(navigation image)
Home American Libraries | Canadian Libraries | Universal Library | Community Texts | Project Gutenberg | Children's Library | Biodiversity Heritage Library | Additional Collections
Search: Advanced Search
Anonymous User (login or join us)
Upload
See other formats

Full text of "Transactions of the Royal Society of South Australia, Incorporated"

VOL. 99, PART 1 



28 FEBRUARY, 1975 



TRANSACTIONS OF THE 

ROYAL SOCIETY 
OF SOUTH AUSTRALIA 

INCORPORATED 



CONTENTS 

Kott, Patricia The Ascidians of South Australia III. Northern Sector of the 

Great Australian Bight and Additional Records 1 

Bnonaiuto, M. F. Notes on the Genus Pseudomalaxis Fischer (Mollusca: Gastro- 
poda) and its Fossil Species in Australia - - - 21 

Gradwell, N. The Clinging Mechanism of Pseudophryne bibroni (Anura: 

Leptodactylidae) to an Alga on Glass - - - - 31 

Schmitt, L. H. Genetic Evidence for the Existence of Two Separate Populations 

of Rattus fuscipes greyii on Pearson Island, South Australia - 35 

Mawson, Patricia M. Two New Species of the Genus Cloacina (Nernatoda: 

Strongylida) from the Tammar, Macropus eugenii 39 

Bullock, D. A. The General Water Circulation of Spencer Gulf, South Australia, 

in the period February to May ------ 43 



PUBLISHED AND SOLD AT THE SOCIETY'S ROOMS 

STATE LIBRARY BUILDING 

NORTH TERRACE, ADELAIDE, S.A. 5000 



THE ASCIDIANS OF SOUTH AUSTRALIA III. 

NORTHERN SECTOR OF THE GREAT AUSTRALIAN BIGHT 

AND ADDITIONAL RECORDS 

by Patricia Kott 



Summary 

KOTT, PATRICIA (1975).- The Ascidians of South Australia III. Northern Sector of the Great 
Australian Bight and Additional Records. Trans. R. Soc. S. Aust. 99(1), 1-20, 28 February, 1975. 
An account is given of 58 species of the Ascidiacea from South Australia, of which 7 species are 
new, including two assigned to new genera in the sub families Euherdmaniinae and Botryllinae. 
Records of 22 species from the northern part of the Great Australian Bight are the first from that 
area and suggest that the ascidian fauna there has a considerable endemic component. Many of the 
species common in other parts of the Flindersian marine faunal Province have not yet been recorded 
from this location. 



1HE ASCID1ANS OF SOUTH AUSTRALIA III* 
NORTHERN SECTOR OF THE GREAT AUSTRALIAN BIGHT AND 

ADDITIONAL RECORDS 

by Patricia Kott* 



Summary 

KoiT, Patricia (1975). — The Ascidians of South Australia TTI + Northern Sector of the Great 
Australian Bight and Additional Records. Trans. R, Soc. S. Aust. 99(1;., 1-20, 2H Feb- 
ruary, 1975. 

An account is given of 58 species of the Ascidiacca from South Australia, of which 
7 species arc new. including two assigned to new genera in the suh families Tuherdmaniinae 
and Bolryllinae. 

Records of 22 species from the northern part of the Great Australian Bight are the first 
from thai area and suggest that the ascidian fauna there has a considerable endemic com- 
ponent. Many of the species common in other parts of the Flindersian marine fatmal Province 
have not vet been recorded from this location. 



Introduction 

This account of the ascidian fauna of South 
Australia is based on the following collections: 
( I ) from Ihe northern part of Spencer Gulf 
(made in connection with environmental 
studies in that area); (2) from the northern 
part of the Great Australian Bight (made in 
connection with an experimental Prawn Trawl 
Survey, Explorer); (3) from Investigator Strait 
(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). 1 lie report is 
supplementary to previous papers on the South 
Australian ascidian fauna (Kott 1972a, 1972b). 
It includes records of 57 species, including 7 
that are new to science. Two of these new 
species have been assigned to new genera in the 
sub-families Kuherdmaniinae and Botryllmae. 
Four species are newly recorded from South 
Australia. 

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 in. Australian 
museums as indicated by the abbreviations AM 
(Australian Museum), NMV (National Mu- 
seum of Victoria), QM (Queensland Museum), 
and SAM (South Australian Museum). 



All available collection data for the speci- 
mens discussed arc given in the Appendix. 

Order ENTEROGONA 

Suborder APLOUSOBRANCHIA 

Family CLAVEL1N1DAE 

Subfamily ciavelininah 

Podnclavelta cylindrica (Quoy & Gatmardj. 
Kott, 1972a: 5 (synonymy); 1972b: 167. 
New Records: Tipara Reef (Spencer Gulf T: 
on reel" NNVV Douglas Bank (upper Spencer 
Gulf). 

Subfamily HOLOZOINAE 

Distaplia anstraliensis Brcwin. 1953: 6L Kott. 

1957: 95. 

New Records: upper Spencer Gull (Stn D5). 

Previous Records: Tas. (D'Entrecasteaux 

Channel and southern Tasmania). 

Description: Colonies consist of a rounded head 

on a short cylindrical stalk. There is a single 

terminal common cloacal aperture and the 

zooids are arranged along cither side of the 

cloacal canals that radiate from this aperture 

and extend down the length of the head, There 

are about 12 flue longitudinal thoracic muscles 

Ten stigmata arc present in each of the four 

rows and these are crossed by fine parastig- 

mutic vessels. There are 8 rounded stomach 

folds, The gonads extend, from the pole of the 



* Queensland Museum. Gregory Tee., Fortitude Valley. Qld, Australia 4006. 



PATRICIA KOTT 



gut loop, /olo a short posterior abdominal 
extension separated from the abdumen by a 
shoa neck. Seven to 8 elongate testis lobes are 
nrrnnged in a circle with their long axes paral- 
lel to tme another to form a barrel shaped mass. 
the VftfS deferens, extending from the distal end 
of the centre of this mass, passes around it 
into the abdomen. There is also an ovary in 
the posterior abdomen. 

ftfnmfa; The colonies arc identical wilb those 
previously assigned to this species. The zooids 
differ from those described by Brewiu in the 
lesser number of stigmata in each row. Koti 
(1957) has reported some variation in this 
character and the differences are not regarded 
as significant, The presence of a paradigmatic 
vessel crossing the rows of stigmata has not 
previously been observed, but since this is very 
delicate it could have l*em overlooked. 

Syro/oa pcdunculata «juoy & Caimard). Kott, 
iy72b: 170; I972d; 234 (synonymy). 
rVt'vv Record: upper .Spencer Gulf (vStn IH). 

Ataposoa uiarshi Brcwih, VJHr. 31. Kott, 
1972b: 163. 
New Recants. Investigator Strait (Stlls Y5, 

m, 

bi'.icriptiotf The specimens arc of the usual 
form with a lonu cylindrical head terminating 
»n a rounded point. The shorter llcshy stalk is 
almost the same diameter as the head. The 
colony from Stn Z6 is the largest yet recorded, 
measuring 17 cm of which the stalk is ouly 5 
cm, The minute zooids are present in the sur- 
face layer of test with long posterior abdominal 
stolons penetrating into the centre of the lobe. 
I here is the usual brown pigment patch over 
the anterior cn<| of the cndostyle. 

Family POLYCITOK1DAK 
Ptilycilor gigantctnn tHerdman). Kou, 1972a; 9 
(synonymy |:. 1972c; 244. 
Mew Records: northern dear Australian 
Bight; West T. | Amphitheatre RocfcK 

Family FOLYCLIN1DAK 

Suhfamily uuhurdmanunai-. 

Eulierdnifcinia aiisimlfs Kott, 1957: 103; 1972b: 
172. 

New Rec/nds 1 Elliston Bay (outside bar>-; 

Investigator SlraiL (Sin YS), 
Di'stripriim; The colonies are iormed of the 
usual long club-shaped lobes joined basully. 
1'itch lobe is composed o\ a single zooid 
covered by its own separate sheath of sand- 



stilTencd test. 'I here are 9-1 1 rows of 27-2S 
stigmata, each row crossed by a paradigmatic 
vessel. A pointed papilla is present in the 
middle uf each primary transverse and para- 
stigmaric vessel oo both sides of the body. The 
internal wall of the siomaeh is arranged in 
longitudinal and transverse slandulaf ridges 
rather than folds. 

RiUerella hcrrimauia Koll; 1972a; II fsyno- 
nymy); 1972f>: 172; 1972c: 24<v 
Nrw Recant: Elliston Bay (outside bar). 
FIG. 1 

Description: The present colonics are suntlhjr 
than usual and sometimes each iobc contains 
only a single zooid. The lobes are the usual 
spatulate, long, narrow-stemmed form. Each 
zooid has 5 rows of about 5 sugrnara but there 
are no parastigmatic vessels, and a single 
papilla is present in the middle of each trans- 
verse vessel. There arc only single rows uf 
testis follicles in the posterior abdomina. There 
are 1 -4 embryos in the peribronchial cavity. 
Larvae ate very small, 0.3 mm long. They have 
1 median ampullae that alternate with the 
papillae, and double, rows of Vesicles lhal 
extend around the anterior aspect of the larvae 
on either side ol the papillae and ampullae and 
extend posteriorly along either side of the 
dorsal mid line Tlicie is also a paueu -series o| 
vesicles that extends postero-ventratly (Fig. I ). 
Remarks: This species has been taken from 
Elliston Bay (Kott 1972b) in May 1971. and 
the present colonies were collected in the pre- 
vious February. Only the latter arc sexually 
mature and contain larvae. It is not clear 
whether rhe colonics taken in May were newly 
settled forms, yet to reach reproductive matur- 
ily, or whether they were older colonics that 
reproduced earlier in the year. "However, the 
species appears to leproduee sexually at the 
vai\ of summer, Collections from I'ort Hacking, 
N.S.W. (Kott 1972c) indicate that Iheie. 
although new lobes were being added to ttw 
colonics at the end of Augusl. the species dis- 
appeared during the summer and did not return 
until autumn. Kecolonising- stock must there 
lure exist olf Port Hacking, which reproduces 
sexually at *he end of summer or early autumn. 
i.e. a similar seasonal cycle to that occurring at 
Elliston Bay. 

MATRIDIUM n. gen. 

/.ooidv completely embedded with both 
ripcrturcs opening separately to the exterior 
and withuut colonial systems. Internal lonei- 



ASCIDIANS OF SOUTH AUSTRALIA 1TI 




Fig. U Ritterelta hcrdmunia. Larva. 

Figs 2-3. Patridhim pulvinawrfu Fig. 2. — Portion of branchial sac showing internal longitudinal 

vessels. Fig. 3; — Thorax and abdomen of adult zooid. 
pigs 4_6. Aplitlium foliorum. Fig. 4. — Dorsal aspect of interior portion of thorax showing tripartite 

atrial lip. Fig. 5* — Stomach. Fig. 6. — Larva. 
Fig. 7. ApJidittm pronum. Thorax and abdomen of adult zooid. 

Fig. 8. Aplidium disihuum. .Portion of colony. 

Figs 9-10. Leptoclinides voh'Us, Fie. 9. — Colony. Fig. 10. — Thorax, 



tudinal vessels are present in the branchial sac. 
The stomach is folded. Gonads are present in 
the thrcad-lrke 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 
Euherdmaniinae 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 Protopo- 
lycUnum Millar, in which 3 species are known, 



PATRICIA KOTT 



viz. P, pedunculatum Millar. I960, Irojn Now 
Zealand; P. sabutoxa (Millar, 1963), from Port 
Phillip, Victoria; and P. claviforme Kolt. 19i>>. 
from Eden, rV.S.VV. 'I ylohrtwrliwn retains some 
primitive characters in the presence of the 
heart half way down the posterior abdomen, 
and a large ovary posterior lo the hunched 
testis follicles In ProropolycUmtm the stomach 
does iiol have longitudinal folds and the testis 
follicles are bunched in a short posterior abdo- 
men as in Polyclinum spp. "I he present genus. 
bears the same relationship lb ApUdtum us 
Protopolyclbutm hears to Polyclinum. It differs 
front both Protopofyclbwm and Pofyclintim in 
ihe presence of stomach folds and id its long 
thread-like posterior abdomen in which the 
testis follicles arc arranged in tows; it is these 
characters that relate it to ApUdtum. Jt differs 
from Ritterelhi, also in the subfamily Eubcrd- 
maniinnc, in the presence of ?hc longitudinal 
vessels, and the absence of parasUgmutic vessels 
in Ihe branchial sac, Riueretlo is usually further 
distinguished by the presence of 5 primary rows 
ofstigmala, although these arc often subdivided 
by paradigmatic vessels. The restricted num- 
ber of primary rows of stigmata suggests that 
Rittcrella may be more specialised than Pntri- 
tliiwi, which demonstrates primitive affinities 
■ j r the presence of it large number or rows, of 
stigmata a^ well as in the retention ot the inner 
longitudinal vessels, 

PatrMiiim puWlnatuw n. sp 

Type Uyrmum: northern Uteat Australian 
Bight t$t*WSk 133 v 10'Fh 42 ru deep. 
5.v.ry73. P S\woruL Httto/vpe: SAM. 
E ID35. 

FIGS 2. 3 
Description: The hololype o«Uy is available II 
is a circular cushion, h cm in diameter and 2 
cm high, mure or less flat topped and with 

rounded borders. Ir appears to have been ses- 
sile and attached by a small area jn the centre 
of the basal surface although there could have 
been a short stalk in this position. The test is 
very soft and semi-iransparcnt, generally with- 
out sand or other adherent foreign particles 
except for a small sandy urea at one side ot the 
basal surface. The zooids are thread like, the 
thoiax and abdomen together arc 1.5 mm long 
and the posterior abdomen about A mm They 
open all around the upper surface and the pos- 
lerior abdomina ptojeei down into the centre 
of the colony. The apertures are both 6 lobed 
Fine longitudinal muscle bonds extend along 



both sides nl the zonid for its whole length. 
I here arc about 25 rows of 16 short, oval, stig- 
mata; rather tall papillae are present on Ihe 
liarisverse vessels and these support longitu- 
dinal vessels running the whole length of the 
branchial sac. The longitudinal vessels are 
crowded, being .separated from one anolher by 
an interval equivalent, to Ihe width of about 
one anil a half stigmata. The oesophagus is 
J'airly long and there is a voluminous stomach 
about halfway down the abdomen with 25 con- 
spicuous longitudinal folds, The proximal pari 
ol the posterior abdomen does not contain 
gonads bin this region is often contracted. The 
ovary b present just anterior lo the double tow 
of testis follicles that occupy the greater part ot 
the posterior abdomen. 'I 'he heart is a wide U- 
shaped tube in the distal tip of the posterior 
abdomen. 

Pseiulutlixfurmi cere una Michaelsen. Kolt, 
1972a: 12 isynonyroy); ly72b: 173. 
New Record: Marxarel Brock Reef (Cane 
Jaffa). 

Remarks: Specimens in the present collection 
measure up to 12 cm of which the pointed or 
roimdcd head represents half of Ihe total 
length. The zooids, opening all around the 
head, are small, the contracted thorax and 
abdomen together measuring only 2 mm (Koti 
1972a. 2 cm ste), 

Subfamily r»oj,YcuNJN.\i: 
I'oJvclinnm neptunium Hartmcycr Kofi. 

I f /72b; 175 (synonymy). 

Ti& Record: Invexiiguior Strait (Stti VII, 
Description: The present colonies are small. 
with rounded heads only about 3 mm in dia- 
meter on thin, branching stalks. Each head 
contains about 6 zoo ids surrounding a central 
common c'oacal opening. The test h very deli- 
cate. There iv no sand inlernally but externally 
there is a heavy encrustation. 

The zooids are minute. The atrial lip is 
typical of the genus (Koti 1963) and has A 
longitudinal muscle hands It arises from above 
the upper dm of the opening and appears to 
close down over the aperture which is produced 
to point directly anteriorly. There arc 7 muscle 
bands radiating from the hranchial aperlurc 
but these do not extend onto the posterior part 
of the thorax. There are 5-8 small oval bJis- 
mata in each of the 7 rows, and papillae on ihe 
transverse vessels coincide with the sligmata. 
The gonads ufj! nol developed The stomach is 
smooth. 



ASC1DIAN5 OF SOUTH AUSTRALIA III 



Remark*. Although Lhe number of muscle 
bands in the atrial Up, and the number of rows. 
of stigmata and the number in each row. arc 
very much less lhan that usually reported for 
this species. The arrangement of the branchial 
papiHue is the same as that usually rcpoUcd 
(Kotl 1972b}. and it is possible that The colo- 
nies arc juvenile. 

A pit (Hum foliomm n. sp. 

hire Loatrion; northern Great Australian 
Bight (3Z C 24'S. U3':V>'U» ( 42 m deep ft 
Sytnond. 5.V.1973. lioloiype. SAM, h 1036. 
FIGS 4-b 
Dei(ripfiorv The colony is a circular cushion 
6 cm in diameter anil 2 cm high, forming a low 
dome, sljohilv conceive baldly where the zooid 
hearing surface layer of the test on one side has 
grown around onto the hasal surface to form 
a crescent shaped pocket invaginated towards 
the border of the colony where the surface 
zuoid bearing layer of test has grown to over- 
lap it. The test is soft, gelatinous and semi- 
transparent. There arc about i< common ctoacal 
apertures scattered over the surface of the 
colony, about 1.5 to 2 cm apart. Canals radiate 
tijit from lhe opening* lined on each side by 
rows of r.onids. These radiating canals sub- 
divide many rimes and zooid* lining them on 
each side crowd the test Zounds are at right 
angles to lhe upper surface. 

The branchial aperture is terminal with the 
opening surrounded by a circular sphincter. 
The atrial lip rising from the upper border of 
the aperture is very variable and may be simple 
or tripartite, while the lobes may be large and 
foliate or small and pointed There is a band 
ot muscles down the centre of each atrial lobe. 
There are 14 rows of about 15 siigmnta. The 
stomach is large with 18 to 25 narrow longi- 
tudinal folds. 

The zuoid* are long and thread-like, the pos- 
terior abdomen comprising the great part of 
rheir length while the thorax and abdomen To- 
gether are only 2 mm long. There arc one or 
two embryos at very different stages ot 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 
lestis follicles are present in two rows m the 
posterior .abdomen. The ovary is present ante- 
rior to the testis (in tbc usual position for this 
genus). 

/ iflwaw The larvae are 0.7? mm long with a 
long tail 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 or 
the cudostyle and to the post ventral aspect of 
the larval body. 

Henmrkx: The species is distinguished from 
Aplidiufn plidierum by the larger size of the 
colony, the very distinct radiating double rows 
Of yno'uU which comprise the systems, and by 
the characteristic foliaccous muscular atria* 
lobes that are present on many of the zooida. 
The colony does resemble thai of A. aitsrra- 
{ienxix which has similar systems and in which 
the branchial sac is the same. In A. utistw 
Uensis\ however, there is a lesser number of 
stomach folds ami they ate sometimes irregular 
and oblique, while the zoo ids lack the distinc- 
tive ainal lobes of the present species. The 
larvae of A plidferunu A, anxi rath nxis and the 
present species arc, however, identical. There 
are slight variations in we (e.g. larvae from 
the holotypc of A, australicttitx arc 0.9 mm 
long) and in the length ot 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 are similar to those described for A. 
datum Koit. 1972b, which however differs con- 
siderably from the present specimen in colon} 
form, 

Aplidiurn flavolineaturn iSluiter) Kott. JV72h: 
176 (synonymy). 
New Record- northern Great Australian 
Bight. 

Description' The colony is mushroom shaped, 
4 cm in diameter across the flat upper surface 
and 2 cm high. The flattened znoid-beanng 
head narrows very suddenly Jo a snort 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 
hut soft. The zooids are crowded in the lest 
and it was not possible io distinguish the form 
of the systems. Zonids open only onto the 
upper surface. They are 6 mm long, of which 
the thorax is 2 mm and abdomen only 1 mm. 
1 he posterior abdomina cross one another in 
the internal lest although the Iho/accs arc 
parallel ai the surface. The atrial lip is divided 
into 3 very pointed lobes from the upper border 
of the opening. There are 12 fine Jongiludinal 
muscles on the thorax. There aic 12 rosvs of 
10 long rectangular stigmata. The stomach is 



PATRICIA KO'I I 



especially small with 17 tlistinct rnngitudinal 
folds, there are 2 rows of testis follicles in the 
posterior abdomen. 

Larvae; Up to 5 developing cmbryus sit pre- 
sent in (he atrial cavity, They arc 0.75 mm 
long, have 3 median papillae alternating with 
Single median ampullae and corresponding 
lateral ampullae develop from the lateral ridge. 
The median ampullae arc narrow and in some 
cases appear to be bifurcated. 'I here are also 
clusters of ampullary vesicles both above the 
endo.stvlc and ventral to the body of the larva. 
'I he tail winds about three quarters of the way 
around the body of (he larva. 
Remarks: The colony and zooids arc of similar 
form to tho.se described previously for /T flavo- 
IffUititWn with the exception of the stomach in 
which there nre only 17 folds. Ihe size and 
shape of the stomach and the course of the 
longitudinal folds are similar to ihnsc described 
previously for (his species. 

Aplidium tonifenim Korr, 1963; 102. 

New Records. Ellision Reef. Pn-vlatL\ 
Records: N.S.VV, (near Twofold Bay, 10-70 
m deep: Montague North, I 3 m deepj — Koit 
1963. 

Liescripiion; Sessile* rounded lobes about 4 cm 
ii greatest diameter, the lest is sandy inter- 
nally, but the external layer of tesi is tree pf 
sand and is smooth and gelatinous. Zooids are 
long and narrow and open all around the head. 
There is a small pointed atrial lip from the 
body wall anterior to a muscular siphon Irum is 
present aboui one third of the distance down 
Ihe dorsal surface. The thotax is long and 
narrow with about 15 rows of 10 stigmata. 
There arc 5 stomach folds. 

Remarks; The specimen agrees with those pre- 
viously described. The clear external layer of 
test without sand hm] the form of the colony 
-arc apparently characteristic of the species. 

Aplidium amorphatum Kott, 1963; tfll. 

New Record; Klliston Buy. P *rt viatts Record: 
Vic. <38"5l'S, U6 B 55'E1— Koit 1963. 
D? sen prion; The colony is soft, sessile and 
dome shaped. The test h scmi-trarispajeiu. 
Without sand Zooids open all around the upper 
suiface and no systems arc evident. The zooids 
are very small and iirc-gulurly oriented in the 
test so that they cross one another. There are 
10 longitudinal thoracic muscles. The atrial 
jperturc is on a short siphon. The upper rim 
of the apcttUie i*. produced into a pointed Up. 



There are 12 rows of about Hi stigmata. Each 
row is crossed by a paradigmatic vessel. The 
stomach is small with 5 folds. 

Larvae: The atrial cavity is occupied by a single 
birgc embryo. I mm long. It has the usual 3 
median, stalked, papillae and numerous ampul- 
lary vehicles ate developed from the attterioi 
part of ihe body. 

Remarks: The specimen is identical with that 
previously described for the species, ihe larva 
fe the same as that of W. pantiles htiwi (see KoU 
1963). The shape and consistency of the 
colony differ, however, and tremble A. pw- 
tmems (Hcrdinan); Kou IM63, (n the latter 
species, however, the zooids arc larger* the 
branchial sac larger, there are more thoracic 
muscle bands and the paradigmatic vessels are 
not present. 

Aplidiuin proiium n. sp. 

Type Location: Investigator Strait (.Sin XI). 
( J in deep, Watson, Jan 197 J. Hoiotvpe: 
NMV. H 2s7_ 

FIG. 7 

Description; The colony consists of small, Hal- 
topped lobes united basally. The lest ts-*ofc and 
there is very little sand internally. There ft ;i 
single common cloacal aperture in the centre 
of each fobe. The zooids arc more than 1 cm 
long and thread-like. The thorax and abdomen 
me of equal length, and about one third of the 
total length of the zooid, The atrial lip is bifid 
or trifid terminally and extends from the upper 
border Of Ihe atrial opening which is on a short 
siphon, There are 1 I rows ot 12 .stigmata and 
8 very weak stomach folds. There arc 20 longi- 
tudinal thoracic muscles. 

Remarks: The colonics resemble those of A> 
novaezehmdiue lire win and A. cotfrelli (Brewiti) 
from New Zealand, although the Hal-topped 
lobes Biewin (1952, 1957 respectively) des- 
cribed for both these species have several sys- 
tems, only .1 stomach folds, and the atrial 
siphon is not produced. The atrial siphon of A. 
hiaritinttHH (Brewiti, 1958a) is. in fact, pro- 
duced in the same way as in the present 
species, and ihe stomach has the same ill- 
derined folds. However, the longitudinal tho- 
racic muscles are more plentiful in Ihe present 
specimens, there arc fewer stigmata in each 
rnw, the posterior abdomen is longer and mure 
thread-like, and there is only a single system in 
each lobe. Further. In Brcwin's species the 
Jobcs arc separate and do not appear tO be con- 



ASCTDIANS 0¥ SOUTH AUSTRAUA III 



7 



liituous In their ba*ui hall as in the present 
species. 

\ptiilium dtgitatum n. sp. 

Type Locatfon: northern Great Australian 
Hight ('32 l 74S, 133 C 3(VE), 49 m deep, 
s v. 1 973. /'. Sytnond. Holotype: SAM, E 
1030. Potorypes: QM, G 7508; AM. Y 
(982. 

tig. 8 

Description; The colonies are long, branched, 
cylindrical stalks 2- 3 cm long, terminally 
rounded and slightly expanded to overlap the 
stalk. The zooids open onto these terminal 
expansions. Sand is absent only from the sur- 
face rest where the zooids open on the 
expanded terminal portion of the lobe*. The 
stalk is densely encrusted with sand. The test 
is firm, especially in the stalk, and is impreg- 
nated with sand throughout. Zootds are minute, 
hui long and thread-like, crowded in Lhe test 
and extending parallel to one another down lhe 
sialk. There arc J 5 rows of about 8 stigmata. 
There is a long, pointed, atrial lip from the 
upper Haider of the opening. The stomach has 
12 longitudinal folds. There is a double row of 
testis follicles in the long posterior abdomen, 
there is a large common cloacal opening in the 
CCIlire of each lobe. 

iistn/irks: 'the colonics resemble those of the 
Antarctic- Species Apluiinm rccurnhfttx; Kott. 
19n9. nut are distinguished by the large num- 
ber of di.%tinct longitudinal folds in the sto- 
mach The zooids are especially delicate and 
narrow. 

ApUdfuiu colelloidc* (Merdrnan;. Kott, 1972b: 
176 (synonymy). 
AVh' Rcvoui northern Great Australian 

Bight. 

Family DIDEMNIDAE 
Poh.sviicraton aspieubUum Tokioka, 1949: 2. 
Kott 1962: 301. 

fvtvyynvrvtan ma^n'tlnrvum Millar, 1961: 13 
(nomen nudum): 1962: 165. Kott, 1972b; 
17*, 
Xt'w Reronf.x: northern Great Australian 
Bight; ?!nvcsligutor Strait. Previous Recants: 
W. Alls'. (Rnitncsi I , l*i Peion) — Kou 
1962. S. Aust. < Investigator Strait) — Kou 
1972b. Qld i.Mncfcay)— Kott 1902. S. Africa 
— MiMat 1962. Mozambique— Millar 1961. 
Japan — Tokioka 1949, 
Description. The colony from the Great Aus- 
tralian Bight is a soft jelly-like cushion. The 



common cloacal jvvstem consists of narrow 
canals at oesophageal level, radiating from 
common cloacal apertures and lined on either 
side by zooids. There are no spicules. The 
zooids arc of moderate -size, with 4 rows of 
stigmata and a long hifiiTcaicd atrial tongue 
The oesophageal neck is long. Gonads are. not 
mature. 

The colony from Investigator Strait (that is 
doubtfully assigned to this species) consists nf 
2 large flattened lobes rising from a fleshy 
cylindrical common basal stalk. The zooids are 
embedded in the surface layer of test. The com- 
mon cloacal canals extend between clumps of 
zooids beneath an especially thin layer of sur- 
face lest. There arc vcrv extensive cloacal 
spaces between the central soft test that forms 
the central core of each lobe and the surface 
that is only occasionally joined with the central 
core, hv solid test connect ives. Secondary canals 
extend between the zooids beneath a very thin 
layer of surface test, The lest is colourless and 
transpatent. and the zooids show through it as 
white dots. There are no spicules. Zooids have 
a long oesophageal neck, and the usual 4 rows 
of stigmata. Gonads are not mature in this 
colony. 

Remarks Specimens assigned to both P aspi- 
cnkvnm Tokioka, 1949 and /'. tnugmfarvum 
Millar. 1962, are soft and rounded, stalked or 
sessile, from 3 to 7 mm thick> with variable? 
spicule distribution to an aspicufar condition. 
The atrial lip is long, and often spread or bifur 
caLe at its tip. There are also, til both, a large 
number of testis follicles (8-12) and large lar- 
vae (over 1 mm long) with up to 15 pairs of 
lateral, finger-like ampullae and precocious 
buds, Zooids of both species are also charac- 
terised by a 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 cloacal 
canats that demarcate rounded zooid-free swell- 
ings of the surface of the colony. In neither 
species has the common cloaca been described 
as posterior-abdominal. The shape of the pre- 
sent colony I torn InvcstigatCu Strait is identical 
with others from this area that arc assigned to 
P. atpiculatitm (%P* nia^nilarvitm. Kott 
1972b). The posterior abdominal cavities in 
this colony, however, do nut occur in those pic- 
viously described. Positive identification is nut 
possible Owing to the lack of mature gonads, 
and the relationship of the extensive cloacal 
system wilh the simple canals that have been 



8 



PAT RIC1A KOTT 



previously described is not known. Ihe cloacal 
system is the same as that of Dtdetunum fam- 
ht/utti and Pofysymnttort cftondrWa but both 
these species have only a single aperture and 
arc not known without spicules, 

I.efKoclintdes voh'tts n. $p, 

Type Locution: northern Great Australian 
Bight (32°24'S, 133 30' fc), 42 m deep. P. 
Symond. Holotype; SAM. E 1034. PiVa- 
types; SAM. F. 1033; QM, G 751 1 

figs y. 10 

Description: The specimen designated as the 
holotype is a flattened sphere 5.0 em in dia- 
meter and 3 cm thick, with a thick but very 
short stalk, constricted to 2 em in diameter 
where it joins the body. The paratypes arc 
entirely spherical, about 3 cm in diameter, ami 
;irc without stalks although the base ol the 
colony is identified by the absence of zooids 
$n<l by foreign m<iiier lhal is included in the 
levt material which has overgrown the area. 
There is a single, apical, sessile* and inconspi- 
cuous common cloaca! aperture, opposite the 
base of the colony at the junction of several 
common cloacal canals. Zooids are arranged 
jlong both Mdcy of narrow cloacal canals at the 
abdominal level oh the zooids and the surface 
ol 1he colony is depressed above these canals 
These depictions demarcate rounded swellings 
ol 0w surface of ihe test corresponding to 
zooidTrcc areas. The tesc is very firm gela- 
iiunus and translucent- There is a layer of blad- 
der cells .superficially. A sparse layer of spi- 
cules is present in the zonid layer of test and 
Ihese are most dense around the zooids, thus 
indicating their position through Ihe test. The 
spicules jre minute. 0.015 to 0.02 mm in dia- 
meter, some stellate and others with needle- 
like rays. There are minute, spherical, brown, 
piemen! cells scattered throughout the test. 
Zooids arc small, with about 8 stigmata per 
row. The branchial siphon is of moderate 
Jengih, The atrial siphon is very short, rising 
Irom the mid dorsum, opposite the space 
between the second and third rows of stigmata, 
It is surrounded by a circular sphincter muscle 
and is posteriorly or laterally diieeicd. There 
arc 5j coils of the vas deferens and tip to 10 
testis follicles. Large ova are presenl in the test 
at abdominal level but none of these appear to 
be developing embryos. 

fteuu/rLv: The spherical body, constricted stalk 
and single common cloaca! opening arc unusual 
in litis 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 patt of the 
colony was fixed to the substrate, hrokc tree 
and was overgrown by the surface test. The 
constriction where the broadening stalk joins 
the head in the holotype also supports the sug- 
gestion that the spherical head may break 
away. Certainly the configuration of the ■sur- 
face, with the projecting swellings of solid gela- 
tinous test, while the zooids, their openings, and 
the common cloacal aperture are depressed into 
the surface of the test, would all accommodate 
U free living habit in which the colony is able 
to roll over the sea floor as in some coral spe- 
cies (see Glynn 1974, Pichon J 974). 

The limited nature of the common cloacal 
system is unusual m this genus where extensive 
posterior abdominal spaces are usually deve- 
loped. It differs front species in other genera, 
in which the /uoids are arranged along both 
sides of narrow common cloacal canals that 
extend around circular zooid-frce areas, in ih.tt 
the canals are at the abdominal rather than the 
thoracic level (see Polysyncwron aspiculatton, 
above; and Didemnmn fmudnm; Kott l*>72a), 

vStaJkcd species are also unusual in this genus. 
L. ftint?Sfonnis Kott, 1^7 2b being the other that 
is known. It is distinguished from the presenl 
specie^ by its undivided testis follicle. 

lA'ptoclinioVs reticularis (Sluiten. Kelt, l r >u2. 
285 (synonymy); 1972a: IS; 1972b- ISO 

New Record: northern Great Australian 
Bight. 

Oescnpjjon: A large colony, investing a speci- 
men of Hcrdnmnict momu\. There are streaks 
of orange-brown r stellate, pigment cells scat- 
tered amongst the spicules in Ihe surface test. 
The spicules have 7 rays in optical transverse 
section and are 0.03 to 0.05 mm in diameter. 
There ix a superficial layer of bladder cells 
rinsed with spicules. Spicules are dense at the 
zooid level but are absent basally. below ah- 
dorainat level. Common cloacal cnnnls weie 
not found and the thoraces oi the zooids 
appeared to be partially dislintegroted although 
the abdumina were in good condition with 5| 
coils of the vas delcrons around 5 ro *> testis 
follicles. Small vegetative buds are present in 
the oesophageal legion. 

Reworks: The absence of common cloacal 
canaW ;imi the condition of the zooids suggests 
that Ihe- colonies may present a ijinescenl winter 



ASUDJANS OF SOUTH AUSTRALIA 111 



co.idiliun. The presence of vegetative buds gjk} 
mature gonads suggests ihe onset rather than 
ihe end of ihis quiescent phase, 

Lepttnliniiks rofiw (Sluiier). Kott, 1962: 286 

(synonymy); Hldredge, 1907: 221. 

Ltt>joefinidvs ttmts; Millar, 1963: 704. 
Sew Records: Llliston Bay (outride bar. and 
25 n\ deep). Previous Records: S. Aust. 
i Port Noarl unga ) . Tas. (Maria I. ) . Vic. 
S Shoieham )— Kott 1 962. N .S, W. I Port. 
Jackson — Knu 1962, Millar 1963. Qld (Bar- 
jpra. Herun i.. Low Isles) —Hastings 1931; 
Kott J<>62. Indonesia U'alernosler 1., Ara- 
, „ra Sea l— Sluilcr 1 909. 1 91 V Tokioka 
1952, New* Zealand (North Island) — 
Michadsen 1924: Brewm 1958b; Millar 
I960. Hawaii (Otthu) — Hldredge I9u7. 

Description: Living colonies are reddish brown 
or grey with orange around the siphons. They 
axe firm and investing, with common cloacal 
opening* in the centre of evenly spaced 
rounded swellings. The common cJoacal aper- 
tures are about 1 cm distant from one another. 
There js a surface layer of bladder cell* and 
spicules are especially dense below this layer. 
They gradually become less dense and arc 
absent altogether from the basal halt of the 
colony, Although they ate present in a layer at 
the base of the common cloac&l cavity, they are 
stellate. 0.02 lo 0.04 mm in diameter with 
pointed conical rays. The spicules arc especi- 
ally densely accumulated around lobes ol the 
branchial apertuies. 

The common cloacal cavity is posterior 
abdominal, extending into circular chambers 
beneath the common cloacal apertures. 

The zooids are about 2 mm long with up to 
\2 longitudinal thoracic muscle bands from 
which fibres branch and anastomose with adja- 
cent bands. There is a minute circular lateral 
organ opposite tbe 4th row nf stigmata. The 
atrial aperture is directed posteriorly as is usual 
for the genus. There are up to 10 stigmata in 
each row. The oesophagus is. long. The gut 
ioop may be curved, although it also occurs 
as a long straight loop. The gut is clearlv dif- 
ferentiated into duodenal and mid intestinal 
'eeions and a posterior stomach swelling. The 
stomach is smooth and rounded, longer than 
its diameter. There -ire 9 to 10 testis follicles 
and 1\ coils of the vas deferens. 

Remarks; The synonymy of f . /furu Hastings, 
from Low Isles, with L. tufas originally des- 
cribed from Indonesia and the Arufura Sea to 



the north, within the same biogcographical 
region, was arrived at after comparison of 
Hastings type material (AM. GIJA49) vvrih 
other specimens Irom 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. (liemetiewis 
has not been examined, although I okiolcu 
(1952), Millar (1960). and kott (1962) have 
not been able to identify any character that 
could distinguish the two species. Its synonymy 
with L. ru/us is here maintained, The range of 
/ rtiftts is, therefore, similar lo that of many 
wide ranging species of this family in boih 
tropical waters <Kotl 1974) and in Antarctic 
waters <Kott 1969a). The other related New 
Zealand species, L. \htireri, L. uunmticus and 
L. tunae-zeia/uliae, are distinguished only hy 
the larger number of coils of vas deferens, the 
retalively shorter oesophagus and the smaller 
zooids. Further specimens are needed to ade- 
quately determine the punmeteis of each of 
these species. L. ruftts is characterised hy its 
relatively large zooids with distinct longitudinal 
thoracic musculature but no retractor muscle; 
by its long oesophagus and gut loop .ind the 
clearly demarcated 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 Ihe 
posterior part of the thorax. The number of 
testis lobes in the present specimens is greater 
than the maximum of 7 previously recorded. 

Didcmiuim candidum Savigny. Kott. 1972a: is? 
fsynnnymy); 1972b: 179. 
New Revon): northern Great Australian 
Bight. 

IMdemnum moscleyi (Herdman). Kntt, I97?a: 
19 (synonymy); 1972b: 179. 
New Record: northern Great Australian 
Bight. 

Trkliilenmurti sarigiili (Kerdmaivl s. sp. sawpnii 
Herdman, Kott, 1966: 2K5 isynnnvmv); 
Eldtedge 1967: 179. 
New Record- Sellick Heacti (south of Ade- 
laide), S, Aust. 

Description: Extensive investing colony with 
round, smooth, margins. There is a spicule-free 
surface layer of bladder cells. Spicules arc 
sparse in the zooid layer and are absent alto- 
gether trout the basal half of the colony. The 
common cloacal canals are deep, emending the 
whole length of the zooids, bur they arc not 



ID 



PATRICIA KOTT 



posterior abdominal. The spicules are 0.04- 
0.014 mm in diameter, They arc stellate with 12 
conical rays in optical transverse section- The 
ionuls arc surrounded by black pigment par- 
ticles Ihat are often but nut always accumulated. 
into the Usual pigment patch at the anterior end 
of the endusryle. There is a distinct alrial 
siphon which I* laterally rather than posteriorly 
directed in these colonies. There is a distinct 
retractor muscle. The testis is not mature and 
ihc vh< deferens was not distinguished. 

Remarks.' No further evidence is available from 
the examination of these specimens that could 
clarify the relationship between this lndo 
Pacific subspecies and the Atlantic Ocean form 
T. vavignii subsp. a/roctnwm Van Name (sec 
Kott l%b). it should be noled that the Pacific 
Ocean specimens (Eldrcdgc J 967) hnvc the 
7— 8 coils v\' ihc vas deferens that is associated 
with (he Indo-Pacific form (Kott t966). 

Tridiftemmim ccrebriforiue Hartmeyer. Kott. 
1972c: 47 (synonymy) 

Tcididemnutn stivt'tjnii; Tokioka, 1%7; 80; 

vt(r. j<A*nn?: t*2. 
New Record: Sellick Beach (south of Ade- 
laide). S Aust. 

S>ej.crtption: The colonies are investing and of 
variable thickness. Conspicuous common 
cloaeal apertures with frilled lips are distri- 
buted randomly over the surface of the colony. 
Posterior abdominal cloaeal 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 common cloaeal 
cavity, they are entirely absent from the basal 
layer of test. The spicules arc stellate, 0.02 to 
n. (14 mm in diameter, wilh about 6 rays in 
optical transverse section. 

The body wall of the thorax is covered with 
black pigment particles although these arc 
absent from the abdomen. The pigment par- 
ticles arc accumulated in a patch over Win 
anterior end of the cndostyle. The atrial siphon 
W posteriorly directed. There are 6i coils of 
the vas deferens around an undivided testis 
follicle. 

Remu/ks: The relationship of this species To '/ . 
.?<nv?w'i is perplexing since its three-dimensional 
common cloaeal system provides the principal 
distinction T, terebrifonne acquires great com- 
plexity in its common cloaeal system with 
prowth. but juvenile colonics must necessarily 
display a cloaca! system identical with that of 
T. sovignii before its subsequent prolifcratiot ,, 



as the colony thickens and surface folds deve- 
lop. In the present specimens, both taken fuun 
Sellick Beach, each species has spicules of dif- 
ferent sizes although this siac difference W&4 
not observed in specimens previously described. 
The number of spicule cays as previously 
reported, howevei. is greater for T. savignti 
than for T. cerebrifortne. 

Diploouma [ransUiculuin l.Hanmeyer) 

lwpio\'hnum I Leju'uclbumt ) trunsltKidum; 
Kou. 1962; 306 (synonymy | 

N,*w Record. Investigator Strait (Stn X1j. 

Previous Records: Indonesia- Sluiler 1909. 

North Western Australia — Hartmeyer 1919. 

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

Description: The colony K irregularly lobed, 
investing weed or other ascidians.. Each lobe is 
flattened, about 2 cm wide, 0.5 cm thick and 
up to 3 cm long. One colony completely enve- 
lopes a specimen of Pyura atisiralfs in which 
only the apertures are exposed. The surface K 
smooth, and the zooids show through as while 
dots. They are small and crowded at the sur- 
face of the colony in groups of about H. 

There is a large ovum present in most zooids 
but the testis is not mature and no coils of the 
vas deferens were detected. 
Remarfo: The tough, firm transparent test and 
the extensive cloaeal system is characteristic of 
this species. 

Family ASCID1IDAE 

Asrictia tbouipsoni Kott; 1972a: 27 (synony- 
my); J 972b: 181. 

New Records: upper Spencer Gulf. 
Dcsaiption: Specimens have a gelatinous test. 
sometimes thick and furrowed. Both apertunrv 
arc present on siphons, usually both directed 
dorsally or anteriorly and sometimes very long. 
The animal is fixed ventrally and by most of 
the letl side. There is sometimes a coating of 
sand encmsting the body, but sand is never pre- 
sent on tlic siphons. The body wall has the 
usual mexhwork of muscles on the right side 
of the body. The anterior part, of the dorsal 
lamina is a double membra ne. ribbed on tl>e 
outer sides but not in the centre. The ribs of 
rbe dorsal lamina extend into pointed projec- 
tions on the free edge of the membrane. 

The neural ganglion is about one-third of 
the body length from the dorsal tubercle. 

Rntuvks; The specimens from Station D3 on 
the flour nl the channel are encrusted with n 



ASUDJANS OF SOUTH AUSTRALIA III 



M 



layer of sand absent only from the siphons 
which project upward from the middle pf the 
Upper surface. In this specimen, the right side 
of lhe foody is narrower than the left and com- 
prises the right side of the upper surface, while 
the base, by which the animal is fixed, is the 
ventral surface and a large part of lhe left side 
of the body. In specimens from G4, also on the 
floor of the channel, the siphons are especially 
long, both directed upwards, and the right side 
of the body is similarly short, 

The very long external siphons directed up- 
wards, and ttie sand encrustation, are unusual 
in this species. Axcidia aclara is the only spe- 
cies of the genus in which a similar sand en- 
crustation hardens the lest. The long, cylin- 
drical extension* of lhe test that, in A, aehtra* 
CWlte a canal or tube from the sessile *per. 
tutcs extending upwards from the animal, are 
analogues rather than the homologues of the 
long siphons in the. present forms. It is aJso of 
interest that, in specimens from stations D> 
and K4> where the siphons extend dorsally, 
their central position, from the middle of the 
upper surface of the body, is achieved by rela- 
tive narrowing (i.e. between the dorsal lamina 
and the endostyle) of the right side of the 
hotly; whereas in A. actaru it is the left side 
of the body that is narrower than the right. The 
base of lhe present specimens is the ventral and 
two-thirds of the left surfaces of lhe body, 
while in A. achtra it is two-thirds of die light 
side, between the dorsal lamina and endostyle 
(Kott I972d). 

The specimens appear to have adapted to 
their free existence on a shifting sandy sea 
Hoar by these morphological variations in the 
position an<l length of the siphons. 

Ascidia aclara Kott; L972dr 236 (synonymy). 
New llftord: Goal I (on" Ccdunu)- 

Family STYELTDAE 
Stolonka carnosa Millar, 1963: 734. Kott. 

New Record: Investigator Strait (S*tn Yl). 

Reounks: The present specimens are small and 
sandy individuals joined by stolons. There are 
only 3 stfjrmara per mesh and the longitudinal 
vessels in the branchial .sac are slightly fewer 
lhan previously reported, viz. DL0(5)2(4)1E. 
It is probable that thk is a young colony, which 
is. characterised by a pyriform stomach wiih 
narrow folds and a Jong curved caecum that 
extends into lhe gut loop from lhe suture alone 
the lateral aspect of the stomach. 



Amphirarpa Hiptyehu (Hartmeyer). Kott, 1972c 
(synonymy). 
New Record' northern Great Australian 
Bight 

Ocutircuia auslralis Gray. Kot1, 1 972a: 29 
Synonymy); Kolt, 1972b: 18-1- 
New Record: ENiston Bay. 

Symplegnia viride Hcrdman. Kott, 1952: 252 
(synonymy); 1962; 129, Millar, 1966: 3ftS. 
Plantc & Vasscur, 1966; 149. ToKiofca, 
1967? Itf? (synonymy). Viisseur, 1967: 
111. 
New Record: EDiston Reef 

Subfamily kotryi unai. 

PARABOTRYLLUS n. gen. 

Colonics are elongate branching stalks 
slightly expanded terminally. Otic 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 cloaca! 
aperture. The terminal ampullae and conspi- 
cuous blood vascular system that arc present in 
other genera of this subfamily are tthscnt The 
rim of the branchial aperture is smooth. The 
in rial aperture hits a single anterior lip. There 
arc only 2 internal longitudinal vessels in the 
branchial sac. Eggs are endogenous. 
Remarks. The zooids are not conspicuous^ 
different from those of the genera Botrxllus and 
Botryijoides except jp the presence of only 2 
internal longitudinal vessels in the branchial 
sac. 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 lobe. Buds art- 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 clove 
connection with the parent until a late stage 
in their development as they do in other genera 
of the sub-family. The zooids art endogenous 
and have three io four developing ova on each 
side as in Botrylhts, while Botrylhides produce 
only single ova on each side of the body, 

Pwrabotryllos neraorus n. sp. 

Type Location: upper Spencer Gulf (Sin Ci), 
m deep, floor of channel. 5.ix.l973. Holo* 
fv/xvSAM, E 1031. Paratopes; AM < Y 1981; 
OM. Q 7507. 



12 



PATRICIA KOI I 



FIGS 11-15 
Description; The colonics consist of narrow, 
sandy lobes, 1-1.5 cm long, usually branched. 
I he superficial layer of test is encrusted with 
sand but is neither stiff nor brittle. Internally 
sand Is absent and the test is very soft. There 
are circular systems of zooids opening onto the 
upper free end of the lobes surrounding the 
central common cloaca! apertures that are 
.slightly depressed into the surface. Generally 
there is only a single system in each terminal 
branch of the colony although occasionally 
ihere are 1 or 2 smaller additional systems. The 
blood vessels in the lest are short and relatively 
tew for rhis subfamily. They terminate in elon- 
gate rounded bulbs at the base of the zooids. 
The zooids arc about 2 mm long. The rim of 
the branchial aperture is smooth and the atrial 



aperture, in the anterior 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 fas in Polyclhutfrt), There are 
10 rows of stigmata with about 10 stigmata in 
each row and two internal longitudinal vessels 
on each side of the body. The gut forms a 
single lighl loop on the left side of the bran- 
chial sac. The stomach is pyrifonn wiih about 
8 distinct longitudinal folds and there is a short 
curved caecum, of moderate length and ex- 
panded into a terminal bulb, from the pyloric 
end of Lhe stomach. There is a connective ex- 
tending from the pyloric region of the stomach 
to the intestine. There is a single, flat, testis 
follicle with lobed margins on each side or 
the branchial sac just anterior to the gut loop. 




Figs 11-15. 



Fit 16. 
Fig. 17. 
F;c. 18. 



ParabairyUm rwmorus. Fig. 11. — Portion of colony showing branching stulks. Fig. 12. — 

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

(dorsal aspect) shoeing endogenous ova. Fig. 15. — Testis. 

Polycttrpa tincior. Aberrant individual with atrial siphon produced anlvTinrly 

Pyura tendata. Section through the body wall, test and sandy coating. 

Mtcmtosmite plaints. Gut and gonad on left side of the body. 



ASCIDIANS OF SOUTH AUSTRALIA Til 



13 



The vas deferens, arising from the middle of 
the mesial surface of ihe testis, is very priori. 
There arc three or four ova in (he botly wall 
anterior to the testis lobe. These are endo- 
genous and project inlu the peribronchial 
cavity. Developing butls arc present in the test 
on either side of the posterior end of the adult 
zooids. These contain tour large ova on each 
side of the body and a clump of large cells 
dorsal to the ova. It is possible thai these may 
he precociously differentiating buds. The buds 
at this slaec of development are 0.2.5 -0.5 jnrn 
long- There is a blood vessel extending from the 
posleriot end of each bud in the region of Ihe 
oesophagus. 

Remarks: The species is distinguished fmm 
others jn the sublamilv by the large number 01 
oVvi and internal longitudinal branchial vessels. 
lbs relatively limited blood vascular system in 
the test, and the form of the colony and the 
limited development of colonial systems The 
internal tes! is also very soft in comparison 
v\ith that ot other species in the subfamilv The 
buds appear to undergo the major parr of their 
development in the test in connection with the 
colonial blood vascular .system and in the pre- 
sent colonies there wete no buds found directly 
associated with the parent zooids, The testis 
of this species is reminiscent ol lhal in Sym- 
piegma, while the multiplicity of ova resemble 
Rotryftas and ihe endogenous nature uf their 
development and the small circular systems in 
the colony are also reminiscent ot the latter 
genus. 

Subfamily srvi t.iNAU 

1'olycarpa tine tor (Quoy & Gaimardi Kott. 
1964; 134 (synonymy); lV72h: 1X6; 
1972c: 254; I972d; 242. 
Nrw Record; upper Spencer Gulf I.Stn 
BIOL 

FIG. 16 

Remarks: One of the specimens is highly modi- 
fied. It has the usual short branchial siphon 
irom the anterior end of the body. Ihe atrial 
siphon, however, extends anteriorly from the 
posterior half of the dorsal surface parallel with 
the anterior half of the body and opens at u 
point more or less level with the branchial 
aperture, so that the individual is U-shaped. 
The lower half of the body is encrusted with 
large particles of sand but the upper half has 
only fine sand encrusting it, and it appears that 
ihe animal had been half buried in the floor of 
the channel and that the atrial siphon was pro- 



duced upwards so that it opened above the sea 
floor. In another specimen theic are long root- 
like processes from the ventral border ot ihe 
body, which is otherwise typical of the specie* 
(Fig 16), Jn the U-shaped specimen there is n 
single row (17) of long polycarps around 1bc 
ventral border of the body and only occasional 
pulycarps, representing a second row. scattered 
dorsal to these* 

Polyesirpa perhinculata iHclIct't. Kott, 1972a; 
35 (synonymy); 1972b; I8fi 

New Records: upper Spencer Gulf; northern 

Great Australian Bight; Investigator SftraU. 

For Previous Records, Description, see Knit 

1972a, 1972b. 
Remarks: From the upper pari of Spencer 
Gulf, arenaceous and naked specimens are 
laken, sometimes growing side by side attached 
to the same shell or stone. There are a large 
number ol specimens and they are either 
stalked or sessile. Naked specimens vvilh a 
leathery test were a bright yellow colour in life 
but pinkish in preservative. Living arenaceous* 
specimens were a sandy colour with a reddish 
tinge There arc small, smooth, black indivi- 
duals in the preserved material and some that 
are larger with rough leathery and rather thin 
lest. In nature such specimens with a smooth 
<c$l arc bright yellow IS. A. Shepherd, pers. 
corn in. i. 

Polycarpa papillate (Stutter). Kott, 1972a: 34 
(synonymy). 
New Record: upper Spencer Gulf (Stn B7l 
Remarks: 2 specimens only are available. One 
is juvenile with a whitish coloured test and 
without developed polycarps. The oiher mature 
individual has a row of eight long polycarps 
around the ventral aspect of the left body wall. 
with the ducts directed towards the atrial open- 
ing. There are live rounded anal lobes slightly 
bifurcated and only two polycarps are present 
in the middle of the right body wall. The speci- 
men otherwise conforms with previous des- 
criptions of this species. 

SUi'la pedata iHcrdman). Kott. 1972b; 185 
fsynnnymy). 

New Records: jtorthem Great Australian 

Bight 

Slyela plieuia fl.eseuer), Kott, |$?2b; IH5*. 

1972c: 254: 1972d: 239 (synonymy). To- 

kioka, 1967. Abbott & Johnson, 1972: 95. 

New Record: Pofl Rivet (St Vincent Gull). 



N 



PAIRICIA. KOTT 



Remarks: The individuals arc small and the 
rounded swellings of the lest tire objured by 
epiphytic growth. However, the species is 
readily distinguished by Ihc short v$$$ elTeren- 
ua, branched testis folliecs, undulating (some- 
iinies branched) ovarian tubes, lorn/ oeso- 
phagus, long recluni, deep secondary cut loop, 
lomi voluminous stomach with internal folds 
and the small leaf-like endocarps that cover 
the body wall and the gut loop distal to the 
stomach. 

