger

(1967). —Aspidomorphus, a genus of New

Guinea snakes of the family Elapidae, with

notes on related genera, J. Zool., Lond. 151,

497-543,

McDowe ct, S, B. (1970)—On the status and

relationship of the Solomon Island elapid

snakes. J. Zool., Lond, 161, 145-190,

MeRRILEES, D, (1968),—Man the destroyer: late

Quaternary changes in the Australian mar-

supial fauna. J-R. Soc. W. Aust. 51, 1-24.

Raw.inson, P. A. (1969).—The reptles of east

Gippsland. Proc, RK, Soc. Vict, $2, 113-128.

SIEGEL, 5. (1956) —“Nonparametric statistics for

the behavioural sciences.” (McGraw-Hill:

New York.)

Storr, G. M. (1964).—Some aspects of the geo-

graphy of Australian reptiles. Senck. biol. 45,

577-589.

Worrett, E. (1963),--"“Reptiles of Australia.”

(Angus and Robertson: Sydney.)

THE ONTOGENY OF THE VOCAL SAC OF THE AUSTRALIAN

LEPTODACTYLID FROG, LIMNODYNASTES TASMANIENSIS

BY M. J. 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, LIMNODYNASTLS TASMANIENSIS

by M. J. TYLeRr*

Summary

Tyner. M. J, (1975).—The ontogeny of the vocal sac of the Australian leplodactylid frog

Liniadynastes taymaniensix, Trans, R. Soc. &, Aust. 99(2). 85-87, IL May; 1975.

The ontogeny of the vocal sac of Limmeadynastes tasmaniensixy proceeds from_ initial

bilateral evaginations of the mouth floor, through median fusion to a wnilabular, suhmandi-

bular structure. The acquisition of pigmentation by the submundibular skin is a concomitant

process. It is guzgested that the vocal sac evolved by a path now reflected by onlogeny, and

involving progressive bilateral herniation,

Introduction

Vocal sacs occur only in male anurans and,

in most species, comprise inflatable, epi-

thelium-lined chambers Jocated between the

hyoid plate and the superficial mandibular

musculature. Data on the ontogeny of vocal

sacs are limited to studies on only a few,

mostly African, species (Inger 1956; Inger &

Greenberg 1956). The available information

is so inadequate that the possible contribution

of ontogeny to an understanding of the evolu-

uion of vocal sacs has not been assessed, At

present there is no published information on

the ontogeny of the vocal sacs of any of the

300 (approx.) species of frogs found in Aus-

tralia and New Guinea.

The Australian leptodactylid species Limno-

dynastes tasmaniensis represents an ideal

initia] subject for studies of vocal sac onto-

geny. This is because the superficial mandi-

bular musculature of adults has already been

described in detail, and there are published

observations on variation in the position occu-

pied by the vocal sac when it is inflated (Tyler

1971).

Material and Methods

Of 646 specimens af Limnodynastes tas-

maniensis in the South Australian Museum,

one series was found to exhibit ontogenetic

variation in the extent to which the vocal sac

intrudes above the superficial mandibular mus-

culature. This series comprises 18 male speci-

mens from a group of 68 males coflected at

West Beach near Adelaide on 1 September

1963 (SAM, R5290). The snout to vent length

of the 18 males ranges from 30 to 34 mm,

Dissections. were performed with the aid of

a low-power binocular microscope, and

measurements made with a pair of dial cal-

lipers. The muscles were stained with the re-

versible iodine/ potassium iodide stain des-

cribed by Bock & Shear (1972), in order to

differentiate the vocal sac from the surround-

ing striated museles. Muscle and vocal sar

terminology. follow that of Tyler (1971).

Observations

Secondary Sexual Characteristics

The secondary sexual characteristics of

Limnodynastes taxsmaniensis are as follows:

males possess a unilobular, submandibular

vocal sac, yellow pigmentation of the sub-

mandibular skin and glandular nuptial pads on

the first and second fingers. Females bear

broad lateral fringes io the first and secon

fingers, and sometimes to the third.

Focal Sac Ontogeny

The carliest step in the progress towards the

development of vocal sacs. involves the forma-

tion of a shallow and efongate involution of

the floor of the mouth on one side of the

tongue. There is a slight clliptical, ventral

depression with w mediad inzlination (Fig.

1A), and the Jateral margin of the depression

is level with the lateral border of the «anterior

cornu of the hyoid.

ee Sa

+ South Australian Museum, North Terrace, Adelaide, S. Aust. 5000.

#6 MJ, TYLER

Fig. 1. Selected progressive stages in ontogeny of

vocal sac. A. Single, elliptic ventral de-

pression. #, Bilateral expansion moediad.

C. Further mediad and initial cuudad

development. D. Media! unity of separate

sucs.

In four specimens, development. was con-

fined to such an evagination on the left side;

in a fifth there were bilateral evaginations.

From these slight folds the vocal sac develops

bilaterally, and as far as could be determined

quite concomitantly, into roughly circular bags

intruding between the superficial, ventral,

mandibular musculature and the deep inter-

mandibular muscles situated above them (Fig.

1B). At this stage of progress mediad develop-

ment is more pronounced than antecior or pos-

terior development, Simultaneously, that area

of the mouth floor between the anterior cornu

and the mandible becomes depressed, render-

ing the aperture 10 each portion of the sac

more conspicuous.,

As the two halves of the vocal sac approach

one unother, their posterior margins extend

further caudad (Fig. 1C). This posterior en-

fargement is accompanied by comparable

enlargement of the interhyoideus muscle into

a slight lobe, extending beyond the post-articu-

lar extremities of the mandibles. Ultimately

the yocal sac occupics the entire muscular

lobe, becoming united medially by loss of the

common medial wall (Fig. 1D).

The presence and extent of submandibular

dermal pigmentation was found to provide an

accurate external index of the presence, and

stage of development, of the vocal sac struc

ture, In specimens lacking the initial evagina-

tians in the mouth floor, the submandibular

skin Was cither entirely unpigmented or else

bore # few scattered chromatophores at the

petiphery of the mandibles. Development of

the evaginations was accompanicd by an in-

crease in the density of pigmentation and of

its medial limit. The pigmentation, and the

appearance of the bright yellowish background

color of the submandibular skin, progressed

in an ideiical 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

situation (he progression of the vocal sac from

paired, Jateral evaginations ta a single, large

sac could readily be regarded as a recapitula-

tion of evolutionary history, However, the

nature of the progression also indicates why

the anuran voral sac origimates in the way that

it does,

The floor of the mouth is supported by the

hyoid plate and its processes, and by muscles

communicating between the hyoid and the

mandibles, These supporting structures provide

what can be visualised as a broad and complex

sling in which the only gaps arc a narrow

lateral zone om each side of the tongue, so

situated between the anterior cornua and the

mandibles. In all anurans possessing vocal

SACS, apertures originate within these ‘unsup-

ported’ zones.

The probuble steps that Jead to the evolution

af vocal sacs in this species, or ily ancestral

stock, can be reconstructed quite readily.

Assuming an increase in the pressure of the

buccal cavity during vocal activity, the existing

sites of Ihe vocal sac apertures wre probably

the areas of least resistance: in these regions

the superficial tissue is of a rather elastic

nature, and presumubly subject to the greatest

distension. Jt follows that the first stage in the

ontogeny of the yocal sac of Limmodynastes

tasmaniénsis is precisely that initial event, Sub-

sequent stages could well have arisen from

litle: more lhan progressive bilateral hernia-

tion.

ONTOGENY OF FROG VOCAL SAC 87

References

Bock, W. J., & Sugar, C. R. (1972).—A staining INGER, R. F., & GREENBERG, B. (1956}.—Mor-

method for gross dissection of vertebrate phology and seasonal development of sex

muscles. Anat Anz. 130, 122-227. characters in two sympatric African toads.

J. Morph. 99, 549-574.

Incer, R. F. (1956).—Morphology and develop- TyLer, M. J. ¢ 1971).—Voluntary control of the

ment of the vocal sac apparatus in the shape of the inflated vocal sac by the Aus-

: 2 tralian leptodactylid frog Limnodynastes tas-

African frog Rana (Ptychadena) 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

BY R. J. DODSON*

Summary

DODSON, J. R. (1975).-The Pre-Settlement Vegetation of the Mt Gambier area, South Australia.

Trans. R. Soc. 8. 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. Dopson*

Summary

Dowson, J. R. (1975).—The Pre-Settlement Vegetation of the Mt Gambler area. South Aus-

tralia. Trans. R, Soc, S. Aus}. 99(2), 89-92, 31 May. 1975,

There are some conflicts as to the nature of the pre-settlement vegciation formations

around Mt Gambier and Glencoe. European settlers long ago cleared the areas of their

vegetation cover.

Pollen analysis of Brownes Like sediment reveals that the most likely formation around

Mi Gambier consisted of open. grassland with perhaps u sparse cover of woody taxa.

Introduction

Crocker (1944) teft gaps in his vegetation

map in the afeds around Mt Gambier and

Glencoe, in south-eastern South Australia, as

the original vegetation was no longer evident

at the time of his survey. Crocker (1944) and

Tiver & Crocker (1951) hypothesized thar the

areas were occupied by lightly wooded gijss-

land, and the trees cleared after settlement.

Woads (1862) recorded the vegetatian tn the

Blue Lake crater at Mt Gambier as thickly

wooded with several varieties of Melaleuca.

The vegetation is sparse in the photograph in

Hill (1972, p. 108) taken in 1860 and yet Hill

when describing an early record (1861) of the

first road between Mt Gambier (then Gam-

hiertown) and Port McDonnell on the coast

states. (p. 109)

“the route to the ‘Bay’ was through dense

bush country, mud and slush in winter, dust

it} summer, und tenanted by thousands of

kitngaroos at all times.”

Specht (1972, p. 203) in his vegetation map

of the South East simply records the areas

around Mt Gambier and Glencoe as cleared.

Brownes Luke occupies portion of one of

the craters which formed in the volcanic erup~

tions at Mt Gambier after 5000 years B.P,

(Fergusson & Rafter 1957, Blackburn 1966).

Tt ig 4-5 m deep, sits on collapsed volcanic

debris and its water surface, like thase of the

other three crater lakes, is an expression of the

regional water table (Bayly & Williams 1964,

1966). Hill (1972) recounts some of the early

records of water level changes of the Mt Gam-

bier lakes and it appears that the most spec-

tucular observed were in Browaes and Leg of

Mutton Lakes. These are the shallowest and

thus changes may have been more obvious.

Henty's hut, the first building in the area, is

said to have been erecfed in 1541 on the site

of Brownes Lake. Brownes Lake and Valley

Like filled and joined and the new lake

reached its maximum depth in the 1890's,

This paper provides data on early vegetation

at Mt Gambier. Pollen analysis is un ideal

method for tackling this problem as the crater

walls are mostly steep-sided and cat support

little local vegetation, Therefore a significant

proportion of the pollen rain is probably

regionally derived, Today the craters are

heavily exploited for recreation purposes and

most of ihe vegetatian within them is intro-

duced. The change from native vegetation

should be recorded in the fossil pollen record.

Methods

Before the Brownes Lake core site was

selected for investigation, areas of Brownes

Lake, Valley Lake and Leg of Mutton Lake

were checked for undisturbed sediment, Access

to sediment at Blue Lake was impossible with

the equipment at hand. A 160 cm sample of

peat was collected from Leg of Mutton Lake.

The dry periods in Leg of Mutton Lake (Hill

1972, pp. 112-114) which are unfavourable

for pollen preservation rendered this core un-

* Department of Biogceugraphy and Geomorphology, Australian National University, Canberra, A.C.T

2600. Present address: Geography Department, University of Canterbury, Christchurch, New Zealand,

Depth om

Restionaceas

Cyperaceag

Moriaphs dum

ri6l NOSGOGHYT

Triglochin

Paturnageton

Ruppia

Tiphe

Lepitaena

Hystrichospheres

@ S6 06

Poin exer

J. R. DODSON

fir

fad

Poaceae “ F

= r

a 6

a S

rr ab

5 ef

5 a

0 L

3 Unidentified pollen oo a |

<a Pacricdiimn

Gleivhenet Se

Loyraportiin ———T~

Monolete spores —_,

AL THUR EON

Lol

L66E

Fig. 1.

Leptosperaiett

Depth cm

Lucutypius Type 1

Lucalypiyy lype?

ducalyptes Typed

Mytaceae Type 1

Myrtaceae Type?

merrsinotde y

Cupressaceae

Cravacartns

(228um)

Casuariia €28uin)

ededelds

Pinus radiate

Dodonaca

eloaena

Chanepodiaceas

Asteraceae {Tuniliflorae)

Asteraceae (iiguliflarae)

Epacridaceae

Planiago lan volate

Brassicaceae

Lillaceae

Polygonaceae

Haloragaceae

Pimolica

Gyrastemonaceae

Malvageas

Swvlidium

Rastionaceas

Pollen diagram for Brownss Lake.

Soe 0 Ol OG

oS § § O5r OF

—

pnw aye]

WnS N3110d

(ODVINVId + SNMNId Suipnjoxe) WaTTOd IWIYLSIHHSL IVLOL

FARLY MT GAMBIER VEGETATION S|

guitable for pollen analysis. No sultable sites

were found ab the uccessible ureas in Valley

Lake nor in much of the Browtes Lake area.

In FPeuruary 1974, a 45 com core was

collected with a Desection sampler from the

core site on the eastern shore of Brownes Luke.

Tt consisted of 30 cm of black luke mud over-

lying 15 cm of pale brown (straw coloured)

material which was. largely clay-sized particles

of silica. This is probably reworked volcanic

debris, ‘The core was sampled on-site and

pollen analysis was carried owt in the Jabura-

tory using the standard hydrofluoric acid,

athali and Erdtman’s acetylosis methods as

described by Faegri & Iversen (1964). Resi-

dues containing pollen were dehydrated.

mmimnted in silicone oil, and counted until at

least 200 pollen grains of woody taxa hud been

recorded. Relative percentages for terrestrial

pollen and spore taxa were calculated against

a pollen sum of total lund plant pollen exclud-

ing the recorded introduced taxa (Pinus and

Plontazo lanceolata). Prequencies for aquatic

vascular plant pollen and Hystrichosphere

remunins were calculated agains! a pollen sum

of total aquatic pollen The results were

plotted on a pollen diagram (Fig. 1). Ecolo-

sical information for taxa in the study area

and details of pollen identification have been

given hy Dodson (1974).

Results and Discussion

The short pollen diagtany (Fig. 1) has not

heen divided into zones, but the presence of

intreduced taxa divides the diagram into post-

settlement (0-10 cms) and pre-settlement

phuses (20-45 cms).

The pre-settlement phase was dominated by

Asteraceae (Tubulifiorue). Posceae and

Preridium, with small numbers of herb pollen,

Tree pollen was. virlually absent, although

small and inereusing frequencies of scrub taxa

(Casuarina (<28um)—probably CL paludose,

wid Myrtaceae Type I-Leptospermuny jrai-

perinin ond Leplospernuan niyrsindides) were

obtanicd. Assuming that regional pollen

dominates, then the assemblage most likely

represents open (treeless) vegetation in the

region with a heath fringe around the lake. The

local vegetation was mainly Amhaceres,

Cyperaceae, and Myriophyllunr, representing a

shallow water environment al the core site.

This pollen assemblage iends to confirm the

Crocker hypothesis that trees were few, rather

than the early report recounted by Hill, unless

the scrub described by the latter consisted of

low pollen producers such as Banksia or

Acacia. There is no evidence tn the form o£

remnant vegetation to support this, The organic

und pollen and spore content of the sediment

was low (except for Anthoceros spores which

must have been derived locally), suggesting

cither slow sedimentation during alternating

wet and dry conditions (which could result in

the Joss. of pollen through oxidation) ar fairly

rapid in-wash of inorganic material from the

steep crater walls. On the evidence presented

hete if is not possible to favour either explana-

tion,

The post-séttlement phase of the pollen dia-

gram is dominated hy Poaceae, herbs and

Pinus radiata, and also shows increasing Euca-

Ivprus frequencies, Anthocervs and Cyperaceac

decrease in importance and Mysiaphyllam

dominates the aquatic spectra, indicating a

change to deeper water at the core sile, Since

the core site is near the edge of the lake, it

follaws that water would nor be deep there

until the level rose above its present position.

Since the rise accompanies the increase io

pollen of exotic plants, it seems likely that the

rise is the one recorded for the latter part of

the 19th Century when Brownes Lake and

Valley Lake were joined. The increase in

Eucalyptus pollen is undoubtedly due to the

plantings established in the craters for recrea-

tion facilities and the wildlife Teserve and not

fo any change in the nalive vegetation.

Acknowledgments

It is a pleasure to thank Gurdip Singh and

Joan Guppy who critically read the manu-

script, and the Australian National University

for supporting the work with finance anct

equipment,

References

Bayiy, f. A, By & Wirtiams,, W. DL (1964).-—

Chemical und bjologica!l observations o4

some volcanic lakes in the south-east of South

Australia, Ano 4. Maro breshwar Res. 15,

123-132,

Bayie, LAL CL, & Witiams, Wo DD. (1966) —

Further chemical observations on same vol

cunic lukes i the south-cast of South Aus-

baits: vtust. J. Mar, Freshwat. Res. 17, 229-

J, R. DODSON 92

BLACKBURN, G. (1966).—Radiocarbon dates relat-

ing to soil development coastline changes, and

volcanic ash deposition in south-east South

Australia. Aust. J. Sci. 29, 50-52.

Crocker, R. L, (1944).—Soil and vegetation

telationships in the Lower South-East of

South Australia. A study in ecology. Trans.

R. Soc. S. Aust. 68, 144-171.

Dopson, J. R. (1974).—Vegetation history and

water level fluctuations at Lake Leake, south-

eastern South Australia. I. 10,000 B.P. io

Present. Aust. J. Bot. 22, 719-741.

Fagor, K., & IVERSEN, J. (1964).—‘“‘Textbook of

Pollen Analysis.” Edn 2. (Munksgaard:

Copenhagen.)

Ferocusson, G, J., & RAFTER, T. A. (1957).—New

Zealand 14C age measurements—3. N.Z. J.

Sci. & Tech. 388, 732-749,

Hitt, L. R. (1972).—“Mount Gambier—the City

around a Cave.” (Investigator Press: Ade-

laide. )

Srecut, R. L. (1972) —‘“The Vegetation of South

Australia.” Edn 2, (Government Printer:

Adelaide, )

Tiver,, N. S., & Crocker, R. L. (1951)—The

grasslands of south-east South Australia in

relation to climate, soils and developmental

history. J. British Grassland Sec. 6, 29-80.

Woops, J. E. T. (1862).—‘Geological Observa-

Hons in South Australia.” (Longman: Lon-

ion. )

AUSTRALIAN LEPTODACTYLID FROGS OF THE

CYCLORANA AUSTRALIS COMPLEX

BY M. J. TYLER*® AND A. A. MARTIN}

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.

AUSTRALIAN LEPTODACTYLID FROGS OF THE

CYCLORANA AUSTRALIS COMPLEX

by M. J. Tyner* and A. A, Martint

Summary

lyter, M. J., & Martin, A. A. (1975).—Australian leptodaciylid frogs. of the Cyelorana

australis complex: Trans, R, Soc. 8. Aust, 99(2), 93-99, 31 May, 1975,

Cyclorana australis vs now defined is shown lo comprise two closzly related species:

C.. australis confined to norihera 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

Ulvergence is so limited that hybridization is considered possible in sympatry-

Introduction

In recent years, examination of the biology

and morphology of several geographically

widespread “species” of Australian frogs has

revealed that each comprises a complex of

species, For example, Crinia signifera as recog-

nised by Parker (1940) is now known to be

a complex of seven species (Moore 1954,

Littlejohn 1957; Main 1957; Straughan &

Miin 1966; Tyler & Parker 1974): Mixophyes

fasciolaius is now four species (Straughan

1968) .and Linmnedynastes dorsalis is also four

(Martin 1972).

The most neglected leptodactylid genus 1s

Cyclorana, of which the type species is Alytes

australix Gray (1842), described from material

collected in the Northern Territory. This

4pecies, us currently defined, extends from

northern Western Australia to northern New

South Wales: a geographic tange of approxi-

mately 3500 km. The conspecificity of indi-

viduals from the extremes of this extensive

range is obviously suspect, and even the most

cursory comparison of specimens of C. aus-

tralis from the Northern Territory and

northern Western Australia with those from

Quecnsland reveals striking differences between

them. The northern individuals (end to have

a tather elongated head, a distingl, dark rostral

stripe and a narrow subocular bar. In con-

trast, most individuals from Queensland are

particularly robust animals with a broad head,

and frequently obscure head markings: a

population described as Cyclorana novae-

hollandiae by Steindachner (1867), and as

Phractops alutaceus by Peters (1867). Both

names were referred to the synonmy of ais-

tralix by Boulenger (1882),

We have assembled and cxamined large col-

lections of Co australis (sensu lato) from

various sources. Here we report our findings

and propose the recognition of a complex of

two. species.

Methods

The specimens reported are deposited in the

following institutions: National Museum of

Victoria (NMV): Naturhistoriska Riksmuseet.

Stockholm (NR); Department of Zoology,

University of Melbourne (MUZD); Northern

Territory Museum, Alice Springs (NTM);

Queensland Museum (QM); South Australian

Museum (SAM): and Western Australian

Museum (WAM)

Measurements of specimens (to 0.1 mm)

were obtained with a pair of Helios dial

callipers. Abbreviations employed in the text

and tables are as follows: F = foot length (the

distance between the proximal end of the tar-

sus and the distal tip of the fourth toe); HI.

— head length (the distance between the an-

terior extremity of the snout and the posterior

margin of the tympanic annulus); HW = head

width (the maximum width of the head,

usually taken at the posterior extremity of the

mandibles); TL — tibia length (obtained by

placing the tibia between the callipers}); S-V

= snout to vent length (the distance hetween

* South Australian Museum, North Terrace, Adelaide, S$. Aust. 5000.

+ Department of Zoology, University of Meibourne, Parkville. Vic. 3052.

o4 M.3. TYLER & A. A. MARTIN

the anterior tip of the snout and the anterior

margin of the cloaca).

Ratios caleulated and subjected iu the Stu-

dent t-test were TL/S-V, HL/HW, F/S-V,

F/TL and S-V/HW. Larval stage numbers

follow those employed by Gosner (1960).

Mating calls wete recorded in the field using

at Uher 4000 Report portable tape recorder

and Beyer M69 dynamic microphone, al a tape

speed of 19 cm/sec. Calls were analysed by

usc of a sound spectrograph (Kay Model

h061-A Sona-Graph) with the overall reg-

ponse curve maintained in (he FL-I position,

Three calls of each individual were analysed

and mean values calculated. Each call was.

analysed {wice; a narrow-band (45 Hz band-

pass) analysis at recording speed to determine

duration and dominant frequency, and a wide-

bund (300 Hz handpass) anjlysis at half

recording speed to resolve fundamental fre-

quency.

The Cyclurana australis complex

Frogs of the Cycelorana australis complex

ate relatively Jarge animals; the snaw. to vent

length of adult males ranges from 61,4 ta 1,4

mm, and that of females from 69.9 to 102

mm. They are all generally robust with a broad

and frequently bloated body and’ retatively

short limbs (TL/S-V range = 0,34-0,46)

All members of the complex exhibit exostosis

Stam AN

ie af |

4 *, é ay

pe Poy e Se } “|

b 3 1 hy « «

° hd | “i < }

' : i es

7 a * “iL

¢ : 4

ee oe * a

\ ‘ _ e + By 4

‘ ‘ : a

. Pit RET Y /

ad a Cansreaus ‘a ' ony if

ce CoWOVAE RIL, ANDIAL “ery, “d

Fig. 1. Geographic distribution of the frogs of the

Cyeloraia australis complex. Circles —

C. dustralis; triangles = C. noveehollan-

diae. Closed symbols indicate sites of the

materia] examined, and the open circles

one of the following literature references:

Brattstrom (1970), Loveridge (1935),

Moore (1981), Parker (1940), Slevin

945),

of the maxillary, premaxillary, nasal, fronto-

parietal and squamosal bones. On the dorso-

lateral body sutfaces there are continuous or

disrupted, longitudinally orientated skin folds

commencing behind the skull, and terminating

ubove the groin.

Nothing is known of the breeding biology

of the members of the complex, but they are

probably opportunistic breeders. The egys ate

small and pigmented (ovidiameters of ovi-

ducal eggs range from 1.1 to 1.3 mm), and the

tadpole (one species) is of the hylid type with

{wo upper and three lower’ rowg of Jabial teeth.

an acumitate tail tip and a median or slightly

dextral anus.

The geographic range af the complex ex-

(ends from northern Western Australia to

northern New South Wales (Fig, |), O£ what

we demonstrate to be twa component species;

C. australis (sensm stricto) accurs in northern

Western Australia, the Notthern Territory and

northern Queensland (o the west of the Divid-

ing Range; C. rovaehollundiae is found

throughout Queensland and extends cas far

south as the northern part of New South

Wales

Cyclorana australis (Gray}.

FIG, 24

Alytes australis Gray (1842),

Cliroleptes. australis, Gunther (1858); Boulen-

ger (1882) (part).

Phractops dustralis, Fry (1914).

Cheirolepies wustralis, Spencer (1901) (replace-

ment name for Cliralepies)

Cyclorana austratiy, Parker (1940) (part),

Hype locality: “North coast of Australia’. ¢ Port

Essington, Northern Territory. )

Material examined: Western Australia—WaAM,

R8732, Carlion Reach, Ord River; WAM,

R43067, Crystal Ck; WAM, R1558-59, Drysdale

River Mission; WAM, R21233, Fossil Downs;

WAM, 1377, 43282-86, 42399-42422, Kotum-

buru; WAM, R22369-75, Kimberley Rescarch

Stn; SAM, R4769-70, RS070, Kununurra; WAM,

R1654-57, Lundor Sin; WAM, R42387, 80 km S

of La Grange; WAM, R42536-40, 42381, 43478,

43491, Mitchell Plateau; WAM, R42530-35, Main

Ord River Dam Site (spillway); WAM, 42424,

Mt Hart; WAM. R32099, Mt Anderson; WAM,

R32291, Mi Barnett; NR, 1362, Mowla Down;

WAM, R13726, Oscar Ranges; NMV, D2354-5S,

Port George IV; WAM, R32149, St George Range.

WAM, RI1/208, RIi894, R1IZT332, Wotljulurn;

WAM, R26769-70, Point Springs, Webher Range;

WAM, R323S51A, Wyndham; WAM, R25093, 40

km SE of Wyndham; WAM, R20307, Yeda Cross-

mg. Northern Territury—SAM, R14332, Adetnide

River, NM¥V, DI2702, Karrow Ck, NMY,

D8307, D8315, DS8327. QM, J17K8S. 2985, SAM,

FROGS OF THE CYCLORANA

AUSTRALIS COMPLEX 95

20mm

Fig. 2. A. Cyclorana australis from Kununurra, W.A. B. Cyclorana novaehollandiae, 15 km

N of Goondiwindi, Qld.

R8968, Darwin; SAM, R13453, Elsey; SAM,

R13450, Howard Springs; WAM, R1935-36,

R21318, SAM, R14330-31, Katherine; SAM,

R4877, Mt Bundy Stn; SAM, R13349 A-G, Smith

Pt, Coburg Peninsula; WAM, R24007, Snake Ck;

NMY, D12704-08, SAM, R13275 A-L, Tennant

Creek; NTM, 498, 525-26, 50 km N of Tennant

Creek. Queensland—SAM, R5010, R5070, Doom-

adgee Stn; NM'V, D8437-38, SAM, R4934, Morn-

ington I.

Description; The diagnostic characters of this

species are: size large, males 70.8-78.0 mm

and females 71.0-81.0 mm in snout to vent

length; S-V/HW ratio high (mean 2.31);

head width only slightly greater than head

length (mean HL/HW ratio 0.89); TL/S-V

ratio moderate (mean 0.32); foot relatively

long (F/S—-V mean 0.40).

6 M. J. TYLER & A. A,

Development and exostosis of the super-

ficial skull bones are moderate in this. species,

The dorsal limit of the squamosal is such that

there is a very broad gap between the squamo-

sal and the frontoparietal On the fronto-

parietal exostosis is confined to the lateral bor-

ders of the bone. The sub-orbital portion of the

maxilla slopes steeply to the Jabial margin and

is not expanded there into a lateral ridge.

Cyelovana australis. is usually pale olive or

grey in preservative and bears a naccow and

very sharply demarcated dark brown rostral

bar and a narrow sub-ocular bar which ter-

ininates far above the labial margin. The

lateral body surfaces are commonly heavily

suffused with darker pigment. The backs of the

thighs are darker and densely variegated with

light pigment.

Geographic variation; The presence of darker,

irregular patches on the dorsum varies through

the range, Dorsal spots are absent from a series

of ovec 100 specimens from Kununurra. Speci-

mens from Tennant Creek have light suffustons

of pigment, and those from the north-eastern

portion of the range are heavily pigmented

with dark stippling. Immaculate and marked

specimens occur on Mornington Tsland.

Variation in some of the pertinent body pro-

portions is summarized in Table 1.

kegs: A gravid femaie from. Kununurra con-

tained approximately 1000 eggs varying from

1.1 to 1.3 mm in diameter. The eges have

black animal poles.

Larval morphology; A series of tadpoles was

obluined at Kununurra on the Ord River by K.

MARTIN

Fig. 3. ‘Tadpole and tudpole

Cyclorana diestralis.

mouthparts ol

Cole in February, 1963. The following notes

are based on four specimens from this series

at stages 36-39, All specimens are poorly pre-

served and badly distorted, so that their total

length range of 50-65 mm is only an approxi-

mation.

The spiracle is sinistral and the anus. median

or very slightly displaced dextrally. The over-

all appearance (Fig. 3) is similar to that of C’.

cultripes and C. platyeephalus (Watson & Mar-

tin 1973).

TABLE L

Geographic variation in proportions of Cycloruna species

(Ranges are give With meany and standard deviations in parentheses)

Speciey and locality N HL/HW TL/S-V S-V/ HW F/TL Fis-V¥

C, aastralix

Rurnunurra, WoA, 7 0,81 — 0.93 0.38 — 0.45 2.43 —- 2.33 O8L - 1.00 0,38 UAd

(0.86 ob 0.04) (0.42 + 0.03) (2.23 0,09) (0.97 = 0.03) (0.41 = 0.02)

Smith Point, N.T, 7 (89 — 0.93 0.40 0.45 2.27 2.44 0.92 — 0.98 0.39 — 043

(O91 > 0.01) (O43 #0,02) (2.46 + 0.06) (0,95 + 0.02) (0.47 a D.01)

Tennant Creek, NT. 7 0.87 — 0.94 0.40 — 046 2.25 — 2.49 0.91 — 0.98 0,38 — 0,42

(0.90 + 0.02) (0,42 + 0.02) (2.36 + 0.08) (0,94 + 0,02) (Q,39 ae QOH}

Mitchell Plateau, W.AL 4 0.95 — 0.99 (44 — U.A0 2.26 — 2.51 Q.8R — LOS O38 (),43

(0.97 = ON) (0,46 = 0.02) (2.41 2 0.12) (0.95 = 0.07) (0.41 + 0.01)

C navachollaudiae

Cooktown, Old 7 0.81 0.83 039 O43 1.93 2.08 0.87 — 0.96 0,36 — 0.41

(0.83 + 0,02) (O44 + 0.01) (2.02 + 0.04) (O81 + 0.037 (0,38 se 1)

Calliope, Qld 3 0.82 — 0.88 (1.39 — 0.40 2.03 — 217 0.89 — 0.94 0.34 — 0.39

(O86 ate 0,04) (0.39 —& 0.01) (2.12 + 0.01) (0.92 + 0.02) (0.36 0.01)

Cunamulla, Qld 3 0.78 — 0.85 0.34 — 0.41 (97 — 2.13 0.93 1.02 0.32 0.39

(0.82 = 0.03) (0.37 = 0.02) (2.05 <b 0.06) (0.96 = 0.03) (0.36 > 0.03)

FROGS OF

The mouth is sublerminal (Mig. 3) with a

large horny beak and papillae around [he sides

und back of the mouth dise. There ate two

upper rows of labial tecth, the second Uis-

rupted medially, and three lower labial rows

of which the first is similarly disrupted,

Mating Call: Calls. of five individuals were

recorded on 13.xi11971, 14 km E of Daly

Waters, N.T, The frogs were calling on land

heside a small water-filled channel. Wet-hulb

air temperature at the calling sites was 241°C.

The call is a short, well-tuned note repeated in

long sequences. The mean. cull duration is 152

msec (range 122-204), The fundamental fre-

yuency 18 199 He (ringe 183-209), but con-

tains little energy: most of the energy is in the

third, fourth und filth harmonies (about 600,

300 and 1.000 Hz) which are approximately

equally emphusized,

Geographic Range: Cyclerana australis extends

trom the Kanberley Districe of northern Wes-

tern Anstralia to the GulP District of Queens-

land. dn che Kimberteys it is clearly widely

distributed, and in the Northern Territory it

extends us far inland as Barrow Creek.

Absence in the north-western portion of the

Northern Territory may simply reflect inade-

quite sampling,

Cyclorana novaechollandiae Steindachacr.

Cyeloaruna novaehollandine Sicindachner (L867),

Plrractops alutacens Peters (L867).

Chiraleptes australis, Boulenger (1882) (parv)-

Phraclops australis, Laveridge (1Y35)-

Cyclorana australis, Parker (1940)

Mocre (1961), plate 35, Fig. 2,

type locealitt: Rockhampton, Qld.

Marerial exuntined: Queensland —NMV, D13049,

SAM, R9SI7, ROR3S, Balle Cump; MUZD, 90-

92/70, 7 km SW of Calliope: QM, 1431. Colos-

seurn: QM, J18062, J1SD66, Condamine River,

Cecil Plains; NMV. D13049, SAM, R11523-24_

Cooklowns OM, 120685-91, 8 km W of Cooktown;

SAM, R9690, Fdward River Sin; OM, F20R4-85.

1/2%44.. Bidsvold; OM, Jt4383-84, Gilruth Plains.

Cunnamulla; MUZD, 56-58/70, 75/70. 9 & 15 km

E of Goondiwindi; QM, J5611-I7, Mackay: SAM,

R4743, Mupoon Mission Sing SAM, R9734. Mary

River; QM, J14159-67, Mitchell R. Mission; SAM.

R10419, Prestwood, Gilbert Rivers SAM, R3640,

Rockhampion:; QM, #10482, SAM, R3686. St

George: OM. 1218-89, Stannary Hills. SAM,

R1935, Stewart River; SAM. R9691, Strathgordon

ALS.; QM, 122227-29, Surat; NMYV, D7542, OM,

14644. Townsville: OM, 13480, Victa, Coengoola;

OM, J18063-65, Waratah Stn, Cunnamulla.

Deccrintiony Snour to vent lenath of males

A1.4-$1.4 mm, females 74.8-101,2 mio; head

nificeahly hroader than Jong (HL/ WW mean

O83): S-V/HL rane low (mean 2.05).

fpart);

THE CFKCLORANA

AUSTRALES COMPLEX x7

TL/S-V ratio rather low (mean 0.40); foot

short ¢F/S-Y mean O37),

There is exireme exostusis of the skull

hones, The squamosal in large specimens Is

usually so heavily overlain with secondary

bone thal tt is visible through the skin, forme

ing humps resembling parotad stands, and

extends so far suportorly that it approaches

the margins of the frontoparietals, The fronto-

parictal is cnlipgly exostosed, but the lateral

margins are raised by bone depusition. so pro-

ducing « deep, median furrow. ‘he suborhital

portian of thy muxilla projects, forming a high

und often concave shelf.

The constricted pupil in stk living specimens

from Cooktown approximated a rhomboid

shape (see discussion).

In preservative, C. noevaehollandide is pale

brown or grey, and is immaculate, Tightly

marked with scattered dark brown or blackish

markings, or-else very densely pigmunted wilh

such markings. The suborbittl murking is

broad and uaually reaches the labial base of

the maxilla, The hacks of the thighs are usuatly

very dark leaden grey and lack lighter vermicu-

lations.

In life the series from Caoklown were af

immaculate dull sandy yellow dorsally The

rostral stripe was dark brown, and similarly

colored, small disrupted patches occurred on

the inferior margin of the manilli. The iris was

polden and suffused with dark brown laterally

and inferiorly. The posterior surfaces of the

thighs were leaden grey, whilst the ventral sur-

face of the body wall was a dull pearl bewring

faint grey vermiciulations on the throat,

Geographic variation: There is considerable

variation in skull structure and coloration of

the dorsum of this species. Comparison of

small samples of extreme vanants led us to

conclude amitially thar [Wo Species were in-

volved, Examination of larger scries, however,

has revealed the ovcurrence of forms of inter-

mediute appearance, The variation may be

summarized as follows. All specimens feon

central and southern Qucensland have high

skulls with 4 gently sloping maxilla and a

densely pigmented dorsum. There is siriking

variation in individuals from northern Queens-

land, Some are densely pigmented whilst others

are immaculate, The skull of the immaculate

individuals is either similar to that of the pig-

mented frogs, or is spatulate and distinctly

Hattened. Unfortunately we have been unable

to devise a means of objectively estimating

skull depth with any degree of accuracy, We

oR M1. TYLER & A, A- MARTIN

hold the opinion that the high and the spatu-

late forms of the skull represent diferent evo-

lutionary trends of development.

Whereas the extremes are clearly different,

assessment of the significance of the observed

Variahion ts. compligated by the existence mn

northern Queensland of a number of inter-

mediate forms that cannot he referred to either

form. In addition there are animals in which

the terminal partion of the skull is more élon-

gated. This variant occurs only on the Cape

York Peninsula and at lacalities at the bare of

the Gulf of Carpentaria. Morphornetric data of

Iwo small series are summarized jn Table |.

Eggs: Ovaducal exgs of three gtavid females

ranged from 11] to J,3 mm in diameter, An

estimate of the number in une individual

exceeded 1.000, The cazs have black animal

poles.

Mating Call; The mating calls of two indi-

viduals recurded 5 km SW of Calliope. Queens-

land, on 187.1970, are very similar to those

of C. australis. The frou were calling on land

beside a rain-filled roadside ditch, with a wet-

bulb wir temperature of 24.6°C. The spectral

structure of the calls of the two species is

essentially identical, with C. moveehollandiac

also having emphasized harmonic bands at

bout 600, 800, and 1,009 Hz. However, its

call duration is considerably longer (mean 249

msec: range 235-262 msec). Judged on the

hasis of the levels of difference in mating call

structure of sympatric anuran spevies, this

diference in duration dogs not represent diver-

gence of sufficient magnitude to achieve repro:

ductive isolation, Hence if C. australis and

C. nevaehollandiae occur in sympairy (as they

may in the Gull District) we would expect

them to hybridize, A similar pattern ol

mirked morphological differentiation, accom-

panicd by very litthe mating call divergence,

characterizes the Western Australian Janimody-

nastes dorsalis and the eastern L. duenerifi

(Martin 1972),

Gvoeraplle Range: Cyelorana nevachollandiae

ninges From the Cupe York Peninsula of

northern Queensland to the New South Wales

border,

KEY TO ADULIS

Buck of thighs variegated, suborbital bar narrow

and not reaching hasz of maxilla; maxilla gently

Yopiig and nol cxpanded laterally C. australis

Back of thighs not variegated; suborbital bar (if

present) usually broud and reaching base of

maxilla: muxila expanded tw form laters! shelf

Co novavhollandiag

Discussion

The morphological complexity of C. neve

tieiandive as defined here is unparalleled

amongst Australian anurans, We believe that

a study of species isolaling mechanisms, such

as male mating call, in north-casterit Australia

could reveal the existence of two or possioly

even three species, Our action of resurecting

C. nevachollundiue From the synonymy of C.

australis is therefore onky the first step towards

an understanding of the CC. anstraliy complex,

The biological dita, eg. mating call structure,

that are mecessary for final resolution of the

problem may be extremely difficult to obtain.

The northern Queensland populptions jire

apparently opportunistic breeders which may

call at a locality on only one of two nights

each year (C, Tanner, pers, comm.), It seems

justifiable, therefore. to treat the complex al

this preliminary level,

In terms of skull structure, C, australis 1%

clearly the most siraplified and primitive mem

her of the complex, exhibiting lintited develop-

ment of skull bones and the least extensive

exostosis, All the variations in the form of the

skull and exostosis of the cranial hones such

us the maxillary and squamoasal in €. #ovar-

hollandiae can reasonably be devived from ©.

australis,

The concept of C. aastraliy being the prumi-

tive member may be acceptithle morphologi-

cally, but it is more difficult to conceive zooged-

graphically in view of the absence of any mem-

ber of the genus Cyclorwna in New Guines.

The georraphic area occupied by the members

of the C. australis complex includes both high

rainfall and relatively arid areas; ie, it does

not appear to be limited clitnatieully. Thus. if

the complex orginated in northern ar north.

western Australia it is surprising that it should

he absent from New Guinea. Jennings (1472)

estimates that the most recent land communi-

cating with New Guinea at “Vorres Strait ter-

minuted only 6,500-8,000 years ago: thus it

is. possible that colonization of north-eastern

Australia by the complex occurred sirbse-

quently.

The gross differences in morphology charac-

terizing the members of the C. australiy com-

plex represenis a situation unique among Aus-

tralian anurans. Differentiaiun in other

species complexes, ¢ven those with extensive

disjunctions of range (é.g. south-eastem and

south-western Australig) is accompanied by

anly slight morphological divergence. IL scenes

FROGS OF THE CYCLORANA AUSTRALIS COMPLEX iced

probable, therefore, that the complex is of

cojisiderable antiquity. Two further circtim-

stances lend support to this suggestion, One is

the affinity of Cyclorana with the Hylidae

demonstrated by Tyler (1972) and Watson

& Martin (1973). The other stems from our

observations on the pupil shape of C. novae-

hellandiae. As stated above, the constricted

pupil of C. navaehollandiae is almast rhom-

boid. In fact, the ventral margin is an obtuse

angle and the upper a broad curve. This

curvature is diffleult to detect during extreme

constriction, Lynch (1971) considers the ver-

tically oventated pupil to be the primitive and

the herizontal pupil the derived state. How-

ever, from the fact that Nyecrinysres (a genus

that can only be derived trom Litoria, a hori-

zontally-pupilled stack) has a vertical pupil, iL

is clear thal. vertical arientalion can be a

derived state. The trend to one or other orien-

lation of pupit shape could be accomplished

most readily fram a rhomboid. That such #

structure ocetlrs in living Cyclorana is con-

sistent With our hypothesis of its artiquity,

Acknowledgmeiits

For the donation of specimens or the lown

ul those in their care we are indebred ta Mv K.

Cole, Miss J. Covacevich. Mr J. Coventry, Mr

D. Howe, Dr B.S. Low, Dr Gi. Stor’, Me C.

Tanner and Dr G. Vestergren. Mr G. F. Wat-

son assisted in the field. Phatographs were pro-

vided by Mr B. N. Douetil and the tape of the

mating call of C. australis by De B Law and

Dr D. Gartside.

Our thanks are also due to Dr J. Ling for

reading the manuscript, and to the Trustees of

the Science and Industry Endowment Fund

for providing travel funds to M,J.7, permitting

visits to vanous Australian muscums to

examine collections.

References

Boutencen, G. A, (1882).—"CalaJogue of the

Batrachia Salientia s. Ecaudata in the collec-

ion of the British Museum.” 2nd Edn.

(London, )

Brattsraom, B. H. (1970),—Thermal acclima-

tion in Australion amphibians, Comp. Bio-

chen. Pliysial. 35, 69-103,

Fry. D. B. {1914)-~On a collection af reptiles

and batrachians from Western Australia. Kec.

WW, Ausi. Mus. 1, 174-210,

Ciosver, K. L. (1960).—A simplified table tor

staging anuran embryos and larvae with notes

on identification, Herprtologica 16, 183-190.

Geav. J. EB. (1842).—Descriptlan of some hither-

to unrecorded species of Australiaa reptiles

and batrachians. Zoli. (disc. 51-57,

Guntuer, A. (1858)—"Catalogus of the Bat-

rachin Salientia in the collection of the

Brilish) Museum.” (London, )

Jewarnes, J, N, (1972).—Some altribukks of

Torres Strait. /a: Walker, D. (Ed.) "Bridge

and Rarrier: The natural and cultural his-

tory of Torres Strait”. Dept of Biogeography

and Geomorphology, Publ, BG/3, Australian

National Wniversily, Canberra.

Liriteso“n, M, J. (1957)—A_ new species of

frog of the genus Crinie. W. Aust. Nat. 6,

18-23,

LOVERIDGE, A. (1935).—Australian Amphibia in

the Museum of Comparative Zoology, Canm-

hridge, Massachusetts. 8x/l, Mas, Comp,

Zool. 78, 1-60.

tyson, J. BD. ti97l).—RFvolutionary — relation.

ships, ostedology, and zoogcography of lepto-

dactylid frogs. Mise. Publ. Mus, Nat. Hist.

Univ, Kansas (59), 1-258,

Materin, A. A. (1972).—Studics in) Australian

Amphibia U1. The Cimnedynasres dovselic

complex (Anura: Leptodactvlidac). Amit. 4,

Zool, 2, 165-27 1-

Moore, J. A. (1954).—Geoeraphic ad genetic

isolation in Australian Amphibia. Amer. Nad.

88, 65-74,

Parker, EE W. 11940)—-The Australasian Frogs

of Use family Leptodactylidae, Nov? Zool.

42, 3-106,

Pecers, W.

Aforiarnh, K. Press.

1867, 13-37.

Siovin, J. R. (1955).—Notes on Australian

amphibians. Prac. Calif. Avud. Sci, 28, 355-

392,

Spencen, B. (1901). -lwo mew species of frogs

from Victoria, Proc, R. Soe, Mie, New Ser.,

13, 175-178.

STEINDACHNER, K. (1867) —Amphibien. Jit: “Reise

der Osterreichischen Fregate Novara um dic

Erde in den Jahren 1857-1859." Zoologische

Theil, 1(4), 1-70. (Vienna).

S5rRAUGHAN, I. R. (1948)}—A taxonomic review

of the genus Mixephyves (Anura: Leptodacty-

lidae), Prog. Linn. Soe, N.S AV. BSL 1). 52-39.

StwaAvcdan, LR, & Main, A. R. (1966}—

Speciation and polymorphism in the gcrius

Crinfa Tschudi (Anura; Leptodactylidue) in

Queensland Pree. R. Soc. Qh TCA), 11-28,

Lvirre, M. J. (1972)—Superficial mandibular

musculature, vocal sacs and the phylogeny of

Australo-Papuan Jeptodiuctylid frogs. Rec. WS.

Aust. Mus, 16(9), 1-20,

Treen, M.J., & Parker, F. (1974). New species

of hylid and leptodactylid frogs from southern

New Guinen. Trans. R&R. Sov. S. Aust, 982),

71-78.

Watson, G. F.. & Martin, A. A, (1973) —Life

history. larval morphology and relationships

of Australian leptodactvlid frogs, rans, #

Soe. &. Aust. Y70L)}, 33-49.

notizeny

£1867) —Herpetalogische

Berliri,

Akad. Wiss.

VOL, 99, PART 3 30 AUGUST, 1975

TRANSACTIONS OF THE

ROYAL SOCIETY

OF SOUTH AUSTRALIA

INCORPORATED

CONTENTS

Mahoney, J. A. The Identity and Status of Thomas’ “Lectotype” of Leportiius

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' “LECTOTYPE” OF LEPORILLUS

APICALIS (GOULD, 1853) [RODENTIA: MURIDAE]

by J. A, Manoney*

Summary

Matoney, J. A. (1975).—The idenuty and status of Thoms’ “lectotype” of Leporillus apicalis

(Gould, 1853) [Rodentia: Muridae].. 27rans, R, Soc. S. Aust 99(3), 101-104, 30 August

1975,

The specimen selected by Thomas as the lectotype of Leportllus upicalis (Gould, 1853) was

misidentifizd by him and belongs to Leporillus conditor (Sturt. 1848). It does not belong io

the lype series of L. apicalis, therefore Thomas’ lectotype selection for that species is invalid.

The type material of F. apicalis and L. conditor 1s missing, Thomas’ “lectotype” of L, apicalis

und a second specimen of L. conditer in ihe British Museum (Natura) History) could belong

to the type series of 1, cenditor, Evidence for the occurrence of L. apicalis in Tasmania is

lacking.

Infroduction

The name Hapalotis apicalis was proposed

by Gould (1853a) for a new species of rodent

from Australia. He did not state in the original

description if one or more specimens were

heing described nor did he give any locality.

Later, Gould (1853b) stated that he possessed

# single example procured by Mr Strange in

South Australia, and he illustrated the external

features,

Thomas (1906a) nominated Hapalotis api-

calls as the type species of 4 mew genus,

“Leporillus, and subsequently {Thomas 1921a)

he selected British Museum (Natural History)

specimen 1853.10.22.15! as the leciotype of

Leporillus apicalis, describing it as a female

from §. Australia. Explaining his leciotype

selection, Thomas (192lc) stated that

although Gould had in his collection two speci-

mens of that species, he seems to have done

his describing from only one of them (BM,

$3.10.22.14 (sic)\—lapsus for BM, 53.10.22.15)

—the worst of the two, young, and with an

imperfect igil, Thomas concluded with the

remark that probably from memory, and cer-

tainly wrongly, Gould stated that the species

had a white-tipped tail, but his overlooked

second specimen fadult with nearly perfect

skull* and quite perfect tail (BM, 53.10,22.14))

has the Jatler organ uniformly blackish or

brownish above and dull white below, and

there is no indication of the white tail-tip

found in so many Australasian Muridae.

A study of three Australian Muscum speci-

mens of L. epicalis, and the literature, enabled

Troughton (1923) to confirm that Gould was

correct in attributing a white tail-tip to the

species, Troughton stated also that Thomas’

remurk that Gould seems to have done his

describing fram only one of his two specimens

means that that specimen must he accepted as

the holotype.

Taie (1951) treated specimens 1853,10,22.14

and (853.10.22.15 as ‘‘cotypes" of L, apicalis

” Department of Geology and Geophysics, University of Sydney, Sydney, N-S.W. 2006,

' The first rwo digits of British Museum (Natural History) registration numbers of mammals are fre-

quently omitted from pubiications. Thus Thomas. uses. $3.10.22.15 for 1853.10,22,14-

* This skull is registered as 1854.10.21.1 and the Register entry mentions a stuffed specimen and refers

to 53,.10.22.16. I

Eritish Museum (N.H,)

have been unuble io find a

therefore 1 am following Thomas’

1853.10,.22.16 in the

specimen numbered

that 1853.{0.22.94,

conclusion

1$53,.10.22.16 and 1854.10.21.1 belong to the one individual; but it is possible that 1853.10,22,14,

identified as Hapalotis apicalis in the Register, is Jost and the skin now numbercd 1853.10.22.14 iy skin

1853.10.22.16 with an incorrect number. A note in Thomas’ handwriting attached to skin

(853.10.22.14 and stating that this specimen was considered to be the type seems io refer lo ithe

Museum Register where “type” has been written opposite the number 1853.10.22.16. A portion of the

posterior half of the cranium, and the Jeft mandibular ramus, are missing from the skoll.

102 J,

TABLE |

Skall measurements (in mim) of Themes’ “lecto-

type" of Leporttiue apicalis (Gould), Bd, (NAL),

853102215,

Miisimum width across nasals , 5

Minimum width across igh 2ygo-

matic plate, ' 44

Length of right M7 _ . - 9.7

Width of left M4 - , . 25

Wirth of left M* ‘ . ' 27

Width of left M3 : i : ' 2.4

Length of left Myen - . - . 9.1

Width of left My - ; - . A

Width of loft Ma & ' v ' 2.7

Width af left Ms . ’ ; ' 32.4

The teeth mensutements are For the crowns of the

seth.

and referred to them as adult and young

temales. from “South Australia’. collected by

FP. Strange. He briefly descmbed the skin and

skull of each and recorded measyrements of

them. He does not tefer, in his account of L.

apicalis, to Thomas’ lectotype selection or to

‘lroughton's recognition of a holotype for the

species.

Identity of the “Iectotype”

British Muyeum (N.H_) specimens

1853,10,22.14 and 1853.10.22.15 are examples

of Leporillus conditor (Sturt, 1848) and not

gpucimens of Leportiins apiealiz (Gould, 1853)

as believed by Thomas, They do not agree with

the original description of £. apiealis (Gould)

and because of this 1 do not accept that cither

of them belong to the type series of thal

species. Consequently, I regard as invalid

Thomas’ lectotype selection for L. apicalis,

Thomas (19216) stated there is no specimen

of L. conditer in the British Museum, and

turther demonstrated his unfamiliarity with its

characters by suggesting that it possibly

belongs to Noiomys, a genus of Australian

hopping mice and rats.

Measurements of the badly damaged skull of

Thomas” “lectotype” are given in Table {. The

skull is illustrated in Figs 1-4.

Discussion

The type matenal of hoth L apiesliy and

L. ecenditor is Jost. Vhe lstter species was

tlescribed by Sturt in 1848 as Afus conditer

from specimens observed and collected on his

A, MAHONEY

1844-6 Expedition ta Central Australia. The

original description was published in the

Narrative of the Expedition and Sturt did not

say where his matenal was deposited, Lt seems

likely however that at least one specimen,

Nlustrated by J, Gould and H. C, Richter ain

a plate’? accompanying Sturt’s description of

the species, would have gone into Gould’s col-

lection, and perhaps from there into a Museum

collection, A collection of mammals made by

the Expedition was presented to the British

Museum by Sturt in 1846, This collection is

noted by Thomas (1906b) and docs not con-

lain Specimens of Leporilius,

Specimens 1853.10,22-14 und 1853.10.22.45

were registered in the British Museum on Octo-

ber 22nd, 1853, and were acquired from

Gould. Labels attached ta them refer to 5.

Australia and F. Strange. The entries for them

in the Register mention neither 9 locality nor

Strange. S, Austruha could be an abbreviation

of either South Australia or Southern Australia

and F. Strange presumably is Frederick

Strange, a collector and dealer in natural his--

tory specimens who accompinied Sturt on

some of his carly surveys (but not the 1844-6

Expedition), and was an early settler in South

Australia and later, in the 1840's, w resident

of New South Wales (Whittell 1947), Gould

(1849) docs not mention Strange and South

Australia in bis account of L. conditer, Subse-

quently (Gould 1863) he gives only the

interior of New South Wales and Victoria as

Jocalitics for the species. Tt is possible that the

inscriptions. on the labels are interpretations. of

the orivins of the two apecimens based on

Gouid's uccount of L. apicalis. 1 they are not

interpretations, their significance is uncertain

singe the citation of S. Australia is ambiguous

and Strange might not be the collector ol the

specimens.

British Museum (N.H.) specimens

1853.10.22.14 and 1853.10.22,15 could have

been collected On Sturt’. 1844 6 Expedition

and might belony to the type serics of L. canedi-

tor, This is so even if they cume trom South

Australia, Sturt (1848, Vol, 1, pp, 120-L21)

referred in his accaunt of the Expedition’s pro-

gress wlong the Darling River in New South

Wales to an individual of L. conditor secured

by Mr Browne and te one, a male, obtained

by himself from uw mative. However, Sturt (Yol,

2, Appendix, p. 4) noted also that the last nest

?Gould’s name is printed on this plate and the s

Sturt,

species name Mus conditey is attAbuted mo him hy

nevertheless Sturt is the wuthor of the nume Mus conditar.

THOMAS’ “LECTOTYPE” OF LEPORILLUS APICALIS 103

CUORUREU REAR RARER RRR 4

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

muddy lagoon to the north of the Pine Forest

(N.S.W.), and the Expedition explored por-

tion of South Australia before reaching the

muddy lagoon.

Although the whereabouts of the type

material of L. apicalis and L. conditor is un-

known, there is no uncertainty about which

species of native rodents were named Hapalotis

apicalis and Mus conditor by Gould and Sturt

respectively, and neotype selections for them

are unwarranted.

Gould (1853b) commented in his account

of L. apicalis that an animal in spirits in the

104 J.

British Museum, presented by R. C. Gunn,

from Van Diemen’s Land, accords very closely

with it in the colouring of the fur and in the

rat-like form af the tail. He added that it is of

much smaller size than L, apicalis and in all

probabihty will prove to be a new species,

Gould’s listing in 1863 of Van Diemen’s Land

as a possible locality for L. apicalis could be

based on that material. Tasmanian rodent

specimens in the British Museum (N.H.) and

zttributable to Gunn are recorded in the Regis-

ter. The identities of these specimens and their

registration mumbers. are Rattuy ratty

(Linnaeus, 1758) (1837.6.10.56). Ratius nor-

vegicus (Berkenhout, 1769) (1838.1,15,17),

A. MAHONEY

Rattus latreelus (Gray, 1841)

1852.1.15.16, 9 1852.1.15.17),

fuscus Thomas, 1882 (1852,1.15.15) and

Pyeudomys —higginsi (Trouessart, 1897)

(1852.1.15.18). None of these are L. apicalis,

und evidence for the occurrence of this species

in Tasmania is lacking.

(1845.5.2.3,

Mastacomys

Acknowledgments

This study was carried out in the British

Museum (N.H.) by arrangement with Dr

G. B. Corbet of the Mammal Section. The

photographs were taken by Mr F, Greenaway,

British Museum (N.H.) Photographie Unit.

References

Goup, J. (1849).—"The mammals of Australia",

Pr 2, pL 8 and text (1863, vol. 3, pl 6 and

text), (J. Gould: London.)

Gourp, J. (1853a)—Remarks on the genus

Hapaloris, Proce. zool. Soe. Lond. \85\, 126-

127. [The approximate date of publication of

this work in 1853 is 29 April—see Prac. zool.

Soc. Lond, V7, 81 (1937).]

Goutp, F. (1853b),—*The mammals of Aus-

tralia". Pt 5, pl. 12 and text (1863, vol. 3, ph

2 and text). (J. Gould: London.) [The date

November Ist, 1853 is printed on the cover of

Pt 5 of Gould's work,]

Goutp, f. (1863), —"The mammals of Australia”,

Pr 13, Introduction, pp. xi-x) (1863, vol. 1,

Introduction, pp. xi-xl). (J. Gould: London.)

|The date May Ist, 1863 is printed on the

coyer of Pt 13..of Gould’s work. An uncor-

rected proof-sheet version of Pt 13, pp, xi-xl

of “The mammals of Australia” 1s included

in pp. 1-51 of « work entitled “An introdtuc-

tion to the mammals of Australia’ by J.

Gould and published in 1863. A copy of this

work in the Australian Museum library has

x date of presentation to Reverend John

Barlow F.R.S.. May 6th, 1863, inscribed on

the title page.]

Sturr, C. (1848) —"“Narrative of an expedition

into Central Australia, performed under the

wuthority of Her Majesty’s Government, dur-

ing the years 1844, 5, and #. Together with

a notice of the Province of South Australia,

im 1847." (Boone, London.) Vol. 1, xX. iv,

5-416 pp. Vol. 2, vi, 1-308, 1-92 pp. [The

final 27 pages, i.e. pp. 66-92, of Vol. 2 are

a botanical appendix by R. Brown.) [Although

this Work has the date 1849 printed on the

title pages, its date of publication is 1848.

Thus it is cited in a “List of New Books" in

The Athenaeum (Journal ef English and

foreign Nierainre, sctence, and the fine arts),

no. 110), December 2nd, 1848, p- 1207, col.

2, and in a list of “Publications Received” in

The Spectator, for the weck ending December

23rd, 1848, no. 1069, p. 1237, col. |. Further>

more, p. 1243, col. 2, of The Speetaror, for

the week ending December 23rd, (848. con-

tains a publisher's notice which states that

Sturts work is “Now ready”.]

Tate, G. H. H. (1951).—Results of the Arch-

bolt Expeditions. No. 65. The rodents of

Australia and New Guinea. Bull. Am. Mus.

nat. Hist. 97, 183-430.

THomas, ©. (1906a).—On the generic arrange-

ment of the Australian ruts hitherto referred

10 Conilurus, with remarks on the structure

and evolution of their molar cusps. Ann,

Mag. nat. Hist. (7) 17, 81-85.

Tromas, O. (1906b).—Mammuals. Pp. 3-66, Ja

“The history of the collections contained in

the Natural History Departments of the

British Mussum". Vol. 2. Separate historical

accounts of the several collections included

in the Department of Zoology. (British

Museum: London.)

Thomas, O. (1921a).—Notes on Australasian

rats, With au selection of Jectotypes of Aus-

tralasian Muridac. Ann Mag. nat, Hist. (9)

8, 425-433.

THomas, O, (1921b).—Notes on the species of

Notemys, the Australian jerbou-raw, Ann.

Mag. nat. Hist. (9) &, 536-541.

Tuomas, O. (1921c).—On three new Australian

rats, Ann, Mag. nat. Flist, (9) 8, 618-622.

TroucuTron, E. Le G, (1923).—A revision of the

rais of the genus Leporillas and the status of

Alapalatiy personata Krefft. Ree. Aust. Mus.

14, 23-41.

WHITTELL, H, M, (1947).—Frederick Strange. A

biography. Aust. Zool. 11, 96-114,

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,

two 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 (adamsi, bakeri, carnabyi, carteri, dedariensis, fraseriensis, globosus, goerlingi,

macmillani, minutus, obscurus, polli, robustus, smythi, tamminensis, 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 repperi Blackburn (of jansoni van de Poll).

Three names are newly synonymised (the synonym first): splendens van de Poll = mastersi

MacLeay; simplex Blackburn = masterst MacLeay; and strandi Obenberger = dilutipes van de Poll.

REVISION OF THE GENUS ASTRAEUS LAPORTE & GORY

(COLEOPTERA: BUPRESTIDAE)

by S. BARKER*

Summary

BARKER, S, (1975).—Revision of the genus Aytrucuy LaPorte & Gory (Coleoptera:

Buprestidae). Trans, R. Soe. 8. Aust, 99(3), 105-141, 30 August, 1975,

A revision is presented of the Australian Buprestid genus .dstraens LaPorte & Gory.

Following van de Poll (1889), but extending the status given by him to two separate groups

within the genus, two new. sub-geneéra ure 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 (adamsi, bakeri, carnabyi, carteri, dedariensis,

fraseriensis, globosus, goerlingi, macmillani, minatus, obscuras, polli, rohustus, smythi, tammi-

nensis, and watsomt) ure described. Two names, major Blackburn and mastersi MacLeay,

previously regarded as synonyms are revahdated. Two names previously regarded as synonyms

ure confirmed asx such. They are us follaws: mevricki Blackburn (of badeni van de Poll) and

teppert Blackburn (of janson’ van de Poll), Three names are newly synoaymised (the synonym

first): splendens van de Poll - mastersi MacLeay; simplex Blackburn = musterst Macleay;

and strand? Obenherger = dilutipes van de Poll.

Introduction

There wre 27 described species of the genus

Astvraeus from Australia and one from New

Caledonia. Originally established as. Asthraeuys

hy LaPorte & Gory (1837), the genus was

referred to by that name until Gemminger &

de Harold (1869) used Astraens, thus correct-

ting an incorrect transliteration from the

Greek. This hus been followed by all subse-

quent authors. Barker (1964) requested the

International Commission for Zoological

Nomenclature to validate the emendation of

Asthraeus to Axtracus and this was eventuilly

carried out (LC.Z.N, 1966, Opinion 795). The

genus was last revised by van de Poll (1889)

who gave excellent descriptions and illustra-:

lions Of all species known at that time, Pre-

sumably van de Poll never saw a live speci-

men of the genus, so wuld have been

unaware of their unique spring mechanism

involving whe release of the elytra from the

closed position. When the living beetle releases

the spring, the clytra flick open with such force

that the insect can be projected for up to

several metres, The effect is reminiscent of the

sternal spring of the Elatcridae excepr that the

same result is achieved i wv different way. It

has no doubt evolved as an escape mechanism,

One other feature has apparently not been

commented on in the literature and that is the

fact that most of the species exhibie classical

warning colouration of yellow markings on a

dark background. Whether any of the species

are noxious to predators or whether they are

Batesian mimics. has yet to be determined,

Since van de Poll’s excellent monograph,

further species have been described and more

discovered so that the genus is again in need

of revision. This poses some difficulty to an

Australian worker as most of the type speci-

mens are housed in overseas museums. Also,

through habitat destruction throughout Aus-

tralia, some of the species are not abundant

over their former range. and unfortunately the

genus ts very poorly represented in museums

and in most private collections in this country.

The adults of more than half of the species

occur on Casrarina sp., but some are also

associated with species of Jacksonia, Banksia,

Xanthorrhaea, Callitrix, Melaleuca, Acacia,

Daviesia, and possibly Dryandra and Haken,

Adults of the early emerging species can he

* Department of Zavlogy.. University of Adelaide, Adelaide, 8. Aust. 3000.

if 5,

collected in August and adults of same species

are présent as late as March, More than half

of the Known species ure found in the southern

half of Western Australia. The others occur

over the rest of the continent excluding the

desert woud northern-most tropical arcas, One

species occurs on New Caledonis. The penus

is unknown from Tasmania.

In this paper mule genitalia have not been

used to diagnose the species, but have been

used as one character to place the species

into related groups.

This revision is based mainly on collections

made by the author in New South Wales.

South Australia and Western Australia between

1962-1975, now housed in the South Austra-

lian Museum. The remaining specimens are

scallered throughout the collections listed

below. The abbreviutions used in the tear for

museum und private collections are as follows:

BA Collection of Mr E. E, Adams,

Edungalba, Queensland,

AM The Australian Museum, Sydnev

ANIC ‘The Australian National Insect Col-

lection, C.S.L.R.0., Canberra

BPBM ‘The Bermice P, Bishop Muscum.

Honolulu.

BM The British Museum {Natural His-

tory), London.

KC Collection of Mr and Mrs K, Car-

naby, Wilga, Weslern Australia.

JH Collection of Mrs J. Harslett.

Amiens, Queensland,

MM The MacLeay Museum. Swdney

(types on permanent loan to ANIC).

MNHN Muscum National d'Histoire

Naluretle, Paris,

NM The National Museum of Victoria.

Melbutrne.

NMP National Muscum. Prague,

OM The Queensland Museum, Brisbane.

SA The South Australian Museum, Ade-

laide.

WADA The Western Australian Department

of Agricullure, South Perth-

WAM The Western Australian Museum.

Perth.

The Australian Buprestidae have been

thvided imto six sub-families (Britton 1970),

Asrraeuy belongs tn the sub-family Bupres-

linge, the members of which have mouthparts

oot produced downwards to form a rostrum,

In the sub-tribe Birprestes the metathoracic

cpimera are completely exposed whereas in

the Anthaxtae they are Lotally or partially

vovered by the lateral extension of the whdo-

BARKER

men, Astraeusy i placed in the sub-tribe Bup-

Testes with nine other genera found in Aus-

tralia. Aytraewy anct Neobupresris both have an

indistinct scutellum, but Neobuprestis has an

exposed pygidium whereas in Astracus it is

covered hy the elytra. All of the other Austra-

fian members of the sub-tribe have a distinet

scutellum, They are: Nascivides, Nascio, Neo-

bubasies, Notohubasres, Bubastes, Buprestine,

Buprestodes and Euryspilus (Carter 1929).

Taxonomy

Genus ASTRABUS LaPorte & Gory

Asthracns LaPorte & Gory, 1837: 1, pl 1.

fig. 1. (Invalidated, Opinien 795.) Imhoff,

1856: 46. Lacordaire, 1857: 43. Saunders.

1868: 10, pl, 1, fig. 12,

Astraens: Gemminger & de Harold, 1869, 5:

1380. Suwunders, [871]: 43. Masters, 187):

124. Macleay, 1873: 239-240. Kerremans

IRRS: 136, van de Poll, 1886: 176-178, Mus-

ters. 1886: 71. van de Poll, 1889; 79-110, pls

2, 3, figs 1-19. Blackburn, 1890: 1256-1258,

1891: 496, van de Poll, 1892: 67-68. Kerre-

mans, 1992; 10]-102. Blackburn, 1892: 211-

713; 1894: Ol; 189S: 45-46. Kerremans,

L900; 295-296; 1902: 148-149, Fhuvel,

1904: 116, Carter, 1925: 224, fiz, 1. Oben-

berger. 1928: 204-205. Carter, 1929: 265,

282, 302, pl. 33, fig, 43, Obenberger, 1930:

265-367, Carter, 1933: 41. Obenberper,

1936; 133, Barker, 1964: 306-307. LC.Z.N..

19662 269-270.

Astreeys: Carter, 1929) 270.

Type species, Axtraeus. flavopicnia LaPorte

& Gory, 1837 (by monotypy).

Generic description

Head. Medium size or large; the surface

pitted; with a median longitudinal impressed

line on the basal half} clypeus. short, the apical

margin with a bow-shaped indentation; labrum

incised; antennal cavities small and rounded.

situated near the lower internal corner of the

eyes,

Antennac, The first segment longer than any

of the resi, rounder and thicker at the apex

than at the base: 2nd segment short and

obconic;, 3rd segment longer than the 2nd but

not as long as the first. alse: obconic; 4th seg-

ment slightly shorter than the 3rd, dorsoven-

trally flattened, the front edge projecting out-

wards to form a tooth; the remaining segments,

except the last, similar in structure to the 4th

and cach slighly smaller than the succeeding

segment; the lasL stgment flattened but not

REVISION OF BUPRESTID GENUS ASTRAEUS LAPORTE & GORY

toothed. Jn specses belonging to Astracuy

(sensu stricto) the antennae are sexually dimor-

phic, being longer in males than in females,

duc to the segments ufter the 3rd being of

approximately equal length, whereas in females

they become progressively shorter, In Asrraceus

{Depollus) the antennae do not show sexual

dimorphism and are short in both sexes.

Pronetum, Surface pitted; wider than long;

dorsally convex; rounded at the sides anil

darrower at the apex thun at the base; a deep

notch occurs on either side of the middle, at

the bare. In all but one species a small spindle-

shaped depression is present at the base of the

median lobe. | have called this structure the

basal crypt (Fig. 24).

Scutedlum. Invisible.

Elytra. Convex dorsally, each side with a

crescent-shaped lobe fitting over the notches

in the base of the pronotium; the apex of each

clyltron ends (with rare exceptions) in a sharp

point, thy sutural spine, usually formed by a

crescent-shaped lateral emargination which

forms on ibe inner edge a smaller lateral

tooth, usually called the marginal spine; if the

lateral emarginalien is abseni, sa is the mar-

ginal spine; below the shoulder the external

margin is diluted, sometimes the dilation is

rounded posteriorly, sometimes strongly pro-

jected backwards and angled forming a point,

the whole being folded jnwards and covering

part of the metastermmul coxae—this structure

is called’ the humeral fold (Fig, 1); punctate—

striute and In many cases also costate; there is

a curved lip on the anterior undersurface of

each elytron which fits over an apposite curved

projection on the mesotharax, together form-

ing the catch of a spring escape mechanism.

Undersurface. Variably pitted! antetior edge

of prosternum straight or crescent-shaped and

a large prostemmal process: mesesrernum short,

melasicinum approximately equal in length to

the prostemum; posterior margins of coxae

and abdominal sternites glabrous:

Legs. Femur slightly dorsoventrally flattened

but robust; tibia almost cylindrical but longer

than the femur; lst tarsal segment longer than

the remuining segments which become pto-

gressively shorter.

Body outline, Laterally teardrop-shaped, the

apex being constricted: dorsally convex; ven-

trally straight at the anterior end and curving

upwards at the end of the ahdnmen. rarely

concave.

14)7

Key to the sub-genera of Astraens

1. Lateral Jobes of pronotum projecting from the

elywal edge Jine; elytra striate-punctate but

never with longitudinal costae, humeral fold

rounded — _ . Astracus (Depollus)

-Laterul lobes of pronotunt confluent with the

elytral edge line; elytra suriate-punctate with

well or poorly developed longitudinal costae;

humeral fold angled (Fig. 1) ti

Asiraeus’ (sensu stricto)

Sub-genus Depollus sub-gen. nov.

Type species Astraeus abérruns van de Poll,

1886,

Head medium sized. Antennae short, Pro-

notum notched on both sides at the base, with

a Median lobe pointed and projecting, the

lateral [obes with the posterior margin curved

upwards and outwards, Elytra striate-punctate,

the edges of the striae never costate. The

suture ehds posteriorly in a short or very short

spine which may be blunt or pointed and

curved outwards, With no marginal spine or

in one species with a very short, blunt marginal

spine. The humeral fold is rotinded and poorly

developed, Only known to occur in Western

Australia.

Key to the species of Astraeus (Depollus)

[. Basal colour brown L pollt ca Tov,

|, Basal colour black , -2

2. Elytra with 4 longitudinal ted. vittae

, 3. lineatux van de Poll

2, Elytra withour tongitidinal red vittoe 3

3, Head, pronotum and undersurface very hairy

—

3, Head, pronduizt and undersurface nel very

” hairy elias: ' Lien

4. Pronotum with clumps ‘ot “hui “emerging

from depressions towards the sides; no mar-

ginal spine _. BR rabustis sp. nov,

4. Pronotun with hair emerging singly, not

clumped; small marginal apine

4. aberrans van de Poll

5, Head and pronorum with | deep punctures closely

packed together . damminensis sp, Nov.

5. Head and pronotum wiib shallow, veparated

punctures ,

6. Pronotum laterally inflated,

Numerous strigiform spots

6. multinatans von de Koll

6, Fronotum pot laterally inflated, elytral pat-

tern smal] und large spots and Jongrbucinal

vittae }

7. Sutural spine well developed and outcurving

7, ireegularis van de Poll

7. Sutura! spine poorly developed and not out-

curving ‘ if 8, deduriensiv sp, mey-

1. Astraeus (Depollus) polli sp. nov.

FIGS 2, 224A

elyiral nil

Types

Holotype: 4d, Wongan Mills, W. Aust. on

Casuarina campestris. GAt9T1, S. Barker,

SAM. T 20940.

108 5

—J mm—

Fig. |. Outline diagram of the left humeral fold

of 4 species of Astraeus, The head end of

each is to the top of the page and the Jeft

side is uppermost. Note that the specimens

figured are all females and that (a) and

(b) were sinailer specimens than (c) and

(d). Rach structure is categorised as in

the text, (a) A. mustersi MacLeay — well

developed, acutely angled. (bh) A.

samouelli Saunders ~— well developed,

angled. (c) A. intricatus Carter —

moderately developed, angled, (d) A.

carnabyi sp. noy. — poorly developed,

slightly angled

Allotype: %, Wongan Hills, W. Aust. on C.

cumpestrix, 9.1971, S. Barker, SAM, I!

20941.

Paratypes: 1 ¢ & 19,2 km W of Wongan Hills,

W. Aust, on C. cantpestris, 20.17.1973, §. Bar-

ker, EA, 5 3 & 4 9, Wongan Hills, W. Aust.

on C, campestris, 94.1971, S. Barker, ANIC

(| found 1 9). BM (1 3 & L 2), MNHN

(if & 1 2), SAM (25 & 1 9) 1 By Deduari,

- BARKER

W. Aust., 12.17.1948, J. A. Douglas, NM; 1 2,

Dedari, W. Aust., WADA; 2 o, South Tam-

min Flora Reserve, W. Aust. on C. campes-

tris, 64.1971 & 941.1971, §. Barker, WAM,

Colour. Shiny. Head brown with the following

yellow markings: two lunettes from the inner

corner of the eye, concave towards the base of

the antennae but not reaching the middle of

the head; behind euch of the former at the

base is a single small spot; « 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 blue reflections and the following yellow

markings: close to the anterior margin on each

side of the centre, but nol 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 is 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 anterior

margins of the elytra are black but elsewhere

brown. Undersurface brown: the mesosternum

and metasternum each haye lateral yellow

patches and each of the abdominal segments

have lateral yellow spots; the coxal plates each

haye a yellow spot near the centre but not

touching it; hairs silver, Legs: femur and tibia,

brown with blue reflections: tarsi dark brown

with blue reflections.

Shape and sculpture. Head with shallow punc-

tures, except for a median longitudinal glab-

rous line commencing between the eyes and

confluent with the basal impressed line; with

short sparse hairs. Pronotum with shallow

punctures in the middle becoming confluent ut

the sides, except for lwo small round glabrous

areas in the centre of each half; a short

impressed line projecting forwards from the

basal crypt; sides roiinded and narrowed from

base to apex. Elytra. with the intervals between

the striae slightly convex and transverscly

strigose with shallow punctures: parallel-sided

until after the middle then rounded to the

base; with a small sutural spine projecting

laterally. Undersurface with shallow punctures,

REVISION OF BUPRESTID GENUS ASTRAEUS LAPORTE & GORY 1b

denser at the sides thai in tlhe middie: with a

Jew short hairs.

Size. Males 10.5 + 0.

{18}, Females 115 4

{18).

Distribution, Western Australia,

General reertarks. Named after the late J. R. H.

Neervort van de Poll,

Specimens etatnineel Wo Aust.: Types; § a & 43

2.13 km N of Guomatliog, on Ceswarine campes-

res, BAIT, S. Barker; 1 ao & 1 2, Waddington,

on CL ciwnpesteis, S197), & Barkers 1 FR 2 4

c. campestris, 20.83.1972, 5, Bar

15 x 3.8 = 0.07 mm

0.27 x. 4.0 =0.09 mm

Dedufi, on c. :

herr 3d X39, 2 km W of Wongan Hills, an C

campestris, 201.1973, $8. Barker.

2, Astracus (Depollus) tamminensis sp. mov.

FIGS 3, 228

Type.

Holotype; ¢, South Tammin Flora Reserve. W,

Aust. on Casuarina campestris, 44.1971, S.

Barker, SAM, T 21438,

Caleur, Read and pronotum dull, elytra shiny.

Head black, bottoms of most of the punctures

yellow with the following yellow markings:

two small elongate spots each between the

eve and above the base of the antennae;

laterally a small] round spot between the eye

and the base of the head; ventrally all yellow,

almost continuous under the eye with the first

mentioned spot; antennae dark brown with

blue rellections, Pronotum black, bottoms of

most of the punctures yellow, with the follow-

ing yellow markings; a thin median longitudinal

line from base lo apex covering all hut the tip

of the median lobe, two longitudinal stripes on

each side of the first, curving inwards from the

base but ending before the apex, each capped

with a circular spot near the upea; a thicker

lateral stripe runs from the base to the apex:

underside with alternate irregular stripes of

black and yellow, the yellow predominating.

Elytra with allernate black and yellow tongi-

tudinal stripes broken irregularly, their breadth

conforming to the intervals between the striie:

predominantly black along the suture and the

outer margin, yellow in the middle, Under-

surface mainly yellow with irregular black

markings along the middle of euch side across

the outer margins of the abdominal sclecites

and at the apex, the black markings all with

brassy metallic gleams; hairs.silver. Leys dark

hrewn with blue reflections; each femur with

elongate yellow matkings on both surfaces,

Siape wand sculpture, Head with dense, uni-

fonmly deep purctures. Pronotum with «nr

lormly dense plinctures; more or less parallel-

sided but abruptly rounded at the apex. Elytra

with the intervals between the striae trans-

versely wrinkled, and deeply and densely pune

tured; parallel-sided from base to behind the

middle then uniformly rounded to the area

where marginal spines occur in the other

suh-genus, then straight sided to ihe apical

spine which is minute. Undersurtace closely

and evenly punctured; with y few shart buirs.

Size. Male 11.3 x 4.f mm (1).

Distriburion. Western Australia,

General remarks. ‘Vhis species shows closest

allinily with Astreeus polli.

Specimens examined. Type only.

3. Astra¢us (Depollus) robustus sp. nov,

FIGS 21, 22C

Types.

Holotype: 3. Payne's Find, W Aust, on

Casuarina divisiane, '7a~.t970, S. Barker,

SAM, I 20842.

Allolype: Y Payne's Find, W. Ausl, on

C. dielsiana, 17,ix,1970, 3, Barker, SAM, I

20943.

Paratypes: | @, Puyne’s Find, W. Aust. on C.

dielsiana, I7AXASTO, S. Bake’, ANIC: 1 9,

Payne's Find, W. Aust, on C, ¢divtsiana,

17,1x.1970, 4, Barker, BM; 1 cf & 1G, 385

km ulony Payne’s Find Rup W. Aust. on CC.

dielsiama., V7ix.1970, S$. Barker, MNHWN; |

3, Payne’s Find, W. Aust. of ©. dielsiana,

(7.ix.1970, S. Barker, WAM.

Colour. Shiny. Head, pronolum and clytra

black wtih purple reflections. Antennae black

with blue and purple reflections, Elyira with

the following yellow markings: o longitudinal

spot in the 2nd outermost inierstice near the

shoulder and another just before the middle; 3

similu’ brit larger spot lies on the Sth outers

most interstice after the middie but not reach-

ing the apex; a large round spot before the

middle with the width of the 3rd-6th inter-

stice from the suture; a variable number of

amall spots at the base of the elyura in the

middle, the maximum numher is four per side

but they are absent in some specimens. Under-

suffuce black with purple reflections; hairs

silver. Legs brown: the joint between the femur

and tibia black on the upper surface; Ist tarsal

scament brown at the base and black at the

apex as are the remainder,

Shape and sevlprare. Head with dense shullow

punctures; with wu glabrous longitudinal medinn

ling between the eyes confluent with the basal

impressed line; bairy, Pronotum sparsely and

shallowly punctured in the middle where ihe

depressions are small but greatly increase m

size Jutevully where the foit-depressed areas

5. BARKER

“a

fat

Hed |Aiv

e TAU

RS 1) Ai

My ‘ is

Fig. 4. Astraeus dedariensiy sp. nav,

sirdeus fasimincusigy 8p. Ov.

ig, 3. A

Astraciis palli sp, nov,

ig, 2

REVISION OF BUPRESTID GENUS ASTRAEUS LAPORTE & GORY Lit

are glabrous; sides gradually rounded from

base to apex; hairy, with hairs emerging singly

in the middle and in clumps from the lateral

depressions. Blytra with intervals between the

striae slightly convex; more or Iscs_ parallel-

sided from base to just after the middle, then

rounded gradually to the apex, with sharp

sulural spines; hairy. Undersurface covered

with punctures, shallow in the middle, deeper

und denser at the sides; hairy, the hairs on the

Presternal pracess and prosternuni noticeably

longer than clsewhere.

Size. Males 15,9 & 0,31 x 53.6 — 0.16 mm (4).

Females 17.0 + 0.90 x 6.0 = 0.25 mm (3},

Distebution, Western Australia.

Specimerty examined. Types anly.

4, Astraens (Depollus) aberrans van de Poll,

1RBG: 176; IRSY: B4, 90, B1, pl. 2, fig.

4, 44. Kerremans. 1892; 101; 19027148,

Carter, 1929: 282. Ohenherger, 1930: 365.

Axtraces aherrans var. pieticnliis van de Poll,

1889-91, Kerremans, 1892;101, 1902;148, Cur-

ter, 1929:282, Obenberger. 1930:365,

FIG, 22D

Yvpe. MNHWN (not seen hy author),

Colour, Shiny. Head, pronetum and elytra

hluck with purple refiections, antennae bluck

with blue and purple reflections, Elytra with

the Follawing yellow markings: a small irrégue

lar number of vlongate markings each being

restricted to the width of one interval between

atriac and occurringly mainly in three areas:

near ihe sulurs; in the middle of the elytra;

text fo Lhe yoterior part of the outer margin:

Undersurface metillic purple, the lateral mart-

gi of the Ist, 2nd and 3rd abdominal seg-

ments each with u yellow spot which is vartable

in size and ubsent from the 3rd segment in

some specimens; hairs silver, Legs dark with

blue and purple reflections,

Shepe and sctlprere. Head deeply punctured,

in parts in the middle less so than at the sides.

the areas involved glubrous; with a longitudinal

glabrous fine between the eyes merging with

the basal impressed lime: very hairy. Prono-

tum with large deep punctures dispersed but

mote so in the middle with the elevated parts

glabrous; mostly purallel-sided but abruptly

rounded and narrowed at the apex: anterior

margin projecting forwards in the middle: very

hairy at the sides, the hairs emerging singly.

not in clumps, Elvira with the intervals

herween the striae convex und slightly

wrinkled, each with two longitudinal rows of

small halts) parallel-sided, rounded after the

middle ta the small marginal spine; sulurial

spine stall. Undersurface evenly and shallowly

punctured; hairy.

Size. Males 14.0 + 0.2 x 4,9 >= 0,09 mm

(25). Females 16.6 = 0.43 4 59 = 0.17 mm

(11).

Distribution, Western Australia,

General remarks. This species shows closest

affinity with Asiraeus rohusius.

Specbrens examined. W, Aust- 1 4, Carnamah,

30.14.1965, B. Baker; 12 # & 6 G S km W of

Payne’s Find on Casuarina sp., 07.ix,1970, 8, Rar

ker; 2 3, 106 km 8 of Payne’s Find on Casicerine

corniculata, 18,ix, 1970, 8. Barker; 2.9, Wialki on

C. corriculata, 19.ix.1970, S. Rarker; 1 oy Wialkt

on Casuarina campestris, 21.ix.1970, 8. Barker: |

3, South Tamnun Flora Reserve on C. carmpestris,

30.ix.1970, 8. Barker; 3 &@ & 4 9, South Tammin

Flora Reserve on ©, campestris, 8.xi.1970, 5,

Barker; 3 do & 1 9, Quairading on C. cayipestets,

TXLEITO, S. Barker,

5, Astraeus (Depollus) lineatus van de Poll.

1889; 84, 87-89, pl, 2, figs 2, 2a, Kerre-

manos, 1892: 102; 1902; 149. Carter,

1929) 282. Ohenberger, 1930; 366.

FIG, 22

Type, Holotype, MNHWN (net seen by the author}

Colour, Head snd pronetym dull, elytra shiny.

Head black with brenze-green reflections i

the middle at the base, the Test with purple

reflections; with or without 4 red spot under-

neath ench eye; antennae black with purple

refiections. Pronotum black with bronze-green

reflections in the middle, the rest with purple

feflectrons: a thin lateral red stripe en each

side, with or without a small red spot on each

side of the median Jobe neur the base, Elytra

black with blue reflections and the following

red markings: 'wo broad vittae, the first near

the margin, but hot touching it, running from

the shoulder to the preapical area} the second

in the middle commencing newt the front cuge

running parulle] ta the suture, but not touch-

ing it, lo the preapi¢al area. Each of the vittae

may be entire or broken into two or a number

of elongate marks, Undersurfauce metallic

purple with lateral red marks on the pro-

sternumt, 3rd coxal plate and lst, 2nd and 3rd

abdominal segments, The prestertal and 3rd

abdominal spols are absent from some speci-

mens} hairs silver. Leys: femur and tibia

metallic purple; tarsi brown,

Shape and sculpture. Head evenly punctured

with a longitudinal glahrous line between the

eyes! hairy. Pronotum evenly punctured, the

punctures deeper at the sides than in the

112

middie: sides narrowed and alightly rownded

from the bitse to near the apex where they are

more noticcably rounded: the Front edge pro-

jecling furwards in the middle; hairs longer

and more dense at the sides than in the

middle. Elytra with the ttervals between the

struc conyex with shallow punctures and

faintly transversely strygose; laterally shghtly.

concaye from the shoulders until after the

middle, then rwunded to the small sutural

spine; hairy. Undersurface: prosternum with

large punctures. the resr with small shallow

punctures, fess.dense in the muddle; hairy.

Sizes, Males 110 © O48 x 3.8 + 06.22 mm

(3). Females 13.0 + 0.29 » 4.6 + 041 mm

(6),

Oistevuition. Western Australia.

Specimens examined. W. Aust.: Lay 5 Am W of

Payne's Find on Cusuarine: sp, 17.ix.1970, 8, Bur-

ker; 1 oo, South Tatnmin Flora Reserve on

Casuarina campestris, 30,1%,1970, 8. Barker: 2 5,

Quairading on C. campesrris, 7.x1,1970, 8S. Barker:

1 of & 2 9, South Tammin Flora Reserve on €’,

campestrty, BXL1970, S. Barker; 2 2, 13 km N of

Goomalling on C. campestris, 94.1971, &, Barker,

6 Astraeus (Depullas) multinotatus van de

Poll, 1889; 84, 89, 90, pl. 2, fig. 3, 3a.

Kerremans, 1892; 102; 1902 149. Carter.

1929; 282. Obenherger, 1930. 366.

FIG. 22F

Type. Holotype. MNHWN (not seen by the author).

Colore. Head and pronetim dull, elytra shiny.

Upper surface and antennae black with blue

reflections. The whole upper surface except the

witennae speckled with small yellow spats,

Undersurface black with yellow spots larger

than on the upper surface; hairs silver, Legs

dark brown; cach femur with a single elomeate

spol; the tarsi black,

SMeape and sculpture, Head with shallow even

punctures and a small glabrous spot between

the eyes. Pronotum evenly punctured, the

punctures larger at the sides than in the

muddle; slightly inflated after the middle,

rounding and narrowing, quickly to the apex:

the front edge projects forwards in the middle,

Elytra with intervals between the striae con-

vex, punctured and wrinkled; laterally slightly

concave, rounding well after the midkle to the

svlural spines which are turned out. Under-

surface cvenly punctured, but the punctures

larger at the sides than in the middle, the

differences pronounced on the thorax. Apart

frum ihe legs, apical edges of the abdomen and

pronotiim, the body fs devoid of hair.

S&S. KARKER

Size. Males 12.3 + 0.73 x 4.9 = O34 mm (7),

Females 15.) + 0.95 x 6.0 £0.35 mim (6).

Distribution. Western Australia,

Specimens examined, W, Aust. 1 @y Leonora,

AL OM, Brown, coe er WAM, 34-51: | o an

km § uf Coolgardie, J a6. 1937, ALM. Dawgs,

WAM, 37-722; 1 2 & 1 9, Tarvin Ruck, 24.1940,

AM, Duiglas. WAM, 40-32, 40-31: 2 o, Wat-

ning, 20,ix.1950, R. P, McMillen, WAM, 73-384,

73-386, 2 a & L G& Watning, 26.xli- 1950, R. f

MeMillan, WAM. 73-385, T3388; 1¢ &

WAM, 73-375/7: 1 3, Red Gum Pass, Sistine

he on Casaeina sp., 26.1971 BL & K, Cor

nahy

7. Astraeus (Depollas) irregularis yan de Poll,

I889; 84. 86, 87, pl, 2, fig. 1, la. Kerre-

mans, E892; [02; 1902; 149. Carter,

1929: 232. Obenberger, 1930: 366,

FIG, 22G

type, Holotype, MNHWN (not seen by wuthor.)

Colour, Head and thorax dull, elytra shiny.

Male. Head black with blue reflections. with

the Following yellow markings: a continuotts

hand ander the lower half of the eve; un elon-

gate spol in the centre of the head which may

he broken inta several spots; two elongate spots

lateral und basal to the central spot, each of

which may be broker into two spots: a small

spot. on either side of the gular, Except for the

central spot, all other spots may be recuced

or ubsent. Aniennge dark brown with bluc

reflections, Pronotum hlack with blue refec-

tions and the fullowing yellow markings: a spot

on the margin of the apical edge on each side

of the middle; 4 spot on each side below the

first, towards the busel two spots along the

sides, a small one at the apex and an clongate

larger one ut the huse—the first may be absent

and the second broken into two spots. All of

these spots muy be reduced and sore or all of

them absent. lytra black with blue reflections,

each elytron with the follawing yellow muark-

ings: the basic pattern is of three spots in the

middle but not touching the suwlure; dhe lirst

is 4 small elongate spot at the base: the second

4 large spol before the midule; the third » vitta

after the middle and extending to the pre-

upieul area; there are also lwo small clongate

spots at the margin, one at the shoulder oxtend-

ing in the region of the humeral fold and a

smaller one just behind it which is sometinies

absent; an wregnlar number of small spots are

present along the busal edge between the villa

und the suture, Undersurface dark brown with

bie reflections: with yellow spots at the base

of each leg and single laleral spots um the pro-

REVISION OF BUPRESTID GENUS ASTRAREUS LAPORTE & GORY

sternum, third coxal plate aud abdominal sez-

ments 1—4+, |—3 or |—2. decreasing in size fram

Airst ro lasts hairs silver. Lees lark brown with

bie reflections.

Female, Head. pronotum and elytra basically

similar to the male except that the yellow

markings aré larger and the irregular spots are

more numerous. Undersurface and legs as in

the male except that lateral spats are present

on all the abdominal segments.

Shape and sculpture. Head broad; with shallow

plinctures larger it the sides than in the middle

and a longitudimyl glabrous ling between the

eyes. Pronotum with shallow punctures latger

at the sides than in the middle; the lateral mar

gins rounded and narrowed gently at the hase,

more abruptly at the apex. Elytra with the

intervals betweeti the striae slightly con-

vex, with a shallow row of punctures and

faintly transversely wrinkled, purallel-sided

until after the middle, then rounded to the

sutural spine which is well developed and

turned out, Undersurface more closely punc-

tured at the sideés than in the middle, with a

few short hairs.

Size. Males 11.2 + 0,24 x 4.) = 1.08 mm

(11). Females 13.5 + 0.22 x 4.7 + 0.08 mm

(20),

Distribution. Wester Ab pa!

Specimens examined. Wo Aust: 32 fd & § 9. Coy

nells on Casuarina sp, 24x. inert &. Barker; 1

cd &2 9, Gosnells on Casuarina sp., 27,xii, 1967.

S. Harker; 2 3 & 3 9, Red Hill Rd near Midland

Junction on Casuarina sp., 44.1968, 8. Barker; 2

9. 77 km along main York Rd on Caswarine sp.,

Si.1968, S. Barker; 3 do & 4 9, South Tammin

Flora Reserve on Casuarina heegeliaqna, 64,1971,

S, Barker; | ® & 22, t42 kin E of Norseman on

C. huegeliana, 19.x1.1972, §. Barker; 1 3, 38 km

W of Tammin on York-Tammin Rd on C.

Aueveliqna, 23.0i,1972, S, Barker; 1 ¢ & 1 3,3

kn Eo Gosnells on C. huegeliana, IT. 1972,

A, Barker,

8, Astraeus (Depollus) deduriensis sp. nov.

FIGS 4, 22H

Vy pes.

Holotype: ¢'. Dedyrl, W, Aust, A, W. Brown,

WAM, 71-1779,

Allotype: 9, Dedari. Wy, Aust, Ay He Brown

WAM, 71-1778.

Puritypes: 2 9, Dedori, W. Aust, WAM,

40-1207, 40-1208; 1 % Borden, W. Aust.,

SAM! | 2, Dedari, W. Aust, ANIC; 1 7%,

Dedari, W. Aust, on Canimrina Larnivulata,

214.1936, 77. HW Brown, MINHN,

Colour, Shinw.. Head black with purple reflec-

tons and the following yellow markings; an

ovoid spot touching the underside of cach cye

\a3

and pointing downwards but not touching the

mouth; a single circular spot in the middle.

Antennae brown with blue and purple reflec-

tions except fur the first segment Which is pre-

dominantly black, Pronotum black with purple

and blue reflections and the following yellow

markings: a small ovoid spot an each side at

the apex; 4 larger ovoid spot in the middle of

each side at the base, ‘slightly angled outwards

on the apical side; laterally there is a line on

tach xide from the apex to the hase where it is

extended by the flaring outwards of the lateral

murgin of each lobe exposing the yellow inner

surface, Efytra black with purple and blue

fellections and the following yellow markings

which do not form a symmetrical pattern on

each side: a large basal spot; behind and in

line a large elongate spot, commencing before

the middle and ending al the middle, close te

the suture but not touching it; am elongate spot

commencing after the middle and running te

the apex, close to the suture but not touching

it; an elongate spot at the shoulder, not touch-

ing the outer margin but close to it) behind the

last a much smaller spot not touch

ing the margin: after the middle there are

several elongute spots ettending almost to the

apex but not tariching the outce margin; several

small eccentric oval spots. Undersurtace black

with purple reflections; the first three

abdominal segments have lateral vellow spots

which progressively diminish im size frem in

front backwards; hairs silver. Leys: meso-

sternal and metastcrnal coxae with yellow spots

on the outer margins; the rest brown with

purple reflections; ihe upper surfaces of the

tarsi are black.

Shape and sculpture. Head with shallow puoc-

tures and without hair. Pronotum with shallow

punctures; a short median impressed linc pro-

jects forwards from the basal crypt; parallel:

sided at the base, abruptly rounded at the

apex; with short hair on the anterior edge only.

Elytra with the intervals between the striae

slightly convex and transversely strigose:

parallel-sided unt? after the middle then

rounding off to the apex; each ending in #

thick shart spine projecting backwards; no

marginal spine but 4 small mound is present in

the region where they occur in other species.

Undersurface with shallow punctures: with a

few short hairs,

Size, Males 11,2 44.2 mm (1). Females 17.0

+ 0,15 x 50 = 0,1 mm (6)-

Disieifviloa. Western Australla-

114 S. BARKER

General remarks, This species shows closest

affinity with Astraeus frregularts.

Specimens examined, Types only

Sab-genus Astraeus (sensu stricto) sab-gen,

nov.

Type species Astracus flavapictus LaPorte &

Gory, 1837.

Description. Head medium sized; with ar with-

wut a median longitudinal raised ridge between

the eyes, the median kee] (Fig. 25). Antennae

long. Pronotum deeply notched on both sides

at the base, with the median lobe and the two

lateral lobes pointed, Elytra striate-punctate

and/or costute, the Guter edge of the intervals

between the striac costate, either down the

entre length or towards the apex only. The

suture ends posteriorly in a spine and in all

but three species these are sharp and curved

outwards, The marginal spines are smaller thon

the sutural spines, The humeral fold is variably

developed and angled (Fig, 1).

Key to the species of Astraeus (sensu stricto)

|. Head with a median keel jcc ee 2

1. Head without a median keel — 0, -.,-..- 15

2. Hairs silver 3

2. Hairs yellow . = 14

3. Part or all of the anterior Sndenmi Paces red brown 4

3. None of the anterior undersurface red-brown 6

4. Gular, prasternum, meso~- und mibasternitit, “Ind and 3rd coxue and Ist abdominal sczment

red-brown ae Pia eth .- * hakert sp, nov,

4. Proslernum and coxae ‘red-brown ryt cm ictaeet 10, 7M hake sp. nov,

4. A red-brown ‘urea on either side of the ‘prosternal Process. 5

$. Usually shorter than 7.5 mm; humeral fold well developed and acutely angled .

I. frasevi iensis sp. nov.

6. Most of the leg testaceous - .

6. Tips of the tibia and Ist tarsal segment tESIACEOUS ne

6, None of the leg testaceaus _

for the femur

Usually longer than 7.5 mm: humeral fold moderately developed and angled ,

15. obscures: sp. noy-

7. Ist and 2nd legs testaceous exent 54 the. citer marein 5 OF the e formu, ard Re leataceous exce 3

7. Tibia, 1st and 2nd tarsal seyments AESLACEQUS os. ccs ccecescsssevsessveseevsereantvveven

8. Usually shorter than 7 mm

8. Usually longer than 7 mm .

_ 19. dilutipes van de Poll

coo VEL sontythe ep mwy.

2. pygirraens van de Poll

9 Humeral fold well developed, acutely "ampied” 17. mastersi Macleay

4 Humeral fold well developed and angled... eg ON Evita ae cabbie bitte Cathie 18. semnouelli Saunders

10. Humeral fold moderately developed and angled. oa . wie II

10. Humeral fold poorly developed, slightly angléd riiveot 1air Levee a vovtowtonoe 13

tl. Head and pronotum green and | coppery: “purple

11. Head and pronotum black -

12. Broad and rounded SPECIES ooo seers herp ssl he ldaldec ae ll eta

12. Elongate species

; 21, aulsibeites Carter

ps gee |

16. plabositi Sp. nav.

35. watsoni sp. nov.

13. Black species; elytral mar kines. a number of yellow Spots. ; r 22. crasyuy van de Poll

13. Blue species; elytral markings two yellow fascia , brome anes 25, fratercylas van de Polt

14, Elytral markings three yellow fascia and red areas 23. mujor Blackburn

(4. Elytral markings two. yellow fascia withont red areas... ....... veo 24. navarchis (Thomson)

15, Body clongate and cylindrical - 16

15. Body not elongate and cylindrical ., 17

16. Pronotum conically elevated in the “rhiidale

16, Pronotum not conically clevated in the middle

17. Sutiral spine with a rounded internal edge .....

17, Suturai spine with a straight internal cdge - _,,,.

18. Legs a red-brown colour oo. occeecicree seen iiiies

18, Legs not a red-brown colour _

19, Elytral markings spots and fascia ......,

19. Elytral markings two vitlae on etich elytron ale a eae

26. proshiarenp fous win de Poll

I elangats van de Poll

40, maciittiet sp. NOV.

28. vittatus van de Pall

20. Head, pronotum and legs metallic brown or brome = 5 29. flavonictus LaPorte & Gory

20. Head, pronotum and lees not metallic brown or bYONZE 2 ie ee BE

21, Humeral fold well developed and angled — - 22

21. Humeral fold poorly developed, slighlly angled .. ee . eld 23

22. Head black or coppery-purple; undersurface ceppery- purple ae 20. ‘alarnat Sp. Tey.

22, Head blue or green; undersurface blue-green

14, simulator van de Pall

REVISION OF BUPRESTID GENUS ASTRAEU'S LAPORTE & GORY 115

23, Basal spot expanded along tis front edge of the el¥lON jc. cen

23. Basal spot not expanded along the front margin pf the elytron

o 31. carnaby? sp. nov.

24, BElytral markings elthe: two spots and iwo [usclu ov the firet fascia ttiely ‘be broken muking

four spots and a fuscia on each elytron .

32. badent van de Poll

24, Flytral markings either six spots and a sascit | or thet Exch b mak be broker Makiie eight

spols On cach elytroo ..

24. Elytral markings 7 spots on each elyiron _

, 33. jansant van de Poll

25_ Pronotum parallel-sided from buse to the middle then sfropely. rounded and narfowed ta the

apex; dorsally convex in lateral profile

34. oherthuri van de Poll

24, Pronotum gradually roumided at the sides and natrowed from ‘base to apex; shrsal} flattened in

lateral profile ...,......

26. Head slightly akcapnted beiween the eves .

26, Head deeply excavated between the eyes .

27. Mead excavuted mainly at the base; propotom laterally inflated with an 1 oval patch of henti>

onal cells in the Middle o...0.. Corfe cifpo regy free

36, carter! sp, Tov,

_ 37. everlingt sp. nov.

38. svyanens Kerremans

27, Mead excavated mainly al the apex; pronotum not laterally inflated and without hexagonal cells

9. Astracus (Astracus) bakeri sp. nov.

FIGS 5, 227

Fynes.

Holotype: od, (Dryandra, W. Aust. on

Casyarina huegeliana, §&.xii 1970, §, Barker,

SAM, 1 20045,

Allotype: 9, Dryandra, W. Aust. on Cy Jiuege-

lian, 8.xi1,1970, §.. Barker, SAM, 1 20946,

Paratypes: 1d & 1 9, Dryandra, W. Aust. on C.

dimegeliana, 1211973, S. Barker, EA: 2 2.

Toompup, W. Aust. on Casnarina sp., F. &

K, Carriethy, KC: 2 9, Dryandra, W. Aust. on

canarias sp., 2.11968, §. Barker, SAM; 4

& 4 9, Dryandra, W, Anst. on Cy huege-

ty &xli1970, 8. Barker, ANIC (1 J & |

2), BM (1 f & 1 2), MNHN (1 @ & 1 9),

WAM (1 a & 1),

Colon, Shiny, Head and. pronotum black with

violet ur green reflections; antennae black with

metallic bie reflections. tips of first and

second segments brown, Elytra black with

violet reflections, each with the following

yellow markings: a spot at the base; a fascia

above the middle, covering the humeral fold

and ending near the suture but not touching it,

broken into spots in some specimens; wu lurge

spot below the middle, Undersurface: gular,

Ppra- meso- and metasternum, second and third

coxae and median anterior part of the first

abdominal scgment all reddish-brown with

light blue reflections on the Jateral part of

the above; remaining abdominal segments

black with metallic blue reflections: hairs sil-

ver. Legs brown with metallic blue reflections

on the upper surface, the end of the first tarsal

segment and all of the rernaining tarsal segs

ments metallic blue,

Shape and sevlprere, Head evenly punctured;

small median keel; hairy. Pronotum deeply

And evenly punctured. with a mediin longi-

ludiNal impressed line at ihe apex; median

39. caledonicus Fauyel

lobe without 4 basal crypt; sides rounded and

narrowed from base to apex; haiey, Elytra

coslaic, the Intervals between flat with a pro-

minent row of punctures; at the sides gently

rounded from the base toa just before the

middle then rounded and narrowed to the

strong marginal spine: sutural spine robuscy

humeral fotd well developed and angled.

Undersurface sparsely punctate in the middie,

more closely at the sides; covered with fine

hair which is denser at the sides than in the

middle,

Size. Males 7.5 = 0.07 x 3.0 + 0,03 mm (44).

Femaics 7,9 = 0.12 « 3.2 + 6.06 mm (23).

Distribution, Western Australia,

General remarks. Named after Dr F. H. Uther

Baker,

Specimens examined, W, Aust: Types; 18 d & 8

2 Dryandra. on Casuarina Anegeliana, 8.x. 1970,

S. Barker; 6 f & 1-9, 13 km F of Ongerup, onc.

huegeliant, 114.1973, S, Barker; 8 4&3 9, Dry-

andra, on C. huegcliana, $2.4.1973, &. Berkers 1

Ponier rockhole 70 km $8 of Balladonia, on €-

huegeliana. S.xi.1974, S. Barkers 3° &. Juranda

tockhole 104 km § of Balladanfa, oa C, Aaege-

fiena, 9.x1i.1974, S. Barker; 1d. 15 km §& of

Balladonin on Casuarina humilis. SoxiL9T4. 8.

Barker: L od SW valley of Me Raped, on ¢

fureceliana, UW.xii.1974, 8. Harker} 1 2°, I km S of

Mt Ragged an Teraclite Bay road, on C. fusrariis,

1Ch, xti. 1974, S. Barker.

10, Astrseus (Astracus) minutus sp. nov.

_ FIGS 6, 22)

Types.

Holotype: 3, South Tammin Flora Reserve, W.

Aust. on Casuarina huevekana, 6.4, |O7E, x,

Barker, SAM, T 20459,

Allotype; 9, South Tammin Flora Reserve, W.

Aust. on C. huexeliona, 23-21-1972, §. Bar-

ker, SAM, T 204460.

Paratypes! 2 4', Soulh Tamunin Flora Reserve,

W. Aust on C. fwegeliana, 23,xii.1972,, 3,

Forkes, ANIC (1), MNHN (1); 1 3, South

S. BARKER

{hee

raat mop

mud abi ==

5mm

}

1énsIS Sp. NOV.

Fig. 7. Astraens fraser

(us Sp. nov.

eus mind

Fig. 6. Astra

ert sp. nay.

Asiracus bak

. 35,

REVISION OF BUPRESTID GENUS 4STRAEUS LAPORTE & GORY

Tammin Flora Reserve, W, Aust, om, C,

Aeeseliana. 6.LI9TL 8. Barker. gehts ! ¢,

Sowth Tammin Tlard Reserve. W. Aust, on

CO. haegeliana, 9i.1974, 8, Batae BM; 2 2,

South Tammin Flora Reserve, W. Aust. on

C, cnmnpestets, BUIITS, SL Barker, WAM.

Coley, Shiny. Head black with purple reflec-

trans; antennae black with blue reflections

except for the tips of the first and second seg-

ments which are brown. Pronotum black with

purple reflections in the mdidle, red-brown at

the sides. Flytra hlack with purple reflections:

each with the following yellow markings: a

large basal spot; a fascia before the middic

covering the humeral fold but not. reaching the

suture; a large median spot after the middle.

Undersurface: prosternum anc coxal plates

red-brown, the rest black with metallic purple

reflecuons; hiirs silver. Legs; upper fernur red-

brown, the rest ol the femur and upper part

of the tibia metallic purple; apical part of the

tibia and first tarsal segment testaceous)

second, third and fourth tarsal segments. black

with blue reflections.

Shape and yenlpture. Head broad; deeply and

evenly punctured: with u median keel; hairy,

Pronatum deeply and evenly punctured, with

a short impressed lite projecting Torwarls

From the basal crypt; sides gradually rounded

from base ta apex; hairy. Elytra costate

the miervals flat cach with a row of very

Shallow punctures; laterally tapering gradirally

from base until after the middle, then rounded

to the well-developed marginal spine; a very

elongate crescentic cmargination between the

marginal xpine and the well developed sutural

spine; humeral feld well developed and

acutely angled, Undersurface evenly punc

tured; hairy,

Size. Males 3.7 © 0.22 & 2.3 + 0.10 mm (7}.

Females 6.4 & 2.7 mm (1).

Oistribulian. Western Australia,

Crneral remarks. The smallest species known

at this time.

Speeimens examined. Types only,

Ll. Astraeus (Astraeus) fraseriensis sp. nov.

FIGS 7. 22K. 24. 25

Types,

Holotype: 3. Fraser Range, W. Ast, on

Casuarina haégeloua, 19xil,1972, 8. Barker,

SAM, 1 20051,

Allolype: 2) Fraser Range, W, Aust on C.

Hinegelona, }9xii1N72. 8. Barker, SAM, 1

20952.

Paratypes: 2 ¢ & 2, Dryandra, W. Avs. on

C. hinesellapies 12,1.1973, S. Barker. EA (td

& 12), MNHN (1 Pk 1 222 G42 9,

Fraser Range, W, Aust. on ¢' hwepeliana

117

19,011.1972, 5, Barker, ANIC (| ¢ & Ll @),

BM (1 3&1 2)5 1 A, Lake Risin, W. Aust,

911972, E. & &. Carnaby, KC: 3 fy Fyre

Highway 142 km E of Norseman, We Aust.

on CC. fueweliana, 19.xi0.1972, 8. Barker,

SAM; 2 ¢ & 9, Somh Tammin Flara

Reserve, W. Aust. on C. Auegeliana, 64.1971,

§. Barker, WAM.

Colour, Shiny. Head. antennae and pronotum

black with vieler and blue refiections. Elytra

black with purple refiections, each with the

followitiy yellow markings: a large basal spot

in the middle; a fascia covering the humersl

fold. running towaruls the suture bul not touch

ing it, und slightly concave towards the base? a

large spot after the middle nol touching the

suture or the margin: sometimes there is a

preapical spor. Undersurface metallic purple

except for a red-brown patch on either side

of the prosternal process at the anterior edge

vf! the prosternum; hairs silver. Legs: femur

metallic purple. tibia and first tarsal seement

predominantly testacequs, second. third and

fourth tarsal segments dark with blue reficc-

tions.

Shape and sculprare. Head closely and evenly

punctured: with w median keels hairy. Prono-

tum closely and evenly punctured with a

slightly excavated median longitudinal line

projecting forwards from the basal crypt;

rounded ut the sides und narrowed Frum hase

Io apex; hairy, Elytra costate, the intervals

between flat, cach with a row of punctures;

parallel-sided until after the middle then gently

rouruled anil narrowed lo the marginal spine;

both spines well developed; humeral fold well

developed and ucutely angled, Undersurface

shallowly punctured in the middle. punctures

deeper at the sides: hairy.

Size. Males 6.7 = 0.05 x 2.6 = 0.02 mm (72).

Females 7.0 © 0.08 ¥ 2.8 = 0,04 mm (28),

Distribution. Western Australis.

General remarks. The three species 4 yraeus

hakeri, Astraeus minurus and Astravus fraser

iensis show a ose affinity to each other,

Specimens examined. W. Aust.; Types: | 2, 88 km

W of Yammin on York—Tanumin Rd, on

Cusuarindg haegeliuna, 23.20.1972, S. Barker; 1 2

$$ kot W of Turnmin on York-Tammin. Rd, on C.

huegeliane, 91.1974, S. Barker, 2 3 South Tam-

min Flora Roserve, an C. buegeliana, 23.xii1,1972,

§& Barker; 4d & 2 9, South Tammin Flora

Reserve, on C. Awegefiune, 9.1.1974, §. Barker;

31 ¢ & 6 Y, Corugina rockhole 66 km 'S of Balla-

donia, on ©. huegeliana, S.xil1974, S$. Barkers

1d & 13 F, Ponier rockhole 70 km § of Batla-

donia, oan C. Auegeliana, §,xi1,1974, 8. Barker;

5 3. Juranda rockhole 106 km 8 of Balladonia,

on C) hiueveliana, 9.xi.1974, S. Barkers 1 3, 19

118 S. BARKER

kot W oof Balladonia. on -C. /unilis, 2341-1975,

8. Barker,

12, Astracus (Astracus) pygmaeus van de Poll,

I88f 178—THO; PRS9: BS, 14, 105, pl. 3,

figs 16, 16a, Mic, Blackburn, 1891: 496,

vah de Poll, 1892: 67, 68. Obenberger,

1920; 366, Carter, 1931: 107; 1933; 41.

Avivuens pygniaews var. subfasclatas van de

Poll, [B86 178-180: 1889: 104. LOS, pl 3,

fig. léb. Obenberger, 1930: 366.

Astravas knareuelle’ var. pygnidens: Bhickburn,

ISut: 1256.

Astrieus sarnouvtli var pypniuens:

1892; 102.

Kerremuns,

Anriens pxynacas: Kerremans, 1902: 148.

Astracys pyynidens suc. subfesciatns: Kerre-

mans, 1902: 144,

Asiracus pyymueus; Carter, 19295 28

Astrucns pyeniacuys var. : ubhateiats Carter.

1929: 282,

FIG, 221.

Type. Holotype: 9 Queensland, MNHN (seen by

author).

Colour, Male.. Turquoise with goltlen-green re-

flections at the apex, dark at the base with

purple reffectionsy antennae dark with blog

reflections. Pronotum dark in the middle with

golden-greein and purple reflections, turquoise

at the sides with polden-green reflections.

Elytra bluck with blue reflections. each with

the following yellow markings: a spot at the

base) a fascia covering the humeral feld,

slightly concave towards the base but not

touching ihe suture; a spor afier the middle,

not touching the margin or the suture. Under-

surfuce metalhe turquoise in the middle. metal-

lic blue pt the sides; hairs silver Legs metallic

bine: tips of tibia and first tarsal segment

testaceous;, tarsal segments 2, 3 and 4 dark

brown with blue reflections.

Female, Head blue at the apex. dark at the

base with purple reflections; antennie black

with blue reflections, Pronotum blick in the

middle with purple reflections, blue at the

sides Elytra as in the male execpt that the

Fascia is always broken in the mriddle te farm

iwo spots, and pill of the yellow murkings are

smaller than those in the mule. Undersurface

deep metallic blue; hairs silver, Legs deep

metallic blue: tips of the tibia and first tirsal

scement testaccous,

Shape and sculpture. Head evenly and closely

punctured; with a median keel; hairy. Prono-

tim closely punctured, the punctures larger at

the sides than in the middle; parallel-sided

from the base until just before the middle then

rounded and tapered to the apex; hiiry, Blytta

custute. the intervals Hat) cach with w row of

punctures und transversely wrinkled: parallel-

sided until afier the middle then rounded and

tapered to the marginal spine; both spines well

developed; humeral fold well developed and

ungled. Undersurface: prosternal punctures

larger at the sides than in the middle, punc-

tares uniform on the abdomen: hairy.

Size. Males 6,2 = 0.05 x 2,5 + 0.02 mm (66).

Females 6,6 = 0,08 x 2.6 + 0.04 mm (40).

Distribution, Queensland and New South

Wales.

Speciniens etentined. Qld: Type: 1 dt & | 2, Edun-

galba on Casdarina sp, 14.xi.1971, BE. BE. Aedarisy

q Jd & 4%, Edungalba on Casuarina sp, 9.xi.1974,

ELE. & §. Adams; | fp & 1 9 Edunealba on

Comey Spi. 'F.xit. Ors, ky Ek 3, Adams.

NSA Ld & 19 %, Oranmeir on Gagharind

littoralis, en 1962, 5 Barker; 2 § & 2 Y,

Majors as on €. Jittoralis, 26.4,1963, 8. “Rare

ker, ACT. 3 ad & 1B, Mt Ainslis on Casmarine

stricta, 29.x1.1962, S. Barker: 6 cf & 2 2 Me

Ainslie on C. sfricta, ‘26.xi1.1962, S. Burkey; 4 2

& 3 9, Tuggeranong on C. stricta, 14.1963, 3, Bar-

ker,

13_ Astraeus (Astraeus) smythi sp, nov,

FIGS 8, 22M

Types.

Holotype; oy Mar parouEl ald,

1956, Eo Smith, WAM, 73-55.

Allotype: % WAM, 73-56.

Paratypes! 2 2, Qld, E. Sutton, QM: | ¢. SAM,

Colour. Male. Head: apex blue with golden-

green reflections, base dark blue with purple

reflections: antennae black with blue and

purple reflections. Pronotum black in the

middle with purple reflections, bloe ut the

sides with golden-green reflections, Elytra

black with purple reflections each with the

following yellow markings: a spot at the base;

a broad fascia covering the humeral fold. rug-

ning transversely towards the suture but not

touching it; a fascia commencing after the

middle at the margin, running transversely to

the suture but not touching it, Undersurface

metallic blue: hairs silver. Legs metallic blue:

tibia and first and second tatsal segments

legtaceous; third and fourth tarsal segments

dark brown. A small preapical spot may or

may not be present.

Female, Head: apex turquoise. base green;

antennae black with blue and green reflections.

Pronotum grecn in the middle, turquoise at the

sides. Elyira black with blue and purple reflee-

tions, yellow markings as described in the male

but Jes. prominent. Unelersurface and legs as

in (he male,

Navember

REVISION OF BLUPRESTID GENUS ASTRAEUS LAPORTE & GORY Hy

Shape and sculpiare. Head evenly punctured;

au median keel: hairy. Pronotum with punctures

larger at the sides than in the middle; a faint

longitudinal impressed line commences at the

middle and runs to the apex: parallel-sided

from the base to just before the middle then

rounded and tapered (o the apex. Elytra

eostate, the intervals between flac each with a

row of punctures and faintly transversely

wrinkled; parallel-sided until before the

middle, then rounded and tapered to the mar-

ginal spine; both spines well developed:

humeral fold well developed and angled.