The gut loop is reminiscent of thai of S. 
tvtmftcaw (Kott 1972d) although the rectum 
and oesophagus arc longer in the present spe- 
cie^ and although the gonads resemble those 
of S, partita (Stimpson), in the present spe- 
cies the oesophagus is shorter and the vasa 
efferent ia -are shorter (Vasseur I9b7>. 

Inoniiluuirpn «chcriduii (Hodman). Kott, 
1972a: 31 (synonymy); 1972c: 253, 

iVev Record: northern deal Austratum 
Bighl. 

Family PYDRIDAK 
Pyuratendata Kott, 1972b: 186. 

New Reco?'d; south of Goat I. (off Ceduna } 
FIG. 17 

Description: A single specimen only is avail- 
able. It is more or less a half circle in outline, 
one cm in diameter. The external siphons ex- 
tend from the anterior and posterior ends of 
the more or less straight dorsal surface. The 
branchial aperture is directed anterodonally 
and the atrial aperture is directed poslerodor- 
villy. The external test is covered by a thick 
coating of sand held in place by hair-like exten- 
sions from the test, There is a thin space 
between the sandy coating arid the surface of 
the test, traversed only by xhc base of the test, 
hairs. There is also n coating nf very fine sand 
on the surface of the test itself. The apertures 
are lined with it very tough invagination of the 
test. The branchial siphon is especially mus- 
cular and appears to be cversihlc. The bodv 
wall is also very muscular. The atrial siphon is 
muscular but not eversible and its aperture is 
protected by a well developed vehnn at the 
distal end of the siphon. Beyond this velum the 
tvsi is produced into a cylindrical fibrous 
extension for a short distance. There are seven 
hranchial folds on each side of the body with 
14 strong internal longitudinal vessels very 
closely placed on each fold. There are no in- 
ternal 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- 
gucts. The gut forms a narrow Mraifiht loop 
and there is q mass of branched liver tubules 
in the gastric area. Gonads, are divided into 
separate paired polycarp-bkc sacs extending 
along both sides ot the central common duct. 
Remarks: This specimen agrees in most aspect* 
with those described from Investigator Snail, 
although the sandy coating is not as thick in 
the present specimen. Nevertheless, the nature 
and orientation of the siphons are identical £S 
arc the internal organs* viz. the branchial sac 
the gut and the gonads, The atrial velum in the 
specimen from Investigator Strait was present 
at the base of the atrial siphon rather than 
more terminally as in the present specimens, 
and this is possibly related to thickness of the 
sandy coating. The test beyond the velum is 
apparently produced to accommodate the thick- 
ness of the sand surrounding the animal, 

Pyura pachydcrmattna (Herdmau) s. sp guV 
bosa (Heller). Kott, 1972b: 187. 
Cynthia gihhow (Heller). 1878: 27, 
Pyura mbftow; Michaelssn & Wartmeypr, 
1928: 410. Non P purhydrtrtwtitut vur. j|fA« 
bosa: Kott, 1952; 265 (</>. pachvderrnaHita 
dras-vhii; Kott, l*J72b: IH7), 

PyUrrt parhyilermatu Vara intvrmvditi; Kali, 
1952: 264 (synonymy) (part) Non P £lh* 
host* Mh'rtacdia Miehaelson, 1922.* 3<)l. (</*, 
xpitiijcni: KotL I972h: 187). 

New Record: northern Cheat Australian 
Bight For Previous Records; Descrip ton. 
see Kutt 197 2b ( P pachydenmtlno 
dtavchii) ; 1 9$2 ( P. pachydermatina inter- 
media ) 

Remark*: The present specimens have the 
typical curved spines in the branchial siphon 
and the uiius is bordered with shallow rounded 
I ones, 

P> lira spiiitfera (Quoy & CaMMrd), Kott, 

1972b: 186 (synonymy). 

A/r w Record: northern Great Austral tan 
Bight 

Pyura aiistralis (Quoy &. Cat ward). Koit, 
1972b: 186 (synonymy). 

New Rerord: reef, Douglas Bank (upper 
Spencer Gulf). 

Pyura scorcsbicnsls Kott, 1972a: 36; 1972b: 
1*7. 

New Record, upper Spencer Gulf rfStnv }'\, 



ASCIDIANS OF SOUTH AUSTRALIA ni 



13 



Pyura vtttaila (StimpsonJ. Kotl, 1972a: 37 
(synonymy); lV72d; 243. 
Net*. Record: upper Spencer Gulf (Stn Al). 
Remarks; The spines lining siphons are typically 
long (0.1 mm) and ncedlc-likc and ovcilappng. 
"I he anal hordcr is smooth and two-lipped, 

Pyura Irregularis (Herdman). Koit. 1972a: 38 
(synonymy); 1972b: 187. 
New Record; upper Spencer Gulf (Stn B7). 
Remarks: The perttuhercular area has the usu- 
ally blister-like appearance tha? is characteitstic 
of this species. The large dorsal tubercle, how- 
ever, is at (he top of that area rathei than, as 
has been previously described, at its base. The 
lough leathery test and strong branchial &ac 
characteristic of this species are present. 

Pyura slolnuifera (Heller) s. sp, praepijtialfe 
Heller. Kott, 1952: 274 (synonymy); 1964: 
141, 

Pyura prae put talis; Millar. 1966. 372. 
bfiip Record: Outer Harbour (St Vincent 
Gulf). Foi previous Records and Descrip- 
tion .see Kntt, 1952; Millar 1966. 
Remarks: This location apparently represents 
the western extent of this species, which has ;i 
continuous distribt ilion from Queensland flown 
the eastern Australian coast. The specimens 
here at the apparent end of its range axe 
smaller lhan have usually been recorded from 
other locations, 

No consta ni dilference has been detecttul 
hciwecn ihe South African P. uolonifera s. sp. 
itotorujera and ihe Australian P. xtotomfera s. 
sp. praepuriaUs and most characters demon- 
strate a remarkable and overlapping ran.ee of 
variation in the two populations. The rounded 
fold of test enclosing ihe 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, (he different frequency with 
which these characters occur in each popula- 
tion suggest that suhspectfic rank is appropriate. 
This matter is discussed more fully by KoH (in 
press). 

Halocyiithia hispida (Hcrdman). Kott« 1968: 7(i 
(synonymy). 1972a: 41, 1972b; )S9, 
rVw Record: upper Spencer Gulf (Stn G\. 

Hcrdmonia mourns tSavigny). Kott, 19 72a: 4] 
(synonymy); 1972b: 1*9, 
New Record; upper Spencer Gt)1l (Sin G); 
northern Great Australian BighL 



Mieroeosmus planus n. sp. 

Type Lotulity: south of Goal L (olf Oduna), 
31 m deep. I7-25.xii.1967, Hewlett. Hok). 

type; NMV. 4284. Paraivpes. SAM. £ H>32. 
QM, G 7510. 

FIG. 18 

Description; The individuals are circulat in out- 
line and laterally flattened, with both apertures 
close together on the upper surface. The test is 
ihm and completely encrusted with sand, so 
that the specimens resemble hardened discs of 
sand. The sand is maintained around (he ani- 
mal by hair-like extensions of the test, and 
posteriorly these are longer, so that a flattened 
sandy keel is developed which interrupts the 
circular outline of the body, ft is apparent, 
therefore, that these laterally flattened indi- 
viduals are embedded upright in the sand 
rather than lying on their side on the surface 
of the sea floor. 

The apertures arc 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 arc 
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 sac has 
7 folds on each side of the body with six 
internal longitudinal vessels on each fold. 
Parastigniatic vessels are present. There arc 
rectangular stigmata but no internal longi- 
tudinal vessels between the folds. The dorsal 
lamina is a wide, plain-edged, membrane. The 
gut forms the usual long, narrow, curved loop 
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 alnat aperture, forming a deep 
curve vvilh the gut loop. The anal aperture is 
bi-labiatt!. The gonads are present in the sec- 
ondary gut loop and consist of a long ovarian 
tube often forming deep, close, curves, Ihe 
ovarian tube has dense male follicles along its 
posterior border and on the lateral aspect nf 
the ovarian tube against the body wall. As the 
curves of the ovarian tube develop, the male 
follicles appear to mingle with the ovaiy. 
Owing to the curving of the ovarian tube, it 
often appears to project back into lobes 
bordered by male follicles. The gonad on the 
right side of the body is in a corresponding 
position to that on the left. 



I* 



PA UK I A KOI'I 



R*»iark\- The species is unusual in the semis 
ilt thai the gonads do nol cross mlo ihe 
primary gut loop. Id this respect only, it rc- 
semhles Microco\tnus stotoniiera. the lorm ot 
the gonads in the present specimens is, how- 
evcr, distinctive and the laterally Hnllened body 
is also diagnostic of the species. 

VWcrocosmns squamigcf (MichacKcnl, (Cult, 
1972a: 43 (synonymy). 

New Retards upper Spencer Gulf <Stn G); 

Outer Harbour (Si Vincent Gulf). 
Remarks, The present specimens have U thick 
lest that ft impregnated wlh sand, and in some 
cases numerous specimens form aggregates. 
The siphons arc a rosy pinkish colour and 
lltere are the characteristic flattened scales 
lining the ouiur part of the siphons. 

Mu-roeosmiis nichollsi Kott; 1972a: M (synony- 
my): I972U: 245 

New Retards, northern Great Australian 
Bight. 

Miixoeosmus .stoUinif'era Koll; 1972;*: 43 

(synonymy), 1972d; 245. 
New Records: upper Spencer Gulf (Stn El), 
Remarks: The position of the leli gonad in Ihe 
secondary gut loop ts characteristic of the 
species. 

Family MOt.nL'LIDAK 

Molgula mollis (Heniman). KoU, 1052; 298; 
1964: 144; 1<>72a: 45 ('synonymy). 
Mult>t/ttt mhitlo.ui; Kott, 1 97 2d: 248- 
New Records: upper Spencer Gulf (Stn 
BIO); Investigator Strait (Stn YD. For 
Previous Recortk, Description, see Kolt 
1972a (Molijula sabtdosa). 
Remarks: I lie specimens are small, more or 
less laterally flattened, spheres. The apertures 
are close together on the upper surface and a 
ridge of slightly thickened lest extends Between 
them, 'there are line hairs on the lower part 
of the test which is completely encrusted with 
scjnd. Ihe rim of the aperture is lobed but 
there arc no hollow test expansions surround- 
ing I hem as in M, xabidosa (sec below), The 
apertures are directed away from one another. 
there are 7 branchial folds on each side of the 
body, with up to 9 internal longitudinal vessels 
nu each distributed over both sides of each 
fold, There are no internal longitudinal vessels 
between the folds, The testis follicles form a 
complete* chele at the end of the ovary, wirh a 
ligumwl extending through the centre of this 



circle. On The lateral aspect of Ihe ovary, it is 
apparent ihat the testis follicles embrace the 
end of the ovatian tube, hut on ihe mesial 
aspect the U is completed to form S circle by 
the jerowth of the male fotlieles acmss Ihe sur- 
face of the ovary. The male follicles are long, 
and on their lateral aspe-cl lie along trie body 
wall dire:ted from the periphery of the circle 
into Ihe eenlre. On the mesial surface of the 
gonads, the testis follicles can be seen to be 
more or less lan-shuped and tightly packed 
with their outer border divided into separate- 
lobes, The vas deferens extends from Ihe 
centre of the circle of testis follicles and. 
viewed from the outside of the body, has two 
vasa clTerenlia connecting ducts from each 
individual testis follicle on each side. The vas 
deferens then extends mesially jind along the 
surface of the ovarian tube and opens above 
the opening of the oviduct. The proximal part 
of the vas deferens is expanded into a dentinal 
vesicle. 

Remarks The lenglh of the oviduct and the 
absence of hollow test extensions around the 
apertures disrjmjuish this species from the veiy 
similar AY. sahtdosa (see below) which has 
alien been confused with it. 

Morula fflbtilostl (Quoy & Gainiard). 1834; 
613. Miehaetsen & HaHmeycr, 1928: 449 
(synonymy).K.ott. 1952: 29X (parti: 1 972b: 
19U. Millar, 19o6; 374* 

Remarks: I here are no new record* lor this 
species: however, a small specimen from Elhs- 
tou has made it possible to compare the species 
characteristics with those of M. mollis with 
which it has been confused. It is clear that the 
differences in the two species arc not associated 
with maturity. M. sabulosa is spherical with a 
sandy test ihat is hard and brittle, while M. 
mollis, although encrusted with test, has fine 
hairs to which the sand adheres, the test itself 
is more flucekJ, and the preserved specimens 
are laterally flattened. Ihe branchial aperture 
is always protected by 6 pointed lobes from 
the sui rounding test a fiille dislance from the 
opening while the rim of the aperture itself is 
produced into 6 smaller pomled lobes, thai an? 
covered in the closed position by the rim of 
the larger lobes. 'Ihe atrial aperture is pro- 
tected by two flattened, wide tongues and theii 
border is separated into three rounded lobes 
that arise from the test at the dorsal and 
ventral sides of the opening. The rirn of ihe 
aperture itself is putduccd mto 4 small, 



ASC1D1ANS OF SOUTH AUSTRALIA Ul 



<• 



pointed, sandy lobes and these are covered by 
the larger lips in the closed position. All these 
extensions from the test around the apertures 
are hollow and have prolongations of the body 
wall extending into them They are characteris- 
tic of the specie* and are never present in M. 
mollis. The gonad in the present species, >vhile 
superficially resembling that of M. tnollii,; has 
a very short vas deferens I hat opens at the 
proximal end of the ovary on its mesial surface. 

Uio&eO£r»pliy 

The 22 species rccoidcd from the northern 
part ol the Great Australian Bight (including 
Ccduna), a* far as 32"24'S, l-i3°30'E. can be 
divided into the following groups: 

1. Possibly endemic to the Great Australian 
Bight. 

Patridium pnivbtatutn n. sp.; Aptidinm digi- 
uxtum n, sp.; Aplldium foiiontm n. sp.: Lep- 
toclinides volvus n. sp,; Parabotryiltts ncmo- 
rus n. sp.: Pyura tendata Kott; Mierocosmm 
planus n. sp. 

2. Southern temperate ( recorded aWo I rom 
South Africa), 

Aplhtium flavolineatum (Sluiler); Aplidtum 
t'oIeUoides (Herdman). 

3. Cirt tt tn -an sf rattan , 

Pojycitor xixunteutn ( Herdman ) ; Lepto- 
clinides reriadutus (Sluitcr); Leptocliitides 
cuius (Sluitei ) ; Potysyncraton ospictdatum 
Tnkiokn; Didemnum candidum Savigny: 
Uidemnnm moselcyi ( Herdman); Atupht- 
ttirpa dipfycha (Hai'tmcycr): Polycarpa pe- 
dwtcuhita (Heller); Cncmidoctupa ciheridxii 
(Herdman) (absent only from tropical Aus- 
tralia); tlerdmania momus (Savignyl 

4. Southern and eastern Australian species. 
Ascidia aclara Kott; Sryeht pedata (Herd- 
man ) i Pyura pachydermatina (Herdman) 
nihhosa (Heller); Pynra spinileta I Quoy ami 
Gaimard ) ; Microcosmus nicholhi Kott 
(absent only from tropical Australia). 

I he apparently endemic species comprise a 
considerable comporem (3l'# ) of the fauna 
in the northern part of ihc Great Australian 
Bight. The ciTcum-Auslialian forms comprise 
almost 5() f 7. of ihc species, while species with 
a range to Port Jackson {Pyura p. #ihhoia* P. 
Aptnifera) or Morvton Bav (Ascidia adam. 
Styeitt pedidtt. Microcosmus nicholUi) also 
occur, the three latter records extend the 
known range lo the west, although the first 
two species are already known to occur in 
southwestern Australia. The data thnt are 



recorded here do nut therefore disagree with 
previous information (Kott 1972b) Bftl Wlffc 
ports Ihe existence ut* a marine fauna! Fioviuce 
extending from Cockburn. Sound (or further 
to the north) un the -western Australian eoa.st 
to the vicinity of the eastern coast of South 
Australia- 

There is no evidence that would suggest lluil 
the sample thai is available is not typical ttf 
the fauna of the Great Australian Bight, This 
fauna, however, dom not, on the basis ol avail 
able data, appear to be Typical of the Flln- 
dersian marine launal Province. Apart from 
the large endemic component, the species oc- 
curring there have n wide distribution around 
the Australian coast, especially along the east- 
ern seaboard. The ipectes that terminate then* 
lange at the eastern end of the Flindeisian 
Province have not been taken in these collec- 
tions from Ihe northern part or' I he Great Aus- 
tralian Bight CKott 1972b), although they have 
been recorded at more easterly locations off 
South Austialia. 

Other species in this colled ion taken from 
other locations of! the South Australian coast 
may be grouped according la ihe Soulh Aus- 
tralian limits o\ their range in the following 
way: 

J. Species ifmr do not extend eastwards into 
the Muageati marine Provi/tce. 
Podociavelta. cylindrica I Quoy & Gaimard); 
Pycrtoclavella dimittuta ( Kott ) ; Atapozoa 
tnarvhit Brcwin; Diplosonm trausiuetdiuv 
Hai'tmcycr). Polyclinum neptunium (Hait 
meycr; Stafonhv carrwsa Millar; Afatguh 
salntlusa Kott. 

2. Species that do not extend westwards jnto 
the Flittdetsian murine Province. 
Distaplia australicnsis Brcwin; Eidterdmania 
australis Kott; Aplidium coni/eruoi Kott; 
ApUd'tutn amorphatum Kott; Ascidia thowp- 
soni Kott; Polyandrocarpa iapidosa (Heid- 
man): Pyuta irregularis (Herdman): Poly- 
carpa tinctor (Quoy & Gaimard); Polycarpa 
papilhtta (Sluiter). 

3. Species jor which the f 1 in der start/ M andean 
boundary does not comprise a harrier, 
Sycozoa cerehriformis {Quoy & Ciairnard): 
Sytozoa ptduHculuta (Quoy & Gaimard): 
fiitterella herdmania Kott ; 1 rididemnutv 
stivignii f Herdman): 1 rididctnnutn cerebri- 
forme (Hartmeyer) ; Symple^ma virtde 
Herdman; Pyura australis (Quoy & 
Gaimard); Microcosmus smiomjera KolL 

4. Ccdf faiota. 

PvtOa scoreshiensis Kotl. 



IK 



PATRICIA KOTT 



References 



Aiibott, D. P & Johnson, J. V_ (1972). — The 
Asvidians Styela harnharti. S. pircata, S. ctava, 
and S. monterevensis in Californian Waters. 
But! So, CrMjnhiitt Acad. S$l 71 {Z) y 95-105. 

BRF.WIN, Bl.RVI I. U952).— Ascidians of New- 
Zealand, Pari VII, Ascidians from Olago 
coastal waters. Part Jl. Trans, R. Soc. N.Z, 
TlM^i). 452-458. 

BnPWiN, ftp.RVf. T. <J9S?) — Australian ttseidians 
of the sub-family Holo^inac and a review 
Df the Mib-familv. Trans, H. Soc, N.Z, 8i( I ), 
53-64. 

BreWuv, hFRVL I. ((95ft). — Ampozoa nuirshl, a 
compound uscidian from Wexlern Australia. 
J, R. Soc. M'',.4. 40(1), 31-32. 

Bur win, Bi-kyi \, (1957). — Ascidians of New 
Zealand. Part X. Ascidians from North 
Auckland. Trans. R. Soc N.Z, 84(3), 577- 
580. 

Bkiiwin, Bkkyl 1. (1958a). — Ascidiuns of New 
Zealand, Part XI. Ascidfans of the Stewart 
bland region. Trans. R, Soc. N.Z. 85(3), 
439-453. 

BBEW1N. BtRYl. T. (1958b). -Aseidians ul New 
Zealand. Part XII, Ascidians of Ihc Hauraki 
Gulf Pt. III. Trans, R, SoC. N.Z. 85, 45^-45*. 

r.uwhoOK, L. O. (1967). — A taxonomic review of 
Indo-Pacifie didemnid ascidians and descrip- 
tions of twenty-three central Pacific Specie*. 
Mieronena I 1 61-261. 

Glynn. P. (1974). — Rolling sttsnes among ihe 
Scleractinia Prqc. Second lale*nationol Coral 
Reef Stkiposfom. Vol. II. 183-198. (Great 
Barrier Reef Commiliee: Brisbane). 

Hart\u;yf.r. R. *IV19). -Ascidfevi. Results of Dr. 
K. Mjoberg's Swedish scientific expeditions to 
Australia 1910-13, K, wrnska Veim^k-Akatl- 
ifandl. 60(4), 1-150. 

Hastings, Anna B. { 1931 ).— "I nnicnta, Sclent* 
Rep. Gt.. Barrier Reef Expcd. 4(3), 69-109. 

IIillCR, C (I67K). — Beitriige zur naheren 
Kenolnts der Tunicate i. S/nr. A Add. WLys. 
IVien. 77(1). 83-110, 

Kan, Patricia (1952)— The ascidians of Aus- 
iralia. I. Stolidobranchiatn and Phlebo- 
hriuichiafii 1ltst. A mar, Frcshh; Rv$. 3(3), 
206-333. 

Kott, Patricia (1957). — The ascidians of Aus- 
1 nilf;i . H AplouNohranchiala I ahille; Clave- 
Imidac H'orbes and Hanly and Polyclinidac 
Vorrill, Aust. J, mat, rfityrV 1 ' Rc\, X( I t. 
64-110. 

Kott, Patricia (1962)- -The ;e.adians of Au>- 
1 ralin- |Tfi AploiiSiihriiDchiaia 1 ahille: Didcnv 
hidac Oiard, Aast. J. mar. treshw. Res. 
13(31, 265-334 

Koit, P.vnuuA. (1963). — The asvidiaus or Aob- 
iralia. IV, Aplousobranchiata LariiUe; Poly- 
clnudae Vern'll (continued ). Anst. J. ntar. 
Freshw. Res. 14(1), 70-118. 

Kott, PAiRtrtA (1964). — StoliJohianch and phle- 
bobianeh ascidians of the Queensland coast. 
Pap, Dep. loot. Univ. Qd 2(7), 127-152. 

Knrr, Patricia (1966) — Ascidians. of tiotlh Aus- 
tralia. Pap. i)ep. loot. Univ. Qd 2fl5). 
279-304., 



Koru, PAiKiCiA 196K).--A review of ihe genu-; 
Hoiocv>ifhia Veridl. I«79, froc. I.ittu. Sec. 
A',\ ) |V, V3r 1 1, 7fV89. 

Ken, pAffticiA (JMA9). — AriUrclie Ascidiacea. A 
monogiaphic uccnuni uf the known species 
based on upecinicns colleclcd under US 
Governmcnl auspices 1947 to 1963. Antarcl. 
ftef. Sfir. 13, i*XV, t-239. 

Kon t Pathicia (J972a> — The 3*eManS of Sooth 
Anslralta I: Spencer Guir. Hi Vincent Gulf, 
atiJ Encounter Bav. IrvjU*. /?, 5'V. S. -las) 
9rt(l), 1-52 

Kott. Patricia (1972b).— The ascidians of South 
Australia II. Eastern ncctor of ihe Great 
Australian Bieht and Investigator Stmir. 
7Yew.v. R. Soc. S. Attsl. 96, 166 196. 

Kott, Patricia (1972c). — Notes on *omc sisci- 
dians from Port Jackson. Botany Bay. and 
Port Hacking, New South Wales. Proc. Lh*n. 
Soc, N.S.W. 97, 241-257. 

Koti. Patricia ( I972J) — Some suhlatoral wsx-t- 
dians in Moreton Bay, Queensland. Mem. Qd 
Mas. 16(2). 233-260. 

Kott. Patricia (1972e).— -Paiina of the Gulf of 
Carpentaria; 2. Asctdiacca (Chordala: Tuni- 
c-atal. W^fA. Notes t\\S. 2(3), 39-57- 

KQTT, Patricia (in piess). — The ascidian foutni 
of Wcslcrnport Bay. Victoria^ and a com- 
parison with that 61 Porl Phillip Bay. Mem. 
Natn. Mas. Vict. 

MroTuri.sTN, W. (1922). Ascidiae Ptycho- 
Sranchiae und Diktyohranchiae von Neusee- 
land und den Chatham-lnseln. (Papers from 
Dr. Tlr Morelensen's Pacific Fxnedilion 
1914-lft. No, XI). Vtdfftk, McJd>, ttmsk 
ttartirh, t'orfti, 73, 359-498. 

Mu'.iiAr.LS.T.N. W, (1924). Ascidiac Kriko 
hranchiae von Nenseelond, den (_'hatham- hm\ 
den Auckland-lnsclu. (Papers from Or. *l'h. 
Mortemcns Pacific Expedition 1914-16. No, 
XXII >. Viiicask. Mctfttr, tlaask. natuth. f'oren. 
77, 263-434. 

MltHAMSrN. W,, & HAHIMlVfcR. R. (I92K|.~ 

Asdditic diktyobianchiae u.*d ptycho- 
branchiae, Fauna SnJncsf-Audl. S, 251-4^0. 

MiLiAR. K. H. f i960). — Asciduceu. Phcon'r* 
Rep, 30. 1-60. 

Millar. R. H. (1961). — ^Avcidians from Mozam 
hi<|ue. Ann. M#x. Nat. Hht. t*3)4. 11-16 

NfiLLAR. K. II. 11962) — FuTther description* ol 
South African aockhan^ Ana, V, A jr. Mas. 
46(7), 113-221. 

Miiiar, K H. (I9f»^),— Australian ascidians In 
Ihe British Museum (Naliual History I. Pew. 
Zttol Soc. Land. 141 1 4). 6X9-746- 

Miliar, Is*. H. (1966).— Ascidlamt P<»rt Phillip 
Survey. Mem, natn. Mus, I'ict. 27, 357-375. 

Nott. J. T, (1892). — On the composire asciditins 
of the North Shore Reef. Trans. N.Z. fust. 24. 
305-334. 

Prhon, M (1974). —Free living sclcractinian 
coral comnuinilies in the coial iccfs of Inlear 
''Madagascar). Ptr.v. SvcotaJ JitTernniionai 
Loral Herf Symposium. Vol It. 173-182 
iGre.jt Barrier Reef Commilteo: Brisbane. 1 



ASCIDIANS OF SOUTH AUSTRALIA III 



iy 



Pi.ante, R-. & VA&snm, f\ (1966),— Sur unc col- 
lection d'ascidies de la region de Tulcar (cote 
sud-ouest de Madagascar). Ann. I'Univ. 
Madagascar. Scric Sciences de la Nature et 
Mathematiques, no. 4. (Tanarivc: Imprimcrie 
Nalionale. ) 

Qtjoy, J., & Gaimard, P, (1834). — Voyages de 
decouvertcs de (Astrolabe 1826-29. MoL 
lusqucs. Zoologie 3, 559-626; 4 7 304-306. 

Slliter. C. P. (1909).— Die Tunicaten der Siboga 
Expedition Pt. 2. Die Merosoraen Ascidten. 
Siboga Exped. 56B, 1-112, 

Sluiter. C. P. (1913). — Ascidien von den Am- 
Inseln. Abh. sr.nckenh, nulurforsch. Ges. 35, 
65-78. 



Tokioka, T. ( 1949) . — Contributions to the 
Japanese ascidian Fauna II. Notes on some 
ascidtans collected chiefly along the coast 
of Kii Peninsula. Pubis Seto mar. biol. Lab. 
1(2), 39-64. 

Tokiora, T. (1952 )— Aseidians collected by 
Messrs, Renzi Wada and Seizi Wade from 
the pearl oyster beds in the Arafura Sea in 
1940. Pubis Seto mar. bwl. Lab. 2(2), 91-142. 

Tokioka, T. (1967), — Pacific Tunicata of the 
United States National Museum. Bull. US* 
nam. Mus. 251. 1-242. 

VASSEim, P, ( 1967 ) . — Ascidies de Nouvelle- 
Calcdonie. Edition de la Fondation Singer- 
Polignae. 127-146, 2 pis. 



Appendix — Station List 



NORTHERN GREAT AUSTRALIAN BIGHT 

(32°24'S. J33 : '30'E). May 1973. Experimental 
Prown Trawl. Explorer (Coll. P. Symond). 
42 m; Poly ci tor gigajiwum 

Patr'tdium puhinatum n. gen. n. *p 

Aplidium cole I hides 

Aplidium foliorum n. sp. 

Leptoclinides votvus n. sp. 

He rt I mania momus 
49 m; Aplidium colelloides 

A plidium flavoltneatum 

Aplidium digitutum n. sp. 

Lcptoclinidex reticulums 

Polysynvraton nspiculuium 

Didemnum candidum 

Didemnum moseleyt 

Amphlcarpa diptycha 

Polycarpa pedunculata 

Sty flu pedutti 

Cnemidocarpa etheridgii 

Pyitra pachydcrmntina gibbosa 

Pyura spinifera 

Tlerdmania mom Us 

Micrr/citsmus nichollyi 

GOAT ISLAND, off Ccduna, Great Australian 
Bight (Coll. P. Howlett). 
32 m; Ascidia acjara 

Pyura tendafa 

Mkrocosmus planus m sp, 

CLLISTON BAY. Feb. 1971 (Coll. S. A. 

Shepherd). 

Outside bar: 

Fulierdmania australis 

Ritterelln berdmunia 

Aplidium ttmorphnlum 

Leptoclinides rttfus 

Oculinaria australis 
Reef; Sytnplegrna viridc 

Aplidkurt caniJi'MM 
Vertical faces (25 m) : Pyura pachydermatina 

Molgula sabulosa 

WFST ISLAND; Amphitheatre RocV 

7 m deep, 13.vii.1972; 

Pplycitor gtganteum 
17 m deep", 12,vh\1972: 

Sycozoa cerehriformis 

CAPE JAFFA; Margaret Brock Reef (3-4 m deep 
and in caves). 28.xi.1972. 

Pseudodistoma cereum 



January, 1971 (Coll. 



INVESTIGATOR STRAIT, 
J. E, Watson), 

Station XI (depth 19 m); 

AplidiUm pronum n. sp. 

Diplosoma translucidurt\ 

Pyura australis 
Station X3; 

Pyura australis 
Station X7: 

Hwdmaniti momus 

Stations X8, X9, XI 0: 

Pxura scoresbiensis 

Station Yl: 

?Polyclinum neptunium 

Stolanicfi carnosn 

Polyundrocarpa lapidosa 

Molgula moth's 
Station Y5: 

Atapozoa marsh't 

Euherdmania australi'i 

?Pulysyncruton usphulatutn 

Pyura australis 
Station 76: 

Atapozoa marshi 

UPPER SPENCER GULF, September. 1973. 
Transects and stations of S. A. Shepherd, Depart- 
ment of Fisheries, S. Aust. 

Transects A-D: 

Polycarpa pedunculata (arenaceous and 
naked specimens, sometimes growing 
side by side, attached to the same .shell 
or stone, stalked or sessile). 

Station A1 on Pinna, depth 0-1 m: 
Pyura irregularis 
Pyura vittata 

Station A5, depth 17 m: 
Ascidia tlwmpsont 
Polycarpa pedunculata 

Station A7. depth 10m- 

Polycarpa pedunculata (2 spec.) 

Station B4. depth 17 m: 

Sycozoa pedunculata 

Station B7. depth 5 m: 

Polycarpa pedunculata (some naked, 
black; a few leathery) 
Polycarpa papillata (single specimen) 
Pyura irregularis (one small aggregate) 



20 



PATRICIA KOTT 



Station BH), depth 10 m, channel: 

Polycarpa iinctor 

Molguhi mollis 
Station C4, depth 12 m: 

Polyrarpa pedunculaia (naked and arena- 
ceous) 

Station D3, depth 18 m: 

Ascidia thompsoul 
•Station D5, depth 15 m; 

Disfaplia austraiiensis 
Station D9, depth 10 m; 

Pyura scoreshiensis 
Station Eh depth 7 m: 

Polycarpa peduucalata 

Microcosmus mchoflsi 

Microcosmus stolon if era 
Station E3. depth 9 m: 

Ascidia tbompsoui 
Station E4. depth 5m: 

Pyura irregularis 
Station FL depth 19m; 

Polycarpa pedunculata (small stalked. 

arenaceous) 
Station F3, depth 19 m: 

Pyuta scoresbiepsis {without stalk) 
Station F4. depth 16 m: 

Ascidia thompsoni (with barnacles) 
Station G, depth 9 in: 

Ascidia thompsoni 



Parabofrylins nemorus n. gen., ti. sp. 
Polycarpa peduuculata 
Pyura scoresbiensis 
fialocytithia htspidu 
Microcosmus squamiger 
Hcrdmania momus 

Reef 4 km NNW Douglas Bank: 

Podoclavclla cyl'mdrica 
Pyura aaMralis 

SPENCER GULF. 

Tipara Reef, depth 11 m, 24jx.197J: 

Pycnoclavella dimiuitta 

Pyura irregularis 
Under stones; 

Hcrdmania momus 
Depth 5 m, 2/V.1972: 

Podoclavclla cylirtdriva 

ST VINCENT GULF. 

Port River (near Ele.tricity Trust), depth 3 m T 
muddy bottom, 9.vi.l972: 

.Slycia plicatii 
Outer Harbour: 

Pyui'n stolonifera 

Microcosm us sqttam fgfiir 
Selliek Beach (S of Adelaide), Feb. 1972 (Coll. 
K. Hammond) : 

Trididemnum savignii 

Indidemnum ccrebriUtrtnc 



Index to Genera and Species 



Amphicarpa diptycha 
Aplidtum amorphatun) 
Aplidium colclloides - 
Aplidium eoniferum - 
AplkUum digitatum - 
Aplidium llavolincatum 
Aplidium foliorum 
Aplidium pronum 
Ascidia aclara - 
A.scidia thompsoni 
Atapozoa marshi 
Cnemidocarpa etheridgi 
Didemnum enndidum 
Didcmtium moseleyi - 
Diplosoma translucidum 
Distaplia ansfrnliensis 
Euherdmania australis 
Halocynthta hispida - 
Herdmaiiia mom us 
Leptoclinides reticulatus 
Leptoclinides rufns - 
Leptoclinides volvus - 
Microcosmus nichollsi 
Microcosmus planus - 
Microcosmus squamiger 
Microcosmus stolonifera 
Molgulu mollis 
Molgula sabulosa 
Oculinaria australis - 



II 
6 
7 
6 

7 
5 
5 
6 
11 
10 

7 

14 
9 

9 

10 

1 

j 

15 

IS 

8 

9 
8 

16 
15 
16 
16 

16 
16 
11 



Parabofrylins ncmortis 
Patridium pulvinatum 
Podocbvella cylindrica 
Polyandrocarpa lapidosa ■ 
Polycarpa papillala - 
Polycarpa pedunculate 
Polycarpa tinctor 
Polycitor gigantetim - 
Polyclinum neptunium 
Polysyncraton aspiculatum 
Pscudodistoma ccrcum 
Pycnoclavella diminuta 
Pyura australis 
Pyura irregularis 
Pyura pachydermatina 
Pyura scoresbiensis 
Pyura spinifera 
Pyuta stolonifera 
Pyura tendata - 
Pyura vittata 
Rittcrella herdmania 
Stolonica carnosa 
Styela pedata 
Styela plicata 
Sycozoa cerebriformls 
Syco/oa pedunculata 
Symplegma virt'dc 
Trididemnum savignii 
Trididemnum cerebriforme 



II 
4 

I 
19 

13 
13 

13 

2 

4 

7 

4 

20 

14 

15 

14 

T4 

14 

15 

14 
15 

U 
I* 
13 

19 

iT 

to 



NOTES ON THE GENUS PSEUDOMALAXIS FISCHER 
(MOLLUSCA: GASTROPODA) AND ITS FOSSIL SPECIES IN AUSTRALIA 

byM. F. Buonaiuto* 



Summary 

BUONAIUTO, M. F. (1975). -Notes on the genus Pseudomalaxis Fischer (Mollusca: Gastropoda) 

and its fossil species in Australia. Trans. R. Soc. S. Aust 99(1), 21-29, 28 February, 1975. 

Two fossil species of Pseudomalaxis Fischer are discussed; P. asculpturatus Maxwell 

(Late Eocene) and P. praemeridionalis (Chapman) (Early Middle Miocene). The former is a new 

discovery in Australia and is one of a few forms common to both Australia and New Zealand; the 

latter is poorly known and is redescribed herein. The taxonomic position of Pseudomalaxis Fischer 

is reviewed and the genus is restored to the Architectonicidae. 

The possible synonymous or subgeneric relationship of Mangonuia Mestayer to Pseudomalaxis 

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

are considered. 



NOTES ON THE GENUS PSEUDOMALAXIS FISCHER 
(MOLLUSCA: GASTROPODA) AND ITS FOSSIL SPECIES IN AUSTRALIA 

by M. F. Buonaiuto* 

Summary 

Buonaiuto, M. F. (1975). — Notes on the genus Pseudomalaxis Fischer (Mollusca: Gastro- 
poda) and its fossil species in Australia. Trans. R. Soc. S. Aust. 99(1), 21-29, 28 Feb- 
ruary, 1975. 

Two fossil species of Pseudomalaxis Fischer are discussed: P. ascidpturatus Maxwell (Late 
Eocene) and P. praemeridionalis (Chapman) (Early Middle Miocene). The former is a new 
discovery in Australia and is one of a few forms common to both Australia and New Zealand; 
the latter is poorly known and is redescribed herein. The taxonomic position of Pseudomalaxis 
Fischer is reviewed and the genus is restored to the Architectonicidae. 

The possible synonymous or subgeneric relationship of Mangonuia Mestayer to Pseudo- 
malaxis Fischer, of Calodisculus Rehder to Awarua Mestayer, and of Claraxis Iredale to 
Torinista Tredale, are considered. 



Introduction 

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 Bijrontia 
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 Otnalaxis 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, Bijrontia Deshayes, and Pseudo- 
malaxis Fischer; he placed Bijrontia 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 Euomphalinac, 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. 



22 



M. F. BUONAIUTO 




NOifcS ON eSUJtHhUAl 4X1S FISCHER IN AUSTRALIA 



23 



or [U leant a subgenus of 1 PseitdoffteltixLs 
Fischer, 1885". Mcstaycr (1930, pp. l44-5> 
refers P. meridianalis (Hedley) to Manftpntda* 
In fact, the recent Mangontfin h**llon.*j Mefr 
layer, the type of this genus, displays the 
general pattern of spiral ornaments, the quad- 
rangular outer and the subcircular inner shape 
o( body whorl characteristic of the Neogene- 
Reeent Pscudomalaxts a, str (l described below, 
and the .same type of pscudoplanispiral coiling 
ILlf this group, 

Render further comments. "Calodhados 
Render, 1935, is very close to Awanta Mes- 
taycr, 1930, and is probably its synonym". 
Wenz f 1939. p, (567) considers Awanta a sub- 
genus of Mangonuia. However, Calodhcuh** 
and Awanta display the same kind of speciali- 
zation in spiral ornaments, in particular a 
marked development of heavily bended circum- 
umhilical cords, Awanta ttmoena (Murdoch &. 
Suter) (Sitter 1913, p. 318; 1915, p] 15. jig, 
21 ah) displays also a, torinioid operculum 
iMcstayei 1930. p. 146). 

"Claraxix and Torinistti Tredale are very simi- 
lar, possibly synonymous, and might Ik sepa- 
rated as a distinct genu?; whicb may be closer 
to Hchucus d'Orbiyny". Wenz (1939. pp. 666. 
bhH) considers the former two as subcenem of 
Mungonuia and the latter as a synonym of 
Tnritt'ttt Gray. In this ca«e, Cbitttxis should fall 
into synonymy with lorttuxta because although 
Iredalc (1A3& p. 327) published their initial 
dingrio;se-s on the same page, ihe former is des- 
cribed after the latter. From the original des- 
cription and drawings of the type species, it is 
impossible to find substantial generic differ- 
ences and Iredale did not specify any. 
(h) The genus PseudotmdaXis Fischer is often 
placed in different families hut generally is 
referred to the Architcctonieidae. In introduc- 
ing the genus, Fischer referred it to the Sotor- 
ridac f — Architcctonieidae) on the basis of the 
conical torinoid operculum (Fischer 1885, p. 
714. see also Earncs 1952, p. 37), observed in 
the Recent form P. (Spirolaxis) centrifuge 
Montcrosato fMonterosato 1890) and in P. 



nobitts (Verxill) (Vcrrill IXR5, p. 421, pi 44. 
fig. (2). 

Ircdalc (1936, p. 326) instituted a new 
family Mungonuiidae lor the Australasian 
genera MctrtgonHiti, Awthna, Tnn'tmta and 
Chwaxis, Apparently, Wenz (|939| considered 
Mangonuiidac synonymous with the Architcc- 
tonieidae. Because of a certain affinity heiween 
the opercula of P. balesi Pilshry & McGinty 
and Parvitarbo zacallcx (Motycl;), Pilsbry & 
McGinty (1945, pp. 9, 57. pi. 6. figs 2-2a. 5) 
placed Pseadomafaxis in the Cyclostrcmatidae. 
Latei, Abhott (1954, p. 138) included Pxeudo- 
rtialaxit in Vitrincllidae on the basis of <* vague 
and incorrect reference to a revision of the 
family by Pilsbry. Maxwell ( IV66, p. 4441 fol- 
lowed Abbnii. 

Rehdei (1974, pers. comm.) gave the 
opinion: '1 have examined young specimens of 
both Pscudo.vtu'aAtx and A rcliiieetoniea (Psl~ 
laxh) and I can find no essential difference in 
their protoeonehs: both show a heterostrophic- 
anastrophic protoconch. 1 feel, therefore, that 
Psfmdomalaxix should remain in the Archilec- 
tonicidac". There is support for this statement 
boih on the basis Of the torinioid conical oper- 
culum and the protoconcli coiling of Pseudo- 
waiaxis. Therefore the author* in agreemetii 
with Render, regards the restoration of Pxeudo*- 
malaxi* Fischer to the Architectonicidue to he 
justified. 

Collections 

The specimens of P. astulpturutus Maxwell 
here studied arc kept in the Pal aeon tologicai 
Seciiun of the New Zcatarul Geological Survey 
(NZGSi, and in the Department of Mines of 
South Australia (GSSA). The figured specimen 
of P. praemeridiomilis (Chapman) is kept in 
the Palaenntologicnl collection of the South 
Australian Museum of Natural Hisiorv 
fSAM). 

Spccuncas M329S and M3299 were found 
by the author and specimens M3300 M3303 
were found subsequently by J. M. Lindsay. 



h»g. 1. Pwtuhrfmla.\fx (PsvudomuhtMS) asvalplanilus Maxwell. MeCullouah's Btiuee, New Zealand. 
NZGS. 9508 2; a, ad apical; b, abapical; c. axial views (all X 17.50). 

He. +JP. (PswHkmuitaxis) ascnlpiurattts Maxwell. McCullou^lt's Bridge. New Zealand. NZCS, 9508-1: 

iL, adapicul: b, abapical; c, axial view* (all X 17.3). 

Ffe 3. P {Pwutlontahais ) asadpturatus Maxwell. Adelaide Plains Sub-Basin, Adelaide Children* 
Hospital Bure No. 5; GSSA, M3299; a, ad apical <\ 161; n. abapical IX lfil; p, axial views 
IX 15) 

l'l£. 4. P. (Pjicudo/nataxis) pwmcridiojudis Chapman Clifton Bank. Muddy Cre^k; SAM. PtH^i: 
t t t adapicnl tX 106); h t abapical 1X10,6); < axial views (X 12.3) 



24 



M. F. BUONAIUTO 



Synopsis oj the history of tlie genus Pseudomalaxis Fischer on the basis oj the major authors, 



Author 
Fischer (1885. p r 

714) 
Sacco (1892, p. 75) 
DaJl (1892, p, 331) 

Iredale (1911. 
pp. 253-7 ) 

Monterosalo 
(1913, pp. 362-3) 

Cossmann (1915, 

pp. 122, 14L) 
Mestaver (1930, 

p. 144) 



Rehder (1935, p. 
128) 



Tredale (1936. 

p. 326) 
Wen/ (1939. p. 

668) 



PiKbrv & IVfcGintv 
(1945. p. 9.) 

Fames (1952, 

p. 37) 
Ahbolt ( 1954, 

pp. 138-9) 

Pchelintsev& 

Korobkov (I960, 

pp. 137-8) 
Maxwell (1966. 

p. 444) 
Gilbert (1973, 

p. 30) 
This paper 



Family 
Solariidae 

Solariidae 

Solariidae 

Solariidae (?) 



Solariidae 



Euomphalidae 

Arcbitcctonicidae 
(syn. Solariidae) 



Architcclonicidae 
(?) 



Mangonuiidae 

Arcbitectonicidae 
(syns Solariidae, 
Mangonuiidae) 



Cyelostrematidac 
Architcctonicidae 
Vitrinellidae 
Solariidae 

Viirinellidac 

Architcctonicidae 

Arcbitectonicidae 



Subfamily Genus 

— Torinia Gray 

Discohelix Dunkcr 
Discohelix 

Pseudomalaxis 
Discohelix 

— Pseudomalaxis 



Euomphalinae Pseudomalaxis 

Mangonuia 

Mestayer 

Awarua 

Mestayer 

Pseitdomufasis 



Mangonuia 
A wurtui 
Manexmuia 

Pseudomalaxis 



Subgenus 
Pseudomahtxis 

Fischer 
Pseudomalaxis 
Pseudomalaxis 
Discosolis 'OaU 
Pseudomalaxis 
Discohelix 
Discosolis 

Pseudomalaxis 

Spirolaxis 

Monterosato 

Pseudomalaxis 



Pseudomahixis 

(syn. Discosolis 
Dall) 

Paurodiscus Rehder 
Calodiscttltis Rehder 



Systematic Descriptions 
Phylum MOLLUSCA 

ClaSS GASTROPODA 

Order Mp.sogastropoda 

Superfamily cerithiacea 

Family ARCHITECTON1CIDAE 

Genus PSEUDOMALAXTS Fischer, 1SK5 

Diagnosis; Shell discoidal, pscudoplanispiral; 
nbapical side concave to suhconeave, adapical 



Mangonuia 
A wdrua 

Pseudomalaxis 
Spirolaxis 
Paurodiscus 
Calodisculus 

Pseudomalaxis Pseudomalaxis 

Pseudomalaxis Pseudomalaxis 

Pseudomalaxis Pseudomalaxis 

Pseudomalaxis Pseudomalaxis 

Pseudomalaxis Pseudomalaxis 

Pieudomahnis Pseudotmdaxis 

Pwudtumduxis Pseudomalaxis 

(?«yn. Mangonuia) 

lAlangomtia 

Spirola.xis 

Paurodiscus 

A warua 

( ?sy n . Calodisculus ) 

'^Calodisculus 

flattened to subconcave; outer shape of the 
body-whorl rectangular to quadrangular, inner 
shape subcircular to similar to the outer; thin 
margins, two carinae at abaxial-adapical and 
abaxial-abapical ends. Straight growth lines 
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). 



NOTES ON PShUDOMALAXlS FISCHER IN AUSTRALIA 



25 



Subgenus Pseudomalaxis *. itr. 

Diagnosis. Shell medium-large to very small; 
margins thin: pseudoplanispiral with tangent 
whorls: suture Hush to subimbricated; two ada- 
pical and two abapical keels: the four keels 
can be smooth or crenulatcd; growth lines a 
little irregularly packed; operculum flat (after 
Wcnz 1939). * 

Observations: Pseud omul axis s. str. appears lo 
bo represented in the Tertiary by two main 
forms. The first predominant in the Eocene- 
OHgocene, is commonly characterized by 
growth lines and the four carinae as the only 
ornaments; a few species bear also spiral orna- 
ments of fine riblets as in the Eocene P. dixoni 
(Vasscur) and the Oligoeene P. italica (Saceo). 
or of beaded cords and keels as in P. texana 
(Aldrieh); the outer shape of the body whorl 
is rectangular to quadrangular, the inner is sub- 
circular to subquadrangular. the inner is sub- 
circular to subquadrangular or subrectangular. 
depending on the thickness of the margins ((he 
latter characteristic could be ontogenetic and 
related to age) 

The second form, predominant in the Neo- 
gene-Recent, is characterized by the develop- 
ment of a main spiral rib or cord on ihe abaxial 
region of the adapical margin, and often of 
spiral costellae on the abaxial margin. The four 
carinae and the adapical rib usually bear beads 
or short spines. The outer shape of the body 



whorl is commonly quadrangular to subquad- 
rangular, Ihe inner shape is swbcircular. 

Pscudomalaxis (Pseudomalaxis) a*cul|>luralits 

MaNwelt. 1966 

FIGS 1-3 

J9n6 Psendomtduxis metdpttiratus Maxwell, 
p. 444, figs 11-13, 

Material: 4 specimens very well preserved with 
(he peristome slight! v damaged (GSSA, 
M 3298-9; NZGS, 9508-1, 9508-2) and an- 
other 4 juvenile or damaged (GSSA. M3300-3). 
Description: Shell very small \riL\ ihin, bicon- 
cave, nearly planispiral; protoconch hctcrO- 
strophic-unastrophic; whorls increasing far 
more in diameter than in height and overlap- 
ping entirely in relation to the coiling axis and 
very scarcely in relation to the normal to it- 
Suture flush to subimbricated. Body-whorl 
shape: outer subquadrate-subtrapezoidal; inner 
subcircular-ovoidal, wider than high. Whorl 
regions : adapical and abapical subconvex ; 
abaxial flattened or subconcave. nearly vertical. 
Body-whorl regions connected by prominent 
smooth abaxial carinae. Abapical side more 
concave than the adapical one. 
Ornaments: Growth lines and rugae, prosocline 
in the adapical and abaxial regions, opistho- 
cyrt in the abapical. Four carinae: at abaxial 
adapical, abaxial-nbapical, adaxial-adaptcal. 
and adaxial-abapical ends. 



DIMENSIONS (in mm): 
Specimen No, whorls 


(see Fig. 
Dw 


5) 

N 


Nis 


Hw 


Lis 


Hgc Dcg 


M329S 4 


2.55 


0.95 


0.40 


0.50 


—0.10 


0.60 0.65 


M3299 4 


2.50 


0.92 


0.39 


0.55 


—0,10 


0.65 0.60 


NZGS 3 

950S-1 


L55 


0J0 


0.30 


0.40 


—0.05 


0.45 0.50 


N2GS 3 

9508-2 


2.15 


0.90 


0.35 


0.5S 


0.00 


0.58 0.50 


RATIOS: (see Fig. 51 
Specimen 


HwDw 




K - I is/Hw 


9 Nis/N 


Hgc/Dgc 


M3298 


0.1961 




—0.2000 




0.421 1 


0.9231 


M3299 


0.2200 




—0.1818 




0.4239 


1.0833 


NZGS 9 SOS -1 


0.25*1 




—0.1250 




0.42 Bfi 


0.9000 


NZGS 9508-2 


0.2698 




0.0000 




0.3889 


1.1600 



Localities: Adelaide Plains Sub-Basin (St Vin- 
cent Basin, S. Aust); Adelaide Children's Hos- 
pital, North Adelaide, hundred of Yalala, 
"I own Acre 717, Bore 5, at 20.42-20.75 m 
IM329S) and 21.64-2 1..95 m (M3299); Bore 
2, at 22.25-22,56 m (M3302, M3303); Ade- 
laide Metropolitan Subway, Bore 3, north bank 



of Torrcns Lake, opposite Kintorc Avenue, at 
20.10 m (M3300) and 24.50 m (M3301) 
New Zealand: McCullOuglrs Bridge. 
Sfraiigraphic Distribution: Adelaide Plains Sub- 
Basin : Lindsay ( 1 966 ) gave a brief strati- 
graphic summary of Adelaide Children's Hos- 
pital Bores 2 and 5. "The interval 20.42-21.95 



lb 



M. F. BUONA1UTO 



rn in Bore 5 is low in Blanche Point Marls 
( Aldingan, Late Eocene). The Han/kt-nina 
primitiva zone occurs in this bore at .19.20- 
19.51 m, i.e. slightly above P. usculpturants. In 
the type section at Maslin Bay, Hamkenina 
ptim'ttiva is present only in a restricted band 
0,80-1.15 m above the base of Blanche Point 
Transitional Marls. 

Other specimens from Adelaide Children 7 s 
Hospital, Bore 2, are within the Hantkenina 
primitiva zone. Although H . primitiva has not 
yet been found in Adelaide Metropolitan Sub- 
way Bore 3, the specimens of P. a sculptural its 



are certainly from near, and probably from 
slightly above, the //. primitiva zone. 

Thus the vertical distribution of all the spe- 
cimens of P. asculptttratus found in the Ade- 
laide area spans a narrow interval from slightly 
below to probably slightly above the H. pr'mti- 
tiva zone, in Blanche Point Transitional Marts, 
Late Eocene/' (J. M. Lindsay, Dept of Mines, 
pers. comm., 1974.) 

New Zealand: Upper Waihao Greensand, 
Kaiatan. Late Eocene. 

Ohsnvfitionv: The two Australian specimens 
show an inversion in the coiling, variable in 



i ) f ) feq Hw Htr^. 



\V^ '-O.jCfl 

I — N'S — I 

I r N _ — . ^ 

„ Dsv 1 



r : Li 

Hw < Hgc JV 

lis <0 




) !2& 



Hgc. n HW 



T < 
Hw - H 3 c r 
Us = Q 



Fig. 5. 'the paiumcteis measured in piaiiispirai 
(a) and in pseudoplanispiral (b) 
gastropod shell arc here defined an 
follows: 

No iv ; the total number of the whorls, 
aw the total number of revolutions 
of the generating curve around the coil- 
ing axis. 

Hw; height of any whorl (in thus cas* 
the last one) as the projection of lh? 
foiling axis of the distance bcrween the 
adapical point of the .generating curve 
at the initial position and ils abapieal 
point at the final position after a 2t 
revolution in the given interval In*" — 
2<n-M)*\ 
L\S\ inlersulural heiyhl of any whorl us the projection on the coiling axis of the distance 
between the imlial and final position of a given point placed on the adapical line of the body 
whorl ;ifter a 2^r revolution* where the adapical tine V s the geometrical locus of any point 
placed at the adapical end of the generating curve. 

//gr: the height of the generating carve as projection on the coiling axis of the distance 
between its adapical and its abapieal points. 