Undersurface: prosternal punctures shallow im

the centre, deeper at the sides; punctures on

the ahdominal sternites sparsee and shallower:

hairy.

Size. Males 6,5 = 0,08 \ 2.6 + 0.15 mm (2).

Females 6.5 + 0.18 x 2.6 = 0.01 mm (4)

Divrthytion, Queensland. 3

General yemarks. A sinvilii shows close

affinity with A, pygmaeus. L plaice these species

nlongside the previous group of three species

Named after my late colleague, Dr M. Smyth.

Specimens examined. Qh: Types: | 2, One ‘Tree

Will, Brisbane, 1920, #*. Muir, BPBM.

14, Astraens (Astraeus} simulator van de Pall.

1889: 85, 102, 104, pl 3, fius 15, 154,

15h, Kerremans, 1892; 102, 1902; 148-

Carter, 1929: 282. Obenberger, 1920; 366.

FIG. 22N

Vype. Holotype: 2, Pewk Downs, MNHWN (seen by

author}.

Colour. Male. Shiny. Head green at the apex

with golden-green reflections, hlick at the hase

with blue reflections: antennae black with

volden-green and blue reflections. Pronalwm

black with bluc and purple retlectinis. Flytra

black with turquoise reflections, cach with the

following markings: a latge basal spot. 4 Lascia

covering the humeral fold, running transversely

Towarels the suture but not touching il. concave

towurds the base and clubbed wt the end; a

fascia after the middle runing transversely

from the margin but not touching the suture,

Undersurface metallic blue, hairs silver. Legs

metallic blue; tips of tibia and first tarsal see-

ment brown; second, third and fourth tarsal

seements black with blue reflections,

Female. Head black with blue Teflections:

antennae black with blue-green reflections, Pro-

potun black with blue reflections. Elytra black

wilt blue and purple reflections. The rest us

in the male.

Shape and seulprure. Head with close uniform

punctures; no median keel; hulry. Pronorum

evenly punctured: basal third parallel-sided

then slightly rounded and narrowed fo the

apex; anterion margin projecting forward in

the middle, that general area glabrous! hairy.

Elytra cCostate. the Intervals between flat but

slightly wrinkled; parallel-sided from the base,

routded before the middle then tapering to the

marginal spine; both spines well developed:

humeral fold Well developed and angled.

Undersurfuce evenly punctured; hairy,

Size. Males 6-7 + 0.10% 2.5 = 0.05 mm (11).

Females 7.0 = 0,24 x 2.7 = 0,09 mm (6).

Distribution, Queensland.

General rentarky. OF the 17 specimens I have

examined, 7 3 and 2 ¥ have preapical spots

on the elytra. the other eight specimens have

no preapical spots, Although male genitalia

(Fig. 22N) is similar in shape to the previous

two groups of species, the external morphology

of A. simuluter is dissimilar to those five

species, Because of this 1 do not group iL with

any of the above mentioned species.

Specimens examined. Qld: ‘Type: 1 &. 3.x0.1971.

E. BE. Adams. KC; 4 ¢ & 1 Y, Edungalba on

Casuarina eqaiselifalia, V1xii.1873, &. &. Adams,

SAM (1 8 & | 2), EA (8s t Qo Fdungalby on

C. equisetifatia, VS. siL1973, & BE Adantws. FA, 2

4, Edungalba on Casuarina sp, | on cach of

23.41.1974 and B.2i/1974, E. BE. d& S. Adams, BAs

1? Edungalba an Caswarina sp.,.30.xi1974, EE.

& 8 Adams, EAs 2 ¢ & 2 9, Edungalba on

Casarina sp., 401-1974, BE. dt 8. Adants, BA:

2 ¢@, Edungalba on Casvarind sp. 15.xii. 1974.

FE. E. & §. Adams, EA.

1S. Astraens (Astraens) obscurus sp. nov.

FIGS 9, 220

J ypes.

Holotype: of, Fraser Range, W. Aust, on

Casuarina heaseliana, 19.x0,1972, & Barker,

SAM, T 20957.

Allolype: %, Fraser Range, W. Aust, on ©.

huegeliana, U9.xit,1972, 8, Barkes, SAM, ?

20958.

Paratypes; 2d & 2%, Eyre Highway 142 km E

of Norseman, Wo Aust. on C2 feegeliana.

19.x11.1972, S. Burkey, BA (1 So & FT Ql,

ANIC (1 3 & 19,20 &2 9 Fraser Range,

W. Aust. on C. huegeliana, 19.xid872. 8.

Barker, BM (1 Go & 1 2), MNHN (1 od & 1

9). 1 4, 24 km NE of Lake Grace, W. Aust.

on C. huegeliana, 241.1973. 8. Barker, KC

1 3, Tammin, W. Aust. J7, W. Broven,

WADA; | 2, 2 km N of Needilup, W. Aust.

23x11. F972. KR, Newhy, WADA; 1 & 30 km

W uf No. 8 pumping station (Kalgoortic

water omain) on Casing canipestris

247,095, WAM, 71-1773; 1 2, WAM, 7t-

1777: | Y, South Tammin Flora Reserve oi

C. hucgellana, 23,.x01,1972, & Barker, WAM

S. BARKER

ees

Fig. 10, Astruews globosus sp, nov.

Fig. 9. Astraeus obscurus sp. nov.

ig, 8. Asirdeus smythi sp. Nov.

REVISION OF BUPRESTID GENUS ASP RAEU/S LAPORTE & GORY (21

Colour Shiny. Head purple or black with

metallic purple reflections at the apex, metallic

blue reflections at the sides and underneath;

antenhae black with blue and purple refiec-

tions, tips of first, second and rhird segments

dark brown. Prosiotum purple or black with

metallic blue reflections at the sides. Elytra

black with blue and purple reflections

each with the following yellow markings: a

large round basal spot; a broad fuscia belore

the middle concaye towards the base commenc-

ing at the outer margin, enclosing the humeral

fuld,. running towards the suture but not touch-

Ing iGo. broad fascia after the middle but not

touching the outer margin or the suture; a very

small preapical spot in lhe form of a Junette

sometimes missing. Undersurface black with

purple fefiections, except, for a red-brown

patch on either side of the prosternal process

extending to the lateral margin; huirs silver,

Legs: femur, basal half of the tibia and

second, third ind fourth tarsal segments dark

hrowa with bluc and purple reflections, apical

half of the tibia and first tursul segment brown,

Shape and sculpture. Head evenly punctured;

a large median keel; hairy. Pronotum deeply

punctured; a short impressed line projects for-

ward from the basal crypt, becoming a groove

and extending ta the apex; short with rounded

sides gradually narrowing from base to apex;

harry. Elytra costate, the intervals betweem flat

each with a row of shallow punctures; more

or less parallel-sided until before the middle

then rounded and narrowed to the marginal

spine: both spines well developed; humeral fold

moderately developed and angled. Undersur-

face evenly and shallowly punctured; hairy.

Size, Males 9.1 = 9,05 x 3.4 = 0.02 mm (105).

Fetnales 9.2 + 0.08 x 3.4 + 0.03 mm (64).

Ojpsersinition. Western Austyalia.

Specimens exumined. W. Ausl: Types 2 25 & ? 3.

edari, HWW Arnwe, WAN, W714, 1 A

Spencer's Brook, 26 x11.1944, R. FP. MoAlillan,

WAM, 71-1764; i 9, Bejoording, 146,i,1949,

WAM, 71-1764! 3 4&1 F, Bejoording, 29.x1.1950,

RP. McMillan, WAM, 71-1765/8:; t @ 1!) km E

of No, § pumping station (Kalgoorlie water mam)

on Casnertva campestriy, 204.1953, WAM,

TI-A776; 33 2 & 2 Y Coragina rackhole 66 km

S of Balladonla on Casuerina smegeliana,

Sxu,1974. S. Barker; 21 d & 13 oe Ponicr rock-

hole 70 km S of Balladonia on Aeegeliagea

Ssii.1974, S. Barkers 1d do & FY, Torands rock-

hole ibs km S of Ballaloniss on C. hivegeliana,

ORULI974, §. Barker; 3 oT & 29 15S km § of

Bevioduiaae on Crearine hwnnitis. Doi l97T4. S.

erker

16_Astracns (Astracus) glubosus sp. tov.

FIGS 10, 22h

Types.

Holotype; ¢, 77 km along mail) York Rd. W.

Aust. on Coswurinu beezeliuga 41.1968, S,

Barker, SAM, T 20953,

Alfotype: 9. 77 kim alone main York Rd, W-

Aust, on C. huegelfana, 141968, 4 Barker,

SAM, | 20954,

Paralypes: 2 5 & 1 9, Dryandca, W, Aust on

C. Agegeliane 1241974, 8. Barker, BA: 3 of

& 2.9, Dryandra, W- Aust. da C. haegellane,

Rxii.1970, S, Barker, ANIC 11 & & 1 9),

MNHN (i @ & 12), WAM (1 dds Lb od, 77

kin along main York Re W. Aus. on C

finegeliane, Todi 1970, S. Barker, BM; 2 3,

Toompup, W. Aust. on Caspartia Spi,

23.5.1972, &. & K, Cariaby, KC: 1 3, 77

kni along main York Rd, W,. Aust, on

Casuarina sp.. 14,1968, 5, Barker, SAM} 1 eh

oe Reserve, W. Aust. on Casuarina

Lilg68, §. Burker, SAM; 1 9, Spencer's

Brook, W, Anst., 26.xi1.1945, R.P, McNillun,

WAM, 71-1763.

Colour. Shiny. Head wnd pronolum and elytra

black with coppery reflections; untennue black

with coppery anc blue reflections, Elytra: cach

elytron with the following yellow murkings; a

large spot at the base: a broad fascia before

the middle, commencing at the shoulder and

ending near the suture bul not touching it,

concave towards the base; a second faseta after

the middle, commencing al the margin but

not reaching the suture; an oval preapical spot;

@ single very stall spot between the lwo fascta

and close to the suture. Undersurface metallic

copperyy hairs silver. Leas: femur metallic

cappery, tibia and tarsi brown with varying

amounts of metallic blue on the femoru and

tibia and on the upper surface of the tarsi.

Shape and sculpture. Head closely punctured;

with a median keel; hairy. Pronotum deeply

punctured but more closely at the sides than

in the middle; short with rounded sides

narrowed towards the apex; a shor} impressed

line projects forwards from the basal ervypt

hairy. Elytra costate, the intervals flut each

with a row of shallow punezures: rounded,

diverging slightly fron) the base to 4 miximum

width before the middle, thereafter rounding

off to the marginal spine; both spmes well

developed; humeral fold moderately developed

and unsled. Undersurface: finely and sparsely

punctured in the middle of the sternal sce-

ments, mote coarsely and closely punctured at

the sides; abdominal sternites finely and closcly

punctured; hairy,

Size, Males 9.4 + 0,13 x 3.6 © 0,05 mm (27),

Females 9,3 = 3% 5,9 = (05 nin £7).

122

Distribution, Western Australi,

General remarks. This species shows closest

affinity with 4. obscuris

Specimens exantined Wo Aust ‘lypes: 1 @, 77

km along, main York Rd on Casdarina sp.,

51.1968, 5. Burker> 2 4.77 Km along main York

Rd on Casuarina huegetiana, 7.xii.1970, 8. Barker;

1 3) Dryandra on C. hweveltana, 7.xi.1970, 8,

Barker; 9 fd & 1 9% Dryandra on CL huegetiana,

Bx, 1970, SW. Barker,

$7. Astracus (Astracus) mastersi Mac cay,

1873; 239, Kerremans, 1885: 136.

Masters, 1886: 71. Kerremans, 1892; 102,

Astracus sumouelli: van de Poll, t886: 176,

1889: 80, 107-109, pl. 3, fiz. I8b. Kerremans,

1907, 148 Carter, 1929: 282. Obenherger,

1930: 366,

Asraeus splendeas van de Poll, (889; 6, 108,

109, pl. 3, fies 19. 194, Kerremans, 1892: 102:

1902: 148. Carter, 1929: 252, ph. 3, tig. 43.

Obeiiberzer. 1930; 367 (new synonym).

Astravus simplex Blackburn, !892; 211, 212

Kerremans, J902: 149. Curter, 1929: 282.

Obenberger, 1930: 366 (new synonym),

FIGS La, 22

Type. Holotype; 9, Gayndah,. AM. X32094 (seen

by wuthort.

Colour, Mele. Shiny, Heal anil antennae

golden-coppery, Pronotum copper coloured in

the middle al the apex, with u dark blue hewrt-

Shaped) areca in the middle, outlined by «a

copper culoured margin; al the sides, blue at

the base green at the apex. Elytra black with

blue or purple reflections, each elytron with

the following yellow markings: a large spot at

the base; a broad fascia running from the

shoulder covering the humeral told and run-

ning transversely towards the suture but not

reaching it, after the middle i broad fascia

trom the margin running towards the suture

hut nol reaching it) w preapieal spat—in some

specimens the preapical spot is ahsent and

usually in these the first fascia is Iroken in the

middle forming two spots, Undersurfice blue;

hairs silver. Legs: blue: the end of the tibia and

fitst tarsal segment teslaceous.

Female. Shiny, Head blue-green a} the apex,

dark at the base and sides. with purple reflec-

lions; antennae black with blue or purple re-

flections. Pronotum dark blue ia the middle,

metallic blue at the sides, Elyira black with

purple reflections, cach with the following

yellow markings: a large spot at the bise: a

lateral fasciy commencing at the margin and

covering the humeral told, running towards the

surure bur interrupted to form a spot near the

S$. BARKER

suture: a small spot mear the base, at the

shoulder, may of may not be continuous with

ihe base of the Fascia; afler the middle wu fascia

tunning transversely from the margin towards

the suture but nat touching wy a preapical

spot. Undersurfuce blue with metallic gleams;

hairs silver. Legs metallie blue: tips of tibia

and first larsul segment testaceous.

Shape and sculpture, Head shallowly, closely

but uniformly punctured, with a median keel;

hairy. Pronotum shallowly punctured with

punctures larger at the sides than in the

middle; a median longitudinal excavate line

from buse lo apex; straight sided from the

base to the middle then rounded and narrowed

lo the apex; hairy. Elytra costate, the inter-

vals flut but slightly wrinkled each with a

row of punctures; more or less purallel-sided

to before the middle, rounded at the micille

then tapering to the marginal spine; both spines

well developed; humeral fold well developed

und acutely angled. Undersurface with shallow

uniform punctures except in the middle and

on the outer edges of the abdominal stennites,

both ureas being glabrous; hairy.

Size, Males 7.9 %F O.16 x 2.7 + 0.22 mi (17).

Females 8.4 = 0.26 x 3.2 + 0.10 mm (10),

Distribution. Queensland and northern New

South Wales.

General remarks, This is the only species of

Astraens that his been greatly confused (sce

van de Poll 1889, p. 79). The basic reason for

this ts that the [enyles of A. metstersi and A.

senouell? ace very similar in pattern although

the males of these species are quite distinct.

Both species were described from female types

The lack of field collecting experience was the

most likely reason for van de Poll redescribing

the male of A, mrestersi us A. spfenidens, as he

woulll nolL huve had aceess to an associatet

series of males and femmes of Ihe two species.

Specimens exantined. N.S.W > Type; | od purutype.

Ash TL, ANIC: 1 a & 1 2 paratypes, Gayndah,

ANTC. Qld: Type; dA. xplendens van de Poll,

Rockhumpton; 3 9 Dalby, Mrs, F. A. Mabler,

SAM: | 9, Stomburpe. 4. Geenmell; 14 ob & 5 9%

Edungalba on Clasuatina squisetifolia, 24.1971,

E. EL Adamy. Type 2, 4. simplex Blackburn NM

(locality of collection uncertain),

18. Astyaeus (Astraeus) samouelli Saunders.

i868; LO. pl oT, fiz, 125 187i: 43.

Masters, 187L: 124. Ketremans. [8R5-:

136. van de Poll, 1886+ 176, 178: 1889:

$6, 107, pl, 3, figs $8, 18a, Blackburn,

INO); 496. Kerrenians, 1892: 162. van de

REVISION OF BUPRESTID GENUS ASTRAMUS LAPORTE & GORY

Poll, 1892; 67, Blackburn, 1892; 212,

Kerremans, 1902+ 145. Carter, 1929: 282,

Astraeus samouellei: Germminger & de Hyrald,

1869: 1380. Masters, 1884; 71. Blackburn,

1890; 1256, Obenberger, 1930> 346.

Astravus sasonelli- Kerremuns, 1900: 295,

Astraeus <plendens var. eqibrikiellys Oben-

berger, 1936: 133,

FIGS 1b, 22R

Type. Holotype: 2 BM {seen by suthor).

Colour, Male. Shiny. Head golden-vreen,

golden or coppery; antennac coppery or tur-

queise. Pronotum uniformly golden-green or

golden or coppery in the middle then golden-

areen blending into turquuise at the sides.

Elytra black with blue reflections. each with

the following yellow matkings: a large spot mt

the base, a fascia just before the middle cover-

ing the humera) fotd, extending backwards

then transversely towards the suture but not

touching it; a spot at the shoulder which may

be discrete or continuous with the base of the

firse fascia; a fascia after the middle, carmmenc-

ing al the miurgin and running towards the

suture but not touching it; variably, a small

elongate spot hetween the two fascia closer to

the first and near the sulure but not touching

it, @ preapical spot. Undersurface blucsgteen;

hairs silver. Legs blue-green; lips of the tibia

and most of the first tarsal scgmnent testaceous.

Female, Shiny. Head green wl the apex with

golden reflections, dark blue with purple reflec-

tions in the middle of the base, turquoise at

the sides; antennae green. Pronolum dark blue

with purple reflections in the middle. turquoise

at the sides, Elytra black with blue refiections,

with markings ss tm the mulé except that the

first fuscia is broken in the miiddle to form a

marginal Fascia and a spot near the suture.

Undersurface turquoise: hairs silver. Legs tur-

quoisc: tips of the this ind most of the basal

part of the first tarsal segment testaccous.

SRape and yealpiure, Head shallowly, closely

but uniformly punctured; with a median keel;

hairy. Propotum shallowly, closely but uni-

formly punctured} with a median longitudinal

excavated line; uniformly rounded and

narrowed from base to apex; very hairy. Elytra

costate. the intervals between flat; slightly

lapered from the base to the middle, then

rounded aud tapered to the marginal spine;

both spines well developed. Undersurface

shallowly punctured, more closely at the sides,

less so in the middie; midsurface glabrous;

hairy.

Size, Males 4.6 = 0.07 % 3.0 + 0.03 min (52),

123

Female: 9.2 = 0.14 5 3.3 = 0.06 mm (40),

General remurks, The type uf A, splendens var.

enbrikielus Obenberger is a } specimen of A

swHouelli. This species shows closest allinity

With A. masteryi,

Distribution, New South Wales.

Specimens examined, N.S. Type: 2 co & 3 9,

Bredbo on Cusuarina stricta, 81.1963, §. Barker;

12 ¢ & 20 G Major's Creck on Coswiine lt

rordlis, 26,1,1963, $. Barker; 8 d & 2 2, Khunyunis

Station, Minuma Runge on CG. iitteralis, 9.1i,1963,

S. Burkery 4 3b & 2 2, Captains Flat.to Braidwood

Ru, on C. fitturalix, 931.1963. 8S. Barkers Type 2

A, xplendens var, embhrikivlluy Qbenberger,

Comara, NMP, 21989. 4.0.7: 4c, Mt Ainslie an

C. stricta, 26.xi1, 1962, 5, Barkers 7 & & 12 9, Tue-

weranong on C. seilete, |.xii1962, 8. Barkers 3 2

& 8 GY, Tuggertuong on C. striced, 2.xi0.1962, 8.

Barker: 4 f & 2 2, Tuggeranone on C. strict,

11,1963, S. Barker.

19. Astracus (Astracus) dilutipes vari de Poll,

IS89: 86, INS, 106, pl, 3, fig, 17, |7a.

Hlackburn, 1892; 212. Kerremans, 'S¥2:

1OT: 1902: 148, Carter, 1929; 282. Oben-

berger, 1930: 363.

“Astraens samouell? yar. dilitipes vun dé Poll,

1886: 180.

Astracus strand’ Obenberger, 1928: 205; Carter,

1929; 302. Obenberger. 1930; 367 (new syno

nym).

FIG, 228

Type. Holotype, MNHN (not seen by author}.

Cofour, Mate, Shiny. Head green at the apex

and golden green at the base or golden-green

ut the apex and coppery at the base; anlennie

black with blue reflections. Pranotum divided

in uw transverse direction by an M-shapeul line.

the small apical part being green or golden-

green, the larger basal part being dark hlug ar

black und the line of demarcation, purple or

coppery. Elyira black with blue reflections

each with the following yellow markings: in

the middle but not touching the sulure there

is; a large hasal spot; 2 spot before the middle;

a preapical spot. On the murzin: o Jars spol

covering the humeral fold; after the middle a

transverse fascia not touching the suture,

Undersurtace blue or green: hairs silver Legs:

first and second demora brown on the outer

murgins, third femur totally or partially solid

colour similar io the rest of the undersurface;

remainder of Jeg and tarsi testuccous

Female. Head green at the apex, purple in

the middle, dark blue at the base with purple

reflections: antennae black with blue reflec-

lions. Pronotum black in the centre with

purple and blue reflections, blue at the sides.

Elytra as in the male. Undersurface dark blue;

124 8. EARKER

hairs silver. Legs: outer margin of femora

brown, tibia and tarsi testaceous.

Shupe and sculpture. Head very densely and

evenly coyered with deep punctures: wilh a

thin median keel; covered with long hair. Pro-

nolum evenly punctured, the punctures al the

sides larger thin those in the middle: slightly

rounded but narrowed from base to apes,

median excuvated line from base tw apex;

hairy, Elyira costate, the intervals Hat with 4

row of deep punctures and faintly transyersely

strigose; narrowed from base until after the

middle then rounded uod again narrowed ta

the marginal spine; both spines moderately

develuped,; humeral fold well developed and

angled. Undersurface evenly and shallowly

punctured; with long haits.

Size. Males 839 = 121 ¥ 3.1 | 0.08 mm (11).

Pemates 9.4 + O13 x 3.4 = 1.07 mm (7),

Diverdhution. New South Wales ond Queens-

land,

General remarks, The type of A. strandi Oben-

berger is o small & specimen of A. dilutipey.

The external morphology of this species is like

thut of 4, sltasfersi and A. samouelli but male

genitulia (hig. 228) 1s different. 1 do not group

it with the other two species,

Specimens examined. NAW & & & 5 9, Major's

Creek on Cusnarine littoralty, 265.1963, s Barker.

Old: 2 ¢ & 2 9, Paluma on Casuarina sp.,

Wai. 1970, EE. Adams; Type @ A. strand Oben-

berger, NMP, 21991,

20. Astracus (Astracus) adamsi sp. nosy.

FIGS 11, 221

Tyre,

Holotype: 3, Edungatha, ON on Casiwarlna

equisetifolla, BO.x8L 197d EL & §. Adams,

SAM, | 20944,

Allotype; &, Eduugulba, Qld on C. a plincti alla,

ih tendie BLE, Adams, SAM, T 20

Puratypes: 3 fo & 3B Eduvealbi, aia nA Cc.

equisetifolia, VS.xii. ida, Fok §. Adantw, EA

(de &19), ANIC (138419), BMO Se

Ia); | gf & 1S, Bdongulba, Qld on €.

equisetifolla, 23.NiINTA & Bx 1974, Bo SS.

Adams, MNHN: ig &dS Edunsulbu, Cd

on ¢. equisetfaha, (4xii.41974, BE & S&S.

Adams, SAM,

Colour. Shiny. Head black with purple tefiec-

lions or coppery-purple with metallic rellec-

tions. Prongtum black with purple reflections

or bronze in the middle, coppery purple ai the

sides with metallic reflections. Elytra black.

with blue and purple reflections, gach elvtron

with the following yellow murkines: a basal

Spotl; a spot civverme ihe humeral fold but not

teaching the shonlder a large spot in the

middle not touching the suture and just behind

the second spot, these lust two in the form of

a broken fascia; just behind the break, near

the margin but het touching it, there is a small

spot (not present in a third of the specimens

examined): after the middle a fascia concave

backwards commencing at the margin but not

reaching the suture; a small preapicul spot.

Undersurface coppery-purple; legs pale brown,

the basal ends of the tibia darker: hairs silver.

Shape and sculpture, Head closely and evenly

punctured; no median keel; lightly haired, Pro-

notum closely and evenly punctured except for

a short glabrous line in the middle at the apes;

at the sides gradually rounded and narrowed

from base to apes. Blytea costate, each inter-

val with a row of deep punctures and faintly

transversely wrinkled: parallel-sided Lo just

before the middle then rounded and tapered to

the sharp marginal spine; sutural spine well

developed: humeral fold well developed and

angled, Undersurface closely and evenly pune>

tured, the punctures shallower on the proster-

nal process than elsewhere; lightly haired,

Size. Mates 7.1 + (.14 8 2.9 © 0.05 mm (6).

Females 7.7 = O10 x 3.2 + 004 (9).

Distribution, Queensland,

General remarks, Male genitalia (Fig. 22T) of

A, adamsi is similar to that of a. dilutipey but

external morphology is different. T do not

group these Iwo species, Named after Mr E. E

Adams.

Specimens exanyred, Types only.

QL, Astracus (Astrqeus) intricatus Carter, 1925:

229, fig. 1; T929: 282, Obenberger. 1930:

366,

FIGS le, 12, 23A

Type, Holotype: ay iM

author).

Colour. Shiny. Head metallic green al the apex

and sides, purple at the base: antennae black

with coppery-purple reflections. Prowotum

metallic purple in the middle, deep cuppery-

purple at the sides. Elytra black with purple

rellections, each elytron with the tollowing

yellow markings: uw basal spot, a spot at the

middle and a spot after the middle, all elongate

und decreasing in size from in front back-

wares, each with the husal end diverginy out-

wards and all in Jine to give the appearance

of a single, broken vitta “with the basal end

diverging outwards and the apical end con-

verging to the suture, buf not tavching if; i

fuseia Commencing at the shoulder, covering

Monnro, (seen hy

REVISION OF BUPRESTID GENUS ASTRAEUS LAPORTE & GORY

5mm

Fig. 13. Astraeus macmillani sp. nov.

ig. 12.. Astraeus intricatus Carter

Fig. 11. Astraeus adamsi sp. nov-

126

the humeral fold and running along the outer

margin to the middle then eatencding obliquely

towards the suture, reaching slightly below but

not touching the middle of the three median

spots; atter the middle a fascia commencing

al the outel margin and running transversely

towards the suture bat not touching it; an clon-

gale preaptcal spot, close to the suture bul not

touching it. Undersurface deep coppery-purple;

hairs silver. Legs deep coppery-purple, ends

of the tibia, first and second tarsal scements

brown, third and fourth tarsal segnients durk

brown,

Shape and sculpture, Ulead closely punctured;

with «w median keel; hairy. Pronotum more

closely punctured cit the sides than jn the

middie, with a median longitudinal impressed

line vistble ut the buse and apex only; paralicl-

stded at the base. gently rounded until after

the middle, songly rounded and narrowed to

the apex, projecting forward slightly in’ the

ropuldle of the upicul edve; hairy, Elytra costate,

the intervals between flat. cach with a row

of shallow punctures; parallel-sided from the

base, rounded off well after the middle and

narrowed strongly to the mureinal spine; both

spines well developed; humeral fold moder-

ately developed and angled. Undersurface:

stemal segments finely and sparsely punctate

in the middle. deeper ond larger punctures at

the sides: abdominal sternites finely and closely

punctured; hairy.

Size. Males 9.6 © 1.10% 3.5 = (05 mm (501.

Females 9.8 > 0.15 x 3.6 £0.07 mm (35).

Distribution. New South Wales,

Generel remarks. The species was described hy

Carter from a unique specimen In the MacLeay

Museum. Only two of the §5 specimens I

have collected conform to the colour parern

of the lype. Uhe pattern on the elyira is

variable. ranging from the basic putter as

described herein (nine specimens) to patterns

in which all of the yellow markings are -con-

fluent, giving the appearance of yellow elytra

with black borders. Between the extremes, in-

lermediary patterns occur, including the intri-

cate pattern of Carter's type. There is no dif-

ference between the sexes in size or colour

pattern, All of the specimens T collected were

tuken on low Caswerina nana heath in the

Minums and Kybean Ranges, Monaro District,

N.S.W. I do not group 4, intricatus with any

other species.

Specimens examined. WS.W.: Type; § do & (0 9,

Oranmeir on Casuvaring fie, 9xiL1962, 5S. Bar-

$3. BARKER

ker; 23.3 & 9 2 Oranmeir on Co yang, 261.1964,

S. Barker; 3d, 16 km S of Counterany Station

on C. nana, 3aui.1963, 8. Barker) 20 f° & 15 2

snaps Siation on ©, nena, 941.1963, 8. Bars

cer.

22, Astracus (Astracos) crassus van cde Poll,

1889: 85, 95-97, pl. 2, fig. 9, Ga. Kerre-

mans, [892; LO1; 1902: 148. Carter,

1929: 262, Obenberger, 1930; 365.

Astravits flavopictus van de Poll, 1386) 180.

FIG, 23B

Type. Holotype, MNHWN (nor seen by author).

Calour. Shiny, Head and pronotuny black with

purple or blue and purple reflections; antennae

black with blue and purple reflections. Elytra

black with blue and purple reflections cach

with the following yellow markings: the basic

pattern Consists of itregulur spots in two rows;

four spots in the middle tear the suture, but

not touching it, one at the buse, one before the

middle, one after the middle and one in the

preapical area: three spots ulong the margin,

one at the shoulder and covering the humeral

fold, one at the niiddje und one after the

middle; there may also be a few irregular spots

and in two specimens there are spots at the

base of the sutural spine. Undersurface and

legs black with blue and purple reflections;

hairs silver.

Shape and sculpture. Head with a medion keel;

evenly punctured; hairy. Pronotum with small

and sparse punctures in the middle leaving

alahrous areas, lager and more dense at the.

sides with a short impressed line projecting

forwards from the basal crypt; gently rounded

and narrowed at the sides from the base to

two-thirds way to ihe apex, then tapered 10

the apex with the apical edge projecting

slightly in the middle. Elytra punctate-striate

with the intervals flat towards the base ane

slightly concave towards the apex, cach with

a.raw of shallow punctures; parallel-sided until

after the middle, then gently roonded and

narrowed to the small marginal spine; sutuzal

apine well developed; humeral fold poorly

developed but slightly angled. Undersurface

shallowly punctured, more closely at the sides

than in the middle; hairy.

Size. Males 14,4 + O25 x 5,3 | 0.06 mm (%)

Females 16.2 = 040% 6.3 + 0.15 mm (7).

Distribution, Queensland and New South

Wales.

General remarks. This species cannot be

grouped with any other

REVISION OF BUPRESTID GENUS 45STRAEUS LAPORTE & GORY

Specimens examined. Old: 1 dy EB, Suttaw.N. SW

Id &J) 3 Sydney, SAM: | 2, Griffith, SAM;

% “enatun Caves, J. C. Wilbard, SAM;

f. Tupper, SAM; 1 9, Sydney, We. cu Boulas.

SAM: I q, WwW, dh Bolas, SAM; 1 ?, SAM; 2 ¢,

Majors Creek on Casuarina litloralis. 26.1. 1963, Ss.

Rarker, | ¢. Khanyunis Station, Minuma on C.

litravalis, 26.L.1963, S. Barker.

23. Astracus (Astracus) major Blackburn,

189: L257, 1258; 1891: 496. Kerremans,

1892: (02; 1902: 149.

Avtrarey wavarchis var. majars Carter, 1929:

282. Obenberecr, 1930; 366.

FIGS 17, 23C

Type. Holotype, BM {nor szen by suthor).

Calour. Shiny, Head coppery with metallic

reflections; antenmae coppery-green or blue.

Pronotum dark blue with purple reflections.

Elytra dark blue with blue and purple refice-

tions, each elytrots with the follawing yellow

Markings” a fascia running from the shoulder

Jransvetsely across the base towards the suture

but not touching it; before the middle w thick

fascia running from the margin transversely

to the suture; after the middle there ts a fascia

commencing al the margin, running towards

the suture but mot reaching it. There 1s an oval

re] spot in the region of the marginal spine.

The outer margins of the fascia and humeral

fold are red, Undersurface dark blue with

metallie purple reflections; haits yellow. Legs

red-hrown; tarsi blue.

Shape and sculpture. Head with sparse shallow

punctures at the base. wrinkled towards the

apex; with a thin median keel; hairy, Prone

tum with small shallow punctures jn the

middle, larger deeper ones ut the sides with an

inregular glabrous longitudinal area in the

middle; there is a short median longiiudinal

impressed line projecting forwards from the

hasal crypt; short with the sides gradually

riunded from byse to apex. Blytra punctate—

sirfate from the base for most of the length, the

intervals between the striae convex. costae at

the apex only; parallel-sided until just after

the middle then gradually touhded ta the mal-

ginal spine: both spines well developed:

humeral fold poorly developed und slightly

angled, Undersurface densely punctured at the

sides, sparsely and shallowly puactured in the

middie: hairy.

Size. Males 15.3 = 0.70 x 5.9 = 0.22 mm {4)-

Females 15.5 + 0.98 x 6,3 + 0,91 mm [2).

Divribistion Western Australia, South Aus-

tralia and Victoria.

General remarks. | have ne evidence of oaver-

fap in the distribution of this species. which

127

is found in low rainfall areas of three states,

ind Astreeus navarchis, which is presumably

found in higher rainfall areas in Victoria,

Because of differences in distribution and

appearance I have treated the two as distinct

specivs,

Speeimeny exaniined, W. Aus: | 8) WAM, 73)

62; 1 dy dung, SAMs 1 4) Wialki on Acacia eoal-

qurdiensis, [8.ix. 1957. S. Barker. WAM, 73-60: L

2, Culham, fix-t9G1, RL PO Metillai, WAM,

73-61, 8. Aust: | 2, Monarto, JG. OG. Tepper,

SAM. Pies 1 9%, Sea Lake, Gordie, NM.

24. Astracus (Astracus) nayarchis {Themson)

Conoxgnatha navarchis Thomson, 1856:

115, 116, pl, 6, fg. 2. Masters, 1886: 79,

Asteeus nayerchie: Saunders, 1371: 43.

van de Poll, 1886: 176. Masters, 1886;

71. van de Poll, 1889: 84, 91. 92, pl. 2

fig, 3, Sa, Blackburn, 1890; 1257. Kerre-

mans, 1892: 102. Blackburn, 1892; 211.

Kerremans, !902: (L48. Carter, 1929:

282, Obenberger, 1930: 366.

FIG. 25D

Type. Holotype, MNHWN (not seen by author}.

Colour. Shiny. Heal coppery at the apex, dark

blue anc purple at the base; antennae dark

brown, segments one and two with golden-

green reflections, the cese with blue and purple

reflections, Pronotum coppery al the apex and

sides, dark blue and violet at the base, Elytra

bluck with violet reflections; each elytron with

two yellow fascia, the firat commencing at the

margin und covering the humeral fold, slightly

concuve towards the base and touching the

suture, the second after the middle, commenc-

ing, at the margin and running transversely

towards the suture but nat touching it. Under-

surface: prothoracic sterites metallic golden-

green, abdomen metallic violet; hairs yellow

Legs red-brown with vieler reflections.

Shape and seulptuse. Head closely punctured

at the hase, grooved and wrinkled al the apex;

Slightly excavated between the eyes towards

the base, but with a thin glabrous median

keel commencing anterior (0 this; hairy, Pro-

notum with small punctures in the middle, Bul

mainly lacking in the centre, forming an in-

definite median longitudinal glvbrous fine, the

puncttires larger and deeper al the sides; a

median longitudmal impressed line projects

forwards from the basal crypt to the middle,

short, rounded at the sides and narrowed from

base lo upex, Elytra punctale-striate. the in-

tervals convex at the basc, flat at the sper,

those in the centre with a shallaw row of punc-

tures, those at the sides with decp punctures;

128

parallel-sided until after the middle. then

rounded ane nurrowed to the marginal spine;

both spines well developed; humeral fold

poorly developed and slightly angled, Under-

surface: anteriorly the punctures are small in

the middle and fargér at the sides, on the

ahdomen uniformly small; hairy.

Size. Males 14.4 0.41 x 3.5 = 0.13 mem (7).

Females 16.2 | 0.3466 = 0,08 mm (5).

Distribution. Vietaria-

General renwrks. This species shows close

aflinuy with A.. major

Speeimeny examined Vier 2d, SAM; 1 9, Jan

Jie, NM) 35 d & 42, NM.

25_ Asterneus (Astraeus) fraterculus van de Poll,

L889; 84, 92, 93, pl. 2, figs 6 6a. Kerr

mans, 1892: 182: 1902: 149, Carter,

1929; 282. Obenberger. 1930: 365.

FIG. 23E

(tpe. Holowpe MNHN (not seen by wuthor)-

Coleur. Shiny. Head antennwe and pronoluns

blue-blick with blue reflections, Elytra blue-

black, each elytron with two yellow Euscia; the

first commencing at the shoulder and cover-

ing the humeral fold, then running trans-

versely towards the suture but noc touching it,

concaye (Owards the base; the second after the

middle, conmmencing at the murgin und runs

ning transvetsely towards the suture bul not

touching it. Undersurface deep metallic blue;

hairs silver, Legs deep metallic blue,

Shape and sculputre. Heuc shallowly and

sparsely punctured; with a small median keel:

hairy. Pronotum sparsely hut evenly pune-

tured, the punctures at the sides larger and

deeper than those in the middle: a short

median longitudinal impressed line projects

forwards from the basal crypts short, the sides

tapering acutely from Tase to apex. slightly

romnded in the middle, Elytra costate, the in-

tervals flat with regular shallow punctures;

purallel-sided until after the middle, then

rounded and murrowed to the marginal spine;

buth spines well developed; humeral fold

poorly developed und slightly angled. Under-

sutface with smal! shallow punctures in the

nididle, larger amd deeper punctures ar the

sides; hairy.

Size. Males 9.3% 0,30 x 4.2 + 0,10 mm (7).

Females 8.8 = 0.30 x 4.0 £0.11 mm (3).

Onverihiuion, Western Australia,

General remerks, She external morphology of

this species is very similar to that of 4. diajer

§, BARKER

and A, aavarchis, however male genitalia is

different (Fig. 23E). Therefore | do not group

the three species together but place A, frater-

culus Next to4, major and A. navarchis,

Spectiiens evamined, W. Aust 3 && 1 2, Borden

on Ffakea trifurcata, November 1939, H. W.

Brown, WAM, 73-378/381; { 2, Bushmesi,

18.x1.1939, RP. MeMillan, WAM, 73-383; 2 4

& | 2 Hopetoun, 18.x,1946, Ages. Morris, WAM,

96-1910/1912; 1 of, Wembly on Daviesin diveri-

ea Sux. 1970, 8, Barker; 1 3 Burden, WAM,

73-382,

26, Astraeus (Astraeus) prothoractcus van de

Poll, 1889: 85, 98, 99, pl. 3, fig. 11, 112.

Kerremuns, 1892: 102) 1902: 149, Carter,

1929; 282. Ohenberger, 1930: 366.

FIG. 23F

Type. Holotype, MNHN (not seen by author):

Colour, Shiny. Read, untennae and pronomm

brunze, Elytra dark brown with bronze reflec-

tions. each with the following yellow mark-

ings: along the margin there is a vitta, com-

mencing at the shoulder and ending in the

preapical area which muy he broken inte

several elongate spots; in the middle bul not

touching the suture there is a vitta commenc-

ing on the anterior edge and ending in the

preapieul arca, this is usually broken into a

hasal spel, a spot before the middle and an

clongate spot afler the middle. Undersurface

bronac; hairs silver. begs red-brown,

Shape and sewlpture. Head closely and evenly

punctured; No median keel; hairy. Pronotum

with punctures wrinkled ac the front and sides,

projecting conically in the middle where the

punetures conlesce and beeome — strignse;

gently rounded at the sides from the base to

the middle. then tapered.and narrowed to the

apex. the apical edye browdly convex in the

middle; ‘airy, Elytra costae, the intervals Ast

with a deep row of punctures: parallel-sicdect

unlil alter the middle then very gently rounded

to the miurginal spine; both spines poorly

developed, blunt and close tovether; humeral

fold poorly developed and slightly angled.

Undersurface; punctures closer ut the sides

than in the middle: legs and whole under-

surface densely covered with hai.

Size. Mules 10.4 © 0.335 x 3,7 + 0.13 mm (4).

Females 10.5 = 0.25 % 3.9 + 0.17 mm (6),

Distribution. Western Australia and Queens-

land,

Ceneral remarks, van de Poll (1889) listed the

two specimens he hid as voming from

Clarence River and Champloi Bay, and Cpr-

REVISION OF BUPRESTID GENUS ASTRAEUS LAPORTE & GORY 120

ter (1929) gave the range as Clarence River,

N.S.W, It is naw known that in Western Aus-

tralta the species is associated with Banksia

privrores which ogcurs on yellow sandplain

jn a wide area between Shark Bay and Esper-

ance, van de Poll's reference to Champion

Bay would be to the Geraldton area. I have

not located a specimen from N.S.W. Overall

the body shape of this species is elongate and

more or less cylindrical.

Specimens examined. Qld> 1 9, Myall Park. Glen-

mergan, Noy, 1962, Dorothy Gardin, TH. W,

Avy: 2 4 & 1 2% Tammin, 20.x1,1939, AL MM.

Danelus, WAM, 39-2668/70; | of & 3 7, Goomal-

fing, 14.13.1950, R. P. AfteMillan, WAM, 73-371.

374; 1 2, 19 km N of Northampton, 14.1972,

K. T, Richards, WADA,

27. Astraeus {Astraeus) elongatus van de Poll,

1886 177; 1889: 85, 101, 102, pl. 3, fig.

14, I4a, Ketremans, 1892: 101; 1902:

148. Carter 1929; 282. Ohenberger,

I95lls 365.

FIG, 23G

Type. Holotype, MNAN (not seen by author).

Colvur. Shiny, Mae. Head and pronotum

green; &ntennac blue-green, Elvtra black with

purple-blue reflections with the tollowing

yellow markings; a single spot al Lhe base; a

spot at the shoulder covering the humeral folkd:

a spot on the outer margin at the middle; 4

spot between the previously described one and

the apea; near the suture but not touching it

there is a large spot before the middle, a

smaller one after the middle and a smaller pre-

wpical spot. Undersurface and legs grcen: hairs

silver.

Foytale. Head and pronotum golden-green with

golden reflections) antennae blue-green, Elytra

as ia the male, Undersurface and Jegs golden-

gteen, hairs silver.

Shape and sculpnire. Head deeply and evenly

punetured: fo median keel) hairy. Pronotuimn

with punctures closer at the sides than in the

middle: with a shore median longitudinal im-

pressed line projecting forwards from the basal

crypt; laterally dilated and with the apical edge

projecting in the middle; hairy. Elytra punc-

Eate-striate at the base costate towards the

apex, the intervals flat and slightly trans-

versely wrinkled; more or less parallel-sided,

rounded after the middle to the marginal spine:

horh spines well developed: humeral fold

poorly developed, slightly angled. Undersur-

face closely punctured; hairy,

Size. Males 10.5 = 0.4 » 3,7 = 0,13 mm (8).

Females 11,8 & 1.28 « 42 = 0.J1 iin (12),

Distribution. Western Australia.

General remarks, A single specimen collected

at Dryandia hus a bright blue bead and pre-

thorax. Overall the body shape 1s elongate and

cylindrical. This species has closest affinity

with A. protharacicus-

Specimens examined, W. Aust: 7 a & 9 FB

Quairading on Nanthorrhoea «sp. 19x 1970, 8,

Barker; | & & 2 G, Quairading on Aaniliorrhora,

7.xi.1970, S. Barker; 1 2, Dryandra en Casuarina

hueyellana, 8.xi1.1970, 8. Barker.

28. Astracus (Asiracus) vittalos van de Poll,

18B9: 85, 99, LOO, pl. 3 fig 12, Ia.

Kerremuns, 1892; 102, Carter, 1929; 282,

Gbenberger,, 1930) 367.

Alstraens vittalus: Kerremans, 1902: 148.

FIG. 23H

Type. Holotype. MNHWN {not seen by authort.

Colour. Shiny. Head and pronotum black wilh

purple reflections; antennae black with blue

teflections, the apex on the first, second and

third segments dark brown with purple refice-

tions, Elytra black with violet reflections, each

with the following yellow markings: a vitta

along the margin from the shoulder to the

preupical region, broken just after the middle;

4 Vitta jn the middle but not touching the

suttire from the base fo near the apex, broken

near the middle, Undersurface metallic: purple;

hait's silver. Legs brown.

Shape and sculpture. Head brond and closely

punctured; no median keels hairy. Pronotum

with small and shallow punctures in the

middie, larger and deeper at the sides;

gradually rounded from base to apex; the

anterior margin projecting forwards in the

middle, hairy, Elytra punctate-strlate at the

base, faintly cosiate al the apex amd slightly

wrinkled, the intervals slightly convex at the

base and flat at the apex, cach with a row of

punclures; parallelesided inti] just after the

middle, then tapered to the very short marginal

spine, sutural ypine very broad, humeral fold

poorly developed, slightly angled. Undersur-

face closely und densely covered with fine

punctures; hairy.

Size. Male 9.9 x 3.7 mm {1)-

Distribution. Western Australia.

Specimens exainined. W. Aust.) 1 oy Watnine,

10.xiL 1950, RO P. MeMillan, WAM, 71-171.

29. Astraces (Astracus) flavopictus LaPorte &

Gory, 1837: 2, pl. 1. fig. 1. Imhoff, R565

46, Lacordaire, 1857: 43. Getimimuer &

de blarold, 1869 1380, Masters, 187):

130

124. Saunders, I871; 43. Kerremans,

L885; 136, van de Poll, TS86: 176, 177.

Masters, 1886; TL. van de Poll, tsa9:

§5, 97, 98, pl. 2, fig. 10, 10 Kerremans,

1892; [O1; 962° 148. Carter, 1929: 282.

Obenberger, 1930: 365. Barker, 1964:

306, 307.

FIG. 235

Trp. Holotype, MNUHN (nar seen by author),

Celonr. Shiny, Head and pronotum brown with

variable green and/or purple = reflections;

antennae black with blue reflections, Elytra

brown with variable bronze and violet refiec-

lions, with the following yellow markings im

two rows, one in the middle but not touching

the suture, the other along the margin; in the

middle; an elongate basal spot; a transverse

bar before the middle; un clongate spot afler

the middle; a long thin preapical mark. Along

the margin: an elongate spol from the shoulder

covering the humeral fold; a spot in the

middle; a spot utter the middle. The three

spots wong the margin may coalesce forming a

single villi of be separated into two or three

spots, Undersurface and legs bronze; hairs: sil-

ver,

Shape and sculpture. Head densely punctured;

uo median keel; hairy. Pronotum closely and

evenly punctured; basul half parallel-sided,

Iherealler rounded to the apex; apical edge

projecting forward in the middle, a short im-

pressed line projects forwards from the basal

erypt: hairy. Elytra costate, the intervals flat

but deeply pimctured and slightly wainkled;

purallel-sided from base until ufter the middle,

then narrowed to the small marginal spines

sutural spine short; humeral fold moderately

developed and angled. Undersurface closely

and evenly punctured: densely hairy as are the

fegs,

Size. Males 10.9 ~ O38 % 99 1 O17 mm

(12), Females 12.0 £0.27 x 4.4 + 0.10 min

(15),

Distalburion. Western Australi.

General vemarks, This species shows closest

alfinaty with A. vitendus.

Sovcimens ¢ximined. W, Aust; L 9, Porongvoups,

28.0 1969, FL HW, Uther Baker; | 2 & 1 9 64 km

Mong main York Rd, on Jacksania sp., 1.1, 196%, 8,

Barker; 1) a & 13 2, 64 km along main York Rd,

on Jucksenia sp. 6.x1,1970, S. Barker,

30. Astraeus (Astraeus) muacmillani sp, nov,

FIGS 13, 23]

8. BARKER

Types,

Holotype: 2, 77 km xlony main York Rd, W.

Aush on Casuarina haeveliana, 2VXA9T0, 8,

Barker, SAM, 120985,

Allutype: 2, 77 km along main York Rd, W.

Aust. on C, fivegeliana, 5.41968, 8. Barker,

SAM, £ 20956,

Paratypes: | of & | 9.77 km sloug muin York

Rd, W, Aust. on Casuarina sp., Lil968, &,

Berker, EA; 2 ¢ & 2 9,77 km slong main

York Rd, W. Aust, on ©. huegeliana,

21.81.1970, 8. Barker, ANIC (1 & & JB),

BM (1 3), MNIIN (1 9); 1 9, 77 km along

min York Rd, W. Aust. on C. Aueyellana,

5.4,1968, §. Barker, BM; 6 & 2 9,77 km

long main York Rd, W,. Aust, on ©,

huegeltana, 7.Xi.1970, S. Barker, MNHN (1

d), SAM (40 &2 9), WAM (1d), 15 &

2°39) UW km S of Walebing, W. Aust.

6.x.197L, KL T. Richards, WADA! 1 9, 77

km along main York Rd, W. Aust. on C.

hiegeliuna, 54.1968, 8, Barker, WAM.

Celaur., Shiny, Head black with purple reflee-

lions: antennae black with metallic blue

refiections, cach of segments [—4 with dark

hrown apex, Pronotum black with blue-green

reflections in the middle, purple reflections

in the front and at the sides. Elytr black with

purple reflections, each with the following

yellow markings: 4 large spot at the base; a

fascia commencing al the shoulder, covering

the humeral fold und running transversely

towards the suture but not touching it, con-

cave towards the base (sumetimes broken inte

a marginal and 4 medial spot): a small spol

after the middle on the outer margin; an oval

preupical spot; between the last twa pols a

short fascia, running from the margin, hulf

way to the suture. Undersurfate metallic

purple and bronze; hairs silver, Legs red-

brown.

Shape and seulpture. Head closely punctured:

no median keel; hairy. Pronotum deeply and

closely punctured at the sides, sparsely in the

middle; 9 median longitudinal impressed line

proicets forwards from the basal crypt to the

middle; parallel-sided at the base then founded

and narrowed to the apex. Rlytra costate, the

intervals Mut ench with a shallow row of punc-

tures; more or Jess parallel-sided until the

middle then rounded gna tapered to the mar-

ginal spine: both spines well developed;

humeral fold moderately developed and angled.

Undersurface: thoracic sternites finely wnil

spatsely punctured in the middle more closely

at the sides; abdominal segmerits finely and

closely punctured; hairy.

Size. Males 10.5 + 010 x 39 + 0.04 mm

(47). Females 11.7 = fhtS & 4.3 + 0.06 mm

(34)

REVISION OF BUPRESTID GENUS ASTRAEUS LAPORTE & GORY 13}

Distrthution, Western Australia.

General remarks. In some species Wie marginal

ypot is reduced or lost. in others (here is an

additronal spout close uy the suture und between

the two fascia. Named nfter Mr R. P.

MeMuallan,

Specimens examined. W. Ausi.c Types; Soc & 4 9,

77 km alone main York Re on Casuarina sp.,

51.1968, §. Barker; tl & Dryandesa oo Casuarina

sp., 21.x.1970, §. Barker; 6 ¢ & & 2, Dryandra on

Casarina huegeliana, 11,.x.1970, §, Barker; 2a &

2°, 6 km E of North Bannister on C. Anvereliana,

19.xi,1870, S. Barkers 6 3 & 4 9, 77 km along

main York Rd on C. dinevelfany, ZIxLI9TO, S.

Barkers | d & 3 2 77 km along main York

Rd on C. huwegeliana, 7.xi1.1970, & Barker, Sa &

7 9, Dryandra on C. hweveliana, B.xii.1970, 5. Bar-

ders Lo dy Dryandra on C, divegellana, 12.1,1973,

§. Barker.

31, Astraeus (Astraeus) carnabyi sp. nov.

FIGS Id. 14, 23K

Types.

Molotype: 2, 16 km NE of Lake Grace, W.

Aust, on Casusrina fiuegeliana, 94,1972,

F. & K. Carnaby, SAM, 1 20947.

Allotype; 2, t6 km NE of Lake Grace, W.

Aust. on ©. /mepelivna, JLI87L. Be de KR.

Curmaby, SAM, 1 20948.

Paratypes. 1 4 & 1G 1h km NE of Lake

Grace, W. Aust. on C. hiaegeliana, 91.1972,

£ & K, Carnaby, ANIC. L 2, 16 km NE of

Lake Grace, W. Aust. on C. Awevelfana,

251.1973, BE. & K. Carnahy, RM: | df & 1 9,

16 km NE of Like Grace, W. Aust. on C.

huegeliana, E, & &, Carnaby, KC, 1 dg & |

9, 16 km NB of Lake Grace, Wi Aust. on

C. hucgeliana, 25.0.1972, &. & K. Curnuby,

MNHN (1 3), WAM (t?)]; 1-9, 24 km S

of Lake King, Wo Aust. on C. Jitegeliona,

25.1.)973, 5. Barker, MNHN; 4 da | Gy 14

km N of Needilup, W, Aust on C. hege-

Jian, 14.xit. 972, KT. Richards, WADA.

Colour. Shiny. Head, pronotum and elyira

black with purple reficctions; antennae

black with blue and purple reflections. Each

clytron with the following vellow markings: an

elongate spot at the hase reaching the anterior

margin; a fgscia commencing, at the anterior

lateral margin, covering the humeral fold then

tunning upwards tawards the suture but not

touching it. coneave forwards; a small fascia

just after the middle, commencing at {he mar-

gin and at right angles to the suture but reach-

ing only hill way lo it} midway belween the

two fascia are two small spots, one on the

margin and the other near the suture but not

touching it: a small preapical spot. Under-

surface metallic purple; hairs silver. Legs dark,

upper sides with metallic blue reflections,

undersides metallic purple.

Shape and senlpture. Head with medium sized

punctures on basal hall merging into regular

longitudinal chantiels en upieal purt; uo

median keel; hairy, Pronotum evenly pune

tured, with a median glabrous line at the base

ung apex; sides rounded gradually from bise

to apex; hairy. Elytra costate, Ue intervals

between flat, each with a row of shallow pune-

tures; parallel-sided until just after the middle,

then rounded off to the strong marginal spine;

well developed suturgl spine; humeral [old

poorly developed but slightly angled. Under-

surface evenly and shallowly puncttired, hairy,

Size. Males 9.8 + 0.17 x 3.7 + 0.06 mm (8),

Females 10.6 = 0.23% 4.07 0,11 mm (7)-

Distribution, Western Australia,

Ceneral remarks. Named after Mr K. Catnaby,

Specimens examined, Types only.

32, Astraens (Asiraeus) budeni van de Poll,

1889: 84, 93, 94, pl. 2. fig. 7, Fa. Black-

burn, I891: 496. Kerremans, 1892: 101.

van de Poll, 1892: 67. Blackburn, 1895:

45, 46. Kerremans, 1902: 148. Caster,

1929: 282. Obenberger, 1930: 365.

Amvacus badeni Var. disjuncinas Obenberoer,

7928; 204; 1930: 345,

Astraces meyricki Blackburn, 1890; 1254, 1257,

Keeremang, 1892; 102. Blackhurn, |895-: 45,

FIG, 23L

type. Holotype: ? MNHN (seen by author)-

Colour, Shiny, Head, pronotum and elytra

black with hlue reflections; antennae black with

blue reflections, tips of first and second sep-

ments brown. Elytra: each elytron with the

following yellow markings: a spot at the base;

a fascia commencing at the margin on the

humeral fold, running obliquely upwards and

backwards uid then at right angles to the longi-

tudinal axis of the bady, just before the middle

anil nat touching the sutare (this is frequently

broken to form two spots wide var. disjunetas

Obenberger, bul this has no taxonomic signifi-

cance); just after the middje there is a small

fuscia commencing at the margin, slightly con-

cave to the apex and not tostching the suture.

Undersurface and legs dark with metallic

purple reflections; hairs silver,

Shape and sevlpiure. Head evenly punctured;

no median keel; hairy, Pronotum with shallow

punctures larger at the sides than in the

middle; at the sides gradually rounded and

narrowed from base te apex: hairy. Elytra

costate, the intervals between flat each with i

row of shallow punctures; parallel-sided until

after the middle then rounded off lo the mar-

ginal spine. strong sutural spinc; humeral fold

veil

8. BARKER

5mm

ingi sp. nov.

Fig. 16. Astraeus goerl

Fig. 15. Astraeux carter? Sp. nov.

Fig. 14. Astraeus carnabyi sp. nov.

REVISION OF BUPRESTID GENUS 4s7RALUS LAPORTE & GORY

poorly developed but slightly angled, Under-

surface, plinctufes sparser in the middle than

at the sides; hairy.

Size, Males 8.8 + (tad x 3.4 + 6.07 mm (16).

Females 8.9 = 0,16 x 3.5 = 0,07 mm (17).

Distribwtion. All of mainland Austfalia except

the Northern Territory.

Geleral remarks, This species was described

almost simultaneously by van de Poll as A.

fwdent and by Blackburn as A, meyricki,

Blackburn (1891) recognised 4. meyrické as

a syhonym of A. tadeni, but later changed his

mind and called A. meyricki a good species

(Blackburn 1895). Distance between the twa

populations Was the main argument Used by

Blackburn (1895) in favour of calling the

castern and Western representatives pwo dis-

tinct species. I have been unable to separate

specimens collected in Western Australia from

those collected in South Australia.

Specimens examined, S. Aast.: Type\ 1 ad &l %,

Morgan, A, M. Lea, SAM: 1 2, Murray Brilee,

Oct. 1911, SAM; 8 d & 6 3, on Melalenra sp.

Dera Pass (probably the same as Pultapa Gap),

71 km S of Copley, 25.%.1969, No AveFarlaned,

SAM; 7 f & 9 G, Puttapa Gap. Flinders Ranges

on Melaleucu vlomeraia, 21.%.1971, 8. Rarker.

W. Aust: Paratype of A. meyrick? Blackburn,

SAM; 1 9, 18 km SW of Three Springs on Dry-

nadra cirsioides, 8.x.1968, N. McFarland, SAM:

3.5 & 3 8, Turanda rockhole, 106 km S$ of Balla-

dania an Colfitety preissti, 9.xii.1974, 5S. Barker.

NSW 1 2, W. du Boulay, WAM, 73-54. Olid:

Type 2 A. badent var, disintetus Obenberger,

NMP, 21 990,

33, Astraeus (Astraens) jansoni van de Poll,

[889; 84, 94, 95, pl. 2, fig, 8 Ba; Black-

burn, IS9L: 496, Kerremans, 1892: 102,

van de Poll, 1892: 68. Blackburn, L894:

101; 1895: 46, Kerremans, 1902: 148,

Carter, 1929; 282. Obenberper, 1930:

366, Carter, 1933; 42,

Astraens teppert Blackburn, 1890; 1258, L249

FIG. 23M

Type. Holetyps, MNHWN {not seen by author}.

Colour, Shiny, Head, pronotum snd elytra

bronze-green or black with green or purple

reflections: antennae black with blue or purple

reflections. Elytra: each elytran with the fol-

lowing yellow markings; along the margin, a

spot at the shoulder half covering the humeral

fold, a spot before the middle, a Fascia after

the middle running transversely towards the

suture bet not touching it; in the middle near

the suture buy not touching it, a spot at the

base. 7 spol betore the middlc, a spot at the

133

middle, a preapieal spot. Undersurface and Jegs

metallic branze-green; hairs silver.

Shape and sculpture, Head evenly punctured,

slightly excavated in the middle between the

eves; no median Keel: hairy, Pronotum evenly

punctured; a short longitudinal median

impressed line projects forwards from the basal

crypt; at the sides rounded basally, then

tapered and narrowed to the apex; hairy, Blytra

costate, the intervals between ilut, cach with

a deep row of punctures and faintly trans-

versely wrinkled: parallel-sided until after the

middle, then rounded and narrowed to the

Marginal spine; both spines well developed:

humeral fold poorly developed and slightly

angled. Undersurface with small shallow pune-

tures in the middle, larger and deeper at the

sides; hairy.

Size, Males 8,1 = 0,08 x 3.2 + 0.04 mmm (37),

Females $.6 + 0.15 = 3.4 = 0,06 mm (32),

Disiribution. Mainland eastern Australia from

South Australla to Queensland,

General reniarns. The yellow pattern is.

variable, there are cither six spots and a fascia

on cach elyiron or the fascia may be broken

in the middle giving a total of eight spots.

Specimens examined 8, Aust.) 2 9, MeDonald

Ferries N.P. on Cuilitriv pretssil, 2.%i 4467, §, Bare

ker 1 J & 1 9, 2 kim E of Hartley on C. preissii,

15.53.1969, 5. Barker; 7 df & 7 9, McDonald

Ferries N.P. on C.. preissii, 15.x1.1969, S, Barker;

2¢&2 9%, on road N of Parra Witra N.P. on C,

prelssi 10.11.1969, S. Barker: 7 & & 11 B, Tothill

anges near Brady Creck on C. preivsit,

12.xi1..196%, §. Barker’ | 2, Onkaparingn Gorge

near Hackhum on Callitris — rhomboideus,

27.21.1969, 8S. Barker; tl dg & 5 3 Mt Remark-

able N.P. on C. preisdi. 30.01,1969, 8 Barker;

24% 2 9 Alligator Gorze N.Po on C. preessii,

30.x10.1969, § Barker; § o& 2 2, Mt Remarkable

NLP. on ©, preissti, 72.%.1971, 3 Barkers 1d,

MeDonald Ferries N.P. on C. preisni, 14.%7.1971,

S. Barker; 2 ¢d. 16 km N of Mannoum on C.

preisyl, 20,5).1971, S. Barker; 6 & & 7 §. Salter

Springs on C. pretssii. 244x,1972, §. Barker, 4

fo & 3 2 Alligator Gorge N.P. an ©. preissii,

B.x. 1972, §. Barkers \ &, 10 km W af Penneshaw,

Kangaroo 1. on -C. preissti, 24.%1,1972, 8. Barker:

1q& / 9, near Rocky Point, Kangaroo 1. on C.

preixsit, 21.%,1974, S. Barker. Olds 7 2, Stan-

thorpe, &, Sutton, OM

34, Astraens (Astraeus) oberthuri van de Poll,

B89: $5, LOO, 101, pl. 3, fig. 13, 13a.

Kerremans, 1892: 102, Blackburn, 1892:

211, Kerremans, 1902: 149. Carter, 1929-

282, Obenberger, 1930; 36f,

FIG. 23N

Type. Holotype 3, MNAN (seen by suther),

Coleur. Shiny, Head, antennae and pronotum

134

black with purple and blue rellections, Elytru

black with purple reflections, cach with the

following Yellow markings: a busal spot: a

fascia commencing at the shoulder, covering

the humeral fold and running tewards the

suture but not touching it, slightly concave

towatds the base; a) spot al the middle touch-

ing the margin; a preapical spol, a spot mid-

way between the previous two which may or

may not touch the margin: # spot midway

he(ween the fascia and the preapical spot,

near the suture but not touching m1, absent in

some specimens, Undersurface and legs metal-

lic purple: hairs silver.

Shape and sculptare. Head closely and evenly

punctured at the base and sides, punctures

coalescing and wrinkled at the apex; ne niedian

keel; bairy, Pronojum closely and evenly punc-

tured in the middle. punctures coalescing and

wrinkled jl the sides; a short median longi-

tutinal impeessed line projects forwards from

the basal erypl: paralicl-sided from the base

unt! after the middle. then rounded and

narroweil ta the apex, trent edge projecting

slightly in the middle: dorsally convex in

Jateral profiles hairy. Elytra. costate, the inter-

vals between flail, each with a row of punctures

and faintly transversely strigose: parallel-sided

fo the middle then rounded and tapered to the

small marginal spine; sutural spine well

develuped; humeral fold poorly developed and

slightly angled. Undersurface with shallow

punctures closer at the sides than in the

middle; hairy.

Size. Males 9.7 = 0.10 x 3.6 £ 0.04 mr (58).

Females 9.8 = 0.34 «3.4 = 0-15 min (18).

Distribuon. Western Austfalia.

Specimens evamined. Wo Aust: Type; a ct. Yan-

chep, 71.7962, F. A. Uiiver Boker; 3 9, Goysnells

on Casuariwe sp.. 7.4.0967, §. Barker; 19 @ & 1 2

8 km W of Reverly 1.0. from Hrookton Rd, on

Casvering $p.. 14.1967, 8. Burker: Loa, 77 km

along main York Rd, on Casuarina sp., 111968,

S. Barker, & & & 1 2, Red Hill Rd, neue Midland

Junction on Casuarina sp., 41.1968. 8. Barker; 2

3, 77 km along main York Rd, .on Casuarina sp.

5.1.1968, 8. Burker; 6 3 & 2 2, 77 km along main

York Rd, on Casuarina huegeliana, 7,xi.1970, S,

Barkers 32 & 2 4, 7 ko B of North Bannister on

C. Auegeliana, '9.4i.1970, §. Barker; & J & 9 2,

14km F oof North Bannister on Co dbeevelianes,

19.51.1970, S. Barker; 5 o & 3 G77 kim along

main York Rd. on C. hwegeliana. 21.x3.1970, 8

Barkers | 4, 77 km along man York Rd, on

C. fieveliana, T.xiil970, S. Barker; 3 Fd & 2 2

Wannamal ow C. Auegellana, Wx 870, S. Bere

kers 2%, 135 kin slong Albany Highway on CC.

fegeliana, WA1973, S. Barker.

S. BARKER

35. Astraews (Astracus) watson! sp, Nov,

FIG, 13

Types:

Holotype; 2 Renter, W. Aust. é.xi.19$7-

RP. McMillan d& J. ALL. Watsou, WAM,

TI-\7é6l.

Paratynes: 1 9, ANIC; 1 9, Goomalling, W.

Aust, SAM; | 9, W. Aust. SAMs 1 & Bor-

den, Wo Aust. 16.xi. 1957, 2. PL MeMillan

a. AL. Watson, MNAN; | 9, Borden, W

Aust, 16.x11.1957, 2, Po MeMillan & J. AL.

Watson, WAM, 71-1760.

Colour, Shiny. Head, pronotum and clytra

black with blue and purple reflections; anten-

nac black with blue und purple rellections., the

base and apex of the first segment and apex

of the sccand seyment hrown, Jilytra; each

elytron with the following yellow markings; a

large basal spot; a broad lascia commencing

at the margin and covering the humeral fold,

running transversely towards the suture bute

broken in the middle forming a large spot near

the suture bul not touching it: after the middle,

a broad fascia concave butkwartds and nat

touching the margin or the suture: a stmall

preapical spot, a smull spot between the fascia

near the margin but oot touching it. Under-

surface black with blue and purple reflections;

hairs silver, Legs brown: top edges of the

femur and top surfaces af the tarsi dark brown.

Shape and seulprure. Head shallowly hut

evenly punctured; with a median keel; covered

with very long hair, Pronotum with punctures

evenly dispersed; # medion longitudinal

impressed line projects forwards from the

basal crypt to the middle; sides gently rounded

and narrowed from base to apex; haity. Elytra

costale, the intervals between convex at the

basal half, flat at the apical half, each with a

row of shallow punctures und slightly trans-

versely wrinkled, parallel-sided until just

before the middle, tapered gradually to the

marginal spine which is moderately developed;

strongly developed sutural spine: hurneral fold

moderately develaped and angled, Undersw-

face evenly and shallowly punctured; hniry.

Size, Perales 12,9 7 0,73 x 4.6 + 0.30 mm

(6).

General remarks. Named after Dr I, A. Le

Watson,

Specimens exaviined. Types anty-

3h. Astraeus (Asfrueus) carteri sp, nov.

FIGS 15, 230

Types,

Hiclalyre: 3, 383 km along Payne's Find Rd,

W. Aust., an Casuarina dtelsiana, IT AR AITUY

5. Barker, SAM, 1 20049,

REVISION OF BUPRESIID GENUS 4SiPRARUY LAPORTE & GORY

Allotype: &, Lake King, W. Aust, [8.xi/-1970,

Bo d& K, Carnaby, SAM, T 20950.

Paraiypes: 3 of, 383 kms along Payne's Find Ra,

W. Aust, on C. dieleana, 17.ix.1970, S- Bar-

ker, ANIC (1 9), BM ti 4), MNHN tL dds

3 ¢6& | &, Borden, W. Aust, On Casrarina

glaneu, 16.4,1939, H.W, Brawn, ANIC (1

9), WAM (3 of), WAM, 73-8870: 2 2, Lake

Cirace, W. Aust., MNHN a 2) WAM [1 9}.

WAM, 73-66; 3 a, Lake Graces, Hw

Brown, WAM, 33-735/6, 73-45; 1g & 1 2,

Southern Cross. H.W. Brows, WAM, 73-67,

73-63; 2 ¢@, Tallering Station, Pindar on

Casuarina Sp,, 22.in.1988, S. Barker, WAM,

73-72/3,

Colour. Shiny. Head black with purple reflec-

liens; antennae black with blue-green and

purple reflections. Pronotum black with blue-

ercen refleclions on top and purple reflections

at the margins. Elytra black with purple reflec-

tions with the following yellow markings: a

spot al the base; a spot originating at the

shoulder coyvenng the humeral fold; a spot

abuve the middle touching the outer murgin; #

spot belaw the middle, near the outer margin

but not touching it; three spots clese to the

suture but not touching it, the first the largest

above the middle, the next smaller below the

middle and a longitudinally clongate preapical

spot (the first of these sometimes coulesces

with the spot covering the humeral fold to

farm a fascia, concave towards the base).

Undersurface and legs dark metallic purple;

hairs silver,

Shape and serdprare. Hea closely punctured;

no median keel; hairy. Pronotum evenly punc-

tured; with «# short median longitudinal

imptessed line projecting Forwards from the

basal crypt, running forwards from the

impressed line is a glabrous line formed by

lack of perforgtions, better defined tn females

where it reaches the anlerior margin than in

males where i runs only ro the middle: sides

rounded and narrowed to the strong marginal

apex; dorsally Nattened in lateral profile; hairy

at the sides but Jess so in the middle, Elytra

custate, the intervals between flat with a tow

of shallow punctures; sides at first diverging

slightly outwatds from the base then parallel-

sided to beiore the middle, then gradually

rounded and narrowed to the strong marginal

spines; well developed sutural spines: humeral

fold poorly <leveloped but slightly «angled.

Undersurface finely and sparsely punctured in

the middle more closely at the sides: densely

hairy,

Size. Males 112 + 0.16 «x 44) + 01.05 mm

135

(13). Females 12.0 + 0.924 4,3 = O40 mm

{3}.

Distetbusion. Western Australia.

Genera! remarks. A. carter? shows clusest

aflinity with the following group of species; A,

macmillani, A, carnubyi, A. badeni, A. fansoni,

A. oberthurt and A. wetsant. Named after the

late Mr H. J, Carter,

Specimens exuminead. Types only,

37. Astracus (Astraeus) goerlingi sp, nov,

FIGS 16, 23P

Types.

Holotype: gd, Marloo Stn, Witrargs, W.. Attst.

1931-1941, 4. Goeeling, ANIC.

Allotype: 9, Marloo Sin. Wurargu, W, Aust,

1921-1941. 4. Goerling, ANIC

Paratypes: 4 ¢ & 3 9, Marloo Stn, Wurarga,

W. Aust. 1931-1941, 14. Goerling, ANIC

2&1), BM (1 0&1 2), SAM (1 2).

WAM (Eo & 19); 3 9, 106 km 8 of Payne's

Find, W. Aust., on Casuarina acutivalvis,

remedy S, Barker, SAM; 2 ¢ & 1 Y

Wurarga, W. Aust. on. Casuarina principiana

16.1%,193%, A, W. Brown, NM

Colour, Shiny, Head, antennae and pronotum

black with bronze, blue or purple reflections,

or a combination of these colours. Elytra black

with purple reflections, each with the following

yellow markings: a basal spot; a fascia com-

mencing at the shouldet and covering the

humeral fold. running towards the suture but

not touching it, concave towards the base; a

spot ut the middle touching the margin; a pre-

apical spot; a spot midway hetween the twa

previously mentioned marks, elongate and

touching the murgin; a spot near the sulure,

bul not touching i, midway hetween the fascia

and the prewpical spot, Undersurface and legs

purple, coppery and bronze: hairs silver.

Shape and sculpture. Head closely and evenly

punctured, slightly excavated in the middle

hetween the bases of the eyes; no median keel;

hairy, Pronotum with punctures larger at the

sides than in the middle. median longitudinal

glahrous line from base to apex formed by the

absence of punctures; gently rounded at the

sides from base to apex; hairy. Blytra costale

at the apex, punctate-striate at the base, the

intervals between flat ut the upex and convex

at the base, each with a row of punctures;

parallel-sided to the middle then rounded and

narrowed to the marginal spine, which 15 well

developed: sutural spine sharp but shortened

by the sutural margin being straight and super-

ficially appearing to be broken; humeral fold

poorly developed and slightly angled, Under-

surface evenly punctured, the punctures

136 ae, 4 «0 BARKER

Fig. (7. Agtraens major Blackburn

deeper at the sides than in the middle; hairy.

In the male the last abdominal sternite has. u

marginal indentation in the centre.

Size. Males 10.8 = 0,12 % 3.8 = 0.05 mm

(26). Females 11.7 © 0.19 x 4.2 = 0.07 mm

(22),

Distribution, Western Australia,

General remarks. A. goerlingl shaws features

in common with the preceding group of species

(Fig. 23P) and also with the following group

of two species, andl because of this [ place it by

itself berween the two groups. Named after the

late Mr A, Goerling,

Specimens examined. W. Aust.: Types; 20 4 & 14

Rane ee Stn, Wurarga, 1931-1941, A, Geerling,

ANIC.

38, Asteaews (Astrueus) ¢yaneus Kerremans,

1900; 295; 1902: 148. Carter, 1929: 282.

Obenberger, 1930: 365.

FIG. 19

Type. Holotype: 3, Standing, N.S.W., BM. tseen

by author).

Fig. 18. Astraeus watsoni sp, nov,

Colour. Shiny, Head and pronotum blue-green:

antennae black with blue reflections, Elytra

black with blue-green reflections, cach witb the

following, yellow markings: a Jarge elongate

basal spot not reaching the unterior margin or

suture; a fascia at the middle, expanded

towards the apex near the lateral margin but

nor touching it.or the suture; a large spot after

the middle, not touching the margin or the

suture, a Spot covering the humeral fold; a

small elongate spot in the form of a lunetic

near the preapical margin and ending at the

Marginal spine (present in the illustrated speci-

men, absent an the holotype). Undersurface

blue-green: hairs silver. Legs metallic blue,

Shape and sculpture, Head evenly punctured:

deeply excavated between the eyes, muinty al

the base: no median keel; sparsely covered

with long, fine hair. Antennae strongly serrate,

Pronotum deeply punctured at the sides,

towards the middle shallow punetures, with aj

central ovoid area consisting of hexagonal

RFVISION OF BUPRESTID GENUS 4STRAEUS LAPORTE & GORY 137

Simm

Fig. 19. Astraéevs cyaneus Kerremans

depressions, cach with a small central punc-

ture; inflated at {he sides just before the

middle, then straight sided and strongly

tapered to the apex; the lateral lobes with their

apices turned downwards; convex at the apex.

flattened at the base; covered with fine hair.

Flytra flattened; pimctate-striate anteriorly.

costate posteriorly, the intervals between con-

vex towards the base and flat at the apex, each

with a single longitudinal row of shallow punc-

tures and slightly transversely wrinkled; paral-

Jel-sided to the middle, then rounded and

narrowed to the small marginal spinc: sutural

spine shortened by the sutural margin being

straight and turned slightly upwards; humeral

fold poorly developed but slightly angled.

Undersurface evenly but shallowly punctured

in the middle: lateral prosternum and abdomi-

nal -sternites Jongitudinally grooved; sparsely

haired.

Size. Mules 11.6 x 3.9 mm (1). Females 13.9

x 48mm (1).

Divtelbution.. New South Wales and Queens-

land.

5mm

Fig. 20. Astraeus caledonicus Fanvel

Specimens exantined, N.S.W.c Type. Qld: | 4,

Acucia Creek via Killarnay, Jan. 1948, Mrs J.

Harstett, SH,

3Y. Astraeus (Astraeus) caledonicus Fauvel,

1904: 116, Obenberger, 1930: 365,

FIG, 20

Type. Holotype: 9%, Baie du Sud, N. Caledonie,

Delauney, MNHN [seeh by author).

Colour. Upper surface glabrous. Head and pro-

notum black with green reflections; antennae

purple, Flytra black with yellow reflections,

each elytron with the following yellow mark-

ings: a large basal spot; a small spot after the

middle near the suture but not touching 4; a

spot at the margin covering the humeral fold:

a spot after the middle at the margin but nol

touching it; and slightly behind the last a spot

near the suture but not touching it. Undersur-

face black with green and purple reflections;

hairs silver. Legs red-brown with purple reflec-

tions; tarsi dark-brown with blue reflections.

Shape and sculpture. Head shallowly but

evenly punctured: excavated between the cyes

mainly at the apex; no mediam keel; without

hairs. Pronotum shallowly but sparsely punc-

138

S. BARKER

Fig, 21. Astraeus robusius sp. nov

REVISION OF BUPRESTID GENUS ASTRAEUS LAPORTE & GORY 139

UO vu dogg

Odd

Fig. 22. Outline diagrams of the parameres of

male Astraeus (Depollus) species (A-H)

and Astraeus (Astraeus) species (I-T),

dorsal surface uppermost. A-—polli; B—

tamminensis; _C-robustus; D-aberrans;

E-lineatus; F—multinotatus; G-irregu-

laris; | H-dedariensis; \—bakeri; J—minu-

tus; K-—fraseriensis; L-—pygmaeus; M-—

smythi; N-simulator; O-obscurus; P-—

globosus; Q—mastersi; R-samouelli; S—

dilutipes; T-adamsi.

tured; a short but deeply impressed median

longitudinal line projects forwards from the

basal crypt; parallel-sided at the base, before

the middle rounded and obliquely narrowed to

the apex; median lobe short and blunt, apices

of lateral lobes sharp and turned downwards.

Elytra punctate-striate, the intervals between

convex with a few faint transverse wrinkles at

the base, without hairs or punctures; parallel-

sided until after the middle, then gently

rounded to the strong marginal spine; apical

Ogg

boo

109006

K L M N

eImmo

Fig. 23. Outline diagrams of the parameres of

male Astraeus (Astraeus) species, dorsal

surface uppermost. A-intricatus; B-—

crassus; C—major; D-navarchis; E-

fraterculus; F—prothoracicus; G-—elonga-

tus; H—vittatus; 1—flavopictus; ay ace

lani; K-carnabyi; L—badeni; M-—jansoni;

N-oberthuri; O-carteri; P-goerlingi.

spine with a straight internal edge; humeral

fold rounded and barely obvious. Undersurface

shallowly and sparsely punctured; lightly

haired.

Size. Females 11.5 x 4.1 mm (1).

Distribution. New Caledonia.

General remarks. A. caledonicus shows close

affinity with A. cyaneus.

Specimens examined. Type only.

140 S BARKER

Fig. 24. Scanning electron micrograph of

the median Iobe of the pronotum

Fig. 25, Scanning electron micrograph of the

head of A. fraseriensis showing the

of A. fraseriensis showing the median longitudinal keel.

basal crypt.

Index

To THE VALID SPECIRS AND SYNONYMS OF goer Fig at WITH THE NUMBERING SYSTEM ApoPTED IN

THE TEXT

Valid Species

aherrans van de Poll 4

adamsi sp. nov. 20

badeni yun de Poll 32

bukeri sp. nov. 9

caledanicus Fauvel 39

varnabyi sp. noy. 31

carieri sp. nov. 36

crassus van de Poll 22

cyaneus Kerremans 38

dedariensis sp. nov. 8

dilutipes yan de Poll 19

elongatus van de Poll 27

flavopictuy LaPorte & Gory 29

[raseriensis sp. noy. 11

fraterculus van de Poll 25

globosus sp. nov. 16

voerlingi sp. nov. 37

intricatuy Carter 21

irregularis van de Poll 7

jansoni van de Poll 33

lineatus van de Poll 5

macmillani sp. nov. 30

major Blackburn 23

masterst Macleay 17

Synonyms

= meyricki Blackburn

— strandi Obenberger

= lepperi Blackburn

= splendens van de Poll

= simplex Blackburn

minutus sp. Wov. 10

maltinotatus van de Poll 6

navarehis (Thomson) 24

oberthuri van de Poll 34

obscurus sp. nov. 15

polli sp. nav. 1

prothoracicus van de Poll 26

pygmaeus van de Poll 12

robusils sp, NOV. 3

samonelli Saunders 18

simulator van de Poll 14

smythi sp. nov. 13

famminensis sp. nov. 2

virtatas van de Poll 28

watson sp, nov. 35

Acknowledgments

1 would like to thank the following people

for their assistance: Mr G, Gross, Dr E.

Mathews and Mrs B. K. Head, South Austra-

lian Museum; Dr E. B. Britton, C.S.LR.O.,

Division of Entomology; Mr K. Dahms,

Queensland Museum; Mr A. Neboiss, National

Museum of Victoria; Dr C. N. Smithers and

Dr D. K. McAlpine, Australian Museum; Mr

L. Koch, Western Australian Museum; Mr B.

Levey and Miss C. M. von Hayek, British

Museum; Monsieur A. Descurpentries, Paris

Museum; Dr G. A. Samuelson, Bernice P.

Bishop Museum; Dr J. Jelinek, National

Museum of Prague; Mr K. ‘T. Richards,

Western Australian Department of Agriculture:

Mr A. George, Western Australian Herbarium;

Mr €. Slater. Canberra: Mr E. E. Adams,

Edungalba; Mrs J. Harslett, Amiens; Dr F. H.

Uther Baker, Applecross; Mr R. P. McMillan,

Cottesloe; Mr and Mrs K. Carnaby, Wilga.

Dr K. Bartusek, Dr S. J. Edmonds, Dr E.

Wollaston, Miss R, Altmann, Miss B. Jones

and Mr P. G. Kempster all of the University of

Adelaide, The National Parks Board of

Western Australia for permission to collect in

Flora Reserves; the Director, National Parks

and Wildlife Service of South Australia for

permission to collect in National Parks. The

Royal Society of South Australia provided a

grant to cover the cost of one illustration. The

Australian Biological Resources Committee

provided a grant-in-aid of research.

REVISION OF BUPRESTID GENUS ASTRAEUS LAPORTE & GORY 14|

References

Barker, S. (1964) —Asthraeus LaPorte and Gory,

1837 (Insecta: Coleoptera): Proposed emen-

dation to Astracus, Bull. zool, Nomencil. 21,

306-307,

Bi.ackpurN, T. (1890)—Notes on Australian

Coleoptera, with descriptions of new species,

part v. Prec. Linn. Soc. NSW, 4, 1247-1276.

Buackutrn, T, (1891).—Notes on Australian

Coleoptera, with descriptions of new species.

Proc. Linn. Soe. N.S AW. 5, 479-550.

Bracksurn, T. (1892).— Further notes on Aus-

tralian Coleoptera with descriptions of new

genera and species. Trans. KR. Sac, S. Ausi.

15, 207-261.

BLAackBuRN, T. (1894).—Notes on Australian

Coleoptera with descriptions of new species.

Prac. Linn. Sac. N.S.W . 8, 85-108,

Biacksurn, T. (1895).—Further notes on Aus-

tralian Coleoptera, with descriptions of new

Benera and species, Trans, R. Sac, 8. Aust

19, 27-60.

Britton, FL B. (197(),—Coleoptera. J “Tnsects

of Australia’. Ch. 30. (Melbourne University

Press: )

Carter. H. J. (1925).—Revision of the Austira-

lian species of Chrysobothriy (Fam. Bupres-

tidae), together with notea, and descriptiors

ot mew species of Coleoptera. Proc. Linn,

_ Soc, N.S.W. 50, 225-244.

Carter, H, J, -(1929)—A check list of the Atts-

tralian Buprestidae. Ausx!. Zoal. 5, 265.304.

Cartier, H. J. (1931)—Check list of the Atus-

tralian Buprestidac, Corrigenda. Ansi, Zool.

6, 107-108, ;

Carter, H. §. (1933)—"Gulliver in the bush.”

(Angus. & Robertson: Sydney.)

Fauvrt, <A, (1904).—Faune analytique des

Coléoptéres de Ja Novuvelle-Calédonie, Rev,

d'Ent. 23, 113-208.

GEMMINGER, Dr. & DE Haron, B. (1869).—

“Catalogus Coleoptorum hucusque descrip-

torum synonymicus et systematicus.” Vol, 4,

(E. H. Gummi, Monachii,)

LC ZN. (1966).—ASTHRAEUS La Porte &

Gory, 1837 (Insecta, Coleoptéra): Validation

of emeéndation to ASTRAEUS, Bull. Zeol.

Nomenc.. 23, 269-270,

TIunorr, L. (1856).—“Dersuch einer Einfuhrung

in das studium der Koleoptern.” (Basel.)

Kerremans, C. (1885)—Enumeération de Bup-

reslides deécrits postericurement au Catalogue

de M. M, Gemminger & dle Harpld 1#70-

1883, Ana, Soc. ent, Bely, 29, 119-187,

KenremMans, C. (1892) —Calalogus synonymique

des GBuprestides décriis de 1758 & 189D-

Mem, Soc, ent, Belp, 1, 1-304,

KFuremMans, ©. (1900),—Buprestides nouveaux et

remarques synonymiques: nn. Sac. ent. Belg,

44, 282-351.

KerkemMans, C. (1902).—Genera Insectorum 72.

Coleoptera; Serricornia Fam, Buprestidae.

LacorpalreE, T.. (1857).—"Histoire naturelle des

Insectes.” Genera des Coléoptéres, Vol. 4.

(Librairie Encyclopédique de Roret: Paris.)

LaPortr, F, 1, & Gory, H. (1837).—“Histoire

Nulturclic et Iconogruphie des Insectes

Coléoptérces.” Tome | Monographie des spi-

locera, eurydera, nycteis, ennostus, Bupres-

tides (Chrysochroites et Agrilites). (P-

Dumenit: Paris, )

MaclLaay, W. (1873).—Noles on a collection of

insecta from Gayndah. Trans. ent. Sac,

NSW. 2, 239-318,

Masrers. G, (1871).—"Catalogue of the des-

cribed Coleoptera of Australia.” (Sydney.}

Masters, G. (1886).—Cuatalogue of the des-

cribed Coleoptera of Australia. Part JIE

Pree. Linn. Soc. N.S.W. 11, 1-106.

Ovenaprapr, J. (1928)—Opuscula Bupresto-

logica I. Arch. Naturgesch, 92, 113-224,

Onennercer, f. (1930),—Tn “Catalagus Coleop-

terus.” Vol. 12, Buprestidae 2, pp. 365-367.

QBENHERGER, |. (1936)—Festschrift zum 60

Gebuctst Embrik Strand, Riga’ Vol, |, pp.

97-145,

VAN DE Pouce, J. R. H, N. (1886).—Description of

three mew species and a synopsis of the Bup-

restid genus Astruews, C. & G. Notes Leyden

Mus. 8 176-180.

van DE Pou, J, R. H. N. (1889).—Monographi-

cal essay on the Australian Buprestid genus

Astraeus C. et G. Tjselir, , Ent, 32, 79-110,

vaN DE Poi, J. R, H. N, (1892).—Noite sur quel-

ques especies d“Asiraeus, Tjschr. v. Ent. 3h,

67-68.

Saumpers, B. (1868). A revision of the Aus

tralian Buprestidae described by the Rey,

F. W. Hope. Trans. ent, Soc. Lond. 1868,

1-68.

Saunpers, E, (1871) —“Catalogus Buprestidarum

synonymicus et systematicus.”

THomson, J, (1856),—Description de dix Coléop-

téres, Rev, Mag. Zool. 2, 112-118,

FACTORS AFFECTING THE DISTRIBUTION OF THE LEPTODACTYLID

FROG GEOCRINIA LAEVIS IN THE SOUTH-EAST OF

SOUTH AUSTRALIA

BY R. 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 (Ginther), 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

GEOCRINIA LAEVIS IN THE SOUTH-EAST OF SOUTH AUSTRALIA

by R. G. BEck*

Summary

Breck, R, G. (1975).—Factors affecting the distribution of the leptodactylid frog Geacrinia

luévis in thé south-east of South Australia, Trans. R. Soc, S. Aust, 99{3), 143-147, 30

Angust, 1975.

Geacrinia laeviy (Gonther), a species of leptodactylid frog previously recorded from

Victoria and Tasmania, was first reported in Sonih Australia in 1966, A survey hus revealed

that this new population is an extension of that known to exist in south-west Victoria. The

frog ig extremely rare in South Australia, occupying only a fraction of fhe potential habitat

for which it shows preference. Some thoughts on this are tendered as a basis foc further

Studies,

Tutroduction

The Jeptodactylid frog Geocrinia laevis

(Gunther) has been described in detail by

Littlejohn & Martin (1964). The specimens

found in the south-east of South Ausiralia

have been up to 25 mm long, with dorsal

skin dark grey and slightly warty, ventral skin

smooth and paler grey, All had distinctive

pink markings on the groin and thighs and

some specimens showed this colour under the

forearms as well.

Prior to 1966, Geocrinia lnevis was. known

to exist in four disjunct populutions in Aus-

tralia: Tasmania, King Island, the Grampians,

and the area in south-west Victoria from

Dartmoor to Port Campbell (Fig. 1). With the

discovery (Woodruif & Tyler 1968) of a speci-

men from Marsh's Swamp near Mé Burr, some

80 km west of Dartmoor, it was desirable to

establish whether this was a fifth isolate or an

extension of the Victorian population,

Methods

The study area comprises the following

Hundreds in County Grey: Mt Muirhead,

Mayurta, Riddoch, Hindmarsh, Grey, Young,

Nangwarry, Mingbool, Blanche and Gambier,

and the adjoining area east of the state border

to the Glenelg River.

The survey was undertaken from 1968 to

1974, and most areas were visited in both

summer and winter. In the early stages of the

"Lynwood Park. Mil Lel, S. Anst, 5291.

survey, field work was concentrated around

Marsh’s Swamp (Site 3, Fig. 1. Grid reference

353362, Australian Army Survey Map, Penola,

1:250,000 Sheet SJ/ 54-6). From here the sur-

yey extended to the north-west, following the

general vone of influence of the Reedy Creek

drainage system, and to the south-east along

the Dismal Swamp complex to the Glenelg

River. Most field work was carried out during

the daylight, but night road surveys were con-

ducted in likely areas, yielding one specimen

only.

When six specimens had heen found,

detailed botanical surveys were made of the

surrounding arcas, particularly of the oearest

probable breeding site. Following the estublish-

ment of definite ecological patterns, soil sur-

veys. of ihese areas were undertaken.

Typical habitats

Geocrinia laevis normally lays eggs in areas

which later become flooded, Breeding sites

may be the edges of permanent swamps, or

non-permanent swampy areas, often situated

to the east of sandy Tises, ranging from a few

to 200 m away. The soils of the rises are

podsolised sands, usually Mt Burr sands as

described by Stephens et a/. (1941). The

swamps occur in Wandillo sands, and there

may be several intermediate soil types between

the rises and the swamps, For this reason, the

Natural dry sclerophyll forest may vary in type

144

R. &:. BECK

SOUTH. AUST.

e eg,

Millicent

% “*>. Tarpeena

Gambler

Kilometres

Fig. 1, Map of the lower south-east of South Australia and adjacent south-west Victoria, showing main

collecting sites of Geocrinia laevis in relation to the Reedy Creck-Dismal Swamp Corridor, ‘The

inset shows known Australian distribution prior to 1966. (After Littlejohn & Martin 1964,

1965.)

and consist of any of Eucalyptus baxteri, E.

obliqua, E. huberiana or E. ovata according to

the soil type. However, the understorey in the

vicinity of the swamps is remarkably con-

stant, and four species have been found in all

areas, viz. Acacia rmelunoxylon, Leptospermurn

juniperinum, Melaleuca squarrosa and Heli-

thrysum denidroides.

The soil map of Stephens et a/. (1941) plus

the plant indicator species greatly facilitated

later survey work and most potential areas in

the south-east have now been examined,

Results

A total of 20 specimens of G. /aevis haye

now heen found in South Australia, and a

further 10 in Victoria between the State bor-

der and the Glenelg River, which, prior to

1966, represented the known westeri limit of

distribution. The species is cammon at Dari-

moor, where on one oceasion, 12 were found

under a log in the bed of the Glenelg River.

Details of the findings are given in Table 1.

Representative specimens have been lodged

with the S.A, Mttseum,

Of particular interest 1s the specimen col-

lected at Grid ref. 376346 (site 7, Fig. 1), in

a deep pil dug lo observe pine lree root growth

at the Forest Rescarch Station north of Mt

Gumbier. This site is two km from the nearest

potential breeding area, giving an indication of

the actual mobility of the species, which is

regarded as sluggish compared with other

local species.

The distribution is shown in Fig. !, and

with one exception corresponds with normal

dispersals from the Reedy Creek and Dismal

Swamp complexes. It is suggested that the

Canunda specimen (site 1, Fig, 1) is from a

community established from eggs or larvac

washed down one of the many man-made

drains which cross the area between the Milli-

cent Hills and the coast,

GEOCRINIA LAEVIS IN THE SOUTH-EAST OF SOUTH AUSTRALIA

145

TABLE |

Recorded distribution af Geocrinia laevis in south-east South Australia and rearby Victoria

Refer A.A.S, Man, Penola, 1:250,000, Sheet SJ /54-6

Grid Site No. No. of

Reference (Fig, 1) Specimens Collector Remarks

353362 3 ! H. Minchan & First S. Aust. specimen, SAM,

C. Taylor R8118

353362 3 rT 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. Kiem Marsh’s Swamp. One = specimen,

SAM, R10583

354364 3 1 F, Aslin 2 km NE of original site at

Marsh's Swamp

381351 6 1 C. Taylor Earl's, 16 km N of Mt Gumbier

381353 6 ! R. Beck Hein’s scrub, 17 km N of Mt

Gambier

378351 6 F. Aslin Telford's scruh

378349 6 J D. Kiem 2 km § of Telford's scnib

376346 7 1 D. Klem Forest Research Station, Soil Pit,

SAM, R13974

368355 5 1 D. Klem Hogarth's scrub

326360 | 1 J, Aslin Canunda Reserve, SAM R13975

342366 2 iT F. Aslin Night Road Survey, 5 km NE of

Millicent

361361 4 ] A. Rowley Lake Leake, SAM, R14199

Refer A.A.S. Map, Hamilton, 1:250,000, Sheet S1/54-7

414325 8 6 P. Roach 16. km E of State Border on High-

way No. 1, SAM, RLO780

414325 8 1 R. Beck Same locality

427324 9 Matiy R. Beck Common in Glenelg River 2 km

up and downstream from Dart-

moor

Refer A,A.S. Map, Portland, 1:250,000, Sheet SJ/54-11

428311 10 i F, Aslin East of Glenelg. River near Jones*

Lookout

409313 12 2 F. Aslin Lower Glenelg River

424310 11 2 J, Aslin Lower Glenelg River

The major geological and physiographic

features of the region have been described by

Sprigg (1952). The Reedy Creek and Dismal

Swamp complexes are separated geologically

by the Gambier Upwarp, and flow in wet years

to the north-west and south-east respectively.

However, the watershed gradient ts so gradual,

being only a few cm per km, that in extremely

wet years such as 1896 and 1946 there was an

almost continuously wet corridor from the

Kingston district to the Glenelg River. Even in

years of normal rainfall, swamps are close

enough to provide ready access for frogs to the

lower south-east from the Glenelg River:

The last occasion on which the Dismal

Swamp actually flowed was in 1946, emptying

into the Glenelg River just north and south of

Dartmoor by way of the Scott and Ardno

Creeks.

In the study area, G. laevis is restricted to

areas receiving an average annual rainfall of

700 mim or more, whereas in Tasmania and

Victoria it is found where rainfall is greater

than 500 mm (Martin 1967). If the species

in South Australia followed the Victorian rain-

fall pattérn, it would be teasonable to expect

the distribution to extend laterally about 100

km, Likewise, if it occupied all sites considered

suitable on the basis of soil and vegetation

patterns, an extended distribution pattern

could be expected to the extent of about 50

km.

146 RB, G,

Discussion

Threw tiets emerge from the survey:

J While it is ceriain that more specimens will

be discovered within and beyond the present

Known maige of distribution, @. laevis is

extremely rare in south-east South Australia.

Even at Marsh's Swamp, whete most speci-

mens have been found, I have not positively

identified its calls during the April-May-

June breeding season. when calls are cam-

monly heard at Darlmonr, Victoria. How-

ever, Woodruff & Tyler (1968) have re-

ported the recording of a mating call al

Marsh's Swamp.

. According to present known records, the

species occupics only a fraction of the poten-

tial habitat for which it shows preference

with respect to soils, Vegetation and tain-

fall,

3. G. laevin was always found under the shelter

of jogs, lither, or stones during the day. As

aA result, the specics has not been found in

wreus Cleared for agriculture or pasture pro-

duction,

Some suggestions for the reason for this

restricted distrinition are tendered as a busis

fer further work by someone wilh more time

and resources than the present author. Ruin-

full and associated weather patterns are

probably the major factors influencing the

spread of any frog species. In 1967, an extreme

drought was expetienced in the lower south-

east of South Australia. ‘The average annual

rainfall at Mc Gambier is 776 mm, but in that

vear only 402 mm fell. and unofficial figures

from the Dismal Swamp area were as low as

280 unck 331) mm respectively, In this. single

dry year. many of the loval swamps previously

considered permanent, dried up completely,

and most of the non-permanent swamps stayed

dry throughout the winter. As a result, none of

the spring breeders bred, and only u few of

the auiumn and winter breeders actually

spawned.

Since Crocker & Wood (1947) first presen-

ted cyidence for a recent arid period, many

workers have commented, and they are about

equally divided in their acceptance or rejec-

tion of the concept (Mulvancy & Golson 1971:

Littlejaho 1967).

Gentlli (1961) had already offered an

explanation for this diversity of opinion: “Aus-

tralia is a large land, spanning several major

climatic belts, and may have experienced dif-

ferent climatic changes im various parts af the

tab

BECK

continent al the same time.” This is suppaoned,

on a one year basis iat least, by an ¢xamina-

ion of the Commonwealth Bureau of Meteord-

logy rainfall figures from major Australian

muintand centres for 1967, Perth, Beoume,

Darwin, Cairns. Brishane and Sydney bad

greater than average rainfall, Geraldton and

Alice Springs wete only slightly lower than

average, whereas Adclaide and Melbourme

received approximately half their average

umount, The coastal strip from the hed of

the CGireat Australian Bight to cast of Mel-

bourne obviously was the worst uffected arew.

Churchill (1968) has shown that, in

Western Australia, during the past S000 years

there have been several fluctuations in climate

of sufficient magnitude to cause the replace-

ment of jurrah forests with karri and vice

versa. Similarly, recent work by Dodson

(1974) in the study area indicules variations

in climate, with relatively dry periods between

5000 and 2000 B.P., and again since 1300

B.P.

Gentilli (1972) also supports the concept

of changing climate. “It must be stressed that

climate. being the result of numerous variables

variously combined in spuce and time, can

vary, fHuctuate, oscillate, or just change.” It

follows that changes such as these must pro-

duce ¢qually dramatic changes in the local

fauna,

With pluvial conditions prevailing in

southern Australia during the Jast glacial

period, Bassian species extended their range

(Litdejohn 1967) and it is teasonable to

expeot that G. Jeevis occupied much of the

lawer south-east of South Australia. With the

retum of present-day weather conditions, ts

range would have decreased but not to the

limits found today. Tt is therefore suggested

that im the recent past, possibly ouch more

recently than that postulated by Crocker &

Wood (1947), there has been a period suffi-

ciently arid to cause the withdrawal of Cr.

Inevis ynd perhaps some other anuran species

to the more faveured parts of south-west Vic-

toria or even the Grampians. It nisi be

emphasised that in this context the term “arial”

is relative rather than absolute.

Reoecupation of the south-east of Sowh

Australia by G, Juevis would have occurred. by

way of the Cilenelg River and the Dismal

Swamp when weiter conditions returned. This

may have takes place even as recently as with

in historic times,

GEOCRINIA LARVIS IN TRE SOUTH-EAST OF SOUTH AUSTRALIA 147

Since the settlement of the southeast of

South Australia by white man, the area

reached its physically wettest state in 1896.

This information wus obtained from S.E,

Drainage Board records and alsa, sume years

ago, from old residents who remembered the

district during the nineties. They claimed “you

could row a boat across country from King-

ston to the Glenelg River", While this is no

doubt a slight exaggeration, it is surely signifi-

cant to the distribution of frogs,

Colville & Holmes (1972) attribute the in-

Crease of wetness during the second half of

the nineteenth ceritury to the clearing of

natural scrub. Subsequent widespread plant-

ing of pines and establishment of drainage

schemes have greatly reduccd surface waters

Very recent reoccupation of the south cast

distribution has been prevented by a combina-

tion of clearing the nutural hubitat, and the

drying out of the district by drainage, pine

plantations and the establishment of better

pastures. Tt is obvious that much more study

is required to explain satisfactorily this limited

distribution.

Acknowledgments

The uwthor gratefully acknowledges the

collection of specimens by the following mem-

bers of the South East Field Naturalists’

Societies: Mr and Mrs F. Aslin, Messrs D,

Klem, P. Roach, C. Taylor, and Miss A. Raw-

ley. Also thanks are duc to Mr M. J, Tyler and

Dr M, Littlejohn fur adyice und encourage-

ment, and to Mr D. Lewis from the 8.A. Dept.

of Agriculture for his, help with soil identifica-

tion. The Royal Society of South Australia

by G. /aevis as outlined ahove would account Inc. kindly granted $50 towards petrol

for the limited spread of the species. Further cxpenses.

References

Current, D. M. (1968)—The distribution

and prehistory of Evealypius diversicolor F.

Muell., E. margiteta Donn, ex Sm. and E,

calophylla R.Br, in relation to rainfall. Aust.

J. Bat. 16, 125-51,

Cocvitee, J. S., & Houmas, J. W. (1972),—Water

table fluctuations under forest and pasture

in a karstic region of Southern Australis.

Ifydrology 17, 61-80,

Crocker, R. L.. & Woon, J. G. (1947) —Some

historical influences on the development of

the South Australian Vegelational communi-

ties and their bearing on concepts and classi-

ee in ecoloxy. Trans. R, Soc. §, Aust. 71,

-136.

Despsun, J. R, (1974)—Vegetation history and

water Jevel fluctuations ut Lake Leake. south-

eastern South Australia, [. 10,000 Bw. to

Present. “vst. J. Bot, 22, 719-741.

GEnNTILLK, J. (1961) —Quaternary climates of the

Mopecratiites region. Aan. N.Y. Acad. Set 95,

65- .

ChN ia J, C1972) —“Australian Climatic Pat-

terns.” (Nelson: Melbourne.)

LimiLEroHnN, M. J, (1967).—Patterns of Zoo-

geography und Speciation in South-eastern

Australian Amphibia, fv A. H. Weatherley

(Fd.), “Australian Inland Waters and their

Fauna, Eleven Studies,” (Australian National

University: Canberra.)

Litttesgoun, M. J., & Marvin, A. A. (1964)—

The Crinja laevis complex (Anura;: Lepto-

dactylidae) in south eastern Australia, Ars.

ft. Zool, 12, 70-83.

Litttesoun, M. J., & Marin, A. A. (1965).—

The vertebrate fauna of the Bass Strait

islands: 1. The Amphibia of Tlinders and

King Islands, Proc, R, Sec, Vier, 79, 247-236,

Martin, A. A. (1967).—Australian Aouran Life

Histories: Some Evolutionary and Eeological

Aspects, Jn A. H. Weuatherley (Ed.), “Aus-

tralian Inland Waters and their Fauna, Eleven

Studies." (Australian National WUniversiry:

Canberra.)

Murvaney, D J, & Gorson, J. (1967).—

“Aboriginal Man und Enyironment in Aus-

tralia.” (Australian National University:

Canberra.)

Spricc, R. C. (1952)—The Geology of the

South-East Province, South Australia, with

Special Reference to Qnaternary Coastline

Migrations and Modern Beach Developments.

South Australia Department of Mines,

Bulletin 29.

StepHens, C. G., Crocker, R, L,, BUTLER, B., &

SmtrH. R. (1941)—A Soil and Land Use

Survey of the Hundreds of Riddach, Hind-

marsh. Grey, Young, und Nangwarry, County

Grey, South Australia. Coan, Set. fid, Res,

Aust. Bull, 142,

Woopwurr, D. S. & Tynek, M. TF. (1968). —

Additions to the Frog Fawna of South Aust.

Ree. S. Aest. Muy. 15(4), 705-709.

THE THALLUS AND SPORE DEVELOPMENT OF LOBOSPIRA

BICUSPIDATA ARESCHOUG (DICTYOTALES: PHAEOPHYTA)

BY TIKVAH EDELSTEIN* AND H. B. S. WOMERSLEY{

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 BICUSPIDATA

ARESCHOUG (DICTYOTALES; PHAEOPHYTA)

by TiKVAH EDELSFEIN* and H. B. S. WoMERSLEYT

Summary

EDELSTEIN, Tikvan, & Womers.ey, H. B, §, (1975).—The thallus and spure development of

Loboespira hicuspidata Areschoug (Dictyolales: Phaeophyta), Trans, R, Sec. 8S. Ages.

99(3), 149-156, 30 August, 1975.

The apical growth of Lobospira hicuspidata, release of tetraspores, and growth of the

Spores in culture to plants up to | cm agioss, are described. Both development of the axes

and growth of the sporelings is from a marginal row of apical cells, and the thallus is mono-

podialy developed. Lohuspira is therefore placed in the Zonarieze group of the Dictyotales.

Introduction

Labospira hicuspidata Areschoug is a dis-

tinctive brown alga referred to the Dictyotales.

It accurs from Nickol Bay, Western Australia,

around southern Australia to Eden, N.S.W.,

and around Tasmania (Womersicy 1967. p.

215) and is frequently abundant in regions of

moderate to strong water movement, from

just below low tide level to 35 m deep.

The alga (Fig. 24) is easily recognized by

its spirally twisted axes, with a phyllotaxis of

about 1/3, bearing laterals with. bicuspid, de-

terminate tramuli (Harvey 1858, pl, 34), and

with lower branches bearing recurved attach-

ment tendrils. Kjellman (1897, pp. 295, 297)

and Oltmanns (1922, p. 185) considered that

the thallus develops fram an apical cell, with

aympodial branching, and. Lohospira has thus

been considered as a member of the Dictyo-

teae. The sporangia, about 100 gm in diameter,

occur scattered over the thallus (Fig, 2B); they

are developed from cortical cells and sunken

in the thallus (Fig. 1B). Neither division of the

sporangia. nor release of spores has been pre-

viously reported. and release was only obtained

by the present authors on the one oceasion,

Sexual reproductive cells also have never been

observed. While Lohaspira has usually been

placed in the Dictyotales, and the Dictyoteue

(Womersley 1967, p. 215), its relationships

haye not been established.

This paper reports observations on apical

development, spore relcase and early yrowth

of the thallus, made in 1972. while the firs:

author was on leave at the University of

Adelaide.

Methods

Plants (ADU, A42264) were callected in

drift at Aldinga reef, South Australia, on 27

May, 1972, and transferred to the laboratory

in sea water. The specimens (Tig. 28) bore

mature sporangia, many of which releused

tetrads of spores. Fertile branches were placed

in a glass jar with Provasoli ES medium

(Starr 1971, p, 359) in a 15°C culture room.

and spores allowed to settle on slides. during

the next two days. On day 3 the slides with

attached sporelings. were transferred to petn

dishes ($5 cm in diameter), and germanium

dioxide at a concentration of 5 p.p.m. added

to the medium. Single sporelings frony the

slides were detached to be grown in free cul-

ture; they were washed several times in a well-

slide and inoculated into a new set of dishes

each with 15-18 gporelings. After 4 weeks,

cultures were maintained in SWM 3 medium

(Chen, Edelstein & McLachtan 1969).

As the sporelings developed, considerable

difficulty occurred with bacterial (and ut one

stage fungal) contamination, Addition of peni-

cillin to the Provasoli medium had liule effect,

but streptomycin (100-150 mg streptomycin

sulphate/| of seawater) eliminated most of the

bacteria, and a commercial fungicide, Myco-

statin-Dusting Powder (1,000,000 — units)

(E. R. Squibb & Sons Ltd, Melbourne) proved

* Atlantic Regional Laboratory, Nations! Research Council of Canuds, Halifax, N.S., Canada. [NRCC

No, 14511.)

y Department of Botany, University of Adelaide, Adelaide, S. Aust. 5000.

150 TIRVAH EDELSTEIN & H. B, 5. WOMERSLEY

lessee nse

<4 QT ISS.

‘i a = Sa ce

AVEDA sorte. Y 6 ,

& Shock el eS sf ase

re a acieee Bs Xe \ é A

th Oy ay. Lm, XC A,

i 2 KE 2 —y Y 4 q

eA ace

Fig. 1. 4. Apical development, with two young laterals present, the older one (right) hecoming hicus-

pid. The axis meristem (a.m.) continues growth of the branch, and the lateral meristems

(l.m.) may or may not develop further inte laterals (ADU, A42264).

B, Cross section of ramulus bearing sporangia (ADU. A42264).

C. Various sporelings 1 week ald, with a rhizoid 3-5 cells long and early stages of the “cell-

mass”.

J, 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 “Mycostatin” per 100 ml of sea ters of braiiches up to | cm across, but from

water for 20 hours. Repeated treatments with which it was impossible ta clean the epiphytes.

streptaemycin and Mycoslatin were used and In general, cultures were maintained at 15°C

frequent cleaning of the sporelings with glass under a light intensity of 610-800 lux provided

needles was carried oul, While this damaged by 40 watt white fluorescent lights, and a

some sporclings, the majority survived and con- regime of [4 hrs light/10 hrs dark. ‘The me-

THALLUS AND SPORE DEVELOPMENT OF LOBOSPIRA 151

4 its 4

;

152 TIKVAH EDELSTEIN & H. B. S. WOMERSLEY

THALLUS AND SPORE DEVELOPMENT OF LOBOSPIRA 153

‘2

Het

154 TIKVAH EDELSTEIN & H. B. S. WOMERSLEY

TRALLUS AND SPORE DEVELOPMENT OF LOKOSPIRA 155

Fig. 2 A, Thallus of Lehospira, showing basal entangled branches with tendril-bearing laterals, and

erect branches with ultimate bicuspid ramuli. Stanley Bench, Kangaroo 1, 5. Aust,

(Edelstein & Kraft 126, 25.iv.1972).. ; _

B. Surface view of a branch showing tetrasporangiay (undivided) and released tetraspores

(arrow), (ADU, A42264,)

Fig 3. 4. A multicellular sporcling, with rhizoid. about 2 weeks old. , t

BLA Plant about 3 weeks old, showing development of a flat thallus with 4 meristematic

margin.

Cc. Plant 6 weeks old,. with two lubes. ;

O Plant 6 weeks old, showing development of lobes from the margin and especially from

the rhizoidal region,

Fie. 4. A, Plant 7 weeks old, with 3 main lobes.

#. Plant 74 weeks old, becoming convolute.

C.. Plant 122 weeks old. with several irregular lobes. ’

®. Plant 174 weeks old, consisting of many irregular branches, each with. marginal lobes af

Various sizes,

Fig. 5, 4. Plant 244 weeks old: general view of meristematic apea, with degencrating cells in centre

of each lobe.

. As in 4, with two darkly-staining cells (arrows) separating in centre of margin of Iobe,

c. Asin A, with degeneration of cells between two darkly-staining cells.

dium was changed every 3-5 days for the first

6 weeks and thereafter at weekly intervals.

Results

Development of the thallus

Dissection of apices (Fig, 14) of mature

field plants of Loboxpira shows that develop-

ment is from a row of apical cells, from which

lateral groups of apical cells separate off alter-

nately and differentiate rapidly to form a

Targer, abaxial, spinous wr bicuspid process

with » proup of meristematic cells on its adaxial

side, Growth is thus monopodial and the affi-

nities of Lobospira ure with the Zonarieye, In

uctively growing upices, the young laterals

develop rapidly and overtop the apex. The

meristematic group of cells on the lateral muy

persist indefinitely. in which case a long lateral

develops: or il may persist ta give a short

lateral with only a few pointed ramuli: or it

may oot deVelop turther. resulting in only a

peinted or bicuspid ramulus, at the apices of

which a single cell remains prominent (Fig.

!4). Alternate series of these ramuli fringe the

longer axes or branches.

The indefinite uxes twisl spirally sa that the

branches. or ramuli become arranged with a

phyllotaxis of about 1/3,

The mature thallus consists of a cortical

layer of cells which are arranged more or less

in Jongitudingl lines and tend to fadiate up-

wards. The medulla consists of cells of similur

size but rather inegularly arranged (Fig. 1B),

and ts 2-5 cells thick; in older branches a

slight midrib is present where the medulla is

thicker. Older axes are ovoid to round in sec-

tion and a central core of narrower and more

elongate cells may be present. Hair groups are

o£ frequent occurrence on the thallus.

Sperangia and spore development

Division of the sporangia into four. ietra-

hedrally arranged, non-motile spores (Fig. 24%)

appears tg occur only shortly before their

Telease, und no divided sporangia have been

observed in any herbarium material. Fertile

collections have been made mainly in gutumn

(Apri! to June),

The spores germinated on slides within two

days of their release, forming a short rhizoul

which was cut off by a cross wall when slightly

longer than the spore. The rhjzoid became 3

or 4 cells long before the sporc-residual cell

enlarged and divided. By day 7, a variety of

cell arrungements (Fig, IC) was present

amongst the sporclings, which, during the next

2 weeks, developed into. clongate-ovoid masses

of cclls (Figs 12, 34), their arrangement

depending on the early cell divisions.

From this cel] mass, which was attached by

a felulively long rhizoid of several cells, a flat,

ovate-spathulate disc of cells developed, mostly

two cells thick and with a distinct apical raw

of meristematic cells (Fig, 38, plant 3 weeks

old). During the next 4—5 weeks, further rhi-

zoids developed from the basal cell mass, and

the flat, erect, frond developed further trom the

apical meristem, usttally becoming lobed (Fig.

3C), Smaller lobes developed both from the

136

basal cell mass and lower parts of the erect

fronds (Fig. 3D).

By & weeks, the plants had developed several

fronds (Fig. 4A) of varying sizes and often

the main frond was. becoming convoluted

(Fig, 48), Tufts of long, colourless hairs dif-

ferentiated at this stage of development. By 12

wecks, numerous fronds were present (Fig.

4C). usually branched or lobed, and thalli 174-

244 weeks old formed a cluster of fronds (Fig.

4D) up to } cm across. One plant reached

almost 2 cm across after 34 weeks but showed

no further morphological development. At this

stage, all thalli were heavily overgrown with

epiphytes and died. Apart from the meri-

stematic and lateral margins, the fronds were

mostly two cells thick, increasing to 3 or 4

eells thick in the older parts.

The apical marginal row of meristematic

cells was prominent in all branches and lobes.

giving a typical “zonarioid’ appearance (Fig.

5.4). Plants 244 weeks ald showed a furthet

apical development of possible significance, m

that centrally along the meristematic margin

of each lobe, two cells became more promin-

ent (Fig. 58) with denser protoplasm, and

breakdown of tissue occurred between them

(Fig. 54, C). Whether this was only a break-

down feature before death is uncertain, but the

two cells concerned, which when first notice-

ahle were densely protoplasmic and appeared

healthy. could possibly correspond to the single

TIKVAH EDELSTEIN & H. B. S. WOMERSLEY

cells which are prominent at the apex(ices)

of the single or bicuspid ramuli of the mature

plant,

Conclusions:

The division of sporangia to give four non-

motile spores, and the occurrence of a mar-

ginal row of apical cells in both the adult axes

and in juvenile stages, indicate that Lobospira

is correctly placed in the Dictyotales, but be-

longs in the Zonaricae and not the Dictyotcac.

The distinctive morphology of Lobaspira separ-

ales iL generically from all other genera of the

Zonarieae.

While the sporeling and juvenile stages ure

now known, further studies are necessary to

show how such stages develop to the mature

laterals which cut off pointed or bicuspid

ramuli, Since sexual plants are still unknown,

cytologcial studies on the division of the spor-

angia are desirable to indicate whether meiosis

accurs at this stage.

Acknowledgments

The second author acknowledges a grant

from the Australian Research Grants. Commit-

jee for technical help, and the assistance of

Mrs E, L. Robertson and Miss C, G. Anderson

with the culture work and ulustrations. Dr

G. T. Kraft assisted in carly stages of the

study.

References

Cuen. L. C-M,. Epesstein, T., & McLacuian, J,

(1969).—Bornemuaisonia hamifera Wariot in

nature and in culture. J. Phycol. 5, 211-220.

Hagvey, W, 11. (1858)—"“Phycologia Australica”.

Vol. 1, Plates 1-60.

Kutiiman, F. R. (1897).—Phoeaphyceae. /n A,

Engler and K, Prantl, “Die Naturlichen Pflan-

zenfamiglien™, Th. 1, Abt. 2, pp. 176-297.

OLTMANNS, F. (1922).—“*Morphologie und Bio-

logic der Algen.” Vol. 2. (Jena.)

Starr, R. C. (1971).—The culture collections of

alpae at Indiana University—additions to the

collection July $966-July 1971, J. Phycol. 7,

350-362.

WomersLeyY, H. B, S, (1967).—A critical survey

of the marine algae of southern Australia. 1,

Phaeophyta. 4ust. J. Bot. 15, 189-270.