Dw: the maximum diameter of any whorl as the projection on the normal to the coiling axis 
of the pari of the planispiral or hclicoid cone produced by the generating curve after a p 
revolution and included in the given intervat (2n+I,)7f — 2(n-H )tt. 

/V: as the projection on the normal to the coiling axis of the distance between the adaxial 
point of the generating curve at 2mr and ils abaxial point at 2(n I l")*i position. 
Nis: intersulural distance as the projection on the normal to the coiling axis of the distance 
between the initial and the final position of a point placed on The adaxial line after a 2t 
revolution, where the adaxial line is the geometrical locus of any point placed at the adaxial 
end of the generating curve. 

D#c: diameter of the generating curve as the projection on the normal to the c. axis of the 
distance between its adaxial and its abaxial point. 

I he generating curve is here considered the projection on a plane, containing entirely the coil- 
ing axis, of the outline of the growing edge of a gastropod shell. The body whorl here repre- 
sents the part of shell cone generated by the growing edge in a given interval 2n* — 2(n H)**, 
T is the translation of the generating curve along the coiling axis per revolution. K and i arc 
respectively Ihe indexes of overlapping of any whorl; K, parallel to the coiling axis: r, uotmal 
to the coiling axis, and arc defined by the ratios K — Lis/Hw and v — Nis/N, (After Raup 
J966. Observations consistent with this theory will be presented in a subsequent publicalion.) 



NOTES ON PSEUOOMALAXiS FISCHER IN AUSTRALIA 



27 



amount, from a more hclicoidal protoconcb to 
a more planisptral lelcoconch. Undescrihed in 
Australia, this form was directly compared with 
two topotypos of tho only coeval species ul 
Pxeiuhnuihtxis known in Australasia, P. asoulff- 
(iirarux Ma\*ell. These two topotypes, more 
juvenile than the Australian ones, display a 
somewhat higher spire, producing a decreasing 
concavity and consequent flattening of the 
adnpicjil side, as a result o( the same inversion 
of the coiling; fainter carinac. but the younger 
of the two has them more marked. Protocoach, 
growth lines, outer and inner shape of the body 
whorl are the same. Because toe .amount of 
inversion m the coiling of the New Zealand 
farms appears to differ only slightly from that 
of the Australian forms, and is also variable in 
!he Australian and New Zealand specimen*. It 
is very difficult to distinguish fA the specific 
level between these two tormx. Hence, they are 
heris considered couspecific and representing 
probably different geographical morphs. 

hrom P. iaculptunuux Maxwell, the Anglo- 
Parisian Eocene P. dixoni (Vasseur) (Coss- 
manu 1D15, pp. 142-3), differs by its higher 
spire and presence of spiral riblets; the Prnto- 
Atlrialic Priabonian P. hcyrlrhi (Oppcnhcim, 
1896) by its much higher spire and an abaxial 
margin anapically more convergent lo the axis: 
the Ljguria-Picmonte Middle Oligoccne P. 
hatirux Sacco. IS92, hy its lower spire and by 
if ic presence of spiral ribiets. The Indian 
Eocene P- ptmjohcnxix Eames, 1952, seems on 
the coniiary >ery close, but the holotype being 
very juvenile and the original illustration very 
poor, it is impossible to determine the actual 
differences and affinities between them, 

The American Claiborne Kocene P. rotetla 
(li*a) (Palmer 1937. p. 176) differs by Us icct- 
anj*utar aperture, higher spire, and lesser num- 
ber Of whorls; P. texana (Aldrich) {Palmer 
1937, p. I7K) hy crenuhiled a basin I Veels and 
SplcaJ cools ou ihc adapical margin; and P. 
plrtmwrr/jfi Palmer (Palmer 1937, p. 37R. by 
subclinical margins. more imbricated sutures, 
and the stdrfral part of the last whorl twisted to 
the adapical. 

Otber Eocene species (Cownann 1915) are 
* i i- Anglo-Parisian P. puieUatus (Sowerby), 
and tbc Egyptian P. lyhicvs (Oppenbeim). 

Maxwell (Nov. 1974, pers. cotnm ) states: 
'two specimens of a Punuhntahxis have re- 
cently been obtained from n Mangaorapan 



(Lower Eocene) sample from North Canter- 
bury. The material is not well preserved but rhc 
shells closely resemble P. tistnlpturutus except 
tor their much smaller size (Ihc larger speci 
men is only 1 mm in diameter), i don't think 
that the lower Eocene shells are [uve-nites ol 
P. Q$atl#Mtr#ilt£, however, as (hey also have 
much smaller proioconchs". 

He quotes also the occurrence ol P. ef, 
wulpmrntm Maxwell from Wharckuri, South 
Canterbury ( Duntroonian, Ohgoccnc ) [ Max* 
well 1969, p. 16!) but The sole specimen is 
broken", however it *'i.s certainly close to the 
Eocene species'*. 



P*eudomaJaxb (Paeudomulaxb) 
praenieridionalis (Chapman, 1912) 

FIG. 4 

1912 Homahtxtx praemeritiinttulix Chapman,, 
p. 186, pi. 12, figs4-ri. 

Muieihil: 1 specimen Very well preserved 
(SAM, PI 83421. 

Dexvription: Shell small, thin, planoconcave, 
pseudoplanispiral, with the adapical side flat- 
tened and the abapical concave; whurl.s increas- 
ing far more in diameter than in height and 
overlapping entirely in relation lo the coiling 
axis and very scarcely to the normal tn it. 
Heterostrophic-anastTophic protoeonch with 
three smooth whorls, the inversion o\ the coil- 
ing already displayed in the third whorl. Suture 
grooved. Body-whorl shape, outer subquad 
rate - trapezoidal: inner subcireular - ovoidal; 
slightly wider than high. Body whorl regions: 
adapical convex in the adaxial part, concave in 
tho .abaxial; abaxial flattened; ahapical convex; 
aperture with subquadrate peristome. Lip*. 
adapical elliptical with a gutter-shaped reflec- 
tion in the middle; parietal and abaxial straight: 
ahapical elliptical. I .ip and region connections 
angular, marked by carinac. 

Orrutmvrttx; Four spiral beaded carinas, the 
abnxiul ones with very short aVmlly elongated 
spines: adaxial-adapical. adaxial-abapicaL aba* 
xial-adapical. abaxial-abapical; a spiral cumu- 
lated cord on the abaxial part of the adapical 
region and a smooth costella on the adapical 
part of the abaxial region. Growth lines: proso- 
cllne in the adapical region; prosocline in the 
abaxial; orthocyrt in the abapical. 



28 



m. r buonau.to 



OlMHNSrONS I in mm)- (tope Fig- 5) 



Nil wfaftte 
4 


Dw N 
2 6 13 


Nis llw 
0.45 0.88 


Ms 

—0.02 


Hoc 
0.90 


Dgc 
0.9S 


RATIOS: 
Hw Dw 

0.3381 


K Lis.'HW 
—0.0227 


v— Nis/N 
0.3461 


Hce Dgc 

U.9KS4 







Localities and Smitigntphic Distribution: 
Muddy Creek Maris, Clifton Bank. Muddy 
Crock, 6,4 km west of Hamilton; "blue days". 
Newport Formation, AUnnu Bay Coal Shall. 
Vic. 

Stnti (graphic Range: Balcombian, Early Middle 
Miocene (Ludbrook 1973). 
Observations: P. praenu-rklttJtjah's (Chapmnn) 
was referred to Darragh (1970. p. 1 SS) to 
Xfangotntia Mcstayer. The specimen here des- 
cribed represents the second discovery of this 
species. From P. praenwndionaft's, the Palco- 
Mcditerranean Neogene /*. ranclaca (Philippi) 
iWenz 193?, p, 668. fig. 1906) differs by a 
slimmer body whorl, higher spire, adapical 
spiral costa closer to the adapical-abaxiai 
carina, absence of spiral rihlets on the abaxial 
margin; from the Paleo-Mediterranean Pliocene 
P afdrovandi < Forest i) (Saccn 1 K92, n 75. pi. 
2. rig. 65 a-d, 65 bis a-d) by smooth adapical- 
abaxial carina, absence of spiral abaxial ri biers, 
higher spire, and adapical costa Closer lo the 
adapical-abaxial carina; from the Parathetys 
Miocene P. bocttgeri Cossmann (Cossmann 



dtotnthv iHedley) (Medley 1 903, p. 351) dis- 
plays, on the other hand, i very close affinity 
to P. ptaemeruiiOnalis ( Chapman ) , differing 
trom it only by two more abaxial riblels and 
the abapical-adaxial carina with heavier beads. 

The New Zealand Recent ?P, boftonsi (Mcs- 
taycr. 1030. p. 144) differs by more whorls 
two beaded abaxial cords, broader abapicul- 
abaxial carina and slightly opisthocline growth 
lines on the adapical region. 

I'he American Recent P. nohilis (Verrili) 
iVcrrill 1885, p. 423, p]. 44. fig. 12) differs by 
its shorter spire, a greater number of whorls. 
slimmer body-whorl, adapical cord closer to 
the abaxial-ndapical carina, five spiral ribs on 
the abaxial margin, and fine spiral ribs on the 
abapioil margin. 

Acknowledgment* 
I dm very grateful to Dr P. A. Maxwell trf 

the Geological Survey of New Zealand for his 
assistance, particularly in lending topotypes of 
Psrudontaiuxis asatlpturatus; to the Director of 
Mines. South Australia, for the loan of male- 
rial; to Dr H. A. Render, National Museum of 



1915, p. 143) by a larger body-whorl, lower Natural History. Smithsonian Institution, tor 



spire, lesser number of whorls, lack, ot spiral 
ornaments except rarely, beaded abaxial 
carinae, a semi-circular inner shape of the body 
whorl; from the Mediterranean Recent P. 
actoni Monterosato (1913, p. 362) by a greater 
number of whorls, slimmer body-whorL lower 
spire, more marked growth lines, more abaxial 
riblets, and an abaxial margin more parallel to 
the coiling axix- The Australian Recent P. meri- 



the information on taxonomy; to Dr H Silcock. 
Department of Pure Mathematics, University 
of Adelaide, for verifying the definitions of 
the pnramenlers; to Dr N. H. ludbrook for 
encouragement and for reading the manuscript. 
The work was carried out in the Department 
of Geology and Mineralogy, University of 
Adelaide, during tenure of a University Re- 
search Grant, 



References 



Aa&OTT, R. 1, iiyMl. "American Scashclls/ 
(Van Nofctraud: New York.) 

Chapman, F. (1912). — New or lilllt; known Vic- 
torian fossil* in the NutionaJ Museum. Pi 
XVI. Some Tertiary Gasteropoda Prcc. H. 
Soc. Vict, 25. 186-192. pis 12-13, 

CossMTan, M. (1915). — "Essais de Paleoconcho- 
logic comparei?." Vol. 10. (Paris,) 

Dall, W. H. (J 892). —Contribution to the Ter- 
tiary £aun3 of .Florida with special reference 
to the Miocene Silex Beds of Tampa and the 
Pliocene Beds of the Caloosahutchie River. 
Pt. 2. Strcptodont and other Gastropods 
Trans. Wagner free faith Sci. Phi fad, 3i'2l. 
201-473, pis 13-22. 



Darkagh, J. A. (19/0).— Catalogue ot Au&lialian 
Terliarv Mollusoa (except Chitons). Mem. 
Nalti. fats, Via. 31. 125-212, 

IEamhs, F. E (3952). — A canti ibtuioti lo the study 
of the Eocene in Western Pakistan and West- 
ern India. C. The description of the Scapho- 
poda and Gastropoda from standard sections 
in the Kakhi Nala and Zinrta Pir areas of the 
Western Punjab and in ihc Kohat District. 
Phil. Trans. R. Soc. London, B, 236(63 1), 
1-168, pis 1-6. 

FlSCUcft. P. (1880-87). — "Manuel de conchyologic 
ct de Pideontologie conchyologique on fnstoire 
nalurctle de?. mullusques vivunta el iWalts/' 
(Savy: Paris.) 



NOTES ON PSEUDOMALAXIS FISCHER IN AUSTRALIA 



29 



Glirfrt, M. (1973). — Revision des Gastropoda 
du Danien et tin Montien de la Belgique. Pt I. 
Les Gastropoda du Calcaive de Mons. Mer&. 
Inst. R. Sot\ Nat. Betg. 173, 1-116. 11 pis. 

Hedley, C. (1903).— Scientific results of the 
trawling expedition of H.M.C.S. ''Thetis 1 ', 
Mollusca. Pt II. Scaphopoda and Gastropoda. 
Mem. Austr. Mas, 4, 327-402, pis 36-8. 

Irt:dai.f. T. (1936). —Australian Molluscan Notes. 
No. 2. Rec. Austr. Mus. 19(5), 267-353. pis 
20-24. 

iREini e, T. (1911). — On some misapplied Mol- 
lusca generic names. Proc. Malac. Soc. T.ond. 
9, 253-265. 

Lindsay, J. M. (1969). — Cainozoic Koraminifcra 
and Stratigraphy of the Adelaide Plains Sub- 
Basin, South Australia. Bull. geol. Sun-. S. 
Aost. 42. 

Li/obrook, N. H. (1973). — Distribution and strati- 
graphic utility of Cenozoic Molluscan Faunas 
in Southern Australia. I ohoku Univ., Set. 
Rep., 2nd Bejr: ( Geol ) , Special Volume. 6 
(Hatiti Memorial Volume), 241-261, pis 24- 
28, 

Maxwtll. P. A. (1969).— Middle Tertiary Mol- 
luscs from North Otago and South Canter- 
bury, New Zealand. Trews. H. Soc. N\Z„ 
GeoL 6(13), 155-185, pis t-3. 

Maxwell, P. A. (1966). — Some Upper Eocene 
Mollusca from New Zealand. N.£. //. Geo!. 
Geophys, 9. 439-57. 

Mi:.stayer, M. K. (1930).— Notes on New Zea- 
land Mollusca. No. 5. Trans. N.7.. Inst, 61, 
144-46, pi. 26. 

Monterosato, Marquis de (1913). — Note on the 
genus Pseudomalaxis Fischer and description 
of a new species and subgenus. Proc. Mulai\ 
Soc. Land. 10, 362-363. 

Moore. R. C. (I960).— "Treatise on invertebrate 
Palaeontology." Part I, Mollusca, 1. (Geol. 
Soc. Amer. & Univ. Kansas Press: New 
York.) 



Oppenheim, P. (1896).— Das Alttertiar tier Colli 
Rerici in Venetian, die slcllung dcr schiclcn 
von Priabona und die oligocane Transgression 
in Alpinen Europa. Z. Dt. geol. Ges. 48. 27- 
152. pis 2-5. 

PALMbR van Winkle, K. (1937)-— The Clairbor- 
nian Scaphopoda, Gastropoda, and Dibran- 
chiate- Cephalopoda of the Southern United 
States. Bull. Am. Paleont. 7(32), Pt I. 1-548; 
Pt 2, Alias. 

PCULLIKTSEV. V., & K.OROBKOV, I. A, ( 1 960). — 

Osnovy Paleontologii. Spravochnik dlia pale- 
onlologov i geologov SSSR. Mulluski-Bruko- 
nogie. (Moscow.) 

Pilsbrv, H. A,. & McGinty, T. L. (1945).— 
Cyclostrematidae and Vitrinellidae of Florida. 
Pt. I & II. Nautilus 59, (1) 1-13, pis 1-2; (2), 
52-59, pi. 6. 

Raup. D. M. (1966). — Geometric analysts of shell 
coiline: general problems. /. Paleont. 41, 
43-65. 

Rrnor.R, H. A. (1935). — "New Caribbean Marine 
Shells." Nautilus 48(4), 127-30, pi. 7. 

Sacco. F. (1892).— T Molluschi dei Terreni Ter- 
ziari del Piemonte e delta Liguria, Part 12. 
(Clausen: Torino.) 

Sutlr, H. (1913-15),— "Manual of the New Zea- 
land Mollusca." (MacKay: Wellington.) 

Vkrrill. A. E. (1885),— Third catalogue of Mol- 
lusca recently added to the fauna of the New 
England Coast and the adjacent parts of the 
Atlantic, consisting mostly of deep-sea species. 
with notes on others previously recorded. 
Trans. Connect. Acad. Arts Set, 6, 395-452. 
pig 42-44. 

Wenz, W. (1939).— Gastropoda- In O. H. Shinde- 
wolf (lid.), ^Handbtich der Palaozoologic". 
Vol. 6, Part 3(4). (Gebruder Borm'tracgcr; 
Berlin.) 



THE CLINGING MECHANISM OF PSEUDOPHRYNE BIBRONI 
(ANURA: LEPTODACTYLIDAE) TO AN ALGA ON GLASS 

byN. Gradwell* 



Summary 

GRADWELL, N. (1975). -The clinging mechanism of Pseudophryne bibroni (Anura: 
Leptodactylidae) to an alga on glass. Trans. R. Soc. S. Aust 99(1), 31-34, 28 February, 1975. 
Despite the absence of an oral sucker, tadpoles of all stages from 26 to 40 (of Gosner 1960) were 
found to be capable of clinging by their jaws to an alga on vertical glass. When the glass was wiped 
clean of the alga, Phyllobium sp., tadpoles were no longer able to attach themselves. Therefore 
substratum algae are necessary for the clinging of the tadpoles to glass. 

As the nares appear to suffice as inhalent channels, the dental apparatus of tadpoles is adapted to 
maintain a firm grip on the alga. There is an absence of peripheral papillae adjacent to the most 
posterior of the tooth rows of the lower lip. Therefore this tooth row can bend farther forward and 
the security of its grip on the alga is probably increased. 



THE CLINGING MECHANISM OF PSEUDOPHRYNE BIBRON1 
(ANURA: LEPTODACTYLIDAE) TO AN ALGA ON GLASS 

by N. Gradwell* 



Summary 

Gradwell, N. (1975). — The clinging mechanism of Pseudophryne bibroni (Anura: Leplo- 
dactylidae) to an alga on glass. Trans. R. Soc. S. Aust. 99(1), 31-34, 28 February, 1975. 
Despite the absence of an oral sucker, tadpoles of all stages from 26 to 40 (of Gosner 
I960) were found to be capable of clinging by their jaws to an alga on vertical glass. When 
the glass was wiped clean of the alga, Phyllobium sp., tadpoles were no longer able to attach 
themselves. Therefore substratum algae are necessary for the clinging of the tadpoles to 
glass. 

As the nares appear to suffice as inhalent channels, the dental apparatus of tadpoles is 
adapted to maintain a firm grip on the alga. There is an absence of peripheral papillae adjacent 
to the most posterior of the tooth rows of the lower lip. Therefore this tooth row can bend 
farther forward and the security of its grip on the alga is probably increased. 



Introduction 

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 ( 1 963 ) 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 Staurois 
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. 



32 



N. GRADWELL 



B 



■ 



w» 



% 







Fig. 1. A. The tadpole of Pseudophryne hibroni (stage 36) clinging to algae on the vertical glass side 
of an aquarium. B. Oral apparatus of a clinging tadpole as seen through a thin transparent layer 
of the alga Phy/lobium sp. growing on the aquarium glass. As the lower beak is enclosed 
within the" upper beak, only the latter is visible. The lower labial teeth are inclined forward 
to grip the algae. Adduction is accomplished by a slight lateral compression of the lips and 
teeth. C The oral apparatus dissected free from a tadpole to show the beaks in an open 
condition. Calibration bars = 1 mm. 



SUBSTRATUM CLfNOlNG IS rSFUDOVHRYSE 



u 



River Crossing, near Jenolan Caves. N.S.W. 
The maximum depth of water was 0,5 Ffl and 
the bottom consisted of mud and fallen Euca- 
lypws leaves and bait. At 3.00 p.m. the Wait* 
temperature was 6 to ft C. Five hours later^ the 
tadpoles were placed in an aerated aquarium 
at approximately the same temperature. 

For initial identification, the mouth pads of 
the tadpoles were examined and compared with 
the description of Martin (1965). Beeau.se the 
absence of a sucker was noted, it was surprising 
to observe that all of the tadpoles clung to the 
vertical glass sides of the aquarium (Fig. 1A). 
This attachment was insecure, for these tad- 
pole* were sometimes displaced from their 
clinging sites when other tadpoles collided with 
them. Examination of the oral region of a 
clinging tadpole, by stereomictoscope through 
the "lass, showed that the lower beak, was kept 
closed within the edge of the upper beak (Fig. 
IB) and that the upper and lower tooth tows 
were held adducied, though not touching one 
another. The nates were the only inhalent chan- 
nels for gill irrigation. 

During preparations for photography it was 
found that the. inside of the glass aquarium wan 
covered with a thin layer of algae which pre- 
vented sharp focussing on the mouth and teeth. 
Therefore, half of one side of the aquarium 
was cleaned of its algae, and ■ camera conven- 
iently positioned to await the settling and cling- 
ing of a ladpole on the cleaned glass Although 
tadpoles attempted to cling to the cleaned glass, 
they were unsuccessful. However, they clung 
readily to the uncleaned glass, demonstrating 
the presence Of algae on the glass to be neces- 
sary foi xhe clinging of these tadpoles. 

Apart from clinging to the glass, the tad- 
poles also showed frequent feeding movements 
over the elaas dunng which their jaws opened 
and closed rhythmically in a scraping action. 
Tigure IC shows the jaws its an open condition 
and both the upper and lower beaks ate visible 
The predominant alga growing oil the glass was 
Phvlfobium sp. t identified from Pntscolt 1970 1. 
which has a thallus of branched stiands. At the 
ends oi the strands are swellings called aki- 
nctes, which contain chloroplasts. It was of spe- 



cial interest to find akinetcs m the manicotto 
("stomach"! of tadpoles, which suggests 1h«t 
thev hud been grazed "If the glass. II would 
also seem that the bulbous akinetes on their 
strand-like thallus could easily be gripped by 
the minute sharp teeth of tadpoles, thus provid- 
ing anchorage. It is possible that other filamen- 
tous algae may also permit the clinging of 
these tadpoles to substrata* 

There are no papillae behind the most pos- 
terior row of lower labial teeth. Therefore 1 his 
row of teeth can bend farther forward than if 
papillae were present here and so the leeih can 
grip substratum plants more firmly. However. 
some other suckcrlcss tadpoles (for example, 
Lhoiki vemw(xi) can cling feebly to vertical 
substrata in similar fashion even though they 
lack a posterior gap in, their peripheral papillae. 

Four tadpole* were raised into Juvenile i rogs. 
Until stage 40. the tadpoles < entire length, 16 .U 
mm; snout-to-vent length, 17.4 mm) were ahle 
to cling by their teeth to the substratum- Alter 
this stage the beaks and teeth were shed prioi 
in widening of the mouth. 

It is proposed here that Psettdophryitr 
bihroni ladpoJes are able to cling to vertical 
substrata by biting their teeth into algae and 
perhaps other vegetation. As the naruil inflows 
appear to be Suffi&lcnt for respiration, the juw« 
need not open and close rhythmically to admit 
additional water, but can maintain their hue 
on plants. However* I have not had the oppor- 
tunity to compare these findings with the ability 
of P. bibroui in their natural habitat to cling 
to .subslrata. The absence of an oral sucker 
in these tadpoles confers greater flcviHiUty on 
the faws and perhaps it broadens their food 
resources. On the other hand, an oral sucker 
in lotic tadpoles facilitates grazing on algae 
in swift currents (Gradwell 1971 ), where more 
tenacious gripping is needed. 

Acknowledgements 
I am grateful to Miss P. McDonnell and Mr 
B. Wilson Tor collecting the tadpoles, and KO 
Miss Vf. Anstis for identifying them. Mr M. J. 
Tyler kindly tend the manuscript and ofTefed 
helpful suggestions, 



ttiiApLiu. J. L. (1935). — The unatomy 

adhesive apparatus in the tadpole of Rana 
ajshatiu Gunthcr. with special reference to the 
adaptive modifications. Tranx. R. Sov. Edfft- 
tmrgh 58. 339-349 

GnsNErt. K- L. (I960). — A simplified table for 
staging aoucan embryos and fofV*c wiih noies 
on identification. HerpehAogica Xh, 183-190 



Reference* 

of the Giunwri c, N- (' 197 1 ).- Astaphux tadpole: experi- 
ments on the suetiou WW gill rnigniuui 
mechanisms. Cun. I, ZooL 49, 307-332. 



GR.vnweiL, N. < 1^73 ). — On the functional mor. 
phology of taction and gill miration in die 
tadpole of AsqaphUfo njvd miles <vn nihermt- 
lion. HotpetofofiicQ 2*>, 84-03 



34 



N. GRADWELL 



Gradwfxl, N. (1975). — Experiments on oral suc- 
tion and gill breathing in five species of Aus- 
tralian tadpole (Anura: Hylidae and Lepto- 
dactylidae). /. ZooL, in press. 

Hora, S. L. (1922). — Animal life in torrential 
streams. /. Bombay Nat. Hist. Soc. 32, 111- 
126. 

Martin, A. A. (1965). — Tadpoles of the Mel- 
bourne area. Vict. Nat. 82, 139-149. 

Noble, G. K. (1931).— "The Biology of the 
Amphibia." (McGraw-Hill: New York. Re- 
printed 1954, Dover, New York.) 



Orton, G. L. (1953). — The systcmatics of ver- 
tebrate larvae. Systematic ZooL 2, 63-75. 

Prescott, G. W. (1970).— "How to know the 
freshwater algae." (W. C. Brown Co., 
Dubuque, Iowa.) 

Tyi.er, 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. 11. Anura: Ranidae and 
Hylidae. Trans. R. Soc. S. Aust. 86, 105-130. 



GENETIC EVIDENCE FOR THE EXISTENCE OF TWO SEPARATE 
POPULATIONS OF RATTUS FUSCIPES GREYII ON PEARSON ISLAND, 

SOUTH AUSTRALIA 

byL. H. Schmitt* 



Summary 

SCHMITT, L. H. (1975). -Genetic evidence for the existence of two separate populations of 
Rattus fuscipes greyii on Pearson Island, South Australia. Trans. R. Soc. S. Aust. 99(1), 35-38, 28 
February 1975. 

A study of genetic variation of the enzyme glutamate oxaloacetate transaminase (GOT) reveals the 
presence of two distinct populations of the southern bush-rat (Rattus fuscipes greyii) on Pearson 
Island. Two allelic genes, Got-f and Got-l are present in the animals collected from the middle and 
southern sections of the island, while Got-f is absent in animals taken on the northern section. 
This is discussed in relation to the Pearson Island wallaby which was, until recently, restricted to the 
northern section of the island. 



GENETIC EVIDENCE FOR THE EXISTENCE OF TWO SEPARATE 
POPULATIONS OF RATTVS FUSCfPES GREY I! ON PEARSON ISLAND, 

SOUTH AUSTRALIA 

by L. H. Schmitt* 



Summary 

Schmitt, L. H. (1975).— Genetic evidence for the existence of two separate populations of 
Rattus fi/.scipps gXtyii on Pearson Island, South Australia. Traits. R. Sov. S. Aust. 99(1), 
35-38, 28 February 1975. 

A study of genetic variation of the enzyme glutamate oxaloacetate transaminase (GOT) 
reveals the presence of two distinct populations of the southern bush-rat {Rattus fuseipes 
greyii) on Pearson Island. Two allelic genes, Got-l« and Got-l b are present in the animals 
collected from the middle and southern sections of the island, while Gnr-f f > is ahscnt in 
animals taken on the northern section. This is discussed in relation to the Pearson Island 
wallaby which was, until recently, restricted to the northern section of the island. 



Introduction 

Pearson Island, which lies 60 krn off South 
Australia's west coast, is divided into three dis- 
crete sections (Fig, 1 ) , The southern and 
middle sections are linked by a causeway, while 
the middle and northern sections are separated 
by a narrow sea channel. The total area of the 
island is approximately 325 hectares. The Pear- 
son J, wallaby, Petrogafe sp. (see Thomas & 
Dclroy 1971, for a discussion on its taxonomic 
status) and the native rat, Rattus fuseipes 
greyii, are the only terrestrial mammals which 
are known to inhabit the island. 

The native rat, which is found on all three 
sections of Pearson I., was first described by 
Thomas (1923) who named it Rati us murrayi. 
Tredalc & Troughton (1934) reclassified it as 
Rattus greyii murrayi recognizing its close rela- 
tionship to Rattus greyii greyii, the native 
bush-rat of mainland South Australia. It was 
later included in the species Rattus fuseipes by 
Ellenrmn (1949) as a distinct .sub-species, /c. 
fuseipes murrayi, along with the mainland form 
R.f. greyii. Recently, Taylor & Horner (1973a) 
have considered it to he subs peri fie ally indis- 
tinguishable from RJ. greyii. 

Until I960, 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 



Tig. 1 . Map of Pearson Island. Areas where ani- 
mals were captured arc indicated by shad- 
ing. 



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



36 



I. K SCHMM I 



migration between the northern and middle 
sections. In I960, six wallabies, including either 
four or live females, were accidentally released 
on the middle seetiort and the species is- now 
abundant on the middle and southern sections 
(Thomas & Del toy iy69). 

This paper describes a genetic difference in 
the fl.f. xreyii population ol Pearson I., appa- 
rently caused by a restriction in migration 
across die channel separating the northern and 
middle sections*, 

Methods 

Specimens of H.L grttyti were caught in 
"Shei-marT traps and transported alive to Ade- 
laide. Tissue homo&euatcs were prepared and 
subjected to starch gel electrophoresis. The 
Llcclrophorelic buffers (pH SO) were essen- 
tially the same as those described by Scfander 
i-' (tl, (1971 >- The methods used to detect glu- 
tamatc oxaloaeetate transaminase (GOT) acti- 
vity and determine the subcellular locality of 
the isozymes were modified from those given 
by De Lorenzo & Ruddle (l a 70). Heart ex- 
tracts were used predominantly, except in lhe 
latter procedure when liver extracts were used. 

Results and Discussion 

Animals were caught from the areas shown 
ih Fig. 1. Seventy-five animals were caught in 
?iy ftpp nights, yielding a capture rate of 
34'% . This is coasiderably higher than the 
2J&% return obtained by Taylor & Horner 
I 1973b) for R.f. greyii on the mainland of 
Australia and Kangaroo 1. 

Two main regions of GOT activity were pre- 
sent in gels, one migrating eathodaJly, the other 
anocially, The cathodal area of activity con- 
sisted of a 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-IA, GOT-1B and 
GOT-IAB were observed. This variation Is 
consistent with the active enzyme heing a 
duneiic molecule and is similar to that found 
in man (Chen & (iihlett 1971 ) and Lire North 
American old-field mouse. Fetomyncits pollo- 
notuK (Selander ft af. 1971). Genotypes can 
he assigned to each phenotype. presuming that 
the difference is under the control of an auto- 
somal locus, with two co-dominant alleles. This 
Incus has heen designated Got 1 and the alleles 
Gat-J" and Got-i h , 

Laboratory matings of R,f 4 greyii individuals 
from different, areas of South Australia, includ- 
ing Pearson Li have been successful. Family 



TAuLII 1 

Family dura on GOT variation 



»iF parentis 


N timber of 


Number and OOT phc 
ul oUs'Ting 
A AR 




Ax A 
A at A* 

AxB 

A x AK 
HxAB 


2 

lf> 
3 
1 
1 


3 

44 H 
9 

2 (1 

tl 4 


1) 






(. 



* One parent was nut scored for GO'I phenotype. 
However, in each of the ten matings ii was 
known tu have come from a population appa- 
rency monomorphic for the Uot-J" allele. 

daia on the inheritance of the GOT variation 
(Tabic 1) »s cosistenr with a 1 locus, 12 
allele mode of inheritance. 

I he H.f* greyii population on the northern 
section is monomorphie tor the Got /'' allele, 
while hoih frV>r-/" and Got l h are present in 
the population on the southern and middle 
sections (Table 2). The genotypic frequencies 
in the latter population 111 the Hardy-Wcinbere 
equilibrium frequencies (P>0.05). If the 
Got-jf* allele is present in the population on 
Lhe northern section, then there is a V5% pro- 
bability that its frequency is less than 3%. In 
any case, the Irequeney o! the Got~i h allele in 
lhe northern section is significantly different 
from its value in (he muldle and southern **o- 
ttot* <P< <(X00T). 

TABLE 2 

GOT vfirnntypi* distribution in animals 
ctwpjtt on Peaf'KttH I 



Alijft i:anj»Tn 

Nortlurn section 
Middle afTds&tnftiMn 
sve (ion 



Numhcr ftttfl GOT r>hcni*uyr>e h'tjUPtfnC] 
A AB B of Oot-lb 



I. DO 
0*3*5 



It seems unlikely rhat the marked difference 
ii) allelic frequencies is maintained by selec- 
tion. All three sections of the island appear to 
provide very similar habitats fur H.f. grt^ii. 
The observed absence of Gtyf-/* from the 
northern section indicates a severe restriction in 
gene flow between the areas sampled on the 
middle and northern sections. The most 
obvious point of demarcation is the channel 
separating the two sections. This surprisingly 
low level of migration between the northern 
and middle sections is similar to rhat found in 
the Pearson I. wallaby 

It would appear improbable that the rats are 
not physically capable of crossing the channel 
When the sea is calm and the tide low, ft is 



GFNETICS OF THE PEARSON ISLAND RAJ 



37 



iltM 



A AB A B A AB 



Origin 



Fig. 2. GOT variation found in /?./. greyii from 
Pearson I. The pattern of variation is in 
agreement With a Uimeric molecule being 
the active enzyme. 

easy for a man lo wade or step from rock to 
rock between the two sections. However, the 
eonfoimalion of the channel is probably in g 
state of continual change. The relationship be- 
tween sea level changes and the occurrence of 
one or more species of small macropod mar- 
supials on small islands off the Western Austra- 
lian coast has been discussed by Main ( 1961 ). 
From Main's dala it appears that Pearson I. 
has been isolated from the mainland for at least 
10.500 yearv 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 since 
the isolation of Pearson Island from the main- 
land, the mean sea level on two or more occa- 
sions has been about 6 metres above ils presenl 
level (Twidalc. pcrs. comm.). During these 
limes of high mean sea level, the channel would 
have been considerably larger than at present. 
Thomas & Delroy ( 1971 I suggested that the 
wallahy did not cross the channel because it 
found the sea water distasteful. Another possi- 
bility is thai, because ot the action of the sea. 
there may be strong selective pressures against 
small animals which wander too close to the 
edge of the water. Under such an hypothesis. 
genotypes would be favoured which predis- 
posed animals to an aversion to moving close 
to the water's edge. This would discourage the 
animal from crossing between the middle and 
northern sections. 

Various suggestions may he Offered to 
account for the present distribution of the Iwo 
alleles at the Got-I locus. All other popula- 
tions of R.f greyii studied {Greenly I.. Hop- 
kins I.. Kangaroo I., Eyre Peninsula and Mount 
Lofty Ranges) have been found lo be mono- 
morphic for the Got-l 1 * allele (Schmitl. unpub- 
lished). 1 he polymorphism on Pearson L may 
have been present before the channel was 
formed and at that time, or some time afier- 
wards, the Got-h 1 allele was lost from the 
northern section. Alternatively, one of the 
alleles could have arisen by mutation, since the 
channel was formed. If the Got-l' 1 allele is the 
more recent mutant, then it has lo be postu- 
lated lhat it subsequently migrated across the 
channel. 

The channel separating the northern and 
middle sections has not only acted as an effec- 
tive barrier to migration lor the Pearson I. 
wallaby, but from the genetic evidence pre- 
sented here, has also had a similar effect on 
R ./. xreyii. 

Thirteen other gene loci, for which about 70 
specimens of R.f. greyii from Pearson I. have 
been scored, are monomorphic on Pearson 
Island and show no differences across the chan- 
nel. However, all Pearson I. animals have 
haemoglobin and malate dehydrogenase pro- 
teins that arc electrophoretically distinct from 
all other populations studied (Schmitt, unpub- 
lished). 

Further studies on the situation described 
here would be a useful part oi any future expe- 
ditions to Pearson I. 



38 



L. H. SCHMITT 



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 



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 



Chen, S. H.. & Giblett, E. R. < 1971).— Genetic 
variation of soluble glutamic-oxaloacetic 
transaminase in man. Am. J. Hum. Genet. 23. 
419-424. 

De Lorenzo, R. J.. & Ruddle, F. H. (1970).— 
Glutamate oxalate transaminase (GOT) 
genetics in Mus musculus: Linkage, poly- 
morphism, and phenotypes of the Got-2 and 
Got-1 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. /. 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 systematica in 
the genus Peromyscus. 1. Variation in the old- 
field mouse {Peromyscus polionotus). Stud. 
Genet. VI, 49-90. (University of Texas Pub- 
lication 7103.) 

Taylor, 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. 21, 437-475. 

Thomas, I. M., & Delroy, L. B. (1971 ).— Pearson 
Island expedition. 1969. 4. The Pearson Island 
wallaby. Trans. R. Soc. S. Aust. 95, 143-145. 

Thomas, 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 EUGENII 

by Patricia M. Mawson* 



Summary 

MAWSON, P. M. (1974). -Two new species of the nematode genus Cloacina (Nematoda: 
Strongylida) from the Tammar, Macropus eugenii. Trans. R. Soc. S. Aust. 99(1), 39-41, 
28 February, 1975. 

Cloacina smalesae and C. kartana are from the stomach of Macropus eugenii from Kangaroo I. 
C. smalesae is distinguished by the shape of the leaf crown elements, postoesophageal excretory 
pore, swollen cuticle at the anterior end of the body, and a long vagina. C. kartana is distinguished 
by the very small cephalic papillae and the presence of two oesophageal teeth situated shortly 
behind the nerve ring. 



TWO NEW SPECIES OF THE GENUS CLOACJNA (NEMATODA: 
STRONG YLIDA) FROM THE TAMMAR, MACROPVS ElIGENU 

by Patricia M, Mawsoic* 

Summary 

Mawson, P. M. (1974). — Two new species of the nematode genus Cloucina (Nematoda: 
Strongylida) from the Tammar, Macropus eugenii. Trans. R. Soc. 5. Aust. 99(1), 39-41, 
28 February, 1975. 
Claacim smalesae and C. kartana are from tbe stomach of Macropus Fugenii from 

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

phayeal excretory pore, swollen cuticle at the anterior end of the body, and a long vagina. 

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

oesophageal teeth siLuated shortly behind the nerve ring. 



Introduction 

The 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 analysis by Mrs 
Lesley SmaJes of the nematodes present at dif- 
ferent times of Ihe year in the stomach of this 
host, undertaken as part of work for a Ph.D. 
degree in the Department of Zoology. 

Types of the new species will be deposited 
in the South Australian Museum; paratypes are 
in the Helmintbologicul Collection of the Zoo- 
logy Department, University of Adelaide. 

Measurements of the two species arc given 
:n Table 1. 

Qoacioa smalesae A. &p. 

FIGS 1-7 
Host and Locality; Mtxropus cugeni: ( Dcs- 
nurest). from Kangaroo T. f S. Aust. 

This is a medium-sized species of Cloncinu, 
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 like a platform. The tnouth collar 
is well developed and slightly lobed anteriorly; 
it bears the four submedian cephalic papillae 
and two amphids. The distal segment of each 
cephalic papilla is longer and slightly thicker 
than the proximal segment. Each clement of 
the leaf crown is domed anteriorly and is with- 
out the unguiform anterior projection usually 
present to species of this genu*. 



The buccal ring is circular and stoutly built. 
Its anterior and posterior walls are sligluty 
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 dis- 
tinct, appearing in the whole mount as three 
wavy longitudinal lines throughout the length 
of the oesophagus. In transverse section the 
lumen appears tri radiate with a thickening of 
the lining In the midlength of each arm. The 
nerve ring lies at about midlength of the oeso- 
phagus. The excretory pore is postoesophageal. 
and the cervical papillae lie a little distance in 
front of the nerve ring. 

The bursal lobes are only slightly divided. 
The arrangement nf the rays (Fig. 6) is 
typical for the genus. The genital cone is not 
prominent; it bears a small ala on each 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 or tbe anus. The ovejectors unite 
6-7 tail lengths in front of the vulva, and the 
vagina curves forwards before passing hack, 
with one or two twists, lo Ihe vulva. 

The rounded elements of the leaf crown 
seen in this species have been described in only 
one other species. C. tongifttbiata Johnston & 
Mawson, 1939b (syn: C. minor J. & M., nee. 
Davey & Wood, 1938). However, in C Jongi- 
labiata the vagina is shorter, Ihe cephalic papil- 
lae are of a different shape and there is no 
cuticular inflation at the anterior end. 



/Oology Department, University of Adelaide, S Aust. 5000. 



40 



PATRICIA M. MAWSON 








Figs 1-8 



Figs y iA 







Claacina sttuttesae. Fig. 1. — Anterior end, lateral view. Fig. 2. — Anterior end, .showing 
leaf crown in everted position. Fig. 3. — Anterior end, en face. Fig. 4.— Oesophageal region. 
Pig, 5. — Transverse section of oesophagus, anterior to nerve ring, Fig. 6. — Posterior end 
of female. Fig. 7. — Bursa, flattened out and viewed from the inside. Fig. 8. — Lateral view 
of genital cone. Figs t, 2, 3, 5 and 8 to same scale, 

Cloacina keirtana. Figs and 10. — Sublateral and en face views of anterior end. FJg. It. — 
Oesophageal region. Fig. 12. — Transverse section of oesophagus, showing one oesophageal 
tooth Fig. 13. — Posterior end of female. Fig. 14. — Bursa. Fig. 15.— DorsaJ ray. Figs 10 
and 12 lo same scale, 



Cloacina kartana n. sp. 

FIGS 8-14 

Host and Locality; Macropm eugenii (Desmn- 
rcst), from Kangaroo I., S. Aust. 

This is a medium-sized species of Cloac'ma 
with a well developed mouth collar bearing the 
four small submedkun cephalic papillae and 
the amphids, The cephalic papillae are small; 



the proximal segment is longer and slightly 
thicker than the distal one. The buccal ring is 
stoutly built, and is thicker posteriorly than 
anteriorly. The six elements of the leaf crown 
do not, in the resting position at least, project 
above the collar. 

The oesophagus is cylindrical for most of 
its length, with a terminal bulb. There axe two 
small subvcntral oesophageal teeth, one a little 



TWO SPECIES Ob CLOACiNA 



41 



TABLE 1 
Measurements of Cloacina sirutlesae and C. kartana; in mi unless otherwise stated. 



C. smalesae 



C. kartana 



Length (mm) 

Oesophagus 

Antr. end — nerve ring 

— cerv. pap. 

— excr, pore 
Spicule length 
Length/spic. 1. 
Length/oesoph. 
Tail 

Postr. — vulva 
Egg length x breadth 



9.5- L 1.4 
690-720 
360-400 
200-230 
790-S70 

4200-4700 
2.2—2.5 

lli-16.0 



13.0-15.8 
710-790 
360-490 
160-200 
910-1000 



16.8 21.8 
220-260 
350-490 

170-175x85-90 



8.0-10.9 
680-790 
300-350 
220-300 
600-740 
1400-1600 

5.7-7.1 
10.6-16.0 



12.4-17.0 

810-900 

340-400 

250-300 

700-800 



J 4.3-20.9 
200-300 
380-500 

180-185x85-90 



posterior to the other, at about midlength of 
the oesophagus, and shortly behind the nerve 
ring. 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 



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, 1939a, 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 



Davxy, D. G., & Wood, W. A. (1938).— New 
species of Trichoneminae (Nematoda) from 
Australian kangaroos. ParasitoL 30, 258-266. 

Johnston, T, H„ & NfAWsON, P, M, (1938). — 
Strongyle nematodes from central Australian 
kangaroos and wallabies. Trans. R. Soc. S. 
Am. 62, 263-286. 



JoHNsroN, T. H., & Mawson. P. M. (1939a). — 
Strongylate nematodes from marsupials in 
New South Wales. Proc. Linn. Soc. N.S.W. 
64, 513-516. 

Johnston, T. H.. & Mawson, P. M. ( 1939b). — 
Sundry nematodes from eastern Australian 
marsupials. Trans, R. Soc. S. Ausi. 63, 
204-209. 



THE GENERAL WATER CIRCULATION OF SPENCER GULF, 

SOUTH AUSTRALIA, 
IN THE PERIOD FEBRUARY TO MAY 

byD. A. Bullock* 



Summary 

BULLOCK, D. A. (1975). -The general water circulation of Spencer Gulf, South Australia, in the 
period February to May. Trans. R. Soc. S. Aust 99(1), 43-53, 28 February, 1975. 
Historical temperature and salinity data are presented together with the three-dimensional velocity 
structure of the general water circulation of Spencer Gulf, deduced from the data using a numerical 
model. In the southern area of the Gulf a cyclonic gyre is found which exchanges Gulf water with 
the ocean water outside the Gulf. On the western coast of the Gulf, a moderate boundary flow, 
which is called the "Port Lincoln Boundary Current", is evident at all depths. The water in the 
northern portion of the Gulf circulates both in the vertical and the horizontal and appears to form an 
almost closed system separate from the system in the southern region. 



THE GENERAL WATER CIRCULATION OF SPENCER GULF, 

SOUTH AUSTRALIA, 
IN THE PERIOD FEBRUARY TO MAY 

by D. A. But. lock* 



Summary 

BULLOCK. D, A. (1975) — The general water circulation of Spencer Gulf, South Australia, in 
the period February to May. Trans. R Soc. S. Aitst. 99(1), 43-53, 2S February,. 1975. 
Historical temperature and salinity data are presented together with the three-dimensional 
velocity structure of the general wa tcr circulation of Spencer Gulf, deduced from the data 
using a numerical model. In the southern area of the Gulf a cyclonic gyre is found which 
exchanges Gulf water with the ocean water outside the Gulf. On the western coast of the 
Gulf, a moderate boundary flow, which is called the "Port Lincoln Boundary Current", is 
evident at all depths. The water in the northern portion of the Gulf circulates both in the 
vertical and the horizontal and appears to form an almost closed system separate from the 
system in the southern region. 



Introduction 

Ao investigation of the general water circu- 
lation in Spencer Gulf, and the region outside 
it. is made 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-south movement of the Gulf's waters are 
excluded. Tidal currents can be superimposed 
on the more sustained flows caused by thermo- 
haliue (and wind) forcing which arc 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 watcr 
entering Ihe Gull', and in part by variation in 
chc effect of evaporation over the Gulf. 

All available data for the region, obtained 
On C.S.I.R.O. cruises, arc shown in Fig. 1, The 
cruises took place between 1961 and I9o6 dur- 
ing the months indicated in the caption. No 
direct current 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 FR.V- Inves- 
tigator (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 watcr column. 
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 
gradient 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 Id evaporation and mixing. The 
density section (Fig. 2b) has a similar form to 
the salinity section, indicating that salinity is 
the dominant factor in determining thermo- 
haline flow. 

the most important feature of the hydrology 
of Spencer Gulf r evident from all data (Figs 
10, 11), is lhat the watcr is approximately 
homogeneous from top to bottom, and hence 



♦ The Flinders Institute for Atmospheric and Marine Sciences, Flinders University of South Australia, 
Bedford Pari:, S. Aust. 5042. Present address: Dept of Civil Engineering. University of Melbourne. 
Parkvillc, Vic. 3052. 



44 



D. A. BULLOCK 




■s 


A 


8 

it y 








/ B 


o 






/ 








$ 








/ ° 




"F 


/ 


' ^ 


:i /. 






o 


^ 




/ A 


* ^. 


r j>~ w & 






b* 




/ 


' * ^ 


^ .*■ 




/ 




tj 


^ 



Fig. 1 



WATER CIRCULATION Oh SPENCER GULF 



* 



STAT I Oh 1(0. 

'0? |W }Q\ tbt i? 3| 

"ttt^ r — r 
7/f 




HijL 2. Vertical sections across Spencer Gulf near 
the month, Marked — ■ " — on Figure 1. 
Jala taken from CSIRO Aust. 1968, samr> 
ling depths indicates by dots, 
(a) Salinity f;,,), <b) <r ( . 

it is reasonable that the quantities be repre- 
sented by their depwVaverage. Fig, 3a-c shows 
horizontal depth-average distributions of salin- 
ity, temperature and density drawn from data 
obtained in the period 4-7 May 1964 
<C.S.l.R.O. 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, which spans only a short duration, so 
that seasonal and shorter term variations m 
properties do no| cause misleading interpreta- 
tions to be made. 



The distributions shown in Fig. 3 are con- 
sistent with those in Fig. 2 and give additional 
information A How up the western shore is 
evident and it can be seen that the salinity in- 
creases eastward over the whole southern area 
of the Gulf indicating that the movement nf 
water is basically clockwise. It appears that the 
advective exchange with the water outside the 
Gulf is limited to the area below a latitude of 
about 33'45'S. Above this region where the 
Gulf narows. the water appears u blocked-in M . 
Along the eastern and central areas of the 
upper part of the Gulf, the high salinity water 
seems to flow down the Gulf until it meets the 
lower salinity water taking part in the advec- 
tive process described above. 

The data from the northern pari indicate thai 
the water moves as a mass in phase with the 
tidal currents. At 34 r 00'S, 137'MO'E, two sfct- 
lions (76 and 89) were tKeupied, separated by 
a period of 29 hours during which the depth- 
average salinity changed from 38.40';,, to 
38.14',,'-,.. At 33'57'S, I37°25'E (stations 77 
and $5) t the aalinitites changed from 38.35^.;,. 
to 38.60'r, over a period of 21 hours. Using 
the horizontal salinity gradients at this location. 
Fig. 3 shows that this salinity variation corres- 
ponds to a tidid excursion of a few kilometres 

The mechanism responsible for the homo' 
geneity of the water columns is based on ver- 
tical 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 thU 
observation is the following. Since, in the north 
of the Gulf, the advective exchange of pro- 
perties is small and the water shallow,, the 
effects of evaporation ate more marked than 
further south, as is reflected by the high salini 



P«i£. ji Location of (Jeeanographic Stations in the South Australian Gulf System until W3. 
key to Swnhots 

a H.MA.S. Gascoyne (G2/61) Feb.-March 1961 (CSIRO Au<t 1966i 
b H.MA.S- Gascoyne (G2/63) March 1965 (CSIRO Aust. J%7ai 
H.M.A.S. Gascoyne iG5/€S) March-April 1965 (CSIRO 1967b) 
H MA.S Gascoyne (G2/62) June-Aug. 1962 (CSIRO Aust. 1967c) 
H M.A.S. Gascoyne (G2/64) Feb. 1964 (CSIRO Aust. t967d> 
H.M.A.S. Gascoyne (G2/65) Feb. t965 (CSIRO Aust. !968a> 
HA1.A.S. Diamanlina (Dm2/66) April 1966 (CSIRO AuM. 1969) 
F.R.V. investigator 1963 (CSIRO Autf. 1968b) 
F.R.V Investigator 1964 (CSIRO Aust. !96Rc) 
F.RV. Investigator Dec. (962 (CSIRO Aust. 196ifci) 
F.R.V. Investigator Feb. 1965 (CSIRO Aust. 1968c > 
r.V. Kstellc Star April-Mav 1965 (CSIRO Aust. I968f> 
F-V .EsteHe Star April 1966 /CSIRO Aust. I968g) 
F.R.V. WceriUia Jaru-March 1961 (CSfRO Ausf. 1968h) 



4(i 



D. A. BULLOCK 



KM SD 




lig. X Horizontal depth-average distributions in Spencer Gulf. Data taken from CSIRO Aust. t%8c. 
(a) Salinity {#*}, (b) Temperature ( U C), (c) <. 



ties. The process of evaporation requires the 
extraction of latent beat of vaporisation which 
in turn lowers the temperature and nullifies the 
effect of direct healing. The following rough 
calculation shows that there is an approximate 
balance between the heat input from the atmos- 
phere and the heat output due to evaporation. 
From Haney (1972), the heat flux/ unit area 
is proportional to the difference between the 
apparent air temperature and the sea tempera- 
ture (denoted by T\,* and T H respectively). 
Assuming this flux is all available for evapora- 
tion, we obtain 

E-MV-T.) 

where h is the evaporative heat flux/unit area, 
and approximately * 34 Wm--(°C)~ l (Haney 
1972). Taking the values, T a * ~ 2TC (the 
mean temperature for early April), and B = 
100 Win 2 , and substituting in equation (I) 
yields "C -^ 1 8 U C. which is approximately the 



observed value (Fig. 3b). The neglect of advee- 
tive exchange is justifiable in view of the ob- 
served small variation of temperature and 
closed nature of the circulation in the northern 
Gulf. Vertical mixing maintains the tempera- 
ture difference between the atmosphere and the 
sea surface. 

Locations 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 accurately. 

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 SPFNCFR GULF 



47 



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 1 . This is done by solving the finite 
diilcrcncc representation of the component 
transport equations written in a form applicable 
to the terrestrial situation-. 



£ » *-*$*•-*¥ 


3c t 

3x 


£-*»-*$-*•* 


3y 


where Ihc following notation is used; 





sx 



sy 



- k|ub|u b 

- k|ub|v b 



(2) 

(3) 



U 
V 

z 

f 

H 
g* 

i 

V 

K 



u 



udz, V 



— H 



vdz 



H 



eastward component of velocity (east = x direction) 

northward component of velocity (north = y direction) 

x & y components of bottom velocity (u B ) 

distance measured perpendicular to earth's surface, positive upwards 

(* — 1 ) a l(K Pin kg!- 1 

Coriolis parameter = Z^cos^ w — angular speed of the earth's rotation. *P = latitude 

depth 

ICH *ar. ii.ng. "sr-DH =- specific volume of the standard ocean 

gravitational acceleration 

sea surface elevation 

bottom drag coefficient 

density of a standard ocean — 

«i!nll,(j 

x & y components of surface wind stress 



The terms in equations (2) and (3) have the 
following physical meanings: the left-hand 
sides arc the linearized accelerations, the first 
terms on the right-hand sides are the Coriolis 
accelerations, the second terms are the hori- 
zontal pressure gradients due to elevation of 
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 another computer pro- 
gramme from the observed density distribution 
(Fig. 3c) and the depths were interpolated 
from those given on a naval hydrographic 
chart. A typical value of the x component of 



the thermohaline terms was 300 mmV-. This 
can he compared with the typical magnitude of 
100 mm-s 2 for the forcing term corresponding 
to wind stress, so that the circulation in Spencer 
Gulf due to thermohaline causes can be ex- 
pected to be quite significant. The bottom drag 
coefficient K = .0025. 