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

Dulbunty, J. A. Shoreline Shingle Terraces and Prehistoric Fillings of Lake

Eyre - - - - - - - - - - 183

Latz, 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 - - - . “ - - ~ - - - = 238

PUBLISHED AND SOLD AT THE SOCIETY’S ROOMS

STATE LIBRARY BUILDING

NORTH TERRACE, ADELAIDE, S.A. 5000

HYDROIDS OF BRUNY ISLAND, SOUTHERN TASMANIA

BY JEANETTE E.. WATSON*

Summary

WATSON, Jeanette, E. (1975).-Hydroids of Bruny Island, southern Tasmania. Trans. R. Soc.

S. 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.

HYDROIDS OF BRUNY ISLAND, SOUTHERN TASMANIA

by JEANETTE BE. WatTson*

Summary

Warson, Jesnette, E: (1975).—Hydroids of Bruny Island, southern Tasmania. Trans. R. Soc.

S, Atist. 99(4}, 157-176, 30 November, 1975.

A systematic collection of the sublittoral hydroids of Bruny Island, southern Tasmania,

using SGUBA, 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 shelicred micro-

habitats, Few species of Sertulariidae are recorded, and Amphishetia 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 ave recorded for the first time from one

locality. Two species newly described, Halecium hruniensis and AH. luteum, ace each closely

related to endemic New Zealand species. The occurrence of these, and the first record of

Salacia farguhari outside New Zealand waters (where it also occurs south of 43°S) suggests

active progress of speciation and dispersal across the Tasman Sea,

Introduction

Bruny Island (43°25'S, 140°20'B) is situated

off the cast coast of Tasmania, 25 km south of

Hobart. The island is approximately 50 km

long and is separated [rom the Tasmanian

mainland by the narrow waters of the D’Entre-

casteaux Channel, The indented coastline of

Bruny Island provides a range ot environ-

mental conditions varying from the sheltered

but swift Mowing tidal waters of the D’Entre-

casteaux Channel to the rough-water eastern

coustine of Adventure Bay and Penguin

Island, open to the Tasman Sea,

Systematic collecting of the sublittoral

hydroid fauna was. undertaken during two

weeks in February. 1972. Sampling, using

SCUBA equipment, was carried out over the

entire depth range (0-20 m) presented by the

tocky subhttoral along the coastline at Satellite

Island, Great Taylor Bay, Simpsons Bay, the

adjacent D'Entrecasteaux Channel, and at

Fluted Cape, Penguin Island and Adventure

Hay on the eastern coastline of Bruny Island.

No collection was made of hydroids from

the littoral zone, as these localities comprise

either steep rocky cliffs facing the Tasman Sea

ev the sandy beaches of the D’Entrecasteaux

Channel, It is unlikely that either of these bio-

topes. would make any significant contribution

to the hydroid fauna of Bruny Island.

In his revision of the hydroid fauna of Tas-

mania, Hodgson (1950) listed 64 specics known

from the deep and shallow waters of the Tas-

Manian coast and Bass Strait. His list includes

22 species from the D'Entrecasteaux Channel

and the adjacent Derwent Estuary, and one

species from Adventure Bay,

The present survey yielded 34 species (in-

cluding two identifiable only to genus) and

2 varieties of one species. There are 11 new

records for Tasmania, including 3 new records

tor Australia, and 3 species are newly described.

Only 19 species of Hodgson’s list appear in the

present collection.

Holotype and paratye microslides of new

species, and other microslides and maternal, are

lodged in the National Museum of Victoria,

Melbourne (referred to as NMV).

No athecate hydroids were recorded from

Bruny Island. The Campanulariidae is repre-

sented by 5 species, Lafoeidae by 1, Halecimdac

6, Syntheciidae 1, Sertulariidae 9, and the

Plumulariidae by 12 species, including 2 yaric-

ties of one species,

LIST OF SPECIES

* Denotes a new record for Tasmania

+ Denotes a new record fot Australia

* National Museum of Victoria, Russell Street, Melbourne, Vic, 3000,

158 JEANRTTR, F. WATSON

THECATA

Family CAMPANULARIIDAR

* Campanilaria ambiplica Mulder & Trebilcack.

Campanularia pulcratheca Mulder & Trebileock.

Clytia sp-

Orihopyxixs celiculata Hincks,

Silicularia rosea Meyen,

Family LAFOEIDAE

* Hebella ?furax Millard.

Fumily HALECIIDAE

Halocinm delicatulam Coughirey.

Halecium 8p.

: Halecivin heanit (Johwston),

Halecium braciensis Wasp,

Halecium luteum nap,

Phylactotheca armita Stechow,

Family SYNTHECIIDAR

Syntheciune patulum Busk.

Furntly SERTULARIUDAE

+ Salagia farquhari (Bale).

Stereatheca elongata (Lamouroux}.

Sertularella robusta Coughtrey

Swnplectoseyplius pygmaeus Bale.

Sertularia acuta (Stechow).

Sertalaria macracarpa Bale.

Amphisbetia minima vat, intermedia Bale.

Amphishetia operculata (Linnaeus).

Amphishetia asia Tsp.

Family PLUMULARINDAE

Halicornopsis elegaris (Lamarck). ‘

* Anpenella campanuliformis (Mulder & Trebil-

cock).

Pyenotheca mirabilis var. mirabilis (Milman).

Halepteris canpanula var. campanila (Busk}.

Plumularia filicaulis Kirchenpauer.

Plumularia hyalina Bale.

Plumularia anpusta Stechow.

Plumtdaria crateriformis Stechow-

Plamutaria wilson’ Bate.

Axglaophenia plumosa Bale.

Thecocarpus divaricatus var. typica (Busk)-

* Thecocarpuy divaricatus var. hriegsi Bale.

Halicernaria langirostrix (ircheapauec),

ws tee

Systematic Section

amily CAMPANULARIIDAE

Cumpanularia ambiplica Mulder & Trebilcock,

1914a; pl, 2, figs 3, 4. Shepherd & Watson,

1970) 140,

Paravalix «mbiplica Stechow, 1925; 209, fig. E.

Record: Penguin I, on a ted alga and sponge

in crevices on rough-water side of island,

16-20 m deep.

Material; Colonies infertile. Hydrothecae with

7 fairly sharp teeth, the embayments hetween

wider than the teeth.

Remarkx: Although occurring on the rough-

water side of the island, this small delicate

species oceupicd a microhabitat in sheltered

crevices al a depth below turbulence due to

surac.

This is thy first record of C, qmbiplica from

Tasmania, Other localities: Victoria; Champion

Bay, W, Aust.

Campunuluria puleratheca Mulder & Trebil-

cock, 1914a: 11, pl. 2, figs 1, 2, Blackburn,

1942; 105.

Paracalix pulcratheca (Mulder & Trebilcock,

19140). Stechow, 1923a: 3

Record: Satellite I. (no depth recorded), on

red alga Delivea.

Material) Colonics infertile. Hydrorhiza tubu-

lar, Stems 0.83-1,33 mm long, 0.06-0,09 mm

diam... perisarc thiek, a spherule between stem

and hydrotheca, //ydrothecae long and tubular,

perisare thickening distally, a distinet dia-

phragm near base and a flexure almost two-

thirds the distance up the hydrothecal wall from

the base, Margin with 8-10) teeth. Diam. of

hydrotheca at margin 0.320.441 mim, depth to

diaphragm (including teeth) 0.77-0.92 mm.

Remarky: The Tasmanian material compares

well with the holotype of C, pwleratheca (in

NMV), although the present specimens have

fewer marginal teeth.

This is the first record of C, pulcratheca from

Tasmania. Other localities: Victoria; S. Aust,

Clytia sp.

FIG. 1

Record: Adventure Bay, 10 m deep on sfem

of Thecocarpus divaricatus var. typice.

Material: A few infertile stems. Stevts of vati-

able length, 0.70-1.90 mm, irregularly undu-

lated, in some places smooth. Mydrothecae

campanulate, expanding from base fo margin,

perisarc fairly thick, and a well defined dia-

phragm with a thickening of the thecal wall

helaw. Depth to diaphragm 0,30-0.40 mm. A

small spherule between hydrothecae and pedi-

cel. Margin 0.20-0.32 mm diam., with 12

bluntly pointed teeth, the embayments between

slightly wider than the teeth.

Remarks: ‘Vhe hydrotheca. of this relatively

small species corresponds in some. respects with

C. hemisphaerica (Linnaeus, 1767), but the

stem Jacks the typical proximal and distal an-

nulations of this species, In the ubsence of

gonosome it is not possible ta further identify

the material.

Orihopyxis caliculata (Hincks, [853). Bale,

1914b: 74, PL 11, fig. 1, pl. 12, fig. bs

HYDROIDS OF BRUNY ISLAND, SOUTHERN TASMANIA

1924; 232, Hodgson. 1950: 7, figs 14-16.

Shepherd & Watson, 1970: 146.

5 uinpunlarig calicwlura Hineks, 1853: 178, pl.

>, Ne, 3,

Euespelia caliculates (Hincks). Hirohite, 1969;

fig. 6.

Record: Penguin 1, 15-20 m deep. on a red

alga and on sponge in a crevice.

Materia: A tew fertile colonies. Nydrothecae

very shallow and expanding; hydrothecal pedi-

cels spirally annulated. a few with smooth

regions. Hydranth with 24 tentacles, Genotheca

smooth with thick perisarc, containing mature

fonophores.

Remarks: The specimens. from sponge possess

longer pedicels and have a thinner persgare

than those epiphytic on algae.

O. caticulate was not abundant at Bruny

Istand, occurring only in a sheltered crevice on

the rough-water side of the island at a depth

below major turbulence. ‘This accords wilh

previous findings on habitat preferences of this

species (Shepherd & Watson 1970; Watson

1973),

Silicularia rosea Meyen, 1834; 204, pl. 35. figs

1-11. Millard, 1968; 259.

Siliculavia: bilabiara (Coughtrey, (875). Ralph,

1957; 842,

breopella campanyhiria vou Lendenfeld, 1883,

Bale, 1888; 751, pl. 13, figs 9-15,

Siticularia campanulurla (von Lendenfeld,

TRS3), Hodgson, 1950: 6, fies 12. 13.

Record; Fluted Cape (no depth recorded) on

the brown algae Seirococeuy uxilluriy and Scvio-

thalia doryearpa.

Material: Luxuriant fertile mate and female

colonies on the algae, Developing gonephores

present,

Remarks: The present material conforms to

descriptions given by Ralph (1956, 1957) for

Silieularia bilabiata forma svbrrapica (demon-

strated by Millard (1968) (a be a synonym of

S. rosea Meyen), a form oceuting anly north

of the Subtropical Convergence, and typical of

the southern Australian coastline, including

Tasmania.

Family LAFOEIDAE

Hebella ? furax Millard, 1957: 200, fig. 8:

1964: 10, figs 2B-D. Millard & Bouillon,

1973: 59,

FIG, 2

Record: Penguin L, on stem of Thececarpus

divaricatus var. typiva, depth 20m.

Material: One smalt colony. Aydror}iza 3

loosely winding tube. Hydrothecae large, cam-

154

panulate, perisare delicate, smooth, Hydrotheca

asymmetrical, one side convex. the other

straight or slightly concave. this side always

inclined to the hydroid host. Margin entire, with

a thin slightly everted rim, Pedicel of variable

length, perisare thick. strongly undulated to

smooth, widening distally to pass inio base

of hydrotheea below diaphragm, Immature

hydrothecae truncated with a thin cap-like

vperculum,

Dimensions. (mm): South

Krony L. Africa

Hydrothecn—

depth to diaphragm 0,80-0,92 0.74-0,84

dian. at margin O.S0-0,68 0.55-0.64

diam. at diaphragm 14018 = 0,22-0.28

Pedicel .

length from diaphragm = 0.20-0.29 —-W,52-0),72

minimum dian. 0,05-0.08 0.08-0.10

Remarks: Comparison of the Bruny T. material

with paratype niicroslides of Mebella furay

Millard from False Bay, South Afnea (pro-

vided by Dr N. A. H. Millard) shows that

although similar in shape to the South African

specimens, the hydrothecae of the Tasmanian

material ure generally deeper and narrower at

the diaphragm, lack the distinct thickening of

the lower thecal wall, and have a mare pro-

nounced eyersion of the margin than the South

African specimens, However, in the absence

of gonophores it is difficult to determine the

specific status of hydroids of simple morpho-

legy, hence the present specimens are pro-

visionally assigned to H. furax,

This is a new record for Australia.

Family HALECHDAE

Halecium delicatulum Coughtrey, 18763; 29.

I876b: 26, pl. 3, figs 4, 5, Ralph, 1958:

344, figs 11, 12 {synonymy}.

Halecivm flexile Allman, 1883; 11, pl, 5, fiz. 2

Hodgson, 1950; 16, files 25-27.

Records: Adventure Bay, Penguin £., Satellite

L, on the kelp Macrocystis pyrifera, 2-T m

deep; Fluted Cape, 10 m deep, on Theeararpus

divaricatus var. typica (Busk),

Material: Luxuriant fertile. colonies. Stems ta

L cm Jong, simple and branched. Many hydro.

phores with marginal replications: secondary

hydrephores. arising trom the pedicel oi

primary hydrophores. Colonies dioecious with

mature gonophores ansing from proximal parts

of stem and on hydrorhiza. Distal parts of

blastostyle capshaped in both sexcs. Colour,

trophosome yellow, gonophore bright orange.

Remarks: Hodgson (1950) described and

figured A, flexife Allman (= H. delicatulum)

160)

Fig,

Fig.

1. Clytia sp, Hydrotheca.

JEANETTE FE. WATSON

0-5 mm

2, Hebella ?jurax Millard. Hydrotheca epizoic on Thecocarpus divaricatns.

Figs 3,4. AHalecium sp. Fig. 3—Whole colony, Fig. 4—Distal part of colony enlarged.

Figs 5,6. Hulecium beanii (Johnston). Fig.

phores.

from Eaglehawk Neck, but did not record the

substrate.

The colonies of H. delicetulum im the pre-

sent collection, while yery abundant on Mucro-

cystis and other brown algae, were, however.

strictly epizoic, always growing on the surface

of the crustose bryozoan Membrinopora mem-

brinacea, a common epiphyte on the aging

fronds of Macrocystis.

Halecium sp.

FIGS 3, 4

Record; Penguin 1, on a red alga in sheltered

water, no depth recorded.

Material: One infertile stem. Sfem 3 mm high,

lightly fascicled at the base, branching irregu-

larly sympodial. Stem internodes of variable

length, 0.30-0,40 mm, narrow, perisare fairly

5,—Whole colony. Fig. 6—Part of stem showing hydro-

thick, with one proximal annulation and a distal

apophysis giving rise to the suceceding intcr-

node, Hydrophore fairly deep, expanding to

margin, slightly asymmetrical, adcauline wall

more expanding than abcauline wall, Depth to

diaphragm, 0,06-0.07 mm, depth to base of

hydranth, 0.05-0.06 mm. Margin 0.12-0.14

mm diam., with a distinct outwardly rolled rim.

Diaphragm concave, approximately 0.0] mm

below line of attachment of hydranth, usually

a strong thickening of the inner wall immedi-

ately below diaphragm, best seen in older

hydrophores, often absent in younger terminal

hydropheres, Punctae not visible; if present.

obscured by hydranths. Pedicels of hydrophores

of variible length; shorter pedicels usually

annulated, longer pedicels smooth. Hydro-

phores regenerated 1-3 times, regenerated pedi-

HYDROIDS OF BRUNY ISLAND, SOUTHERN ‘TASMANIA

cele otten with a deep proximal constriction,

Hydraeeh short and stubby, with approximately

14 tentaelcs,

Remarks: The single specimen from Bruny 1.

resembles H. tenclivin Hinks, 1861, and Atile-

cium sp. recorded from Pearson 1, (Watson

1973. p. 167). However, H. tenellem: is mono-

siphonic and the dimensions given by Millard

(1957, p. 193) and Ralph (1958, p. 340) for

BH, terelium are grewter than those of the pre-

sent material. The specimen from Pearson L.,

while similar in habit, is monosiphonic, the

margin of the hydrophore is more everied, and

the overall dimensions are smaller, Without

adequate fertile material it is not possible to

make a decisjon on the specific status or rela-

tionships of the specimen fram Bruny L

Walecinm $= beanit (Johnston, 1838). Millard,

1957: 188; 1966: 464; 1968; 256, fig,

9A-F. Ralph, 1958; 332, fig. 10a, b, e-k.

FIGS 5, 6

Records: Adventure Bay {no depth recorded)

on encrusting sponge on underside of the red

alga Sonderaphycus australis: Penguin 1,, 15 m

deep, on sponge tn crevice.

Matevialy Busby infertile colonits to 15 mm

lung. Proximal parts of the colonies fascicled,

becoming monosiphonic distally, usually where

regrowth has occurred from broken poly-

siphonic tubes. Branching irregular, occasion-

ally tendrils given off distal ends of branches

through the orifice of the termina! hydrophore.

Stent internodes of variable length, 0,28-0,52

mm, narrowest at node, 0.08-0,12 mm, widen-

ing distally to 0.18-0.23 mm to accommodate

hydrophore. Nodes distinct, with a slightly

Oblique slope alternatcly nehe and left, occa-

sionally straight, Hvdrapkores alternate, shal-

low, saucer-shaped, adnate to internode, diam,

at margin 0,12-0,14 mm. Diaphtagm distinct,

6.02-0.04 betow margin, tilted towards node,

marked by a thickening of perisare, a ring

of punctue (frequently not well seen) sbove.

Secondary hydrophores given off on a short

pedicel from diaphragm of primary hydro-

phore. adcauline wall convex, abcauline wall

straight or very slightly bulged. Body of

hydranth delicate, with approximately 14-16

long filiform tentacles borne on a long

peduncle, a deep constriction between

peduncle and hyposiome,

Remarks; Since the Bruny J. specimens con-

form to descriptions, and fall well within the

range of dimensions given for HW, beanit by

1ét

Millard (1957) and Ralph (1958), the speci-

mens, although infertile, are asgi¢ned to this

spevies.

An unusial microhabilat is the epizoie

growth on Crustase sponge on the underside of

the thick. plate-like thallus of Sonderophycus.

H. beanii is « cosmopolitan species, not

previously recorded from Australian waters. It

i3 rare at Bruny I

Halecium bruniensis a. sp.

FIGS 7-15

Type material wad records; Holotype, NMV,

G2494—microslide; G2495-preserved maicrial,

remainder of holotype cotony; Penguin 1., 20m

eep, On sponge and bryozoa in crevice,

Description from holetype- Erect stem J om

high, growth habit arborescent, in one plane,

stems sparingly fascicled at base, woody, the

polysiphonic tubes running up stem, forming

the branches, Branches monosiphonic distally,

with markedly sympodial growth. Stem inter-

nodes of ultimate branches of fairly constant

length, 0.48-0.64 mm, narrower proximally,

widening distally to 0.10-0.12 mm at node.

Nodes oblique, well defined, sloping alternately

left and right, often 1 or 2 annulations above

node. Pedice! of hydrophore given off distally

from a well defined apopkysis 0.04-0,14 mim

long, at the same level as the node. Pedicel

tubular, 0.13-0,20 mm long (node to dia-

phragm), perisare of younger parts smooth, in

older regenerated parts heavily internally

ridged, a deep fold just above apophysis giving

pedicel an offset appearance. Hvdrophore fairly

deep, 0,06-0.08 mm margin to diaphragm,

slightly expanding with an everted margin,

0,15-0,18 mm in diam,, and distinct rim, Dia-

phragm weil defined, thin, with a fing of

Punctac just above and a pseudodiaphrapm

below, frequently only marked by a thickening

of the adcauline wal), Secondary and tertiary

regencrations of the pedicel from thé orifice of

the preceding hydrophore common, each suc-

ceeding pedice! usually shorter than the last: in

some instanctcs regeneration is reduced to a

mere replicition of the hydropkore. Secondary

branching of pedicels rare. Mydransh elongated,

slender, with 10-16 long filiform tentactes,

Femaie gonothece very large, flattened, lenti-

culur, but somewhat variable in shape, usually

slightly longer than wide: greatest width 0,98—

1.33 mm; length (excluding pedicel) 1.20-1,43

mm, tapering proximally into a short pedicel

arising at base of hydrophore, usually at junc-

von with main stem, Perisare very delixite,

JEANETTE E. WATSON

162

Figs 7-15.

HYDROIDS OF BRUNY ISLAND, SOUTHERN TASMANIA ms

distal extremity puckered, a small circular

Ovifice sealed by an operculum situated in the

Wistal Vhird of the abcauline wall, Malv gono-

theca small, sausage-shaped, 0,25 mm lenge,

0.10 mm wide, arising from a short pedicel ut

base of hydrophore. Buth sexes arising only

on younger, monosiphonic pars of colony, the

males more distal than females. Colour, stems

straw coloured, female gonophores pink.

Reiarks: Halecium &raniensiy ts closely allied

to Al, Jenticulare Trebileock, 1928, in sym-

podial habit, shape of fernale gonotheca and

hydrotheca, However, HW. lenticulare as known

at present (Ralph 1958, p, 331) is a mono-

siphonte species with smaller stems {less than

L am high), «a considerably smaller female

gonotheca, and) a male gonotheca of somewhat

differen? shape.

The type material of H. Aruniensix shaws

growth stages of the female gonophere from

ecarfiest development to maturity (Figs 12-15).

Development of the yenophore begins with

formation of a hook-shaped hlastesiyle sur-

rounding a central body (Fig, 12). Further

growth and dilfercntation into 6-8 large bodies.

possibly larvae (but material insufficiently well

preserved for positive ilentificalion), then oc-

curs, filling yonotheca (Figs 13. 14}. A small

circular aperture with slightly thickened rim

then develops in the distal third of the ab-

caulme wall through which the reproductive

products escape (Fig. 1S). The orifice of the

now ¢mpty gonatheca then becomes Tesealed

hy a very thin operculum.

Only one group of colonies of A. bruniensiy

was lound These were oplzoic on sponve and

encrusting bryezoa in a sheltered crevice.

Myleciom lutemmn n. sp.

FIGS 16-18

Type marerial and Recerds: Helotype, NMV,

Gl4%6—microstide {KOH cleared preparation);

G2497—-preserved malerial, remainder of holo-

type colony; paratype G24%8—microslide;

Penguio L. 15 m deep on sponge and rock.

Description from fiolofype and’ paratype:

Colonies to 2.5 em high, growth arborescent,

Main stem strongly fascicled, woudy and stiff,

Branching more or less in one plane, the ulti-

mate branches in any one part of the coloiv

all directed anteriorly: in other paris they may

face in other directions, Srem jnrernudes ot

variable Jength, 0.40-0.65 mm, expanding

distally, with 1, occasionally 2, extra nodes.

Nodes distinct, oblique, parallel in each inter-

node, sloping alternately left and right in sad-

jacent internodes; perisare indistinctly internally

fidged. Width at proximal node, measured

parallel to node, 0.12-0.14 mm, Hydrophore

stated on distal third of distal segment of inter-

node, well below node. Hydrophore sessile,

very flat and shallow, walls. thin and strongly

constricted just above diaphragm, abcauline

wall more concave than adcauline. Margin cir-

cular, 0.14—-6.16 mm in diam,, with outrolled

tim. Depth from margin to punciae, 0.015—

0.02 mm; depth from margin to diaphragm,

0,025-0.035 mm. Diaphragm very strong, with

a distinct ring of punctue above, and a wedge-

shaped thickening of the perisare of the infer-

node below. Below the wedge the wall of the

internode thins into a large circular fenestra-

tion from which the apophysis of a branch

may arise, Wydranth large with an annular

hypostome and 25-28 tentacles. Colour, bright

yellow, Gonotheca, absent.

Remarks: Fidlectuni luteum is superficially

similar to A. corrugatissinum Trebilcock, 1928.

from New Zealand, us it has strongly ridged

internodes and shallow hydrophores charucter-

istic of this species, In the latler species, bow-

ever, the nilges of the stem are merely strung

annular constrictions. ot definite oblique

nodes as tn 4. fyrewen, Furthermore, the hydro-

phores, while shallow, are somewhat deeper

than those of the new species. Also, H- corru-

felissinuen is a monosiphonic species, while

A. tntewm has & strongly fascicled hahit,

H. luteum displays several uttusual morpho-

logical features. One is |he presence of supple-

mentary nodes, so well defined that the stems

could almost be described as being alternately

athecale and thecate; another feature is the

extraordinarily shallow hydeophore. which

seems tu offer negligible suppomt to the very

Figs 7-15, Halevium bruniensis hsp. Fig. 7.—Holotype colony, natural size. Fig, &—Distal part of

colony, showing fasciculution of stem and emply female gonothecue. Pigs 9, 10—Hydro-

phures enlarged, showing offset pedicel and replications of the hydraphore. Fig. 1!—Male

gonotheca. Figs 12-1§—Development of the female gonophore. Fig.

12.—Barly stage

of development showing hook-shuped blastostyle with developing central mass. Hig. 13.—

Tater stage of development. Fig.

14.—Mutlure gonophore. Fig. 13.—Gonotheca afrer clis-

chatge of reproductive products. Note residual mass tetnaining at site of the circular orifice

through which cuntents kave been discharged,

164

JEANETTE E, WATSON

imm

Figs 16-18, Hulecium luteum nsp. Fig. 16,—Paratype colony, twice natural size. Fig. 17—Distal part

of branch. Fig: 18.—Stem internodes with hydrophores enlarged.

Fig.

bulky hydranth, The extreme thinning of the

wall of the internode in the fenestration below

the hydrophore must produce a serious struc-

tural weakness of the hydrocaulus. This is,

however, offset by the support given to the

hydrophore by the very strong wedge of peri-

sare extending across the base of the hydro-

phore from the stem.

The group of colonies were both epizoic and

epilithic, growing down from the roof of a

cavern in sheltered conditions.

Phylactotheca armata Stechow, 1924: 59; 1925.

204, fig, C. Blackburn, 1942: 106. Hodg-

son, 1950: 17, fig, 31, Watson, 1973: 166.

Ophiodissa frayiliy Blackburn, 1937: 365, fig. 1.

FIG, 19

Records: Penguin 1. and Adventure Bay: epi-

zoic on solitary ascidians, bryozoa and sponge;

epiphytic on crustose coralline algae and on

19. Phylactotheca armaia Stechow. Gonotheca with developing male gonophorc.

holdfasts of P/ryllospern comoca,

deep.

10-22. m

Material; Luxariant colonies, some fertile.

Siems of variable length, 4-15 mm, usually

simple, occasionally branched, Calonies dioe-

cious, gonophores borne thickly on hydrorhiza

at base of stems, gonothecae large, flatly ovate,

both sexes of same size and shape, widest at

middle or top, perisarc thick, slightly undulat-

ing, borne on a very short pedicel, length 1.14—

1.56 mm (excluding pedicel), maximum width

0.90-1.17 mm Gonophores mature, of creamy

white colour, almost filling gonothecal cavity,

male surrounded by a thin blastostyle, female

packed with mature ova. Mydranths with a

single row of large lenticular nematacysts

(probably stenoteles) alternate with the ten-

tucles surrounding the hypostome. These also

occur in the nematocyst batteries in the capi-

HYDROIDS OF BRUNY ISLAND, SOUrHERN TASMANIA

tulum of the retracted dactylozooids, and are

scattered throughout the hydrocaulus in some

stems, No discharged nematocysis were seen.

Remarks: Blackburn (1937) descried the

gonophore of “Ophtodissa fragilis’ (= PL ar-

mate) as being “subspherical, arising at the

junction of stem and peduncles, as well.as stem

and hydrorhiza", Blackburn’s type microslide

of “OQ. jragiliy’ (NMYV collection) shows three

extremely delicate structures which appear to

be either directly attached to, ar enveloping

the hydrocaulus. Although two of these contain

a central mass which could possibly be a

developing gonophore, they zesemble neither

io shape or structure the immature and mature

gonophores of P. armaza as seen in the present

material. They do, in fact, closely resemble

egg capsules of certain minute gastropods.

P. armata shows # Wide range of substrate.

The epiphytic colonies, particularly those from

the rough-water sites among Phyllespora hold-

fasis. were usually short and robust. with

heavily vidged cauline perisarc, Epizoic

colonies, particularly those from more shel-

tered situations under ledges in deeper water,

were lax, branched, with a more delicate peri-

sare and fewer intranodal ridges. All stems,

however, show a tendency towards thickening

of the perisare and increase jn cauline ridges

With age.

This is the most abundant occurrence of P.

armata so far recorded In Australian waters,

and demonstrates. a greater variability in stem

characteristics than formerly known,

Family SYNTHECIDAE

Synthecium patulum (Busk, 1852), Hodgson,

1950: 18, figs 32, 33.

Sertuferia patela Busk, 1882: 390.

Recards: Satellite L., 14 m deep, under ledge;

Simpsons Bay, 11 m deep, on scallop Equi-

chlamys fifrons; Penguin 1, 20 m deep, on

sponge.

Material: A few infertile coloniés, Sienzs to

25 mm long, some stems immature. Proximal

internodes of stems short, with 1 pair of oppo-

site hydrothecue, followed by an internode

with 2 pairs of opposite hydrethecae and

uistal hydrocladia; distal internodes with either

| pair or 2 pairs of hydrothecae. Aydrothecae

on older stems with thick perisarc and replica-

tions of the margin, Colour, purple and white,

Remarks; The present material conforms to

descriptions of Syntheciam patulunt yiver by

Bale (1914a, p. 5) and Hodgson (1950). How-

ever, it is very difficult (Watson 1973, p, 167)

165

to distinguish between Synthecinmn elegans ¢.

subventricesum and §. petulum on characters

of the hydrothecae alone, The present material

is thus provisionally assigned to §. parulam.

Further work may cventually prove that the

two are conspecific.

No morphological differences could be

detected between stems vrowing in lhe environ-

mental extremes of exposure to current (Simp-

sons Bay), sheltered situations under ledges

(Satellite f.) or exposure to surge [Penguin f.).

Family SERTULARNIDAE

Salucia farquhari (Bale, 1924), Ralph, 196La:

769, fig. 7,

FIGS 20-22

Thuigria farqriart Bale. 1924: 244, fie, 1D

Trebileack, 1928; 19, pl. 8, fig. 4

3 m deep, on sponge under ledge,

Material: Two infertile colonies. Stes mond-

siphonic, to 18 mm long, simple and branched,

the simple stems shortest. Stem internodes

tapening proximally and distally, with one pair

of opposite hydrothecac adnate in front. widely

separated behind. Branching regularly alternate

from an apophysis of the internode 0.25-0.4

mm Jong, usually 2 pairs of hydrothecae

between branches, Branches giver off at an

upward angle with a distinct proximal genicu~

Jation at the apophysis and a V-shaped distal

joint. Some secondary branching, but whefe

developed, these branches carry few hydro-

theeac.

Remarks; The specimens compare well with

microslides of Téuiarie farquharl Bale (NMV

collection) and with the redescription of S.

farquhari given by Ralph (19614). The present

material docs, however, exhibit certain dif-

ferences from the species as described from

New Zealand, These are the greater length and

the proximal geniculation of the first branch

internode, as well as the tendency towards

thickening and Joss of the stem Internodes

in older parts of the colonies, features recog-

nized by Ralph as being more characteristic of

Salacia bicalycula (Coughtrey, 1876a) than §,

farquhari. Since ibe present material agrees in

most respects with the latter spectes, particu-

larly in the size of the colonics, it is assipned

to §, farguheari,

This is the first record of 5. farguiari for

Australia, and the first record of the species

owlside New Zealand waters, where it occurs

only south of 43°S, the same latitude as

Briny I.

16a

Stereotheca elongata (Lamouroux, 1816).

Ralph, 196la;: 762, fig. 4, Watsun, 1973:

170,

Sertularia elongata Lamouroux, 1816; 1839, pl,

5. Bale 1884: 75, pl, 6, figs 7, 8, pl. 19. fig. 7;

1915: 277. Hodgson. 1950: 23, figs 3R, 39

Record; Penguin b., 15 m deep, on rough-water

side of island, on a red alya,

Material; Several lertle colonies. Stems short,

2.5 cm. Gonathecae long and narraw, horned

pracesses very much elongated.

Remarks: Hodgson's Oyster Bay material Fron

storm-drifted Weed is the “long-stemmed” form

af §. elongata. The Bruny 1. material is. the

“short-stemmed” ocean form frequently asso-

ciated with red algae.

Sertularella robusta Coughtrey, 18768; 300, fig.

2, Hodgson, 1950; 33, fig. 58. Ralph.

6la: 824, fig. 22a-d. Watson, 1973:

191, fig, 21.

FIGS 23, 24

Reécorey: Satellite L. 3 m deep, on sponge

under Jedges, and 6 m deep on the red alga

Sanderophycus — custralis; TY Entrecasteaux

Channel, 11 m deep on dead seawhip Priv-

roella ausiralustae, and on old scallogy shells,

Eguichlamys bifrons; Adventure Bay, 5 m deep

on sponge under ledges: Fluted Cape (no depth

eecorded) on stem of a brown alga.

Material: Stems 3-4 mm long, the longer stems

fexuous, with Jong internodes, occasionally

branched, Shorter stems robust, with short

intetnodes, each stem type occurring in

separate colonies, Hydrothecae of the “long

internove” form large with smooth, very faintly

undulated walls, bhydrotheeae of the “short

internode” form distinctly smaller (see dimen-

sions), with thinner perisare. heavily ridged

with 3—4 annulations and a strong submiat'ginal

constriction of the thecn! neck on. the abcauline

side, All material infertile except for one stem

of the “short internode” form,

Dimensions (mm); _ long shor

imernode = internode

form form

Length, abcauline wall 0.55-G.40 133-0945

Length, freeadcauline wall Q.35-0.43 0,28-0.38

Remarks; Specimens of S. robusta from Brony

I, show a complete range of variability between

the extremes of (he long and short internode

forms, and large and small hydrothecuc. Some

correlation uppears to exist between stem type.

environmental conditions and habital pre-

ferences, the long Mexuous stems growing ept-

JFANETTE FE. WATSON

zoically on dead shell and other material in

the D'Entrecasteaux Channel in situations of

good current flow, those af intermediate stem

length being from cryptic habitats beneath

ledges and on the underside af Senderophyeus,

while the more robust stems occurred on the

lower parts of brown algiuc in moderately tur-

bulent open water. Dimensions given by Hodg-

son (1950) for his material correspond to the

“short internade™ form, although the stems of

his material were 15 mm long, and were epi-

phytic,

Except for the greater thickness of stem

and very Faint theeal undulutions, the “long

internode” form of the Bruny J. material cor-

responds very closely with Serratarella simplex

(Hutton, 1873) deseribed from Peurson L

(Watson 1973). A speeitic distinction based on

thickness of perisare and faintness of undila-

tions of the thecal wull as defined by Ralph

(19G61a, p. 820) scems to be somewhat art-

ficiyl. Thus, if further muterial with smooth

hydrotheeae and thin perisare is found, 3.

robusta must be referred to the synonymy of

S. sirmplex.

Syniplectoscyphus pygmaeus

Watson 1973; 176..

Sertularella pygmaca Bale, 1881; 25, pl. 12, fi

9: PRR4: 108, pl 3, fig. B. pl 19, fiz. 19.

Hodgson, 1850: 36. fies 63, 64.

Recard; Penguin L, 16 m deep on bryozoa in

sheltered situations,

(Bale, J88L)-

Marerial: Sparse infertile colonies. Stemy to

7 mm long. simple, or with one branch. A

delicate 3-flapped operculum visible in most

hydrotheeae.

Remarks: The line of smull dots in the thecal

wall diagnostic of S. pygmaeus is obscured by

the. hydranths. However, the material cozres-

ponds closely with specimens of S. pygmaens

in the Bale collection (NMV) and for this

reuson is assigned to this species.

Sertularia acuta (Stechow, 1921). Millard, 1958:

192, fig. 8. Shepherd & Watson. 1970: 140,

Tridentata aenta Stochow, 1921: 231

Seriularia doculoss Bole, (884: 91, pl. 4, figs

5, 6; 1913: 121 pl. 12, figs 7, 8; 1915: 272.

Hodgson, 1950: 25, fies 43, 44.

Reena: Adventure Bay, 1U-22 m deep_ on ted

algae.

Material: Luxuriant fertile colonies. Steet to

7mm tong. Gonethecae with 4 stronz annu-

lations, arising from stem tnternode below

proximal hydrotheca. Gonophores mature,

HYDROIDS OF BRUNY ISLAND, SOUTHERN TASMANIA

Renurks; Hodgson's infertile spectmens came

from storm-drifted Macrocystis. S. acuta was

no| assocjated with this kelp at Bruny I.

Sertularia tmacrocarpa Bale, 1884: 80, pl, 5,

fig. 2, pl 19, fig. If; 1914a: b4; 1915:

277. Mulder & Trebilcock, 1914hb: 42,

Hodgson, 1950: 27, fig. 47. Shepherd &

Watson, 1970: 140, Watson, 1973: 177.

Recerd; Penguin I, sheltered side, 1¢-22 m

decp, among algal holdfasts.

Marerial> Rare colonies, comprising a few

stems each, Stems to & em long, infertile.

Remarks: The internal submarginal tooth is aot

as Well developed in these specimens as in those

from the Australian mainland.

This dark brown species with «distinctive

white-tipped hydrocladia has previously been

recorded among the holdlast tuna of red algae

at Pearson L. (Watson L973}.

This is the first record of 5. macracarpa from

Tasmanian waters.

Amphisbetia minima var, intermedia Bale.

1915. Watson, 1973; 179, fig. 29

FIG. 25

Seridaria minima Thompson, 1879: 104, pl.

U7. fig. 3. Bale, 1881: 21, 45, pl 2, fig, 2;

1884; 89, pl. 4, figs 9, 10, pl. 19, fies 12, 13;

Mulder & Trebilcock, $914b> 39, Hodgson,

1950; 23, files 41, 42.

Recerds: Adventure Bay, 4-6 nv deep, on red

alga Rhodymenia and on sponge under ledges:

Penguin I., 16 m deep, on red alguc and in

crevices in rough water; Satellite I., 1 m deep,

on Laurencia.

Material: Luxuriant fertile colonies, mainly on

red algae, same on sponge. Stems to 5 mm

long, internode length 0.30-0,38 mm; diam, at

node 0.03-0,06 mm_ Aydrotheca 0.19-0.28 mm

long, Tabular nematothecae present in the base

of proximal infrathecal chamber.

Remarks; Although the hydrothecae are larger

than those recognized as var. intermedia (Wal-

som 1973, p. 181), the present material ts

referable ta this variety on the basis of the

shape of the hydrothecae and the presence of

the characteristic tubular nematothecae. How-

cver, several of the slems have rather robust

hydrothecae and the wedge of perisarc hetween

hydrocaulus and hydrotheca considered typical

of the var, pumiloides, Furthermore, one stem

(Fig. 26) from sponge. shows distinct transi-

tional features between the pumniloides and

(mtermedia types, ‘The stem of this specimen

hus four broad, robust. proximal hydrothecae

1467

of “pupniloides” type, and a distal regrowth of

three hydrothecae of unmistakubly “intermedia”

type following stem breakage. The basal pair

of “intermedia” hydrothecae have well

developed tubular nematothecac. Dimensions

of the proximal and distal groups of hydre-

thecae on this stem are given for comparison:

Dimensions (mm); infer-

pumiloides nicdia

type type

Internade—

length 0.33-0.35 0.30-0.34

width at node 0.06 0.04

Hydrotheca—

length 0,20 0.21

width across base 0.24-6.25 0.18-0.20

width across margin D43-0.52 0.32 0.37

The finding of two varietics of A. minima on

ane stem lends support to evidence {Watson

1973) that these varieties are in reality eco.

morphs, the development of which may be

dependent upon the type of substrate available,

or environmental conditions prevailing during

growth,

Hodgson (1950) did not differentiate be-

tween the varieties of 4. mint in his collec

tions; however, according to his measurements.

and the absence of nematothccac, as well as

the material having come from Macrocystis

and “drift” (brawn?) aleac, the material may

be ascribed to the “var, paniloides”.

Amphisbetia operculata {Lannaeus,

Ralph, 1961a; 775, fig. 8.

Sertilaria operculate Litnacus, 1738: 808. Bale,

1884: 67. pl. 6, fiz. 1, pl. 19, fig, 3, Hodgson,

1950: 22, figs 36, 37.

Record; Simpsons Bay. 11 m deep on stem of

dead seawhip Primaoella ausrraliasiac,

Material; One small infertile colony of a few

short stems.

Remarks: Although Hodgson remarks that

“this species is verv abundant in the D'Enire-

casteaux Channel, being constantly taken in the

form of large tangled masses in scallop

dredges”. oniy one colony was found in the

present survey, This apparent rarity may be

due to seasonal erowth (Hodgson's material

was collected in August) or to permanent

changes in the environmental equilibrium

eaused by scallap dredging. The substrate of

old shells preferred by A, operculata (pers,

observ.) is no longer available as the seafloer

of the estuary has now heen invaded by enar-

mous numbers of the New Zealand gastropod

Maoricolpis roseus, Since no colonies of A,

operculata were found associated with MM

rosens, it may be concluded that the smooth

175K).

168 JEANETTE E. WATSON

HYDROIDS OF BRUNY ISLAND. SOUTHERN TASMANIA

shell of this pastropod is unsuitable for settle-

ment of larvae of 4. opercuiaia,

Amphisbetia avia o. sp-

FIGS 26, 27

Type Material and Records: Holotype, NMV,

62499—microslide; G2500-presetved material,

remainder of holotype colony; Adventure Bay:

paratypes. G2501, G2502—microslides; Satellite

1. all colonies on the brown alga Carpogiossam

confiuens, 3 m deep,

Description fram halatype and pardtypex:

Aydrortiza tubular, 0.09 mm diam., reticulate,

very loosely wound on algal surface. Srey

arising at stofonic junctions, simple, un-

branched, to 5 mm long, beginning with 1 or 2

twisty, then a V-shaped proximal joint, follawed

by first thecate internode, Perisate of stem and

hydrothecue thick and very brittle. Stenz inter-

nodes O.38-0.46 mm long, with one pair of

hydrothecae, nodes slender, 0.06-0.08 mm

wide, distal node coilat-shaped, proximal node

V-shaped and socketted into the collar of pre-

ceding internode; if noe absent, it is replaced

by a narrowing of the internode. Hydroiecae

opposite, on distal half of internode. tubular.

narrowing tO margin, adnate for one third af

length, proxtmal adcauline wall more or less

purallcL to axis of internode, in contact or

slightly separated, base of hydrotheca hori-

zontal; a very deep notch. and occasionally a

short oblique intrathecal fold about one third

distance up abcauline wall from base of hydro-

theca, and a corresponding, hut not so deep

inflexion (sometimes niissing altogether) of the

adcauline wall, opposite, but more distally

situated, just behind margin. Width of inter-

node just below hydrotheca, 0.24-0.29 mm:

length of fixes! adcauline thecal wall (measured

diagonally) O.15-0.19 mm, length of free

adcauline wall (to end of tooth) 0.12-0.18 mm;

length of abcauline wall (measured diagonally

from base fo end of tooth) 0.26-0.3t mm,

Margin horizontal, facing upwards, with two

long, sharp, laterally placed teeth with a deep

hortontal embayment between, connected to

the internode by a thick wedge of perisarc,

—_——-_ x rr eee

169

Width across margin between teeth 0.07-0,09

mm; width across paired hydrothecae (outer-

most teeth) 0.52-0.69 mm, A delicate internal

sheath aften present within margin. Goro.

thecae large, obovate, 1.15-1.33 mm long (in-

cluding pedicel) expanding from base to sum-

mit; max, width 0.75—0.83 mm, tapering evenly

into a narrow pedicel arising from the infes-

thecal chamber of hydrotheca on lower stem.

Aperture circular, 0,35 mm in diam., centrally

situated af distal end, with a slightly raised

collar, a ring of minute denticles within, and a

Nat operculum. Gartorhecae identical in both

sexes, only onc borne on each stem: male and

female gonophore on same colany, Female

gonophore narrowly elliptical, not filling gono-

thecal cavity, with 16-20 eggs; mule gonophare

of same shape and size as female, spermato-

genic mass surrounded hy a thin blastostyle.

Remarks: Amphisbetia avia i8 closely related

10 |he Amphishetia minima group in size,

colour, habit, and preference for algal sub-

stfate, and jis not easily distinguished from the

varicties of 4, minima in the field, However.

the deep retroflexion of the abcauline wall, the

horizontally directed distal part of the hydro-

theca, and the long marginal tecth immediately

distinguish A. evia from A. mininie.

A. avia wis associated only with one species

of alga. Carpogiesswm confluens, growing in

shallow water.

Family PLUMULARIUDAE

Halicornopsis elegans (Lamarck, 1816). Bale,

1914a: 56, 1915; 303; Briggs, 1914: 299;

T9715: 309, Blackburn, 1942; 107. Hode-

son, 1950: 48, fig, 79, Watson, 1973; 195,

Flamataria elegans Lamarck, FRI6> 129.

Record: Adventure Bay. 15 m devp, epilithic

on Yertical face.

Marerial: One fertile colony. Stenrs shart. tc

7 em, Gonotkecue borne prolifically on main

stems and occasionally at base of branches on

the apephysis of the hydrociadium. Gono-

thecae mature. imegularly ovate, Aattencd dis-

tally or slightly flattened on onc side. aperture

closed by 4 thin membrane. Only female gono-

Figs 20-22. Sulecia faryxhari (Bale), Fig, 20.—Part of stem. Fig 21.—Stem imternodes, enlarged.. Fig.

22.—Single hydrotheca, anterior view. ;

Figs 23,24. Serilarella robusta Coughirey. Fig. 23.—“Long internode"

farm with large hydrothecae

and smooth thecal walls. Fig. 24.—*Short internode" form, showine smaller hydrothecse

with undulated walls.

Fig-

25, Aniphishetia minima vat. intermedia Bale. Aberrant stem with beth “pumileides” (proxi-

mal part of stem). and “intermedia” (distal) hydrothecue,

Figs 26, 27.

internodes, enlarged,

Amphisbetia avia n.sp, Holotyps. Fig. 24.—Stem with

fermile gonophore. Fiz 27 —Stem

phores present, spherical. half filling gano-

thecal cavity, packed with maiure ova sur-

rounded by a thin granular blastostyle.

Remarks: Only one small colony of A. elegany

was found growing in a relatively exposed

situation,

Muture gonopheres of Ff. elegrins have not

previously been described,

Antennella cumpauuliformis (Mulder & Trebrl-

cock, 1909). Watson, 1973: 182. figs 43.

44.

Flumwlaria campanuliformis Mulder & Trebil-

cack, 1909; 31. pl. 1, figs 6,9, 10; 1910: 115,

Record: Satellite 1, on Lenormandia marginata

and other red algue: no depth recorded,

Material: Luxuriant infertile colonies. Erect

stems to & mm _ high, Colour, trophosome

pinkish-yellow, stolons dark brown.

Remarks! This material conforms reasonably

well with the description of Mulder & Trebil-

cock (1909). However, the hydrothecae are

slightly more campanulate and delicate, with

no thickening of the abcauline thecal wall as in

specimens described from Pearson 1. (Watson

1973),

The hydroid was found on one side only of

the algal fronds.

This is the first record of A. cantpanulifarmis

from Tasmania, Other localities: Victoria; S.

Aust.

Pycnotheca mirabilis (Allman, 1883) var, mira-

bilis Stechow, 1925; 241, Ralph, 1961b:

50, fig, Ta, b.

FIG, 28

Pyenatlieca mirabilis (Allman, 1883). Hodgson,

1950: 50, figs 81, 82.

Diplocheilus mirabilis Allman, 1883: 49, pl. 8,

fizs 4-7,

KRirchenpauerta mirabilis (Allman, 1883), Bale,

1894: J09. pl. 6, figs 4-7. Briggs, 19151 308,

Blackburn, 1942: 106.

Records: Satellite 1., 9 m deep; Adventure Bay,

10-22 m deep, on red algue.

Warerial: Fertile stems to 5 cm high. Stems

monosiphonic, arising singly from bydrorhiza,

lower stems.devoid of hydrocladia, Gonothecae

large, adnate to lower stem. hydrorhiza, or

algae, Perisare very thick, strongly ridged with

up to 9 ridges, more prominent on abeauline

side. Colour of gonotheca, deep red brown,

Gionophores, female,

Reniarks: Ie is difficult to distinguish between

infertile stems of FB. mirabilis and P. producta

Bale, 1881. Examination of fertile miterial of

hoth species in the Bule collection (NMV),

JEANETTE E. WATSON

shows that the Bruny I. specimens correspond

closely to a microslide at P. mirabilis from

Buss Strail. Gonothecae of PL praducta col-

lected at Port Jackson, in 1886, are smaller

(1,561.86 mm long, 0.75-0.84 mm wide),

have a thinner perisarc, and are only faintly

undulating. These are similar to Hodgson’s

(1950) figure and dimensions of “P. smurabilis”.

Ralph (1961a) noted the discrepancy between

Hodgsen’s description, figures, and the actual

dimensions of the gonotheca of P, mirabilis

and sugwested that it may represent a new

varietal form of P, producta. It scems more

likely, however, that the numbers of the two

figures of the gonothecae of P.. producta and

P. intrabifis have heen confused in Hodgson’s

poper,

Halopteris campanula var, campanula (Busk,

1852). Ralph, 1961b: 47. Watson, 1973:

184,

Plumularia campanuta Busk, 1852: 401. Bale,

1884; 124, pl. 10, fig. 5; 1913: 133, Hodgson,

1950: 40.

Record: Satellite 1, 14 m deep, under ledge.

Material; One infertile colony, Stems to 2,5 cra

Jong, sparingly branched, fax, Colour, yellow

Remarks: H. campenula var. eampanula. is

represented in this collection by only one sparse

cpilithic colony. which was growing in reduced

light in 4 crevice.

Plumolyria filicaulis Kirchenpauer. 1876; 28,

pl. 5, fig. 6. Bale, 1884: 134, pl LI. figs

6. 7. pl, 19, figs 41, 42. Léloup, 1934: 4

Hodgson. 1950; 42, fig, 72, Millard, 1958:

209, fig. 13D, EB. Shepherd & Watson,

1970) 140.

Record: Adventure Bay, 10-22 m deep, on red

algae.

Marerial: Luxunant infertile colonies, Myadre-

rhiza pegged, forming a reticular network on

the algal frond, Stemi short, simple and pine

nate on the one colony. Simple stems to 2 mm,

pinnate stems to 4 mm high.

Remarks; PB. filicaulis is 9 common epiphyte on

several species of red algae (Shepherd! &

Watson 1970).

Pluniularia hyalina Bale, 1881: 41. pl. 15, fig, 9:

1884: 141, pl. 12. figs 4, 5, Ralph, 196th:

41.

FIG. 29

Revord: Fluted Cape, 16m deep, on a red alga

Marerinl: Abundant, sparingly fertile colonies.

Stems 10 3 mm long, stems and hydrocladial

HYDROIDS OF BRUNY ISLAND, SOUTHERN 1 ASMANIA 171

Timm

Fig 28.

female gonophore.

Imm

Pycnotheca mirabilis var, mirabilis (Allman). Gonotheca with heavily ridged perisare and

29, Plumularia hyalina Bale. Stem with ripe female gonophore. -

Fig,

Figs 30, 31.

Plumiularia angusia Stechow. Fig. 30—Purt of hydrocladium, Fig. 31.—Gonothecu.

Fig. 32. Plumularia craterifarmis Stechow. Part of hydrocladium,

Figs 33, 34.

downwardly directed growth habit,

internodes strongly ridged, hydrocladia with

one terminal hydrotheca, Gonothecae large.

top-shaped, greatest widih distally, 0.7 mm

wide, 0.9 mm long; gonophores. mature, female

anly, completely filling gonothecal cavity,

Colour, stems white to yellow; gonophores

bright yellow.

Remarks: This is the first record of mature

P. Ayalina in Australian waters. Ralph (19616)

described much longer (1.3-1.7 mm) and

slightly narrow (0.5—-0,65 mm) gonothecae for

her mature specimens from New Zealand.

Judging by this considerable difference in size

of gonothecue, it seems that an as yet undocu-

mented tunge of geographic variants of P,

hyalina may exist.

Plumularia. wilsari Bale. Fig. 33.—Part of hydrocladium. Fig. 34.—Gonotheca, showing

Some stems of the material from Bruny I.

have a well developed “stolonic plate” identical

with that described for Plumularia epibracteo-

losa Watson, from Pearson I.

This is a new record for Tasmania, Other

localities: Yic,; S$. Aust.; New Zealand.

Plumularia angusta Stechow, 1923b:

Blackburn, 1942: 108,

FIGS 30, 31

Plumularia setaceoides vars 4a, b, d, Mulder &

Trebileock, 1910; 117, pl. 2, fig. 9, pl. 3, figs

Records: Penguin I., 15 m deep, on Macro-

evstis holdfast and sponge; Adventure Bay, on

Macrocystis holdfasts and the brown alga

226.

Myriodesma quercifolium.

172

Material: Abundant fertile stems to 1 ¢m long,

Hydrothecae pitcher-shaped, but variable, some

with a pronounced concavity of the adcauline

wall and.a consiriction behind margin, others

with a thickening of the abcauline wall. Colour,

straw-coloured, gunothecae bright orange.

Remorks: The hydrothecae are very variable

in shape even on the one hydrocladium, the

thecal walls ranging from pitcher-shuped to

almost straight, often closely approaching the

shape of the hydrotheca of P. sefaceoides.

Plumularia craterifarmis Stechow, 1923b; 227,

FIG. 32

Plamularia setacevides yar. —_cratertfarmis

Mulder & Treblleock, 1910: 118, pl. 3, figs 8,

Raz 1918: Sl, pl. 7, figs 3. Ja.

Reeord: Advenlute Bay, 4-6 m, on the brown

alga Niphophora gladiata,

Marcrial: Infertile stems to 1 em long,

Remarks on P. atgusta und P. crateriformis:

Stechow (1923b, pp, 226, 227) raised Mulder

& Trebilcack’s. varieties of P. setaceoides (the

unnamed vars a, b, d, and var. ereferi/ormls)

to specific rank, his distinction between the

two species resting largely on the shape of the

hydrotheea, a character of considernble varta-

bility in this group and thus of doubtful diag-

nostic value. The material from Breny 1,

although showing some intergrudation, can,

however. be fairly readily assigned to one or

other of these two species,

This, together with the fact that the two

species occur together in the one locality,

although at different depths. and on different

algal substrates (and one, P. angusta, was fer-

tile, while P. crateriformis was not) indicates.

thal these are valid, although closely related

species radiating from the central P. setaceoides

stock.

Neither P. angusra and P, crateriformis have

previously been recorded from Tasmania. They

may have been confused with P. sefacéoides

P. angusta is Known from Victoria and §, Aust.;

P. craterifermis from Victoria.

Plumularia wilvent Bale. 1926: 21.

196th: 31, figs 2, 3,

FIGS 33, 34

Plumularia delicatula Role, (881: 28, pl. 15,

fig. 2; J884: 137, pl 11, fig. 5, Mulder &

Trebileock, 1910: 115, pl. 2, fig. 2.

Records; Penguin 1; Adventure Bay: Satellite

J,, 3 m deep, on sponge.

Muserialr Sparse fertile slems to 2 cm long.

CGonethecae vop-shaped, | or 2 arising on a

Ralph,

JEANETTE LE. WATSON

very short pedicel from proximal part of stem,

distal end directed downwards towards suh-

strate. Perisare delicare, very faintly undulated,

no operculum. Gonophorcs—male,

Remarks: The \tophosome of the Bruny [.

specimens corresponds very closely with Bale’s

material of PL wilsent from Griffiths Point,

Victoria, The submarginal constriction behind

the hydrothecae of the present material is

rather variable, being more pronounced in same

hydrothecae than in others even on the same

hydrocladium. The hydrothecae with a shallow

constriction closely approach those forms of

P. setaceoides with more recumbent hydro-

thecae.

The gonothecae of the Bruny 1, specimens

ate identical with those of Mulder & Trebil-

cock's (1910) microslide from which they

described the gonotheca of P. wélseni. Their

figure is, however, misleading, as the walls of

the figured specimens, like those of the present

material, are only faintly undulated, not heavily

ridged, as may be inferred from their figure.

A new record for Tasmania, Other Jocalities:

Victoria; New Zealand,

Agtaophenia plumosa Bale, 1881: 25, pl. 14,

fig. 6: 1884; 153, pl. 14, fig. S, pl 17,

fig. 12; 1924: 257. Blackburn, 1942; 110,

Hodgson, 1950; 56, fig. 37. Shopherd &

Watson, 1970: 140.

Record: Fluted Cape, 20 m deep, on sponge.

Masertal: Rare infertile colonics Stems to

1 cm tong.

Remarks: This is the robust form of 4. plti-

nose with short hydrocladial and stern Inter-

nodes, Hodgson notes thai his material cel-

lecied from Macrecystis and drift had shorter

stems (Le. the short internode farm) than thase

stems collected from the scagrass Zostera,

which grows in sheltered waters.

Thecocarpus. divaricatuy var. typica (Busk,

1852). Shepherd & Watson, 1970: 140.

Agluophenia divaricata Busk, 1852: 398.

Records: Adventure Bay, 1-10 m deep, on

horizontal faces among holdfasts of brown alga

Phyllospora comosa; (0-22 m deep, epilithic

in sheltered situations.

Material: Stents infertile, to 10 cm in height.

branched, Cauline nematotheeac similar in

shape to those of 7. divaricatus var. cystifera,

but much smaller, Hydrocladia close-set, hydro-

theeae crowded on hydrocladium, no oblique

intranudal septa on thecate hydrocladial imler-

ioule, Colour, dark brown.

HYDROIDS OF BRUNY ISLAND, SOUTHERN TASMANLA

Thecocarpus divaricatus (Busk) var. briggsi

Bale, 1926- 22, fig, 5, Watson. 1973; 194,

Record; Satellite L., on the red alga Thamno-

clonivn dicietomam. No depth recorded,

Material; Stragpting, irregularly branched.

stems to 15 cm long, Colonies infertile, Colour,

black.

This ts the first record of var. brigesi from

Tasmanian waters. Other ‘localities, Port

Jackson, N.S.W. (Bale); Pearson 1, S. Aust.

(Watson) -

Remarks on the varieties af T. divaricatus at

Bruty Island: The two varieties of T. alvari-

cutus tecorded at Bruny J, are easily distinguish-

able by the presence of an intranodal ridge in

the hydrocladitum of var. driggsi, as well as in

differences in habit and substrate preferences

of cach.

The var, friggst was found only in sheltered

recfs near Satellite 1, in the D'Entrecasteaux

Channel, where it was a common epiphyte on

Thamnoctanium, The more robust var 7) pica

occurred on rock faces and as a coimmon epi-

phyte on Phyllospora holdfasts in situations of

moderate exposure to surge.

Although showing affinities with var. cysri-

fera, the latter form dors not display the dis-

tinctive planar growth habit recorded for this

variety ¢Watson 1973), nor the enlarged

cauline nematothecac. In microstructures it

most closely fesémbles a fragment of Busk’s

type Of Aglaophenia divariceta from Bass Strait

{NMV collection) and is thus recoynized here

as Var. typica,

T, divericetks is a common and variable

Spectres of the southern Australian toast,

Further study is necessary to elucidate the sys-

tematic status and ecological relationships of

the yarletiés of this species.

Halicornaria longirastris (Kirchenpayer, 1872),

Bale, IBS4: TRI. pl. 13, fiz, 7, pl. 16,

fig. 3, pl 19, fie. 30, Hodgson, 1950: 51

fig, 83, Watson, [973+ 197,

Records; Penguin [., 10-20 m deep, on Dhera-

carpus divaricatus: Adventure Bay, 10-20 m

deep, on red algae and epilithie on vertical

faces: Satellite 1.12 m deep, on 7. divarlcatus

var. briggsd.

Material: Luxuriant colonies. stems 3-8 cm

long, unbranched, infertile.

Remarks: The material from Bruny lL, displays.

the same range in substrate as wlready noted for

H. lougirestris 4¢ Pearson 1, (Watson 1973),

Epilithic colonies from faces exposed 1% surge

have the longest and most robust stems, while

epizoic colonies on PF. divaricuius var. brigest

from both Penguin ft. and Satellite f (ice, from

Protected and relatively rough-water situations)

had somewhat shorter, lyx stems, given off

singly from a stolon creeping on the stem of

the hydroid host. The cauline internodes of

these latter stems are long, with distant hydro-

cladia, the hydrocladia themselves having long

internodes, and the mesial nematotheca extends

well over the mouth of the hydrotheca, Rare

epiphytic colonies on red algae at Adventure

Bay had the shortest stems, There were no

discernible clifferences in microstructures be-

tween these three ecomorphs. Bripge (1915)

Mentions that his specimens from Storm Bay

were “86 mm in height (and) were fotind asso-

titted with Agiaophenia divaricata”, Hodgson

(1950) did not record the substrate of his

material from Blackman's Bay in the Derwent

Estuary,

Discnssion

Ecology

Campanulariidae were recorded from all

depths, being epiphytic on red algae growing

mostly in sheltered places, One exception, Sifi-

cularia rosea, was associated with the brown

algae Seytothalta darycarpa and Sviraceceu;

axillaris in situations of modergte water move-

ment.

The only representative of the Lafoeidae,

flebella furax, provisionally recorded for the

first time in Australian waters, is a small form

epizoic on the stems. of Thecacarpus divarice-

tus var, briggsi Bale. The majority of species

of the Lafoeidae known from Tasmanian and

maintand Australian waters are larger forms

from the deeper continental shelf, hence are

unlikely to be found in a shallow water collec-

tion such as the present one from Bruny I.

With the exception of Phylacrorhece armata

and Haleciun p., all haleciid species in the

collection are epizoic forms. Hulecium delicatu-

fum Coughtrey [H, flexile of Hodgson (1950)]

is one of the mast abundant hydroids at Bruny

J,, the luxuriant orange-yellow colonies growing

on the ¢rustose bryozoan Membrinopora mem-

brinaeca epiphytic on old stipes of the large

kelp Macrovystis pyrifera, The two species

newly described. Halecivim bruniensig and

FAY, lateum, as well as H. beanii, are of eryplic

habit, the two new species growing on sponge,

bryozoa, and rock in crevices, while H. heanii

occurs on the underside of the large plate-like

ales Sanderaphycus australis, While fertile

tolonies of Phylactotheca armata Were tecarded

abundantly a all rough-water siles. they in Eact

also occupicd « relatively shellered milerchabitat

among the holdfasts of the brown alga Piyile-

SPOra COMUSE..

Remarkably few species of Sertulariidac were

recorded. Of greatest interest 7s Uhe first record

of Salacie farguhert (Bule) outside New #ea-

land waters (see further discussion beluw). 4.

fargultari was. however, tare at Bruny I, Seree-

Jaria acute Stechow, while one ol the most

abundant epiphytes in the collection, was asso-

ciated only with red algae, and not with

Macracystiy as were Hodgson’s spetimens.

Hodgson (1950) also recarded very ubundant

colonies of Amphishetia aperealata on ald

scallop shells from the D’Entrecasteaux Chan-

nel, Careful search in [he wrea of the D'Untre-

casteaux Channel covered In this suryey pro-

duced anly one attenuated colony. Although

A. operculate is known to display strong sea-

sonal growth, some of the rootstock and puts

of the colonies usually persist fram one scason

to another (pers. obsery.}, The virtual absence

of this species fram a former habitat may be

explnined by permanent chinges in the cca-

svstem brought about by invasion of the gastro-

pod Maoricvolpuy raseus, following, the callapse

of the scallop dredging industry. Probably the

shell of M. roxseay does not offer an attractive

substrate to the larvae of A. operculuta,

Of particular interest is the occurrence in

une locality of three species, Plurnularia an-

gusta, P. erateriformiy and P, wilyoni, all of

which we known from Victoria. P. angusta is

4)so recorded {rorm South Australiv (Blackburn

1942) and P. wilsont from New Zealand

{Ralph 19616), where it is. however, ture.

These species are closely related to onc another

and ro Plemuleria setaceoides Bale, endemic to

southern Australian and New Zealand waters,

from which central stock they may have

Tadiated,

None of the larver Plumularians were of

common occurrence. Halicornapsis elegans and

Thecoearpus divavicatus vat. lypica were Found

in shellered situations, The association of T-

divaricarus var. briggsi with a bluish coloured

sponge investing the warty surface of the alga

Thanmoctonium dichatomun bas not pre-

viously been observed. Halicornaria longirastrixs

was moderately abundant at all sampling sites.

the three ecomerphs of this species displaying

an identical choice uf substrate with that

reported from Pearson I, (Watson 1973).

Zoogengraphy

The Bruny & material, allhough collected

from a testricted locality, yielded 11 sew

JEANETTE FE, WATSON

records for Tasmania, including 3 new records

for Australian Waters. Seven of these species

are ulready known trom the Victorian coast-

line of Bass Strait, so would be expected ta

occur among the Tasmanian faung. Thecocar-

pus divaricatus var. briges, cow kooawn from

N.S.W. (Bale 1925). Tasmania and South Aus-

tralian (Watson 1973) has not yet been recorded

from Victoria,

Of the 3 new records for Australia, fMuale-

cium beanil js cosmopolitan, Hebelle furax is

known anly from South Africa, while Salmeis

farauharl is a New Zealand species recorded

only from the South Island, where it does. nv,

however, occur north of 43°S, The absence of

S. farquhari from maintand Australian waters

may thus be attributable to this southern distr-

hution, as Bruny §. also fies close to 43°.

The profound influence ol the Bust Australia

Current and the West Wind Drift in the dis-

persal of species and the biological and zoo-

geogtaphic relativoships of the trans-Tasmun

hydrold Jauna have been discussed at Jeneth

by Ralph (1961c). In this regard, three spectes

recorded from. Bruny J, are of considerable

interest. These are the two haleciids newly

described, and the occurrence of 3. faryuhari.

Haleciiua Aryniends bears a strong resemblance

to H. fenticulare, a species known from the

South Island of New Zesland and Cook Stra,

while H. Juteum displays close aflinives with

H. corrugdissimun, a tare species recarded

from hoth North and South Islands, ‘The strik-

ing similarities in stem maiphology between

the members of these two pairs. as well as the

gross differences in hub (beth Australian

species are polysiphonic, and the colonies are

largert, strongly suggests ywetive progress ul

specialion from a common siock in addition

to dispersal across the ‘Tasman Sca.

Acknowledgments

fo any indebted t9 Professor H. B. 5.

Womersley, Butany Department, University of

Adelaide, for identification of algae; to my

diving compunions, §. A. Shepherd, R. Baldock

and N. Coleman, for assistinice in the ficld

und to Miss Rhytlis Plant for some af the

drawings, T gratefully acknowledge financial

support in the form of a research grant from

the Trusteey of the C.S'L.R.O. Science and

Industry Endowment Pund.and a field research

grant from the Royal Society of South Aus-

tralia which partly met the costs of this study.

HYDROIDS OF BRUNY ISLAND, SOUTHERN TASMANIA

175

References

ALioan, G. J. (1872).—A Monograph of the

Gymnoblastic or Tubularian Hydrnids, Ray,

Soc. Lond. Part (. (874; Part T1872,

ALLMAN, G. J. (1885)—Report on the Hydroida.

Part I. Plomulariidac. Rpt. Sci. Res, HAS.

Challenger. Zoolagy 7(2), 1-55, Plates 1-20.

ALLMan, G, J. (1888).—Report on the Hydroida

dredged by H.M.S. “Chullénger” ducing the

years 1873-1876. Part WW. Uhe Tubwlarinae,

Corymorphinse, Campinularinge, Sertu-

larinae and Thalamorpha, Rp. Vey. Challen-

ger 1873-1876, 23( 70). 1-90.

Bave, W. M, (1881).—On the Hydroida of south-

easter Australia, with descriptions of s\yp-

posed new species, and notes on the venus

Aglaophenia. J. micr, Soe. Vier. 2(1), 15-47,

Pls XI-XV.

Bare, W. M, (1884}.—Catalogue of the Aus-

tralian Hydroid Zoophytes," (Aust. Museum;

Sydney.

Bare, W. M. (1888),—On some new und cnre

hydroids in the Australian Museum ¢ollec-

lien, Proc, Linn. See. NSW. 3, 745-799,

Pls XU-XXI1.

Bate, W, M. (1894).—Further notes on Aus-

tralian Hydraids with descriptions of some

new species. Proc. R. Soc Vici, (0.s.) 6, 93-

(17, Pls ILL-VI,

Bath, W, M, (1913)—Further noves on Ausira-

lian hydreids. Th. roc, RR. Sov. Viet. tne.)

26, 114-147, Ply XDL XU

Bate, WM. (19/448),—Repor on the Hydroida

collected In the Greal Austratiam Bight and

other Jocalines, Biol. Res. FILS, “Endesvour”

2(1), 1-62, Pls T-VII,

Bate, W. M. (19t4b).—Further notes on Aws-

traltun hydroids, Ul, Proc. Ro Soc. Viet.

(ms.) 27, 72-93, Pls XI-XII.

Bark, W, M, (1915)—Report on the Hydroida

collected in the Great Australian Bight and

other Jocalitles. Part TT. iol, Res PLS.

“Endeavour” 35), 241-336, Pl XLVI-

XLYVII,

Bare, W. M. (1924) —Report on some hydroids

from the New Zealand coast with notes on

New Zealand Hydroida generally, supple-

menting Farquhat’s lis Tras. NZ. Inst. 35,

225-268,

Bare, W. M. (1926)—Puorther notes on Austra-

Tain hydroids. V. Proc, Ro See, Vier. (n.s.)

_ 38, 13-23.

BLAckKsuitn, M. (1937),—Reparis of the expedi-

tion of Ihe McCoy Society for field investiga-

tiun utd research, I, Cuelenterata, Froc. R.

Sec. tes, (na.) 49. 364-371.

Biackeurn, M. (1942),—A systematic list of the

Ilydroida of South Australis with a summary

of their distribution in other seas. Trans, R.

doc. S. Aust, 66(0), 194-118.

Bricss, FE. A. (1914)—Hydrozos from one

hundred fathoms, seven miles east of Cape

Pillar. Tasmania. Ree. Awst, Mus. 10(10),

2R5-301_. Pls 25, 26,

Bripss. EA. (19{5).-Noles on Sasmaniun

Mydrozoa, J. Proe, R, Soe. NSW. 48, 3W2-

318, Pls X. M1,

Busk, G, (1852),—An account of the Polyzou

und Sertularian zoophytes. Jv J. MacGillivray,

"The Natrative of the Voyage of the H,M.S,

“‘Rattlesnake’", Appendix LV, 385-402.

(Bonne: London.)

CouGgHTREY, M_ (1875).—Notes on the New Zea-

land Hydroidene, Trans, Pres NuZ inst. 7,

381-293, Pl, 2

CoucHTReY, M. (1876a)-—Criti¢al jioltes on th

New Zealand Hydrolda, Trans. Proc, NZ

fest. 8, 298-302,

CouGHTRer, M. (18766),—Critcal notes on the

New Zealand Hydroida. Ann, Mog, Net, Hist.

scr, 4, 17, 22-32, Pl, 3,

Hincks, T. (1853).—Further notes on Hritish

Zoophytes, with descriptions of new species.

Aan. Mus. Net, Hist, sec. 2, 2, 178-185, Pls

5, 6

Hinairro, Emperor af Japan (1969),—Some

hydroids of the Amakuga Islands. Hial, Lad.

Mnperial Household, Tokyo %, 1-32.

Hobcson, M. (1950),—A revision of the Tus-

munian Hydroida. Pup. Proe, R, Suc. Tasm,

1949. 1-65,

Jitisen, F, W. (1873)—On the New Zeuland

Ser tularians. Trans, Proc, NuZ. Inst. 5, 256

Jounstox, G. (1838) —*A history of the British

Zoophytes.” ist ed. (W. H. Lizars: Edin-

burgh, 1

RIRCHENPAUER, G, H. ({872)—Uber dit

Hydroidenfamilie Plumularidae, — cinzelne

Gruppen derselben und ihre Fruchtehaiter. [.

Aglaopheniu, Abh, naturw. Ver. Hamburg §.

1-52, Pls 1-VIUL,

Lamarck, 7. B. (1816)—"Hisioire naturstle des

Animuux sans Vertebres, U1, Polypes.*

LaMouroux, J, VY. F. (t8)€)—*Histoire des

Polypiers Corallizénes ficxibles. vulgairement

hommes Zoophytes." (Caen.)

LeLour, E. {1934).—Tmis hydrapolypes de la

bale de la Table. Afnque, Avstrale, Ball.

Mus, Fis), nat. Belg. WCE9), 1-7,

LENDENFELY, R- von, (1883)—Uher Coelen:

teraten det Sdusec. IV, Miutheiing Eucopetla

campaniuléria nov. gen. Zelisetir, fowine “owl.

Leipzig 3804), 497-582, Pls 27-32.

Lyynatus, C. (1758) —"“Systema Nawurac,! 10th

ed. (Lipsine.4

Meven, F. J. F. (1834) —Beitrige zur Zoologie,

kesammelt aul einer Reise um die Erde. V-

Ucber das Leuchten des Meeres und Beschrei-

bung einiger Polypen und anderer niederer

Thiere. Nova Arta Acad, Caes. Leepold-

Curof. suppl. 16, 125-216.

Miraam No A. H. (1957)—The Hydrozoa of

False Bay, South Africa. ava. 8. Afr. Mus.

$3, 173-243.

Mrtcarp, N. A. H (1958), —Hydrazoa from the

cogsis of Natal and Portuguese Esst Afrieg.

Part 1, Calvproblustea. Ann, S Afr. Mus.

44, 165-226,

Mittaro, N. A. H. (71964),—The Hydrozoa of

the south and west coasts of South Africa.

Part U1. The lafoeidae, Syptheciidae ond Ser-

Tulariidae. Avi, $. Afrf. Mivs, 48, 1-56.

176

MiLLarp, N, A, H. 41966)—The Hydrozoa of

the south and west coasts of South Africa.

Part UI. The Gymnoblastea and small

families of the Calyptoblastea, Ann. 5S, Afr.

Mus, 48, 427-487.

Mittarp, N. A, H. (1968).—South African

hydroids from Dr. Th. Mor'tensen’s Java-

South Africa expedition, 1929-1930. Fidensk.

Meddr. dansk naturh, Foren, 131, 251-288.

Minztarp, N. A. H., & Boumuon, J. (1973).—

Hydroids from the Seychelles (Coelenterata).

Annis Mus, r. Afr. ceni. ser. 8, 206, 1-106.

Mutoper, J. F., & Tresmcock, R, £. (1909).—

Notes on Victorian hydroida with descrip-

tions of new species. Geelong Nat. 4, 2nd

ser., 29-35, Pl |.

Muxper, J. F., & Tresircock, R. E, (1910).—

Notes on Victorian hydroida with descrip-

tions of new species. Geelong Nat, 4, 2nd

ser., 115-124, Pls Jt, IIL

Mutoper, J. F.. & Trepitoock, R. E. (1914a).—

Victorian hydroida with descriptions of new

species, Part II. Geelany Nat. 6(1), 6-15,

Pls I-Iil.

Mucper. J. F., & Trearmcock, R. E. (1914b).—-

Victorian hydroida with descriptions of new

species, Part IV. Geelong Nat. 6(2), 38-47,

Pls 1V-V1,

Mutper, J, F., & Trepitcocx, R. E, (1915).—

Victorian hydroida with description of new

species. Part V. Geelong Nat. 6(3), 51-59,

Pls VIV-LX,

RAvpH, Patricia M. (1956).—Variation in Obélia

geniculata (Linnaeus, 1758) and Siticularia

bilabiate (Coughtrey, 1875) (Hydroida, TF.

Campanulariidae), Trans. R. Soc. NZ.

84(2), 279-296.

Rate. Patricia M. (1957)—New Zealand

thecate hydroids. Part I. Campanulariidae and

Campanulinidae. Trans, R. Soc. N.Z. 84(4),

811-854,

RALPH, Patricia M. (1958)—New Zealand

thecate hydroids. Part Il. Families Lafecidae,

Lineolariidac, Haleciidac and Syntheciidae.

Trans. R. Soc, N.Z. 85(2), 301-356.

JEANETTE E. WATSON

Rapa, Patricia M. (196la).—New Zealand

theeate hydroids. Part IL. Family Sezrtu-

jatiidae. Trans. R. Soc. N.Z 88(4), 749-

835.

Rarpa, Patricia M. (1961b)—New Zealand

theeate hydroids, Part TV. Plumulariidae,

Trans. R, Soc. N.Z, (n,s.) 1(3), 19-74.

RALPH, Patricia M. (196lc).---New Zealand

thecate hydroids. Part V. The distribution of

the New Zealand thecate hydroids. Trans, R.

Soc. N.Z., Zool. 1(7), 103-111.

SHEPHERD, 8S. A., & Watson, Jeanette E. (1970).—

The sublittaral ecology of West Island, South

Australia. 2. The association between

hydroids and algal substrate. Trans. R. Soc.

S. Aust. 94, 139-146.

Srecnow, E. (1921)—Ueber Hydroiden der

Deutschen Tiefsee—Expedition nebst. Bemer-

kungen liber cinige andere Formen. Zool.

Anz, 53, 223-236.

Srecuow, E. (1923a)—Neue MHydroiden der

Deutschen Tiefsee—Expedition nebst Bemer-

kungen tiber einige andere Formen. Zool.

Anz, 56, 1-20.

Srecnow, E. (1923),—Zur Kenntnis der

Hydroidenfauna des Mittelmeeres, Ametikas

und anderer Gebiete. Zool. Jb. (Syst.) 47,

29-270.

SrecHow, E, (1924-),—Diagnosen never

Hydroiden aus Australien. Zool. Anz; 59,

57-69,

StecHow, E. (1925).—Hydroiden von West- und

Siidwestaustrulien nach den Sammiungen von

Prof. Dr Michaelsen und Prof. Dr Hart-

meyer, Zool. Jahrb. Abt, f, Syst. 50, 191-269.

Trompson, D’A, W. (1879).—On some new and

rare Hydroid Zoophytes (Sertulariidae and

Thuiariidae) from Australia and New

Zeslund. Ann. Mav. Nat, Hist. (s2r. 5), 3,

97-114, Pls 16-19,

TREBILCOCK, R. E. (1928),—Notes on New Zeu-

land Hydroida, Proc. R. Soc. Wier, (s,)

41(1), 1-31, Pls 1-7.

Watson, Jeanette E, (1973)—Pearson Island

Expedition 1969.—9 Hydroids. Trams. R. Soc.

S, Aust. 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 myrtifolia, 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.

rostrata and H. rugosa are rounded spreading shrubs.

SHOOT AND FOLIAGE PRODUCTION OF FIVE SHRUB SPECIES OF ACACIA

AND HAKEA IN A DRY SCLEROPHYLL FOREST

by J. R. Maconocnie*

Summary

Maoonocime, J. R, (1975).—Shoot and foliage production of five shrub species of Acacia and

Aakea in a ary selerophyll forest. Trans. R. Soc. 8. Aust, 99(4), 177-181, 30 November,

1975,

An examination has shown that shoot growth of Acacia myrtifolia, A. pycnantha, Hakee

rostrata, H, rugosa and I. wlicina is distinctly seasonal, Growth commenced in spring, finished

by mid-summer 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 patiern and size of new shoots along

4 parent shoot. A. alicina, the species which showed a tendency toward apicul dominance is.

ah erect, several stemmed shrub, whereas H, rastrata and H. rugesa are rounded spreading

shrubs,

Introduction

This study on shoot production of the five

temperate shrubs, Acacia myrtifelia (Sm.)

Willd., A. pyengntha Benth.,, Hakea rostrata F.

Muell, ex Meisn., H. rugosa R. Br. and H.

ulicina R, Br., was part of a project in which

shoot growth was measured on other shrubs

from the arid and semi-arid areas of South

Australia (Maconochie & Lange 1970,

Maconochie 1973), to obtain basic phenologi-

cal data in a range of habitats.

Study Site and Methods

This study was carried out on the boundary

of the Para Wirra Reserve in the Mt Lofty

Ranges, South Australia from April 1965 until

January 1967. The occurrence and size of new

shoots produced on about 120 tagged shoots

of each species of Hakew and A, pyenantha,

and of 25 tagged shoots on two bushes of A.

myrtifolia, were recorded monthly. The tech-

nique has heen described —_ previously

(Maconochie & Lange 1970).

From 1965 until 1971, samples of shoots

were collected at the end of the growing

season from the three Hakea species and data

recorded on size and position of new shoots,

These data for axillary and terminal shoots

were separated and the annual mean shoot

sizes compared by analysis of variance.

Climatic data were supplied by the Austca-

lian Bureau of Meteorology,

Results

Oudalitative Aspects

The percentage cumulative gains and losses

of foliage for A. pycnantha and A. myrtifolia

are presented in Fig. 1, A: myrtifolia did not

produce any few shoots. in the first year of the

study but the burst of growth during the

second year was in phase with that of A.

pycnantha, The “stepped” effect displayed by

A. pycnaniha was also produced by H.

rostrata, H. rugosa and FH. ulicina. New shoat

growth was. distinctly seasonal.

Figure 2 presents the relative rates. of gain

and loss for the five species. Climatic data are

presented in Fig. 3; soii moisture values were

taken from Martin & Specht (1962).

Shoot growth reached a peak during Octo-

ber when duily temperatures were increasing

and soil moisiure was available (Figs 2 and

3). In both 1965 and 1966, flawering, which

commenced in early August and finshed by the

* Animal Industry and Asricullure Branch, Arid Zone Research Institute.

tory, Alice Springs, N.T. 5750.

2pt. of the Northern Terri-

178

500

400

300

% GAIN

200

100

LOSS

200

300

1965

J. R. MACONOCHIE

1966

Fig. 1. The percentage cumulative leaf gain and loss of A. pycnantha (11, total new positions; 4,

end of September, preceded shoot growth.

Immature flower buds on the Acacia species

were observed io develop as early as January.

Both axillary and terminal flower and shoot

buds on the Hakea species were enclosed in

bracts and these buds developed as the new

leaves matured, This observation suggested

that the size of new shoots for the next seyson

may be predetermined by conditions at the

time of bud development.

Although the A. myrtifolia bushes did not

produce new shoots in 1965, flowering did

take plice.

The peaks in rates of foliage loss coincided

with or immediately followed the peaks of

production, with loss comprising both mature

senescing and soft immature foliage. Some

further loss was caused during a severe wind

and hail storm in December 1966 in which

trees and branches were felled, and also by

bird pruning during July-August 1966, Parrots

were observed pruning all the trees and tall

shrubs, apparently at random, in the area and

in some cases clipping off branches up to 5

mm. thick.

Some losses of H, ulicina and H. rostrata

during January-February 1966 were a result

of the death of several plants. These bushes

were not examined for cause of death, but

since rainfall in 1965 was 223 mm_ below

average it is probable that localised drought

during summer was the most likely cause of

death,

Generally once new foliage had matured,

the rate of leaf loss on the Hakea species

declined to almost zero. The needle-like, rigid,

sclerophyllous leaves of these species are

obviously more resistant to physical damage

than the leaves of the more mesomorphic

species of the community.

Quantitative Aspects

The mean number of leaves per shoot for

cach species and for the years 1965 to 1971

inclusive are presented in Table J. Analysis

of these dala showed that for some years

SHOOT AND FOLIAGE PRODUCTION IN ACACIA AND HAKEA

Wop

PRs Way

4 bow @ lp then oa Ss go ows

Fig. 2, The relative rates of gain (block) and loss

of foliage of (a) 4. myrtifolig, (b) A.

pyctiantha, (c) H. alicina, (a) H. rostrata,

and (@) 7. rugosa, for the years 1965 and

there was a significant difference between the

sizes of shoots produced in successive years.

In other years, the variation in shoot size was

so large that no significance could be attached

to the differences.

A sei of correlation coefficients (r) and

regression equations were computed between

total rainfall (x) for the period Sepiember to

November and the mean shoot sizes (y) for

successive years during which there were sig-

nificant differences (P<.05) between the

means. This rainfall period was selected

because it was the time of active growth,

A. ulicina—axillary shoots

r=+0.74 010> P >.05

y-= 50+ 0.014x

H, rastrata—terminal and axillary shoots

r= +052 P>0.10

y= 5.1 + 0.008x

H. rugosa—terminal and axillary shoots

r=+0.83 .05>P>.02

y> 41+ 0,029x

H. rugosa was the only species showing an

acceptably significant corielation between

mean shoot size and rainfall during the grow-

ing season,

Calculation of the correlation coefficient

between the mean shoot size and rainfall

period Septeniber-November of the preceding

year gave the following: A. ulicina—0.65, H,

roxtrata—O.07, and H. riugosa—o0.55,

Analysis of the data for 1966 on the rela-

tive positions of shoots showed that there were

no significant differences in shoot sizes

hetween 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

und three. It would appear, therefore, that A.

ulicina is the only one of these three species

which has a tendency towards apical

dominance,

Discussion

Cambage (1918, 1927) measured the height

growth of A. pyenantha at Sydney Botanic

Gardens and his study showed the normal

pattern of rapid growth during the juvenile

stage followed by a decreasing rate ag the

plant matured. The plants studied at Para

Witra were mature and the rates of growth as

reflected in both Figs 1 and 2 were of a

“steady state’ nature, as occurs at the plateau

of the sigmoid growth curve.

Martin & Specht (1962) measured the

moisture relationships in a dry selerophyll

forest, of which these species ate shrub com-

ponents. Their studics showed that the more

mesic community of this vegetation type had

a higher index of evapotranspiration and could

be subjected to a drought period during the

mid-summer. The thtce species of Makea and

two Acacia species all show a distinct season-

ality of shoot growth with a cessation occur-

ring in mid-summer probably during the

drought period or when soil moisture is only

sufficient to maintain a dormant growth phase.

The negative correlation between mean

shoot size and soil moisture (as reflected by

180 J, R. MACONOCHIE

200

150

100

TEMPERATURE °C

RAINFALL (mm)

J F M A M J J A $ 0 N D

Fig. 3. Climatic data of Parra Wirra study site; soil moisture ex Martin & Specht (1962), other data

tial evapotranspiration (P/E 0.75); , florescence and duration of active growth.

TABLE 1

Mean number of leaves per shoot for seasons 1965 to 1971

Shoot Year

Species Posn. 1965 1966 1967 1968 1969 1970 1971

Hakea ulicina Total 7 A* 8.1* 8.2 8.7 9.2 9.4 98

Terminal 9.6 10.2 9.A* 9.4 10,9 10.6 11.2

Axillary 5.9T TAY 4.92 8.1* 6.77 8.9 8.4

Hakea rostrata Total 6.47 5.87 5.2 5.45 6,5 6.74 9.0

Terminal 6.1 6.1 5.7 5.4¢ 7.0 6.54 10.6

Axillary 6.5% 5.7 5.2 5.4¢ 6.4 6.8" 75

Hakéa rugosa Total 7.24 GA 4.64 8.7 8.47 9.24 13.7

Terminal 7.3% 94 8.1 8.8 8.8 9.5 15.9

Axillary 6.94 9.0% 4.42 8.8 8.3* 8.9t 12.7

Significant differences between mean and mean of succeeding year are indicated.

*p < 0.05; f P< 0.01; 1 P < 001.

Meun nuinber of leaves per sheot for terminal Se cticds positions distal from shoot tip for year 1966

Relative

Position Term. AX, AXa AX: AXy AX; AX, AX; AXe

A. ulicina 109* 78 5 7.2 7.6 74 6.6 1.6 5.6

H. rostrata 6.2 6.0 53 5.6 5.3 55

H. rugosa 93 8.6 9.6 10.2 103 8.2

* Indicates significant difference (.05 > P > .01) between mean and that of succeeding position.

SHOOT AND FOLIAGE PRODUCTION IN ACACIA AND HAKEA

tuinfall) during the growing period of the

previous. year suggests that shoot size in the

following year is not necessarily determined

at the Eud formation stage, Rather these

results suggest that shoot size is more likely

to be determined by the soil moisture during

the period of active growth.

The time of shoot growth for these species

contrasts with that of heath studies of Specht

(1957), Specht & Rayson (1957) and Groves

(1965). who recorded that the shoot growth

commenced im December when ‘soil moisture

was decreasing. Specht (1957) showed that

drought conditions occurred in both December

and Jamiary. Groves (1965) noted that shoot-

growth continued throughout the summer of

his study period; however a recharge of soil

moisture from a mid-summer rain was

recorded.

Maconochie & Lange (1970) and Macon-

ochie (1973) have reported the seasonality of

shoot growth on 4. sowdenti, A. ligtdlata and

l, murrayana, and possible non-seasonal and

seasonal shoot growth responses on A. anenra

and A. kempeana, Wetherell (1966) recorded

flushes of growth on A, Aarpophylla in Quecns-

land during spring and summer, and during

ane period of early winter. A. pyenatha, by

181!

contrast, actively grew only during the spring

period at the study site, and although shoat

growth en A. myrtifelia was only recorded

during the secand year, it appears. that from

this slender evidence that the shoot srowi{h

follows a scasonal pattern alsa,

Both A. rostrara and H. regasu have a more

spreading bushy habit than AH. uficina which

has a tendency to be more erect, The habit

of these plants is reflected in the sizes of new

shools on a parent shoot. Both H. rostrata and

H. rugosa did not produce significantly smaller

axillary shoots in comparison to the terminal,

but the terminal shoots of HW. ulicina were sig-

nificantly larger than shoots on the three suc-

ceeding positions below the apex. This suggests

a tendency towards apical dominance and

thus explains the more erect habit of this

species.

Acknowledgments

The author is indebted to Dr R. T. Lange,

Botany Department, University of Adelaide,

for his encouragement in the first part of the

project. Dr R. W. Rogers, now of the Univer-

sity of Queensland, supplied samples during

the years 1967 to 1970 and his assistance is

gratefully acknowledged.

References

Campace. R_ H. (1918).—The vertical] growth of

Irees. I. J. Proc. R. Soc. N.S.W, 52, 377-384.

CamaBace, R. H, (1927).—The vertical grawth of

trees. I. Proc. R. Soc. N.S.W. 61, 279-284.

Groves, R, H. €1965)—Growth of Heath Vege-

tation, (1. The seasonal growth of a heath on

ground-water podzol at Wilson’s Promontory,

Victoria. Aust. J. Bot. 13, 281-289.

Maconocuig, J. R... & Lance, R. T. (1970),—

Canopy dynamics of trees and shrubs with

particular reference to the arid-zone topfeed

species. Trans. R- Soc. §. Aust. 94, 243-244,

Maconocuie, J. R. (1973)—Leaf and. shoot

growth on Acaeia kempeana F, Muell. and

selected other arid-zone species, Traplcal

Grasslands 7(1), 49-35.

Martin, Helene A., & SpecHr, R. L. (1962).—

Are mesic communities less drought-resistant?

A study on moisture relationships in dry

sclérophyll forest at Inglewood, South Aus-

tralia. Aust. J. Rot. 10(2), 106-118,

SPEcCHY, R. L., & Rayson, Patricia. (1957).—Diurk

Island Heath (Ninety-Mile Plain, South Aus-

tralia). T. Definition of the ecosystem. Ams.

J. Bot, §, 52-85,

Sercur, R, L. (1957).—Dark Island Heath

(Ninety-Mile Plain, South Australia). V. The

water relutionships in heath vegetation and

pastures on the Makin Sand. Aust. J. Bat. §,

51-172,

WETHERALL,, A, J. (1966).—Leaf growth of Briga-

low (Acacia harpophytia) suckers in relation

to seasonal conditions. Qld J. agric, anim,

Sei, 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.

DeLkunty, J. A, (1975).—Shoreline shingle terraces and prehistoric filling of Lake Eyre,

Trans. R. Soc, §, 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) ofd shoreline shingle

terraces built by prehistoric fillings of the Recent salina to levels above thal of 1974.

Evidence indicates at Icast three prehistoric fillings to levels of 280, 160 and 70 cm

above that. of the 1974 fillmg. No precise evidence of the ages of the prehisloric Nilings way

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 Jake, and late Recent subsidence of its sonth-

western areas, was also inferred from field observations.

Introduction

Lake Eyre consists of a large northern area

—Lake Eyre North, connected by a narrow

channel to a small southern area—Lake Eyre

South (see 1:250,000 Topographical Sheets.

Noolyeana, Lake Eyre, Curdimurka). It is a

salina forming the “sump” of a large infernal

drainage system (Bonython 1955; Mason

1955: Wopfner & Twidale 1967). Aridity and

high rate of evaporation, over the greater part

of the drainage area, normally prevent river

water from reaching the lake. On infrequent

oceasions when rivers flow into it, part or

whole of the lake bed is covered with water

which cannot escape and must evaporate. The

bed of Lake Eyre North slopes gently from

north to south, and evaporation of brines in

the southern bays has produced salf crusts

(Dulhunty 1974; Bonython 1956).

When the lake contains water, its shores are

actively eroded by wave action, producing

well-defined shorelines with shelving strands

or low cliffs rising abruptly from its almost

flat bed. At some places, cliffs up to 10 m or

more in height have been cut in soft, partly

consolidated sediments. At other places shore-

lines lie along, or around the ends of, longi-

tudinal sand ridges rising steeply to as high as

LO m above the lake bed.

Around Babbage and Hunt Peninsvlas and

in Belt Bay (Fig. 1A). shores are being

eroded in hard dolomite, silerete and ferru-

vinised siltstone (Williams 19731), Wave

action, during periods of filling, has. produced

quantities of pebbles or shingle consisting

largely of slightly flattened ond rounded

pebbles from 2-10 om in diameter. Single

strands or beaches have developed, and break-

ing waves have washed up actoss the beaches

long parallel dcposits of shingle, These

deposits, formed during the present salina

episode of the lake’s geological history, as dis-

cussed later, are Recent to coniemporary in

age. They are wave-built, strand, shingle ter-

races forming ridges, terraces, ramparts and

banks, closely associated with the present

shoreline of the salina. For the purpose of the

present paper they are referred to as “the

shoreline shingle terraces of Lake Eyre salina”.

Gravel deposits wer¢ observed and recorded

by Williams (19737, 1975) on Hunt and

Babbage Peninsulas and on the western side

of Belt Bay. He described them as “strand

gravels” and “old shoreline deposits”. They

* Department of Geology and. Geophysics. The University of Sydney, N.S.W. 2006.

‘Williams, A. F. (1973).—Explanatory Notes for the LAKE EYRE 1:250,000 Sheet. S. Aust. Dept.

Mines, R.B, 72/93 (unpubl,),

184

include the Recent shoreline shingle terraces

described in this paper, and wther gravel

deposits. possibly at higher levels, and more

remote from the presenf shoreline, which may

be older and related to pre-Recent history of

the Lake. Williams (pers. comm. 1975) also

noted shorcline shingle jerraces along the

southern shores of Lake Eyre South, which

he regards as probably similar in age and

origin to those of Lake Eyre North described

in this paper, He also found and cxamined

more remote termiees up to 20 km from the

present lake shore.

The filling of Lake Eyre in 1974 to the

greatest depth known since European settle-

ment, provided a unique opportunity for the

study of contemporary shingle terrace develop-

ment in relation (6 lake levels and wave action,

This was undertaken to gain a better under-

standing of the nature and origin of the old

shoreline terraces and enable interpretation of

their significance regarding the maximum

depth to which the lake had been filled during

its. role us the salina of the present internal

drainage system, and the time since it was

filled tu, or above, the level of the 1974 filling.

Such investigations and results, recorded in

this paper, ate confined to the significance only

of terraces along present shorelines. and do

not represent a comprehensvie study of all

shingle dennsits and their regional disteiburion

around lake Eyre,

Investigation of shingle terraces

Conremporary wean lake level

During the latter half of 1974, water level

around the shores of the lake provided a hari-

zontal datum, of any one time within certain

limits af variation, which was used extensively

in determining levels of shereline shingle ter-

races in relation to each other and ta mean

lake level, Short term varialions in water level

were caused by wind tides, currents and

possibly ulher factors, Wind tides were most

pronounced on windward shores where water

level rose and full, within a maximum range of

about 45 em, very quickly with rise and fall

of Wind velocity wnd wave height. When wind

dropped, wave action usually ccased and water

fevel fell to “normal” within 5 10 6 hours,

“Nornval” water level or the level free of wind

effects during periods of calm, varied as a

result of ather factors within a range of about

20 cit, To view of this il was ussumed. for

the purpose of the present investigation, that

J. A. DULHUNTY

water level ut any point along the shore,

meusured after a calm peria¢d of not less than

6 hours, could be regarded as mean lake level

plus or minus 10 cm,

Contemporary shingle terraces

Studies of shingle terrace development

associated With the 1974 filling of Lake Eyre

were carried out in August of that year. Mean

lake level reyched its maximum during May.

It rose by only about 20 cm during April,

remaining steady during May and fell by

abour 20 cm during June and July. Therefore,

the shingle terraces studied in August had been

built Over a period of four months during

which mean jake level rose and fell through

only about 20 cm.

Well-defined shingle terraces were built

along all shelving shores with shingle beaches

around Hunt and Babbage Peninsulas, and in

Belt Bay. They were built during periods of

strong wind, by waves breaking and washing

shingle up across the beaches to form terraces

above mean lake level. It was evident that

shingle moved slowly up the beaches as the

lake filled, then formed into fully-developed

terraces during the maximum mear fake level

petiod, and finally remained stranded when

water fell, providing evidence of the highest

level reached,

The !974 contemporary shingle terraces

varied in with up to 500 cm, and in depth

or thickness up to 150 cm ahove pre-existing

beach material. The height of their tops above

maximum mean lake level attained during

their development, reached a general maxe

mum of 90 cm (+ 5 cm) on exposed shores.

On sheltered shores, terrace tops were lower.

All the terraces formed al any one time, at a

given mean lake level, could be regarded as a

group, although the height of their lops above

mean lake level varied from place to place

depending on degree of exposure to wave

action. It Was found that the general Tevel of

the tops ef the highest terraces, on exposed

shores. Was a uniform feature of a group, and

the most significant feature in determination,

after the water level had fallen, of the mean

luke level at which they had lormed, There-

fore, it was concluded that the maximum pcan

luke [evel of any pase filling would be very

close 10 Of) em below the general level of the

original tops of the highest shingle terraces

produced during that filling,

Greater depths of water during prehistoriy

fillings ate net likely to have produced waves

SHINGLE TERRACES AND FILLINGS OF LAKE RYRE

ear

=Q)- 4

WALIGAN

BL +

ANY THON HES,

PREHISTORC “=SRACES -—

RIGHEN |

. 330UF —MIDRLE

ain GRE

SuWwesT

Gyour

Un CMAIRARY

WS?0)TERqaLe

-—= 7" 500"

SANKY SHINGLE

LAKE

FILLINGS =

oa -—- "3000" +h BF

=== ane" YR BF

TR BF

> >> PRESENT (1941

QUATERNARY

SFBIMENTS

PREHISTORIC SHINGLE TERRACES (\ CONTEMPORARY (:g7a)

SHINGLE TERRACE

POWEST GRauP

Maa Gay

CULE

WEDL = SR OUP

HIGHEST ONE

be Weiter

SKETCH PLAN SHINGLE TERRACES un WILLOW Sav

PSPHUSTORIC TERRACE ot HiGPEST GROUP

SUBSEQUI NT

CRET RK

fel SONGECUEN'

CREER CHANNEL

a a

— “BXTENDP)

f] i

D SECTION F~ CIR EXTENSION) ACHUSS SHINGLE TERRACES

qa5 -loozamire — [--|—I

elisa Nea

LF Ld

C SECTIONS Ae

i—}

5 f

—& 31=C ON FLAN B

Fig. 1. Wave-built shoreline shingle terraces in Lake Eyre North.

of greater amplitude than in 1974, or terraces

with tops more than 90 cm above mean lake

level, as the 1974 water depth was more than

one half the wave length of average strong

wind waves. This was sufficient ta allow

development of waves as high and long as the

limited fetch across the lake could produce.

Increases in depth of water, within the limits

of beach heights, would not appreciably in-

crease distances of water across the luke as the

shores are relatively steep. If average strong

winds during prehistoric fillings were similar

to those of today, heights and lengths of waves

must have been similar ta those which built

the present day terraces.

Old shoreline shingle terraces

At many places along the shingle-strewn

shores, old partly eroded shingle terraces occur

at levels above that of the 1974 terraces, fol-

lowing contours across sloping terrain behind

and above the 1974 beaches. Excellent

examples occur along the eastern shoreline of

Hunt Peninsula between Willow Head and

Campbell Point, immediately north of the

Elliott Price National Park fence, at numerous

places on Babbage Peninsula and Bonython

Head in Belt Bay, and on Silerete Island off

Bonython Head (Fig. 1A), Although eroded,

the old prehistoric shingle terraces all bear

close resemblance in general form and nature

to the 1974 terraces, and follaw contours con-

forming to the present shoreline. They were

formed in & manner similar to the 1974

terraces, by previous fillings of the lake to

depths greater than that of 1974 which was

the deepest known filling in white man's his-

tory. Wave action which formed the prehis-

186

lyric Jervaces would seem to have been similar

io that of the 1474 Niling, as already discussed.

Lower rales of evaporation in orchistoric mid

io lale Recent! time could have resulted in

greater duration of peak lake-level periods and

Songer tines of terrace formation, than in

1974. However, this would not necessarily

have produced terraces higher than the equilib-

rium height, at a given mean lake level, for a

given wave height, From observations,

vquilibrium terrace height appeared to have

beer teached after 4 months in 1974, So the

terrace top height of 90 em above mean lake

level, altained in I974, is assumed to have

heen the equilibrium height attained in the fowr-

mation of the prehistoric terraces, Whilst

longer time of terrace Formation is not likely

io have produced highet terraces, it could well

have built wider terraces in association with

wave cut benches. However, after allowing

for erosion, the old terraces do not seem to

have been wider than the contemporary ones.

and there 1 no general evidence of asaoctated

wave cut benches

Meusurements of the relative levels of old

shingle terrace tups at many places indicated

the ocvurrence of three groups at successively

higher levels above the 1974 terraces and

naXximino mean lake level, They are described

as the lowest, middle and highest groups for

the purpose of this paper (Fig. 1B, D). Each

is considerably more eroded and obviously

alder than the one beneath it, They represent

three prehistoric fillings of the lake to levels

higher than that of 1974. No higher terraces

were found on the presen! shoreline slones of

Hunt and Babhave Peninsulas, or on Silerete

Island, Strand gravels recoriled by Williams

(1973!) at distances up ta 8 km from the

present Izke shore, lying outside the field of

this investigation, contain Cexlella which is

absent in the recent shoreline terraces, They

may have been disturbed by fectonism, but

could represent older, possibly Pleistocenc.

water levels of an carlier phase of lake history

as discussed later.

Tt was concluded that the highest group of

old shingle terraces on the present shorelime

slope was built by the deepest filling of Lake

Fyre during, the Recent ycological history of

the salina as we know it to-day. Terraces of

this group were studied tn grcatest detail to

ascertain as precisely as possible the height

and age of the deepest prehistoric filling of

the present salina.

5. A. DULHUNLY

Mostly the old shingle terraces of the

highest group have been greatly eroded or

completely removed. However, in protected

places, where less erosion has occurred, their

tops reach uniform maximum levels which

vary in different areas fram 280 em to 350

om ahove the 1974 maximum mean lake level.

They reach Icvels of 351) cm along both

eastern and western sides of Hunt Peninsula,

320 em on both sides of Babbage Peninsula.

and 280 em on Silcrete Island. Comparison

with the shape and yeneral nature of the 1974

terraces sugvests that the least eroded of the

old terraces, which were those selected for

measurement, have fost unly about 20 cm from

their tops. This means that the original maai-

mum level of the old terraces was 370 cm on

Hunt Peninsula, 340 cm on Babbage Peniu-

sula and 300 cm on Silcrete Island. It is

evident that the highest group of old terraces,

in each of the three areas, is of the sume age,

having been formed during the same prehis-

foric Alling. Therefore, the present differences

between the terrace-lop levels of the highest

group, in the three areas, suggest subsidence,

increasing to the west. since the filling which

formed the terraces, but investigations over

witler areas of the Take must be carncd oul

before any general conclusions can be reached

about Recent tectonic movements. The pur-

pose of the present investigation was to gain

some knowledge about prehistoric fillings,

Therefore, Hunt Peninsula was selected as the

most centrally placed area of old shingle ter-

races, ahd likely ta have heen least affected hy

icctonic movements knawn to have been

prevalent along the western side of Lake Eyre

(Williams 19734; Wopfner & Twidale 1967)

V'rehistoric Fillings

As iliscussed above, the terruce-top level of

the highest group of old shoreline shingle

terraces on Hunt Peninsula was 370 cm above

the peak mean lake level of the 1974 filling,

and the contemporary 1974 shingle terracc-top

level was 90 cm above the mean lake level al

which the terraces were formed (Fig. 1D).

Therefore, the peak mean lake level of the

prehistoric Alling which built the highest group

of old shingle terraces must have been very

close ta 280 env above that of the 1974 filling

This means that the peak mean lake level of

the highest prehistoric filling of Lake Eyre

éalina, in Recent lime, would have reached a

depttr of about 640 cm above the lake hed

along the southem shores of Madigan Gulf.

SHINGLE TERRACES AND FILLINGS OF LAKE EYRE (87

about 850 cm in the deepest parts of jhe gulf,

and a level of approximately 286 em above the

1974 Water loVel in Lake Fyre Semth and along

the Central Australian Railway line.

The levels of the terrace tops in the middle

and lowest of the three groups of terraces on

Hunt Peninsula measured 230 cm and 140-cm,

respectively, above the 1974 peak mean lake

level. Allowing for 20 cm of erosion, and 90

cm for the height of the terrace top levels

above the mean lake levels at which they

were built. the two groups would represent

fillings of the Jake to depths of 160 cm and

70 cm ubove the 1974 peak mean take level

(Fig. 1D),

The three groups of old shingle terraces,

described above, were built at widely spaced

intervals of time, during which many other

major fillings must have built terraces, ar inter-

mediate and lower levels, which were prabably

destroyed or modified by redistribution of

shingle during subsequent deeper fillings. Any

teraces built by a filling to a level intermediate

between the depths represented hy the lowest

and middle groups, during the interval

between the building of the middle and

highest groups, were partly or wholly

destroyed by the filling which built the middle

group. Similarly terraces built by fillings to

depths less than thal of the 1974 filling, includ-

ing terraces of the 1950 filling, have been

modified or removed by the 1974 filling,

Antiquity of fillings

The ages of the three prehistoric fillings of

Lake Eyre, to depths preater than that of the

1974 filling. are very difficult to ascertain pre-

cwely. The only available evidence is very

vague and indefinite, and ihe only conclusions

possible are of the order of magnitude of age

tather than exact periods of time,

Extensive searches were made for

carbonaceous material within the old shoreline

shingle iectaces, sujlible for carbon dating, Nu

sheil material or aniinal bones were found, No

plant material such as driftwood was found,

although sticks and fragmentary pieces of

wood were plentiful in the contemporary 1974

lerraces, Plant roots were found in varying

degrees of decay, but it was evident that they

belonged to plants which grew on the terraces

after they were built. The environment of

burial in the old shmgle terraces was evidently

one of maximum aeration, oxidation and pene-

tration of rai) water, leading 1 complete des-

tructton and loss of all plant material

originally |neluded in the terraces.

Shell inaterial Was absent from both con-

temporary and old terraces, sugeesting that the

Jatter were built by iAtermittent fillings

between which aridity and dryness of the

salina prevented establishment of shell fish,

The more remote and higher travel deposits

described by Williams all contain small gastro-

pod shells (Williams 1973!', and pers. comyn.

1973) supporting the suggestion that they were

deposited during an earlier pre-salina stage

when the Take was permanently filled with

Water varrying 2 shell fish community, Studjes

by King (1956, }960) suggest that gypseous

lacustrine clays with Coxiella occurring along

the southern shores of Lake Eyre are of

Pleistocene age, and that younger shoreline

deposits and sand ridges ogiginated in associa

tion with recent. deffation of the lake and

development of the present salina,

Fluvial erosion of the old shingle terraces

bears evidence of their antiquity, The highest

and oldest group has been extensively eroded,

and completely removed in many places, whilst

the middie and lower groups have cach suf-

fered successively less erusion. Rainfall is very

low, bul this is offset by high runoff {rom arid

surfaces. Fluvial erosion woder these condi-

tions could be comparable with thar of higher

rainfall areas where ronoff is seduced by

vegetation. Wind erosion is not effective as the

shingle is too large to be temaved,

Where shingle terraves of the highest group

have not been compleiely removed, they have

been breached in places by flowing Waler duor-

ing Fare occasions of heavy rain. Terraces

built across pre-existing or antecedent creeks,

trend upstream Jn U-hends. as would contours.

Where breached, their eroded ents still turn

upstream (Fig. 1B), Terraces built across

gently sloping surfaces with no pre-existing

creeks, have beet breached in places by water

hefd back on the sloping surface, leaving the

eroded ends of the terrace “in-line” and nor

Girned upstream (Fig. 1B, C). Such breaches

are frequently about 8 m wide, and the newly

formed subsequent creek channels running

through the breaches have eroded down into

underlying weathering dolomite to depths of

150 cm below the bases of the terraces, Tribu-

tary consequent creeks carrying water alony

the top sides of the terraces towards the

breaches have eroded channels in dolomite to

50 cm deep (Fig. 1C). Whilst erosion of this

kind ig very difficull to refaie In time, it is

evident (hat the terraces concerned are of can.

siderable antiquity.

tsa

The close proximity and relations of the old

shingle terraces to: the contemporary terraces

indicate very little modification of the shore-

line in general since their formation, suggest-

ing limited age,

Exudation of highly mineralised ground

water, due to aridity, has produced much sur-

face and shallow sub-surface cementation of

Pleistocene and carly to mid Recent sediments,

Gypcrete, silcrete, calcrete and ferruginite have

been formed (Williams 19731; Wopfncr & Twi-

dale 1967). and the processes still appear to

be in operation, No cementation has been

found in the old shingle terraces or the more

sandy and earthy material forming their basal

layers, suggesting limited antiquity,

The evidence outlined above is not at all

precise or satisfactory in attempting to estab-

lish ages in yeays, but it is all that is, as yet,

available. For purposes of geological history,

the evidence is more significant, indicating that

the old shoreline shingle terraces are almost

certainly of mid to late Recent, and most

probably late Recent, age, For the purposes

of physical geography and hydrology of Lake

Eyre, estimates are required for (i) the age

in years of the deepest prehistoric filling of the

salina under existing environmental conditions,

and (ii) the number of years since the lake

had been previously filled to the level of the

1974 filling,

J. A. DULHUNTY

After making extensive field studies, taking

all factors into actount, and balancing the

somewhal conflicting significance of different

aspects of evidence, it was concluded that the

oldest and highest group of shingle terraces

could date from more than 1,000 years before

present, but less than 5,000 years, Purther

narrowing of these Limits must involve inferred

speculation, but in the author's opinion the

age may be of the order of 3,000 years, which

provides um estimate of the age of the deepest

prehistoric filling of the present salina. The

middle and lowest groups of old shingle

terraces are certuinly younger, and could well

be of the order of 1,000 and 500 years, respec-

lively. The age of 500 years for the lowest and

youngest group of old terraces, occurring just

above the contemporary 1974 terraces, pro-

vides an estimate of the time since the Jake

had previously been filled to the level of the

1974 filling.

Acknowledgments

lu is wished to acknowledge (1) valuable

assistance and co-vperation of Mulooring Sta-

tion which made field investigations possible,

(ii) valuable discussion with Mr A. F.

Williams of the Geological Survey of South

Australix, und (i) research facilities provided

by the Department of Geology and Geo-

physics, University of Sydney.

References

Bonytnon, C, W. (1955),—in “Lake Eyre, South

Australia, The Great Flooding of 1949-1950",

The report of the Lake Eyre Committee, R.

geogr, Soc Aust. (S. Aust. Branch), pp- 7-9,

27-56, 63-70, (Griffin Press: Adelaide.)

BonwyrHon, C W. (1956).—The Salt of Lake

Eyre—its occurrence in Madigan Gulf and

its possible origin. Trams. R. Sac. S. Aust.

79, 66-92.

Durpunry, J. A, (1974),—Salt crust distribution

und lake bed conditions in southern arcas of

Toke Eyre North, Lrans. Ro Soc. §. Aunt.

98(3), 125-133.

King, D. (1956)—-The Quaternary stratigraphic

record at Lake Eyre North and the évolution

of existing lopographic forms. Trans. R. See.

S, Aust. 719, 93-103,

King, D, (1960)—The sand ridge deserts of

South Australia and related Aeolian Jand-

forms of the Quaternary arid cycle. Frans, R.

Sov, 8. Aust. 83, 93-108.

Mason, B, (1955).—Jn “Lake Byre, South Aus-

(ruliu. The Great Flooding of 1949-1950".

The Report of ihe Lake Eyre Commitice, R.

geogr. Soc. Aust. (S. Aust. Branch), pp. 1l-

26. (Griffin Press: Adelaide.)

Wiuurms, A. FP. (1975) —LAKER EYRE map-

sheet. Geological Atlas of South Australia,

4:250,000 series. Geol, Surv, &, /lust.

Worrrer, H., & Twipare, C. RB. (1967 ),—Gao-

morphological history of the Luke Eyre

Basin, In J. N. Jennings, & J. A. Mabbutt,

Tds,. “Landform studies from Australia and

New Guinea", pp. 118-143. (A.N,U, Press,

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 RELICT PALM LIVISTONA MARIAE F, MUELL. IN

CENTRAL AUSTRALIA

by P. K. Latz*

Summary

Latz, P. K. (1975) —Notes on the relict palm Livistona mariage 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 io 56 ycars indicate that the relict

palm Livistone mariae may reach an age of up to 30U years. Aspecta of the ecology of the

Palmi are presented.

Introduction

Palm Valley, situated in the Finke Gorge

National Park in the MacDonneli Range sys-

lem, Northern Territory (Fig, 1) is of con-

siderable interest to tourists and scientists alike

lurgely because of the presence of a stand of

the relict cabbage palm Livistona mariae F,

Muell. This species of Livistona is restricted

lo an area of about 60 km? on the Finke River

and its tributaries. It is a relict species separated

by about 1000 km from the nearest Livisfona

to the north, The closest relative to LZ. mariae

occurs on the Fortescue River in the Hamersley

Ranges, W. Aust., a stand in many ways simi-

lar to that at Palm Valley. The relict nature of

the palms is discussed by Keast (1959), Bur-

bidge (1960) and Chippendale (1963).

Although the explorer Ernest Giles was. the

first to discover the palms in the Finke Gorge

in 1872, he almost certainly bypassed Palm

Valley itself, and its discovery is attributed to

a Lutherun missionary from Hermannsburg

Mission some time Jater, The valley was in-

vestigated by members of the Horn expedition

during 1894 (Tate 1896). After a visit to the

Valley, Lothian (1959) discussed aspects of

the palm's reproductive behaviour.

Chippendale (1959) listed 200 plants found

lo occur in Palm Valley. Since this time a total

of 333 plant species have been recorded from

the valley (about 4 quarter of the total number

of species recorded for the whole of central

Australia!), About 10 percent of these species

can be considered to be of rare or restricted

distribution in central Australia, The majority

of these rarer species are restricted to areas in

the valley fed by permanent water seepage.

Recently a large gas field has been dis

covered in the arca adjacent to and north of

the Park, Two gas bearing wells are situated

only a few kilometres north of Palm Valley and

tapping of these wells for commercial produc-

tion is being considered.

The Habitat

The palm has a shallow fibrous root system

and is situated in an arid area. Therefore u

suitable habitat for tts continued survival must

have a permanent shallow water supply over

an area large enough to support a viable breed-

ing population. The grea must also be prco-

tected from severe erosion forces or have a

stable soil environment, as the palm will not

survive scouring of surrounding soil.

Palm Valley appears to be one of the very

few areas in the central ranges which mects all

of these requirements. The permanent water

supply in the valley appears to be associated

with the peculiar stratification of this area. The

gently sloping conglomerate and sandstone

strata percolate water fram a large catchment

area, down to where the valley dissects these

strata. The position of this dissection ensures

a continuous permanent seepage arca along a

considerable distance of the valley floor and

lower slopes. Although other springs and

seepage areas occur in the Ranges, few, if any,

extend over such a continuous urea, The Finke

River bed below Palm Valley has a permanent

* Arid Zone Research Institute, Animal Industry and Agricultural Branch, Department of Northern Aus-

tralia, Alice Sprinus, N.T, 5750),

190

P. KR. LATZ

= oy @

B vete

= a

} ‘, : ¢

4 \ \

I ~~

} K — ny

ty 6 : J }

~.* f

cr e /

J f

mY ~

Mar SOCALITY

a 4

NORTHERN

TERR LOY

Polm vellvy

“yas

cyeat Terge Ul ad If

oe sana Te -~

i Ry!

LFESFHD

@ 50 Fame

#22 Poms

Rare saterbary,

Fig. 1. The distribution and numbers of mature individuals of the cabbage palm Livistona mariae in the

Finke Gorge National Park, Northern Territory,

shallow water tuble for much of its length, but

it is subject to severe water wash and the few

palms found there are restricted to areas

which escape the full force of the hoodwaters.

Palm Growth Rates and Ages

The author was fortunate in obtaining a

number of accurately dated photographs of

vertuin areas of the valley taken as early as

1917. Exact relocation of many of the source

points was made possible by the cliff back-

grounds of most of the photographs and the

fact that the narrow valley is restricted in the

number of photographic vantage points,

Of the 13 photograph sites relocated, three

are presented in Figs 2-4. ‘The palms in Fig.

2 show the least change of any of the palms

investigated and almost all of the individuals

present in 1917 (when the photograph was

tuken) are now still present. The present

heights of the living palms were estimated with

a Suunto clinometer. By using the cliff back-

yround as a scale, palm heights at the time of

the carlier photograph were estimated. Distor-

tion due to angle of viewing and fo Jens aberra-

tion was estimated 19 be Jess than 10 percent,

A mean annual growth rate of 10 ¢m was esti-

mated for this group of palms.

Palms in Fig. 3, however, have grown

approximately 114 m im 38 years, a meun

annual growth rate of 30 cm. Pulms in Fig, 4,

which apparently were absent in the earlier

photographs, taken in 1918, ure now up to 12

m high, a minimum mean annual growth rate

of 22 cm.