FLOWDHN was run with the precalculated 
thermohaline term as the only forcing term and 
with the thermohaline term together with a 
constant wind. After the equivalent of 17 hours 
an equilibrium current was reached (actually 
a small oscillation ahout the solution remained. 
but this was ignored). The initial conditions 
were that the How velocity was zero everywhere 
and the surface elevation was also zero every- 
where except at the boundary of the model 
across the south of the Gulf, which was 
assumed to have an initial slope similar to that 



1 Bye, J. A. T. T 1975. Thallasso-seale nuraericaJ modelling. Computer Report. School of Earth Sciences. 

The Flinders University of South Australia. (Unpublished.) 
-Bullock, D, A. 1973. The general circulation uf St. Vincent and Spencer Gulf, Honours Thesis. The 

Flinders University of South Australia. (Unpublished.) 



48 



ft A. BULLOCK 



developed later itt the Adjacent pan of the 
Gulf. The boundary slope was found to have 
Male effect on the current distribution and so 
the assumption is presumed lo nol have serious 
consequences. 

The I'esuJtaul depth-average current, which 
ii the transport divided by the depth, was ob- 
tained al each point of n grid system formed 
over the Gulf, in polar form so that yjfafc magni- 
tude and direction of the current could be seen 
immediately. The grid length was 6.4 km. 

f £y 



To gain a detailed picture of the velocity 
field in the Gulf, a routine was added to the 
program which printed out the surface anil bot- 
tom currents. By involving the shallow water 
assumption, i.e. that the Ekman layer {the sur- 
face- layer influenced by wind) and the layer 
oli bottom frictionaJ influence ovedap fan 
assumption which is consistent with the waters' 
vertical homogeneity), it can be shown (Bul- 
lock 1973 — footnOLe 2) thai the curreni iom- 
ponents at nny level are given by. 



H 



(z 



b 



f'l) 



v(z) = 



n " f 3x 

Appl> mg c4ua(ions (4) and (5) at z = *i - 
and at z " — H gives the surface and bottom 
currents. 

Since ^ r is independent of z it. can be seen 
Irom equaiions (4) and (5) that the vertical 
homogeneity fit the Gulf implies that the cur- 
vent 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 about a 
line Marking mid-depth (Fig. 4) and give, as a 
I unction ol depth, the current which is directly 
a*latcd to Lhe local distribution of density. The 
lirst terms on the right-hand side of the equa- 
tions, on the other hand, represent the depth- 
average current predicted by the model under 
the constraints imposed hy the Boundaries of 
the Gulf (the shores and bed). 

The vertical components ol velocity were 
also found, using lhe bottom current and. the 
eradicate of the bottom topography, 






c • 



w = 



3H 



31 1 



U B ^ + V B 97 



(61 



Finally, the changes in the sea IcvcT which 
develop from the initial surface condition of 
zero slope are also obtained, 

Results of the Numerical Miukl 

( 1 ) Predicted Velocity Fields due to Thermo- 
halinc forcing only 

Fig. 5 shows the main features of the pre- 
dicted depth-average current. The figure is con- 
sistent with the water ma.ss analysis of the pre- 
vious section, i.e. the circulation in the wide, 
southern pari of Spencer Gulf consists ol a 
clockwise gyre, water entering via the western 
side and leaving on the eastern side and the 
circulation in the northern end appears closed. 



The magnitude of the velocity vectors shown is 
typically about n.07 ms 1 (ranging from zero 
to I? mr 1 ). 

from Figs 6 and 7, which show the surlace 
and bottom currents respectively, it can be seen 
that at both the surface and bottom there is a 
very distinctive flow up the western side of the 
Gulf, which we shall call the "Port Lincoln 
Boundary Curreni'* This current is broader in 
e-.tcnt at the surface than at the bottom wheie 
it tends to peel away to the east down the slope 
of the boiioin. The Port Lincoln Bounary Cur- 
rent is also somewhat stronger at tJie surface 
than at the bottom due to the fractional 
iniluencc of the latter. 

The direction of the bottom current can he 
seen to swing eastward and then to the south- 
east over a broad area before flowing south- 
ward to exit down the -shovel-shaped topo- 
graphy through inc Gulf's mouth. The surface 
current displays a similar pattern; howcvei. 
since the Port Lincoln Boundary Current is 
wider at the surface, particularly near the 
mouth, the centre of the gyre of the surface 
circulation is located further to the north-casr 
than the centre of the bottom gyre. Evidence of 
the eastward movement of water from the 
boundary current is also provided hy Fig, 3a— c 
where the contours bulge to the east in thi.s 
area. 

The differences between the top and bottom 
current gyres implies that the surface and bot- 
tom currents on the eastern side of the Gulf 
are, tn general, at an angle to each other, un- 
like the western boundary. This explains whv 
there is no obvious salinity or ff l tongue indi- 
cating a return* How down the eastern side of 
the Gulf in Fig. 3b and c. 



WA1ER CIRCULATION OF SPENCER GULF 



49 



/ 



/ 



■>■ 



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- 



gences of the currents exist. A large proportion 
of the water in the northern end appears to 
recirculate in a vertical gyre. Fig. 8, which 
shows the vertical components of velocity, 
lends support to this interpretation by the 
downwclling at the top of the Gulf near Point 
Lowly, which corresponds to an inflexion in the 
salinity contours (Fig. 3a). Tn the southern 
part of the Gulf, a complex system of vertical 
cells of typical dimension 20 km is predicted. 
The following changes in surface elevation 
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 
Gull 

(2) Velocity field due to Thermohaline and 
Wind Forcing 

The numerical experiment was now extended 
to include the effect of wind stress in addition 



PORTAUGUSTA 



KM 50 




P0RT\ ^ \ V "" 

.[NCOLN-yr*} ^ 



Fig. 5. Predicted depth-average thermohaline currents in Spencer Gulf. 
Fig. 6. Predicted surface thermohaline currents in Spencer Gulf. 
Fig. 7. Predicted bottom thermohaline currents in Spencer Gulf. 



50 



O. A. BULLOCK 




D DOWNWELUNG 
U UPVVCLUNG 



KM CI 



\ 



\ 



, IP 

fee 





Fig. 8. Predicted regions of up- and down-welling due to thcrmohalinc currents in Spencer Gulf. 
Fig. 9. Predicted currents in the presence of a 14 ms 1 SW wind- 
la) Surface, (b) Bottom. 



to the thermohaline forcing. The resulting sur- 
face and bottom currents predicted by the 
model for the same density structure as before, 
combined with a 14 ms -1 south-west wind, arc 
shown in Fig. 9a and 9b. 

The pattern of circulation is basically the 
same but the velocity of the flow is accentuated, 
being typically about 0,16 mr 1 , some velocity 
vectors having a magnitude over 0.30 ms 1 . 
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 it 
was directed southwards previously. Similarly 
the bottom current flows north adjacent to the 
shores, whereas in the no-wind case ihe bottom 
current was slight here. It 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 rrO with distance 
up the Gulf, i.e. the water is piled up at the 
northern end by the wind. The low in the 
southern half is also accentuated, the level 
being raised by up to SO mm around the shores 



compared with the depression of 10 mm in the 
centre. 

( 3 ) Possible reasons for the Persistence of the 
Density Structure 

In the numerical model, the thermohalinc 
forcing terms remained constant throughout the 
period from zero velocity to equilibrium; i,e, 
wc assume that the observed density field per- 
sists and is not an instantaneous feature which 
subsequently runs down until the Gulf becomes 
a completely homogeneous body of water in 
which water motion of thcrmohalinc origin 
would cease- 
It i.s surmised that the cause of the persist- 
ence of the thermohaline circulation is external, 
i.e. it i.s due to water from the Southern Ocean 
moving up under influences that determine the 
pattern of flow in this region, perhaps the pre- 
vailing westerly winds. Fig. 1 2, 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, basically perpendicular to the Aus- 
tralian coastline. 



WAlEK CIRCLXATION OF SPENCER GUEF 



5! 







■in 






■j 




hi*. K), Vertical sections along Investigator Strau 

to the Continental Slope. Marked 

on Figure 1, data taken from CSIRO 
Aust. 1966 and iy67a. Sampling depths 
indicated by dots, 
(a) Temperature (°C), <b) <r t . 

The other factor which exerts a continuous 
effect on Spencer Gulf is evaporation, which 
rws the effect of maintaining a high .salinity and 
hence a high density at the northern end. ft is 
worth noting that in Fig. 3a the salinity tongue 
extending downward from the Gulf's northern 
end occurs over a favourable bottom gradient, 
whereas this is not so for the tongue which 
corresponds to the Port 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. I. 
Data from this region were obtained primarily 
to aid the fishing industry and suffer from the 
drawback that there are no synoptic surveys of 
the whole area, which would enable horizontal 
distributions of properties to be drawn. Only 
vertical sections and profiles can be satisfac- 
torily constructed. The paths along which sec- 
tions have been drawn are shown by the 
various lines in Fig. 1. 



Fig. 10a and b show temperature and % 
sections drawn along Q line extending through 
Investigator Strait to past the end of the Con- 
tinental Slope, The data were obtained in Fcb,- 
March 1%1 (C.S.1.R.0 Aust. 1966) and 
March 1963 (C.S.I.R.O. Ausl. 1967a), One 
feature which is noticeable is the change in pat- 
tern from a vertically homogeneous situation 
just inside the mouth of Investigator Strait to 
the more complicated horizontal stratification 
in the deeper waters outside the Gulf. 

The most significant feature shown by these 
sections, however, is the gradient in salinity, 
temporal ure and density about Station 122. 
which was located nonh-east of Cape Borda on 
the north-west tip of Kangaroo Island. This 
density gradient implies the existence of tf 
thcrmohaline current, water to the west of Sta- 
tion 122 flowing approximately perpendicular 
to the plane of the section (into the page). This 
direction correlates very closely with the direc- 
tion of the thcrmohaline current entering 
Spencer Gulf which forms the Port Lincoln 
Boundary Current. Conversely there is also a 
southward directed flow between Stations 121 
and 122 which could have its origins in the 
outflow from the eastern side of vSpencer Gulf. 
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 occupied in a different year to the 
adjacent stations (although at the same time 
of the year), so that the surface water tempera- 
tures are not consistent 

The section shown in Fig. Ma and b runs 
roughly in a north-east direction from the Con- 
tinental Shelf up Spencer Gull (C.S.I.U.O. 
Aust. 1967a K The gradient of the lines adja- 
cent to the rapidly deepening region outside 
Spencer Gulf under Station 64 again indicates 
a current inwards across the plane of the sec- 
tion, probably angled up 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 line 
of the section. 

Fig. 12 shows that the direction of the circu- 
lalion in (he Southern Ocean is consistent with 
the How in the region outside the Gulf as de- 
duced from Figs 10 and 1 I 

Conclusion 

The various data presented were mainly col- 
lected in the months of February to May, 



52 



D. A. bullock: 



STft 1 1 D N r-l to. 
5ft 6i 



65 bl 6B 59 70 




23 
J 50 

i ait 

tr. 

10 [I 


2 = 6 / 2SO 

: V// / 


b 



Fig, II. 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 CO, <b) ^ t . 

although during different years, so that it would 
appear that the general circulation deduced 
occurs at least seasonally. However, to gain a 
complete and detailed piclurc 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 need to be made to 
check the results from the model. Since the dis- 







>918/ MH / <•«* <~-^fl*3 \ 




1-3£6 t'Sll 



Fig. 12. Dynamic topography relative to 2000 dh., 
south of Australia (taken from Bve 
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 
Vincent Gulf would also display a pattern 
basically similar to that in Spencer Gulf. 

Acknowledgement 

I am 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 
Augusl lf>73. 



References 



Rowdhn, K. G. (1965). — Horizontal mixing in the 
sea due to a shearing current. J. Fl. Mech. 21, 
83-95. 

Bullock. D. A. ( 1974).— Cruise Report 1. (Flin- 
ders Institute for Atmospheric and Marine 
Sciences, The Flinders University of South 
Australia.) 

Byi:. I. A. T. ( 1971 >. — ^Variability south of Aus- 
tralia. Proc. hul Mat, Set. Sytnp. Institute of 
Marine Science. University oj fitgW South 
Wates, / 6- 17 Aug., 1971. 

CSIRO Aust. (1966). — Oceanographies! observa- 
tions in the Indian and Pacific Oceans in 
1961. H.MA.S. Gascoyne, Cruise G2/6I. 
CSIRO Aust. Oceanog. Cruise Rep. 10. 



CSIRO Aust. ( 1967ti ).— Oceanngraphicai obser- 
vations in the Indian Ocean in 1963. 
H.M.A.S. Gasvoyne, Cruise G 2/63. CSIRO 
Aust. Oceanoxr. Cruise Rep. 22. 

CSIRO Aust. (1967b). — Oceanngraphicai obser- 
vations in the Indian Ocean in 1965. 
H.M.A.S. Goscoyne, Cruise G 5/65. CSIRO 
Aust. Oceanogr, Cruise Rep, 46, 

CSIRO Aust. (1967c), — Oceanographical obser- 
vations in the Pacific and Indian Oceans in 
1965, H.M.A.S. Guseovne, Cruises G 2/62 
and G3/62. CSIRO Aim, Oceanogr Cruise 
Rep, 16. 



WATER CIRCULATION OF SPENCER GULF 



S3 



CSIRO Aust. (1967d). — Oceanographical obser- 
vations in the Indian Ocean in 1964. H.M.A.S. 
Gascoyne, Cruise G 2/64. CSIRO At4st. 
Oceanogr. Cruise Rep. 34. 

CSIRO Aust. (1968a).— Oceanographical obser- 
vations in the Indian Ocean in 1965. H.M.A.S. 
Gascoyne, Cruise G 2/65. 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.V. 
Investigator on the South Australian tuna 
grounds in 1964. CSIRO Aust. Oceanogr. Stn 
List 67. 

CSIRO Aust. (I968d).— Investigations by F.R.V. 
Investigator on the South Australian tuna 
grounds in 1962. CSIRO Aust. Oceanogr. Stn 
List 60. 

CSIRO Aust. (1968c).— Investigations by F.R.V. 
Investigator on the South Australian tuna 
grounds in 1965. CSIRO Aust. Oceanogr. Stn 
List 70. 



CSIRO Aust. (1968f). — Investigations by F.V. 
EstelJe Star in South Australian and New 
South Wales waters in 1965. CSIRO Aust. 
Oceanogr. Stn List 71. 

CSIRO Aust. ( J 968g).— Investigations by F.V. 
Estelle Star in South and Western Australian 
waters in 1966. CSIRO Aust. Oceanogr. Stn 
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. /. 
Phys. Ocean. 1, 241-248. 

Marshallsay, P. G., & Radok, 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. y and Blackburn, D. T. A New Procedure for Mounting 

Cleared Leaves Using Polyester Resin 55 

Womersley, H. B. S., and Conway, Elsie Porphyra and Porphyropsis (Rhodo- 

phyta) in Southern Australia 59 

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

lyier, M. J. The Ontogeny of the Vocal Sac of the Australian Leptodactylid 

Frog Limnodynastes tasmaniensis ------ 85 

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

Australia 89 

Tyler, M. J., and Martin, A. A. Australian Leptodactylid Frogs of the 

Cyclorana australis Complex 93 



PUBLISHED AND SOLD AT THE SOCIETY'S ROOMS 

STATE LIBRARY BUILDING 

NORTH TERRACE, ADELAIDE, S.A. 5000 



A NEW PROCEDURE FOR MOUNTING CLEARED LEAVES USING 

POLYESTER RESIN 

byD. C. Christophel* and D. T. Blackburn* 



Summary 

CHRISTOPHEL, D. C, & BLACKBURN, D. T. (1975).-A new procedure for mounting cleared 

leaves using polyester resin. Trans. R. Soc. S. Aust. 99(2), 55-58, 31 May, 1975. 

Palaeobotanical studies of leaf floras require detailed comparisons with extant leaves. Techniques 

for clearing and mounting extant leaves using either sodium or potassium hydroxide or a 

lacto-phenol solution as a clearing agent have long been known. This paper presents modification of 

this technique using an hydroxide dealing agent and mounting in polyester resin. 

In this procedure leaves are cleared in 15% KOH followed by saturated chloral hydrate. Leaves are 

then dehydrated, stained with safranin and mounted in Mulford EX-80 polyester resin. Use of this 

mountant shortens preparation time by several weeks as well as giving superior transparency to 

specimens. Photographic reproduction using direct printing of specimens placed in an enlarger is 

also discussed. 



A NEW PROCEDURE FOR MOUNTING CLEARED LEAVES USING 

POLYESTER RESIN 

by D. C. Christophel* and D. T. Blackburn* 

Summary 

Christophel, D. C, & Blackburn, D. T. (1975). — A new procedure for mounting cleared 
leaves using polyester resin. Trans. R. Soc. S. Aust. 99(2), 55-58, 31 May, 1975. 

Palaeobotanical studies of leaf floras require detailed comparisons with extant leaves. 
Techniques for clearing and mounting extant leaves using either sodium or potassium hydroxide 
or a lacto-phenol solution as a clearing agent have long been known. This paper presents 
modification of this technique using an hydroxide clearing agent and mounting in polyester 
resin. 

In this procedure leaves are cleared in 15% KOH followed by saturated chloral hydrate. 
Leaves are then dehydrated, stained with safranin and mounted in Mulford EX-80 polyester 
resin. Use of this mountant shortens preparation time by several weeks as well as giving 
superior transparency to specimens. Photographic reproduction using direct printing of speci- 
mens placed in an enlarger is also discussed. 



Introduction 

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 1972 1 , Christophel 1973-, 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 1 967, 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 (1-5 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 Cinnamomitm 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). 

- Christophel, D. C. (1973). —Fossil floras of the Smoky Tower locality, Alberta. Ph.D. thesis, Uni- 
versity of Alberta, Canada (unpublished). 



56 



D. C. CHRISTOPHEL & D. T. BLACKBURN 




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

with a Leitz Aristophot 4x5 plate camera, x 2.5. 
Fig. 2. Same specimen as in Fig. 1. Direct print using a Durst 1000 enlargcr. 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 auloclaving 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, * 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 CLKARKD LEAVES USING POLYESTER RESIN 



57 



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 to absolute alcohol. This senes should 
take less than one day. 

4. Stain in 1 % Satranin in 1:2 absolute 
alcohol/ toluol solution. Stain for three to five 
minutes or until all vascular tissue has taken 
Up colour. 

5. Destain the mesophyll in alcohol/ toluol 
Liking care not w destain the ultimate vena- 
Iron. This step determines the final contrast 
between the mesophyll and th^ vascular tissue 
and is consequently critical. Experience has 
shown thai mounted leaves having an absolute 
absorbance oi' 1.2 to 2.5 at 525 am yield the 
best density for transmitted light photography, 
Absorbanees were measured on a Bcekman 
Acta CIII Spectrophotometer with the leaves 
at the back of the sample chamber. By com- 
parison, leaves having an absorhance of 1cm 
than 1.0 proved loo light for acceptable repro- 
duction while those with an absnrhance of 
more than 2.5 proved too dense. As a com- 
parison, the ab.vorbance of 0.2% saframn 
measured over a 1.0 em light path is t.9, and 
a one cm cuvette of this solution may be used 
as a control while detaining. 

6. Transfer correctly detained [eaves to 
acetone for about ten minutes to remove re- 
maining alcohol/ toluol. 

7. Without allowing the leaf surface to dry, 
transfer the leaf to uncatalysed resin mono- 
mer 11 . 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 
overnight m the monomer. This monomer bath 
is reusable, and consequently this, step is best 
carried out away from direct sunlight and 
under fairly cool conditions to prevent poly- 
merisation of the resin. 

8. Prepare ihe final mounting medium by 
adding about 5% of MEKP catalyst to ftesh 
resin monomer. This is best accomplished by 
mixing in a small glass phiai and Tolling 
between ihe hands to combine the two com- 
ponents. Such gentle mixing prevents ihe for- 
mation of bubbles in the mountant. 



9. Drain the leaf of excess monomer and 
mount, between two glass slides. Setting should 
take about 2 hours at 3(TC. Higher tempera- 
tures give shorter setting times but increase 
the risk of the resin parting from the glass. 

10. When the mount is dried, excess resin 
may be removed with a sharp razor blade or 
scalpel. Small particles or thin films may be 
removed with xylene or acetone. 

Discussion 

Having obtained a permanent mount of a 
leaf with maximum transparency, it is then 
necessary to reproduce the venation pattern 
with as great a degree of contrast as possible 
for comparison with fossil material. For nor- 
mal photographs, a Lettz "Artstophof 4 by 
5 sheet film camera with a Macro-Dia attach- 
ment for transmitted light gave excellent 
results (Fig. 1), With a large 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 js 
obtained. 

Since maximum contrast between Ihe veins 
and ihe mcsophyil tissue is desirable, however. 
the subtle shadings of greys provided by this 
method proved unnecessary. Galavazi {1963) 
made brief reference to the direct use ot an 
enlarger for plant materia] cleared in methyl 
benzoate. Ditcher < 1 974 j also successfully 
used this technique on skeletonized and 
hydroxide cleared leaves. This technique was 
also admirably suited tor our cleared leaves. 
The mounted leaves fit easily uitu the nega 
live 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 partem 
is the important aspect, this reversal b 
immaterial. 

To achieve maximum contrast, a number 
of different stains were tried (Morley 1949). 
Bismarck Brown gave perhaps the greatest 
contrast, but stained very unevenly. Safranin 
gave very nearly equal contrast, and had the 
advantage of staining much more uniformly. 
Grade four photographic paper was used to 
£ive 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 to three 



:1 For mounting the specimens, Mutfoid EX-80 poUester resin was usetf, It is KVmlable from MulfotC 
Hustles Pty Lid.. 25 An*ac Highway. Keswick, S. Alist. 5035. 



58 



D, C. CHRISTOPHEL & D. T. BLACKBURN 



weeks for difficult leaves. Work is in progress 
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 



Dilcher, D, L. (1974). — Approaches to the iden- 
tification of angiosperm leaf remains. Bot. 
Review 40(1), 1-157. 

Foster, A. S. (1952). — Foliar venation in angio- 
sperms from an ontogenetic viewpoint. Amer. 
J. Bot. 39, 752-766. 

Galavazi, G. (1965). — Clearing and staining 
plant material in toto with phloroglucinol- 
HC1 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. /. 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. Iowa 
State L ScL 41, 481-486. 

Morley, T. (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. Slain Tech. 
49(3), 175-176. 



PORPHYRA AND PORPHYROPSIS (RHODOPHYTA) IN 
SOUTHERN AUSTRALIA 



byH. B. S. Womersley* and Elsie CoNWAYf 



Summary 

WOMERSLEY, H. B. S., & CONWAY, Elsie ( 1975) .-Porphyra and Porphyropsis (Rhodophyta) 
in southern Australia. Trans. R. Soc. S. Aust. 99(2), 59-70, 31 May, 1975. 

Two epilithic species of Porphyra, P. columbina and P. lucasii, occur on southern Australian 
coasts, mainly during winter; their seasonal morphology and distribution are described. One 
epiphytic species, the little-known P. woolhousiae, is described from additional collections, some of 
which are reproductive. Porphvrousis minuta sp. nov., epiphytic on Pterocladia capillacea and 
other cartilaginous red algae, is also described. 



PORPHYRA AND PORPHYROPSIS (RHODOPHYTA) IN SOUTHERN 

AUSTRALIA 

by H. B. S. Womersley* and Elsie CoNWAYt 



Summary 

Womersley, H. B. S., & Conway, Elsie (1975). — Porphyra and Porphyropsis (Rhodophyta) 
in southern Australia. Trans. R. Soc. S. Aust. 99(2), 59-70, 31 May, 1975. 

Two epilithic species of Porphyra, P. columbina and P. lucasii, occur on southern Aus- 
tralian coasts, mainly during winter; their seasonal morphology and distribution are described. 
One epiphytic species, the little-known P. woolhousiae, is described from additional collections, 
some of which are reproductive. Porphyropsis minuta sp. nov., epiphytic on Pterocladia 
capillacea and other cartilaginous red algae, is also described. 



Introduction 

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. iimbilicalis (=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. iimbilicalis (=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 

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 jum thick; carposporangialj 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 ^m 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, pi. 265. 
J. Agardh 1883: 59. De Toni 1897: 15; 
1924: 12. Guiler 1952: 84. Lucas 1909: 
20; 1929: 15. 

FIGS L 2 



* Department of Botany, University of Adelaide, Adelaide, S. Aust. 5000. 
t Department of Botany, University of British Columbia, Vancouver, Canada. 

i Although the true nature of the reproductive bodies in Porphyra is not fully understood, the classical 
terms carposporangia and spermatangia are used here to avoid confusion (Conway 1964). 



60 



H. B. S. WOMERSLEY & ELSIE CONWAY 



sio*. vitv 




2 cm 














200/jm 




Tig. 1. Porphyra woalhouxiae. A. Blackman's Bay, Tas. tADU. A44234). B, Thallus margin, from the 
type. C. Thallus with margin (A44234). 



PORPHYRA AND PORPHYROPSfS IN SOUTHERN AUSTRALIA 



61 




B 






"v " % 

!ft. ft. 









^ J * 

'?£' 



f4 > y ^ . 



50>um 



« 



D 



-* 






A CD 



200>um 




I : 
1 * 

•■ * 5 g 

- • ' - ! : 

. f m . • * . . • * " <4lgf 

^f ^ % »" » ( " lit ' * *^k 



.1 



ft / * - S 



i 






" g & 






*■ *^ 



it 



v£$ 




■ft « 



> 









- **•*.• - «v* * ^^ »» „t \ . 

Fig. 2. Porphyra woolhousiae. A. Thallus with area of darkly-staining reproductive cells. (Blackmail's 
Bay, Tas.. ADU, A44234). B. Cell arrangement in older part (A44234). C. Thallus margin 
with liberation of reproductive bodies (St Kilda, S. Aust., ADU, A42722). D. Margin with 
spermatangia (A42722). 



€1 



H. B. S. WOMERSI T-V & ELSIE CONWAY 



Thai I us (Pig. Ml epiphytic on brown algae 
or rnrcly on red algae, to 17 cm high and 4 
cm broad, from irregularly ovate or cuneate 
elongate when young, to broadly expanded or 
elongate with curved margins winch are gently 
convolute in older plants (e.g. the type), deli- 
cate, with one to a few blades arising from a 
»hi/.oidal holdfast, rose-red to rose-purplish. 

Monostromatic, ( 1 2-) 16-24 /jn thick, 
ccUs isodiamcttic to slightly elongate in sec- 
non;»l vie.w, each with a single, laminale or 
stellate rhodoplast, with a pyrenoid. Growth 
hy a marginal meristem (Figs 1B,C, 2A ) 
and by intercalary divisions: margins meris- 
tematic, wilh anticlinal lows of cells; older 
parts with cells often in rows but becoming 
irregular, with cells often polygonal or angu- 
lar (Figs IB. 2B), some rounded to ovoid, 
often with adjacent cells unequal in size. Mar 
ginal cells isodiametric to slightly elongate, 
angular, 6—10 ^m across; cells [ft older parts 
ito-diamctric to twice as long as broad, 10-16 
i' -20) ft.m across. 

Reproductive bodies ( Fig. 1A ,C ) formed 
in marginal areas, apparently singly, spherical 
to slightly ovoid. 12-17 ^m across, usually 
with a stellate rhodopla.st, 

Spermatangia (Fig. 2D) forming well 
defined, irregular, patches near ihc thallus 
margins, the whole patch when matuie tend- 
ing to deliquesce with numerous spermatangia 
irregularly arranged; spermatangia originally 
formed in packets of (16-) 32-64, in two 
tiers, individually 2-4 uTO across. 

Type locality; Tasmania. 

Type: Herb. Harvey, TCD (presented by 

Miss Wool house of Sheffield). 

Distribution: As well as the type, this species 
is known from St Kilda, S. Aust., on Go;ar- 
tina{'!) on Poxidftniu, I] m below Jow tide 
level (S. Lewis. 23.Viii.I972; ADU. A42722): 
Mallaeoota, Vic, on Seytothammts auxtralis 
( Pucker and King, 1 5.ii. 1970; MELU. 
20652); I>eal l. T Bass Strait, on Pfrithalia 
cmuhto (Kiw>, 2Xxi,1969; MELU. 21357); 
Curtis I.. Ba&s Strait, on P. cutulata {King, 
8.ii,l97I; MELU, 21358); and from Black- 
man's Bay (S. of Hobart), Tas., drift (Tyler. 
Oct (V) 1^73; ADU, A44234), 

The type of P. woolhousiae is a well 
developed specimen to 15 cm high, as in the 
Blackmail's Bay, Tas M specimen, whereas St 
Kilda specimens are 2-4 .-m huih, the Malla- 
coota specimens are only 1-2 cm high and 
Ihose from Bass Strait less than I cm high. 



the hitter appear to be juvenile All these, 
including the type, show similar cell structuie 
and presence of marginal growth on young 
parts and often also on mature thallL This 
marginal meristem is not apparent on the 
other southern Australian species or on most 
other species placed in Porphvra. A fragment 
of the type studied is not fertile, but the other 
specimens show eharaetcristic spennataogial 
groups which lend to deliquesce, and from 
marginal areas the contents of each cell appear 
to be liberated as a reproductive body. P t 
woolhousjae has n typical Potpfwta base with 
one or a few fronds attached by rhi/oids from 
the basal thallus cells, and in most features 
agrees well with Porphvra. If, however, it re 
confirmed that ihc carposporangia aie formed 
singly, then relationships wilh the genus Por- 
phyrella Smith & Moilcnbcrg (1943, p. 215) 
must he considered, though Conway & Wylic 
< 1972) have shown that the New Zealand 
Porphyra sitbtuntetis does not form packet's 
of carpusporanjjia, 

While it is desirable that mature plants on 
Matrocystix, corresponding to the type, be 
studied in detail, and their reproduction fol- 
lowed in cultuic, the other records arc 
sullicienlly similar to be placed under/*, woof- 
houxiac with some confidence. Most are epi- 
phytic on brown algae, cither on marginal 
spines or on spinous branchlets, and this hahi- 
lal may be characteristic for the species. 

P. w(fo(hoits{ae has been recorded from New 
Zealand (Lcvring 1955 T p. 412), followed by 

D. J. Chapman fl962, p. 129), V J, Chapman 
(1969, p. 20) and Adams (3972, p. 67). 
However Adams, following notes of E. Con- 
way in CHR* expresses doubt as to whether 
the New Zealand records arc not P. colnoj- 
bina in an abnormal habitat. 

The New Zealand specimens growing on 
Macrocyslis (and possibly Etklonia and Scyto- 
thutnnus) need further comparison with the 
Australian plants referred to P. noolhottsUic. 
Ones from Hokio Beach, l.evin. N.Z., on 
Macrocystis i Moore, 1 7 .xi. 1 946; CHR. 
55566). determined by Levring j |y55, p. 412) 
as /'. wnofhnusiac (accompanied by notes of 

E, Conway (1971) that they might be young 
plants of P. cohtmbitm), agree fairly well with 
P woolhousme as now known from the Aus- 
tralian plants. They are similar in form, in 
thickness and cell arrangement, and in having 
a liierisleinatie margin, but their reproduction 
is inadequately known. 



PORPHYRA AND PORPHYROPS/S 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 1 842: 1 4; 
1845: 33, pi. 9, rig. 2. J. Agardh 1883: 
70, pi. II, figs 65-66. V. J. Chapman: 
1969: 22. Dawson, Acleto and Foldvic 
1964: 32, pi. 615. Guiler 1952: 84. 
Hamel 1928: 51. Kuetzing 1849: 693; 
1869: 29, pi. 80e,f. Laing 1928: 39, figs 
1-7. Levring 1953: 464, figs 2-4; 1955: 
410; 1960: 29. Pujals 1963: 8. 

Wildemania columbina (Mont.) De Toni 1897: 

22. 

P. umbilicalis sensu Cribb 1954: 30. 
FIGS 3, 4 

Thallus fairly tough, varying from ribbon- 
like (Fig. 3/4), often with undulate margins, 
to broader forms, sometimes furcate, and 
usually umbilicate (by loss of upper parts and 
basal proliferation) (Fig. 35) 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 uffl thick, cells iso- 
diametric in section and with a single axile, 
pyrenoid-bearing, rhodoplast. Cells 10-15 una 
across in surface view (Fig. AA), isodiametric 
to slightly elongate, more or less in rows but 
becoming irregularly arranged, separated by a 
gelatinous matrix |-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. 4fi, 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 1. (D'Urvilfe). 
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. denticuiata Levring from southern 
Queensland is probably not distinct from P. 
columbina and represents the range extreme of 
P. columbina. 

P. columbina is the commonest intcrtidal 
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, 

7. 

P. umbilicalis sensu Guiler 1952: 84. Womers- 
ley 1950: 162. 

FIGS 5, 6 

Thallus usually fairly delicate, varying from 
lanceolate or ribbon-like (Fig. 5^4) to broadly 
ovate or cordate (Fig. 55), 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 8-1 5^on across in surface view (Fig. 
6A ) , mostly irregularly arranged, separated 
by gelatinous matrix i-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. 65, 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 "jfrisged" following shedding of sperma- 
tangial strips. 

Type locality: Bunbury, W. Aust. 

Type: Herb. Levring, Goteburg. (Isotype in 
ADU, A42700). 



64 



H. B. S. WOMERSLEY & ELSIE CONWAY 




\ 







,t x '«■> i 



C, Pi H.lt l.u*—*M~) 






ftnyVyffc. 


,jU-,1, 


«* i>m 




Uy 


La-»*1, 


S ft«u 






n,* < 


■J/Wrwl 






7 


! t*?J 


toll 


♦u* - 


(,«!»•* 


"'"1 



Fig. 3. Porphyra columbina. A. Ribbon-like forms, early winter. Little Dip, Robe, S. Aust. {W&mers- 
lev, 15. v. 1972; ADU, A42203). B. More umbilicate forms from spring. Cape Lannes, S. Aust. 
iiVomersley, 7.x. 1972; ADU, A42768). 



PORPHYRA AND PORPHYROPSIS IN SOUTHERN AUSTRALIA 



65 




lOOyum 






Fig. 4. Porphyra columbina. A. Vegetative cells. Nora Creina. S. Aust. KWomersley, 3.ix.l97I; ADU, 
A39559). B. Spermatangial marginal area (A39559). C. Area with carposporangial groups and 
young spermatangia (A39559). 



66 

A 



H. B. S. WOMERSLEY & ELSIE CONWAY 





J- 



/ 






*■%* 

* 



1 23*9«7»9W 



K 









B 



* J -a 





: 
y 





1 j ' ■ • ' I ' ' ' ' | ' ' : 1 1 1 1 1 1 : m 1 1 1 1 1 , | m , h 1 1 1 1 1 1 1 1 , . , . , 1 1 1 1 1 , i , t < 1 1 1 1 , r . 1 1 1 1 1 1 1 1 1 : , m 

"123456789 10 



Robe, S.dwlr. 

MCA 4«AtirwA t « V**j 

8 ■ x mz 

0»H "»-j>«a-. h. 8s.*d«?*.*-l*« 



Fig. 5. Porphyra lucasii. A. Ribbon-like forms. Port Stanvac, S. Aust. {Lewis, l.ix.1972; ADU, 
A42637). B. Broadly ovate forms showing spermatangial strips. Robe, S. Aust., in bay 
(Womersley, 8.X.1972; ADU, A42764). 



PORPHYRA AND PORPHYROPS/S IN SOUTHERN AUSTRALIA 67 






100/jm 






Fig. 6. Porphyra lucasii. A. Vegetative cells (ADU, A42764). fl. Edge of a spermatangial strip with 
vegetative and carposporangial cells on the right (A42764). C. Part of a spermatangial strip 
releasing spermatia above and with carposporangial groups below (A42764). 



68 



II D. S. WOMERSLEY & ELSIE CONWAY 



Distribution; From Cotiesloe, W. Aust. to 
Western Port Bay, Vic. and the north and east 
coasts of Tasmania, 

P. lucasii is the common Porphyta of calm 
to moderately rough waters, being replaced by 
P. eolutnbina 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 lucasii is found within calmer bays 
such as Port Phillip Bay and Western Port, 
Vic. ? whereas P. cotumbina occurs on rough- 
water coasts 

Genus PORPHYROPSiS Rosenvinge 
PorphyropsLs minuta sp. nov. 

Thallus epiphytic on certain Rhodophyta 
with a firm surface, developing from a sub- 
parenchymatous holdfast to form a hollow, 
sub-iphcncal to ovoid Madder (Fig. 1A ), later 
opening above to form irregular, monostro- 
matic membranes often with convolute mar- 
gins. Color greenish-brown to purplish, not 
jose-rcd. 

Bladders up to 1 (-2) mm across, torn 
membranes to 5 (-8) mm high and across. 
Cell divisions intercalary. Holdfast 50-11)0 
(-200) fjjn across, formed of irregularly 
arranged cells without rhizoidal extensions, 
Ceils of bladder (Fig. IB, C) often paired or 
in fours following division, and lying in rows 
more or Jess at tight angles, sometimes becom- 
ing irregularly arranged; membrane about 10 
f.tTY\ thick, cells 3-5 ju.m across and rounded to 
somewhat elongate in surface view, slightly 
ovate in sectional view. Rhodoplast apparently 
filling the cell, without a pyrenoid. 

Monosporangia (Fig- IB) formed from the 
whole contents of cells near the margin of the 
membranes, subspherical to ovoid, 5-7 ^m 
across. 

Thallus in Rhodophytis epiphyticus, solido 
superficie. ex subparenehvmato base ortus, 
primum vesicam cavam, subglobosam vel 
ovoideam formans deinde supcrne membranas 
nrcgulares monostromaticasque saepe margine 
convoluto producens. Color brunneo-viridis 
vel purpureas, sed nunquam carneus. 

Vesicae ad 1 (-2) mm latae, membranae 
laccratac ad 5 (-8) mm altae et latae. Divi- 
sura cellularum intercalans. Basis ad 50—100 
(-200) uffl lata. e\ cellulis sine projecturis 
rhizoideorum irregulariter composite fonnata 
est. Cellulae vesicae saepe binae vel quaternae 
post divisionem, seriatim plus minusve rec- 
tangulatae, tnterdum irregulariter compositae: 
merabrana circa 10 ^rn crassa. cellulis ad 3—5 



^ra Jatis et aspectu fromali globosis vel 
aliquantum elongatis, in sectione transversal! 
parum ovoideis. Rhodoplastus ut videtur, cellu- 
Jam complet, pyrenoide absenti. 

Monosporangia subglobosa vel ovoidea. 5-7 
^m lata, in cellulis prope marginem mem 
branarum formata. 

FIG 7 

Type locality: Pearson I., S. Auil.. on Ptero- 

cladia capillacea. upper sub-littoral (Spevht, 

I7.U.1960). 

Type: ADU, A24525. 

Distribution; From Garden I., W. Aust. 
around southern Australia to Bateman's Bay 
N.S.W., epiphytic on Pierocladia capillacea in 
upper sublittoral (and pools) on rough-water 
coasts, and occasionally on Plocamium attxus- 
tMftij P. tnertensii and Delisea pulchra in simi- 
lar habitats. 

P. minuta agrees well with P. coccinea in 
the holdfast, form and development of the 
bladder, and in reproducing apparently only 
by monosporangia. The cctls of P. minuta are 
similar in size to those in P, coccinea but art 
arranged in distinct rows more or less at right 
angles, in contrast to both P. coccinea from 
Europe (Rosenvinge 1909, p. 69, fig. 9) and 
P. coccinea var. dawsonU Hollenberg & 
Abbott (196S, p. 1239, % 5a-c) from Cali- 
fornia, where the cells are more irregularly 
arranged but are grouped into elongate, some- 
what lenticular patches. The life-history' 4tWj 
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 near the margins are 
liberated as monospores, whereas in P. 
coccinea the monospores are cut of! from the 
parent cell by a curved wall, a residual cell 
remaining when the monospore is liberated. 
Also, in P. coccinea some holdfast cells form 
rhizoidal extensions whereas this has not been 
observed in P. minuta. The colour of P. minuta 
is always a greenish-brown-purple, never rose- 
red as in P. coccinea 

The only other southern hemisphere record 
of Porphyropsu is by Adams (1972, p. 63) 
who reported P. coccinea var. dawsonii from 
New Zealand This taxon (e.g. CHR 248053 
from Kaikoura, N.Z.; Parsons, 13.xLl973) has 
numerous ligulate fronds from a clumped base, 
each with descending rhrzoids. It is not a Por- 
phyropsis* but more closely related trj Porphyra 
woolhoustiu*. 



PORPHYRA AND PORPHYROPSIS IN SOUTHERN AUSTRALIA 



69 




r « &gm m 



11 
# 



m m % 
4p #. | 









5 * * 5 S> TT-* a * • • • m J" • 

m m m § 1 , % w ^ i 

^ '■I MM *ZT " ^ & A 



50yji 



inn 







Fig. 7. Porphyropsis minuta. A. Small plants, on Pterocladia. B. Older membrane, with some cells re- 
leasing monosporangia. C. Cell arrangement. (All from the type, ADU, A42525.) 



70 



H. B. S. WOMHRSLEY & ELSIE CONWAY 



Acknowledgments 

We are grateful to Mrs S. C. Ducker (Uni- 
versity of Mclhoume) who rnade available 
collections for study, and 10 Dr P. Tyler (Uni- 



versity of Tasmania) who recollected P. wool- 
housiue. The first author acknowledges pro- 
vision of technical assistance by the Australian 
Research Grants Committee. 



References 



Adams, Nancy M. (.1972). — The murine algae of 

the Wellington area. A list of species. Rec. 

Dom. Mus. 8(5), 43-98. 
Agardh, J. G. U883). — Till algerncs systcmaiik. 

Nya bidrag. Vt Ulvacene. Acta Univ. land 

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

Chapman, D. J (1962). — A check list and key 

to the Rhodophyceae of New Zealand Sec- 
tion A: BangioiUeae. Trans, R. Soc. N.Z. 

Bof, L 127-137. 
Chapman, V. J. (1969). — The marine algae of 

New Zealand. Part lilt Rhodophyceae. 1. 

Bangiophycidac and FlorideOphycidae (Nema- 
lionales, Bonnemaisoniales, Gelidiales). 

I Cramer: Germany.) 
( . onway, Elsie ( 1964).— Autecological studies of 

the eenus Porphvra: I. The species in Britain. 

Brit. Phycol. Bull. 2(5), 342-348. 
Conway, Elsie, & WYUBj Ann P. ( 1972).— Spore 

organisation and reproductive modes in two 

species of Porphvra from New Zealand. Proc. 

VJt Int. Seaweed Symp., pp. TO5-107. (Univ. 

Tokyo Press; Tokyo.) 
Ckibu, A. B. (1954).— The algal vegetation of 

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

Tasm. 88, 1-44, Plates 1-10 
Dawson, E. Y., Acitro. C, & Foi.dvic, N- 

1 1964 J. — The Seaweeds of Peru. Nova tled- 

wit>ia 13, J- i 11, Plates 1-81. 
Da Tom, G. B. ( 1897)— -"Sylloge Algarum 

omnium hucusque CognitariunV\ Vol. 4. 

Florideae. Sect. !, pp. 1-388. (Padua.) 
Hi: Tom, G. B. (1924).— "Sylloge Algarum 

omnium hucusque Cognitarium". Vol. 6. 

Florideae, (Padua.) 
CituLER, E. R. 11952). — The marine algae of 

Tasmania. Check list with localities. Pap. 

Proc. R. Soc. Tasm. 86. 71-106. 
(.hiikr, E. R. ( 1954). — The recolonization of 

rock surfaces and the problem of succession. 

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

1-6. 
Hamel. G. (19281.— Sur quelques Porphyra des 

mers australes. Ann. Crvpt. E.xot. 1(1), 51- 

57. 
HarvlV. W. H. (1863). — "Phycologia Auslralica". 

Vol. 5, Plates 24I-3(K), synop, pp. 1-73. 
HoixtMBtun, G. J.| & Abbott, I. A. (1968). — 

New species of marine algae from Califor- 
nia. Can. J. Bot. 46, 1235-1251. 
KutrziNU, F, T. j 1849 ). — "Species Algarum". 

( Leipzig.) 
Kuetzing, F. T. ( 1 869 ) .—"Tabulae Phyco- 

logicat". Vol, 19. (Nordhausen.) 



Laino, R. M- (1928).— New Zealand Bangiales 

I Bang'ta, Porphyra, Erythrotricfua and ( ? ) 
Lrxtiirocladia). I rat is N^Z.. Inst 59, 33-59, 
Plates 1-15. 

LdVKiNG, T. (1953). — The marine algae of Aus- 
tralia. I. Rhodophyta Goniotrichales. Ban- 
giales and Nemalionales. Arkiv. for Botanik. 
Ser 2, 2(61, 457-530. 

Levrjng, T. ( 1955). — Contributions ft) the marine 
algae of New Zealand. I. Rhodophyta: 
Goniotrichales, Bangiales, Nemalionales and 
Bonnemaisoniales. Arkiv lor Bot. Ser. 2. 
3(11 ), 407-432. 

Lcvring, T. (I960). — Contributions to the marine 
algal flora of Chile. Lands Univ. Arssk. 
N.F. Avd. 2. 56(10). 1-85. 

Lucas, A. H. S. t.J 909). —Revised list of the 
Fucoidcac and Florideae of Australia. Proc. 
Linn. Soc. N.SW, 34, 9-60. 

Lucas. A. H. S. (1929).— The marine algae of 
Tasmania. Pap. Proc. R. Soc. Tasm. 1928. 
6-27. 

MoNVAUNk, C. (1842). — "Prodromus Generum 
Specierumque Phycearum Novarum In 
ltinere and Pol urn Antareiieum." (Paris.) 

Montaone., C. (1845). — "Voyage au Pole Sud 
et dans 1'Oceanie sur les Corvettes )' Astrolabe 
et la Zelce." I. Botaniqu". (Paris.) 

Murray, S. N., Dixon, P. S„ & Scon. J. L. 
J 972) . — The life history of Porphyrop\is 
ctHnnta var. dawsonii in culture. Br. phvcol. 
/. 7, 323-333. 

Pujals, Carmen (1963). — Catalogo de Rhodo- 
phyta citadas para la Argentina. Revta Mus. 
argent. Cienc. nat. Bernardino Rivadavia, Bot 
3(1). 1-139. 

RoshNviNot, L. K. (1909).— The marine algae 
of Denmark. Part 1. Introduction. Rhodo- 
phyceae. 1. (Bangiales and Nemalionales). 
SW/« Danske Videnskah. Sclskab. Biol. Skr.. 
7 Raekkf, AftL 7, M51, Plates 1, 2, 2 maps. 

SKorrsatRO. C. ( 1923). — Botanische Ergebnisse 
der schwedischen Expedition nach Patagonien 
und dem Feuerlande. 1907-1909. IX. Marine 
algae. 2. Rhodophyceae. K. svenska Vetensk- 
Akud. Handi. 63(8), 1-70. 

Smith, G, M., & Holl£NREpg. G. J. (1943). — 
On some Rhodophyceae from the Monterey 
Peninsula. California. Amcr. /. Bot. 30(3). 
211-222. 

WoMBRSitv. IT B S (1950) — The marine algae 
of Kangaroo Island. III. List of Species J. 
Trans. R. Soc. S. Aast. 73(2), 137-197. 

WoMKKStnv, H B. S, & Edmonds. S. J. (1958). 
— A general account of the intertidal ecology 
of South Australian coasts. Aust. J. mar. 
Fresh*. Res. 1(2). 217-260. Plates 1-12. 



THE VERTEBRAE OF FOUR AUSTRALIAN ELAPID SNAKES 

(SQUAMATA: ELAPIDAE) 

by Meredith J. Smith* 



Summary 

SMITH, MEREDITH J. (1975).-The vertebrae of four Australian elapid snakes (Squamata: 
Elapidae). Trans. R. Soc. & AusL 99(2), 71-84, 31 May, 1975. 

In vertebral morphology, the elapid species Pseudechis porphyriacus, Austrelaps superba y Notechis 
scutatus and Pseudonaja nuchalis conform with general descriptions and closely resemble each 
other. Features not previously noted are that epizygapophysial spines appeal* on the first 8 to 10 
vertebrae, and that bilaterally on every precloacal vertebra a foramen opens through the accessory 
process near the anterolateral edge of the prezygapophysial facet. No morphometric feature was 
found to completely separate any two genera, i.e. there was some overlap in the values of all the 
characters (ratios) studied. However, in general P. porphyriacus vertebrae are distinguished by their 
relatively long accessory processes, A. superba by their short accessory processes and lesser width 
across postzygapophyses, and N. scutatus by their greater width across postzygapophyses and 
shorter neural spines. P. nuchalis vertebrae have strong subcentral ridges 



THE VERTEBRAE OF FOUR AUSTRALIAN ELAPID SNAKES 
(SQUAMATA: ELAPIDAE) 

by Meredith J. Smith* 

Summary 

Smith, Mkkedith I. ( I97i)»— Ifia vertebrae of four Australian elapid snakes (Squamata: 
Elapidae). Tarns, H. Soc. $ Aush 99(2) ? 71-84, 31 May, 1975. 

In vertebral morphology, fhfl efapid species Pseudediis porphyriacus, Austreiaps super ba, 
Soiethis scutams and P&cudonuja nuchuiis conform with general descriptions and closely 
resemble each other. Features not previously noted are that epizygapophysial spines appear 
on the first 8 to 10 vertebrae, and that bilaterally on every precloacal vertebra a foramen 
opens Through the accessory process near the anterolateral edge of the prezygapophysial facet. 
No morphoinclric feature was found to completely separate any two genera, i.e. there was 
some overlap in the values of all the characters (ratios) studied. However, in general P> par' 
phyruicuM vertebrae are distinguished by their relatively long accessory processes, A. superha 
by their short accessory processes and lesser width across postzygapophyses t and N, scutuius 
by their greater width across postzygapophyscs and shorter neural spines. P. nuchaUs vertebrae 
have strong subcemr&l ridges. 

Fossil vertebrae from a Pleistocene deposit differ from P. porphyiacus, A. superba and 
N. sc.utatus but resemble Pseudonaja in most features, The fossils differ from modern P. 
nuchalis chiefly in having a thicker zygosphene and relatively wider postzygapophyses. As these 
fire features which develop with increasing size of vertebrae, the fossil vertebrae are assigned 
to the genus Pseudonaja, 



Introduction 

Although snake vertebrae have been 
recorded from Australian Pleistocene deposits 
(l.ydekkcr 1888, Merrilees 1968), the only 
species that has been identified is the Carpet 
Snake, Morel ia spilotes variegata Gray (~- 
Pyihon variegatux) from Marmor Quarry. 
Queensland (Longman 1925). No detailed 
studies have been published of Australian 
snake vertebrae and the diagnostic characters 
have not been established. The need to deter- 
mine the reptile fauna in a Pleistocene cave 
deposit stimulated the present study. 

The sixty-odd Australian species of clapids 
have been arranged in 29 genera (Worrell 
1963) or fewer (McDowell 1967, 1970), but 
only seven genera contain species with a 
recorded maximum length of over 0.9 ro. As 
centrum lengths of some of the Pleistocene 
vertebrae suggested that 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 to 



the whip snakes by Worrell (1963)|. Hopio- 
cephulus, and Oxyuranus been examined as 
iheir range is northern and eastern Australia 
(Worrell 1963). Specimens have been studied 
from the remaining four genera, dustretaps, 
Notechis, Pseudechis and Pseudonaja, which 
arc each represented in southern Australia by 
one or more common species. 

The exact column position of an isolated 
vertebra is impossible to determine (Johnson 
1 955 ) and in elaptds. with well-developed 
hypapophyses on all pre-cloacal vertebrae, 
division of the pre-cloacal column into regions 
is virtually impossible. Nevertheless vertebral 
shape changes along the column, and to iden- 
tify single vertebrae it is essential to know the 
range of variation within individuals and 
species. Auffenbcrg (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 
intracolumnar variation. Although Johnson 
(1955) measured 10 precloacal vertebrae at 
regular intervals along the column, he assumed 



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



72 



MEREDITH I. fiMJTH 



equnl variances i, between specimens) of ratios 
of these measurements and in his comparisons 
he used only the mean for each specimen, 
Here, enure vertebral columns have been 
examined in an attempt to assess the morpho- 
logical and murphornetrlc variation which 
occurs within individuals and species, and to 
find unique specific characters. As most elapids 
have over 200 vertebrae, the available samples 
are large and the time required for detailed 
examination of one individual snake precludes 
sampling many individuals. 

Because taxonomy of species of Fseudtmwt 
and Austrclaps is confused (Rawlmson 1969, 
Storr 1964) comparisons arc made ai the 
I'encric rather than species level. 

Materials and Methods 

A total of 2, 1 23 vertebnie from nine Recent 
specimens (Table I } and 556 Pleistocene ver- 
it:hr.:e were examined. Of the nine modern 
specimens, four specimens of Pxeudcchis 
pnrphyrior\ti, one PseiuhnnU* nachulis 
(RI4064) and one Notechix $cntalus 
(R14059) were collected near Armidale. New 
South Wales: the. one Austrclaps superba was 
collected at Umidla. South Australia. The 
localities of One P. vnchnfix (R 1 4065) and 
one jV. stututus (R 14058) arc unknown. 

The common brown snake. PscutJothij^ tex- 
tile (Dumcril & Bibron), has beco described 
as a separate species Jrorn the western brown 
snake. Psendonaja mtcholis Gunthcr. but It is 
not yet clear (Stnrr 1964) whether they are dis- 
tinct or are merely races of a single species, P. 
tcxtilh. One clearcut diagnostic feature is ihe 
shape of the nasal bones, which are anteriorly 
COnCHtVO on the lateral margins in P. ttuchulis 
but anteriorly convex in F. te.Kiilis (Worrell 
|9o_Vt. In R 1 4064 and R14065 the nasals arc 
anteriorly concave and hence are attributable 
to P nuchaUs. 

The cleaned skeletons were dried and com- 
pletely disarticulated for study. The specimens 
arc now lodged in the South Australian 
Museum, with register numbers as above. 

The Pleistocene vertebrae were excavated 
from an extensive bone deposit in Victoria 
Cave. Naracoorte. South Australia. The age 
of this deposit is unknown, but the abundance 
of extinct marsupials and the absence of 
remains of aboriginal man suggest that Ibc 
deposit accumulated during the Pleistocene but 
was sealed during Recent time. As the verte- 
brae were collected singly in many locations 
in the bone deposit, they probably represent 



at least several individuals. Visual examination 
of the fossils revealed that 454 of then) were 
similar to each other in shape, anil dixlincdy 
different from the rest, which were of seveial 
kind.s, Ihe latter, heterogeneous group will be 
discussed in a later paper. Of the larger group 
of 454 ihe HO most complete were examined 
in detail and measured in the same way as the 
modern vertebrae 

Because reputes grow ihroughuut their life. 
absolute dimensions of the vertebrae are ol 
little use in comparing individuals or species. 
Ratios between dimensions have been calcu- 
lated, the denominator being vertebra length 
in most comparisons. Mean values of ratios 
have been prepared independent ly for each 
individual snake, and arc given as mean, T. 
— standard error, followed in brackets by the 
number of vertebrae measured. The non-inde- 
pendence of measurements of different verte- 
brne of the one individual precludes statistical 
comparison of these samples (Siegel 1956) . 

Kroni a preliminary study of numerous 
characters of snake vertebrae, characters were 
selected that vary between species within the 
tamily Elapidae, exhibit low intracolutnnai 
variation and arc well preserved in fossils. To 
establish variation throughout the column, 
every fifth vertebra was measured in two speci- 
mens of each of P. porphyriucits, P. nuchalis 
and N. sctttatttx, and one A. xttpertuj. 

Descriptive techniques and terminology 
follow Aullcnherg (1961). Measurements were 
made to 0.1 mm with dial Calipers, as in Fie. 
I. 

pr-po Ijcngth between zygapophyses-distance 
between most anterior point of prczy- 
gapohysis to most posterior point of 
postzygapophysis. 
ap-ap Width across accessory processes-dis- 
tance between outermost tips of acces- 
sory processes, 
po-po Width across postzygapophyscs-dis- 
tance between outeimost points of the 
pnst/ygapophyxial facets, 
prl Length of prczygapophysis- the longest 
diameter of the pre/.ygapophysial facet 
feven though this was almost perpen- 
dicular to the long axis of the cen- 
trum). 
pr Width of prezygapophysis-tbe maxi- 

mum diameter at right angles to die 
length of the prezygapophysix (the 
pre/ygapophysial width being almost 
parallel to W*e lOrtg axis of the cen- 
trum y 



VERTEBRAE OF FOUR E1.APID SNAKES 



73 




poi 



po 



po-po 




Fik. J. Dorsal (D). ventral (V) and lateral (L) 
views of an elapid vertebra showing where 
measurements were taken. 



Length of postzygapophysis-the 
longest diameter of the poslzygapo- 
physial facet (even 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. 



nsl Minimum length of neural spine-this 
usually occurred about halfway up the 
spine, as the dorsal edge overhangs 
posteriorly and/or anteriorly. 

zw Width of zygosphene-the maximum 
width of the tenon. 

cw Width of condyle-the maximum dia- 

meter in the transverse plane. 

hyp Length of hypapophysis-the vertical 
distance from the lower edge of the 
condyle to the tip of the hypapophysis. 

Results 

The vertebrae conform with the general des- 
criptions of AulTenberg (1963), HorTstetter & 
Gasc (1969) and Johnson (1956). 

Pseudcchis porphyriacus (Shaw) 

The number of prccloacal vertebrae (Table 
1) is consistent with the number of ventral 
scales (184, according to Worrell 1963). 

In every precloacal vertebra, the width 
across postzygapophyses exceeds the length 
between zygapophyses (Fig. 2). The hypapo- 
physis arises near the lip of the cotyle, extends 
as a low lamella for about two-thirds the length 
of the centrum and then deepens sharply 
before tapering to a sharp point which does 
not much exceed the posterior surface of the 
condyle in any but the most anterior vertebrae. 
The hypapophyscs of the anterior vertebrae 
are very long; they decrease in length fairly 
uniformly along the column (Fig. 2). The 
subcentral ridges are low and rounded. The 
dorsal articular facet of the paradiapopbysis 
projects as a little round dome; the lower facet 
is saddleshaped. The prominent parapophysial 
processes are rounded anteriorly. They do not 
extend closer to the midline than the most 
lateral lip of the cotyle, Interzygapophysial 
ridges are faintly distinguishable. The neural 
spine is a low, laterally-compressed blade, its 
dorsal edge parallel with the long axis of the 
vertebra, its anterior edge almost vertical and 
its posterior edge overhanging. 

The minimum length of the neural spine 
is about half the length between the zygapo- 
physes [x 0.53 ± .0056 (35); x 0.55 ± .0049 
(36)] (Fig. 3). The neural arch 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 vertebrae and 
are distinguishable on the sixth to tenth. The 
cotyles and condyles are nearly round, but arc 
slightly flattened dorsoventrally. The width 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 





SAM 




Total 


Snout-vent 




Number o 


/ vertebrae 






register 




length 


length 


Pre- 




Post- 




Species 


number 
R14060 


Sex 


(mm) 


(mm) 


cloacal* 


Cloacal 


cloaca I 


Total* 


Pseudechis 


no data 


1220 


no data 


181 


8 


>42 


>231 


porphyriacus 


R14061 


no data 


1375 


no data 


182 


& 


52 


240 




R14062 


no data 


1120 


no data 


182 


5 


>30 


>217 




R14063 


no data 


1070 


890 


177 


5 


48 


231 


Austrelaps 


R14066 


2 


515 


433 


143 


4 


42 


isy 


superba 


















Notechis 


R14058 


c* 


930 


775 


174 


s 


S8 


237 


scuta tits 


R 14059 


no data 


no data 


800 


178 


5 


52 


235 


Pseudonaja 


R14064 


3 


1555 


1300 


204 


5 


60 


?70 


nuchal is 


R14065 


no data 


1100 


910 


206 


5 


62 


273 



* Atlas-axis complex not included in this count. 



10 



8 



•••••. 



• • 



• • 



o 



.' o°o°o * 

°i ** °°°£ a* • 

V A ° A AA fl AAAAA^ A AA AA % * . 

fcO A y " 



l=K 



Cloacal region 



** 



A • 



■ i 






■ ■ 



A 

o* a a 

o • 
o 



D DD u o 

a □ a d 



■ ■ I 



DD 



D a 



■ ■ ■ 

D D D D n 



"■",■■■ 

dDdddd 



J L 



50 



J I L 



200 



100 150 

Vertebra number 

Fig. 2. Variation throughout the vertebral column in (a) length between zygapophyses 
(A * J, (b) width across postzygapophyses (O • ) and (c) length of hypapophysis 
<□■) ot two specimens of Pseudechis porphyriacus. Hollow symbols R14060 solid 
symbols R 14061. 



250 



VERTEBRAE OF FOUR ELAPID SNAKES 



75 



20 
10- 

20 

10 




JL 

S3 R^k . 



I 



RU0S1 




■■^^^ 



5VojR14060 



10 



[Qfti 



20r 





JL 



2 or 

10 

30 

20h 



J. 




10" 



L m 



to 



I 1 1 I I I I I I 

1.0 11 12 13 
Rel. width 



' » ' i i ' i ' ' ' i 

1.0 1.1 12 1.3 W 15 
Ant. width: Post, width 



EL 

Sjct RW063 



J^ i 



D 



RU066 



g RK059 




R14Q64 



T^i Fossil 



i't 

03 0A 0.5 06 

Rel. length spine 



Fig. 



3. Distributions of values for the ratios (a) width across postzygapophyses to length 
between zygapophyses, (b) width across accessory processes lo width across post- 
zygapophyses, and (c) minimum length of neural spine to length between zyga- 
pophyses. Left-to-right downward hatching, Pseudechis porphyriacus; open columns, 
Austrelaps superba; right-to-left downward hatching, Notechis scutatits; solid columns. 
Pseudonaja michaiis; fine stipple, Victoria Cave, type A; coarse stipple, Victoria Cave, 
type B. 



7fi 



MEREDITH I. SMITH 



20 
10 

20p 

10 

20 

10- 

20r 

10- 




I 




40 r 

30 

20 

10 



w 



I'll 



I I i 



a* OS 0.6 07 
Zygosphene 




a 







J 



I L 



' I I 



0.2 0.3 M 
Condyle 



ris 



b 



Dfij, 




RU061 



|g RK060 



RU066 



%% RK059 

m 



RUOSfi 



RU065 



R14Q64 




Fossil 



■ '''■' 
0.1 0.2 0.3 
Postzygapophysis 



Fig, 4. Distributions of values for the ratios (a) width of zygosphene to length between 
zygapophyses, (b) width of condyle to length between zygapophyses, and (c) width of 
postzygapophysis to length between zygapophyses. Hatching as in Fig. 3. 



VERTEBRAE OF FOUR fcLAWl SNAKHS 



7T 



TABLE 2 

SiHtff c}}tTrmfm,\ii<* uf ttoC doc/cat /i\\iion ef four 
\)W.itfH'n.\ <>t pfi^rfitchis porphynactis 

Nu other Number 

Joiai with Number iffjtfl 

number of jrlicu- wilh nh* 

cloacal toteU single deeply 

Specimen tosrwbtae ribs hypypophyvib forked 



H 1 4060 


ft 


1 


T 


8 


R 14061 


* 


1 


3 


> 


RM062 


5 


4 


4 


A* 


ft 14063 


6 


2 


3 


51 



* TIii' amcnor-nrasi pair of rite have shallow fark*. 

between zveapophvses \x 0.33 ± 0047 (36); 
JIU? ± ".0067 (36)1 (Fig. 4). The zygo- 
sphcrte. viewed from the front, is drill and 
straight or slightly convex; from abuve it is 
nearly straight or slightly concave with * faini 
median notch on some vertebrae. The width 
of the zygosphene is a little over half the 
length between the zygapophyses [.? 0.53 ± 
.0063 (36); F0.57 * .OU38 (35)] Tfic prczy- 
gapophysial facets are oblong [length/ width 
I.11M.58, xl.40 x .019 1 36): 1.01-1 .73. f 
1.49 ± 025 (36)|. The acute accessory pro- 
cesses extend laterally perpendicular lu both 
vertical and horizontal 3\es of the vertehra, 
<and project well beyond the artiodar facets. 
Hence the ratio ot width across accessory pro- 
cesses in width across postzygapophyses is hieh 
[T I 39 ± ,014 |35); .v 1 .38 ± .Q10 (35)] 
(Fig. 3>. The postzygapophysc< are large 
[widen postzygapophysis/ length between 
2yjttpophy.se*: F0.2G ± .003 (3<i>; "0.20 ^ 
.002 |3b)| and have an anterior notch that 
gives their otherwise ovutc shape a kidney out- 
line. The maximum diameter of the post- 
zygapophysis is almost perpendicular to Ihe 
long axis of the vertebra and exceeds the width 
[length/width 1.03-1,51. T 1.27 ± .020 (3o); 
1.02-1.46. x J 22 = .017 (36) J. 

The typical four pairs of foramina are 
present on all precloacal vertebrae, and in 
many vertebrae they are unilaterally or 
bilatcially doubled, For example, in ihe I S3 
precloacal vertebrae of R 1 4061 the lateral 
foramina are bilaterally double (H vertebrae), 
double on the right only (II) or left only 
(13); the paracotylar foramina are bilaterally 
double (I), double on Ihe right only (5) or 
kit only <oM: rhe subccntral foramen is double 
on the right (2) or left <3) and in each of 
three vertebrae three pairs of subceniral fora- 
mriu appear. A fifth pail' of foramina is 
present in »he precloacal vertebrae of all speci- 



mens, each foramen of this pair opening 
through the accessory process near the antero- 
lateral margin of the prezygapophysial facet. 

Vertebral structure fn the cloacal region 
varies widely between specimens, and the num- 
ber of vertebrae with articulated forked ribs 
may be equal to or less than the number wiih 
n single hypapophysis (Tabic 21 The fork is 
deep on most forked ribs but on the rnosc 
anterior cloacal vertebra, the notch may be 
midway along the rib The lymphapophyses 
project laterally and slightly ventrally, as do the 
pleurapophyses of the post-ctoaeal vertebrae 

The haemapophyses of each post -cloacal 
vertebra arise separately from the ventral sur- 
face of the centrum, and remain completely 
separate, although on the anterior pusi-eloacal 
vertebrae the tips of the haemapophyses con- 
verge slightly. In lateral view the shape of the 
hacmapophysis is similar to thai of the hypapo- 
physe.s of the precloacal vertebrae The pleura- 
pophyses are directed anterolateral^* (viewed 
from above or below) and extend anterior to 
the cotyle. 

Some iiregularities occut in the skeletons. 
In R 14060 the right prezygapophysial facets of 
the J 30th 10 134th vertebras are enlarged lo 
almost double the si^e of the left-side facet. 
and the right poftlzygapophyse.s of these vcrte- 
brae are slightly enlarged. A similar, hut even 
greater, enlargement of the right prezvgapiv 
physis occurs in I Wo vertebrae of K 14062, 
where an outgrowth ot spongy bone has com- 
pletely fused the prezygapophysis to the pro- 
ceeding postzygapophysis. and the accessory 
process is reduced. These abnormalities prob- 
ably resulicd from injury, but R 14063 shows 
Slight congenital abnormalities, firstly on verte- 
brae 91. where a small, but distinct epi/.ygapo- 
physial spine appears on the left side only, and 
secondly in lour posl-cloacal vertebrae. Near 
the beginning of the pusicloacal scries, the 
centra of two consecutive vertebrae are com- 
pletely fused On the left, the outline of the 
posterior edge of the neural arch of the first 
vertebra can be seen, but on the right the 
suture hnc is only faintly distinguishable The 
pleurapophyses ul both vertebrae lie close 
together, apparently fused to the centrum ot 
the firsl verlebia. Two paijs of haem apophyses 
arise separately on the ventral surface uf the 
combined centrum. Later in the post-cloacal 
series two vertebrae are more fully fused and 
share a common neural spine Two pairs of 
haemapophyses are fused at their base near the 
condyle of the second vertebra. An apparently 



78 



MEREDITH I. SMITH 



8 




•••••.. 



•••• 



00 00 °o 00 



A *4 • 



o 
oA 
ft* 



6fe A 
o &6 



9 



* aa 4 a a 



.■■■■■■■■■■ 






°D 



^— I 2_i I D ?°° D PD 9D9 a qODODOD I , , i i I , i , , 



50 



100 150 200 

Vertebra number 



250 



Fig. 5. Variation throughout the vertebral column in la) length between zygapophyses 
(A* ), (b) width across zygapophyses (O*), and (c) length of hypapophysis 
(□W »" two species. Hollow symbols, Ausirdups superba (R14066), solid symbols. 
Notechis scutatia (R14058). 



similar abnormality has been observed in dor- 
sal vertebrae of colubrid snakes (King 1959). 

Austrelaps supcrba (Giinther) 

The low number of precJoacal 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 postzygapophyes 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. porphyriacus and does not extend 
further posteriorly than the posterior surface 
of the condyle. 

The subcentral ridges arc 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 



from above. The ratio of zygosphene width to 
length between zygapophyses (Fig. 4) has a 
mean of 0.50 ± .0038 (27). The prezygapo- 
physial facets 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 hacmapophyses of the posl- 
cloacal vertebrae are long anteroposterior^, 
about half the length of the centrum, and are 
completely double. 

Notechis scutatus (Peters) 

In all precloacal vertebrae, the width across 
the postzygapophyses exceeds the length 
between zygapophyses (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 













1 cm 



Fie. 6. Line drawings, to exact scale, of the 80th vertebra of (a) Pseudechis porphyriacus 
(R14060), (b) Notechis scutatus (R14059), and (c) Pseudonaja nuchalis (R14064) 
in dorsal, lateral, ventral, anterior and posterior views. 



so 



MLRHDrTH I. SMITH 



very long (Tig. 5.1. The suheenirnl ridges arc 
low and rounded: the inter/ygapophysiu) i'(d;ees 
are weak hut distinct. The neural spine is 
higher than in P. porphyriacm (Hip. 6) and jts 
horizontal dorsal edge overhangs boih 
anteriorly ami posteriorly The minimum 
lenglh of the ncur-il spine is generally relatively 
shorter than in the specimens of P, porphf- 
nacus [ratio of minimum length of neural 
spine lo length between ?ygapoph\scs: 7 0.4S 
± .0064 (35): x (X4fl =t .0049 (3n"l| (Fig. 3) 
Hpizygapophysial spines are welt deveJopcd on 
»hc first four vertebrae, faintly distinguishable 
on the fifih to eighth, but absent from all 
others. 

The round condyle is set on a short but 
d i mi net neck fh'ig. 6). The ihin /ygosphenc is 
straight or slightly convex from the front, 
siraighl or slightly concave from .above. The 
prezvgapophysial facets are oblong (length/ 
width I.OO-i.67, .7 1.47 =t .027 135); 1.09- 
1.58. x 1.35 ± .022 (35)]. The accessory pro- 
cesses are obtuse and short, so that the ratio 
of width across accessory processes to width 
across posuygapnphyses is generally less than 
in P parphyitKus (Fig. 3). The posuygapo- 
physcs arc obovate [length/ width 1.10-1.58, 7 
1.35 i= .021 135); 1.00-1.47, J 1.2S t 022 
< 35)1* and their width is about one-fifth of 
the lenglh between the zjgapophyses (Fig. 4), 
Prezvgapophysial foramina arc constantly 
present. 

Jn both specimens, all five cloacal vertebrae 
have fused forked ribs. One has three, the 
other two cloacal vertebrae with a single 
hypapophysis. 

The baemapophyses are large, paddle-shaped 
and completely paired. The plcurapophyses 
extend ventrally more than laterally. The 
/.yganophysial facets extend anteriorly and 
posteriorly, rather than laterally as in the pre- 
cloacal vertebrae, and acute nccessory pro- 
cesses ore distinct on all the postcloacal verte- 
brae. 

Unilateral or bilateral doubling of the para- 
cotylar and/ or subeetUial foramina occurs in 
a very lew prcctoacal vertebrae (no more than 
four In cither specimen). 

Pscudtuutju nucha lis Ciiinther 

Pwudonuiu nuchuJb has more vertebrae 
than any of tire other species studied here 
fTnble IK and maximum length and width 
occur rn sequentially more posterior vertebrae 
than in the other species <I : ig. 71. All pre- 
cloacal vertebrae aie wider tlian lonu The 



hypapophvsjs is well developed on all pre- 
cloacal vertehrae. Thickenings on the rim of 
the eotyle on either side of me ventral midline 
give rise to a low ridge lhai narrows sharply 
in join the laterally-comptessed h>papophysis 
The hypapophvsis deepens from about the 
middle of the vertebra and terminates in a 
rounded point thai extends posteriorly to the 
condyle. Subccntral ridges are strong and inter- 
zygapophysial ridges are distinct, lite para- 
diiipophyscN are well developed, with a pro- 
truding. dors< w cntrally-c!onsated upper facet 
uiid saddle-*haped lower facet. The parapo- 
physial processes appear in ventral view as a 
rial surface nearly as broad as long; the ventral 
projection of these processes is medial to the 
lateral border Of the eotyle. The neural spine 
is low and long (Tigs 3 and 6); its dorsal edge 
overhangs slightly at the front, markcdlv 
bchmd Cpizygapophysiai spines are distinct on 
the ftisi five vertehrae, family visible on the 
sixth to eighth and absent from all others, the 
eotyles and condyles are almost round; in 
R 1 4065 some eotyles are depressed slighily, 
but »n R 1 4064 some are slightly compressed 
laterally. The condyles are relatively much 
larger m the smaller specimen than in the 
larger one (Fig;. 4). The zygosphene is convex 
from the front, 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 [lenefh/ width 
1.09-1.72, J 1,39 - 022 (41); US-1.78. * 
1.35 ± .018 (41)|, and the acute accessory 
processes extend laterally well beyoad the 
articular facets (Rig, 3). Except fur a slight 
notch posteriorly, the outline of the post- 
zygapophysial facet is almost round [length/ 
width 1 OVI .35 5 1.17 ± .011 (41); 1.09- 
1.59. A 1.30 ± .019 (41)1. 

In the cloacal region the ribs arc all fused 
except for the anterior vertebra uf R 14064, 
and a hypapophvsis occurs on one (R14065) 
or three (R14064) cloacal vertebrae. 

The haemupnphyses arc completely paired 
and extend anteriorly as two separate ridges 
to the Jim of the eotyle The plcurapophyses 
are broad and flat and not pointed. In anterior 
view they project ventrolateral!}' but in ven- 
tral view mainly laterally and only slightly 
anlerinriy. 

In RN065, more than the one pair of para- 
coiyh,r foramina appear on many vertebrae — 
15 prccloacal vertebrae have an extra para- 
coiylar foramen on the right *ide, 1R on the 
left, and 34 on both sides. R|4flh4 has only 



VERTEBRAE OF FOUR ELAPID SNAKES 



»l 



10 



8 



• *• 



• • • 



• • 



AA 



• A 

A 



A AA4 A A A44A A*A* A A ## * 

AA A • 



\ 



G- 



Cloacal region 



• A 

A 



oo 



O oo 00oo oo oo 



A« 



ooo 



°0° 



*; 



o° aAA^A^**^ A fl %AAAAA A 
I O A^A 

- 8'° 
s 



*&*&.°0 



4% 



A 



A *■ 



"AA 
• A 



■ ■■ 

a 

D 



■ ■■ 



• ■ 



■ ■ 



on a D Q a° a o° 



DO 



■ » ■ 



oa 



■ ■■ 



■ - ■ 



«■ 



a □ u d d 



D D D D n D 



DD D n a D D 



A * A * 

A Aa 

o 
o »• 

o 

o 

o 



J . I L 



50 



100 150 

Vertebra number 



200 



250 



Fig. 7. Variation throughout the vertebral column in (a) length between zygapophyses ( A A ), 
(b) width across zygapophyses (0«), and (c) length of bypapophysis (D") "* 
Pseudotwja nuchalis. Solid symbols, R14064, hollow symbols, R14065, 



8 precloacal vertebrae with additional para- 
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, J 0.52 ± .0048 (40); B, x 
0.54 ± .0057 (40); .002 < P < .01]. 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. super ba; the ratio of width 



82 



MEREDITH ). SMITH 




I cm 




1 cm 





D 







Fig. 8. Precloacal vertebrae of Pseudonaja nuchalis; left, 80th precloacal of R 14064; right, 
Victoria Cave, Type B (P16126b) in dorsal (A, B), ventral (C, D) and anterior 
(E, F) views. 



VERTEHRAH Or FOtIR ELAPID SNAKHS 



83 



1*1 












« 


31 


- 












2S 














JfJ 










J* 


5*# 


£ .,. 










**£.% 




|U 


- 








a*V<s 




' 










J?i 


. 


TT 










M c 




? 










H <P 




'A 










XV 












ft' 






l' 






»k- 












.a 








a 






•i 














• • 








175 




i 


i 




i i 


1 1 1 1 1 






L 


5 




6 7 


B 9 >Q It (2 



Cer I n urn I ETTjrth 'mm) 

Kg. Double-logarithmic regression of width of 
posslzygapophysfc on length between post- 
zvgapaphyses in Pseudonaja nuchaVts, K 
R 14054; + f R14064; Q, fossiL Type A: 
• . fossil. Type B. 

across accessory processes to width across 
posCzygapophyses is less than in most P. par- 
phyr'iacus and the relative length of the neural 
s»pwe is generally greater than in N. scitiafus 
(Fig. 3). In these ratios, the Victoria Cave 
vertebrae closely resemble P. mtchalis. In the 
relative width of the condyle, the distribution 
of ratios for Victoria Cave vertebrae resembles 
the larger P. rwchalis (R 14064)., though not 
Lhe smaller (RI4065). Finally, in the relative 
width of the postzygapophyses the Victoria 
Cave vertebrae of Type A resemble the larger 
hut not the smaller P- nuchalis, and the relative 
width of the postzygapophysis is generally 
greater in Type B fossils than in any of the 
modern species studied I 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 6 vertebrae differ in the 
thick 2ygosphene and the relatively large ROSt- 



jygapophyses, the possibility exists that they 
are of a species different from Type A. How- 
ever, these characteristics are two which 
develop with age (Auffenberg 1963) and most 
of the Type B vertebrae are larger than the 
Type A (Type A t length 6.0-8,1, x 6.7; Type 
B, 7.1—1 1 . K x 8.9) and also generally arc 
larger than the vertebrae of the modern speci- 
mens. When the correlation between length 
between zygapophyses and relative width of 
postzygapophysis of Type B vertebrae was 
tested by the Kendall Rank Correlation Test 
(Siegel J956) 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 
fossil vertebrae (of both types, A and B) falls 
near the same straight line as those of R14064 
and R14065 (Fig. 9). The similarities between 
Victoria Cave preloacal vertebrae and those of 
modern P. nuchah's so far outweigh the slight 
differences that recognition of the fossils as a 
separate species seems unnecessary at least 
until the limits of variations of modern species 
arc better known. 

The post-cioacaJ vertebrae recovered from 
Victoria Cave resemble those ot P. nuvhulh 
in the laterally-directed plcurapophyses. In the 
other species studied the pleurapophyses pro- 
ject anterulatcraily. 

Discussion 

In all the vertebrae of the four species, the 
hypapophysxs is well-developed (as in all 
elapids) and hypapophysis length decreases 
slowly from anterior to posterior along the 
column. There is no suggestion of two distinct 
regions as in Aehrochordus juvanieus where 
the anterior region ( to vertebra 96 ) has 
hypapophyses long and fairly constant in 
length and the posterior region has hypapo- 
physes short and of constant length (Hotl- 
stetter & Gayrard 1964). 

At family level, the presence of vertebral 
foramina may have diagnostic value [e.g. the 
constant absence of lateral foramina in Achro- 
chordidac (HorTstetter & Gayrard 1964)1. but 
the variability in the number of foramina at 
each position (i,e< lateral, subeentral, etc.) in 
the one snake indicates the need for caution 
in the use of foramina in taxonomy. 

Although middle prccaudal vertebrae may 
be the most constant in their structure (within 
species) and hence best for identification 



84 



MEREDITH !. SMITH 



(Auffenberg J 963), the difficulty of assigning 
an isolated elapid vertebra to a particular 
region of the column precludes 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 
varies. 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 identified, but 
further studies of congeneric species are 
needed to determine whether specific identifi- 
cation is possible. 



Acknowledgments 

Mr R. Shine, Mr G. Whitten and Dr R. T. 
Wells kindly donated the modern snakes. Dr 
Wells and members of the Cave Exploration 
Group ot South Australia helped in excavating 
the fossil vertebrae. I am grateful to Dr R. I 
Wells, Mr I. M. Thomas. Dr T. F. Houston 
and Mr N. Pledge for their criticisms of the 
manuscript. Mr P. Kcmpsler prepared the 
photograDhs for Fig, 8. 



References 



Aufpenblrg, W. (1963). — The fossil snakes of 
Florida TuUme Stud. Zoot. I0 f 131-216.- 

Hoffstjctter, R., & Gasc\ J. P, ( 1 y<H9 ) .— Vertc- 
brae and ribs of modern reptiles, in C. Gans 
(Ed.) "Biologv of the Reptilia" Vol. 1, pp. 
201-310. 

HohFSTKTTKR, R., & Gayrakd, Y. (1964). — Obser- 
vations sur Tosteologie et la classification des 
Achrochordidae (Serpentes). Stilt, Mus. nutn. 
Hist. nat. t Paris (2), 36. 677-696. 

Johnson, R. G. (1955). — The adaptive and phylo- 
genetic significance of the vertebral form in 
snakes. Evolution 9, 367-388. 

Johnson, R. G. (1956). — The origin and evolu- 
tion of the venomous snakes. Evolution 10, 
56-65. 

King, W. (1959). — Vertebra duplication, an ostco- 
logical anomalv widespread in snakes. Her- 
petoloxica 15, 87-88. 

Longman, H. A. (1925). — Ophidian Vertebrae 
from cave deposits at M armor Quarry. Mem, 
Qld Mu.w 8, 1 11-112. 



Lydekxer, R, (1888). —"Catalogue of Fossil Rep- 
tiles and Amphibians in the Uritish Museum 
of Natural History." Part 1. (London.) 

iMcDowir.Lt, 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, Loud. 151, 
497-543. 

McDowell, S, B. (1970). — On the Hiatus and 
relationship of the Solomon Island elapid 
snakes. /. ZooL, Loud. 161, 145-190. 

Mekrileks, D. (1968),- — Man the destroyer: late 
Quaternary changes in the Australian mar- 
supial fauna- 7.7?. Sot- W. Aim. 51. 1-24. 

Rawlinson. P. A. (1969). — The rcptles of east 
Gippsland. Frot, R. Soc. Via. 82, 113-128. 

Siecel. S. (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. 

Worrell, b. (1963). -"Reptiles of Australia." 
(Angus and Robertson: Sydney.) 



THE ONTOGENY OF THE VOCAL SAC OF THE AUSTRALIAN 
LEPTODACTYLID FROG, LIMNODYNASTES TASMANIENSIS 

byM. 7. Tyler* 



Summary 

TYLER, M. J., (1975). -The ontogeny of the vocal sac of the Australian leptodactylid frog 

Limnodynastes tasmaniensis. Trans. R. Soc. S. Aust. 99(2), 85-87, 3 1 May, 1975. 

The ontogeny of the vocal sac of Limnodynastes tasmaniensis proceeds from initial bilateral 

evaginations of the mouth floor, through median fusion to a unilobular, submandibular structure. 

The acquisition of pigmentation by the submandibular skin is a concomitant process. It is suggested 

that the vocal sac evolved by a path now reflected by ontogeny, and involving progressive bilateral 

herniation. 



THE ONTOGENY OF THE VOCAL SAC OF THE AUSTRALIAN 
LEPTODACTYLID FROG, LIMNODYNASTES TASMANIENSIS 

by M. J. Tyler* 



Summary 

Tyi.er. M. J , (1975). — The ontogeny of the vocal sac of the Australian leplodaelylid frog 
Lirntiodynasfes tasmaniensis. Trans. R. Sac. S. Anst. 99(2). 85-87, 31 May, 1975. 
The ontogeny of the vocat sac of /.imnorfynanrs tasmaniensis proceeds from initial 
bilateral evayinations of the mouth Moot, through median fusion to a tmilohular, submandi- 
bular .■structure. The acquisition of pigmentation by the submandibular skin is a concomitant 
process. It is suggested that the vocal sac evolved by a path now reflected by ontogeny, and 
involving progressive bilateral herniation. 



Introduction 

Voca! sacs occur only in male anurans and. 
in most species, comprise inflatable, epi- 
thelium-lined chambers located 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 (Jngcr 1956; Inger & 
Circenberg 1956). The available information 
is so inadequate that the possible contribution 
of ontogeny to an understanding of the evolu- 
tion 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 frog* found in Aus- 
tralia and New Guinea 

The Australian leptodactylid species Limno- 
dynastes tasmaniensis represents an ideal 
initial 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 of Limnodytiastex 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 collected at 



Wesl Beach near Adelaide on 1 September 
1963 (SAM, R5290). The snout to vent length 
of the 1 8 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 ik Shear (1972), in order to 
differentiate the vocal sac from the surround- 
ing striated muscles. Muscle and vocal sae 
terminology follow thai of Tyler ( 1971 ). 

Observations 

Secondary Sexual Characteristics 

The secondary sexual characteristics of 
IJrnnodynastes tasmaniensis 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 finger*. Females bear 
broad lateral fringes to the first and second 
fingers, and sometimes to the third. 

Vocal Sac Ontogeny 

The earliest step in the progress towards the 
development of vocal sacs involves the forma- 
lion of a shallow and elongate involution of 
the floor of the mouth on one side of the 
tongue. There is a slight elliptical, ventral 
depression with a mediad inclination (Fig. 
I A), and the lateral margin of the depression 
is level with the lateral border of the anterior 
cornu of the hyoid. 



* South Australian Museum, North Terrace, Adelaide, S. Aust 5000. 



H 



M J tyler 




Fig. 1. Selected progressive stages in ontogeny of 
vocal sac. A. Single, elliptic ventral' de- 
pression, ti. Bilateral expansion mediad. 
C. Further medtad and initial caudad 
development. D. Media! unitv of separate 
sacs. 

In four specimens, development was con- 
lincd to such an evagjnation on the left side; 
in a fifth (here were bilateral evaginations. 
From these slight folds, the vocal sac develops 
bilaterally, and as tar us 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. 
IB). At this Stage of progress mediad develop- 
ment is more pronounced than anterior or pos- 
terior development. Simultaneously, that area 
of the mouth floor between the anterior cornii 
and the mandible becomes depressed, render- 
ing the aperture to each portion of the sac 
more conspicuous. 

As the two halves of the vocal sac approach 
one another, their posterior margins extend 
further caudad (Fig. 1C). This posterior en- 
farsement is accompanied by comparable 
enlargement of the interhyoiileus muscle into 
a slight lobe, extending beyond the post-articu- 
lar extremities of the mandibles. Ultimately 
the vocal .sac occupies the entire muscular 
lobe, becoming united medially by loss of the 
common medial wall (Fig. ID). 

The presence and extent of submandibular* 
derma) 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 evalua- 
tions in the mouth floor, the submandibular 
skin Was either entirely unpigmented 01 else 
bore a few scattered chromatophores at the 
periphery of the mandibles. Development of 
the evaginations was accompanied by an in- 
crease in the density of pigmentation and of 
its medial limit. The pigmentation, and the 
appearance of the hright yellowish background 
color of the submandibular skin, progressed 
in an identical sequence until, at completion 
of vocal sac development, the skin was entirely 
yellow. 



Discussion 

Beyond the sphere of its intrinsic interest, 
ontogeny can contribute to an understanding 
of the evolution of structure. In the present 
situation the progression of the vocal sac from 
paired, lateral evaginations to 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 vocal sac originates 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 on 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 probable steps that lead to the evolution 
oi vocal sacs in this species, or its ancestral 
stock t can be reconstructed quite readily 
Assuming an increase in the pressure of the 
buccal cavity during vocal activity, the existing 
sites ot I he vocal sac apertures are probably 
the areas of least resistance: in these regions 
the superficial tissue is of a rather elastic- 
nature, and presumably subject to the greatest 
distension. It follows that the first stage in the 
ontogeny of the vocal sac of Limnodynaxtes 
tasnnmiensis is precisely that initial event, Su& 
sequent stages could well have arisen from 
li'.lle more than progressive bilateral hernia- 
tion. 



ONTOGENY OF FROG VOCAL SAC 87 

References 

Bock, W. J., & Shear, C. R. (1972).— A staining Inger, R. F., & Greenberg, B. (1956).— Mor- 

method for gross dissection of vertebrate phology and seasonal development of sex 

, , , , n A i-ti i*v7 characters in two sympatnc African toads. 

muscles. Anat Anz. 130, 122-227. ; Morphm 9% 549-574. 

Inger, R. F. (1956).— Morphology and develop- Tyler, M. J. (1971).— Voluntary control of the 

ment of the vocal sac apparatus in the f sh ^ e ^Zt'^vff^*}*™!}* ^ A , US ~ 

„ . t * trahan leptodactylid frog Limnodynastes tas- 

Afncan frog Rana (Piychadena) porosissima maniensis. Trans. R. Soc, S. Aust. 95(1), 

Steindachner. /. Morph. 99, 57-72. 49-52. 



THE PRE-SETTLEMENT VEGETATION OF THE MT GAMBIER AREA, 

SOUTH AUSTRALIA 



BYR. 7. DODSON* 



Summary 

DODSON, J. R. (1975). -The Pre- Settlement Vegetation of the Mt Gambier area, South Australia. 
Trans. R. Soc. S.Aust 99(2), 89-92, 31 May, 1975. 

There are some conflicts as to the nature of the pre- settlement vegetation formations around 
Mt Gambier and Glencoe. European settlers long ago cleared the areas of their vegetation cover. 
Pollen analysis of Brownes Lake sediment reveals that the most likely formation around 
Mt Gambier consisted of open grassland with perhaps a sparse cover of woody taxa. 



THE PRE-SETTLEMENT VEGETATION OF THE MT GAMBIER AREA. 

SOUTH AUSTRALIA 

by J. R. Dodson* 



Summary 

Dooson, J. R. (1975). — The Pre-Settlement Vegetation of the Mt Gambler area. South Aus- 
tralia. Trans. R< Soc. S, Aust. 99(2), 89-92, 31 May. 1975. 

There are some conflicts as lo the nature of the pre-settlcment vegetation formation* 
around Mt Gambier and Gleneoe. European settlors long ago cleared Ihe areas of their 
vegetation cover. 

Pollen analysis of Brownes Lake sediment reveals that Ihc most Likely formation around 
Ml Gambier consisted of open grassland with perhaps a sparse cover of woody taxa. 



Introduction 

Crocker (1944) left gaps in his veget;uion 
map in the areas around Mt Gambier and 
Gleneoe, in south-eastern- South Australia, as 
Ihe original vegetation was no longer evident 
at the time of his survey. Crocker ( 1944) and 
liver & Crocker (1951) hypothesized that the 
areas were occupied by lightly wooded grass- 
land, and the trees cleared after settlement- 
Woods ( 1562) recorded the vegetation in 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. L08) taken in 1860 ancfyct Hill 
when describing an early record < 1861 ) of the 
first road between Mt Gambier (then Gam- 
biettown) and Port McDonnell on the coast 
states (p. 109) 

"the route to the 'Bay* was through dense 
bush counlry, mud and slush in winter, dust 
in summer, and tenanted by thousands of 
kangaroos at all times." 

vSpecht (1972, p. 203) in his vegetation map 
of the South East simply records the areas 
around Mt Gambier and Gleneoe as cleared. 

Brownes Lake 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). 
It is 4-5 m deep, sits on collapsed volcanic 
debris and its water surface, like those 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- 
tacular observed were in Brownes 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 erected in 1841 on the site 
of Brownes Lake, Brownes Lake and Valley 
Lake filled and joined and ?be new lake 
reached its maximum depth in the 1890*s. 

This paper provides data on early vegetation 
at Mt Gambier. Pollen analysis is an ideal 
method for tackling this problem as the craier 
walls are mostly steep-sided am! can support 
Jiirlc local vegetation. Therefore a significant 
proportion of the pollen rain is probably 
regionally derived. Today the craters arc 
heavily exploited for recreation purposes and 
most of the vegetation 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 w«is 
selecied 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 Biogcography and Geomorphology. Australian National University, Canberra, ACT 
'2600. Present address: Geography Department. University of Canterbury, Christchurch, New Zealand 



90 



J. R. DODSON 



Depth err; 
Restionaceae 



Cyperaceae 



-\Unophyiiwn 



g 



Trigtochm ° r 

Pvfumngetim *n J- 

ftappw ° r 

/.v/j/w ° r 

Z, epitaenct 



E 



Hygtrichospheres 






t 



i — r 



i I 



Poaceae 



Unidentified pollen 
Ph'lhluuii 



n 



l.yuiftotfiitw 
Monolete spores 



htnuiypiui Type 1 ° 
Litaitypiux lype 
l.iii'tiivpius iype4 ^ j 



* it 



Myiaceae Type 1 8 
Myrtaceae Type2 ^ 

Cuptessacaae P r 

;_J1 L 

Cusmiiina (,>28^nVl S r 
J* L 
( '(j\thJritui fe28pm) ° 



:c 



Acacnti ° 
Chenopodiaceae 

As;erac«ae (Tuoiliflorae) s 



Asleraceae (uguhflnrafi) a 

t paendaceae 
Plan/two ititneohhi ^ 



Brassicaceae 



Liliaceae u r 
Polyganaceae 



Haioragaceae 

Pimdia 
Gyrostemonaceae 

M^lva^eae 
Slylidiwn 

Restionaceaft 



*7^ 





m 



H 



-n-n 



:et— 



Fig. 1. Pollen diagram for Brown es Lake. 



I \«LY MT GAMB1ER VEGETATION 



:> 



suitable for pollen analysis. No suitable sites 
we-.ret found &JL the accessible ureas in Vallev 
Lake nor in much or the Brownes lake area. 

In February 1*574, a 45 cm core was 
collected with a D-vection sampler from the 
core site on the eastern shore of lirownes Lake. 
It consisted of 30 cm at black lake mud over- 
ling 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 out in the labora- 
tory using the standard hydrofluoric acid. 
alkali and Erdtmnn's acetylosis methods as 
described by Faegii & iversen (1964). Resi- 
dues containing pollen were dehydrated. 
mounted in silicone oil. and counted until at 
least 200 pollen grains of woody taxa had been 
recorded. Relative peivrentages for terrestrial 
pollen and spore taxa were calculated against 
r pollen sum of total land plant pollen exclud- 
ing ihe recorded introduced taxa (Pimts and 
PlcwtagO hmceolata). Frequencies for aquatic 
vascular plant pollen and Fystrichosphere 
remnins were calculated aguinsl a pollen sum 
of total aquatic pollen. The results were 
plotted on a pollen diagram 'Fig. 1). Ecolo- 
gical information for taxa in the study area 
and details of pollen identification have been 
given hy Dodson (1974). 

Results' and Discussion 

The short pollen diagram (Fig. 1) has not 
been divided into zones, but the presence of 
introduced taxa divides the diagram into post- 
seUlemciu (0-10 cms) and pre-settlement 
phases (20-45 cms). 

The prc-scttlement phase was dominated by 
Astcraccac (Tubulillorae). Poaceae and 
Pteridium, with small numbers of herb pollen, 
Tree pollen was virtually absent, although 
small and increasing frequencies of scrub taxa 
( Ctisnwmct ( ^ 2$p$i ) -pi'obably C. pehtdosa. 
and Mvrtaccac Type i-Lepm$penvitr*i juni- 
perinum and l.eptQspermum myrsineides) were 
obtained. Assuming that regional pollen 
dominates, then the assemblage most likely 
represents open (treeless) vegetation in the 



region wi;h a heath fringe around the lake. The 
local vegetation was mainly Amhoeeros, 
Cypcraceae, and Afyriophylftwh representing a 
shallow water environment at the core site 
This pollen assemblage tends 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 
Acactu. There is no evidence in the form of 
remnant vegetation to support this, The organic 
and pollen and spore content of ihe sediment 
was low (except for Atuhoceros spores which 
must have been derived locally), suggesting 
cither slow sedimentation during alternating 
wot and dry conditions (which could result in 
the loss of pollen through oxidation) or fairly 
rapid in-wash of inorganic material from the 
steep crater walls. On the evidence presented 
here it is not possible to favour either explana- 
tion. 

The post-settlement phase of the pollen dia- 
gram is dominated hy Poaceae, herbs and 
Phws ntdiata. and also shows increasing Euca- 
Ivp/ttx frequencies. Antlwceros and Cyperaeeac 
decrease in importance and Myuophyllu/n 
dominates the aquatic spectra, indicating a 
change to deeper water at the core site. Since 
the core site is near the edge ot' the lake, it 
follow? that water would not be deep there 
until the level rose above its present position. 
Since the rise accompanies die increase in 
pollen of exotic plants, it seems likely that the 
rise is the one recorded for the latter part of 
the i9th Century when Rrownes 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 
to any change in the naiive vegetation 

Acknowledgments 

It is a pleasure to lhank Gurdip Singh and 
Joan Guppy who critically rend the manu- 
script, and the Australian National LTniversity 
for supporting the work with finance and 
equipment. 



References 



Kwtv, 1. A. E., & Williams, W. D. (1964).-- 
Chemical and biological observations on 
some volcanic lakes in the soulh-easi of South 
Australia. Ausi. J, Mar, t-t^sh\fjr. A'c*. 15, 
123-132. 



TUylv. L A. C, & Williams. W. IX <i!>6n) L — 

Further chemical observations on >*ime vol- 
canic lakes t\l the south-east of South Aus- 
tralia. Ausl. J. Mar. Frvthwut. Res. 17, 229- 
237, 



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. /. Sci. 29, 50-52. 

Crocker, R. L. (1 944 ) . — Soil and vegetation 
relationships in the Lower South-East of 
South Australia. A study in ecology. Trans. 
R. Soc. S. Aust. 68, 144-171. 

Dodson, J. R. (1974). — Vegetation history and 
water level fluctuations at Lake Leake, south- 
eastern South Australia. I. 10,000 B.P. to 
Present. Aust. J. Bot. 22, 719-741. 

Faegri, K., & Iversen, J. ( 1964).— "Textbook of 
Pollen Analysis." Edn 2. (Munksgaard: 
Copenhagen.) 



Fergusson, G. J., & Rafter, T. A. (1957). — New 
Zealand 14 C age measurements — 3. N.Z. J. 
Sci. & Tech. 38B, 732-749. 

Hill, L. R. (1972).— "Mount Gambier— the City 
around a Cave." (Investigator Press: Ade- 
laide.) 

Speciit, 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. /. British Grassland Soc. 6, 29-80. 

Woods, J. E. T. (1862).— "Geological Observa- 
tions in South Australia." (Longman: Lon- 
don.) 



AUSTRALIAN LEPTODACTYLID FROGS OF THE 
CYCLORANA AUSTRALIS COMPLEX 

byM. 7. Tyler* and A. A. Martin f 



Summary 

TYLER, M. J., & MARTIN, A. A. (1975). -Australian leptodactylid frogs of the Cyclorana australis 
complex. Trans. R. Soc. S. Aust. 99(2), 93-99, 31 May, 1975. 

Cyclorana australis as now defined is shown to comprise two closely related species: C. australis 
confined to northern Australia and C. novaehollandiae to eastern Australia. Notes are provided on 
the tadpole of C. australis, and the calls of both species are analysed. Call divergence is so limited 
that hybridization is considered possible in sympatry. 



\USTRALL\N LEPTODACTYLID FROGS OF THE 

CYCLORANA AUSTRALIS COMPLEX 



by M. {» Tyler* and A. A, Martin! 

Summary 

lYLtK, M. L & Martin, A. A. (J 975). —Australian Icptodactylid frogs of the Cyvtvnwa 
atistralis complex. Trans. #• Soc. S. Aust. 99(2), 93-99, 31 May, 1975. 
Cyctorana australis as now defined is shown to comprise two closely related species: 
C. australis confined to northern Australia and C. ttovaeholkmdiae to eastern Australia. Notes 
are provided on the tadpole of C. tiustruiis, and the calls of both species are analysed. Call 
divergence in so limited that hybridization is considered possible in sympatry. 



Introduction 

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 signifcm as recog- 
nised by Parker (1940) is now known to be 
a complex of seven species (Moore 1954; 
Lilflcjohn 1957; Main 1957; Straughan & 
Main 1966; Tyler & Parker 1974): Mixophyes 
fasriolaius is now four species (Straughan 
1968) and Limnodynastes dorsalis is also lour 
| Martin 1972). 

The most neglected leptodactylid genus Is 
Cyctorana, of which the type species is A fates 
australis Gray (1842), described from material 
collected in the Northern Territory. This 
species, as currently defined, extends from 
northern Western Australia to northern New 
South Wales: a geographic range of approxi- 
mately 3500 km. The conspeeificity of indi- 
viduals from the extremes of this extensive 
range is obviously suspect, and even the most 
cursoiy comparison of specimens of C. aus- 
tralis from the Northern Territory and 
northern Western Australia with those from 
Queensland reveals striking differences between 
them. The northern individuals lend io have 
a rather elongated head* a distinct, dark rostral 
stripe and a narrow suhoeular ban Tn 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- 



hvlhmdiae by Steindachner (1867), and as 
Phractons ahttaceus by Peters (1567). Both 
names were referred to the synonmy of aus- 
tralis by Boulenger (1882), 

We have assembled and examined large col- 
lect ions- of C 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 arc deposited in the 
following institutions: National Museum of 
Victoria (NMV): Naturhistoriska Riksmuseet. 
Stockholm (NR); Department of /oology. 
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 posterioi 
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 between 



* South Australian Museum, North Terrace, Adelaide, S. Ausl. 5000. 

t Department of Zoology, University of Melbourne, Parkville. Vic. 3052. 



94 



M. J. TYLER A A. A. MAfcTlN 



the anterior tip of the snout and the anterior 
margin of the cloaca). 

Ratios calculated and subjected 10 the Stu- 
dent (-test were TL/S-V, HL'HW. F/S-V, 
F/TL and S-V/HW, Larval stage numbers 
follow those employed by Gosner (I960). 

Mating calls were recorded in the Meld using 
a Uher 4000 Report portable tape recorder 
and Beyer M69 dynamic microphone, af a tape 
vpeed of 19 era /sec. Calls were analysed by 
use of a sound spectrograph (Kay Model 
f»061-A Sona-Graph) with the overall res- 
ponse curve maintained in ihe FL-I position, 
Three calls of each individual were analysed 
and mean values calculated. Kach call was. 
analysed twice; a narrow-band 145 Hi band- 
pass) analysis at recording speed tn determine 
duration and dominant frequency, and i wide- 
band (300 Hz bandpass) analysis at half 
recording speed to resolve fundamental fre- 
quency. 

The Cjclorana anstralis complex 

Frogs of the Cyctorana austratrs complex 
are relatively large animals; the snout to vent 
length of adult males ranges from 61,4 to 81.4 
rnm. and thai of females from 69.9 to 102 
mm. They are all generally robust with a broad 
and frequently bloated body and relatively 
short limbs (TL/S-V range * 0.34-0,46)* 
All members ol rhe complex exhibit exostosis 




Fig. 1. Geographic distribution of ihe frogs of the 
CyLloruita australis complex. Circles - 
C. atosttiilis: triangles = C. tiovaehollan- 
diae. Closed symbols indicate sites of The 
material examined, and the open circles 
one of ihe following literature references- 
Braltsu-om 1 1 970 ) . Loveridee ( 1 93.S ) , 
Moore (!9filr t Parser 1 1940). SloVXn 



of the maxillary, premaxillary, nasal, fronto- 
parietal and squamosal bones. On the dorso- 
lateral body surfaces there are continuous or 
disrupted, longitudinally orientated skin folds 
commencing behind the skull, and terminating 
above the groin. 

Nothing is known of the breeding biology 
of the members of the complex, but they are 
probably opportunistic breeders. The eggs ate 
small and pigmented (ovidiametors of ovi- 
ducnl eggs range from 1.1 to 1.3 mm), and the 
tadpole (one species) is of the hylid type with 
Iwo upper and three lower rows of labial teeth, 
an acuminate tail tip and a median or slightly 
de^tral anus. 

The geographic range of the complex ex- 
tends from northern Western Australia to 
northern New vSouth Wales (Fig. I ), Of what 
we demonstrate to be two component species, 
C. australix (sanm strirta) occurs in northern 
Western Australia, the Northern Territory and 
northern Queensland lo the west of the Divid- 
ing Range; C. ri<n<aehoIhific}uw is found 
throughout Queensland and extends as far 
south as the northern part of New South 
Wales. 

Cycloraiu* australis (Gray). 
FIG. 2A 

Atytt's uusiraHs Gray ( 1842). 

Chitn/eptes tiustralis, Gunihcr (1858); Boulen- 
Ber (1882) (part). 

Phruitvps amtrults, Fry (1914). 
Cheirokptes aitxtralis, Spencer (1901) (replace- 
ment name tor Cftirotepies) 
Cyclortttut austntlix, Parker (1940) (part). 
Type locality "North coast of Australia". <l\>ri 
Essington, Northern Territory.) 

Material examined; Westrrn A ustralia — -W AM. 
R8732. Carlton Reach, Ord River; WAM. 
R4306 7, Crystal Ck; WAM, R 1 558-5*), Dryttdaic 
River Mission; WAM. RZ1Z33, Kownl Down*; 
WAM, R1377. 43282-86, 42399-42422, Knfum- 
buru; WAM, R22369-75, Kimberley Rcscaich 
Stn; SAM., R4769-70, R5O70. Kummurra; WAM, 
M 654-57, Landar Stn; WAM, R423X7, HO km S 
of La Grange; WAM, K4253G-40. 1238 1, 43478. 
43491, Mitchell IMalcau; WAM, R42530-35, Main 
Ord River Dam Site (spillway): WAM, R42424, 
Mt Hait; WAM. R32099. Ml Anderson; WAM, 
R32291. Ml Barnell; NR, 1562, Mowh. Down; 
WAM, KI3726, Oscar Ranges; NMV, D2J54-55, 
Km George IV; WAM. R32I49, Si George Range. 
WAM, Rl 1208, R 1 1894, RI2332, Woljulum: 
WAM, R26769-70. Point Springs. Webber Rang*; 
WAM. R32351A, Wvndftam; WAM, R2*093. 40 
km ST- of Wyndhom; WAM, R2O307. Yeda Cross- 
ing. Northern Territory — SAM, R14.VU, AttMnide 
River; NMV. DI2702. Harrow Ck. NMV. 
D8307, DXJI5. DS327. QM. J17K5. 2985, SAM. 



FROGS OF THE CYCLORANA AVSTRALIS COMPLEX 



95 




20mm 




20mm 



Fig. 2. A. Cycloruna uustralis from Kununurra, W.A. B. Cyciorana tiovaeholliwdiue, 15 km 
N of Goondiwindi, Qld. 



R8968, Darwin; SAM, RI3453, Elsey; SAM. 
R 13450, Howard Springs; WAM, R 1935-36. 
R21318, SAM. R14330-31. Katherine; SAM, 
R4877, Mt Bundy Stn; SAM, RI3349 A-G, Smith 
Pt, Coburg Peninsula; WAM. R24007. Snake Ck; 
NMV, DI2704-08, SAM, R13275 A-L. Tennant 
Creek; NTM, 498, 525-26. 50 km N of Tennant 
Creek. Queensland— SAM, R5010, R5070. Doom- 
adgee Stn; NMV, 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). 



96 



M. I. IYII.R & A. A. MARTIN 



Development and exostosis of the super- 
ficial skull bones are moderate in this species. 
The dorsal limit of the squamosal is such that 
I he re is a very hroad gap between the squamo- 
sal and the frontoparietal. On the fronto- 
parietal exostosis is confined to the lateral bor- 
ders of the bone. The sub-orbital portion of (he 
maxilla slopes steeply to the labial margin and 
is not expanded there into a lateral ridge. 

Cyclvtuna australis is usually pale olive or 
grey in preservative and bears a narrow and 
very sharply demarcated dark brown rostral 
hnr and a narrow sub-ocular bar which ter- 
minates far above the labial margin. The 
lateral body surfaces are commonly heavily 
.suffused with darker pigment. The backs of the 
thighs arc darker and densely variegated with 
light pigment. 