Palms grown trom seed in Miami, Florida,

on the bank of a permanent stream, grew to a

height of 9 to 18 m in 27 years—an excep-

tional growth rate of up to 60 cm per year

(Lothian 1959). A palm in the cooler climate

of Melbourne Botanic Gardens is reputed to

have grown to only 3.75 m in 30 years. a

growth rate of 12.5 cm per year (Australasian

Post, 31 May, 1973).

In its natural habitat, the rate of growth of

the pulm appears to be mainly dependent on

the water supply. The palms showing the fastest

growth rate (Hig. 3) are at Palm Bend on the

bank of the Finke River about 6 km east of

the Valley. This urea has a greater depth of

fertile alluvial soil and a better water supply

than most areas in Palm Valley. Pulms in Fig.

4 are situated on one of the best spring areas

in the valley, whereas those in Fig. 2 occupy

one of the drier areas of the valley, as is

reflected by their slower growth rates.

The tallest palms in the valley have reached

a height of 25 m, Making allowances for dif-

ferent growth rates at various ages, 100 to 300

years seems a reasonable estimate of the age

of the oldest individuals of Livistona mariae,

THE RELICT PALM LIVISTONA MARIAE 191

AGA

Fig, 2, The upper photograph was taken in 1917 and the lower in 1973,

Effect of Fire

The areas of high palm densities are sus-

ceptible to fire as large amounts of plant debris

accumulate around the base of the palms. A

small area of the valley was subjected to a

natural fire in the early part of 1973 and the

area was visited by the author approximately

2 months after the event, The fire appeared to

have been intense; a palm 28 m high was burnt

at the crown and thick ashes (up to 40 cm

deep) were found at the bases of the majority

of the palms.

Ninety-six established palms were affected by

the fire, thirty of which subsequently died. To

ascertain whether certain height classes of the

palms had higher survival rates, the heights

of all affected palms were measured with a

Suunto clinometer. Results indicated that the

palms in the height range of 5-8 m had greater

mortalities than those in the 1-3 m_ height

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-

nificant difference between the two height

populations.

A localised fire was reported in the same

area in the 1940's and this fire was severe

enough for the smoke to be seen from Her-

mannsburg Mission, 12 km distant and on

the other side of the Ranges. After several sub-

sequent rains, regeneration of palms was

reported to be abundant. (A. Latz, pers.

comm.) A fire is also known to have occurred

through the stand in Little Palm Creek early

in 1959, but little evidence of this fire is now

apparent.

It does appear that many of the palms can

survive a fire and subsequent regeneration is

quite rapid. However, an excessive accumula-

tion of debris over a long time period could

provide enough fuel for a more severe and

extensive fire than those previously observed,

and could effect a higher palm mortality. If this

fire followed a few years of poor seed produc-

THE RELICT PALM LIVISTONA MARIAE

193

Fig. 4. The upper photograph was taken in

relocated by using the large rock ri

marker. This rock is hidden by palm

tion, or if a severe flood subsequently removed

the majority of seeds, palm densities could be

severely reduced.

Palm Numbers

Tate (1896) stated, “ . . . Livistona mariae

is known by only one colony estimated to com-

prise not more than a hundred mature indi-

viduals”. A few were also observed by him

along the Finke River as far as Boggy Hole.

Keast (1959) estimated a total of at least 300

1918 and the lower in 1973. This area was

ight of centre in the earlier photograph as a

s in the later photograph.

individuals at all stages of growth were present

in the valley.

An exact count of all individuals in the

valley is impractical because of the high den-

sities of the palms in some areas and the many

younger palms at all stages of growth. How-

ever, individuals of reproductive age (ca. 3 m

or taller) were found to have at least some of

their trunk clear of leaf debris and are rela-

tively easy to count. A count made early in

194

1973. indicated that about 750 palms of repro-

ductive age oceur in Palm Valley,

This is a much greater number than those

previously given. Several recent visitors to the

valley were asked for their estimates of palm

numbers and these ranged from 30 to 800 with

an average of 150, These observations tend 10

show that rough estimations give a lower

number than is actually present and may

uccount Jor the previous Jow estimates. How-

ever. the early photographs, including those

not presented in this article, do show lower

palm densities than those at the present time.

Stirling (1896) reported that the white basal

parts. of the inner leaves of the young palms

were eaten by aborigines in the area. This

would have caused the death of at least some

of the palms. This practice has now ceased, In

the past, aborigines frequently lit fires to hunt

game, stimulate regrowth of some edible plants

and signal their presence to other members of

ihe tribe. Only two Gres of restricted extent

are known to have occurred in Palm Valley

during this century. Before this time the

aborigines may have burnt the area more fre-

quently to stimulate growth of young palms

which could then be used for food,

Contrary to Tate’s observations, there are

as many palms outside Palm Valley as in the

valley itsel€ (Fig. 1), The second largest

colony is in Lite Palm Creck where 550

mature individuals have been counted. Except

for about 6 palms further downstream in the

Finke River, the focation of all individuals of

Livistona mariae known to occur naturally is

indicated ly Fig. 1. All of the palms of repro-

ductive age have been counted vnd total 1500.

Counts of all of the palms in small representa-

tive areas (including established seedlings)

show that the palms of reproductive age con-

stitute less than half of the population. There-

fore the total number of established individuals

of Livistona mariae (the total naturally occur-

ring world population) is estimated to be up-

wards af 3000.

After a goad season, cach mature palm pro-

duces an abundance of seeds with a high

natural rate of germination, Although few of

these seedlings are able to get their roots into

PK, LAT?

the rock fissures before they are washed away

or die of desiccation, there is present in the

populations a wide range of all growth stages.

Seeds of other palms are usually slow to ger-

minute and quickly Jose their viability unless

kept in humid conditions (de Leon 1958), but

seeds of the cabbage palm can germinate within

months and remain viable for two or more

years under dry storage.

Discussion

The cabbage palm is restricted to a small

area and its numbers are small. However, it

has a relatively fast growth rate, high potenual

for reproduction, and a relatively rapid ability

to recover after fire, all factors that indicate

the palm is in no real danger of extinction, pro-

vided the present ecosystem is not substantially

altered. In fact. numbers appear to have in-

creased somewhat in the last 50 years,

The greatest danger to the continued health

of the stand would arise from the lowering of

the water table or general disruption or pollu-

tion of the seepage area, Fortunately the struc-

ture of the sediments probably ensures that this

possibility is remote, The relict nature of this

unique palm warrants close attention to the

general health of the stand and prevention of

any disturbances to the area. The planting of

palms jn other areas of Australia to ensure’

that replacements ate available in case of

natural disaster in recommended.

Conclusion

Palm Valley fs situated on the edge of a

large gas field. ‘The valley is unique botanically,

not only because of the presence of the palms,

but also because of the high number of other

rare plants and the overall diversity of the flora,

A recluse area suich as this, ts by nature fragile,

and therefore uny development in this area

will require peeat care and foresight.

Acknowledgments

f acknowledge the help in interpreting and

dating the photographs lent ta me by my father

Arthur Latz. 1 am indebted to G. Griffin, the

ranger at Palm Valley, for his able assistance

in gathering facts for this report, and to his

employers, the Northern Territory Reserves

Board.

References

Burperince, N. T. (1960)—The phytogeography

4 the Australian region, Aust. J. Bot, 8, 75-

211.

Ciiprenpare, GM. (1959),—Planis of Palm

Valley, Central Australia. Mie. Nat, 75, 192-

199.

Cuippenpae, G. M. (1963) —The relic naqure of

some Central Australian plants. Trans. R. Soe.

S. dust. 86, 31-34,

bE Laon, N. J. (1958).—Viability of Palm Seeds

Principes 2(3), 96-98,

THE RELICT PALM LIVISTONA MARIAE 195

Gites, E, (1875).—“Geographical travels in Cen-

tral Australia, from 1872-74.” (McCarron

Bird and Co,: Melbourne.)

Keast, A, (1959).—Relic animals and plants of

the Macdonnell Ranges. Aust. Mus. Mag.

13(3), 81-86.

Loruian, T. R. N. (1959),—Further notes con-

cerning the Central Australian Cabbage

pi Lice mariae. Principes 3(2), 53-

Tare, R. (1896).—Botany. Jn W. B. Spencer

(Ed.) “Report of the Horn Scientific Expedi-

tion to Central Australia Pt. HI—Geology

and Botany.” (Melville, Mullen and Slade:

Melbourne.)

STIRLING, E. C. (1896).—Anthropology. in W. B.

Spencer (Ed.) “Report of the Horn Scientific

Expedition to Central Australia Pt. IV—An-

thropology.” (Melville, Mullen and Slade:

Melbourne.)

GEOMORPHOLOGICAL EVOLUTION OF PART OF THE EASTERN

MOUNT LOFTY RANGES, SOUTH AUSTRALIA

BY C. 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. 99(4), 197-209,

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.

GEOMORPHOLOGICAL EVOLUTION OF PART OF THE EASTERN MOUNT

LOFTY RANGES, SOUTH AUSTRALIA

by C. R. TwipaLe* and JENNIFER A, BOURNE*

Summary

Twipace, C. R., & Bourse, Jennifer A. (1975)—Geomorphaological cyolution of purt of the

eastern Mount Lofty Ranges, South Australia. Trans, R. Soc. 3. Aust. 99(4), 197-209,

30 November, 1975,

Scattered relicts of a lateritic erosion surface of probable early Mesozoic age occur as

the Whalley Surface high jn 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

dne 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 embuyments eroded in Cambrian racks where major nvers debouch

from the uplands. It carries a veneer of ferruginous grit, Quaternary calcretc, dunes and

alluvia.

The scarps associaled with the Milendella and Palmer faults are primarily of late Meso-

zoic-carly 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 slreams of intensely weathered rocks in the fauit and scarpfoot

zones in Quaternary times.

Introduction

The Mount Lofty Ranges have received

considerable attention from geomearphologists,

but even in studies which ate ostensibly con-

cerned with the uplands as 1a whole it is the

western areas that have been investigated in

detail (see Benson 1911; Fenner 1930, 1931,

1938; Sprigg 1946). There has been passing

reference io particular landforms which occur

in the eastern Mount Lofty Ranges (e.g. Fen-

ner 1931, p, 23); there have been reconnais-

sanee studies of small parts of the area (Saies

19681; Breuer 19687; Forrest 1969°; Frahn

1971+); and specific features or problems such

as the riverine features of the Finnis valley (de

Mooy 1959}, the yranite forms of the Palmer

and Caloote areas (Twidale 1968, p. 95 et seq.;

1971, pp, 5-4), and the Permian glacial

features of the Strathalbyn area (Maud

1972) have been investigated. Although there

has been significant geological mapping of the

region (Kleeman & White 1956, White &

Thatcher 1957; Mills 1965*; Thomson 1969)

there has been no consideration of the evolu-

tion of the ¢astetrn Mount Lofty Ranges as a

whole, no attempt to correlate its development

with that of the western Murray Basin, and ne

investigation of the interplay of tectonism amd

landform development in the area.

* Department of Geography, University of Adelaide, Adelaide, S. Aust. 5000.

VSaies, Tanet (1968).-.“Geomorphology of the Mannum Falls Area”, (Unpubl, B.A. Hons thesis, Univ.

Adelaide.)

“Breuer, Margaret (1968) —‘“The Geomorphology of Harrisons Creck”. (Unpubl. B.A. Hons thesis,

Univ. Adelaide.)

4 Forrest, G. FP. (1969).—"Geomorphological Eyolution of the Bremer Valley”. (Unpubl, B.A. Hons

thesis, Univ. Adelaide. )

'Brahn, D. (1971).—"Geomorphology of the Milendella Area of South Australia’, (Unpubl. B.A. Hons

thesis. Univ. Adelaide.)

* Mills, K. J. (1965).—*"The Structural Petrology of an Area Kast of Springton, South Australia. (Un-

publ, Ph.D. thesis, Univ, Adelaide.)

S.A.

== al

1) 8

a ee |

aIudY nrea

Whalley Surel dun

afobillo

s.-a1emenl sone

Murray _

MunvaySuitiee

plains .

I brome iM tie

= Pate at lO me eben

L [Tae sulfece |

mc omeandsescre NI!

Fig. |. Physiographic regions of part of the eastern

General Description

The area under consideration lies between

the River Marne in the north and Gorge Creck

in the sowth, 1) falls waturally into three physio-

graphic regions: the high plain, the Murray

Plains and the intervening escarpment (Fig.

1}.

The Summit High Plain

* Standing at elevations af 200-300 m above

sea level, the summit high plain or Tungkillo

Surface* fies 120-170 m higher than the nearby

Murray Plains. tt slopes down to the sooth

from some 430 m elevation near the Marne to

330 mi south of Harrisons Creck.

The Surface {s eroded in gneviss, schist. and

granite (Fig, 2) which are esseatially un-

weathered and which give rise to distinctive

landform assemblages. Boulders known locally,

and inadvisedly (Twidale 1971, pp. 14-17), as

‘tors’, bpve developed on the northern parl of

Ife Palmer granite outerop where the rock is

well-jointed hut ant shattered and where split

C. K. TWIDALE & JENNIFER A, BOURNE.

2. MARR Ale

: Mount Lofty Ranges, and (inset) location mip.

The area around Ruetjens Hill and depicted in Fig. Sb is indicated, A-B and C-D are the lines

of seciion shown in Fig. Sa, and X-¥ the section represented in Fig. 7.

rocks, perched rocks or loganstones { Fig, 3a)

together with the many clusters of boulders

form a distinctive landform assemblage. The

foliation of the gneisses has been exploited by

weuthering agents ta produce penitent [Acker-

mann 1962) or tombstone rocks (Fig. 3b), but

in other places, us for instance 6-8 km cast

of Mount Pleasant. the gneiss is more massive

and forms low (up to 10 m) angular blocky

residuals (Fig. 3c) or miniature castle koppics.

Also standing above the tevel of the Tume-

killo Surface ure several scattered Jow (again

up fo 10 tm) mesas. or conical hills which are

vither underlain by luteritic Weathering profiles

carrvipg 4 ferruginous encristatein, of are

capped by alluvial gravels and boulders of pre-

dominantly yuartzilic composition, These

residual hills collectively form) what hus becn

called the Whalley Surface*.

The Tungkillo Surface is deeply incised by

such liiteemittent streams as the River Mame,

and Ssunders, Milendella, Harrisons ar!

GEOMORPHOLOGICAL EVOLUTION OF THE EASTERN MT LOPTY RANGES

VINCENT GULF

smo

Cainozale sediments laterite Paliroer grertite range Palerenic

Fig. 2. Generalised geological map of part of the Mount Lofty Ranges (after S. Aust, Geol. Surv, map

sheet SI/54-9, Adelaide).

Gorge creeks, Their valleys are however sur-

prisingly natrow in proportion to their depths.

and in many places the streams tun through

gorges or defiles,

The Escarpment

The eastern escarpment of the Mount Lofty

Ranges. is, like its western counterpart (Fig.

2). a complex feature. However in broad view

il comprises two north-south trending scarps

which are so gently arcuate that they can be

regarded as linear. The two are offset with

respect to one another by a distance of some 3

km, the northern component extending as far

south as Milendella, the other from the vicinity

of Raetjens Hill and south through Palmer

(Fig. 2).

The scarps are intensely dissected. The

Milendella Scarp 1s. greatly eroded, the zone

of dissection being some 4-6 km wide. How-

ever, dissection behind the Palmer Scarp ex-

tends for only some 2 km. The lutter is also

lower, being 100-145 m high compared to the

Milendella which rises 250 m above the scarp

foot.

Various minor breaks of slope of probable

structural origin can be discerned on the scarp.

Jn some places the upland scarp displays a basal

steepening and scarpfoot depression, as do

some of the ridges within the upland (Pig. 4a).

In some few sites. prominent flats occur at

levels intermediate hetween the Tungkillo Sur-

face above and the Murray Plains below, One,

199

ean Premarin \\ fant

the Millendella Bench, is greatly dissected hy

the present Milendella Creek and its tributaries,

but there are nevertheless broud flats at eleva-

tions between 260 and 280 m, which are inter-

preted as remnants of a once continuous flood

plain. They stand some 80 m higher than the

plains to the east, and descend by way of a

steep slope to the Murray Plains (see Tepko

1:50,000 topographic series (6728-191)

between grid lines 320 and 347. and 440 and

493; see also Pig. 5a and b). The bluff leading

up to the summit surface from the perched

bench is precipitous and arcuate in plan.

Several features suggest that it is an abandoned

meander bluff: its shape, the presence of a

central hill interpreted as a meander core with-

in the horseshoe-shaped valley which forms the

northerly extension of the bench, and the

presence in the valley floor of several metres

of alluvium. It was associated with an in-

trenched meander similar to those found on the

present Milendella Creck but of roughly 5-6

times greater radius. The meander loop and

core developed when the precursor of Milen-

della Creek flowed at a Jevel some 80 m

higher than at present.

A second bench occurs where Harrisons

Creek leaves the uplands to low across the

Murray Plains, where it is known as Reedy

Creek (see Tepko 1:50,000 topographic

series (6728-111) around MR313374; see Fig.

4b). Less extensive than the Milendella Bench,

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, Small scarpfoot depression developed at the base of a gneiss ridge at Raetjens Gap. (C. R.

Twidale. )

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),

the scarpfoot valley (d), and the lower steeper scarp face (e). (C. R. Twidale.)

c. The Whalley Surface (a) and Tungkillo Surface (b) west of Mount Pleasant, Outcrops of the

gneiss country rock are seen in the foreground (c). (C. R. Twidale. )

202 C_ R. TWIDALE & JENNIFER A. BOURNE

= =

A 5 = 5 B

ba % 3 = =

a a

o -

Ty 5 oO 26

G + sf Fy = D

la i rs) z

ado) 2 <3 Ss =

a ol yh # we :

t > Baer Grete => t

= ae =

2 = 5

ut B = Heedy Groek =

c =

r | = 4

— 7

qi ur ba 20

RILOMETFES

Fig, 5. 4. Profiles (A-B and C-D on Fig. 1) across the study area showing the Whalley and Tungkillo

surfaces, the east-facing scarps, and the Milendella and Palmer benches. Drawn from S.A.

Lands Dept. 1:50,000 topographic sheet 6728-111, Tepko,

“

(ye C

‘,

ih contour interval 20 metres

Fig. 5. b. Generalised topographic map of the Milendella Bench showing the old meander loop and the

meander core (spot height 339). Drawn from S.A. Lands Dept. 1:50,000 topographic sheet

6728-11, Tepko.

GEOMORPHOLOGICAL EVOLUTION OF THE EASTERN MT LOFTY RANGES

level, 30 m above the adjacent plains and 130

m below the Tungkillo Surface and in detail

licludes several levels which represent stages

in downcutting. This Palmer Bench (so called

beeduse the site of the original Palmer town-

ship was only a short distance away) carries a

mantle of quartzitic blocks, some of which are

iron-stained. It has been dissected by Reedy

Creek which in this sector displays well-

developed inirenched meanders.

Other. simifar, perched benches have been

noted near the mouth of Pine Valley; south of

Tepko, and particularly west of Strathalbyn

where there ig an e&tensive dissected bench,

but as these are outside the study ares they are

not considered further,

Murray Plaitts

The western Murray Plains comprise a roll

ing Jowland which stands 200-150 m high

tear the scarp foot, to 70-50 m ar the cliff tap

overlooking the River Murray. Ii several sec-

tors there is a Scarpfoot depression up to $0

m deep fronting the Mount Lofty Ranges (Fig.

1).

‘These western Murray Plams are undertnin

by Cainozoic strata 80 m or more thick. In the

west. Quaternary fanglomerates predominate

hut these thin to the east where Miocene

Mating siratz accur in bores, in tributary

valleys, and in the high precipitous bluffs bor-

dering the Murray trench (Fig, 2). Borelog

data indicates that the Miocene strata rest on

an irregular surface cut in acid atid hasic crys-

talline focks. Basic crystalline rocks ate

exposed at Black Hill, where the so-called

‘norte’ has been extensively quarried: similar

recks have heen located in bores near Sedan

and Walkers Flat. Granitic rocks and schists

are exposed extensively in the valley of Reedy

Creek. but also in many other minor outcrops,

How did this landscape evolve? Of what age

and ongin yre the plains? What is the nature

oF the scarp which separates the high from the

low plains?

The lineatity of these scarps and their

association with faults exposed in the gorges of

the River Marne and Saunders Creek or in-

ferred from the displacemens of strata. suggests

that the iopographic forms are fault venerated

and probably fault «carps, ic. due directly and

wholly to. tectonic Uisplacement This is cer-

lainly the view of Mills? who with respect

lo the Milendella Fault, which is of reverse

203

type, argued \hal several stages could be

detected in its development:

1. Initiation of the fuult in the Palacozcic with

the east side upthrown by many hundreds of

metres,.

2. Early Tertiary erosion and peneplanation

with no faulting.

3. Renewal of faulting during the early Tertiary

to Miocene with the east side subsiding by

about 260 m,

4. Miocene marine incursion with strata over-

lapping the eroded faull-scarp.

3, Recurrent movement on the fault since the

Miocene, the east stde being lowered by 60—

90 m,

Several aspects of this chronology can le

challenged, and in particular the date and ex-

tent of faulting are open to question. The proh-

lem involves a consideration of ihe ape and

character of erosion surfaces identified in the

Mount Lofty Ranges and on the sdjacent

Murray Plains.

Age and Nature of the Summit Surface

The summit surface is a complex feature. The

lateritic remnants of the Whalley Surface arc

crucial to the interpretation of the whole up-

land. They are sufficiently scattered to sugges!

that they are relicts of a once-contiguous

weathered surface of low relief, over which

Nowed streams carrying « gravelly and

bouldery quartzose bedload, Remnant deposits

of these boulder beds occur in places. and a

discontinuous mantle of angular quartz frag-

ments forms u veneer on the Jow divides. The

weathering extended only some 10 m beneath

the surface but it is notable that the base of

significant weathering—the ‘weathering front’

of Mabbutt (1961)—is everywhere coincident

with the local level of the Timneakillo Surface.

underlain by intrinsically fresh rock, Thus it

is. teasOnable to suggest that the Tungkillo Sur-

face is an etch plain (Wayland 1934), formed

as a result of the stripping of the lateritic deen

weathering profile and the consequent exposure

of the essentially fresh rock beneath (Figs 4c

and 6), The penitent rocks, small castle koppies

and boulders typical of various areas of the

Tungkillo Surface (Fig, 3) were developed by

differential weathering at the weathering Front

and exposed as a result of the stripping of

weathered material,

The age of the Whalley Surface and of the

Weep weathering is Mesovoic und probably

carly Mesozoic (Triassic), The laterite reni-

nants can be traced to the sovthern Mount

20\4 c

latenle

Fig. 4. Diagrammatic section through the eastern

Mount Lofry Ranges showing the rela-

tionship between the lateritised Whalley

‘Surface underlain by weathered bedrock

und the elch plain, coincident with the

Weathering front, of the Tungkillo Sur

face,

Lofty Ranges where the laterite surface stands

high above: valleys partially fled by Miacene

marine sediments, and whete the fault ungle

depressions resulting from the fault-dislocation

of the summit surface contyin basal marine

sediment of early Tertiary age. and with in-

clusions of laterite blocks (Glaessner 1953;

Glaessner & Wade 1958; Campana 1958).

Thus the summit surface and the laterite

carries must be of curliest Tertiary or late

Mesozoic age However, evidence from Kan-

guroa Island. just across Backstairs Passage

from Fleurieu Peninsula, suggests a much older

age. There the luterite plateau formed on Cam-

brian and Permian strata was purtially dissected

before the extrusion of Middle Jurassic basalts

near and just west of the present site of Kings-

cote. Palaeoclimnatic consideration suggest the

Triassic as the most likely period fur laterite

development (Daily, Twidale & Milnes 1974),

This by short-distance correlation the

Whalley Surface and the associated deep

weathering are considered to be of early Meso-

wc age. The development of the Tungkillo

Surface followed the disruption of the upland,

prohably by faulting. Streams were rejuvenated

und they exploited the contrast in cohesiveness

wnd reststance between fhe regohth and the

fresh rocks beneath, with the result that an

eich surface of low relief, the Tungkillo Sur-

fuce. was initialed, [t is still extending.

Disruption of the Summit Surface

In view of the character of the Milendella

nnd Palmer scarps it is obvious that the reason

for the disruption of the summit surface could

have been a renewal of movement on the two

recognised faults, If this were so, rocks similar

to those of the nearby upland should be found

beneath the Cainozoic strata of the western

Mureay Basin. Granites and giielsses are

exposed widely tn the valley of Reedy Creek,

R TWIDALE & IENNIFER A BOURNE

im minor outcrops throughout the western

Muteay Basin, and they are commonly en-

countered at shallaw depths west of the River

Murray. But there is one feature missing.

numely buried faterite, though weathered

(kaolinised) Cambrian bedrock is recorded in

bores beneath the fresh erystallincs and the

overlying Cainozoic strata. There are also

superficial Ferrugingus encrustations on high

plain remnants and at higher elevations than

the Miocene strata and therefore of later

Cainozoice age, in the Murray Plains, for

example just south of the Mannum Falls.

Further reference 1s made to these oecurrences

below, but no laterite profiles have been found

either i outerep or in bores.

There are a nuntber of possible explanations,

For some reason of reasons al present un-

known, laterite may not have developed on

those rocks of what Was to become the down-

thrawn block; this is rejected as heing

irrationnl, Second, the laterite cowld have been

eroded betore burial by Miocene and fater

sediments. but then some remnants should

surely have been preserved, Third, the laterite

may indeed be preserved beneath the Cainazoice

cover, but has not yet been discovered. This is

unlikely because many bores have been sunk

and though ihere is some reference to

weathered crystallines, there is. none to laterite,

Fourth, the weathering of the Cambrian rocks

could haye taken place beneath the Caingzoic

cover. but it is unlikely that the oxidation rep-

resented by the weathered bedrock could have

developed in such conditions. Fifth, the latertre

could have been buried and iron oxide duricrust

destroyed by groundwater attack in the sub-

surface, Such dissolution of fron oxide by water

charged with organic acids has been shown to

be possible (Bloomfield 1957, Coulson, Davis

& Lewis 1960; Hingston |963),

This Jast explanation seems to be the mast

likely. AIL other indications are that the

Whatley Surface was dislocated by faulting,

with the east side subsiding by a matter of 260

m in the north. Vhe concentration of grownd-

waters in the scarp foot zone (see Ruxtun

1958; Twidale 1962, 1967) would account for

the subsurface elimination of the ferruginous

crust,

Thus jf it is accepted What faulting occurred

ufter the development of the lateritised summit

surface but prior to the Miocene marine trans-

sressiun, a number af geomorphological

changes can be related to this periud:

GFOMORPHOLOGICAL EVOLUTION OF THE EASTERN MT LOFTY RANGES

1. The rejuvenated streams began the stripping

of the laterite regolith and the development

of the Tungkillo Sarface.

2. The rejuvenated streams incised info the wn-

weathered hedrock creating quite deep, mar

row, and in places meandering gorges,

3.As they emerged from the uplands, the

streams eroded quite broad embayments in

the crystalline basemen! rocks of the uplands

contiguous with the then surface of the

Murray Plains, The Milendella Bench stands

considerably higher than either the Palmer

Beneh or bench remnanm at the mouths of

the Saunders and Mame gurges, This may

reflect its development at a stage before

faulting hat! ceased, and its grading to the

Miocene shore. o it may indicate focal fate

Cainuzoic faulting,

When did faulting begin? There is na direct

evidence, but the stratigraphy of the Murray

Basin (O'Driscoll 1960; Ludbrook 1949) sug-

gests that part of jhe region became a restricted

marine embayment during the early Creia-

ceous. The next proven maring transgression

took place during the Late Eocene und marine

influence continued through to the Middle Mio-

cene, However, though strata of Late Crets-

ceous age have not been located. they may

huve been deposited und subsequently been

removed by ¢rasion (Ludbrook 1969, p. 159).

The fault dislocation wis probably gradual and

possibly occupied the whole of this late Meso-

z0ic-middle Tertiary period

Age of the Plains Surface

The broadly rolling plains surface of the

western Murray Basin is being dissected by the

few through-flowing Streams such as the Marne,

Milendella and the Hurrisons-Reedy systems,

which emerge from the Mount Lofty Ranges

and persist to reach the Murray, Uhe River

Murray treach is a Quaternary feature, though

there Was an earlier shallow estuary coincident

with the present Murray valley as far north as

Morgan. In this long narrow and shallow est.

ury. Marine sediments, the Norwest Bend For-

mation, of Late Pliycene age were depustted

(Ludbrock 1958), Remnants can he seen

perched high on the modern cliff-tops in many

places. occupying a shallow valley cut in the

Miocene strata. As the trench now occupied by

the river has been incised below these Pliocenc

strata, it musi be younger and there is much

general argument as Well as specific evidence

{van der Borch 1968) fo suggest that if is of

205

Pleistocene age, being related fo low glacial

Stands of the sea,

However, in Pliocene times the regional base

level on the western Murray Plains stood 40-

70m higher than it does now, and it is to this

higher basclevel that the streams responsible

for the erosion of the plains were graded.

These streams transported fo the then Murray

estuary the deiritus which in part comprises the

Norwest Bend Formation, Thus the Murray

Surface Js in the study area nf Pliocene age.

There is some corroboration from the thin

ferruginous duricrusts [ound on some parts of

the surface, as for instance just south and

South-west of Mannum Falls, for surfaces of

comparable age on northern Yorke Peninsula

(Horwitz & Daily 1958) and Eyre Peninsula

(Twidale, Bourne & Smith 1976) also carry x

ferruginous encrustatian, as indecd do post-

Miocene vallev Aoors in the southern Mount

Lofty Ranges (Horwitz, 1960), and the

Karoonda Surface of the central Murray Plains

(Firman 1973, p. 23).

‘The occurrence of iron staining, as well as

their elevation, suggest that the several perched

benches found at the mouths of major upland

gorges were also formed at this time, even

though they are now separated from the plains

by scarpfoot valleys up to 50 my deep, This is

corroborated hy profiles relating the benches to

the main plains surface (Fig. Sa),

Although little is known of the geometric

relationship between river discharge, channoe!

width and the radius of intrenched meanders

(cf. Bates 1939 and Dury 1954, 1964; with

respect ta flood plain. meanders, also Geyl

1968) the abandoned meander loop preserved

on the Milendella Bench sugpeste rivers of

much greater (? occasional! discharge than

those active at present,

Thus, during the Pliocene, the scarp fout

stond 80 m higher than present at the mouth

of Milendellz Creek and 50 m above present

baselevel where Harrisons Creek dehouches or

lo the pin. Is the post Pliocene scarp tectonic

or efosional, or are there elements of both

origins present?

The Question of Post Late Cainozoic Faulting

The Iower part of the Milendella, and hy

implication the Palmer, fault scarps are seen

by Mills (1965, pp. 436-442) as due to post

Miocene faulting. He discovered several out

fiers of Miocene sediments perched on the

scarp north of Milendella, correlated them with

Mincene strata found in bores in the Cambrat

473 metres

METAMORPHIC

perched

Miocere limestone

162 s

Quaternary alluvium BE

SAUNDERS CREEK

upper Miocene sand

Miocene limestone

Eocene lignitic sands

Sea Levei

Okm i

. TWLDALF. & TENNIFER A, BOURNE

Fig. 7. Section through the Milendella Scarp near Saunders Creek, (After Mills: 1965).

znd Milendella areas, and took the elevational

difference between the two occurrences (i.e.

60 and 90 m) as a measure of post Miocene

dislocation. on the faults (Fig. 7). He also noted

that boulders in the Pleistocene fanglomcrates

abutting the fault plane display onentation sug-

gestive of recent movement along the plane.

This suggestion of recent and continuing dis-

location is borne out by the seismic recotd

(Sutton & White 1968) which shows that the

eastern border of the Mount Lofty Ranges 1s

subject to earth tremors, The distinctness of

the escarpments bath here and to the south

suggests geological youthfulness (see e.g. leune

& Chittleborough 1974), The higher level of

the Milendella Bench (see earlier) can alsa be

interpreted us indicating local late Cainozoie

dislocation.

On the other hand, the scarps have been

deeply and intricately dissected, the Milendella

scarp in particular being greatly cut about yet

retaining its overall lincar morphology. More

significantly, the evidence cited by Mills is

equivocal and is equally well accounted for in

other terms,

The Miocene of the western Murray Basin

comprises two major formations: the strali-

graphically lower or older Mannum Formation

and the higher and younger Morgan Formation

(Ludbrook 1969}. The two are very difficult to

distinguish i the field and can only be differen-

tiated palacontologically. The faunas of the

perched outliers are unfortunately not diagnas-

tic. Mr. J. M, Lindsay reported (pers. comm.)

in 1975: “A sample of recrystallised quartzose

fine-grained limestone, from a perched. outlier

of mid-Tertiary rocks 3.5 km north of Sander-

ston, contained small benthonic foraminifera

but no diagnostic forms and no lithological

features which would distinguish between Mor-

gun Limestone and Mannum Formation.” As

the stratigraphic position of the Miocene cn-

countered in bores has also not been deter-

mined to this date, it is not possible to identify

either the outliers: or the strata. intersected in

bores with certainty and precision. Thus corte-

jation is unwarranted. It cai he argued that the

topographically higher outliers are Morgan

Formation left behind as remnants of circum-

denudation after the removal by solution and

fluvial action of the bulk of the Formation dur-

ing the Pliocene and Quaternary, that the Mio-

cene encountered in the bores is Mannum Vor-

mation in situ, and thus that na post-Miocene

dislocation of any significance is implied by the

evidence (Fig, 8). This crosional hypathesis

receives support from the character of the

Milendella and Palmer scarps. If the scarps

were wholly of tectonic origin, then the searps

ought everywhere to be of similar elevation or

at least to vary systematically and gradually

according to differences in the throw of the

fault dislocations, But the scarps in fact

vary in height according to their location with

respect to scarpfoot streams. Where there are

streams either heading back to the scarp Foot

GEOMORPHOLOGICAL EVOLUTION OF THE EASTERN MT LOFTY RANGES 207

Se ES Pe a Wicceng

a taulting

implied

Miocene

— Morgan

bno post Miocene \

faulting Implied Machu

Fig. 8 Two possible modes of evolution fox the

eastern scarp of the Mount Lofty Ranges.

ur debouching from the hills on. to the plains,

the scarp is higher, there is a faceted scarp with

the lower unit steeper than the higher, and

there is a distinct scarpfoot valley running

parallel to the escarpment (sce Figs, 1, 4b and

Sa), frequently with intensely weathered rock

exposed in the yalley floors. In between these

sectors ate others where there are no scarpfoot

streams, Where the escarpment is of Jesser

amplitude and where the hill-plain junction is

higher and coincident in elevation with the

break of slope on the higher scarps nearby,

This suggests that the upper part of the scarp

is of fault or tectonic origin, though of ¢on-

siderable antiquity (late Mesozoic or early Ter-

tiary), but that the lower part, where present,

is due to erosion by streams eroding the

weuthered strata of the scarpfoot zohe (Twi-

dale 1967); ic. that the lower scarp is of fault-

lime type and that no significant recent disloca-

Cambrian

tion is implied, This is not to suggest that there

has been no recent movement on the fault: on

the contrary, the seismic evidence and the dis-

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

principally crosional and that tectonic disloca-

tion has been a relatively minor factor in its

evolution,

There remains the problem of the contrasted

state of dissection displayed by the Milendella

and Palmer scarps, It is not a matter of can-

trast in rock type or volume of run-off, for the

lithological units of the eastern Ranges run

north-south and are essentially similar behind

both scarps. Ii cannot be a question of climatic

differences. It may be due to the Milendella and

Marne draining a higher scatp, and eroding

regressively more quickly to capture the head-

waters of the Torrens drainage. More likely

however, the dissection of the southern part of

the study area has been retarded by the base-

level control exerted on the Harrisons-Recdy

creek system by the large miass of crystallines

exposed west of Culoote. The lower elevation

of the Palmer as compared to the Milendella

scarp may be a contributary factor,

Conclusion

The development of the eastern Mount Lefty

Ranges and the parts of the Murray Plains is

summarised helow.

Deposition in Adelaide Geosyncline

Early Palaeozoic

Late Palaeozoic

Early Mesozoic

Late Mesazoic—

Middle Tertiary

Pliocene

Qualernary

Orogenesis, granite emplacement and metamor-

phism

Permian glaciaiion (not evidenced if study

area)

Planation and deep weathering. Marine deposi-

tion in lower part of Murray Basin in early

Cretaceous (Thornton 1974)

Faulting, eastern block down. Marine sedimen-

tation Eocene-Miocene, Stream rejuyenution

and stripping of regolith

Essential withdrawal of sea, only estuarine sedi-

mentation. Planation of Miocene sediments and

of embayments in crystalling rocks on face of

scarp

Lowering of seq level. Murray Trench fermed,

tributaries rejuvenated, valleys and scarpfoot

Zone excavated

Whalley Surface

Tungkillo Surface:

Searp formed. Gorges

developed

Perched benches and

Murray Surface

Scarp foot valleys,

abandonment of scarp

benches

20K

Acknowledgments

‘The writers wish to thank Dr Brian Daily,

Geology Department, University of Adelaide,

for a critical reading of the paper, und Dr

BOURNE

Kingsley Mills, Department of Geology and

Geophysics, University of Sydney. for helptul

discussion and continutng imerest in the eVolu-

tion of the area under investigation,

References

AckKERMANN, E. (1962),—Bussersteine-Zeugen vor-

zejtlicher Grundwaaserschwankungen, Z. Geo-

mearph. (N.S) 6, b48- 182.

Bayes, R, E, (1939},—Geomorphic history of the

Kickapoo Region, Wisconsin. Ball. seal. Soe,

Am, 50, 819-879- ;

Henson, W. N. (I911)—Note descriptive of a

sleveogiam of the Mt Lofty Ranges, South

Australia. Trans, R. Soe. 8. Aust, 35, 18-114,

BLouMETELD, C. (1957).—The possible signifi-

cance of polyphenols in soil formation. J. Sei.

Fd, Agric, 8, 389.

Campana, B. (1958) —The Mt Lotty-Olary regan

and Kangaroo Island. 7a M. F. Glaessner &

L. W. Parkin (eds) “The Geology of South

Australia”, pp, 3-27. (Melb, U-P.)

CouLrgon, C, RB, Davis, R. Lb, & Lewis, D, A.

(1960).—Polyphenols in plant, humus and

soul, L. Polyphenols of leaves, liter, and super-

ficial humus trom mull and mor sites. 1 Re-

duction and transport by polyphenols of iron

in model soll eolumns. J. Soil. Sef, 14, 20.

Damwy, B, Twitaue, C. RK. & MIcNes. A R.

(1974).—The age of the loteritized summit

surface on Kangaroo [sland sand adjacent

areas of Souch Australia, J, Geol. Soc. Aus.

21, 387-392.

Dery, G. H. (1934),—Contribution to a general

theory of meandering valleys Am, f. Sev.

252. 193-224, :

Dury, G. H. (1964)—-Principles of underfil

giréams. Prof. Pap. U.S. geal. Surv. 452-4.

1-67,

Fenner, C, (1930).—Phe mujer siructural and

physiographic features of South Australia.

Trans. R, Sow. 8. Aust. 54, 1-36.

Fenner, C, (1931),—"South Australia: A Geo-

prophical Study.” (Whitcombe vad Tombs:

Melbourne.)

graphy of Anstey Hill, South Australia. Trans.

R. Soe. 8. Aust. 63, 79-87.

Pianmtan, J. B. (1973),--Regional stratigraphy of

surficial depositg in the Murray Basm and

Gambier Embayment. Rept. Invest. geal.

Surv. S, Aust. 39, ’

Gevn, W. F, (1968) —Tidal stream action and

seu level chonge as one cause of valley meun-

ders and underfit streums, Aust, Geag, Srad.

6, 24-42, ‘ .

Guaessnér, M, F. (1953) —Some problems of

Tertiary geology in southern Australia JR,

Sar. NUS.A. 87( 2), 31-45,

Geaessyer, M. F., & Want. Maury (1958)—The

St. Vincent Basin. /x M. F. Glocasner & 1. W.

Parkin {eds}. “The Geology of South Aus-

wali”, pp. 115-126, (Melb. U.P.)

Hinustun, F. J. (1963) —Activity of polypheno-

lic constituents of leaves of Fueulyplas and

other species i complexing and dissolving

iran oxide. dust. J, Soil. Rew. O11), 63-73

Horwitz. RoC. (1960).—Géologie de la région

de Mi Compuss (feuille Milang), Australie

meéridionsle, Eclog. Géol, Helv §3, 211-283,

Horwity, R. Cy & Daiwy, B. (958i. Yorke

Peninsula, /n M. F, Glaessmer & L. W, Pur-

kin ¢eds). “The Geology of ‘South Australia”.

pp, 46-60. (Melb. U.P.)

JEUNE, R, F., & CHITTLEROROUGH, D. F. (1974), -

City of Monarto. Preliminary Geotechnical

Investigations, 3, stuse, Pepe Mines Rept, Bk

74/105,

Kinuman, A, W, & Wrrre, A. J. R. (1956).—

The structural petrology of portion of the

eastern Mount Lofty Ranges. J, Geral, Soc,

Aut. 3, 17-31.

Luperook, N. AH. ¢19S88)—The Murray Basin in

South Australia. fa M. & Glaessner & 1. W,

Parkin {eds}. “The Geology of South Ats-

tralia", pp. 102-114. (Melb. U.P.)

Liuipomook. N. H. 11969)—Tertiary Petiod. ?n

L. W, Parkin (ed.). “Handbook of South

Australian Geology”, pp. 172-203. (8, Aust.

Geol. Surv.: Adelaide. )

Mansutt, J. A. (1961).—‘“Busal surface” or

‘weathering front”. Proc. Geol. tw. Lendan.

72, 357-358,

Maun, R. R. (1972).—Geology. geomorphology,

und soils of central County Hindmarsh (Mt

Compaxs-Milang), South Australia. C.S.LA8.0.

Soils Publ. 29,

ne Mooy, C. J. (1959) —Nates on (he seomor-

phic history of the urea surrounding lukes

Alexatidrvina and Albert, Sauith Australia,

Trous, R. Soe. §, Aust. 82, 99-118.

O'Driscon., E. P. (1960) —The hydrology of the

Murray Basin Province in South Australia,

Bull. veal. Surv. S. Aust, 35.

Ruxrox, B. P. (1958)—Weathering ood suobsur-

face erosion in granite at the piedmont ingle,

Halos, Sudan. Geol. Mag. 45, 353-377.

Spauc. RC 1 1946),—Reconnuissance geological

survey of porpon of the wesiern escarpment

of the Mount Lofty Ranges, trans, RK. Sane,

A. Apest. 70, 319-347.

Surton, D. F, & White, Ry, E. | 1968).—The

seismicily of South Avstralin 2 Geal. Soe.

Awst!, 18, 25-32_

THomsor, Bo f (1969),—Adrlaide $A, Geologi-

cul Atlas Series. Scale |}:25D,000 Sheet ST/

54-9, zones 5 & 6, (S. Aust, geo. Surv; Ade-

laide. )

Tirornton, R. C, N, (1974).——-Hydrocarbon

potential ol westera Murray Busi and infra

basins, &, Aust, Geal, Surv. Rept, Invest. al.

Twipace. C. R, (1962)—Steepened margins of

inselberes from northwestern Eyre Peninsulu

South Australia, 2, Geontorpl, ONS.) 6, Sl-

69.

angle as evidenced in South Austral 4.

freo!, 75, 393-411.

GEOMORPHOLOGICAL EVOLUTION OF THE EASTERN MT LOFTY RANGES

TWIDALE, C. R. (1968).—Geomorphology with

special reference to Australia. (Nelson: Mel-

bourne.)

TwIipsLe, C. R. (1971).—Structural Landforms.

(A.N.U. Press: Canberra.)

TwmaLe, C. R., Bourne, J. A., & Smiru, D. M.

(1976).—Age and origin of palaeosurfaces on

Eyre Peninsula and the southern Gawler

Ranges. Z. Geomorph. (in press).

209

VON DER Borcu, C. C. (1968).—Southern Aus-

tralian submarine canyons: their distribution

and ages. Mar. Geol. 6, 267-279.

Way anpb, E. J, (1934).—Peneplains and some

erosional platforms. Geol. Surv. Uganda, Ann.

Rept. Bull., 77-79,

Wuits, A. J. R., & THaTcuer, D. (1957).—Man-

num Geological Atlas 1 mile series, No. 811,

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 WIRRAPOWIE LIMESTONE — TWO

NEW LOWER CAMBRIAN FORMATIONS, ELINDERS RANGES,

SOUTH AUSTRALIA

by P. G. Hascerr*

Summary

Hoasteri, PL G. (1975) —The Woodendinna Dolomite and Wirrapowie Limestone—two new

Lower Cambrian formations, Flinders Ranges, Sonth Australia, Trans, R. Soe. 8. Aiast

99(4), 211-219. 30 November, 1975,

Two new formations, the Woodendinna Dolomite and the Wirrapowie Limestune, are

proposed for two distinctive units of the Lower Cambriun carbonate sequences of the Flinders

Ranges, which have not, hitherto, been differentiated from the presently defined formations.

Roth formations aré widespread in their occurrence, but are best developed in the eastern

Fiinders Ranges. north of the line of the Blinman, Wirrealpa and Frome Diapirs. The Wooden-

dinna Dolomile and Wirrapowie Limestone record a significant period of supratidal and shallow,

sheltered, intertidal deposition respectively within the Cambrian of the Flinders Ranges.

Introduction

Formations. within the Hawker Group (Dai-

garno 1964) of the Lower Cambrian of the

Adelaide Geosynelite characteristically show

frequent changes in both thickness and factes,

This results in complex intertonguing of litho-

logical units. The Ten Mile Creek type section

{Daily 1956), measured within a syndeposi-

tional grahen structure northeast of the Ora-

parinna Diapir, represents onc of the thickest

and most comp{ete sequences of the Hawker

Group in the Flinders Ranges. Mapping by the

S.A. Department of Mines (Dalgarno &

Johnson 1966; Coats er al. 1973) and by B.

Daily und « number of his students (Gehling’;

HBaslett*; Hatcher'; Mount®; Hull’; Daily

1972a) has resulted in the recognition of

several significant lithological sequences not

present within the Ten Mile Creek type section.

The thick basal Cambrian carbonates in par-

ticular exhibit complex and varied facies inter-

relations and it has become obvious that within

the presently established broad divisions, viz.

the Wilkawillina Limestone, the Parafa Lime-

stone and the Ajax Limestone (‘Table 1), several

distinctive lithofacies exist. Not only do the

present formations contain separate and indi-

vidually mappable lithologies of significantly

different depositional origin but in the past the

formation names have in some instances been

loosely applied (Dalgarno 1964, pp, 134-135).

TABLE 1

Existing formation names and their sources for the Lower

Cambrian carbonate sequences of the Flinders Ranges

CENTRAL AND

WESTERN EASTERN

LAKE TORRENS FPLINDERS FLINDERS

AREA RANGES RANGES

PARARA

LIMESTONE

ANDAMOOKA AJAX (Daily 1956)

LIMESTONE LIMESTONE

Vohns 1968) (Daily 1956) WILKAWILLINA

LIMESTONE

(Daily 1956)

Whilst it is extremely impractical and un-

desirable to introduce an excessive amount of

formal stratigraphic terminology for the Lower

Cambrian carbonate rocks, it is necessary that

* Department of Geology and Mineralogy, University of Adelaide, Adelaide, 5. Aust, 5000, — .

' Gehling, J, G. (1971).—The Geology of the Reaphook Hill Area. Honoyrs Thesis, Univ. Adelaide

(unpublished ).

* Haslett, PG. (1969) —The Cambrian Geology North of the Wirrealpa Diapir, Flinders Runyes, South

Australia. Honours Thesis, Univ. Adelaide (unpublished), 2

3 Mount, T. J, (1970}.—Geology of the Mt Chambers Gorge Region. Honours Thesis, Univ, Adelaide

(unpublished).

& Hatcher, M. I. (19706).—The Geology of the Mt Chambers Mine Region, Northern Flinders Ranges.

S.A. Honours Thesis, Univ. Adelaide (unpublished). .

* Hull, K. G, (1973).—The Lower Cumbrian, Puttapa Syncline, Flindery Ranges, S.A, Honours Thesis,

Univ. Adelaide (unpublished).

212

sufficient nomenclauite be available to map and

flescribe, on aw fegional scale, the major rock

relationships present within the depositional

basin, Careful recognition of mappable units

al uniform) or repetitive lithubogieal nature

should facilitate rather than chcumber further

work, Such accurate lithological subdivision,

coupled with thorough — palacontological

evaluation, will result in a more lucid inter-

pretation of the depositional history within the

basin. It is with this intention, that the new

formation names Woodendinna Dolomite and

Wirrapowie Limestone are proposed.

Present Nomenclature of the Lower Cambrian

Carbonate Sequence in the Flinders Ranges

Wilkawillina Limestone

The Wilkawillina Limestone, in its Ten Mile

Creek type section, disconformably overlies the

Bonney Sandstone Member of the Pound

Quartzite, and is extensively dolomitized at the

hase. In general the unit is massive, pure and

pale cream to pink coloured. I consists pre-

dominantly of ooid and skeletal grainstones,

which may be patchily silicified and recrystal-

lized, At abundant tauna is characteristic of

the wpper parts, some beds consisting almost

entirely of broken skeletal remains of archeo-

cyathids, and associated brachiopods. hyo-

lithids. and trilubites. Algal remains are

common throughout and although some

cryplalgalaminates and oncolite horizons are

present at the very base, stromatolites ave vir-

tually abseni, Subaerial desiccation features are

not represented in the type section, Daily

(1956, 1972a) considers the Wilkawillina Lime-

stone fo have heen deposited on a stable shelf

under shallow marine conditions.

Parara Limestone

Flaggy to cubbly dark to black impure lime-

stones with interbedded dark calcarcous shules

{urna the Lawer Meniber of the Parara. Lime-

stone in tlhe Wilkawillina grahen, The Upper

Member consists of Nagey and thin bedded

dark prey to black aphanitic limestones and

thin carearcous shales. Although scantily

fessiliferous throughout, the Parara t,imestone

contains irilobites, hyolithids, brachiopods,

sponge spicules, cunchostracans and rare

archeacvathids (Daily 1956). Stromatolites,

flatpebble conglomerates and desiccation

features are all completely absent,

Ajax Limestone

Daily (1956) propused that the term Ajax

Limestone should be used for all carbonates

P. G. HASLETT

above the Pound Quartzite and beneath the red

clasties of the Billy Creek Formation in the

Mt Scott Runge areca. This comparatively thin

carbonate sequence incorporates a number of

different. Hthafngical types. Daily (1972b) has

subsequently distinguished # lower und Upper

Member, and in view of the thickness of this

unil, any further subdivision of the Ajux Lime-

stone may prove impractical on ua regional

stale, It is Very important to realize, however,

that a large number of lithotypes present in the

Ajax Limestone ure present within the Cam-

brian carbonate formations mapped elsewhere

in the Flinders Ranges: (Daily, pers. comm)

and that the change in stratigraphic ter-

minology in the Mt Scott Range area is in large

part historical rather than geological.

Proposed New Formutions

A significant proportion of the basal Cam-

hrian ¢arbonate sequences, particularly those in

the north-east Flinders Ranges, are not ceadily

wssignable to any of the above stratigraphic

units, Although mapped on the COPLEY

1:250,000 and parts of the PARACHII.NA

1:250,000 geological maps as Wilkawillina

Limestone. the carbonates show only seant

similanty to Wilkawillina Limestone as defined

in the type section. In part, these carhonates

show greater similarity with the Parara Lime-

stone of the type section, but crucial differences

in lithology and depositional environment are

upparent on careLul examination. Tt 3s for these

distinctive and widespread basal Cambriun

units that the new names Wewadendinna Doio-

mite and Wirrapowie Limestone are proposed,

Type Section

The type section is located’ in: Wirrapowic

Creek and one of its tributaries, about |4 km

due east of Paint Well Station homesteat! in the

Central Flinders Ranges (Fig. 1). Unfortunately

(he section presents some difficulties with

respect ts access, which involves 15 km of

travel by four-wheel drive vehicle from a point

on the Point Well-Natrina road to Wooden

dinna Bore, and a further 2 km on foot along

4 creek to the section. 'This disadvantage ts fur

outweighed by the completeness: of the

sequence and the good exposure of both the

ubper and lower boundaries of both forma-

tions,

The combined Wondendinna Dolomite and

Wirrapowie Limestone in the type section can-

sist of 392 m of carbonate rocks which over-

lig, with apparent’ conformity, Dipleacruterion

LOWER CAMBRIAN FORMATIONS, FLINDERS RANGES 213

WOODENDINNA DOLOMITE

WIRRAPOWIE LIMESTONE

TYPE SECTION LOCATION

Ta Natrina

COPLEY 1: 250,000 __

PARACHILNA 1: 250,000

PARARA LIMESTONE

WIRRAPOWIE LIMESTONE

WOODENDINNA DOLOMITE

ace “| PARACHILNA FORMATION

POUND QUART ZITE

Wirrealpa Spring

l 138° 55° All tracks shown are passable co four-wheel drive vehicles only

fr ve at T T

SOUTH AUSTRALIA

CLD WIRRE ALIAS

SPRING 4}

BLINMAN

sd (eSDELA DE

WIRREALPA

138° 55’

Fig. 1. Location of type section.

214 P. G. HASLETT

TYPE SECTION —= wWoobdENDINNA DOLOMITE AND WIRRAPOWIE LIMESTONE

Very recwiarly uedued, strongly nutcropping fine graincd

PARARA Ss Jimestanes with Tignler coloured sqaley interbeds

Sparsaly fosstliferous and lacking desiacation features

LIMESTONE and tlat-gebble conglomerates.

» grained jrey ta black sity livestones and

reve silggtones, with very regular ory

dig fark ffnely lamaeted algal qtrunialol the

beds (5) fd lansicular tla -pebbie conylonendtes are

comion. Minor cross-bedded vuis, uraidstone beds (2)

WIRRAPOWIE aruien Liraughaul tub niecatbohate detritus is

vecudlly absent *1 che type section. The uppermost

LIMESTONE .

inde y oF bhis ues Tg samewhet gradational Ince che

BARBIA LIMESTONE und oo pluced at Lhe tapnast algal

stromatolite bed,

2oOm

velicw-weathoring, Flaqqy vo tanstve dolowsas

ans tolornitic limestores, very ciniliene Glad, Cros

bedded quart sandstones fy)y inkevlormational

conglomerates and day?ceulron iu packs. Nluncanyt

stromacolite beds (s). O&litic Timestones (9)

éckyr seee tha top

1nGm Massive outenuping dull grey dalamisiaed | nwsbenes.

= Usually crystalline, primary capositicnal textires bai

WOODENDINNA difficult to detenmre my the field. driginal limestones

DOLOMITE were yredoninantly grasy-ledded aod ara tastoies

Oe ae Luntatning scattered quartz sard qreins, Nabstv=

lalere Ty Tinka stromatelites are covmin.

Maseive LO Pladgy yes Tow-weatherrng dolomites and dle laedt te:

Tanestn as, = Tio bese at hhe WIRRAPCWTE GIMCSTONE 18

apadeently conformable with Lhe undertying PARACHTENA

POWMALIUN aid a6 places at the reladively abrupt apnedrarce

dP carcunete sediineake. The catbenvtes are comuundy

co phir and niarddbie, Flat-pebble conq loners les,

Wwe-srachs dnd slretinlo' thes (5) are also sresent,

Trin, cress-teddew quarts sandstones (4) are interbedded

witi delen (bes afd yolitic jimestanas Tn upper parte,

pa Base is Tikuologieally identical wich umlurty iia POUND

PARAGHILNA = ; MIAKIZITE, but contains abundant Gplacraterion burrons,

' Dipl serater Le

= The ume L becanes red and inéréasingly clay-rich toward the

FORMATION aa af Lt, Ugleeretur an burvsws bevng present theguqhour

Toy TONS*sts een-qrey, poorly cuteropsing shales

Fig. 2. Type section.

LOWER CAMBRIAN FORMATIONS, FLINDERS RANGES

215

ROCK RELATION DIAGRAM

SOUTH

Wilkawillina

Graben €hr

Woodendinna Dolomite and

(Not to scale )

Wirrealpa

Wirrapowie Limestone

NORTH

Arrowie

WEST

Mt Scott

Range Angepena

WOODENDINNA = DOLOMITE

EAST CAMBRIAN

€he Nepabunna _ Siltstone

Arrowie ef

2 €hd Midwerta Shale

8 €hw Wilkawillina Limestone

F Ehj Ajax Limestone

€hp Parachilna Formation

€u Uratanna Formation

PROTEROZOIC

Pwp Pound Quartzite

Disconformity ~———

PGH 75

Fig. 3. Rock relation diagram.

sandstones and shales of the thinly developed

Parachilna Formation. The entire sequence

dips 25° to the SW, on the western limb of a

large, slightly southward plunging anticline. In

the type section, the Woodendinna Dolomite

conformably underlies the characteristically

flaggy grey limestones of the Wirrapowie Lime-

stone, the formations being 176 m and 216 m

thick respectively (Fig. 2).

Lithological Characteristics

Woodendina Dolomite

The Woodendinna Dolomite in the type sec-

tion is apparently conformable upon the Para-

chilna Formation, and its base is selected at the

abrupt appearance of carbonate rocks above

the shales and Diplocraterion sandstones of the

Parachilna Formation. The basal carbonates

are partly or completely dolomitized oolitic and

pisolitic limestones, interbedded with very

minor green shales. Massive to flaggy yellow

weathering dolomites with pronounced desicca-

tion features are abundant just above the base,

in association with low, broadly domed stro-

matolites (Fig. 4) and, beginning about 36 m

from the base, thin cross-bedded, pure quartz

sandstones. Intraformational conglomerates,

often with dolomite clasts in a quartz sandstone

or ooid grainstone matrix, are also common

(Fig. 5).

Between 69 m and 119 m from the base of

the Woodendinna Dolomite in the type section,

all carbonates have been extensively dolo-

mitized. This results in the development of a

massive dull grey unit which appears, super-

ficially at least, to be lithologically homogen-

eous, but which has pre-diagenetic lithologies

comparable to those directly below and above,

but with a greater number of ooid grainstone

beds. Some of the grainstones are rich in dis-

persed quartz sand. Thick, dolomitized algal

stromatolite beds and intraformational con-

glomerates are also common.

Above this dolomitized zone to the top of

the Woodendinna Dolomite the sequence con-

sists of interbedded flaggy yellow dolomites,

ooid grainstones, stromatolites, flat pebble con-

glomerates and quartz sandstones, all with

beautifully preserved primary sedimentary

structures. Desiccation mudcracks, some up to

8 cm deep, are common in the dolomitic mud-

216 P. G. HASLETT

Figs 4-11.