Geographic variation: The presence of darker 
irregular patches on the dorsum vanes through 
the range. Dorsal spots are absent from a series 
of over 100 specimens from kununurra. Speci- 
mens from Tcnnant Creek have light suffusions 
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 I. 
bggs; A gravid female from Kununurra con- 
tained approximately 1000 eggs varying from 
1.1 to 1.3 mm in diameter. The eges have 
black animal poles. 

fMrval morphology: A series at tadpoles was 
obtained at Kununurra on the Ord River by K. 




10mm 




1mm 
Fig. 3. Tadpole and tadpole mouthparis ui 
Cyctoruna ausiralh. 

Cole in February. 1 963. The following notes 
are based on four specimens from this series 
at stages 36-3*5. 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 ('. 
cttlrt'ipts and f . phvycL-phalus (Watson & Mar- 
tin 1973). 



TABLE I 

Geographic variation in prono) lions of Cyclorana species 
[Ranges are ^ivet) with means and standard deviations n> porenthews) 



Sf/et h's und Ii>iiii't1\ 



HL'UW 



TL/S-V 



S-V.1HW 



l/TL 



FS\ 



C. aiistniliv 
kiinumirru. W.A. 


7 


0.81 - 0.93 
(0.86 _!_ 0.04) 


0.3K 0.45 
(0.42 ±0.03) 


2.13 — 2.33 
(2.23 ± 0,09 J 


0.9 1 1.00 
(0.97 ±0.03) 


0.38- 0.44 
(0.41 ±0.02) 


Smith Point, N.T. 


7 


0.K9 0.93 
(0.91 ± 0.0 1) 


0.40 0.45 

(0.43 ±0.02) 


2.27 2A4 
<2 .36 ±0.06) 


0.92 - 0.98 
(0.95 ±0.02) 


0.39 - 0,4} 
(0.41 ±001) 


lennant Greek. N T- 


7 


0.87 0.94 
(0.90 ±0.02) 


0.40 — 0.46 

(0.42±n.o:'> 


2.25 2.49 
12.36- ±0.08) 


0.91 - 0.98 
(0,94 ±0.02) 


0,38 - 0.42 
(0.39 ±0.01) 


Milctiell Pliileiui, W.A. 


4 


0.95 — 0.99 
(0.97 ±0.01) 


0.44 — 0..MJ 
(0.46 ±0.02) 


2.26 - 2.51 
(2.41 ±0.U) 


0.88 - 1.05 
(0.95 ± 0.07) 


0.38 0.43 
(0.41 ±0.01) 


C\ nayavliidlundiuc 
Cooktown. QUI 


7 


0.8 1 0.8* 
(0.83 ±0.02) 


039 U.43 
(0.41 sfcJXQl) 


I.V3 2.08 
(2.02 ±0.04) 


0.87 - 0.96 
(0.91 ±0.03) 


0.36 — 0.41 
(0.38 ±0.01) 


Calliope, Qld 


J 


0.82 — 0.KK 
(0,86 ±0.03) 


0.39 — 0.40 
(0.39 ±0.01) 


USA - 2.17 
(2.12 ±0.01) 


0.R9 - 0-94 
(0.92 ±0.02) 


0.34 0.39 
(0.36 ±0.01) 


Cunamutlu. Old 


5 


0.78 — 0.85 
(0.82 ±0.03) 


0.34 0.41 
(0.37 ±0.02) 


1,97 - 2.13 
(2.05 ±0.06) 


0.93 102 
(0.96 ±0.03) 


0.32 0.39 
(0.36 ±0.03) 



I- RUGS OF IHh craMH4S>\ .tVSTRAUS COMPLEX 



9? 



The mouth is subicmunal (Tit;. 3) with a 
large homy beak and papillae around the sides 
und bock ol the mouih disc. There are two 
upper rows of labial teeth, the second dis- 
rupted medially, and three lower labial rows 
of which the first is similarly disrupted. 
Mating Cali: Calls of five individuals were 
recorded on 13 xii. 1 971 • 14 km C of Daly 
Waters, N.T, The frogs were Calling on land 
beside a small water-filled channel Wet-hulb 
air temperature at ihe calling sites was 24.1 'C. 
The call is a short, well-tuned nolo repealed in 
long sequences. The mean call duration is 152 
msec (range 122-204). The fundamental fre- 
quency is 199 Hz (range 183-20^>. but con- 
tains little energy: most of the energy is in :be 
third, fourth and fifth harmonics (about 600, 
800 and 1.000 Hz I which arc approximately 
equally emphasized. 

Geographic Range: Cyclomna mtxtralis extends 
I rem the Kamberley District of northern Wes- 
tern Australia In the Gulf District of Queens- 
land \n the Kiraherleys it is clearly widely 
distributed and in the Northern Territory it 
extend*, as fai inland as Barrow Creek. 
Absence in the north-western portion of the 
Northern Territory may simply reflect inade- 
quate sampling, 

Cyclorana novaehollandiue Stcindachner. 

Cychmma novut'ltoUtindine Stcindaclmei MS67). 

Pftractojus ulutuvcHS PtWn l L#G7h 

ChirohpU's auai raits, Bou lenger i 1882) ipani. 

Phravtops uuMnttis, Lovehdj;? (IM35J- 

CvcJoranu attxtruiis Paikcr (]<M0) <part): 

Moore (I96t>. plate 35. Fig. 2. 
Vv/v tocnitrt: Rockhatnpton, Qld 
Matrnal txumitud' QocetisUihd — NMV, Dli049, 
SAM. R98I7. R<>B3S, Battle Camp; MUZD, 90- 
92/70, 5 1cm SW of Calliope; QM, 1431. Gpfcat- 
<eum: QM, 118062. 11*066. Condamine River, 
Cecil Plains; NMV. D13049. SAM, R 1 1525-24. 
Opokbown; QM, !20*8V4I h km W &f rc>oUuv.ri. 
SAM, R%«J0. F.dwurd River Stn; QM. I2I84-H5. 
JI2M4. Eidsvold; OM, J 14383-84, tlilruth Plains 
Cunnnnuilla; MUZD, S6-58/7Q. 75/70. 9 & 15 km 
E of Goondiwimli; QM. J56I1-I2. Mackav: SAM. 
R474?, Mupoun Mission Sin; SAM, R9734. Mtuy 
River; QM. J 14 159-67, Mitchell R. Miwion: SAM. 
KI04I9, Prcstwood, Gilbert River; SAM. JO 640, 
Rockhnrnpion; QM. 110482, SAM, R3686, Si 
George: QM. .12186.89. Slannarv Hills: SAM, 
RW5, Stewart River; SAM. R9"S9l, Strathgordnn 
R.S . QM, 12222-7-2*. Surat. NMV. D7542, QM. 
14644, Townsville: QM, .13480, Vieto, Cooncoota; 
IJM. J 18063-65. Waratah Sin, Cunnamulla. 
Ot' n 1 1 pt ion' Snout to vent length ot mate* 
rVM-SM mm. females 74.8-101,2 mm; head 
noticeably hroadcr than long (HL/MW mean 
0.X3 1 : S -V.' HI. ratio low (mean 2.05 ). 



TL.'S-V ratio rather tew (mean 0.40); foot 
short <F/S-V mean 11371. 

There is extreme exostosis of the skull 
bones. The squamosal In large specimens is 
usually so heavily overlain with seeondaiy 
bone thai (t is visible through the skin, form- 
ing humps resembling parotoi<i glands, and 
extends su far superiorly that it approaches 
the margins of the frontoparietals. The fronto- 
parietal is entirely exostesed, but the lateral 
margins arc raised by bone deposition, so pro- 
du.ing a deep, median furrow The suborbital 
portion of the maxilla projects, forming a high 
and often concave shelf. 

The constricted pupil in six hvmc, specimens 
from Cooktown approximated a rhomboid 
shape (sec discussion). 

In preservative, C\ ttovuthoiUinrfiae H pale 
brown or grey, and is immaculate, lightly 
marked with scattered dark brown or blackish 
markings, or else very densely pigmented with 
such markings. The suborbital marking is 
broad and usually reaches the labial base of 
the maxilla. The backs of the thighs are usually 
very dark leaden giiey and lack lighter vemiicu- 
lations. 

In life the «eries from CookloWn were an 
immaculate dull sandy yellow dorsally The 
rostral stripe was dark brown, and similarly 
colored, small disrupted patches occurred on 
the inferior margin of the maxilla. The iris was 
goWeo and suffused with dark brown laterally 
and inferiorty. The posterior surfaces of the 
Thighs were leaden jircy, whilst the ventral sur- 
face of the body wall was a dull pearl bearing 
faint grey vermiculations on the throat- 

Geographic variation: There is consisfcrable 
variation in skull structure and coloration of 
the dorsum of this species. Comparison of 
small samples of extreme variants led us to 
conclude initially that Iwo species were in- 
volved. Examination of larger scries, however, 
has revealed the occurrence of Jotnw of inter- 
mediate appearance, The variation may be 
summarized as follows. All specimens front 
central and southern Queensland have high 
skulls with a gently sloping maxilla and a 
densely pigmented dorsum. There is sinking 
variation in individuals from northern Queens- 
land, Some are densely pigmented whilst other* 
are immaculate. The skull of the immaculate 
individuals is either similar to that of the pig- 
mented frogs, or is spatulate and distinctly 
flattened. Unfortunately we have been unable 
lo devise n mean* of objectively estimating 
skull depth with any degree of accuracy We 



9H 



M. 1. TV I ^K & A A. MAR11N 



hold the opinion that the high and the spatu- 
laic terms of the skull repiesenl different evo- 
lutionary trends of development. 

Whereas ihe extremes are dearly dilfeTen'., 
assessment of the significance of the observed 
variation rs complicated by the existence in 
northern Queensland of a number of inter- 
nediale fornix that cannnt he referred to either 
form. In addition there aie animals in which 
the terminal portion of the skull is more elon- 
gated. This variant occurs only on the Cape 
York Peninsula and at localities at the hase of 
the Gulf of Carpentaria. Morphometric data of 
iwo small series are .summarised in Table I. 
BggS: Oviducal eggs of three gravid females 
ranged from 3 1 to 1,3 mrn in diameter, An 
estimate ot the number in unc individual 
exceeded 1.000. The eggs have black animal 
poles. 

Mating Call: The mating calls of two indi- 
viduals fcJMj/dcd 5 Km S\V of Calliope. Queens- 
land, on 1 8.5.1 970. are very similar to those 
of C. ouxiralis. The ho\n< were calling on laml 
beside a rain-filled roadside ditch, with a wet- 
bulb air temperature of 24.6*C. The spectral 
structure of the calls of the two species is 
essentially identical, with C. novaahoHaiuiiitt' 
also having emphasized harmonic bands at 
about 600, 800, and 1.000 Hz. However, its 
call duration is considerably longer (mean 249 
m&ec: range 235—262 rmee). Judged on the 
hosts of the levels of difference in mating call 
structure nf sympatrie anuran spe.ies, this 
difference in duration does not represent diver- 
gence of sulTjcicnl magnitude to achieve repro 
duciive isolalion. Hence if C. aust talis and 
C. novaeholiatulioe occur hi sympalry (as they 
may in the Gulf District) we would expect 
them ft) hybridi/.c. A .similar pal tern of 
marked morphological differentiation, accom- 
panied by very little mating call divergence, 
characterizes the Weslern Australian IJmnody- 
uastvs rfonaU.s and the eastern L. <ltuneriH 
/Martin 1972). 

Geographic Ran tie: Cyclorana novaehollandiae 
ranges from the Cape York Peninsula of 
northern Queensland to the New South Wales 
border. 

KEY TO ADULTS 
Duck ol' thighs varicyaled, suborbital bar narrow 
*nd not reaching hnsj nf mnxilla; maxilla gently 
sloping and nut expanded laterally . ... C. austratts 

Back of thighs not variegated; suborbital toil (if 
present) usually broad and reaching base pf 
maxilla; tmixilla '.rxparuled to form lateral ahfiW 



Discussion 

The morphological complexity of C. novae- 
hoUatidiue as defined here is unparalleled 
amongst Australian anurans. We believe that 
a study of species isolating mechanisms, such 
as male mating call, in jjorth-etwtcnt Australia 
could reveal ihe existence of two or possibly 
even throe species. Our action of resurrecting 
C. novtwhoUatuimt- from ihe synonymy of C- 
ausftalis is therefore only the first step towards 
an understanding nf Ihe C tmuratis complex. 
The biological data, e.g. mating call structure. 
that are necessary for final resolution of the 
problem may be extremely difficult to obtain. 
The northern Queensland populations arc 
apparently opportunistic breeders which may 
call at a locality on only one or two nights 
each year (C. Tanner, pers, conim.). It seems 
justifiable* therefore, to treat ihe complex ai 
this preliminary level, 

In terms of skull structure. C. attxtraJh rs 
clearly the most simplified and primitive mem 
her of the complex, exhibiting limited develop- 
ment of skull bones and the least extensive 
exostosis. All the variations in the foTm of the 
skull and exostosis of the cranial bones such 
as the maxillary and squamosal m C fu>va?~ 
hollantiiae can reasonably he derived from C\ 
australi.s. 

The concept of C. uusirahs being the primi- 
tive mem her may be acceptable morphologi- 
cally, but it is more difficult to conceive fcftog&fe- 
graphically in view of the absence of any mem- 
ber of the genus Cytloratm in New Guinea. 
The geographic area occupied by ihe members 
Of the C. aust/alh complex, includes both hieh 
rainfall and relatively arid areas: i.e. it docs 
uot appear to be limited climatically. Thus if 
the complex originated in northern or north- 
western Australia it is surprising that it should 
be absent from New Guinea, Jennings (1V72) 
estimates that the most recent land communi- 
cation with New Guinea at fortes Strait ter- 
minated only 6,500-8,000 years ago; thus it 
is possible that colonization of north-eastern 
Australia by ihe complex occurred subse- 
quently. 

Ihe gross dilTerenc.es in morphology charac- 
terizing the members of the C. mnimlis com- 
plex represents a situation unique among Aus- 
tralian anurans. Differentiation in other 
species complexes, even those with extensive 
disjunctions of range (e.g. south-eastern and 
south-western Australia* is accompanied hy 
only slight morphological divcrpcncc. U seems 



PftOGS OF the cvrto/^iWj avstMUS complex 



y* 



probable, iherefore, that the complex is uf 
considerable antiquity. Two further circum- 
stances lend support to this suggestion. One is 
tfac affinity of Cyclorana wilh the Hyhdae 
demonstrated by Tyler (1972) and Watson 
$ Murtin (J 973). The other stems from our 
observations on the pupiL shape of C. novae- 
hoflandiae. As stated above, the constricted 
pupil of C. novae hoi land lae is almost rhom- 
boid. In fact, the ventral margin is an obtuse 
angle and the upper a broad curve. This 
curvature is difficult to detect during extreme 
constriction. Lynch (1971) considers the Ver- 
tically orientated pupil to be the primitive and 
the horizontal pupil the derived state. Kow- 
cvcj. from the fact that Nvctimysres (a genus 
that can only be derived from Litoriu. a hori- 
zontally-pupilicd stock) has a vertical pupil, it 
is clear thai, vertical orientation can be a 
derived State The trend to one or other orien- 
tation of pupil shape could be accomplished 



rnosi readily from a rhomboid. That such a 
structure occurs in living Cyclorana is con- 
sistent With our hypothesis of its antiquity. 

Acknowledgment* 

For the donation of specimens or the loan 
of those in their care we are indebted to Mr K. 
Cole. Miss J. Covacevich. Mr J. Coventry, Mr 
D. Howe, Dr B. S. Low. Dr G. Storr, Mr C. 
Tanner and Dr G. Ycstcrgren Mr G. )•*. Wat- 
son assisted in l he Meld. Photographs were pro- 
vided by Mr B N. Douetil and the tape of the 
mating call of C. auxttatis by Dr B Low and 
Dr D. Gartside. 

Our thanks are also due to Oi J. Ling tor 
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 various Australian museums ro 
examine collections. 



References 



Boui enokr, Qj A, (, 1882). — "Catalogue of the 
Batraehia Salient iu 8. Ecandata in the collec- 
tion of the British Museum." 2nd Htin. 
(London > 

U*ArrsruoM, B. H. ( L970). — Thermal acclima- 
tion in Australian amphibians. Camp. Hio- 
ehent, Physiol. 35, (S9-I03. 

Krv. D. B. < 1914).- -On a collection of reptiles 
and batrachians from Western Australia. /*»t. 
W. A ust. Mas. 1, 174-210. 

(JneoJbR, K. L. (I960). — A simplified tuble tor 
staging anuran embryos and larvae with note» 
on identification, Hcrpctoloi:u:a 16. 183-190. 

Gray, J. E. ( 1842).— Description of some hither- 
to unrecorded species of Australian reptile* 
and batracfuam. Zoo!- Misc., 51-57, 

Guntiirr, A. (185K>. — ''Catalogue ot the Bat- 
rachia Salientia in the collection of the 
British Museum." (London,) 

JENNtNCS, J. N. (1 972) .—Some attribute oi 
Torres Stunt. ///: Walter, O. I Ed.") "Bridge 
and Barrier: The natural and cultural his- 
tory of Torres Strait"- Dept of Biogeotsranhy 
and Geomorphologv. Pubh I3C. 3, Australian 
National University, Canberra. 

LintEjOHN, M. L (1957). — A new spacies of 
frog of the genus Cr'tnia. W. Aust. Nat. 6, 
18-23. 

LovrjtiDGE, A. (1935) — Austrian Amphihm jn 
the Museum of Comparative Zoology, Cam- 
bridge. Massachusetts. Hull, Mas, Camp, 
ZooL 78, 1-60. 

I vnch, J. D. I 1971). — Kvolutionary relation 
ships, osteology, and zoogeography of lepto* 
dactylic! frogs. Misc. Pahl. Mas .Vu/. Wsi. 
Vni'v K<wsn\ <53>, I-23K. 

Maki'IN, A. A. 0972),— Studhs in Australian 
Amphihia III. Tte Ijmnmivnanu's dorsalis 
complex (Amua. Lcptodacivljdac) Aa\i. /, 
7,ool 20, 165 211. 



MooRr, i. A. (.1954). — Geographic and genetic 

moThJiuu in Australian Amphibia. Anur. Nal. 

»8> 65-74- 
Parklr, U. W- 11940). -The Australasian frou-s 

of lbs tamily LeplodactyJidae. A'oWr Zool 

42, J-IOG. 
Pfcrfcjts, VV. (1867). — Heip2loloi?isctie notizen. 

Afofiarsb. K. PreiiM- Akuci. Ifm. tier (in, 

1S67, 13-37 
Sllvin, J. R. <I955>. — Notes oil Australian 

amphibians. Pmc. Calif. Acad. Sci. 28, 355- 

392. 
SpPNCta, B. (1901). -Two new species oi fttjfia 

from Victoria. Proc, R Spp, vjp n New Scr. 

13, 175-178. 
SrhiNUACHNfR. K (1867). — Amphibien. in: "Keis^ 

der Osterreichischen KtogaltiS Novani um die 

Erde in den Jahrcn I857nj$5fc" Zoologiache 

Theil. 1<4), 1-70. (Vienna). 
Sirax'ghan, I. R (I9n8). — A Uxonomic review 

of the genus Mixoplnts I Aouta: LeptOdacft- 

lidae). Proc Linn. Sac. N.5.W. SQlf), 52-5v. 
Stkaoch^.1. K. & NHiN, A R. (I9fi6).— 

Specintion and polymorphism in the gcrtuS 

Criuia Tbchudi (Anura; Leptodactvlidac ) in 

OueenslancL Prat: /?. Sot: QUI 78U.L H-3& 
IvrrR. M i U97$}< — Superficial mandibular 

muscuJalurc. vocal «ac« and the phytogeny of 

Anstralo-Papuan l^plodactvlid nxies. Rev. S. 

Aast. Mus, 16(9), 1-20. 
Tyllu, M. L, St Parkhr, K (1974). -New Hpecie^ 

ot hylid and leptodactylid frogs from southern 

New Cjuiucu. Trans. R. Sue- 5. Aast, V8(2). 

71-78. 

Watson, G. F.. Sl Martin. A. A, < 1973). —Life 
history, larval morphuloyy and relaliotwhips 
of Aitttraliftn Icptodfictvlid froas. 'thing. H 
Sac.fr Aast. 97(1). 33-15. 



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 1 "Lectotype" of Leporillus 

apicalis (Gould, 1853) [Rodentia: Muridae] . . . .101 

Barker, S. Revision of the Genus Astraeus Laporte & Gory (Coleoptera: 

Bupr estidae ) 105 

Beck, R. G. Factors affecting the Distribution of the Leptodactylid Frog Geo- 

crinia laevis in the South-East of South Australia - - - 143 

Edelstein, T. and H. B. S. Womersley The Thallus and Spore Development of 

Lobospira bicuspidata Areschoug (Dictyotales: Phaeophyta) - 149 



PUBLISHED AND SOLD AT THE SOCIETY'S ROOMS 

STATE LIBRARY BUILDING 

NORTH TERRACE, ADELAIDE, S.A, 5000 



THE IDENTITY AND STATUS OF THOMAS "LECTOTYPE" OF 
LEPORILLUS APICALIS (GOULD, 1853) [RODENTIA: MURIDAE] 

BY J. A. MAHONEY* 



Summary 

MAHONEY, J. A. (1975) .-The identity and status of Thomas' "lectotype" of Leporillus apicalis 
(Gould, 1853) [Rodentia: Muridae]. Trans. R. Soc. S. Aust. 99(3), 101-104, 30 August 1975. 
The specimen selected by Thomas as the lectotype of Leporillus apicalis (Gould, 1853) was 
misidentified by him and belongs to Leporillus conditor (Sturt, 1848). It does not belong to the type 
series of L. apicalis, therefore Thomas' lectotype selection for that species is invalid. The type 
material of L. apicalis and L. conditor is missing. Thomas' "lectotype" of L. apicalis and a second 
specimen of L. conditor in the British Museum (Natural History) could belong to the type series of 
L. conditor. Evidence for the occurrence of L. apicalis in Tasmania is lacking. 



THE IDENTITY AND STATUS OF THOMAS 1 "LECTOTYPE" OF LEPORfLLUS 
APICALIS (GOULD, 1853) (RODENTIA: MUR1DAE] 

by J. A. Mahoney* 

Summary 

MaHONEY, J. A. (1975). — The identity and status of Thomas' "lectotype" of Leporillus apicalis 
(Gould, 1853) [Rodentia: Muridael. Trans, R, Soc t S, Atut, 99(3*, 101-104, 30 August 
1975. 

The specimen selected by Thomas as the; lectotype of Leporillus apicalis (Gould. 3 853) was 
misidentiftsd by him and belongs to Ltparilhts conditor (Sturt. J 848). ft does not belong to 
the lype series of L. apicalis, therefore Thomas' lectotype selection for that species is invalid. 
The type material of f.. apicalis and L. conditor is missing, Thomas 1 "lectotype" of L. apicalis 
and a second specimen of L. conditor in the British Museum (Natural History) could belong 
to the type series of L, conditor, Evidence for the occurrence of L. apicalis in Tasmania is 
lacking. 



introduction 

The name Hapalotis apicalis was proposed 
by Gould (1853a) for a new species of rodent 
from Australia. He did not stale 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 
it single example procured by Mr Strange in 
South Australia, and he illustrated the external 
features, 

Thomas (1906a) nominated Hapalotis api- 
calis as the type species of a new genus. 
Leporillus, and subsequently (Thomas 1921a) 
he selected British Museum (Natural History) 
specimen 1 853.1 0.22.15 1 as the lectotype of 
LepoyUlus apicalis, describing it as a female 
hum S. Australia. Explaining his lectotype 
selection, Thomas (1921c) stated that 
although Gould had in his collection two speci- 
mens of that species, he seems to have done 
hi* describing from only one of them (BM, 
53.10.22.14 (sici— lapsus for BM. 53.10.22.15) 



— the worst of the two, young, and with an 
imperfect tail. 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 [adult with nearly perfect 
skull- and quite perfect tail (BM, 53.10.22.14)1 
has the latter 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 Murtdae. 

A study of three Australian Museum speci- 
mens of L. apicalis, and (he literature, enabled 
Troughton (J 923) to confirm that Gould was 
correct in attributing a white tail-tip to the 
species. Troughton stated also that Thomas 1 
remark thai Gould seems to have done his 
describing from only one of his two specimens 
means that that specimen must be accepted as 
the holotypc. 

Tale (1951) treated specimens 1*53,10.27.14 
and 1853.10.22.15 as **cotypes" of L. apicalis 



'•' Department of Geology and Geophysics, University of Sydney, Sydney. N.S.W. 2006. 

1 The first rwo digits of British Museum (Natural History) registration numbers or mammals are fre- 
quently omitted from publications. Thus Thomas uses 53.10.22.15 for 1853.10.22,15 

"This skull is registered as 1854.10.21.1 and the Register entry mentions a stuffed specimen and relen 
lo 53.10.22.16. 1 have been unable to find a specimen numbered 1853-10.22.16 in tbe 
British Museum (N.H.) therefore 1 am following Thomas' conclusion that IS53.10.22.I4, 
1*53.10.22.1 6 and 1854.10.21. J belong to the one individual; but it is possible that 185V 10,22.14, 
identified as Hapalotis apicalis in the Register, is jost and the skin novv numbered 1853.10.22. 14 is skin 
1853.10.22.16 with an incorrect number. A note in Thomas' handwriting attached to skin 
IK53. 10.22.14 and stating that this specimen was considered to be the type seems \o refer to the 
Museum Register where "type"' has been written opposite the number 1853.10.22.16. A portion of the 
posterior half oi' the cranium, and the left mandibular ramus, are missing from the skull. 



102 



J. A. MAHONF.Y 



TAB Lb 1 

Skull measurements (in mm) of Thomas' "Iccto- 

t\>ve" of Leporillus ypieyli* (Cmuldl 8.M (N Wj, 

185 J. 1 0-22.1 5. 



Maximum width acio .s na 


sals 


5.11 


Minimum width 


across 


right zygo- 




matte plate 




■ • t 


4,4 


Length of tight M 1 


-fl 


_ 


9.7 


Width of left Mi 


- 


t 


2.6 


Width of left Ivt^ 




• 


2.7 


Width of lcftM» 


• 


• 


24 


Length of left Mj- 


i 


- 


9.1 


Widthof teftMj 


. 


. 


%& 


Width DricflM... 


. 


• - t 


2.7 


Width of left M A 




■ 


2.4 



The teeth measurements ate fur the crowns of the 
teeth 



and referred to them as aduit and young 
females, from "South Australia 1 ", collected by 
F. Strange. He briefly described the skin ami 
skull of each and. recorded measurements of 
them. He does not refer, in hi.s account of L. 
ttpkatis, to Thomas' Jectotype selection or to 
Troughton's recognition of a hoiotype for the 
>pccjes. 

Identity of the "leclotypc" 

British Museum (N.H.) specimens 
1853.I0.22.J4 and 1853.10.22.15 are examples 
of Lepohtlus conditor (Start, 1848) and not 
specimens of Leporttlus dpicalh (Gould, 1853) 
as believed by Thomas, They do not agree with 
the original description of L. apicalh (Gould) 
and because of this 1 do not accept that either 
of them belong 10 the type series of that 
species. Consequently, I regard as invalid 
Thomas' leelotype selection fur L. apicalis, 
Thomas ( 1921b 1 ) stated there is no specimen 
of L. conditor in the British Museum, and 
tajther demonstrated his unfamiliarity with its 
characters by suggesting that it possibly 
belongs fo Notomys, a genus of Australian 
hopping mice and rats. 

Measurements ol the badly damaged skull of 
Thomas' "lectotype" arc given in Table L The 
skull is illustrated in Figs 1-4. 

Discussiou 

The type material of both L apicalis and 
L. venditor is Jost. I he latter species was 
described by Stun in IS4S as Mus conditor 
from specimens observed and collected on hi* 



1844-6 Expedition to Central Australia, the 
original description was published in the 
Narrative of the Expedition and Sturt did not 
say where his material was deposited. It seem*. 
likely however thai at least one specimen. 
illustrated by J. Gould and H. C. Richter in 
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- 
tain specimens of LepitrUlus. 

Specimcus 1853. JO.22.I4 and 1853.10.22.15 
were registered in the British Museum on Octo- 
ber 22nd, 1853, and were acquired from 
Gould. Labels attached to them refer to S. 
Australia and F. Strange. The entries for them 
in the Register mention neither a locality nor 
Strange. S. Australia 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 accompanied Sturt on 
some of his early surveys (but not the 1844-6 
Expedition), and was an eaily settler in South 
Australia and later, in the 1840's, a resident 
of New South Wales (Whittell 1947). Gould 
(1849) docs not mention Strange and Soulh 
Australia in his account of L condiU*r. Subse- 
quently (Gould 1863) he gives only the 
interior of New South Wales and Victoria as 
localities for the species. It is possible that the 
inscriptions on the labels are interpretation* of 
the origins of the two specimens based on 
Gould's account ot L. upica/tK. If they are riot 
interpretations, their significance is uncertain 
since the citation of S. Australia is ambiguous 
and Strange might not be the collector of the 
specimen*. 

British Museum (N.H.) specimens 
1853.10.22.14 and 1853.10.22.15 could have 
been collected on SiutVs JK44 ft Expedition 
and might belong to the type series of L, condi- 
tor. This is so even if they came from .South 
Australia. Sturt <IS48, Vol. I, pp 120-121) 
referred in his account of the hxpedition's pro- 
gress along the Darling River in New South 
Wales to an individual ot /-. conditor secured 
by Mr Browne and to one. a male, obtained 
by himself from a native. However, Sturt (Vol 
2, Appendix, p 4) noted also that the last nest 



* Gould's name is printed on this plate and the Bpsciea name Mas conditor is attributed to him by 
Sturt, acveruVJess Sturt is the author of the name Mux conditor. 



THOMAS' "LECTOTYPE" OF LEPORILLUS APICALIS 



103 







iiimimiiiiiiimmiii 



Figs 1-4. Leporillus conditor (Sturt, 1848). British Museum (N.H.) 1853.10.22.15. Thomas' "lecto- 
type" of Leporillus apicalis (Gould, 1853). Fig. 1 — Ventral view of cranium (x3). Fig. 2 
— Occlusal view of left upper molar row (x8). Fig. 3 — Occlusal view of left lower molar 
row (x8). Fig. 4 — Right lateral view of cranium (x3). 



of L. conditor was found on the bank of the 
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. A. MAHONfcY 



British Museum, presented by R. C, Gunn, 
from Van Diemcn's Land, accords very closely 
with it in the colouring of the fur and in the 
rat-like form uf the tail. He added that it is of 
much smaller size than L, apicatis and in all 
probability will prove to be a new .species. 
Gould's listing in 1 863 of Van Diemen's Land 
as a possible locality for L. ttpicalis could be 
based on that material. Tasmantan rodent 
specimens in the British Museum (N.H.) and 
attributable to Gunn are recorded m the Regis- 
ter. The identities of these specimens and their 
registration numbers are Rattus rttttus 
(Linnaeus, 1758) < 1837.6.10.56). Rattus nor- 
ve&cus (Berkenhout. 17b9) ( 1838.1.15.17), 



Rattus lutreotux (Gray, 1841 ) ( 1845.5.2.3, 
1 852.1. 15. 16. 1852.1.15.17), Maswcomys 
fuscus Thomas, 1882 (1852.1.15.15) and 
Pseudomys hifigmsi (Trouessart. 1897 ) 
( IX52J.15.IK). None of these are L. apicalis, 
and evidence for the occurrence of this species 
in Tasmania is lacking. 



Acknowledgments 

This study was carried out in the British 
Museum (N.H ) by arrangement with Or 
G. B. Corbel of the Mammal Section. The 
photographs were taken by Mr F. Greenaway, 
British Museum (N.H.) Photographic Unit. 



References 



(Jouid. 3. ( 1849). — "The mammals of Australia". 
Pi 2, pi. 8 and text (1863, vol. 3, pi. 6 and 
text). (J. Gould: london.) 

(iot'io, J. (1853a). — Remarks on the genus 
Hupalotis. Proc. zoot. Soe. Loud. 1X51, 126- 
127. [The approximate date of publication of 
this work in 1853 is 29 April — see Prac. zoot. 
Sac. Lorut. 107, 81 (J 93 7). I 

Goui.o, J. (1853b). — "The mammals of Aus- 
tralia". Pt 5, pi. 12 and text (1863, vol. 3, pi. 
2 and text). (J. Gould: London.) [The date 
November 1st, 1853 is printed on the cover of 
Pi 5 of Gould's work.l 

Gould. J. (1863). — 'The mammals of Australia". 
Pt 13, Introduction, pp. xi-xl (1863, vol. 1, 
Introduction, pp. xi-xl). (.1. Gould: London. ) 
(The date May 1st, 1863 is printed on the 
cover of Pt 13 of Gould's work. An uncor- 
rected proof-sheet version of Pt 13, pp. xi-xl 
of "The mammals of Australia'' is included 
in pp. 1-51 of a work entitled "An introduc- 
tion to the mammals of Australia" hy 3. 
Gould and published in 1863. A copy of this 
work in the Australian Museum library has 
a date of presentation to Reverend John 
Barlow L.R.S., May 6lh, 1863, inscribed on 
the title page.1 

Sti'itr, C. (1848). — "Narrative of an expedition 
into Central Australia, performed under the 
authority of Her Majesty's Government, dur- 
ing the years I844 T 5. and 6. Together with 
a notice of the Province of South Australia, 
in 1847." (Boone. London.) Vol. 1, X. 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, Us date of publication is 1848 
Thus it is cited in a "List of New Books" in 
The Atttennentn (Jnuttud of English and 



foreign literature, .science, and the fine arts), 
no. 1101. December 2nd, 1848. p 1207. eol. 
2, and in a list of "Publications Received" in 
1 he Spr* tutor, for the week ending Deeemher 
23rd, 1848. no. 1069. p. 1237. col. 1. Furthei- 
more. p. 1243, col. 2, of The Spectator, for 
the wctk endine December 23rd. 1848. con- 
tains a publisher's notice which states that 
Sturls work is "Now ready". 1 

Tate, G. H. H. (1951).— Results of the Arch- 
bold Kxpedilions. No. 65. The rodents of 
Australia and New Guinea. Ball. Am. Mas. 
nat. Hist, 97, 183-430. 

Thomas, O. (1906a). — On the generic arrange- 
ment of the Australian rats hitherto referred 
to C'onilurus, with remarks on the structure 
and evolution of their molar cusps. Ann. 
Mas'- mn Hist. (7) 17, 81-85. 

Thomas, O. ( 1906b).— Mammals. Pp. 3-66. in 
"The history of the collections contained in 
the Natural History Departments of the 
British Museum". Vol. 2. Separate historical 
accounts of the several collections included 
in the Department of /oology. (British 
Museum: London.) 

Thomas. O. (1921a).— Notes on Australasian 
rats, with a selection of lectotypes of Aus- 
tralasian Muridae. Ann Man. not. Hist. (9) 
8. 425-433. 

Ihomas, O. (1921b). — Notes on the species of 
Notomxs, the Australian jerboa-rats. Ann. 
Mag. Hat. Hist. (9) S, 536-541. 

Thomas, O. (1921c). — On three new Australian 
rats. Ann, Afa&. not. fttst. (9) 8. 618-622. 

TbcjGhton, E. Le G. (1923). — A revision of the 
rats of the genus Lcporittus and the status ot 
Htipalotis per.soitata Krefft. Rrc. Attst. Mus. 
14, 23-4L 

Whiiteil, H. M. (1947).— Frederick Strange. A 
bioKiaphv. Attst. Zoot. II, ^6-114. 



REVISION OF THE GENUS ASTRAEUS LAPORTE & GORY 
(COLEOPTERA: BUPRESTIDAE) 

by S. Barker* 



Summary 

BARKER, S. (1975). -Revision of the genus Astraeus LaPorte & Gory (Coleoptera: Buprestidae). 
Trans. R. Soc. S. Aust. 99(3), 105-141, 30 August, 1975. 

A revision is presented of the Australian Buprestid genus Astraeus LaPorte & Gory. Following 
van de Poll (1889), but extending the status given by him to two separate groups within the genus, 
Iwo new sub-genera are proposed, Astraeus (Depollus) and Astraeus (sensu stricto). The genus is 
considered to contain 39 valid species. The species are keyed and a complete description or 
redescription is given of each, together with an illustration of all species not previously figured in a 
publication; an outline diagram is given of the male genitalia of 36 of the 39 species. Sixteen new 
species {aclamsi, bakeri, carnabyi, carteri, declariensis, fraseriensis, globosus, goerlingi, 
macmillanu minutus, obscurus, polli, robustus, smythi, tamminensis y and watsoni) are described. 
Two names, major Blackburn and mastersi MacLeay, previously regarded as synonyms are 
revalidated. Two names previously regarded as synonyms are confirmed as such. They are as 
follows: meyricki Blackburn (of badeni van de Poll) and tepperi Blackburn (of jansoni van de Poll). 
Three names are newly synonymised (the synonym first): splendens van de Poll = mastersi 
MacLeay; simplex Blackburn = mastersi MacLeay; and strandi Obenberger = dilutipes van de Poll. 



REVISION OF THE GENUS ASTRAEUS LAPORTE & C20RV 
(COLEOPTERA; BUPRESTIDAE) 

by S. Barker* 

Summary 

Barker, S. ( 1975).— Revision o£ the genus Axtraeux f.aPorle & Gory (Coteopterui 
Buprestidae). Jrans. R. Soc. $. Aust, 99(3 ), 105-141, 30 August, 1975. 

A revision is presented of the Australian Buprestid genus Axtraenx I a Porte & Gory. 
Following van de Poll (1889), but extending the status given by hinj to two separate groups 
within the genus, two new sub-genera are proposed, Axtraenx {Depoltux) and Axtraenx {sensu 
strtcto). The genus is considered to contain 39 valid species. The species are keyed and a 
complete description or redescription is given of each, together with an illustration of all 
species not previously figured in a publication; an outline diagram is given of the male genitalia 
of 36 of the 39 species. Sixteen new species (adamsi, bakeri, varnubyi, carter/, dedariensis, 
fraseriensis, glohosus, goerlingi, macmiUani, mlnuiux, ohseurux, palli, rahuxtux, stnytht, tammi- 
nensis t and watsom) are described. Two names, major Blackburn and masters! MacLcay, 
previously regarded as synonyms are revalidated. Two names previously regarded as synonyms 
are confirmed as such. They are as follows: meyricki Blackburn (of hiidrni van de Poll) and 
tcppt'ri Blackburn (of junsoni van de Poll), Three names are newly synonymmed (the synonym 
first); splendent van dc Poll • maxtcrxi MacLcay; simplex Blackburn = muxterst MacLeay; 
and xtrandi Obenberger = dihuipes van de Poll. 



Introduction 

There are 27 described species of the genus 
Astmeus from Australia and one from New 
Caledonia. Originally established as Asthraeus 
by La Porte k Gory (1837). the genus was 
referred to by that name until Gcmminger & 
de Harold (1869) used Astraeux, thus correct- 
ting an incorrect transliteration from the 
Greek. This has been followed by all subse- 
quent authors. Barker (1964) requested the 
International Commission for Zoological 
Nomenclature to validate the emendation of 
Asthraeus to Axtraeus and this was eventually 
carried out (I.C.Z.K 1966, Opinion 795). The 
genus was last, revised by van de Poll (1889) 
who gave excellent descriptions and illustra- 
tions of all species known al lhat time. Pre- 
sumably van de Poll never saw a live speci- 
men of the genus, so would have been 
unaware of their unique spring mechanism 
involving the release of the elytra from the 
closed position. When the living beetle releases 
the .spring, the elytra flick open with such force 
that the insect can be projected for up to 
•vevcral metres. The effect is reminiscent of the 
sternal spring of the Elatcridae except that the 



same result is achieved in a 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 exhibit classical 
warning colouration of yellow markings on a 
dark background. Whether any of the species 
are noxious to predators or whether they arc 
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 .in 
Australian worker as most of the type speci- 
mens are housed in overseas museums. Alv>. 
through habitat destruction throughout Aus- 
tralia, some of the species are not abundant 
over their former range, and unfortunately the 
genus is very poorly represented in museum*- 
and in most private collections in this country. 
The adults of more than half of the species 
occur on Casuarina sp., but some are also 
associated with species of Jaeksonia, BanhsUi, 
Ximthorrhoea, CalUtris, Melaleuca, Acacia, 
Daviesia, and possibly Dryandra and Hakeu, 
Adults of the earfy emerging species can he 



* Department of /xiology. University of Adelaide, Adelaide. S. Aust. 5000. 



I Oft 



S RARKFR 



collected in August and adults of some species 
are present as late as March, More than haJf 
of the known species are found in the southern 
hfltf of Western Australia- The oihcrs occur 
over the rest of the continent excluding the 
desert and northern-most tropical areas. One 
species occurs on New Caledonia. The genus 
is unknown from Tasmania. 

In this paper male genitalia have not been 
used to diagnose the species, but have been 
mod 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 (be South Austra- 
lian Museum. The remaining specimens are 
vCaltered throughout the collections listed 
below. The abbreviations used in the text for 
museum and private collections arc as follows; 
EA Collection of Mr K. l\. Adams, 

Edungalbtu Queensland 
AM "I he Australian Museum. Sydney 

ANIC The Australian National Insect Col- 
lection. C.S.I. R.O.. Canberra. 
BPBV1 The Bermce I* Bishop Museum. 

Honolulu. 
BM The British Museum (Natural His- 

tory J. London. 
KC Collection of Mr and Mrs. K. Car- 

naby, Wilga, Wcslern Australia 
)H Collection of Mrs J. Harslett. 

Amiens, Queensland. 
MM The MaeLeay Museum. Sydney 

(types on permanent loan to ANIC) 
MNHN Museum National d'Histoire 

Nalurelle, Paris. 
NM The National Museum of Victoria. 

Melbourne. 
NMP National Museum. Prague. 
QM The Queensland Museum, Brisbane- 

SAM The South Australian Museum. Ade- 
laide, 
WADA The Western Australian Department 

of Agriculture, South Perth- 
WAM The Western Australian Museum- 
Perth. 
The Australian Buprcstidae have been 
divided into six sub-tamilies ( Britton 1970). 
Affrttffus belongs in the sub-family Bupres- 
tinae. the members of which have mouthparts 
not produced downwards to form a rostrum, 
In the sub-tribe Buprestes the metathoracic 
epimera are completely exposed whereas in 
the Anihaxiae they arc totally or pailially 
covered by the lateral extension oj the abdo- 



men. Axtraeux is placed in the sub-tribe Bup- 
restes with nine other genera found in Aus- 
tralia. Axirtteux and Neohupresiis both have an 
indistinct scutellum, but Neobuprextis has an 
exposed pygidium whereas in Astravtix it. is 
covered hy the elytra. All of the other Austra- 
lian members of the sub-tribe have a distinct 
scutellum. They are: Naxciohifs, Nose in, Neo- 
bubaxies, Notobuhasrex, Bubaxtex, Bttprextinti. 
BupreMocles n\\d huryspitux (Carter 1929"). 

Taxonomy 

Genus ASTRAEUS LaPurte A Gory 
Avhractts LaPortc & Gory, 1837: I, pi. 1. 
fiy. 1. (Invalidated, Opinion 795.) Imhoff. 
1 856; 46. Laoordaire. 1857: 43. Saunders, 
IK6S: 10. pi. 1, Jig. 12. 

Axtiwux: Gemniinger & de Harold, 1869, St 
1380. Saunders, (871: 43. Masters, 1871: 
124. MaeLeay, 1873: 239-240. Kerremans 
1885: 136, van de Poll. 1886: 176-178. Mas- 
ters. 1836: 71. van de Poll, 1889: 79-110. pis 
J, 3, figs 1-19. Blackburn. 1890: 1256-1258, 
1891: 496. van de Poll. 1892: 67-6$. Kerre- 
man*. 1892: 103-102. Blackhurn. 1892; 211- 
213: 1894: 101: 1895: 45-46. Kerremans. 
I9Q0: 295-296: 1902: 148-149. PmlveL 
1904: 116. Carter. 1925: 229, fig. 1 Oben- 
bcrgcr. 1928: 204-205. Carter. 1929: 265. 
2S2, 302, pi. 33, fig. 43, Obenbcrgcr, 1930: 
.'65-367. Carter, 193.3 : 41 . ObenbergeT. 
1936: 133. Barker, 1964: 306-307. T.C.Z.N.. 
1966: 269-270. 

Ait>e?ia; Carter. 1929; 270. 

Typa species. A stratus jitrsoplrsux LuPorte 

& Gory, 1837 (by monotypy). 

Generic description 

Head. Medium size or large; the surface 
pined; with a median longitudinal impressed 
line on the basal halt; clypeus short, the apical 
margin wilh a bow-shaped indentation; labium 
incised: aniennal cavities small and rounded, 
situated near the lower internal corner of the 
eyes. 

Antennae, The first, segment longer than any 
of Ihe rest, rounder and thicker a! the apex 
than at the base; 2nd segment short and 
obconic; 3rd segment longer than the 2nd bet 
not as long as the first, also obconic; 4th seg- 
ment slightly shorter than the 3rd^ dorsoven- 
trally flattened, the front edge projecting out- 
wards to form a tooth, the remaining segment*.. 
except the last, similar in structure to the 4th 
ami each sligbly smaller iban the succeeding 
segment; the last segment flattened but not 



Rh VISION OF BUPRESHD GfcNUS ASTKAEUS LAPORTfc A OORY 



107 



toothed. In species belonging to Astrarus 
tsensu stricto) the antennae are sexually dimor- 
phic, being longer in male* than in females. 
due to the segments after the 3rd being of 
approximately equal length, whereas in females 
they become progressively shorter. In Asttaeus 
iDepdlltts} the antennae do not show sexual 
dimorphism and are short in both sexes. 

Pronotum. Surface pilled; wider than lung; 
dorsaJly convex; rounded at the sides and 
narrower a! the apex than <it Ihe base; a itecp 
notch occurs on either side of the middle, at 
the base. In all but one species a small spindle- 
shaped depression is present at the base of the 
median lobe. I have called this structure the 
basal crypt (Fig. 24 >. 

Scutellurn. Invisible. 

Elytra. Convex dorsal ly, each side with a 
cruvcenl-shaped lobe fitting over the notches 
in the base of the pronotum; the apex ol each 
clylron ends (with rare exceptions) in a sharp 
point, the sutural spine, usually lonned by a 
crescent-shaped lateral emargination which 
lormx on the inner edge a smaller lateral 
tooth, usually called the marginal spine; it' the 
lateral enwginalion is absent, so is ihc mar- 
ginal spine; below the shoulder the external 
margin is dilated, sometimes the dilation is 
rounded posteriorly, sometimes siiongly pro- 
jected backwards - and angled forming a point. 
the whole being folded inwards and covering 
pan of the metastemal coxae — this structure 
IS called the humeral fold I Fig. I); punctate- 
striate and in many cases also cosiate. there ft 
a curved lip on the anterior undersurfnee of 
each elytron which fits over an opposite curved 
projection on the mesothoras. together form- 
ing the catch of a spring escape mechanism. 

Undersurface. Variably pitted; anterior edge 
ol presternum straight or crescent-shaped and 
a large prostcrnal process, mesostcrnum <J>ort, 
fucuistcmurn approximately equ.-il in length to 
the presternum; posterior margins of coxae 
and abdominal sternites glabrous. 

Legs. Femur slightly doisovcnirplly flattened 
but robust; tibia almost cylindrical but longer 
than the femur; 1st tarsal segment longer than 
the remaining segments which become pro- 
gressively shorter- 
Body outline. Laterally teardrop-shaped, the 
apex being constricted; dorsally convex; ven- 
trally straight at the anterioi end and curving 
upwards ai (he end of the ahdnnien. rarely 
concave. 



Key to the sub-genera of Astraeus 

I. Lateral lobes of pronotum projecting from rbc 
elytra t edge 'ioc; elytra st date-punctate but 
never with longitudinal costac; numeral fold 
rounded . . Astracus {Depol(us) 

I. Lateral lobes of pronotum confluent with the 
elytra I ed^e line; elytra sLriate-punclaLe with 
well or poorly developed longitudinal costac; 

humeral fold angled (Fig. I) . ,,,. 

Astracu.s (siwu .stricto) 
Sub-genus Depollus sub-gen. nov. 
Tvpe species Aitraeus abefrath van dc 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 lobes with the posterior margin curved 
upwards and outwards. Elytra striate-punctate. 
the edges of the striae never costate. The 
suture ends 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 maiginjl 
spine. The humeral fold A rounded and poorly 
developed. Only known to occur in Western 
Australia. 

Key to the speck* of Astracus (Dcpollus) 

1. Basal colour brown .,1. pwt sp. flow, 
I Basal colour black ... 2 

2. Elytra with 4 longitudinal red vittaa 

5. iineatux van de Poll 

2, BJytra without longitudinal red vitrnc - 1 

3. Head, prunotum and uiidersiirfacc vciy hairy 

4 
3 1 Head, pronotum and undersurfa.ee not verv 

hairy 5 

4. Pronotum with clumps of hair emerging 
from depressions towards Ihe sides; no mar- 
ginal spine 3. robustm sp. nov. 

4. Pronotum with hair emerging singly, not 
clumped; small marginal spine 

4. akermwi van dc Pol) 
5. Head and pronotum with deep punctures close 1} 

packed together 2. tammhtctts'u sp. nov, 

5 Head and pronotum with shallow, separated 

punctures ,,.. .,....., ,6 

6. Pronotum laterally inflated, elvlral patiern 
numerous sfrigifonn Bjiatfi 

6. muhimtaws van de Poll 
ft. Pronotum not laterally inflated, elytra! pat- 
tern small and large spots and longitudinal 
vtttae ... 7 

7. Sutural spine well developed and nuicurving 

7. irregularis van de Poll 

7 SunjraJ spine poorly developed and not out* 

curving , 8. detlortettsis sp. ro>v- 

: \sh.u-iis (Pepullus) polll sp. nov. 
FIGS 2, 22A 

TVfH'S 

Holotype. <£ Wungan Hills, W. Ausi.. On 
Casuurina campesirls, 9.il97l. -V- fair for, 
SAM. T 20940. 



I ox 



S. BARKPR 





« — 1mm 



Pig. I. Outline diagram of the left humeral fold 
of 4 species of A.straetia. The head end of 
each is to the top of the page and the left 
side in uppermost. Note that the specimens 
figured are all females and that (a) and 
<b) were smaller t>peeimens than (c) and 
(d). Bach structure is categorised as in 
the text, (a) A. masiersi MacLeay - well 
developed, acutely angled. (b) A- 
sumaut-lH Saunders — well developed, 
angled, (c) A. iniricatits Carter - 
moderately developed, angled, (d) A. 
camabyi sp. now - poorly developed, 
slightly angled - 

Allotype $?, Wongan Hills, W. Anst. on C\ 
rampeMri.s, 9.L197I, £ Barker, SAM. I 
20941. 

Paratypes: 1 £ & 1 gj 2 km \V of Wongan Hills, 
W. Aust, on C. campvstris, 20.U973, 5. Bar- 
/r?r, EA; 5 o* & 4 9. Wongan Hills, W. Altai. 
on C\ vumpestris. 9.1.1971. 5. Barker, ANIC 
1 1 rf arid I ?).BM(1 ^& I j-V MNHN 
(1 i/Al 2). SAM (2 £ & I S); I ft Deduri. 



W. Aust. T 12.U948, ;. A, Douglas, NM; 1 A 
Oedari, W. Aust, WADA; 2 d\ South Tam- 
min Mora Reserve, W. Aust. on C. catnpes- 
tris\ 6.i.l97l & 9.ii.l971, S. Barker, WAM ( 

Colour. Shiny. Head brown with the following 
yellow markings: tw r o lunettes from the inner 
corner of the eye. concave towards the base of 
the antennae but not reaching the middle of 
the head; behind each of the former at the 
base is a single small spot; a small spot at the 
base behind each eye; an elongate band runs 
from the anterior underside of the eye almost 
to the base; there is a large spot on each side of 
the gular; antennae brown. Pronotum brown 
with bfue reflections and the following yellow 
markings: close to the anterior margin on each 
side of the centre, but not reaching it is 
a laterally elongate mark; in the same line, 
directly behind the eye, is. a smaller laterally 
elongate spot: towards the base, in the middle 
of each side, there is a longitudinally elongate 
spot, and between them a small median longi- 
tudinally elongate spot; laterally there h a 
longitudinally elongate spot commencing just 
after the middle, running to the base, which is 
flared outwards with a dark rim and the 
exposed inner surface is yellow. Elytra brown 
with blue reflections: the intervals between the 
striae are irregularly mottled with yellow, more 
so at the base than at the apex; the sutures, 
apical spines, and the lateral and antcnoi 
margins of the elytra are black hut elsewhere 
brown. Undersurface brown; the mesoslerxuim 
and metasternum each have lateral yellow 
patches and each of the abdominal segments 
have lateral yellow spots; the coxal plates each 
have 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 at 
I he sides, except for two small round glabrous 
areas in the centre of each half; a short 
impressed line projecting forwards from the 
basal crypt; sides rounded and narrowed from 
base to apex. Elytra with the intervals between 
the striae slightly convex and transversely 
strigose with shallow punctures; parallel-sided 
until after Ibe middle then rounded to the 
base; with a small sutural spine projecting 
laterally. Undersurface with shallow punctures, 



REVISION OF RUPRESTID GENUS ASTRAEUS LAPORTli k GORY 



10:1 



denser at the sides tluu in the middle; with a 
lew short hairs. 

Size. Males 10.5 ± 0.15 x 3.8 dr 0.07 mm 

118). Females 11,5 * 0.27 v 4,0 ^0.09 mm 

MS). 

Oisthbutiim, Western Ausl ralia. 

General remarks. Named after the late J. R. H. 

Nccrvort van de Poll. 

Spt.t Unt'n.s e.ifiiniuitl. W Ami.: Types; 5 rf & 3 
?. J3 km N of Guomalliny. Dd Casuaritia campey- 
im. 9J.I97I, S. Barker, I ,? & I ?, Waddirtgton, 
on C. campextri*. 9,1.1971, J. farker; \ J & 2 £ 
Deduii. on C. campffitris, 20.xii.l972. 5. Uar- 
gvrj 3 rf # 3 9, 2 km W Of Wongan 9%, on C 
cam pest ris, 20.U973, A'. Barker. 

2, Astra? us (Dcpnllus) tamniinejiMs sp. nov. 
MCiS 3, 22H 

Holotypc: J, South T&ramiu Flora Reserve. W 
Aust. on Casiturina camju.;trls. 6 J. 1 97 1 , 5. 
POFiTr, SAM, I 20438. 
Colour. Head and pronofum dull, elytra shjny. 
Head black, bottoms of most of the punctures 
yellow with the following yellow markings: 
two small elongate spots each between the 
eye and above fhe base of the antennae; 
laterally a small round spot between the eye 
and the base of the head; ventially all yellow, 
almost continuous under the eye with the iirst 
mentioned spot; antennae dark brown with 
blue rellections. Pronotum black, bottoms ol 
most of the punctures yellow, with ihe follow- 
ing yellow markings; a thin median longitudinal 
Jim; from base to apex covering all bin 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 apex; a (bicker 
lateral stripe runs from the base 10 the apex: 
underside with alternate irregular stripes ol' 
bJack and yellow, the yellow predominating. 
Elytra with alternate black and yellow longi- 
tudinal stripes broken irregularly, iheir breadth 
conforming lo the intervals between the striae; 
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 each side across 
the outer margins of the abdominal sclerites 
anil at the apex, the black markinjra all with 
brassy metallic gleams; hairs silver. Legs dark 
brown with blue reflections; each femur with 
elongate yellow marking?; on both surfaces, 

5/m/k? und sculpture. Head with dense, uni- 
foitnly deep purcture*- Pronoium wilh uni- 
formly dense punctures; more or less parallel- 



sided but abruptly rounded at the opex. Elytra 
with the intervals between the striae trans- 
versely wrinkled, and deeply nnd densely punc- 
tured; parallel-sided from base to behind the 
middle then uniformly rounded in the area 
where marginal spines occur in the other 
sub- genus, then straight sided to 1hc apical 
spine which is minute. Undeisurfaoe closely 
and evenly punctured; wilh a lew short bails. 

fee. Male 11.3 x 4.1 mm (1>. 
Distribution. Western Australia. 
Central remarks. This species shows closest 
attinity with Attraeus polti. 

Specimens examined. Type only. 

3. Astracus (Depotlus) robustus sp. nov. 

FIGS 21, 2ZC 

Types, 

Holutype: <fi Payne's Find, W Aust, on 
Camarina rhehiana, I7.ix 1970. 5. Barter, 
SAM, I 20942- 

AUolype: '$. Payne's Find, W. Ausl., on 
C. dichima, t7,Lv 1970, S. Barker, SAM, 1 
20943. 

Paratypcs: I o. Payne's Find, W. Aust., on C. 
diehuwu.. I7.ix.iy70, S, Hitfke*. ANlC; 1 9» 
Pavnc's rural. W. Aust., on C. dietsiana, 
I7.ix.t970, .S'. Barker, BM; I -rf & I ? t 385 
km along Pavnc's Find Rd, W. AusL on C. 
diehitvia. i7.ix.l970. 5. Barker, MNHN; t 
6*. Payne's hind. W. An si. on C". diefsictna, 
I7.ix.1970, 5. Barker, WAM. 

Colour. Shiny. Hcatl. pronotum and elytra 
black wtih purple reflections. Antennae black 
with blue and purple reflections, Elylra with 
the following yellow markings: a longitudinal 
spot in the 2nd outermost interstice near the 
shoulder and another just before the middle; a 
similar but larger spot lies on Ihe 5lh Outer- 
most interstice after the middle but not reach- 
ing ihe apex; a large round spot before the 
middle with the width of the 3rd-6th inter- 
stice from the suture; a variable number of 
small spots at the base of the elytra in the 
middle, the maximum numher is four per side 
but they are absent in some specimens. Under- 
suilaee black with purple reflections; hairs 
silver. Legs brown: tfie joint between the femur 
and tibia black nn the upper surface; 1st tarsal 
segment brown at the base and black at the 
apex as are Ihe remainder. 

Shape and .sculpture. Head with, dense shallow 
punctures; with a glabrous longitudinal median 
line between the eyes confluent with the basal 
impressed line; bairy Pronotum sparsely and 
shatlowly punctured in the middle where the 
depressions are small Hut greatly increase in 
sue laterally where the non-depressed areas 



1(1 



S. BARKER 




* 



* 




ID 



I 



g 



^ 

m 




1 



H 



K£ VISION OF BUPREST1D GENUS ASTRAEUS LAPORTE ft GORY 



1 J 1 



arts glabrous; sides gradually rounded from 
base to apex: hairy, with hairs emerging singly 
in Ihe middle ;tnd in clumps from the lateral 
depressions. Elytra with intervals between the 
virkie slightly convex; more or Ises parallel- 
sided from base to just after the middle, then 
rounded gradually to ihe apc\; with sharp 
sumial spines; hairy. Undersurfaee covered 
whh punctures, shallow in die middle, deeper 
and denser at the sides; hairy, the hairs on the 
prosiernul process and presternum noticeably 
longer than elsewhere. 

Sfctf Males 15.9 ± 11,31 x 5.6 = 0.16 mm (4). 
Females I 7.0 ± 0.90 x 6.0 ± 0,25 mm (3>. 
Distnbuthm. Western Australia. 
Specimen,? examined. Types only. 

4. Astraens (Dcpollus) aberrant; van de Poll, 
1S86: 1 76; IRSY: R4, 90, 91, pi. 2, fig. 
4, 4a. Kenemans. 1892; 101; 1902:149. 
Carter. 1929: 2K2 . Ohenberger, 1930. 365. 

Asfraeas rt/irvtv>M.c vur pictkolUs van dt roll, 
IK8M;9l. Kenemans, 1S9M01. 1902:148. Car- 
tel, 1929:282, Obenberser. 1930:365. 
FIG, 22D 
Type, MNHN (not .seen by author). 
Colour. Shiny. Head pronotum and elytra 
hlack with purple reflections; antennae black 
with blue and purple reflections. Elytra with 
Iho following yellow markings: a small irregu- 
lar numbec of elongate markings each being 
restricted to the width of one interval between 
striae and occurringly mainly in three areas: 
near the suture; in the middle of the elytra; 
jiexi to the anterior part of the outer margin, 
Undersurfaee metallic purple, the lateral mar- 
gin tif the 1st, 2nd and 3rd abdominal seg- 
ments each with u yellow spot which is variable 
In she and absent from the 3rd segment in 
some specimens; hairs silver I cgs dark with 
blue and purple reflections. 
Shape and satlpntre. Head deeply punctured, 
in parts in Ihe middle less so than at Ihe sides. 
the areas involved glabrous; with u longitudinal 
glabrous line between the eyes merging with 
the basal impressed line; very hairy. Prono- 
tum with large deep punclure* dispersed but 
more so in the middle with the elevated parts 
glabrous; mostly parallef-vided 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, Elytra with the intervals 
between the striae convex and slight! v 
wrinkled, each With two longitudinal rows of 
small hairs; parallel-sided, rounded after the 



middle to the small marginal spine; suiural 

spine small. Undersurfaee evenly and shallowly 

punctured; hairy 

Size. Males 14.0 ± 0.2 x 4.9 ± 0.0* mm 

(25V Females 16.6 ft 0.43 x 5.3 * 0.17 mm 

(11). 

Diwtbutioth Western Australia. 

General remarks. This species shows closest 

affinity with Astravus mhusutx 

Sprritnctw exutitined. W. A list 1 r£, Carnomah. 
30ax.lS65, E. Baker; 12 <? & ft ?, 5 km W of 
Payne*s t'ind on CaM«*>1n(i sp , l7.ix.l970, $, Par 
ker; 2 o. 106 km S of Payne's Find on CttMutnKU 
conwulaia^ 18.ix.l l )70, S Barker; 2 ft Wialki on 
C. cortticatnfa, V)A\.V)10 t S. Rurker; I & Wialkt 
on Casaurtna campestris, 21.ix.l970, 5. Barker: I 
rfi South Taninun Flora Reserve on C, unnptstris, 
JO.ix.WO. S. Barker; 3JS*?, South Tamtnin 
Flora Reserve on C\ twmpesrns, fell .1**70, S, 
Barker; 3 J 1 & 1 % Quairadlms on C. eampesirit, 
7.xr*.l970, 5- Barker. 

5. Astraeu* (Depollus) lineatus van de Poll. 

1*S9: 84, 87-R9, pi. 2, figs 2, 2a. Kerrc- 
mans, 1892: 102; 1902. 149. Carter. 
1929. 2S2 ObenbeTger, 1930: 366. 

FIG. 22E 
Type- Holotype. MNHN (not seen by the author) 
Colour, Head and prunotum dull, elytra shiny. 
Head black with bronze-green reflections u\ 
the middle m the base, the Test with purple 
reflections, with or without ,» red spot under- 
neath each eye; antennae black with purple 
reflections. Pronotum black with bronze-green 
reflections in the middle, the rest with purple 
reflectrons: a thin lateral red stripe on each 
side: with or without a small red spot on each 
side of the median lobe near Lhe ba*e. Elytra 
black with blue reflections and the following 
red m^rking.s: two broad vJtrae, the first near 
the margin* but not touching it. running from 
the shoulder to the preapieul area; the second 
in the middle commencing near the front edge 
running parallel to the suture, bur not touch- 
ing it. to the prxapical area. Each of the vittae 
may be entire or broken into two or a number 
of elongate marks. Undersurfaee metallic- 
purple with lateral red marks on the pro- 
sternum, 3rd coxal plate and 1st. 2nd and 3rd 
abdominal segments, The presternal and 3rd 
abdominal spots are absent from some speci- 
mens; hairs silver. Lejis: femur and tibia 
metallic purple; tarvi brown. 

Shape and sculpture. Head evenly punctured 
with a longitudinal gljhrous line between the 
eyes; hairy. Pronotum evenly punctured, the 
punctures deeper at the sides than in the 



112 



S KAjttCER 



middle; sides narrowed and slightly rounded 
fumi the base to near the npex. where they nre 
more noticeably rounded; the Croat edge pio- 
jecling forwards in the middle; hairs longer 
and more dense at the sides than in the 
middle- Elvira with the intervals between the 
striae convex with shallow punctures and 
lainlly transversely slnguse, laterally slightly 
concave from the shoulders until after the 
middle, then founded to (he smalt sutural 
spine; hairy. Undcrsurface: presternum wit!) 
large puncture*, the rest with small shallow 
punctures. less dense in the middle: hairy. 

Sizes. Males J 1.0 * 0.46 x 3-S st 0.22 mm 

<3). Females 33.0 ± 0.29 >. 4.6 ± 0.U mm 

<6). 

Distribution. Western Australia. 

Specimens examined. W. Auxr.: 1 d\ 3 km W of 
Payne's Find on Cnsuaritnt sp„ 17. ix 1970, \, Bar- 
ker; 1 J, South Tainmiu Flora Reserve on 
Casuarina campestris< 30,ix.l97O, .S. Barker; 2 ?, 
Qutiirading on C. cumpe.stris, 7.\i, 1970. S. Barker: 
J £ & 2 ?, South Tammin Flora Reserve on l 
vamptslris. K.xi.1970. 5. Atflbr; 2 Si 13 km N of 
Goomnlltm; on C. campestris, 9.i.t97I, 5. Rarler. 

6 Wr;uus ( n«*|ji iHnM milHinotaius ■. '. r de 

Poll, 1889; 84. 89, 90. pi. 2, fig. 3, 3a. 
Kerremanx, IR92: 102; 1 902 149. Carter 
1929: 282. Obenberger, 1930. 366. 

FIG. 22F 

t v/h-. Holotype- MNHN (not been by the author). 
Colour. Head and pronotum dull, elytra >hiny. 
Upper surface and antennae black with blue 
reflections The whole uppei surface cccept the 
antennae speckled with small yellow spots. 
Undcrsurface black with yellow spots larger 
than on the upper surface; hairs silver, Legs 
dark brown; each femur vvilh a single elongate 
spur; the tarsi black, 

Shape and sculpture. Head wiLh shallow even 
punctures and a small glabrous spot between 
the eyes. Pronotum evenly punctured, the 
punctures larger at the sides than in the 
middle; slightly inflated after the middle. 
rounding and narrowing quickly to the apex: 
ihe I root edge projects forwards in the middle. 
Clytra with intervals between the striae eon- 
vex, punctured and wrinkled: laterally slightly 
concave, rounding well after the middle to the 
sutural spines which are turned out. Under- 
surface evenly punctured, but the punctures 
larger at the sides than in the middle, the 
differences pronounced on the thorax. Apart 
Inim the legs, apical edges of the abdomen and 
pronotum, the body is devoid of hair. 



Size. Males 12.3 £ 0.73 x 4.9 t 0.3 mp 1 7 1 ( 

Females 15.I ^ 0.95 x 6.0 ± 0.35 mm »6>. 

Dntributitm. Western Australia. 

Specimens examined. W, Aust t A Leonota. 
H W. tfr.ww!. 2.0.1934, WAM, 34-511: I £ W 
km 8 of Coolyardte, ian. 1937, A. A*. P4Utlu& 
WAM, 37-722; 1 j & t fc Tmin ROtf, UJHOi 
A M. UoUiitwr. WAM, 40-32, 40-31. 2 tf t Wat- 
nint;, 20.ix.l950, R. r, MtMUhu % WAM, 73-3*4 t 
73-386; 2 j£ -K I % Waininis, 2b.xiU9S(i. ft. f. 
McMillan, WAM. 73-385. 73-3S8, 1 £ & 2 ft 
WAM, 73-375/7: I % Red Gum Pass. Stirling 
Ranges On Castutrina sp- 2G,i. 1971. £', <£• A', Cor* 
nahy . 

7 Astraeus (Dcpullus) irregularis van de Poll, 
1889; 84. $6, 87. pi, 2, fig, 1, la. Kerrc- 
mans, IS92; 102; 1902: 149. Caller, 
1929: 232. Obenberger. 1930: 36h, 

FIG. 22G 

type, Hnlotypc, MNHN (not seen by author.) 
Colon/, Head and thorax dull, elytra .shiny. 
Male. Head black with blue reflections, with 
the following, yellow markings: a continuous 
hand under |he lower half of the eye; an elon- 
gate spot in the cenfre of the head which may 
he broken into several spots; two elongate spots 
lateral and basal to the central spnr, each of 
which may be broken into two spots: a small 
spot on either side of the gular, Except for the 
central spot, all other spots may be reduced 
oi absent. Antennae dark brown with blue 
reflections. Pronotum black with blue reflec- 
tions and the following yellow markings: a spot 
on the margin of the apical edge on each side 
of the middle; a ^pot on each side below the 
first, towards the base, two spots along the 
sides, a small one at the apex and an elongate 
larger one at the base — the first may be absent 
and the second broken into two spots. All of 
thc.se spots may be reduced and some or till of 
them absent. Hlytra black with blue reflections, 
each elytron with the following yellow mark- 
ings*, the basic pattern is of three spots in the 
middle but not touching the suture; Ihe lirsl 
is a small elongate spot at the base; the second 
a large spot before the middle; ihe third a virta 
after the middle and extending to the pie- 
apical area; Iben: are also two small donjratc 
spots at the margin, one at the shoulder extend- 
ing U> the legion of the humeral fold and a 
smaller one just behind it which is sometimes 
ahsent; an irregular number of small spots are 
present along the basal edge between the vittii 
and the sulure. Undersurfnce dark brown with 
blue reflections; with yellow spots at the base 
of each leg and single lateral spots on the pro- 



REVISION OF Bl.'PRFSl ID GENUS ASTKAKVS T.APOKTE At GORY 



115 



sternum, thud coxal plate and abdominal seg- 
ments I— 4-, 1—3 or 1—2. decreasing id size fTom 
first to last; hairs silver. Legs dark brown with 
blue reflections. 

Female. "Head, pronotum and elylra basically 
similar to the male except thai the yellow 
markings are larger and the irregular spols are 
more numerous. Undersurface and legs as in 
the male except that lateral spots are present 
on all the abdominal segments;. 

Shape (put sculpture. Head broad; with shallow 
punctures larger at the sides than in the middle 
and a longitudinal glabrous line between (he 
eyes. Fronoturu with shallow punctures larger 
at the sides than in the middle; the lateral mar- 
gins rounded and narrowed gently at (he base, 
more abruptly at the apex. Elytra with the 
intervals between the striae slightly con- 
vex, with a shallow row of punctures ami 
taiully transversely wrinkled; parallel -sided 
until after the middle, then rounded to the 
sutural spine which is well developed ai>d 
iurntd out. Undersurface more closely punc- 
tured at the pdfe than in (he middle, with a 
few short hairs. 

Size Males 1 1 .2 ± 0.24 x 4,1 - fi.OK mm 
(II >. Females 13.5 ± 0.22 x 4.7 ± OM mm 
(ID). 
Distribution. Western Australia. 

Specimens examined. W. AuM-: 3 & & 5 ?. Cion- 
nclk on Casuarirui up., 24."V»k19fi7, S Barker; 1 
tJ & 2 9, Gosnclls on Casuavina sp . 27.xJi, 1967. 
■V. 'Darker; 2 S A 3 ?. Red Hill Rd near Midland 
Junction on Casuurina sp.. -4.i.l9rJS, S. Barker: 2 
?. 77 km along main York Rd on Casuatina «p. t 
5.i I06R> S. Barber, 3 4 ik 4 ?, South Tammin 
Mora Reserve on CasuaruHt hne^cliana, C 1,1971. 
S. Barker; I & & 2 8; 142 km E of Norseman on 
C, fuu'Reliarut, 19.xii.1972. S, Barker; 1 ft 58 km 
W of Tammin on York -Tammin Rd on C. 
WeWfVv/w, 23 \x\U97t, S. Barker; 1 rf •& 1 5. 3 
km E of Gosnclls on C. lutegciiana, 27.xii 1972. 
.V. frmm 

K A stratus iOej#ollu^> dednrieiiMs sp. nov. 

FTGS 4, 22M 

Typ?!- 
Holotype* tj? DeuVil, W. Aust , fi t W. Brvwa. 

WAVf, 71-1779. 
Allotype: ?, Dcctari. W. Aust, H, W Browrn 

WAM, 71-1778. 
Pamtypes: 7 ?, Dcdari, W. Aast., WAM, 
40-1207, 40-1208; 1 f, Borden, W Aust., 
SAM: I ?, Deduri. Vv\ Ausu, AN1C; 1 ?. 
Dedari, W. Aust. t on Camanrta Lwmeulatih 
21X1936, fj W, fa own, MNHN. 
Colour, Shiny. Head black with purple reflec- 
tions and the following yellow markings; on 
ovo>ii spol touching the underside of each eye 



and pointing downwards but not touching the 
mouth; a single circular spot In the middle. 
Antennae brown with hlue and purple reflec- 
tions except fur the first segment which is pre- 
dominantly black. Pronotum black with purple 
and blue reflections and the following yel low- 
markings: a small ovoid spot on each side at 
the apex; a 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 
each tide from the apex to the base where it is 
extended by the flaring outwards of the lateral 
margin of each lobe exposing the yellow inner 
surface Elytra black with purple and blue 
reflections 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 befote 
the middle and ending al the. middle, close to 
the suture but not touching it; an elongate spot 
commencing alter the middle and running to 
the apex, close to the suture but not touching 
it; an elongate spof at. the shoulder, not touch- 
ing the outer margin hut close to it, behind the 
last a much smaller spol not touch- 
ing die margin: after ihe middle there are 
several elongate %poK intending almost to the 
apex but not touching the outer margin; several 
small eccentric oval spots. Undersurface black 
with purple TcrtcetLons; the first three 
abdominal segments have lateral yellow spots 
which progressively diminish m size from in 
fronl backwards; hairs silver. Legs; raeso- 
sternal and mctastcrnal coxae with yellow spots 
on the outer margins; the rt$t brown with 
purple reflections; the upper surfaces of the 
tarsi are black. 

Shape and sculpture. Head with shallow punc- 
tures and without hair. Pronotum with shallow 
punctures; a short median impressed line 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 until after the middle then 
rounding off to the apex; each ending in a 
Lhiek short spine projecting backwards; no 
marginal spine but a small mound is preseni in 
the region where they occur in other species. 
Undereurface with shallow punctures; with a 
lew short hairs, 

Size, Males J 1.2 .\ 42 mm 111 Fem.Ves 11.0 
± 0.15 :< ftp m 0,1 mm (6). 

Distribution. Western Australia. 



2 
15 



114 S. BARKER 

General remarks. This species shows closest long. Pronotum deeply notched on both sides 
affinity with Astrueus irrcyttlctris. at the base, with the median lobe and the two 

Specimens examined, Types only, lateral lobes pointed, Elytra striatc-punctatc 

and/ or costute, the outer edge of the intervals 
SflgpgftfH$ AstratMi* (seiisu stricto) sub-gen. between the striae costatc, either down the 

nov# entire length or towards the apex only. The 

Type species Astweus flovopiclus LaPorte & Sl j flirc cn <j 5 posterior | y in a spine and in ai( 

Uory, 1837. but three species these are sharp and curved 

Description. Head medium sized: with or with- outwards, The marginal spines are smaller than 

out a median longitudinal raised ridge between the siitural spines. The humeral fold is variably 

the eyes, the median keel (Fig. 25). Antennae developed and angled (Fig. 1). 

Key to the species of AstraeuS (sensu slrklu) 

I. Head with a median keel ..,,_.. 

l.Head without a median keel ... ,., ....... ..... 

2. Hairs silver . , ... _ 

2. Hairs yellow ., ,., , ,.,. ., )* 

x Part or all of the anterior nndersurface red-brown 4 

.3. None of the anterior undcrsurfaec red-brown 6 

4. Guiar, prosternum. meso- and metastcrnum, 2nd and 3rd coxae and 1st abdominal segment 
red-brown _ , „ .. . .. . . <J. hakeri sp. nov 

4. Prosternum and coxae red-brown ,..,.,,..,. - 10. minutu.\ sp. nov, 

4. A red-brown area on either side of the proslermd process 5 

5. Usually shorter than 7.5 mm; humeral fold well developed and acutely angled , 

II. frasniensis sp. nov. 

5. Usually longer than 7.5 mm: humeral fold moderately developed and angled 

15. ohsaiYux sp. nov. 

6. Most of the leg testaceous . . .... 7 

6. Tips of the tibia and 1st tarsal segment testaceous . .. . ... 8 

6, "None of the leg testaceous .. .. -.,.- .._ .. 10 

7. 1st and 2nd legs testaceous except for the outer margin of the femur, 3rd leg testaceous except 

for the femur . „ . . '. _ ...... 19. dlhttipt's van do Poll 

7. Tibia, 1st and 2nd tarsal segments testaceous , , 13. ,s?nythi sp. nov. 

8. Usually shorter than 7 mm 12. pypnaeus van dc Poll 

8. Usually longer than 7 mm - 9 

9. Humeral fold well developed, acutely angled 17. nmstersi Mad env 

9. Humeral fold well developed and angled !8. samoueUi Saunders 

10. Humeral fold moderately developed and angled II 

10. Humeral fold poorly developed, slightly angled .,., ,. 13 

I I. Head and pronotum green and coppery-purple 21. intrkaius Carter 

IT. Head and pronotum black .. ... ... ... „ .. 12 

J2. Broad and rounded species 16. gloko.\u.s sp. nov. 

12- Elongate species ....... .,_ _ ,,. 35. watxoni sp. nov. 

13. Black species; elytral markings a number of yellow spots 22- crasxits van de Pull 

13- Blue species; elytral markings two yellow fascia 25. fraternrius van de Poll 

14. Elvira! markings three yellow fascia and red ureas 23. major Rlaekbuin 

14. Elytral markings iwo yellow fascia without Ted areas .. 24. navarchis (Thomson! 

t£. Body elongate and cylindrical . I IS 

l5.Body not elongate and cylindrical , . , 17 

16. Pronotum conically elevated in the middle 26". prothoma'ats van de Poll 

16. Pronotum not conically elevated in the middle 27- slnntwttts van dc Poll 

17. Su rural spine with a rounded internal edge . IR 

|/i Suturaf spine with a straight internal edge 26 

J8. Legs a red-brown colour IV 

18. Legs not a red-brown colour 20 

19. Elytral markings spots and fascia 30. mmrnHhmi sp. nov. 

19. Elytral markings two vittae on each elytron ,., 28. vttuittts van de Poll 

20. Hsad, pronotum and legs metallic brown or hron/.e , , 29. ffavopictus LaPorte & Gory 

20. Head, pronotum and legs not metallic brown or bronze 21 

21. Humeral fold well developed and angled . 22 

21. Humeral told poorly developed, slightly angled ...- ,-. 23 

22. H-ad black or coppery-purple; undersurface cc-pp^y-purple • 20. adamsi sp. nov. 

22. Head blue or green; undersurface blue-green . ... 14. Abnuhi/i/t van de Foil 



REVISION Oh BUPRESTID GENUS AStttAEVS LAPORTE k GORY 



115 



•■■• *..- • .. 



31. carnuh\i tip. nov. 
24 



1} Basal spot expanded along the trout edee of ihe elyuou ... 

23. Basal spot not expanded along Ihe front margin of the elytron 

24 l £Jytral mark/rigs eitlici two spots and two fascia oi tbt first fascia may be broken making 

lour spots and a fascia on each etvtron ... 32. bndenl van de Poll 

24 Flytrnl markings either $rx spots and a fascia or the fascia may be broken iuaJcnie eight 

spots on each elytron 33. jansonl van de Poll 

24. Elytra! markings 7 spots on each eJyiron ... 25 

25. Kronotum parallel-sided from bass? to the middle then strongly rounded and narrowed to the 
apex; dorsally convex in lateral profile ... - 34. edwrthuri van <Jc Poll 

25 Vronotum gradually rounded at the sides and narrowed from base Co apex; domily flattened in 
lateral profile , . . .. _, ..... _ _ s6, carter! sp, nnv. 

26. Head .slightly excavated between the eyes , .. ... 37. goertingi sp. nuv. 

26. Head deeply excavated beiwccn the eye* 27 

27. Head excavated mainly at the base; propotOiU laterally inflated with an oval patch of hexag- 
onal cells in the middle .... . . 38. e.yaneus Kerremnns 

27. Head excavated mainly ai. the apex; pronotum not laterally inflated and without hexagonal cells 

39. catedoniciis Fauvel 



9. Astraeus (Astraeus) bakeri sp. nov. 
FIGS 5, 221 

Holotypc: i Dryandra, W. Aust. on 
C'ostfarifiii hucgeliana, 8,xii,l£70 ( S, Barker. 
SAM, 1 20945. 

Allotype: 9, DryandrB. W. Aiixt. on C*. /»«<#•*• 
H«hu> 8 xii.1970, 5. Barker, SAM, I 20946. 

Furalypes: 1 <f & 1 ?, Dryandra, W. Aust on C- 
hueveUann, i2a\l973>S. B#tker t EA; 2 ?. 
Toompup, W. Aust. on Casnarina sp., 7t. <£ 
K, Cartmhy, KC*. 2 ?, Dryandra, W. Aust. on 
Casuariaa sp., 2. i. 1968. S. Barker, SAM; 4 
3 & 4 ?, Drvandra, W t Aust. on C, hong* 
liana, 8.xu\1970. S, Barker. ANIC (1 <j & 1 
2), BM (1 $ &. \ 2), MNHN < \ tf & \ g)', 
WAM (I f& 1 ?). 

Cntottt, Shiny. Head and pronotum black with 
vtolet or green reflections; antennae black with 
metallic blue 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 foJd 
and ending near the suture but not touching it. 
broken into spots in some specimens: a liirge 
spot below the middle, Undersurlutcc; gulaf, 
pro- meso- and meiasternum, second and third 
coxae and median anterior part of the first 
abdominal segment ail reddish -brown with 
light blue rellections on the lateral part of 
the above; remaining abdominal segments 
black with metallic blue reflections; hairs sil- 
ver. Legs brown with metallic blue rellections 
on the upper surface, the end of the first tarsal 
segment and all of the remaining tarsal seg- 
ments metallic blue, 

Shape and tfcttrpwre. Head evenly punctured; 
small median keel; h.iiry. Pronotum deeply 
and evenly punctured, with a median longi- 
tudinal impressed line at Ihe apex; median 



lobe without a basal crypt; sides rounded and 
narrowed from base to apes, hairy, Elytra 
costalc. the intervals between fiat with a pro- 
minent row of punctures; at the sides gently 
rounded from the base to just before the 
middle then rounded and narrowed to the 
strong marginal spine: sittural spine rohust; 
humeral fold we'l developed .ind angled. 
Undcrsurfacc sparsely punctate in the middle, 
more closely at the sides; covered with fine 
hair which is denser at the sides than in the 
middle. 

She. Males 7.5 ± 0.07 x 3.0 as 0,03 mm (44). 
Females 7,9 i 0.12 x 3.2 ± 0.06 mm (23). 
Distribution. Western Australia. 

General remarks. Named after Dr F. H. Uther 
Baker. 

Specimens examined, Wr Aust.: Types; 18 <? & S 
2. Dryandra. on Casuartna h nest?) iana, 8. xii.1970, 
S. Barker; 6 d & 1 ?, (3 km F of Ongcnip. on C 
W\eW/Vuw, 11, i. 1973. S. Barker; 6 g & 3 5, Dry- 
andra, on C. h/texcliana, 12.1.197*. .V. Barker: 1 & 
Ponicr rockhole 70 km S of Balladonia. on C. 
Inwpeliana. S.xiL 1974, 5. Barker; 3 rf. Jurartda 
rockhole 106 km S of Balladomfi, 00 C. huei:** 
liarta, 9\ii.lV74 T S. Barker; 1 ^. 115 km S of 
Bnlladonin on Casnarttut humitis. 9.xiiM974. S~ 
Barker; I 6*, SW valley of Ml Rafieed,' on C 
huepcliumi. I0.xii.l974, S. Barker; » 9, 3 km S ot 
Mt Ragged on Israelite Bay toad, on C. lutmiffx, 
KUii.1974, S. Barker. 

10. Asfraeiis (Astracus) minufns sp. nov. 
FIGS 6, 22J 

Types. 

Holotypc: d\ South Tammin Flora Reserve. W. 
Aust. on Casuartna hue^eltana, 6.i. 1971, &> 
Barker, V SAM, T 20459. 

Allotype; ?, Souih Tammin Mora Reserve, W. 
Aust. oa C. huexetiana, 23.xii.l972, .S Bar- 
ker, SAM, I 204/V0. 

Panjfypcs: 1 .J t Soalh Tammin Flora Reserve. 
W. Aust. on C. huexttinna, 23.xii.1972> Si 
Barker, ANIC (1), MNHN (1); 1 «?♦ South 



116 



S. BARKER 




K 







1 

I 






ft 




5 

^ 



REVISION OF BUPKfcSUD GtNUS ASTRAEUS MPOftTB & GORY 



11*7 



Tammin Flora Reserve, W. Aust, on C 
huv-elkma. 6.LI971. 5. £a/A?r. SAM: 1 jjf, 
South Tammin Tloni Reserve;. W. And. on 
C. tlUdf&IUMA 9.i.l9?4, 5. MrWi SM; 2 <?, 
South Tammin Flora Reserve, W. Aiist. on 
C tfflmgpAfrfo J>.il974> A, ^tifte/i WAM. 
Colour. Shiny. Head black with purple reflec- 
tions; 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 mdldlc, red-brown at 
(he sides. Flytra black with purple reflections 
each with the following yellow markings: a 
large basal spot; a fascia before the middle 
covering the humeral fold bui not reaching Ihe 
suture; a large median spot after the middle, 
Undersurfacc: presternum and coxal plates 
red-brown, the rest black with metallic purple 
reflections; hairs silver. Legs: upper femur red- 
bnwn, the rest of the femur and upper part 
of the tibia melallic purple; apical part of the 
tibia and first tarsal segment testaceous; 
second. Lhird and fourth tarsal segments black 
with blue reflections. 

Shape (ind sculpture. Head broad; deeply and 
evenly punctured, with a median keel; hairy. 
Pronotum deeply and evenly punctured, with 
a short impressed line projecting forwards 
from the basal crypt; sides gradually rounded 
from base to apex; hairy. Trly tra costate 
the intervals flat each with a row of very 
shallow punctures; laterally tapering gradually 
from base until after the middle, then rounded 
to the well-developed marginal spine; a vt*ry 
elongate crcsccntic cmargination between the 
marginal spine and ihe well developed sutural 
spine; humeral fold well developed and 
acutely angled. Undersurfacc evenly punc- 
tured; hairy, 

She Males 5.7 ± 0.22 x. 2J * 0.10 mm (7). 
Females 64 x 2.7 mm (I). 
fiistrihutUm. Wcslern Australia. 
Gaieraf remark*. The smallest species known 
at this time. 

Specimens extfflv'Wii Types only. 

1 1 . Astraeus (Astrueus) fraserfrnHs sp. nov. 

FIGS 7. 22K. 24. 25 
Types, 
Holotypc: £ Vraser Range, W. Aim on 

Casuarina hfitwft'oua, J*>,x;ii.t972, S. Barker, 

SAM, 1 20951. 
Allotype: 2, Frnser Range, W, Aust- on t\ 

km&IImm* l5xiiW2. S. Barker. SAM 1 

20952, 
PaTHivptrs: 2 cf & 2 5, Dryandia, W Aust. on 

C AflteftWtaM 111 1^75, S. Barker. EA (I J 

& I 5). MNHN (I cf & I 2): 2 $ & 2 ?, 

Fraser Ran*e, W. Aust. on C Imey.eliamt 



I9.dii.1972. S. Barker, ANTC f I cf & I 9K 
BM (Id 1 * 1 2}-: 1 A J-ate Grace. W. Aust., 
9.L1972. E. A K. Caraabv, KC: 3 & Fyre 
Highway U2 Krh E of Norseman, W. Aust. 
on C. huetreliana. I9.xii.l972. S. Barker, 
SAM; 2 6 &. I 9, South Tammin Flora 
Reserve. W. Aust. un C hueselitma. 6.U971. 
V. Barker, WAM. 

Colour, Shiny. Head, antennae and pronotum 
black with violet and blue reflections. Elytra 
black with purple reflections, each with the 
following yellow markings: a large basal spoi 
in ihe middle; a fascia covering the humeral 
fold, running towards rtie suture but not touch- 
ing it, and slightly concave towards the base: a 
large spot after the middle not touching the 
suture or the margin; sometimes there is a 
preapical spot. Undersurfacc metallic purple 
e<ccpt for a red-brown patch on either side 
of the proslernal process at the anterior edge 
ui the presternum: hairs silver. Legs: femur 
metallic purple, tibia and first tarsal segment 
predominantly testaceous, second, third and 
fourth tarsal segments dark with Wuc reflec- 
tions. 

Shape and sculptutt. Head closely and evenly 
punctured; with a median keel; hairy. Prono- 
tum closely Hnd evenly punctured with a 
slightly excavated median longitudinal line 
protecting forwards from the basal crypt. 
rounded at the aided and narrowed from has? 
lo apex; harry. Elytra costate* the intervals 
between flat, each with a row of punctures; 
parallel-sided until after the middle then gently 
rounded anil narrowed lo the marginal spine; 
both spines well developed; humeral fold well 
developed and acutely angled, Undersurfacc 
sballowly punctured in the middle, punctures 
deeper at the sides: hairy. 

SSte Males 07 ^ 0.05 x 2.6 ± O.02 mm (72). 
Females 7.0 ± O.ritS \ 2.S =r 0.U4 mm (2S) . 

Disitibnrton. Western Australia. 
General remarks. The three speciei Atfraeus 
bakvri, Asrraeits mfrwrus and Astnwu* fra&et* 
tenxis show a close affinity to each other. 

Specimens' examined. W. Aust.; Types: I V. 58 km 
VV of Tammin on York-Tanuniii Rd, un 
Camarina hue a? litt nu, 23.xii.1972, S. Darter; I o- 
58 km W of Tammin nn York-Tnmmin Rd, on C, 
hu<yyha,ut, Sh|J9H & Worker; 2 3, South Tam- 
min Flora Reserve, on C. bucgrfiana, 23.xii.l972. 
5. Barker; 4 cf & 1 % South Tammin Flora 
Re8crve, on C. tiueseiiutw, 9. 1 1974, S. Barker; 
31 cf & I* y, Gwumh rocVhole riri km $ of Halla- 
doniu, on C huefiviwntt* 5.xii.iy74, S. Barker: 
18 rf & \3 a Punier rockhole 70 km S of Bnlln- 
donia, nn C'. hur^rliatuh 5.xiU974* S. Barker: 
5 S* Ju/anda jockhole 106 km S of Balladouiu, 
on C huecetiana. 9.xii.l974, ffi Barker; 1 & 19 



Ilfi 



S. BARKER 



km W of Balladonia. on C. tuimifis. 23.ii.l97S, 

V. Hotkey, 

12 \fitraeus (Aslraeus) pygmaeus van de Poll, 
1 8Kb I7K-IH0; ISS9: 85. 104. 105, pi. 3, 

figs In*, 16a, loc, Blackburn, IR91: 496. 
van de Pod, 1892; 67, 68. Obenoetgor, 
19*0; 366. Carter, 1931: 107; 1*3,3: *!. 

Astt tti tts pvginui'ii\ var. suhjawiutus van dc 
l»olK 1886' 178-180; 1889: 104. 105, pi 3. 
fig. 16b. ObeiibcrRot. 1930: 366. 
Astrnfus ftrftfrwWfci v^r. pvxmaeitt. filackbarn, 
ISyO: 1256. 

Asnaan stunoitelli var. pvyhuutis. Kerrcrnans. 
1*92: 102, 

Antaeus py^iaaefis: Kerremnns. 1902: MK. 
4sf/v*c(/\ pxt>muens var. sithfasciatt<s: Kerre- 
mrws. 1 002: MH. 

AstriH'H.s p\)>tntu<t(\: Carter. 1929: 282. 
A stracu v f v eutacus var. subfaxrmtus: Carter. 
1920: 282. 



FIG 



Type, liolotype: ^ Queensland, MNHN (seen by 
author). 

Colour. Male. "I urquoise with golden-green re- 
flections at the apex, dark at the base wilh 
purple reflections: antennae datk with blue 
reflections. Pronotum dark in the middle with 
golden-green and purple reflections, turquoise 
at the sides with golden-grc.cn reflections. 
Elvira black with blue reflections, each with 
the following yellow markings: a spot at the 
base; u fascia covering the humeral told. 
slightly enneave towards the base but not 
touching the suture; a spot after the middle, 
not touching the margin or the suture. Under* 
surface metallic turquoise in the middle, metal- 
lic blue at the sides: hairs silver l^egs metallic 
hi tie: tips of rihiu and first tarsal segment 
testaceous; tarsal segments 2. } and 4 dark 
brown wilh bine reflections. 
Fentv!e Head blue at the ape*, dark at the 
base with purple reflections; antennae black 
with blue reflections. Pionotum black in the 
middle with put pie reflections, blue at the 
sidev Elytra os in the male except that the 
fuxeta is always broken in the middle* to form 
two spots, and all of the yellow markings are 
smaller than those in the nude. Undeourface 
deep metallic blue; hairs silver- Legs deep 
metallic blue: tips of the tibia and iirst tarsal 
segment testaceous. 

Shape and St ulpturv. Head evenly and closely 
punctured; with a median keel; hairy. Prono- 
tum closely punctured, the punctures larger at 
the sides than in the middle; parallel-sided 
from the ha.se until just before the middle then 
tounded and tapered to the apex; hairy. Elytra 
custutc. the intervals tint, cadi with a row of 



punctures and transversely wrinkled; parallel- 
sided until after the middle then rounded and 
tapered to Ihe marginal spine: both spines well 
developed; humeral fold well developed and 
angfeo. Undersurface: prosier nal punctures 
larger at the sides than in the middle, punc- 
tures uniform on the abdomen: hairy. 
Size. Males 6,3 '- 0.05 x 2.5 ± 0.02 mm (66). 
Females 6.6 at <j.08 x 2.6 ± 0.04 mm (40). 

Distribution. Queensland and New South 
Wales. 

Specimens ewtiitied. QlcL: Type: I r? & t ?, Edun 
galta on Ltuuur'ma sp., I4.xi.197l, E. E, Adam*.: 
8 <i & 4 ?, Oduitgalba Oil Casaanna sp., 9.xi.1974. 
E. E. cE J. Adams; I rf & 1 9- Eduncalba -on 
f asuarina sp., 1 5.*ii. 1974, E. E it ft Adatus 
ttJLW* 41 r? & J 9 <?. Oranrncii on Casuarina 
iittordit'j. 9.xii.!962. S. Baiktr; 2 cf & 2 ?, 
Majors Creek on C. lUtoralis, 26a. 196"*, S. flit/ - - 
ker. A.C.I.; Le?& I 9, Mt Ainslic on Cuxuanau 
stritta, 29.xl.l962, S. Barker: 6 rf & 2 2. Mt 
Ainslic on C. .stricta, 26.xii.l962, 5. Barker; 4 ^ 
& 3 $\ Tugeeninnng on C sfrirfo, l,i VXi,\, J, Bar- 
ker. 



13. Astraeu* (Asfraeus) stnythi sp. nov, 
FTG8 8. 22M 

Types. 

Holotvpe: rfi Maryborough Qld, November 
1956. C. Smith. WAM. 7.V55. 

Allotype: ?. WAM, 7.V56. 

Paratypes: 2 % Qld, E. Sutton, QM; I rf. SAM. 
Colour. Mate. Head: apex blue with golden- 
green reflections, base dark blue with purple 
reflections: antennae black with blue and 
purple reflections. PrOnottun black in the 
middle with purple reflections, blue at 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, run- 
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 
testaceous; third and fourth tarsal segment* 
dark brown. A small preapieal spot may or 
may not be present. 

Female, Head; apex turquoise, base green; 
antennae black with blue and green reflections. 
Pronotum green in the middle, turquoise at the 
sides. Elytra black with blue and purple rcflcc- 
tioas. yellow markings as described in the male 
but less prominent. UrKtersutface and legs as 
in the male. 



REVISION OF BUPRESTID OFNl-S A5TRAEUS LA PORTE & GORY 



IH> 



Shape and sadputn. Head evenly punctured; 
a median keel; hairy. Pronotum with punctures 
latget 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 lo the apex. Elytra 
costate, the intervals between flat 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. prostcrnal punctures shallow in 
the centre, deeper at the sides; punctures on 
the abdominal stemites sparser and shallower: 
hsiry. 

Size. Males 6,5 ± 0,08 H 2.6 ±0.15 mm 1 2) 
Female* 6.5 ± 0.18 x 2.6 £ 0.01 mm 14) 
pi\?rihnti'->n- Queensland. : 

Genera! remarks, A WW hi shows cfnse 
affinity with A, pygmaean. I place these species 
alongside *he previous group of three species 
Named after my late colleague, Dr M. Smyth. 

SpecitHwt* vxatmnefL Qld : Types; I 2, One Tree 
Hill, Brisbane. 1920. i\ Mwr, nPBM. 

14, Astraeus (Asiraeuil si initial or van dc Poll. 
188°: 85. 102. 104. pi *. rigs 15. |5& 
15l>. Keiremans 1892; 102. 1902; 148. 
Carter. 1929" 282. Obcnbcrgcr, 1930; 366. 

FIG 22N 

J y/if* Hn|otypc 9, Peal Downs, MNHN (seen by 
:i u tlioi I . 

Colour. Malt'. Shiny. Head green at the apes 
with golden-green rcllecions, black at the base 
with blue reflection*: antennae black with 
golden-green and blue reflections. Frenulum 
hlack with blue and purple reflections- Hlytra 
blciek with turquoise reflection*, each with the 
following markings: a large nasal spot", a lasCta 
covering the humeral fold, running transversely 
Towards the suture but not touching it. concave 
towards (he base and clubbed ut ihc end; u 
fascia after the middle running transversely 
from the margin but not touching the suture. 
Undersurface metallic blue: hairs silver. Legs 
metallic blue; tips of libia and first tarsal seg- 
ment brown: second, third and fourth tarsal 
segments black with blue reflections. 
Female. Head black with blue reflections: 
.antennae black with blue-green reflections Pro- 
notum black with blue reflections. Elytra, black 
with blue and purple reflections. The rest as 
In the male. 



Shape and sculpture. Head with close uniform 
punctures; no median keel; hairy. Pronotuin 
evenly punctured; basal third parallel-sided 
then slightly rounded and narrowed to the 
apes; anterior margin project big forward in 
the middle, that general area glabrous: hairy. 
F.l\lra eostale. the intervals between flat but 
slightly wrinkled; parollcl-sided from the base. 
rounded before the middle then tapering to the 
marginal spine; both spines well developed: 
humeral fold well developed and angled. 
Undersurface evenly punctured; hairy, 
5a. Males 6.7 ± 0.10 x 2.5 ± 0.05 mm ( II >. 
Females 7.0 - 0.24 x 2.7 ± 0,09 mm (6)- 
Dhtriluition, Queensland. 

General remark* Of the 17 specimens I have 
examined, 7 c? and 2 ? have preapical spots 
on the elytra, the other eight specimens have 
no preapical ivpots. Although male genitalia 
( Fig. 22N ) is similar in shape to the previous 
two groups of species, the external morphology 
of A, simulator is dissimilar to those five 
species. Because of this 1 do not grnup it with 
any of the above mentioned species. 

Specimens examined. Qld: 'lyre: 1 c?. 3..xii.l 07 1 _ 

E. E. Adams. KC, 4 <f & 1 9, fcdungalbn on 
Casuarhui ccfuisettlolia, 1 l.xii.|973. E. E. Adams. 
SAM (t rf & I 9>, EA !?»: 1 2, Fdunjulba on 
C. equisrtifoth, 15 xiU*>73. t E Adam*. EA. 2 
rf, Edungalba on Caxuar'ma sp.. I on each o( 
23JU.1974 aUtl 8jrf|4$H E, E, <* 5. dtfeWi HA; 

1 <?. Ediingalbfi on Casuarina sp., 30. xi. 1974. E. E. 
*• ,V, Adams, EA; 2 S & 2 t & Edungulbn on 
Casuarina sp., M.xii 1974. E. E. i S. Adams-. EA; 

2 c?» Edtinjealba on Ca&uarirta sp., 15.xii.l974. 
E. E. tfi S. Adams, EA. 

15. AsflnAn (Astraens) onscurus sp. nov. 

FIGS 9. 220 

J ypes. 

Hototype: 3*, Eraser Range, W. Aust.. un 
Casuariua htuKvu'ana, ]9.AJi. 1972. &. BarUvv. 
SAM. I 20957. 

Allotype: ?, Eraser Range, W. Ausl.. on <"*- 
Imefiehww, I9.xir.1972, ,y, timier, SAM, ) 
20958. 

Paratypes: 2 J &. 2 £ Eyre Highway 142 km E 
of Norseman. W. Aust. on C . Iiue^eh'iwu. 
I9.xii.1972, S. Barker, EA U 6* & 1 $K 
ANIC ( I 6* A i ?»; 2 o* & 2 ?» rrftttr Range. 
W. Aust. on C. huik'eliaua. I9.xii. 1972. 5. 
Barker, BM (1 o & I 2). MNHN ( 1 rf &. 1 
¥>; 1 ri\ 24 km NE of Lake Grace. W Auhi.. 
on C. ht/wtiana. 24.U973. 2f, Barker, KC; 
1 ^, Tammin, W. Aust.. //. W. firowih 
WADA; I 9. 2 km N of Nccdilup, W. Auu.. 
23.xiiJ972, K, Kewhy. WADA; I 9, 30 *m 
W uf No. 8 pumping Hlattim ( Kalgoorlie 
vvalcr m;iin) on Casuarhta >'ampesirts 
24.I.IM5X, WAMs 71-1775; I ?, WAM, 7t* 
1777; 1 S; South Tammin Flora Reserve on 
C fmmttmi* 23.xii.l972. .?. BarUt WAM 



1 SO 



S. BARKER 




i 

i 

O 



I 



; 3 ' 




1 



^ 

rf 




£ 
in 



I 
* 
1 



I 



REVISION Of BUPRRST1D GH-KUS 4&T&4&V& LAW1RTE & GORY 



.2. 



Colour. Shiny. Head purple or black with 
metallic purp/e reflections at the apex, metallic 
blue reflections at the sides and underneath; 
antennae black With blue and purple reflec- 
tions, tips of first, second and third segments 
dark brown. Pronotum purple M 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 fascia before 
the middle concave towards the base commenc- 
ing at the outer margin, enclosing the humeral 
fold, tunning towards the suture but not touch- 
ing it; a broad fascia after the middle but. not 
touching the outer margin or the suture; a very 
small preapical spot in the form of a lunette 
sometimes missing. Undei surface black with 
purple reflections, except, lor a red-brown 
patch on either side of the prosternal process 
extending lo the laleral margin; hairs silver. 
Legs: femur, basal half of the tibia and 
second, third and fourth tarsal segments dark 
hrown with blue and purple reflections, apical 
hall' ul (he libia and first tarsal segment brown. 

Shape tmd sculptta?. 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 to the apex; short with rounded 
sides gradually narrowing from base to apex; 
hajry Hlytra costatc, the intervals between 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 ungled. Undersur- 
face evenly and shallowly punaured; hairy. 

Size. Males 9.1 £ 0.05 x 3.4 ± 0.02 mm (105). 
Females 9.2 ± 0.08 x 3.4 ± 0.03 mm (64). 

Distribution. Western Australia. 

Specimens examined. W. Aust.: Types; 2 S & 2 ?. 
Dedari, If. W Bntw/>, WAM, 7M77J/4; 1 J, 
Spencer's Brook. 24.xii.l94A. K. P. McMillan. 
WAM. 71-1764; t S, Bcjoording, I6i,liwy. 
WAM, lirliWX 3 6*& I '?. BefooixSIOfc, 29.xi.J950. 
K. P. McMillan, WAM.. 71-1765 /& V3. 3 1 km E 
of No. 8 pumping, station (KalgOOritfe water mam) 
oil Casuarina eampextrit, 2(J.i. 1958. WAM, 
71-1776; 33 A & 2J 9. Coragina rockhole 66 tan 
S Of BalladOllla on Casuarina hutaeluma, 
5xii,l974. 5. Bar/car: 21 J & 13 9. Ponic? tock- 
hole 70 km S of Balladonia on C. htiegrtiatia. 
S-xii 1974. S. Barker; 14 cf &■ 3 ?, Tummf* rock- 
hole 106 km S of Balladonia un C hut'Kelitwa. 
9X11197*1. S. Barker; i if & 2 <?. 115 km S of 
Balladonia on Cits-uMinn ktwntte, 9.xiLI974. S. 
flutker 



In Astrae-ns (Astraciis) gltitaosus sp. ttov. 
FIGS 10, 221 1 

Holotype: d, 77 km alone main York Rd. w. 

Aasl. on Cusuunna kHetoeJhlNA 5, i. 1968, .V. 
Bar^r SAM, T 2U953 
Allotype: $ 77 km along main York Rd, W. 
Aust. on C hbfg$l(0Jl8, 1 4 196* , V. Horker* 
SAM, J 20954. 

Paraiypes: 2 A & I ?. Dryandra. W. AusT- co 
C. fute&liana I2.JJ973, S. Barter* EA; 3 3 
& 2 $, Dryaiidra. W. Aust. on C. hne#<?f tarts?, 
Hxii.1970, S. Burin, ANIf It a* & 1 ?). 
MNHN fl d & I ?), WAM (I -1). I o. 77 
kin along main York Rd. W- Aim OU C 
ntwltmo, ?jcii 1970. 5. Sarktr, BM; 2 & 
loompup, W. Aust. on Cotywh\a sp., 
23. i. 1072, E A A' T Carnaby, KC; I 8, 77 
km along main York Rd". W. Aiwt. on 
Casuarina sp., J. i. 1968, a\ Barker, SAM; 1 <£. 
luuaning Reserve. W. Aust. on Casuarina 
to., 111968. S. Barker, SAM; I g, Spencer's 
Brook. W. Anst, 26..\ii.l94.S. R, p, McMillan. 
WAM. 71-1763. 
Colour. Shiny. Head and pronotum and elytra 
black with coppery reflections; antennae black 
with coppery and blue reflections. Elytra; each 
elytron with the following yellow markings; a 
large spot at the- bn.se; a broad fascia before 
(he middle, commencing at the shoulder and 
ending near the suture but not touching it, 
concave towards the bn**e; a second fiucia after 
the middle, commencing ,it the margin but 
not reaching the suture; an oval prcapicai spot; 
a single very small spot between the two fascia 
and close to the suture. Undersurface metallic 
coppery; hairs silver. Legs: femur metallic 
coppery, tihia and tursi brown with varying 
amounts of metallic blue on Ihc femoru and 
tibia and on the upper surfcicc of the tarsi. 
Shape and .w.'fiputre, He;u1 closely punctured; 
with a median keel; hairy. Pronotum deeply 
punctured but more closely at Ihc side* than 
in the middle; short with rounded sides 
narrowed towards the apex; H shon imposed 
line projects forwards from the basal crypt, 
hauy. Elytra coslale, the intervuls ilitt each 
with a row of shallow punctures: rounded. 
diverging slightly from the base to a maximum 
width before the middle, thereafter rounding 
off to the marginal \pine; buth sprnes well 
developed: humeral fold modetafely developed 
and angled. Undetsurface: finely and sparsely 
punctured in the middle ot the sternal seg- 
ments, more coarsely and closely punctured at 
the sides: abdominal sternircs finely and closely 
punctured; hairy 

She. Males 9.4 £ 0.13 x 3.6 ± 0.05 mm (2?). 
Females 9.3 i II .3 x 1.9 £ 15 mm (7). 



122 



S BARKER 



Distribution, Western Australia. 

General remarks. This species shows closest 

affinity with A obsenrn:,- 

Saeeitnens examined W. Alls!.: 'types; 1 J 1 , 77 
km along main VorV Rd on Casuaritm sjv, 
5.U968. Si Barker; Z t& 77 km atone main York 
Rcf i>n Cnsuuritm huexefiana, 7.xii.l970, 5- Barker; 
J »;*, Dryandra on C- hta^Hana, 7.xi.lW), t S\ 
Barker; 9 J & \ 9- Dryandra on C". hut^elutna, 
X \M.IS>7(I. .V. Barker. 

17. Astracus (Agtraetts) masters! Max J coy, 
1873; 239. Kerremans, 1S85: 136. 
Masters. 1886: 71. Kcrremans. 1892; 102. 

Astraeus stanoaetli; van dc Holl» 1886: 176, 
IS89: 80, 107-109, pi. 3, fit*. 18b. Kcrrcnmns, 
1 i0? 1 48. Gctfftfi 1 089 : 2S2. Obenherger, 
1930; 366. 