LOWER CAMBRIAN FORMATIONS, FLINDERS RANGES 217

stones (Fig, 6), Ripple marks aud chevran-type

cross bedding are present in the ooid grain-

stones and quartz sandstones (Figs 8 and 9), At

135 m from the base, a thick cross-bedded

quariz sandstone with abundani creamy yellow

dolomitic niudstone clasts is developed. Toward

the top of the Woodendinna Dolomite, ooid

xrainstone beds commonly show a primary

tubbly or cobbled upper surtace, réminiscent of

thar found in modern environments on arens

of subaerially exposed, partially lithified car-

bonate saids (Fig, 7).

Wirrepowie Limestone

The hase of the Wirrapowie Lintestone coin-

vides with the relatively abrupt disappearance

of dolomite in the sequence. Flagev fine-

grained, dark grey, limestone and green grey

calcareous siltstones dominate the sequence.

esiccation features are less abundant hut stro-

matolites, intraformational conglomerates and

minor cross-bedded ooid grainstones are

present throughout (Fig. 11). Quartz sandstones

are virtually absent, As in the Woodendinna

Dolomite, stromatolites are finely laminated,

low, laterally linked domes which, on bedding

plane exposures, show little if any preferred

orientation (Fig. 10),

The Wirrapowie Limestone in the type sec-

tion is overlain by Parara-type limestones, The

topmost limit of the Wirrapowle Limestone is

selected at the top of the uppermost algal

siromatolite bed. Although a superficial simi-

Jarity with the Wirrapowie Limestone remains

above this limit, the sequence lacks ooid grain-

stones, flat-pebble conglomerates and evidence

of subagrial desiccation, [he dark limestones

are far more slrongly outcropping, purer and

more crystalline above the contact. They also

tend to be sparsely fossiliferows, with archeo-

cyathids, hyolithids and various brachiopods

present. This is in contrast to the underlying

Wirrapowie Limestone and Woodendinna

Dolomite which are notably depleted in fossils.

Apart from algal stromatolites, the only evi-

dence of organic activity found to date are very

minor irregular burrows in some dolomitic

mudstones of the Woodendinna Dolomite. and

minor trails and apparent bioturbation of bel.

(ling which may be found in impure limestones

uf ihe Wirrapowte Limestone.

Distribution

The Woodendinna Dolomite and the Wirra-

powie Limestone form the basal Cambrian cat-

donate succession over much of the Flinders

Ranges. They are best developed in the eastam

part of the ranges, north of the line of outerop

of the Blinman, Wirrealpa and Chambers dis-

Pirs (sec Fig. 3). In this region their combined

thickness ranges from 300 m to 500 m and thsy

appear to thin gradually to the west, where

comparable Jithologics are thinly developed in

the Ajax Limestone at Me Scott (Daily, pers,

comm.) and the Andamooka Limestone in the

vicinity of Yarrawurta Cliff (Wopfner 1969;

Daily 19724). To the south, along the above-

mentioned line of diaptrs. the Wirrapowie

Limestone thins very abrupily and is overlain

by, and interfingers with, Wilkawillina Lime-

stone. Syn- and post-depositional tectonism

along this line of diapirs. and the consequent

development of complex local facies variations,

has complicated the zegional facies relation-

ships along this line. Lateral interfingering of

the Wilkawillina Limestone and the Wirrapowie

Limestone is strikingly demonstrated. however,

in outcrops just north of the Wirrealpa Diapir

(Haslett, in press), Abrupt lateral change south-

ward from Wirrapowie Limestone may also be

seen in the Vicinity of Fregunda Creek. In addi-

lion, Hatcher? indicates a similar southward

thinning of a wedge of Wirrapowle-type lime-

Stones within the northern part of an area of

thickly developed Wilkawillina limestone near

Mt Frome.

In the north-eastern Flinders Ranges, the

Wirrapowie Limestone is in all cases overlain

by Parara-type fimestones. To the best of the

author's knowjedge, Wilkawillina Limestone is

absent from the north-eastern Flinders. Those

rocks which sare mapped as Wilkawillina Lime-

stone in this area on the COPLEY 1:250,000

Geological Map are assignable to the Wood-

eee

Fig, 4. Bedding plane exposure of stromafolites, Woodendinna Dolomite,

Fig, §, Dolomite clasts in intraformational conglomeraie. Woodendinna Dolomite,

Fig. 6. Well developed mudcracks on the underside of a bedding plane, Woodendinna Dolomite.

Fig. 3. Cobbled upper bedding plane surface of ovid erainstone, Woodendinna Dolomite.

Fig. 8, Chevton-type cross beds in quartz sandstones, Woodenditma Dolomite.

Fig. 9, Ripple marks on bedding sutface of a thin

Fig. 10. Squat stramatolite bicherms on bedding a

Fig. 1 Impure flaggy-bedded timestones of the

of intra-formational conglomerate.

quartz sandstone, Woodendinna Dolomite,

lane, Wirrapowie Limestone.

irrapowie Limestone. Note the testicular aggregates

TZIK

endinna Dolomite and Wirsapowie Limestone

in ihe lower part and obviously hitherto wndif-

ferentiated, and in the upper part can be much

more justifiably related to Parara Limestone

lithologies than those of the Wilkawillina Lime-

stone,

Thin but well developed Woodendinna Dolo-

mile decs occur south of the line of the Blin-

man-Wirrealpa-Chambers diapirs along the

southern limb of the westward plupging anti-

cline near Mt Lyall (see PARACHILNA

1:250,000 Geological Map). Here it is overlain

with apparent conformity by the Wilkawillina

Limestone. Further south, toward the Wilka-

willina Gorge area, in the Central Flinders

Runges, the Woodendinna Dolomite rapidly

thins in response to erosion, of nondeposition

or both, At the Ten Mile Creck type section

itsell, Wilkawilljna Limestone disconformably

overlies the Bonney Sandstone Member of the

Pound Quartzite (Dalgarno 1964; Pierce").

The very highly dolomitized basal Wilka-

willina Limestone in this section, however,

superficially resembles parts of the Waonden-

chnna Dolomite, but duc to the degree of dia-

genesis, the original nature of these basal car-

bonates hus not been resolved. For convenience

they are considered at this stage to represent

dolomitized Wilkawillins Limestone — litho-

facies.

In the east-central Flinders Ranges, Gehling!

describes approximately 25 m of carbonates,

which are tithologically similar to Ihuse of the

Wooueniitina Dolomite, from the base of the

Cambrian carbonate sequence at Reaphook

Hill.

Along the entire western outcrops of the

Cambrian sequence, fram Parachilna Gorge

south, a thin seyuence of flaggy grey lime-

stones, dolomitic mudstones, stromatolites, fat

pebble conglomerates, and interbedded sand-

stones, o¢cur beneath typically fossiliferous

Wilkawillina Limestones. These beds are

assignable to the Woodendinna Dolomite and

the Wirrupowie Limestone and roughly cor-

respond to what Dalgarno called the !.ower

Member of the Wilkawillina Limestone (Dal-

garno 1964), although as staled above, these

lithologies arc not present within the defined

type Wilkawillina Limestone, Dalgarno &

Johnson (1963) report significant thicknesses

of Wirrapowie-type limestones from just. north

P. G. HASLETT

of Brachina Creek to south of Mt Aleck, and

Dalgarno (1964) records similar rock types in

the vicinity of the Chase Range. More recent

work has confirmed the presence of both

Woodendinna- and Wirrapowie-type carbonate

sequences in these southern anu soulh-westerit

party of the Flinders Ranges (Daily, pers.

connnt.}.

Depositionsal Environment

From the above descriptions, it is apparent

that although the Woodendinna Dolomite and

Wirrgpowie Limestone are distinctively dif-

ferent in gross mineralogy and in field appear-

ance, they share a number of lithological simi-

larities, They are intimately associated with one

another both temporally and spacially and

clearly have formed in closely related deposi-

tional environments.

The Woodendinna Delomite shows

numerous features characteristic of deposition

on emergent high intertidal and supratidal

mudflats, Prevalent desiccation tmudcracks

record repeated periods of suhaerial exposure.

and the highly restricted nature of the environ-

ment is suggested by the paucity of fauna and

dominance of primary dofamitic mudstones or

pene-conlemporancously dolomitized — car-

bonate mudstones. The anly organisms capable

of tolerating the extremes of restriction and

desiccation of the environment were algae, a5

evidenced by the ahundance of stromatolites

in the sequence.

Cross-bedded ooid grainstanes and quartz

sandstones record brief episodes of high cncrey

conditions penetrating the mudflats. These con-

ditions, which probably occurred during

periods of storm activity or times of unusually

high tides, resulted in the spreading of sbeets

of coarser clastic’ from more open marine

preas, over the expased mudflats, Quartz sand-

siones in the Woodendinna Dolomite show con-

siderable variation im their lateral development,

reflecting the local availability of a quaria sund

sauree, which in mony cares probably cor-

responded to areas of exposed, unlithified

Pound Quarizite.

The Wirrapowie limestone has developed in

4 sheltered but less restricted upper intertidal

to laguonal environment, The absence of dolo-

mite and the rarity of desiccation mudcracks

suggest that only limited exposure and restrtc-

tion prevailed, The sequence lacks coarse detri-

en, Etta

8 Pierce. PLR. (1969}.—The Cambriun geology south of the Wirrealpa Duapir, Flinders Ranges. South

Australin. Honours Thesis, Univ, Adelaide (unpublished).

LOWER CAMBRIAN FORMATIONS, FLINDERS RANGES

tus, current bedding and any preferred clonga-

tion of stromatolite bioherms indicating that

energy conditions were in gencral still very low.

As described above for the Woodendinna Dolo-

mite, however, thin grainstone shects attest to

the periodic penetration, probably during

storms, of high energy conditions into the shel-

tered tidal mudflat environment.

219

Acknowledgments

The writer wishes to acknowledge the use of

the facilities of the Geology Department, Uni-

versity of Adelaide, and to thank both Dr Daily

for his generous assistance over a considerable

period of time, and Mr C. R. Dalgurno who

read a draft of this paper and offered valuable

criticisms.

References

Coats, R. P., Cacten, R. A., & WILLIAMS, A. F.

(1973).—COPLEY map sheet, Geological

Atlas of South Australia, 1:250,000 Series.

(Geol. Surv. 8. Aust,: Adelaide.)

Dait¥, B. (1956) —The Cambrian of South Aus-

tralia. /n El Sistema Cambrico, su Paleogeo-

grafia y el Problem dé su Base 2: 91-147.

(xx Congresso Geol. Internacional, Mexico.)

Dawy, B. (1972a).—Aspects of carbonate sedi-

mentation in the Cambrian of South Aus-

tralia. Abstracts, Joint Specialists Groups

Meetings, Canberra, Geol, Soc. Aust. c10-cl4.

Dairy, B. (1972b).—The base of the Cambrian

and the first Cambrian faunas. Centre for Pre-

cambrian Research, Univ. Adelaide, Spec.

Pap. 1, 13-41.

Dairy, B. (1973).—Discovery and significance of

basal Cambrian Uratanna Formation, Mt

Scott Range, Flinders Ranges, South Austra-

lia. Search 4(6), 202-205.

DaLcarno, C. R. (1964)—Lower Cumbrian

stratigraphy of the Flinders Ranges. Trans. R.

Soc. §, Aust, $8, 129-144,

Darcarno, C. R., & JOHNSON, J. E. (1962).—

Cambrian sequence of the Western Flinders

Ranges. Quart. geal. Notes, geol. Surv. 8.

Aust, 4,

Datcarno, C, R., & JoHwson, J. E, (1963),—

Lower Cambrian of the Eastern Flank of the

Flinders Ranges. Quart. geol, Notes, geol.

Surv. S. Aust. 7,

Davcarno, C. R., & Jonnson, I. E. (1966) —

PARACHILNA map sheet, Geological Atlas

of South Australia, 1:250,000 series. (Geol.

Surv. S. Aust.: Adelaide.)

Hastetr, P. G. (1975, in press) —lLower Cam-

brian stromatolites from open and sheltered

intertidal environments, Wirrealpa, South Aus-

tralia. Jn M. R. Walter (ed.) “Stromatolites”,

(Elsevier: Amsterdam.)

Jouns, R. K. (1968).,—Geology and mineral

resources of the Andamooka-Lake ‘lorrens

area. Bull. geol, Surv. §, Aust. 41, 1-103,

Worrner, H. (1970).—Early Cambrian Pateogeo-

graphy, Frome Embayment, South Australia.

cia Odie Ass. Petrol. Geal. 54(12), 2,395-

THE SOUTHERN AUSTRALIAN SPECIES OF SPYRIDIA

(CERAMIACEAE: RHODOPHYTA)

BY H. 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* & SaLty A. CARTLEDGE*

Summary

WomersSLeY, H. B. S., & CartTLepor, Sally A, (1975).—The southern Australian specics of

Spyridia (Ceramiacene: Rhodophyta), Trans, R. Soc. S, Aust. 99(4), 221-233, 30 Novem

ber, 1975,

Four species of Spyridia aré recognised on southern Australian coasts. §. filamentosa

(Wulfen) Harvey (including 8. bianniata J. Agardh, S. breviarfictulata J. Agardh, and S.

spinella Sonder) is common in sheltered to moderately rough water. § dasyoides Sonder

(including S. oppesita Harvey and §. prolifera Harvey) is fairly common on rough-water reefs

and in deeper water, .S. squalida J, Agardh (including §. wilsonis J, Agatdh) is a less common,

usually deep-water, species. §, fasmmanica (Kuetzing) J. Agardh occurs in relatively culm loca-

lities 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 zeneric uniformity of the species

ascribed to Spyridix.

{introduction

While Spyridia 1s a distinctive and easily

recognised genus of the Ceramiaceae, the taxo-

nomy of the southern Australian species. has

been confused. Some 10 species have heen

¢redited to the region, but the only recent

account of the species by Lucas & Perrin

(1947) and a key by May (1965) offer little

help in recognising of separating the species.

The type species of Spyridia, S. filamentosa

(Wulfen) Harvey, has recently been described

in detail by Hommersand (1963, p. 177), who

reviewed earlier studies and clarified tis vege-

tative and reproductive features. Indian mate-

rial of this species has been studied by Krish-

namurthy (1968). Other species referred to

Spyridia agree well with the type in gencral

morphology, preserice of nodal and internodal

cell bands, branches of limited growih (ramelli

or “brachyblasts”) and uhlimited growth, and

in teproductive features. However. the features

which Hommersand (1963, p. 177) used to

distinguish 8. filamentosa from other specics of

Spyridia do not apply satisfuctorily to the Aus«

tralian species. Cortication by tiers of nodal

and internodal cells. is Found in all species (the

cell shape varies considerably in 8. filamen-

tesa). and all species have radially disposed

ramelli (Opposite in §. dusyeidex) which (ex-

cept S. squalida) commonly have mucronate

end cells.

Growth of the uniaxial thallus is from an

apical cell which cuts off a row of short axial

cells which develop into a branch of unlimited

growth, from each cell of which one or more

ramelli develop laterally, The ramelli are of

limited growth, developing rapidly to between

10 and 30 cells long by divisions of their apical

cell, and following cessation of cell division

they expand by cell elongation to their mature

length of generally 1-3 mm. The axial cells

also. cut off in alternating sequence a ring of

periaxial* cells, which form ia band around the

node between two axial cells, Each of these

petiaxial (nodal) cells cuts off two cells fram

its lower end, and these elongate and become

attached by pit-connections ta the nodal cells

of the next lower segment. Thus the thallus

shows bands of shorter and broader nodal cells

alternating with. the bands of internodal cells,

* Department of Hotany, University of Adelaide, Adelaide, 5, Aust, 5000,

* 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 polysiphonous families of the Ceramiales; are still referred to as

pericentrai cells.

199

ane A

which are longer, narrower, afd approximately

twice as many as the nodal cells, Further cor-

lication occurs some distance from the branch

apices, from descending rhizoidal cells deve-

loped from the nodal cells, and this obscures

the regular pattern of nodal and internodal cell

hands, especially in certain species (e.g. 3.

sptallda) -

The cells of the ramelli each cut off a ring of

&—-& cells from their upper end, and these deve-

lop into a nodal band 1-3 cells broad in S.

flamentosa, Spyridia is readily recognised hy

the alternating nodal and internodal bands, and

ramelli with corticated nodes.

Reproductively Spyridia is also distinctive.

especially in that the carposporophyte hecomes

surrounded by pericarpic filaments developed

from the. segments above and helow the one

hearitig the procarp. These filaments can give

the appearance of a cystocarp with a well deve-

loped pericarp wall, being held together by a

muciluginous shealh and some lateral pit-can-

nections, but they disintegrate fairly readily in

preserved material,

Procarps (3-4) are produced on small

lateral branchlets of resiricted growth, Accord-

img to Hommersand (1963, p. 191), three peri-

central (periuxiol) cells are normally fornted

in each fertile seement, one of which (the sup-

porting cell) bears the carpegonial branch, and

each pericentral farms an auxiliary cell, Hom-

mersand reparted that the carpogonitim fuses

with the third cell of the carpogonial branch

which then connects with the auxiliary cells. by

means of connecting cells. Thus two (or rarely

three) gonimoblasts are initiated and the ma-

ture cystocarp is commonly bilobed. Krishna-

murthy (1968, p, 48), however, observed only

two pericentral cells per fertile segment in

material from South India, and considered that

the fertilised car'pogonium divided into two

cells, each of which fused with an auxiliary

cell.

Spermatangia cover several cells in the lower

part of the ramelli, usually excluding the basal

celJ, They are derived fron filaments originat-

ing from the nodal cells, which grow over the

two adjacent cells, then cut off spermutangial

mother cells before forming the continuous sur-

face Jayer of spermatangix

H. B.S, WOMERSLEY & SALLY A. CARTLEDGE

Tetrasporangia occur on the lower cells of

the mumelli, being sessile and mostly on the

vipper (adaxial) side. Hommersand (1968, p.

196) considers that they arise directly as pro-

trusions from the axial cell, while Krishna-

murthy (1968, p. 47) considers that they are

fovried from. periaxial cells.

Some 10 species of Spyridie have been

recorded from southern Australia, This study

recognises unly four species, including 8. filu-

meniose. The thallus development and mer-

phology, and the structure of reproductive

organs, are very similar to those of S,_ file-

mentosd, and the Australian species differ

mainly in vegetative features. “he descriptions

below are therefore confined largely to recos-

nition of the species, with brief notes only on

reproductive details.

Key to southern Australian species

1, Ramelli robust, opposite and decussate, usually

100-150 vm thick with isodiametric cells aud

nodal bands 3-5 cells broau virtue

Sb. dasyeides (p, 231)

1, Ramelli slender, single or whorled but nat oppo-

site, less than 70am thick, Uxnally with cells

longer than broad, wnd nodal bands J—3 cells

broad 1 .a4Ht HPN coat aOR

2. Ultimate branchlets stout (4-1 mm thick),

murkedly basally constrictcd, heavily corti

cated to their apices, bearing slender, irmegu-

larly branched romelli_... S. sqndlida (p. 229)

2. Ultimate branchlets slender (under $ mom

thick), Not or only slighily basally constricted,

corticatinn only on older branches, with

ramelli either one per segment or verseeibine

. Ramelli one per segment, 35-65 4m thick, nods!

hands 2-3 cells brond = S. Mavnentoan tp. 222)

4. Ramelli becoming vecticillnte (1-8), 2040 am

thick. nodal bands 1 cell, broad ,

XN, rasenanica (p. 227)

Spyridia filamentosa (Wulfen) Harvey 1833;

436: 1844: 449; 1846, pl. 46: 1 855a: 337;

1859: 329, 1863. synop.; 42. J. Agardh

IR52: 340; L876: 248: LRAT: 13, Boerge-

sen 1917; 223, figs 222-2726. Feldmann-

Mazayer 1940: 348, Guiler 1952: 98.

Horamersand 1963: 177, figs 4-10.

Hooker & Harvey 1847: 409, Kiishna-

murthy 1968: 42. Newton 1931: 394. fig.

236, Okamura 1932; 130. Reinbold 1897:

60. Sonder L853: 480; 1880: 16. Tate

Fig. 1. Spyeidia flumentosy. A. Pt Stanvac, S. Aust. (Lewis, 14iv.1972; ADU, A41869). B. Fentate

Mant with spermatangial turelli (Aldinga,

Tetrasporangiol plant (A485),

5, Aust, Cortledge, 30.1 19727; ADL, A418IS), B-

223

THE SOUTHERN AUSTRALIAN SPECIES OF SPYRIDIA

nn ei.

Stee gy get

roe ®

Lb “AT 4)

~“\-0

TW Ss “Ryo pores, kL

Fig. 1,

224 H. B. S. WOMERSLEY & SALLY A. CARTLEDGE

THE SOUTHERN AUSTRALIAN SPECIES OF SPYRIDIA 225

1882: 18. Tivdall 1898: SOS. Wilson

1892; 181, Womersley 1958: 157,

Frees filamentous Wulfen 1803: 64,

4S, filamentosa var. arbuscula Sonder TH55: S18,

3. blannlate J. Agaidh 1876) 267: IR9T: 13,

De Toni 1903; 1426. Guiler 1952; 98. Lucas

1909: $3; 19298: 25; 1929: $3. Lucas & Perrin

1947; 363. May 1965: 369. Okamurs 1932:

130, Reinhold 1897: 60; 1899: 50, Sonder 1880:

i6, Tisdall [$98- S05. Wilson 1892: 18),

Wainersley (950: 180.

§. breviarticalaie J, Ayardh L&76: 268: 1897:

13. De Toni 1903: 1427, Guiler 1952: 98,

Lucas 190%: $2; 1929a: 25; 19296; $3. Lucas &

Perrin 1947: 363, May 196S; 369, Okumura

1932: 130, Reinbold 1897: 60; 1899: 50. Sonder

IS80; 16,

4. spinella Sonder 1845: 55; 1846. 168; L880:

16. J, Agardh 1852: 342; 1876: 269: 1897> 13.

Ne Toni 1903; 1430, Harvey 1863, synop.: 472.

Kuctzing 1849; 668; 1842: 16, pl. Sled, Lucas

1909; 52. May 1965: 369. Mazza 1935, no. 824.

Okamum 1932: 130.

FIGS I, 34, B

Thalluy (Fig. 1A) usually 7-18 em high,

epilithic or epiphytic on various larger algae

and seugrasses, lax and soft, irregularly much

branched on all sides with longer and shorter

branches intermixed. with one ta several axes

(often poorly detined) from an originally dis-

coid haldfast, soon becoming fibrous or stoloni-

ferous and entangled, commonly grey ta grey-

red, samelimes red-brown, in colour: Axes and

larger branches corticated, terete; axes !—1

(-1}) mm thick, tapering to branches 300-500

pm thick and branchlets 100-300 jm thick;

laterals arising from periaxial cells or adventi-

liously feam cortical cells. Segments usually

clearly defined on branchicts (Fig. 1B), vagi-

able in length and proportions but usually

(4) 4-1 times as long as broad, with bands. of

shorter noda} cells and longer internodal cells

alternating; nodes with 11-14 periaxial cells,

cach corresponding to two internodal celly ex-

cept far the (usually) larger periaxtat cetl bear-

ing the ramellus. Cortication usually commence

ing a few mm from the apices but very

variuble, consisting of rhizoidal cells lying

belween the internodsl cells and gtadually

forming a4 continuous cortex 1(—2) cells thick.

Ramelli (Figs 18, 34, B) single per segment,

inregularly spirally arranged, $-14 mm long

with 12-20/-27) cells, linear or gently taper-

ing apart from the terminal 2-3 very short

cells (Fig, 434) which taper abruptly to a

Mucronate cell, (35—)40-$5(-65) ym thick

with cells (1}-)}4-24(-3) times as long ss

broad; mucronate end cell often Inst from

older ramelll, ramelli with about 9 nodal cells,

each usually cutting off 1(-2) cells anteriorly,

giving a nodal band 2-3 cells broad.

Cystocarps (Fig. 1B) short-stalked, usually

bilobed, lobes globular, 300-700 aM across,

Spermatangia covering the lower (except

basal} several segments of ramelll (Fig, IC),

forming male organs 75-120 pm in diameter,

Tetrasporangia (Fig, 19) sessile, 1-3 per

cell on Jower cells of tamelli, mostly on the

upper (adaxial) side. spherical, 50-75 jin in

diameter, tetrahedrally divided,

Type locality; Adriatic Sea,

Type: 7?

Distribution: All around the Australian coast

{including Tasmunia) in conditions of ayoder-

ate to slight water movement.

Spyridia filamentosa is recognised ws a

widely distributed species, having beca

recorded from most seas, and many authors

fe.g. Harvey 1846, pl. 46; Feldijinn-Mazoyer

1940, p, 348) refer to it as a very variable

species. J. Agardh (1876, p. 268) in segresst-

ing two Australian species (S. Biannulatea and

S, breviarticulata) from 8, filamentosa, referred

ta their similarity in habit with S. filamentose

and the large number of forms classed as this

species, J, Agutdh apparently regarded his two

stgregate species with some doubt, und a de-

tailed study of extensive collections of Austri-

lian material does nod provide any satisfacsory

Way of segregating S. diannulata and S. brevi-

erticulata fram §. fillamentosa-

The type of S. bignnutata is from Tasmania

(Georgetown, Tas. Gunn, LID, 51300, selected

as lectotype) and was distinguished by J,

Agardh in having more conspicuous bands of

nodal and internodal cells and heiny less cor-

licated above, 8. breviarticnlara is based on

specimens from Whitsunday 1,, Queensland

(lectotype is LD, 51311}, and was distin-

guished by having the nodal und intervodal

bands short and of almost equal Jength, The

vaTauon in 3S, flamenrosa encompasses the

above features of bath S. bienmuata and S,

breviarticalata.

SSS

Fiv 2. Spyridia tasmanica. A, Portartington,

Vic, (Wollaston, V7,vili, 1956; ADU, A20567), B. Female

plant with young cystocarps (investigator Strait, S. Aust., 33 m deep, Haisun, 24,),.1971; ADU,

A41066). C. Male plant with spermatangial tamelli (Tapley Shoal. S. Aust, 15 a deep, Shey

herd, 2.111969; ADU, 433538). D. Tetrasporangial plant (A4 1066),

2246 H. B.S. WOMERSLEY & SALLY A. CARTLEDGE

Fig. 3. tidy S. filamentosa. Apex and mid region of ramellus, with periaxial cells in face view

41815).

C, D. S. tasmanica, Ditto ( A41065),

E, F. 8. tasmanica. Ditto, robust form (A37622).

G. S. lasmanica. Female axis with procarps. (Gt Taylor Bay. Bruny L., Tas., 10 m deep. Shep-

herd, 7.ii1.1970; ADU, A35287). c.br.—carpogonia

{ branch; p.a—periaxial cell; s.¢.—support-

ing cell.

A, ft. &.

J squalida. Apex and mid region of Lamellus, with periaxial cells in face view

(A26375).

J, K. 8, dasyoides. Ditto. (A20170)-

THE SOUTHERN AUSTRALIAN SPECIES OF SP¥RIDIA

S, filamentosa var. arbuscula Sonder (1855,

p. 518) from Wilsons Promontory, Vic., May

1853 (type in MEL, 45181) is typical of the

species and not a distinct variety.

S. spinella Sonder (type in MEL, 502090)

was based on Preiss material from Western

Australia. The type has somewhat broader and

stouter ramelli than most specimens of S.

filamentosa, with cells about as long as broad

and the nodal bands of the ramelli 2-3 cells

broad. Collections from Cottesloe, W. Aust.,

reef pools (Parsons, 14.xi.1968; ADU,

A34072) and from Elliston, S. Aust. (Womers-

ley, 15.14.1951; ADU, A15142) agree well with

the type in having densely aggregated ramelli

400-800 4m long, composed of 15-18 cells,

50-65(-90) ym thick and 1-14 times as long

as wide. This is within the extremes of S. fila-

mentosa and is probably typical of plants

occurring in rock pools subject to moderate to

considerable water movement. These plants

are much closer to typical S. filamentosa than

the Brest specimen discussed below, and are

provisionally placed under S. filamentosa, but

further studies on this “spinella” form and its

variation are desirable.

S. filamentosa has been studied by several

authors, most recently by Feldmann-Mazoyer

(1940, p. 348), Hommersand (1963, p. 183)

and Krishnamurthy (1968, p. 42). The Austra-

lian material agrees well with these descrip-

tions, and also with material from Leghorn,

Italy (Sartoni, 18.viii1973; ADU, A43938).

However, material from Brest, France

(Cabioch, Dec. 1972; ADU, A43050) has dis-

tinctly more robust ramelli (about 100 pm

thick, cells scarcely longer than wide) which

have 2-3 very small basal cells with the parent

periaxial cell not enlarged, in contrast to the

full-sized basal cells and enlarged periaxial cell

of typical S. filamentosa, These two collections

indicate that there may be greater variability

in European S. filamentosa than in the Aus-

tralian material.

Hommersand (1963) and Krishnamurthy

(1968) differ in some details in their accounts

of reproduction in the material they studied, as

mentioned above in the introduction. Austra-

lian material (e.g, Aldinga, S. Aust. Cartledge,

30.ii1.1972; ADU, A41881) shows three peri-

axial cells in fertile segments, but clarification

of immediate post fertilisation stages has not

been possible. No clear stages have been ob-

served of a tetrasporangium arising directly

from the axial cell of a ramellus (Hommersand

1963, p. 194), but it appears more likely that

227

periaxial cells are usually transformed into

tetrasporangia. The number of periaxial cells

in a ramellus does vary slightly, so it is not

possible to state as Hommersand does that

tetrasporangia must arise directly from the

axial cell because the number of periaxial cells

is the same in sterile or fertile segments.

Spyridia tasmanica (Kuetzing) J. Agardh 1852:

342. Gordon 1972: 39. Harvey 1859:

329. Kuetzing 1862: 14, pl. 42 c, d.

Sonder 1853: 680.

S. Jiameniots var. tasmanica Kuetzing 1849:

S. filamentosa var. verticillata Harvey 1844:

Wrangelia setigera Harvey 1859: 309, pl. 1914;

1863, synop.: 27, J. Agardh 1876: 622; 1879:

pl. 32, fig. 3. De Toni 1897: 133; 1924: 149,

Gordon 1972: 39. Guiler 1952: 99. Lucas 1909:

23; 1929a: 16, May 1965: 365. Mazza 1919, no.

678, Okamura 1932: 133. Sonder 1880: 29. Tis-

dall 1898: 511. Wilson 1892: 170.

FIGS 2, 3C-G, 4A-C

Thallus (Fig. 2A) usually 8-25 cm high,

irregularly much branched, epilithic or on

Amphibolis, with a small, discoid holdfast,

grey-red to red-brown in colour. Branching

irregularly alternate, branches terete, with one

to a few main axes and prominent lateral

branches, bearing lesser branches and branch-

lets on all sides, with whorled ramelli. Axes

1-14(-—2) mm thick, branches about + mm

thick, lesser branchlets 200-400 pm thick. Seg-

ments (3—)4—14 times as long as broad (Fig.

2D), with usually 12 periaxial cells, each pro-

ducing two internodal cells; cortication by rhi-

zoidal cells from the nodal cells, commencing

within 1-2 cm of apices and becoming heavy

on axes and main branches. Ramelli (Figs 2D,

3C-F) 1(-3) per node near apices, becoming

whorled (3-6(—8) per whorl with the addition

of adventitous ramelli), arising from enlarged

periaxial cells, 3-2 (-3) mm long with 20-30

(-35) cells of greatest diameter (15—)20-35

(—40) »m and 3-5 times as long as broad, the

end cell mucronate; robust form with ramelli

(12-)15-20 cells long, greatest diameter

30-40(—45) ym and 14-24 times as long as

broad. Ramelli with a single row of 8-14 nodal

cells (Fig. 3D, F).

Cystocarps (Figs 2B, 4A) terminal on a

short branchlet bearing ramelli, 4-3 mm in

diameter,

Spermatangia cover 2-6 cells (Figs 2C, 4B)

within 1-2 cells of base of ramellus, forming a

cylindrical male organ 70-130 ,m in diameter

and 1-6 times as long as broad,

H. B. S. WOMERSLEY & SALLY A. CARTLEDGE

THE SOUTHERN AUSTRALIAN SPECIES GF SPYRIDIA

Tetrasporangia (Figs 2D, 4C) on 1-4 cells

near the base of the ramelli, borne mostly on

the upper (adaxial) side, 60-100(-120) jam in

diameter when mature, tetrahedrally divided.

Type locality; Tasmania (probably Gan, ex

Hooker).

Type: L (941, 301.371).

Distribution: From Elliston, S. Aust. to

Western Port, Vic. and around Tasmania. Gen-

crally in relatively calm localities, offen with

cousiderable current, 2-35 m deep, occa-

sionally in purtly sheltered habitats and shaded

tuck pools on rough-water coasts.

Development of reproductive structures in

S. fasrnanica appears to be very similar ta that

in 3. filamentasa. The female axes (Fig. 3G)

arise as short laterals and bear up to 3 pro-

carps, separated by one or two sterile segments

each with eight periaxial cells, one of which

bewrs a ramellus. Each procarp consists of 3

(rarely 4) periaxial cells, one of which is the

supporting cell bearing the 4-celled carpogonial

branch. Immediate post-fertilisation stages have

nol been clearly followed, but usually two

groups af gonimoblast cells develop, probably

from fusion cells originating from auxiliary

cells cut off from the supporting cell and one

of the other fertile periaxial cells. Sterile peri-

carp filaments arise from the periaxial cells of

segments above and below the procarp-bearing

segment, forming a eystocarp similar to that in

3. filamentoset.

Development of spermatangia and tetraspo-

rangia is similar to that in S. filamentosa.

S, tasmanica is a distinctive species with its

wharled ramelli and single row of nodal cells.

The ramelli are typically slender (20-35 pm

thick), with mature cells 3-5 times as long as

broad. However, some plants from rougher-

water localities e.g. Ellistan. S. Aust. 7 m

deep (Shepherd, 20.x.1970; ADU, A37622)

and Cape Lannes, S. Aust., in shaded pool

(Kraft, 12.41.1972; ADU, Ad41809)] have

more robust ramelli, 30-40(-45) ym thick

(Figs 3E, F, 4B, C) and mature cells 14-24

times as long as broad. Otherwise the latter

specimens are similar to the majority of plants,

and the type, of 8. /asmanica, and the more

robust. shorter-celled ramelli are regarded as

229

more characteristic of plants found under

rougher-water conditions. Further studies on

this farm are. however, desirable.

Spyridia squalida 1, Agardh 1876: 270; 1897:

16. De Toni 1903: 1436. Lucas 1909; $2:

1929b: 53. Lucas & Perrin 1947; 264

May 1965: 369. Okamura 1932: 130.

Reinbold 1897; 60, Souder 1880: 16,

Tate 1882: 18.

5. wilsonis J. Agardh 1897: 16, De Toni 1903:

1435. Lucns 1909: 52. Lucas & Perrin 1947-

36d. May 1965> 396. Okamura 1932: 130.

‘. valida Sonder 1880: 16 (nomen nudum).

FIGS 3H, 1, 40, &

Phallus (Fig. 4D) usually 10-30 cm high,

robust, c¢rect, irregularly and praliferousty

branched, epilithic with one to several axes

from a small discoid holdfast, usually with

long, much branched, laterals on all sides,

grey-red to red-brown io colour and when

dried ofien appearing somewhat farinaceous,

All branches terete and corticated to their

apices, axes and main branches linear, branch-

lets (Fig. 4£) basally constricted and bearing

densely arranged ramelli, especially on their

upper parts, sometimes denuded belaw. Axex

1{-23 mm thick, denuded below or with short,

projiferous branchlets, tapering slightly to

branches 1-14 mm thick and lesser branchlets

{-1 mm thick, Segrrents largely obscured by

cortication, 14-14 as long as broad, with 16

periaxial cells and about twice ws many inter-

nodal cells; cortication commencing within a

few axial cells of apices, pseudo-parenchyma-

tous, 2-3 cells thick on branchlets, several cells

thick on axes. Ramelli (Figs 3H, [, 4E) one

per Segment close to apices and derived from

periaxial cells, but scattered adventitious

ramelli (usually) densely cover the branchlets,

sometimes persisting onto larger branchlets;

ramelli J-1(-14) mm long with (10-) 14-20

(—24) cells of greatest diameter (20-)30-40

(—45) um and (1-)14-2(-24) times ws long

us broad. Ramelli with a single row of small

nodal cells (Fig. 34) derived Crom 5-6 peri-

axial cells each of which cuts off 2-3 outer

cells in the same transverse plane.

Cystocarps short-stalked, globular-bilobed,

+4 mm in diameter,

a SSSSSSSSSSen

Fig. 4, Spyridia tasmanica (robust form), 4A, Femule plant with squashed mature cystocarp (Elliston,

S. Aust. 7 m deep in bay. Shepherd, 20.x.1970; ADU, A37622),

(A37622). C. Tetrasporangial plant (A37622).

tangial ramelli

Spyridia syualida. B. Victor Harbor, 8. Aust,

6. Male plant with sperma

(Wantersley, (8.11966; ADU. 430032). FE.

Branchlets and ramelli (Pt Elliot, §. Anst. Dodd, 12.41.1963; ADU, A26375).

TT eee di

H. B. S. WOMERSLEY & SALLY A. CARTLEDGE

THE SOUTHERN AUSTRALIAN SPECIES OF SPYRIDIA

Spermatangia cover the lower 2-6 cells

(except basal cell) of ramelli, forming a male

organ 30-80 am in diameter,

Teirasperangia borne on the lower several

cells of the ramelli, largely on the opper

(adaxial) side, 1(-2) per cell, sessile, spherical

to slightly ovoid, 40-60 «m in diameter when

mature, tetrahedrally to sub-cruciately divided,

Type focaliry: “Nov. Holland. australem".

Type: Herb. Agardh, LD, 51533.

Distribution; From Geographe Bay, W,

Aust. ta Waratah Bay, 8. Gippsland, Vic.,

usually in deep water (2-24 m deep).

Female axes of S. squalida develop as short,

adventitious branchlets which are more heavily

cortecated than in other species but less so than

in vegetative branchlets of this species, Alter-

naling segments each bear 2 procarp, with the

sterile scements beating ramelli, Usually three

Periaxtal cells occur in fertile segments, one

{the stipporting cell) producing a 4-celled car-

pogonial branch, Two, or probably often 3,

auxiliary cells are formed, Ieading to a carpo-

sporophyte with two of three lobes. The peri-

varp develops similarly to that in other apecies

S. squalida is a distinctive and robust species

of Spyridia, having cortication ta the apices

and thus forming swollen, busally-constricied,

branchiets, bearing ramelli often densely scat-

tered but usually soon denuded. The farina-

ceous appearance js also a common feature of

elder, dried plants.

5. wilsonis J, Agardh is typical 3. squatlida,

The type of the former is from Pt Phillip Hds,

Vic. (J. B. Wilson, 1887: LD 51532), and the

thallus is. not compressed as stated by J.

Agardh (1897) and May (1965).

S. valida Sonder (1880: 16) is a nomen

nudum, based on a specimen in MBL (45195)

from Geographe Bay, W. Aust. (Burberry.

1875), accompanied by Sonders drawings. It

is typical 5S. squatida.

WV, squalida was recorded from Port Alfred

{Kowie), South Africa by Barton (1896, p,

196), and the record repeated by De Toni

(1903, p, 1436) and Lucas & Perrin (1947,

Pp. 364), This record almost certainly applies

lo some other species, probably to S. plamese

Schmitz ex J. Agardh (G. F. Papenfuss. pers.

comm).

231

Spyridia dasyoldes Sonder 1853: 680; 1880:

16. J, Agardh 1876; 272. De Toni 1903:

1437, Harvey 1863, synop.; 42, Lucas

1909; 52. Lucas & Perrin 1947; 364. May

1965; 369. Okamura 1932: 130. Tate

1882: 18. Tisdall 1898: S05,

3, opposita Harvey 1855b. 256; 1860: pl, 158.

Adpens 1972: 83. J, Agardh (876; 270; 1897¢

4. De Tom 1903; 1431; 19242 $02, Guiler

1952; 9%, Lucas 1909: $2; 19293: 25; 1979b:

53. Lucas & Perrin 1947: 343, fig, 182. May

1965> 349 Mazza 1912. a0. 421. Okamura

1932! 130, Reinbold 1897; 60. Shepherd &

Womersley 1970: 135, Sonder 880: 16. Tate

1882; 18. Tisdall 1898) 405. Wilson 1892: 181.

Worersley 1950: 180; 1986: 151.

S. protifera Harvey 1863; pt. 274. J. Agardh

WW76; 26%; L897= L4. De Tont 1903: 143),

Lucas 1909- 52, Lucas & Perrin 1947; 362, fie.

(Rl, May (965. 349, Mazza 19127, no. 422.

Sonder 1880: Lf

FIGS 3/7, K, 5

Thallus (Fig. SA~C) usually 10-20 cm high,

erect, epilithic or epiphytic, much branched

with one to several axes from an originally dis-

coid holdfast which soon becomes fibrous and

stoloniferous, dark red to red-brown in colour.

Axes and larger branches heavily corticated,

terete to angular and becoming four-sided with

thickened cortical flanges in line with the 4

ranks of ramelli, densely branched, Axes 1-2

(-24) mm thick, often denuded but sometimes

with numerous, short, proliferous branchlels.

tapering to branches $-2? mm thick and lesser

branchlets 4-4 mm thick. branching usually

subdistichows (Fig, SC) with laterals arising

from nodal cells. Segrterts largely obscured by

cortication, 1~1 times as long as broad, with 3

perinxial cells producing 16 internodal cells

and the 8 cells soon with interposed rhizoidal

cells giving both nodal and tnternodal rings of

16 ceils (Fig. $9); cortication commencing

within a few segments of apices, nf elongate

cells later appearing pseudo-parenchymatous, a

few cells thick on branchlets, many (especially

on flanges) celis thick on axes, Rammelli (Figs

3J, K, SD, E) arising from an enlarged peri-

axial ceil, in opposite and more or less decus-

sate pairs (Fig. 5D) on successive segments

{often displaced to two rows on each side in

the plane of branching), (1—)14-2¢-24) mm

long with (16-18-22 cells, relatively uniform

=——— SS

Fig. 5, Spyridie dasyaides, A. Holotype (MEL, 45128), B, Robe, S. Aust. (Cartledge, 14.1972; ADU,

A42174)—an irregularfy branched form. C_ Investigator Strait, S, Aust,, 43 m deep {HW eatson.

27,1197), ADU, A38143)—distichously branched form, BD, Branch showing arrangement of

ramelti (Pt Denison, W. Aust. Aree,

14.x1L1971; ADU, A41730). E, Ramelli with tetra-

sporangia (Vivonne Buy, Kangaroo V., S, Aust. Womersley, 301.1956: ADU, A20170),

232

in diameter and tapering fairly abruptly to a

point, (70-)100-150 jm thick, cells about

b(JA) times as long as broad. Ramelli with

(6-20 nodal cells, eavh cutting of 1-2 cells

(which often divide again) on both sides (an-

teriorly first), producing a nodal band (2—)3-S

(-6) cells broad (Fig. 37, K).

Cystocarps short-stalked, irregularly globular

to bilohed, 400-600 pm in diameter,

Spermatangia cover the lower (except hasal)

several seaments of young ramelli of adventi-

tious branchlets lying between older ranvelli,

forming male organs 120-200 ym in diameter.

Tetrayporungia (Fig, JE) sessile, |-3 per

cell, mostly on the wpper (adaxial) side of the

tumelli, subspherical, 50-90 pm in diameter,

tctsahedrally divided

Type fovality: Holdfast Bay, S. Aust. (Py.

Mueller).

Type. MEL, 45128.

Distribution: From Port Denison, W, Aust.

ty Gabo 1. Vic. §. dasyoides usually occurs on

Tough-water coasts from Jow ude level to

depths of 33 m. Deeper growing plants are

usually more delicate than these growing in

turbulent conditions.

The reproductive cells develop very similarly

to those in S. filamentosa or S. rasmanice, Fe-

male axes correspond well with that illustrated

(Fig. 3G) for S. taxymanica, having alternating

sterile and procarpic segments, with the latter

comprising a 4—celled carpogonial branch on

the supporting cell and usually two other peri-

axial cells. The mature cystocarp encloses two

or ihree discrete carposporophyle lobes. and the

pericarp is relatively firm-at its periphery.

S. edasyoides is characterised by its robust,

opposite and more or less decussate ramelli,

with cells about a3 long as broad and nodal

bands 3-5 cells hroad, tovether witlr the largely

dislichous branching. Eight periaxial cells are

formed in branches and snon become separated

H. 8, S. WOMERSLEY & SALLY A. CARTLEDGE

by shizoidial cells, thus forming both wodal and

internodal rings of usually Lf cells, the banus

being about equal in length.

The type of S. dasyoides (Fig, 54) in MEL

is typical of this Species, previously known

mainly as S. oppesita, Sonder’s hand-written

label with the holotype gives the locality as

“Adelaide”, inland from Holdfast Bay. The

type of §. opposite (in TCD) ts From Preserva-

tion Harbour, on the south-west coast of the

south island of New Zealand (Lyall, Jan.

1851), and agrees very well with the southern

Australian plants. Adams (1972, p. 83)

records §. oppesita from several localifies near

Wellington, New Zealund, and although it is

apparently not widely known in New Zealand,

the specimens (cig. CHR, 55775, from Patu-

rau, Nelson) agree well with Australian mate-

rial. The type of S. prolifera Harvey, in TCD,

is From Fremantle, Western Australia (Clifton),

and is a plant with denuded branches bearing

proliferous branchlets. S. prolifera represents

older plants of S. dasyefdes, where the thick

axes and branches are probably cemmants of

the previous years’ growth, from which num-

erous short branchlets have arisen prolifer-

ously. The structure of the ramelii is identical

with that of §. dasyeides, Plants referred to 8.

prolifera are apparently not infrequent on the

Western Australian coast, where younger and

typical plants of S. dasyoides also occur.

Acknowledgments

The first author gratefully acknowledges a

grant from the Australian Research Grants

Committee and the technical usststunce pro-

vided by Mrs Enid Robertson and Miss Cheryl

Anderson. Dr G, Sartoni, Mstrlute Botanica,

Firenze, Italy. and Dr J. Cabioch, Station Bin-

logique de Roscoft, France, kindly made aval-

able material of 4. filamnentose from Curepe.

The Director of the National Herbariiim, Mel-

bourne, is Thanked for the loan of specimens,

References

Apams, Nattecy M. (1972).—The marine algae of

the Wellington urea A list of species. Rec,

Doom. Mus. 8(5), 43-98.

Agar J. Gi. (1852). -“Species, Genera et

Ordines Alzarum™. Vol. 2, P12, pp. 337-720,

(Lund.)

Acaspy, J, G. (1876).—“Species, Genera +t

Ordines Algacum”, Vol. 3, Pt. J, PP. 1-724,

Epicrisis systematis Flosid¢arum, (Liund.)

Aoarpy, J. G. {1879),—Florideernes. morphologi.

K. Wetenskt Akad. Forschandl, 1§(6), 1-199,

Plates 1-33,

Agaron, J. GG. (1897),—Analecta Algologica.

Goat. TVW. Acta Univ. tun, 8, |-L06, Plates

1-2.

Rorton, Gthel S. (1846),—Cape Alene. J. How.

34, 193-198.

BoERonsen, F. (1917).—The murine algae of the

Tamish West indies. Vol, 2. Rhadophyceae.

Part 3. Dansk. bot. Ark. 3, 145-240,

De Torr, G. B, (1897). —“Svlloge Algarum. om-

nium bucusque Cognitatium'. Yol. 4. Tori

dicae. Sect, 3) pp. 1-388. (Packus.)

Dy Tons, G. B. (1903) —"Sylloge Algurum cin-

Hlom hucusqne Cognitarium"”. Vol 4, Flori-

ileae, Sect. 3, pp, 775-1525. (Padua.)

De Toni, G. B, (1924).—“Sylloge Algarnm om-

fin bucusque Cogmlarium™, Vol. 6, Flori-

deuce. (Padus.)

THE SOUTHERN AUSTRALIAN SPECIES OF SPYRIDIA

FELDMANN-MAzoyer, G, (1940).—“Recherches

atir les Céramiacées de la Médireryanée occ

dentale’. (Alger). ;

Gerben, Elizabeth M. (1972) —Comparative

morphology “und taxonomy of the Wrange-

licae, Sphondylothamniege, and Spermotham-

miexe (Ceramiaceac. Rhodophyta! Aira? J,

Bot, Suppl. 4, 1-180.

Gui-re, E, R. (1952),—The Marine Algae of Tas-

mania. Check Jist with localities, Pap. Prov.

R, Saw. Tas, 8. 71-106,

Hawvey, W. H, (1833) —Aleae. in W. J. Hooker,

“The British Flota". Vol, 2, Part J, pp. 248-

401, [Also In W. J, Hooker, “The Rrelish

Flora of James. Edward Smith”, Vol. §, Part

1. pp. 252-405,]

Harvey, W. H, (1844),—Algae of Tasmaiiia.

Lond. J.. Bot, 3, 428-454, ;

Harvey, W. H, (1846)-—"Phycologia Britannica”,

Vol, 1, Plutes 1-72. (Loridon, )

Harvey, W. HH, (1855a)—Some account of the

marine botany of (he colony of Western Aus

tralia. Trams, BR, Irish Acad. 22, 425-566.

Harvey, W. H. (1855b)—Algac. Jn 3. D,

Hooker, “The Botany of the Antarctic Voy-

age". Part JI, Flora Novae-Zolandiac. Vol, 2,

pp. 211-266, Plates 107-121,

Haver, W. H. (1859) —Alpae. Jn J. D, Hooker.

“The Botany of the Antarctic Voyage.” Part

HT, Flora Tasmuniae. Vol. 2, pp, 282-343,

Plates 185-196,

Harvey, W. H, (1860),—"Phycologia Australica™.

Vol, 3, Plates: 121-180,

Harvey, W. H, (1863) —'Phycologia Anstratica”,

Vol, 5, Plates 241-300, synop.; 1-73.

Honmaprsanp, M, H, €1963).—The morphology

and classification of some Ceramiaceas and

Rhodomelaceae. Univ. Calif. Pubts Bot. 35

(2), 165-366,

Hooker, J. D., & Harvey, W. A, (1847) —Aleae

Tasmanicae. Laid, J, Bot. G, 397-417,

KaisNaMURTHY, V. (1968}.—The morphology of

sep tiaia filameniosa (Wuli.) Harvey. Phiykas

, 42-49.

Kurrzine, FT. (1849) —“Species Algartiim’.

(Leipzig. y

Kuetzing, Fo oT. (1862),—"Tabulae Phycolg-

gicac”. Vol, 12. {Nardhausen.)

Lucas, A. H. S. (1909) —Revised Tist of the

Fucgideae and Floridéae of Australia, Pree.

Linn. Soc, NSW. 34, 9-60,

Lucas, A. B.S. (19249a)—The marine algae of

Tasmania, Pap. Proe. R. Soc. Tashi, 1928,

Lucas, A, H, 5. (1929b).—A cepsus of the matine

algae of South Australia. Trans, R. Sac. 5.

Aw 53, 45-53,

T.uras, A. HS, & Perrin, FP, (1947).—“The Sea-

weeds Of South Australia". Part 2: The red

ecg Pp. 109-458. (Govt. Printey, Ade-

wide.)

Mat. V. (1965).—A censtis and key tothe spevies

of Rhodophyceae (red algae) recorded from

ert i Contr. NSW. natin. Herb, 3063,

-429,

23)

Mazza, A, (0912) —Saggio di ulgologia oceanicy,

Nuova Notarisiqa 23, Nos, 415-447,

Mazza, A. (1919) —-Saggin dl algologia oceaiiici.

Nvova Notarisia 30, Nos, 661-683,

Mazza, A, £1925) —Sagpio di alpologia oceanics.

Nos, 810-027 (yp. 1692-2096), (Publ. pri-

valely?}

Newron, T. (1931).—"A Handbook of the Bri-

tish Seaweeds”, (British Museum: London.)

OKAMURA, Ky (1932).—The distribution of marine

algae in Pacific waters. Ree, Oveanapr, Whe

Japan 3, 30-150.

REinnoto, T, (1897).—Die algen der Lucepede

und Guichen Bay. Nera Nofarisia 8 41-62.

REINBOLD, T. [1899)-—Méeresalgeen von Investi-

gator Street (Sud Australien). gegammelt von

Miss Nellie Davey (Waltham. Honiton).

Hedwigia 38, 39-51.

SHEPHERD, S, A.. & Womerszey, H. B.S. ¢ 1970).

—The sublittoral ecology of West Island,

South Australia, I. Environmental features

and the algal ccolugy, Trans. R. Soe, 8. Anil.

94, 105-238.

Sonper, O. G, (1845).—Nova algarum genera ef

Species, Quasx in itinere ad oras occidentales

Novae Hollandiae, collegit L. Preiss, Ph. Dr,

Bot. Zre 3, 49-57.

Sonprr, O. G. (1846).—Algae. I C, Lehmann,

“Plantae Preissianae’, Vol. 2, pp. 148-195,

Sonper. O. G. (1853)—Plantae Muellerianac.

Algae, Linnaea 25, 657-709,

Sonper, O. G, (1855),—Algae annis 1852 ef 1853

collectac, Linnaea 26, 508-528,

Sonpgr, O, G. (1880).—/n EF von Mueller.

“Fragmenta Phytographiae Austrilae™. Sup-

plemonium ad Volumen undecinum: Algue

petrelianae haclenns cogiitae. pp. 1-42, 105-

1u7,

Tare, R- (1882),—A list of the charas, mosses,

liverworts. lichens, fungs, and algals of extra-

tropical South Australia, Trans. R, Soc

Aust. 4, 5-24,

Tispaur, H. T. (1898).—The slgae ot Victoria,

Rep. 7th Meet. Aust. Ass. Adv, Sci. Sydney,

1898, pp. 493-516,

Witson, J. B. (1892).—Cuatalogue of algae col-

lected at or near Port Phillip Heads and

Western Port, Prac, R. Sev. Vies, a, 157-190,

WOLLASTON, Elise M. (1968).—Morphology and

iaxonomy of southern Australian: genera of

Crouanieae Sehmitz (Ceramiscenc. Rhodo-

phyta), Ays/. J. Bar, W, 217-417, Plates 1-10,

Womersiey, H, B.S, (1950).—The marine algae

of Kangaroo Iskind. TIL, List of species J.

Trans, R. Soe. S. Aust, 73, |37-197

Wowmrrsiey, 4, 8, 8, (1958),—Marine algae from

Arnhem land, North Australiu. Ree. “mec.

mart Sc. Exp, Ambheas Lund. Vol. 3, 139-

Womerstey, H. B. S. (1966).—Port Phillip sor.

vey, 3957-1963: Algae, Mem, natn. Mus,

Vict. No, 27, 133-156.

WULFEN, X. (1803).— Cryptogamu aquatica.

Roemer’s Archiv. Bot. (Leipziz) 3, 1-64.

OBITUARY: GRAHAM FREDERIC WHITTEN, M.SC.

BY P. B. WEBB

Summary

OBITUARY

GRAMAM FREDERIC WHITTEN, M.Sc.

16,1V.1924-16. 111.1975

Graham Whitten, former Deputy Director of

Mines in South Australia died on 16 March,

1975, after a long illness, With his passing the

State Jost an outstanding geologist.

Whitten graduated as B.Sc. from the Univer-

sity of Queensland in 1946 and subsequently

joined the Electrolytic Zine Company where he

worked as geologist and later Senior Geologist

at the Rosehery Mines in Tasmania, After

several years at Rosebery, he became respon-

sible for exploration fer the company in

Tasmania, New South Wales and Queensland,

In 1954 he accepted an appointment as

Senior Geologist in the Geological Survey

Division of the South Australian Department

of Mines, He was promoted in 1965 to Super-

vising Geologist and during the next five years

was responsible for all activities of the Explora-

lion Services Division, and in 1970 was

appointed Chief Geologist,

With the untimely death of the then Acting

Director of Mines, Dr K. R. Miles, in March

1972, Whitten was. left in administrative con-

tro] of the entire Department for a period of

cight months. He was subsequently appointed

Deputy Director, the post. which he held until

his premature retirement due to serious illness,

in November 1973,

Whitten made a significant contribution

towards the understanding of the iron forma-

tions of South Australia, In relation to this

work he visited Europe and North America in

1958 and 1967, and was awarded an M.Sc-

degree from the University of Adelaide,

He was elected Fellow of the Society in 1962

and was Secretary from 1970-72. He played a

prominent part in the affairs of the Geological

Soviety of Australia and was also an active

member of the Australasian Institute of Mining

and Metallurgy, He was also a member of the

Society of Economic Geologists and of the

Society of Exploration Geophysicists (ULS.A.),

His energy and drive will be remembered by

all who were privileged to know and to work

with him. His many official reports and

published papers are a fitting record of his

carcer.

To his wife Hetty and to his three daughters,

we offer our deepest sympathy. He will be sadly

missed by his professional associates and by

his many personal friends,

B. P. WEBB.

Ribliography

1963 Investigation of Aeromagnelic Anomalies.

hundreds of Carina, Chandada and Ripan—

Western Eyre Peninsula. Rept. favest,, geal,

Surv, $, dusr, 23.

Ironstone deposits, Radium Hill district.

Min. Rev, Adelaide, 117, 80-87,

Metallurgical testing of Peeralilla HiIl

laterite, Min. Rev, Adelaide, 117. 84-90.

Investigation of Kopi ucramagnetic ano-

malies. Min. Rev. Adelaide, LLB, 116-123,

The use of Aeromagneti: Maps in Geo-

logical Regions! Mapping, Quart. Geel,

Notes, geol. Surv, S. Aust. 14.

Tron Ore ueposits in South Australia out-

sidé the Midileback Ranges. Proc, Elvhth

Comm, Min. Metuil, Congress, Aust. New

Zealand, 1965.1, 309-3114,

Leached QOutcrops, Proc. Eigfih Comen.

Min, Metall, Cangress. dust, New Zealand.

1965. 2, 193-204.

The Geology of same South Australian Iron

Deposits. M.Sc. Thesis, University of Acle-

laide (unpublished).

Suggested Correlation of Iron Deposits

within South Australia, Quart. Geal, Notes.

geal, Surv. §, Aust, 18.

1967 Investigation of Carrow and Neill sero-

magnetic anomalies. Min. Rev, Adelaide,

121, 40-46.

Type Section of Iron Formations, Tarcoolit

District. Quart, Geol. Notes, z2ol. Surv, S

Aust, 26.

Atlas of Technical Data,

Notes, geal. Surv. S_ Angi, 26.

With Risely, B. G. A ground magnetic and

Bravily survey of the Warramboo Aeromag-

netic Anomaly, Central Ayre Peninsula.

Rept. ievest. geal. Surv. S. Aust, 32,

Regional geochemical investigations in the

Clere one-mile sheet areca. Mineral Resour.

Rev., 8. Aust, 127, 34-45.

‘The Investigation. and Exploration of the

Razorback Ridge Iron Deposit, Rep, Invest.,

geol, Surv. S. Aust, 33,

The [nvestigation of the Almanda mine

area, Wineral Resour, Rev, 5. Aust, 13),

25-37.

Annual Report of the Directar of Mines

and Government Geologist—for year ended

30th June, 1972. South Australia,

1968

1968 Quart, Geol.

1968

1969

1970

194

1972