A.tnaru.s spletnh'HS van dc Poll, 188*); gg, 108, 
109, pi. 3, figs 19. 19a. Kctrcmans, IS92: 102: 
1902: 148. Carter, 1*29: 2S2, fjj. 3, tig. 45. 
Obeiiber»er. 1930; 367 (new synonym). 
Astraeas simplex Blackburn, 1802: 211, 212 
kern-man-,. J 902: |4!>. Carter, 1929: 2S2. 
Obenberger, 1930: 3f>h inew synonym"). 

HGS U, 22Q 

r.v/u. Holotype; 2, Gaynclab. AM. X32099 (seco 
by author). 

Colour. Mute. Shiny. Mead ami antennae 
golden-coppery. Pronotum copper coloured in 
the middle at the apex, wiih a dark blue heart- 
shaped area in the middle, outlined by a 
copper coloured margin; at the sides, blue at 
the base arecn at the apex. Elytra black with 
blue or purple reflection*;., each elytron vvitb 
the following yellow markings; a large spot at 
Lhe base; a broad fascia running from the 
shoulder covering the humeral rold and run- 
ning Hansvers^Iy towards the suture but not 
teaching a. after the middle n broad fascia 
from the margin running towards the suture 
hvi not reaching it; a pt'eapical spot — in some 
specimens the preapicM spot \$ absent and 
usually in these the first fascia is broken in the 
inrddle forming two spots, Undersurface blue; 
hairs silver. Legs: blue: the eud of the tibia and 
first tarsal segment leslaceous. 

female. Shiny. Head blue-gicen at the apex. 
dark at the base and aides with purple tcIIcc- 
lions; antennae black With blue or purple re- 
flections. Pronotum dark blue in the middle. 
metallic blue at ibe sides, Elytra black with 
purple reflections, each with the following 
yellow markings: a large spot at the base; a 
lateral fascia commencing M the margin and 
covering the humeral told, running towards lhe 
suture but Interrupted to form a spot near the 



suture: a small spot neat the base, ai the 
shoulder, may or may not be continuous with 
the base of lhe fascia: after the middle a tavcia 
running transversely from the margin towards 
the suture but not touching iu a preapical 
spot. Undersurface blue with metallic, gleams; 
hairs silver. Legs metallic blue; lips of tibia 
and first tarsal segment testaceous. 

Shave 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 excavated line 
from base to ape*; straight sided from the 
base to the middle then rounded and narrowed 
to the apc.\; hairy. Elytra costate, the inter- 
vals flat but slighlly wrinkled each with a 
row of punctures; more or less parallel-sided 
to before the middle, rounded at the middle 
then tapering to the marginal spine; both spines 
well developed; humeral fold well developed 
and acutely angled. Undersurface with shallow 
uniform punctures except in the middle anil 
on the outer edges of the abdominal stcuijtes. 
both areas being glabrous; hairy. 

Steej Males 7.9 * 0.16 x 2.7 * 0.22 mm (17). 
Females 8.4 - 0.26 x 3.2 ± 0,10 mm ( 10). 

Dtxtnbuiiw. Queensland and northern New 
South Wales. 

General temaeka. This Is the only species of 
Astraeus that has been greatly confused (see 
van tie Poll 1889. p. 19). The basic reason for 
this h that the females of A. nutstersi and A. 
Mimouetli are very similar in pattern although 
the males of (hesc species are quite distinct. 
Unih .-species were described from female types 
The lack of hold collecting experience was the 
most likely reason for van dc Poll redesc.ribing 
the male of A. mast erst as A. spteiittens, as he 
would noL have had access to an associated 
scries of males and females ol lhe two species. 

Syeehm-ris examined. N.SAV-' Type; I 6 paratvpe. 
Ash 1.. ANtC: I A & 1 S paratync, CiaynriAh. 
ANTO. Qld; Type; £ A. splendent van de l J oli, 
Kockhumpton, 3 ?, Da [by, Mrs, F. H. Jlokler. 
SAM; I J, Sloiuhorpo. A Cemmetl; 14 dT ft 5 9, 
Edongalba on CwtfitUia rqnisetifolia. 24.x. 1971. 
F. E. Adam*. Type $, A. aimulex Blackburn KM 
(locality of collection uncertain). 

18. Astraeus (Astraeus) samoueHi Saunders. 
1868: 10. pi. 1. fig. 12, 14S7I : 43. 
Masters. 1 871: 124. KcTremans. IRK5: 
136. van de Poll, 1886; 176. 178, 1889: 
«6, 107, pi, 3. fi^s i8 t 18a. WacVburn. 
IN91: 496. Kerremans, 1S02. i02. van dc 



REVISION OF BUPRCST1D GENUS ASJXAtUSS l.APORTF. A GORY 



123 



Pol), 1892; 67. Blackburn, IS92; 2J2, 
Kerremans, i902. US. Cariet, 1929; 282. 

Axtrueas samouelki: Cernmirigcr & tie Harold, 

IH69: 1380. Masters, 1**6; 7J. Blackburn. 

IK90: 1256. Obcnberger 1930: 366. 

Astraetts sufHOticlli: Ker remans, 1900: 295. 

Astmcus .iplenden,\ var roihrikieflas Oben- 

oer$ei\ 1936: 133. 

FIGS lb, 22R 
Type. Hofotype: ? BM fseen by author). 
Colour. Male. Shiny. Head golden-green, 
golden or coppery; antennae coppery or tur- 
quoise. Pronotum uniformly gtildcn-grcvn or 
golden or enppcry in the middle then golden- 
green blending into turquoise ai the sides. 
Elvira hlack with blue reflections, each with 
(he following yellow markings: a large gpol ut 
the base; a fascia just before (he middle cover- 
ing the humeral told, extending backwards 
then transversely towards the suture but not 
touching it: a spot at ihe shoulder which ma> 
be discrete or continuous with the base of the 
lin>t fascia; a fascia after the middle, commenc- 
ing at the margin and running towards the 
suture but not touching it; variably, a small 
elongate spot between the two fascia closer to 
the first and near ihe sulure hul not touching 
it; a preapical spot. Undcrsurfaec bluc-gteen; 
hairs silver. Legs blue-green; tips of the tibia 
and most of the first tarsal segment testaceous. 
Female: Shiny- Head gieen ;il the ^pcx with 
golden reflections, dark blue with purple reflec- 
tions in the middle of the base, turquoise at 
the sides; antennae ^reen. Pionoium dark blue 
with purple reflections in the middle* turquoise 
iii the sides. Elytra black with blue reflections, 
with markings as in the male except that the 
first fascia is broken in The middle to form a 
marginal fascia anil a spot near the suture. 
Underxurface turquoise: hairs silver, ljcgs tur- 
quoise: tips of the tibia and mosi of the basal 
part of the first tarsal segment testaceous. 
Shapt- artel sculpture. Head shallowly. closely 
but uniformly punctured; with a median keel: 
hairy. Pronotum shallowly closely hut uni- 
formly punctured; with a median longitudinal 
excavated line; uniformly rounded and 
nanowed From base to apex; vciy hairy. Elytra 
oostatc. the intervals heiween flat; slightly 
lapered from the base to tile middle, then 
rounded and tapered to ihe marginal spine; 
both spines well developed. Undersurfiiee 
shallowly punctured, more closely at the sides. 
less so in the middle; midsiirface glabrous; 
hairy. 

Size, Males 8.6 ± 0.07 x 3 ± 0.03 mm (52) 



Females 9.2 =: 0.14 x 3.3 ± 0.06 mm (40) 
G+nrrai remarks. The type of A, .splcadens var. 
emhtikieUus Obenbergei is a V specimen of A 
samouetli. This species shows closest ailinitv 
With A. masters!. 

Distribution. New South "Wales. 

Specimens examittrt/. N.$,W., Type: 2 6* & 3 9, 
Bredbu on Ctisttarina striata, 8.1.1963, S. Barker; 
12 o* & 20 % Majors Creek on CvSUttriuti fit- 
Totals 2fr.i,1963, 6'. Barker; 8 d 1 & 2 2, Khanyunis 
Station. Mimima Range on C. Uttatalls, 9,ii.19fiV 
S. Darker, 4 d & 2 ?. Gipttiins Hat to Braidwood 
Rd. on C UitoraHs. 9.ii.l96.V S. Barker; Type 9 
A. splendent Vac, enthr<\teltti> Obenberger, 
Comoro, NMP, 219S9 A.C.T.; V, Mt A indie on 
C strhiu, 26.xii.l962, S. Barkrr; 7 & & 12 ?, Tng- 
jeeranong on C ihfota, I xii 1962. S. Barker; 3 rf 
& 8 *?, Tufigetauong on C. xtricw, 2.xii 1962, ,S\ 
Barker: 4 A St 2 5, Tugsernnotte on C. f/ri*tJff, 
l.i.Wtf. V tiarker. 

19 \ st rums (Astraciis) diluUpe* van de Poll, 
ISS9: 86, ir>5. 106. pi, 3. fig, 17. 17a. 
Illackburn. 1892; 2l2. Kerremans, 18922 
101: J902: 148. Carter. 1929- 282. Obeu- 
berger. 1930: 365. 
Asrraeits samunelli var. ditmtpes vun de PnlL 
1886: 180. 

Astrarat MrumH OberuSerger, 1028: 205; 'Carter. 
1929: 302. Ofccnbcrgcr. 1930: 367 (new syno- 
nym). 

FIG. 22S 
J>0« Hofotyps, MNHN Uiot seen by author). 
Colour. Mate, Shiny. Head green at the apc.\ 
and golden green w\ ihe base or jroldcn-green 
at the apex and coppery at ihe base; antennae 
black with blue reflections. Pronotum divided 
in u transverse direction by an M-shmped line. 
the small apical part being preen or golden- 
.green, the larger basal part bein? dark blue tit 
black and the line of demarcation, purple or 
coppery. Elytra black with hluc reflections 
each wiih the following yellow markings: in 
the middle but not touching ihe sulure there 
is; a large basal spot; a spot before the middle; 
a preapical spot. On ihe margin- a large spot 
covenn^ ihe humeral fold; after the middle a 
transverse fascia not touchine the suture. 
Undersurface blue or green; hairs silver. Legs: 
first and second femora hrown on the outer 
margins, third femur totally or partially solid 
colour similar io the rest of the iindersurfHce; 
lemamder of leg and tarsi testaceous. 
Female. Head green at the apex, purple in 
the middle, dark blue at the base with purple 
reflections: antennae black with blue rcflec- 
lions Pronotum black in ihe centre with 
purple and blue reflections, bine at the sides. 
Elytra as in the male. Undersurtace dark hlue; 



124 



S. BARKfR 



hairs silver. Logs: outer margin of femora 
brown, tibia and tarsi testaceous. 

Sfta/H- and sculpture. Head very densely and 
evenly covered with deep punctures: with a 
thin medial) keel; covered with long hair Pro- 
noium evenly punctured, the puncture* at the 
sales larger than those in the middle; slightly 
rounded bui narrowed from base to apf.v, 
median excavated line from base to apex; 
hairy. Ulytra cost ate* the intervals Hut with a 
row ut deep punctures and faintly transversely 
strigose; narrowed from base until after the 
middle then rounded and again narrowed to 
the marginal spine; both spines moderately 
develuped; humeral told well developed and 
angled. Undcrstirfaec evenly and shallowly 
punctured; wiih long hairs. 
Skt. Males &&--£ 0s2i K~3J ' 0.0R mm (ll). 
Fenuites 9.4 ± 0.13 v 3.4 s= ti.til mm (7), 
Diunhtiwon. New* South Wales and Queens- 
land. 

Ginttal remarks. The type of A. stnmdi Obcn- 
hergcr is a small V specimen uf A. dtitu'tpex. 
The externa! morphology of this species is like 
that of A. Muitet'si and A. sanwuctll but male 
genitalia (Fig. 22S) is different. 1 do not group 
it with the other two species. 

Specimens e\<wiitie<1. NSW- 9 J & 5 2. Major's 
Creek on CasuetHtta littoralis, 26.M963, S. Marker. 
QUi- 2 rf 4fe 2 ° f Pakinu on CoAUwirtu up., 
I.TU970. E. £. Adams; Type 2 A. tfromti Obcn- 
bcrgcr. NMP. 21991. 

20. A*ilracu& (Aslraeus) iidamsi sp nov 
FIGS II. 221 

Type, 

kolotype; rj\ Hdungalba, QW on Cauturtna 
Vifuisetifoiui, 30.xi. 1 974. t". & S. A dams. 
SAM, I. 201*44. 
Allotype; V. tidutt^alba, yjd On C. equisciijolia. 

15.xiUP73, fi, H, Adorns, SAM, I 209V). 
l K aratypes: 3 £ & 3 £, HduueaJba, QUI on C. 
rtfinsrtifolia, l5.Tn.J97*, E. iC S. Adams. EA 
(1 <3 & 1 S), ANIC (T 6' * 1 2), BM (1 2 & 
I c?); I d & i ?. rdtmgulba, QM un C. 
etfaiseltfoliu, ?.?.Ni.l!>74 & S.xii 1974. £ <V ,\\ 
rfdom; MNHN: 1 c? & -J f i Edunsuilbu. QUI 
on (,'. eqaist-njalm, I4.xii.l974. E. & S. 
Aihunz, SAM. 
Colour. Shiny. Head black with purple Tcflcc- 
linns or coppery-purple with metallic reflec- 
tions. Pronotum black with purple reflections 
or brorve in the middle, coppery purple a1 Lhe 
sides with metallic reflections. Elytra black. 
with blue k\nd purple reflections, each elytron 
iVUtt the following yellow marking: a basal 
vpol; -i spot covering the humeral fold bui no* 
reaching the shoulder, a large spot in the 



middle nor touching the suture and just behind 
the second spot these last two in the form of 
8 broken fascia; J4JJt behind the break, near 
the margin but not 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 lhe margin but not 
reaching the suture: a small preapical spot. 
Undcrsurfncc coppery-purple; legs pale brown,, 
the basal ends ol the tibia darker: baits silver. 

Shape and sculpture, Head closely and evenly 
punctured; nu median keel, lightly haired. Pro- 
notum closely and evenly punctured except lor 
a short glabrous line in the middle ai the ape.v. 
ut the sides gradually rounded and narrowed 
from base to apex. Elytra costate. each inter- 
val with a row of deep punctures and faintly 
transversely wrinkled: parallel-sided lu jusi 
before the middle then rounded and tapered to 
the sharp marginal spine, sutural spine well 
developed: humeral fold well developed and 
angled. Undersurfaee closely nml evenly pune 
lured, the punctures shallower on the prostcr- 
nal process than elsewhere; lightly haired. 
Size. Miiles 7.1 ± 0.14 X 2.9 ± 0.05 mm (6). 
Females 7.7 ± 0.10 x 3.2 J* 04 (9). 
Distribution. Queensland. 

General remarks. Male genitalia (Fig. 22T) of 

A. adanlsi is similar to that of A. dihitipes bin 

external morphology is different. I do not 

group these two species. Named after Mr E. L- 

Adams. 

Specimen* exermwed Types only. 

21. Astracus (Asirneiis) intricatuv Carter 1925: 

229, fig. I; 192'J: 282. Obcnbcrgcr. 1930: 

3GG. 

FIGS 1c, 12, 21 A 

lypr, Ilolotype: $, Montiro. MM (seen hy 
author). 

Colotdr. Shiny. Head metallic green ai the apex 

and sides, purple nl lhe base: antennae black 
with coppery -purple reflections. Pronotum 
metallic purple in the middle, deep coppery- 
purple at the sides. Elytra black with purple 
rellections > each elytron with lhe toMuwin^ 
yellow markings: a basal spot, a Kj>Ot at the 
middle pnd a <;pot after lhe middle, all elongate 
and decreasing in size from in front back- 
wards, each with the hasal end diver^ini^ out- 
wards and all in line to give the appearance 
of .a single, broken vitta with the basal end 
diverging outwards and the apical end con- 
verging to lhe sulure, but nol touching it; a 
fascia commencing at the shoulder, covering 



REVISION OF BUPRESTID GENUS ASTRAEUS LAPORTE & GORY 



125 




«* £ 







O 

S 



<* 




I2f> 



S. BARKER 



the humoral fold and running along the ouier 
margin to the middle then extending obliquely 
towards the suture, reaching slightly below but 
not touching (he middle of the three median 
spots; after the middle a fascia commencing 
ai the outei margin ;tnd running transversely 
towards the suture but not touching it; an elon- 
gate preaptciil spot, Close to the suture but not 
touching it. Undersurface deep coppery-purple; 
halts silver. Legs deep coppery-purple, ends 
of the tibia, first and second tarsal segment* 
biown, third ;ind fourth tarsal segments d;*rk 
hrown. 

Shape and sculpture. Mead closely punctured; 
with a median keel; hairy. Pronotum more 
closely puneiured fit the sides than in the 
middle, with a median longitudinal impressed 
line visible nt the base and apex only; parallel- 
stded at the bu*c, gently rounded until after 
the middle, strongly rounded and narrowed to 
the apex, projecting forward lightly in the 
middle uf the apical edge; hairy- Elytra eoslate. 
the intervals between flat, each with a row 
ol shallow punctures; parallel-sided from the 
base, rounded off well after the middle and 
narrowed strongly to Ihe murginal spine; both 
spines well developed, humeral fold moder- 
ately developed and nngled. Undersurface; 
sternal segments finely and sparsely punctate 
in the middle, deeper and larger punctures at 
the sides: abdominal Ntciniies finely and closely 
punctured; hairy. 

Ske. Males 9.6 ± 0.10 x 3.5 ± ffcgjl mm £50*. 
Females 9.8 £ 0.15 x 3.6 ±0.07 mm (35). 
Distribution. New South Wales. 

General remarks'. The species was described hy 
Carter from a unique specimen In the MacLeay 
Museum* Only two of the S5 specimen* I 
have collected conform to the colour pattern 
of the type. I he pattern on the elylr;. is 
variable, ranging from the basic pattern 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 extreme*, in- 
termediary patterns occur, including the intri- 
cate pattern of Carter's type. There is no dif- 
ference between the sexes in si/.e or colour 
pattern. All of the specimens I collected were 
taken on low Casiwrina nana heath in the 
Minuma and Kybcao Ranges. Monaro District, 
N.S.W. I do not group A, intricatus with any 
other species. 

Specimens examined. N.S.W.: Type; 5 <$ & Ml ?, 
Ornnmeir on Cnsuarti^a inula, 9.xii 1962. S. Bar- 



ker; 23 S & 9 V. Orunmeir on C nana, 26. i. 1^63, 
S. Barker; 3 eft lf> km S of Countceativ SUiion 
on C nana, 3.H.1963, £ Barker; 20 rf'A 15 ?. 
Klutnyunis Station on C, nana 9ii.l963. S. Bar 

ker. 



12, Astraoiis (As^raeos) crassus van tie Poll. 
1889: 85, 95-97. p|. 2, %. 9, 9«. Kci re- 
mans. 1892; 101; 1SI021 148. Carter, 
1929: 282. Obetiberger, 1930: 365v 
AstwvuA fUnopictus van de Poll* 3S86: ISO. 



PE 



3B 



Type. Holotype, MNHN (not seen by author 1. 
Colour. Shiny. Rend and pronotum black with 
purple or blue and purple reflections; antennae 
black with blue and purple reflections. Elytra 
black with blue and purple reflections each 
wilh the following yellow markings: the basic 
pattern consists of irregular spots in two rows; 
lour spots in the middle near the suture, but 
not touching it, one at the bu%c, one before the 
middle, one after the middle and one in the 
preapical area, three spots along the margin, 
one ft] the shoulder and coveting ihe humeral 
fold, one ot the middle and one after the 
middle; there may also be a few irregular spnts 
and in two specimens there are spots at the 
base of the sutur.it spine. Undersurface and 
lee.s black with blue and purple reflection*; 
hairs silver. 

Shape and sculpture. Head with a median keel; 
evenly punctured; hairy. Pronotum with small 
and sparse punctures in the middle leaving 
glabrous areas, larger and more dense at the 
sides with a short impressed line projecting 
forwards from the basal crypt; gently rounded 
and narrowed at the aides from the base |o 
two-thirds way to ihe apex, then tapered to 
the apex with the apical edge projecting 
slightly in the middle. Elytra punctate-striatc 
with the intervals flat towards the base and 
slightly concave towards ihe apex, each with 
a row of shallow punctures; parallel-sided until 
after the middle, then gently rounded and 
narrowed to the small marginal spine; suttiTai 
spine well developed; humeral fold poorlv 
developed but slightly angled. Undersurface 
shallowly punctured, more closely at the sides 
than in the middle; hairy. 

Size. Mates 14.4 rt: 0.25 x 5.3 ' 0.06 mm (5) 
FcmaJcs J6.2 ± 0.40 x 6.3 * 0.15 mm (7). 

Distribution, Queensland and New South 

Wales. 

Gcnentf remarks. This species cannot be 
grouped with an} other 



REVISION OF BUl'KESTID GENUS ASTR4EUS t.APORTC & GORV 



127 



Spotirnvns exmwrtcd. Qid: I xi-, £i SuUoh. NSW 
I <jf & J 5. Svdnev. SAM: I ft Griffith SAM; I 
f„ lenolun Caves. /. C Wilbtml SAM; I % 
I. G. Tvpper, SAM; J $. Sydney, IV. d'u Boutaw 
SAM; i 9, IV, ,/>/ Houlay, SAM; 1 ?, SAM; 2 0', 
Majors Creek, on Camariftu titloralis. 26.1.1903, 5. 
Barker. I d. Khanyunis Station, tvlinumn on C. 
tittorotis r 26.1.1963. 5. Barker. 

2^. Astraetrs (Astracus) mnjor Blackburn, 
1S90; 1257, 125S; 1891:496. Kerremans, 
1892: (02; t902: 149. 
A-itiwti* ntworthts var. major- Carter, 1929; 
2S2. Obenbcrgci. J930; 366. 

FIGS I7,23C 
Type. Holoiype, HM inor s:en by author). 
Colour. Shiny. Head coppery with metallic 
reflections; .antennae coppery-green or blue. 
Prouotum dark blue with purple reflections. 
tlytr.i dark blue with blue and purple reflec- 
tions, each elytron with the following yellow 
markings a Fascia running from the shoulder 
iransversely across the base towards the \uture 
but not touching it; before the middle a thick 
fascia funning frv>m the margin transversely 
to the suture; after the middle there in a fascia 
commencing at the margin, running towards 
the suture but uot reaching it. There is an oval 
red spot in Ihe region of thy marginal spine. 
The outer margins of the fascia and humeral 
Told arc red. Undorsurface dark blue with 
metallic purple reflections; hairs yellow. Legs 
rcd-hrown; tarsi blue. 

Shape ami xculpture He^d with sparse shallow 
punctures at the base, wrinkled towards the 
apex; with a thin median keel: hairy. Prnno- 
tum with .small shallow punctures in the 
middle, larger deeper ones at the sides with an 
irregular glabrous longitudinal area in the 
middle; there is a short median longitudinal 
impressed line piojecting forwards from the 
h.tsal crypt; short with the sides gradually 
rounded from base to apex. Elytra punctatc- 
striatc from the base for most of the length, the 
intervals between the striae convex, costate at 
the apex only; parallel-sided unhl just after 
1 he middle then giadually rounded to the n'lar- 
g*na! spine; both spines well developed: 
humeral fold poorly developed and slightly 
angled. Undersurface densely punctured at the 
.\idev sparsely ami shallowly puuciuied in the 
middle; hairy. 

Sizr. Males 1 5.."? ± 0.70 x 5.9 ± 0-22 mm «4)- 
Females 15.5 ± 0.98 x 6.3 ± 0.91 mm 1 2). 
Distribution Western Australia. South Aus- 
tralia and Victoria. 

Gewntl r**mitks. I have no evidence of over- 
lap in the distribution of this species, which 



is found in low rainfall areas of three states, 
and Axtraeus wirarchis, which is presumably 
found in higher rainfall areas in Victoria. 
Because of* differences in distribution and 
appearance I have treated the two as distinct 
species. 

Spvcintcttv CMittiiWii, M-*. Ann.: I o*. WAM 73 
62> I cm hitm, SAM; I <?, Wialfci cm Acaoia '*>ol- 
Rurtlicfuh. JS.ix.1957. £ Barker. WAM. 73-60*. I 
V, Cnllvim, fi.ix.l9tfl. /?. r MMUWi\ WAM, 
73-61. Sf. A urt.: I % Monartu. /. G. O. Ifpfivn 
SAM. I'it :.: 1 B, Sea Lake, Gomiic, NM. 

24 Aslraciis (AsUaeus) navarchis I Phonicon) 
Corto$rtatha rtavunhix Thomson, 1856: 
115, W6> pi. 6, fig. 2. Masters, (886; 79 t 
Astttieus navi/nhif: Saunders, 1S7J; 4J. 
van de Poll. 1886: 176. Masters, 1886; 
71. van de Poll, 1S89: 84, 91. 92, pi. 2. 
fig 5, 5a. Blackburn. 1890: 1257. Kerre- 
mans, 1892; 102- Blackburn, 1892. 211. 
Kerremans, 1902; US. Carter. 1929; 
282. Obenbcrger, 1930: 366. 

FTG. 23D 

Type. Holotype. MNHN l not seen by author). 
Colour. Shiny- Head coppery at the apex, dark 
hluc and purple at the base; antennae dark 
brown, segments one and iwo with golden- 
green reflections, the rest wilh blue and purple 
icflections. Pronotum coppery al the apex and 
sides, dark blue and violet at the base, Elytra 
black with violet reflections: each elytron with 
two yellow fascia, the first commencing at the 
margin and covering the humeral fold, slightly 
concave towards the base and touching the 
suture, the second after the middle, commenc- 
ing, at the margin and running transversely 
towards the suture but not touching it. Under- 
surface: prothoracic steiniles metallic golden- 
green, abdomen metallic violet; hairs yellow 
Legs red-brown with violet reflections. 

Shape and sculpture, Head closely punctured 
at the base, grooved and wrinkled at the apex; 
slightly excavated between the eyes towards 
the base, btit wilh a thin glabrous median 
keel commenciny anterior to this; hairy, Pro- 
notum with small punctures in Ihe middle, bnl 
mainly lacking in the centre, forming an in- 
definite median longitudinal glubrous line, the 
punctures larger and dccpci at the sides; a 
median longitudinal impressed line projects 
forwards from the basal crypt to ihe middle, 
short, rounded at the sides and narrowed from 
base to apex. Elytra punctaic-stnate. the in- 
tervals convex at the base, flat at the apex, 
those in the centre with a shallow row of punc- 
tures, those at the sides with deep punctures; 



m 



S. BARKtK 



parallel-sided until after the middle, then 
rounded and narrowed to i.hc rnargm.il spine, 
boQj spines well developed; humeral fold 
pooily developed and slightly angled. Under- 
surface: anteriorly the- punctures arc small in 
the middle and larger at I he sides, on the 
abdomen uniformly r-matl; hairy. 
Site. Males 14.4 ±0.41 x SJ = 0.13 mm (7). 
Females 16.2 '- 0-34 x (g & - 0.08 mm (5). 
Distribution. Victoria . 

General remarks. This species shows close 
•ifliniiy wilh A. major 

Specimens rvambh'd Vic: '2 rf, SAM; 3 ?, Jan 
lUC^ NM; S df &4S. NM. 

25. A^tiaeiis (Astrueus) fralerculus van de Poll, 
1889: 84, 92. 93, pi. 2, %s 6. 6a. Konc- 
mans, 1892: 102: 1902: 149. Carter. 
1929: 282. Obenhcrger. 1 930. 365. 

FIG. 23E 

type. Rolotyne MNHN (HOC seen by author). 
Colour. Shiny. Head antennae and prnnoi.um 
hiue-black wilh blue reflections. Elytra blue- 
Mack, each elytron wi(h two yellow fascia; the 
first commencing: at the shoulder and cover- 
ing the humeral fold, then running trans- 
versely towards the suture but not touching it. 
concave lowards ihe base; the second alter the 
middle, commencing at the margin and run- 
ning transversely towards the suture bill not 
touching it. Undersurface deep metallic blue; 
hairs silver. Legs deep metallic blue, 

Shape and sculpture. Heat! shallow ly and 
sparsely punctured; with a small median keel; 
hairy. Pronotum sparsely hut evenly punc- 
tured, the punctures at the sides larger and 
deepei than those rn ihe middle; a short 
medinn longitudinal impressed line projects 
lorw.uds from Ihe basul crypt; short the sides 
tapering acutely from base to apex, slightly 
rounded in ihe middle. Elytra cost ate, the in- 
tervals ftai v\ilh irregular shallow punctures; 
parallel-sided until after the middle, then 
rounded and nai rowed to I he marginal spine; 
httih spines well developed; humeral fold 
poorly developed and slightly angled. Under- 
surface with small shallow punctures in the 
middle, larj/er and deeper punctures ai the 
sides; hairy. 

Sir.c. Males 9.3 t 0.30 x 4.2 ± 0,10 mm (7). 
Females 8.8 = 0.30 x 4.0 ± 0.1 1 mm (3). 
Disirihmion. Western Australia. 
General remarks. I he external morphology uf 
this species is very similar to that of A. major 



and A. ttavarchis, however male genitalia is 
different (Fig. 23E). Therefore I do not group 
the three species together but place A. jraler- 
chIus next to A. major and A. navarcbis, 

Spcihthtt\ rwntineti, W. Ausl.; 3 rf & I ?. Borden 
on ffakea trif areata, November 1939. H. W. 
Hrown, VV A M, 73-378/38 1 : I ?, Bnshmead, 
18.xi.1939, /?. P. McMillan, W'AM 73-383. 2 - 
& I 2. Hopetoim, 18x1946, Mrs. Morris. WAM 
$6-191071912-. 1 cf, Wcmblv on Davie.Mu divori- 
cuta, 5.ix.l970. S, Bather; 1 & 4 Borden, WAM. 
73-382, 

26 Astnmi* (Astraeus) ptotttoracicus van de 
Poll. 1889: R5, 98, 99, pi. 3. fig. 1 1. 11a. 
Kcrrcmuns, 1892: 102: JV02; 149. Carter. 
1929: 282. Obenbergcr, 1930; 366. 

FIG. 23 F 

Type. Holoiypc. MNHN (not seen by author). 
Colour. Shiny. Head, antennae and pronotum 
bronze, hlytra dark brown with bronze rclloc- 
tiom. each wilh Ibc following yellow mark- 
in cs: along the margin there is a vitta. com- 
mencing at ihe shoulder and ending in the 
preapical area which may be broken into 
several elongate spots; in the middle buL not 
touching the Suture there is a vftta commenc- 
ing on the anterior edge and ending in the 
preapical area, this is usually broken into * 
basal spot, a spot before the middle and an 
elongate spot after the middle. Undersurface 
bronze; hairs silver. Ixgs red -brown. 
Shape and sculpture Head Closely and evenly 
punctured; no median keel; hairy. Pronotum 
with puncture* wrinkled at the front and sides, 
projecting conically in the middle where the 
puncture coalesce and become siriynve; 
gently rounded at the sides from the base to 
the middle, then tapered and narrowed to the 
apex, the apical edge brood ly convex in the 
middle; Sairy, Hlytra costate, the intervals flat 
with a deep row of punctures; parallel-si tied 
until alter the middle then very gently rounded 
to the marginal spine; both spines poorly 
developed, blunt and close together; humeral 
fold poorly developed and slightly angled. 
Undersurface: punctures close* at ihe sid«s 
than in the middle: legs and whole under- 
surface densely covered with hair. 
Size. Males 10.4 l 0-33 x 3,7 ± 0.13 mm \%\. 
Females 10.5 ± 0.25 x 3.9 *■ 0.17 mm (6). 
Distribution. Western Australia and Queens- 
land 

(tenerr.il remarks, van dc Poll ( 1 889) listed the 
two specimens he had as coming from 
Clarence River and Champion Bay, and Cjif- 



REVISION OF BUPRF.STID GENUS ASTRAEVS LAPORTE & GORY 



I2!> 



ter (1929) gave the range as Clarence River. 
N.S.W. It is now known that in Western Aus- 
tralia the specie* is associated with Banksia 
prttittoses which occurs on yellow sandplain 
in a wide area between »Shark Bay and Rsper- 
artce. van dc 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. QUI: 1 % Myall Parte. Glen- 
nioruan, Nov. 1962, Dutofhv Gtntlatt. JH. W, 
Ait u,; 2 & & J 9, TarnmiTi." 2l.xi.3itt9, A. M. 
Douglas. WAM. 39-2W*^Wj I 4 & 3 ?. Goomal- 
ling, 14J.I9S0, R. P. McMillan. WAM. 73-371/ 
374; I ?, 1^ km N of Northampton, 1.7.1972, 
K. T. Richards, WADA. 

27. Asftaeus (Astraeus) elongatus van dc Poll. 
1*86- 177; 1889: 85, 101, 102. p! 3, % 
14, i4a, Kerremans, 1892: 101: 1902: 
14H. Carter 1929: 2#2. Ohenberger, 
1930: 365. 

FIG. 23G 

Typv- Holotypc, MNHN (aol seen by author). 
Colour Shiny. Male. Head and pronotum 
green: antennae blue-green Klvtra black with 
purple-blue reflections wilh the following 
vellow markings; a single spot ai the base; a 
spot at the shoulder covering the humeral fold; 
a spot on the outer margin at the middle; a 
spot between the previously described one and 
the apex; near the suture but not touching it 
there i* a large spot bctorc the middle, a 
smaller one after the middle and a smaller pre- 
upical spot, Undersurlace and legs green, hairs 
silver. 

Faiiale. Head and pronotum golden-green with 
golden reflections; antennae blue-green. Elytra 
its in the male. Undersurface and legs gotdeu- 
jjreen: hairs stlvef- 

Shapc and sculpture. Head deeply and evenly 
punctured: no median keel: hairy. Pronotum 
with punctures closer at the sides than in the 
middle: with a short median longitudinal im- 
pressed line projecting Forwards from the basal 
Crypt? laterally dilated and with the apical edge 
projecting in the nuddJe; hairy. Elytra punc- 
late-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: 
hoth spines well developed; humeral fold 
poorly developed, slightly angled. Undersur- 
face closely punctured; hairy. 
Size. Males 10,5 =t 0.4 x 3,7 * 0.13 mm <$* 



Females li.8 ± 0.28 x 4.2 ^ 0.J I mm H2>. 
Bistrihwion. Western Australia. 
Genet'ril remark?. A <innle specimen collected 
at Dryandia has a bright blue head and pro- 
thorax. Overall the body shape is elongate and 
cylindrical. This species has closest affinity 
with A* pro thoracic us. 

Specimens examined. W- Ausi.: 7 ,-( Ac 9 }, 
Quairading on Xanthorrhoea to« 19.x. 1970, 5. 
Barker; 1 & & 2 5, Quairading on Xaniharrho>'a. 
7.xiJ970. S. Barker; 1 2. Drvandra on Ctnuarina 
htifxeildrta, 8.xiU970, .S\ Barker. 

28. fateomt <Astrueus) vi<(»t«is van de Poll, 
1889: 85, 99, 100, pi. 5. % 12, 12a. 
Kerremans. 1892; 102, Carter, 1929: 2S2, 
Obcnbcigcr, 1930: 367. 
Astraetts vitiatus: Kerremans, [902: IA9. 
FIG 23H 
Type. Holutype. MNHN inot seen by auibor'l. 
Colouc Shiny. Head and pronotum black with 
purple reflections: antennae black with blue 
reflections, the apex on the first, second and 
third segments dark brown with purple reflec- 
tions. Klytra black with violet reflections, each 
wiih the following yellow markings 1 : a vhta 
along the margin from the shoulder to the 
preitpical region, broken just niter the middle; 
a vittii in the middle but not touching the 
suture trom the base to near the apex, broken 
near the middle Undersurface metallic purple; 
hairs silver. Legs brown. 
Shape arid sculpture. Head broad and closely 
punctured; no median keek hairy. Pronotum 
with small and shallow punctures in the 
middle, larger and deeper at the sides, 
gradually rounded from base to apex; the 
anterior margin projecting forwards in the 
middle, hairy', Elytra pUncUitc-Mriatc at the 
base, faintly costate at the apex and slightly 
wrinkled, the intervals slightly convex at the 
base and flat at the apex, each with 3 row of 
punctures; parallcl-sidcd until just after the 
middle, then tapered to the verv short marginal 
spine, sutural spine very broad: humeral fold 
poorly developed, slightly angled Undersur- 
face closely and densely covered with fine 
punctures; hairy. 
Size. Male 9.9 x 3.7 mm < t )- 
Distribution. Western Australia. 

Specimens exatniftcd. W. Ausl.: I H*. W&lnllftgi 
10.xiL19M), R~ P. McMitlnn, WA.VT, 71-1761. 

29- Astraeits (Astraeus) flavopjetus LaPoTte A 
Gory. 1837 2. pi. 1. fig 1 Imhnff. 1856. 

46. Lacordaire, 1 857: 43. Geinminget A 
de Harold. IK69; 1380 Masters. 1K7I: 



130 



S- RAKKbR 



124. Saunders, 1871 ; 43. Kcrremans. 
1885: 136. van de Poll, ISS6: 176. 177. 
Musters, 1886; 71. van de Poll, 1889* 
85, 97 98. pf. S, fig. 10, 10a. kerremans, 
1892; 101; 1902" J 48. Carter. 1929: 282. 
Obcnbctger. 1930: liS5. Barker, 1964: 
306> 307. 

FIG 23f 

T\pc. Holoiype. MNNN (nor $ocfl by author). 
Colour. Shiny, Head and pronotum brown with 
variable green and/ or purple relied ions; 
antennae black with blue reflections. Elyua 
brown with variable bronze and violet xctfec- 
lions, with the following yellow markings in 
two rows, one in the middle but not touching 
the suture, the other along the margin; in the 
xiuddle; an elongate basal spot; a transverse 
bar before the middle: an elongate spot aiier 
ihe middle; a long thin preapical mark. Along 
the margin: an elongate spot from the shoulder 
covering (he humeral fold; a spot in the 
middle: a spot alter the middle. The three- 
spots along the margin may coalesce forming a 
single vitta or be separated into two or three 
spnis, Understirfacc and legs bronze; hairs sil- 
ver. 

Shape and sculpture. Head densely punctured; 
no median keel; hairy. Pronotum closely and 
evenly punctured: basal half pnralleUsided, 
thereafter rounded to the apex; apical edge 
projecting forward in the middle; a short im- 
pressed line projects forwards from the basal 
crypt: hairy. Elytra cosiate. the iniervals flat 
but deeply punctured and slightly wrinkled; 
parallel-sided from base until after Ihe middle, 
then narrowed to the small marginal spine: 
sutural spine short; humeral fold moderately 
developed and angled. Under.surface closely 
and evenly punctured: densely h<iiry <& arc ific 
tejis. 

Size. Males 10.9 - 0J9J k T«* L ().J7 mm 
(12). Females 12.0 ± 0.27 x 4.4 ± 0.IU mm 
(15), 

Distribution. Western Australia. 

General tematk*. This species shows closest 
.itTiiiiiy with A. vituitm. 

Specimens examined. W. Auat.: I ?, Porougioups, 
2X ii 1063. h H, Utlwr tinker; I rf & I $ 64 Vm 
along main York Rd, on Jacksnnia sp,, \.\ low*, S. 
Harkrr; I \ c? & 13 9, 64 km along main York Rd, 
un Jacksonia sp., ti.xi. (970. 5. Barker, 

30. Astraeus (Astraeu*) iijueruillvmi sp, new, 
FIGS 13, Til 



Koiotypc: tf, 77 km along main York Rd. W. 
Aiisl- on Ca\uaihm hut'fHlfwia, 21.x. 1970. S. 
fittt&rt, SAM. 1 20955. 
Allotype; =?, 77 km along fgAto "York Kcl, W. 
Aiwl. on C liiivnetwtKi, 5 i. 19f>8, S. Batk,r, 
SAM. t 20956. 
Paraiypes: I <J &, I ?. 77 km along main York 
Rd, W, AuM. on Ctnuarhm sp., I.i.i9f>8, .V. 
Bt,rkt<r. EA; 2 tj & 2 ?, 77 km fllQnfl main 
York Rd, W. A list Oil C. luic^eliana, 
2T-*i 1970. S. Barker, ANfO ( I J & J £)i 
BM ( I if, MNIIN II ?); 1 5, 77 km along 
main York Rd, YV Ausi. on C. huemUana, 
.Vi.1968. 5- Barker, BM; 6 tf & 2 <?, 77 km 
along main York Rd, \V. Aust. on C 
hrwtlaua, 7,xi.l970, & Bnrter\ MNHN (1 
t?), SAM (4 rf & 2 ?>, WAM (1 rf); I tf * 
2 ?■ II km S of Walebing, W, Aust, 
6.x. 197 1, K. T. Rivhntth, WAIM: 1 & 77 
fcm alone main York Rd. W. Aust. on t 
iHn'Mthwu, 5.1.1968, ,V. Barker. WAM. 
Colour. Shiny. Head black with purple reflec- 
tions: antennae black With metallic bine 
reflections, each ol segments 1-4 with dark 
brown apex. Pronotum black with blue-green 
reflections in the middle, purple reflections 
in the front and at the sides. Elytra Hack with 
purple reflections, each with the following 
yellow markings: a large spot at the base; a 
fascia commencing at Ihe shoulder, covering 
the humeral fold and running transversely 
towards the suture but not touching h, con- 
cave towards the base (sometimes broken into 
a marginal and H medial spot): a small spot 
after the middle on the outer margin, an oval 
preapical spot; between the last two spois a 
shon fascia, running From the margin, half 
way to the suture. Undeisurface metallic 
purple and bronze; hairs silver. Legs rcd- 
brown. 

Shape and sculpture. Head closely punctured; 
no medinn keel; hairy. Pronotum deeply and 
closely punctured at the sides, sparsely in ihe 
middle; a median longitudinal impressed line 
protects forwards from the basal crypt to the 
middle; parallel-sided at the base then rounded 
and narrowed to the apex. F.lylra coslnte. tire 
intervals flat eaeh with a shallow row of punc- 
tures; more or less parallel-sided until the 
middle t.hen rounded and tapered to the aviar- 
ginal spine: both spines well developed; 
humeral fold moderately developed and angled. 
Undersurface; thoracic stetnites finely and 
sparsely punctured in the middle more closely 
at the sides; abdominal segments finely and 
closely punctured; hairy. 
Size. Males 10.5 ± 0.10 x 3.9 ± 0.04 mm 
(47). Female* 11.7 = 15 >. 4.3 ± 0.06 mm 



KbVTStON OF ttUPROSTID GENUS ASTRABVS LAPORTE & GORY 



i 1 



Pt-itrihufson, Western Australia. 

General rvmurhx. In same specie* trie marginal 

spot is reduced ox lost, in others (here is an 

additional spot close to the suture and between 

the two fascia. Named lifter Mr R. P 

McMillan. 

Specimens examined. W. AuU.: Types; 5 of & 4 ?, 
77 km alone mam York Rd on (.'osuarina sp., 
5.i.l968, .9. Darker; 11 j. Diyandra oq Cusuarirta 
sp, 21 x. 1970. 5. Barker; 6 cf & 6 9, Dryandra on 
Catuarhta httt^lionci t 1 1 xi .1970, 5 fi4/&4 2 -d 1 * 
2 ?, 6 km E of North Bannister on C fytteuc liana, 
19jaJ970, 5". JfVA<7, 6 c* & 4 9. 77 km along 
main York Rd on C. hucxoliutw, 2J.xi 1^70. S. 
/toAe/. I rf & 3 9* 77 km along main York 
Rd on C. httrgetiorta, 7.*ii. 1970, .$ Barker, $ J & 
1 ?, Dryandra on C huexeliami, 8.xii.t970. 5. /?ur- 
At'r; L rfi Dryandra on C, huepxliam, 12t,l973. 

;M. AUraeus tAstraeus) camabyi sp. nov. 

FIGS Id. 14, 23K 
Types. 

Ilolotype: r?, 16 km NE of Lake Grace, W. 
Aust., on Ca\uurhtu ImegeUoiia, 9.1.1972, 
/:. <K- K. Citrmiby, SAM, 1 20947. 

Allotype; P. f6 km NE of Lake Grace, W. 
Aust.. on C Ititepelhma, 3.1 1971- £. <# A. 
Carnaby. SAM, 1 20948. 

Parutytx*; I # & I ?. 16 km NE of Lake 
Grace, W. Aust. on C hueRetiatia, 9.5.1972. 
£ * A'. Cvrnuby, ANiC; 1 ?. 16 km NE of 
Lake Oiace, W. Ausi. on C. huegeliaiui, 
25.LI973. F\ rf AT. Carnahy, KM: I rf & I ?. 
16 km NE ol Lake Grave. W. Ausi. on C. 
huegcliatta, E, & K. Carnaby, KC; I c? A I 
9> 16 km NE of Lake Grace. W, Ansr. on 
C. hut.getimut, 25.1.1972. £. dt K. Caraubv, 
MNHN (1 «?)♦ WAM fl?); 1 9, 24 km S 
of Lake Kjn&, W Aust. on C. knvxelumtt, 
25J.I973. 5. Knftir* MNHN; 4 <? & I 9< M 
km N of Nccdilup. W. Aust on G huvgt- 
liamt, I4.xii.l972, K .7* Richards, WADA. 

Colour. Shiny. Head, pronotum and elytra 
black with purple reflections; antennae 
black with blue and purple reflections. Each 
elytron with the following yellow markings: an 
elongate spot at the base reaching the anterior 
margin; a fascia commencing at the anterior 
lateral margin, covering the humeral told then 
running upwards towards the suture but not 
touching it. concave forwards; a small fascia 
just after the middle, commencing at the mar- 
gin and at right angles to the suture hut reach- 
ing only half way lo it; midway between the 
two fascia arc two small spots, one on the 
margin and the other near the suture but not 
touching jt: a small prcapical spot. Under- 
surl'aee metallic purple; hairs silver. legs dark, 
upper sides with metallic blue reflections, 
undersides metallic purple. 
Shape and sculpture. Head with medium sized 
punctures on basal half merging into irregular 



longitudinal channels on apical part; uo 
median keel; hairy, Pronotum evenly punc 
cured, with n median glabrous line at the base 
and apex: sides rounded gradually from base 
to ape*; harry. Elytra eo-state, the intervals 
between flat, each with a row of shallow punc- 
tures; parallcl-stded until just after the middle, 
then rounded off to the strong marginal spine; 
well developed sutural spine; humeral [old 
poorly developed but slightly -angled. Under- 
surface evenly and shallowly punctured, hairy. 
Size. Male* 9.X * 0.17 x 3.7 ± 0.06 mm (8). 
Females 10.6 ± 23 x 4.0 -i 0.1 1 mm (7). 
Dixtrshution. Western Australia. 
(leneral rentarta. Named after Mr K. Carnaby. 
Specimens examined, Types only. 

32. Astraens (Astroeus) budenj van dc Poll, 
1889: 84, 93. 94, pi. 2. fig. 7, 7a. Black- 
burn. 1891: 496. Kerremans, 1892. 101. 
van dc Poll, 1892: 67. Blackburn, 1895: 
45. 46. Kcrrcmans, 1902: 148. Carter, 
1929: 282. Obeuberger, 1930: 365. 

Asxmens badeni var. disjunctus Obenbertjcr, 

1928: 204; 1 Q30* 365 

Astraeux merricki Blackburn, 1890 125^1257, 

Kerremans. 18i>2: 102 Blackburn, 1895: 45, 

46. 

FIG. 23L 

fXlflfc Tlolotype: ? MNHN ^seen by author). 
Colour. Shiny. Head, pronotum and elytra 
black with blue reflections; antennae black with 
blue reflections, tips of first and second seg- 
ments brown. Elytra: each elytron with the 
following yellow markings: a spot at the base; 
a fascia commencing at the margin on the 
humeral i'old, running obliquely upwards and 
backwards and then at right angles to the longi- 
tudinal axis of the body, just before the middle 
and not touching the suture (this is frequently 
broken to form two spots vide var. disjmuius 
Obcnberger, hut this has no taxonomic signifi- 
cance); just after the middle there is a small 
fascia commencing at the margin, slightly con- 
cave to the apex and not touching the suture. 
Undctsuifacc and legs dark with metallic 
purple reflections; hairs silver. 
Shape and sculpture. Head evenly punctured; 
no median keeh hairy. Pronotum with shallow 
punctures, larger a! the sides than in the 
middle; sit the sides gradually rounded and 
nan owed fiom base to apex; hairy. Elytra 
costate, the intervals between flat each with a 
row of shallow punctures; parallel-sided until 
after the middle then rounded olT lo the mar- 
ginal spiuc. strong sutural spine; humeral fold 



132 



S. BARKER 




•a 

.5 







LO 



9 



S 

g 

©0 
K 




a 



g 



REVISION OF BUPREST1D GENUS AS'i'KAtVS LAPORTH it GORY 



133 



pooxly developed but alightly angled Under- 
.surface, punctmev sparser In the middle than 
at the aides' hairy. 

$&*. Males S.8 i 0.14 x 3.4 ± 0.07 mm (16). 
Females S.^x0,|6x 3.5 ± 0.07 mm (17). 

Distribution. All of mainland Australia except 
the Northern Tctntorv. 

General rework*, This species was described 
almost simultaneously by van de Poll as A. 
htnfeni and by Blackburn as A meyricki, 
Blackburn (1891) recognised A. titcyrieki a* 
u synonym of A. h*tcleni t but later changed his 
mind and cnlled A. meyr'tcki a good species 
(Blackburn 1S95) Distance between the two 
populations was the main argument used by 
Blackburn (1895) in favour of calling the 
eastern and western representatives two dis- 
tinct species. I have been unable to separate 
specimens collected in Western Australia from 
those collected in South Australia. 

Specimens examined. 6". Aust.; Type; 1 c? & X ?. 
Morgan, A, M. Lea, SAM: 1 ?, Murray Bridge, 
Oct. 191 1, SAM; 8 rf 4 6 ?, on Melaleuca sp., 
Dernu Pass (probably the same as Puttapa Gap). 
21 km S of Copley, 25.x.) 969, ,V WT^fe/ttfi 
SAM; 7 3 & 9 ?, Puttapa dap. Flinders Rangev 
on Melaleuca t/iomerata, 21.X.197I, S. ftarkct, 
W. Ausi.t Patalype of A. tnevrickt Blackburn. 
SAM; 1 ?, T8 km SW of Three Springs on Dry- 
andra cirsioides, $.xi.l968, N. McFxvland, SAM: 
3 S & 3 H, luianda rockholc. 106 km S of Balla- 
donia on CaltitrFx preissii. 9xii.l974, 5. Barker. 
K.S.fy,: 1 2, W. du Boutay, WAM, 73-5-1. QUI 
Type £ A. kaleni var disiunetus ObenbergeT. 
NMP. 21 990. 

VV Astraews fAstraeus) unison i van de Poll, 
1889: 84. 94, 95. pi. 2. fig. 8, Ka; Black- 
burn, J891. 496, Kerremanv 1892: 102, 
van de Poll, 1892: 68. Blackburn. 1894: 
10J; 1895: 46. Kerremans, 1902: 148. 
Carter. 1929: 282. Obenberger, 1930: 
3<56. Carter, 1933: 42, 
A sweat Upper* Blackburn, 1890: 1258, 1*259 

FIG 23M 
T\pt Holotype, MNHN (not seen by author). 
Colour, Shiny, Head, pronotum and elytra 
bronze-green or black with green or purple 
reflections: antennae black with blue or purple 
reflections. Elytra: each elytron 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 3fier 
the middle Tunning transversely towards the 
suture bui not touching it; in the middle near 
the suture bu; not touching it, a spot at the 
b*»sc. n spot before the middle, a spot at the 



middle, a prcaptcal spot Undersurface and Ices 
metallic bronze-green; hairs silver. 
Straps and sculpture. Head evenly punctured, 
stichtly excavated in the tnt-ddte between the 
eyes; no median kech hairy. Pronotum evenly 
punctured: a short longitudinal median 
impressed line projects forwards from the basal 
crypt; at the sides rounded basal ly, then 
tapered and narrowed to the apex; hairy, Elytra 
costatc, the intervals between llat. each with 
a deep row of punctures and faintly trans- 
versely wrinkled: parallel-sided until after the 
middle, (hen rounded and narrowed to the 
marginal spine: both spines well developed: 
humeral fold poorly developed and slightly 
angled. Undersurfacc with small shallow punc- 
tures in the middle, larger and deeper at the 
sides; hairy. 

Size. Male*. 8.1 ± 0.08 x 3.2 ± 0.04 mm (37>. 
Females 8.6 ± 0.15x3.4 ±0.06 mm (12). 
Distribution. Mainland eastern Australia from 
South Australia to Queensland. 
Ctnetui remarks. The yellow pattern Is 
variable; there aie either six spots and a fascia 
on each elytron or the fascia may be broken 
in the middle giving a total of eight spots 

Specimens' examined. S. Aust,: 2 ?, McDonald 
Femes N.P. on CaVlttis prei.wii, Z.xi. 1967, A, Bar* 
krf 1 o* & I ?, 2 km E of Hiii (ley on C. un'tswi, 
15.xi.1969, S, Barker; 7 6* & 7 ?, McDonald 
Ferries N.P. on C. prvissii, 15.xi.19f>9, S* Barker; 
2 -? & 2 9, on road N of Pa era Wina N.P. on C. 
pr'fissii, lO.xii.196V, S. Rnrket: 7 <* & 1 1 5, Tothill 
Ranges near Brady Creek on C. preissii, 
IZjuLT969« S. Barker; I 9. Onknparinpu Gorge 
near Hack fi am on Caltitri'i rhomhofdeus, 
27.Xii.1V6V, 8. Barker; II <? & 5 $, Ml Remoik* 
able N.P. on C. preterit, 30.vii.l969. S Barker: 
2 A & 1 9, Alligator Gorge N.P. on C preissii, 
30.x. 1. 1969. ff. Barker; 5 £ A 2 R Mt Remarkable 
N-P. on C. preissii, 22.X.1971. vS. Barker. I tf, 
McDonald Femes N.P on C preissii, 14.xi.197t, 
.V Barker; 2 c?. 16 km N of M annum on C 
pm\y*l, 2fi.jrf.l97l, S Baker; 6- rf & 7 ?. Salter 
Springs °" C. preissii 24.ix.1972. fc Bother, 4 
J &. 3 ?, Alligator Gorge N.P. on C. preissiU 
8.x. 15*72, S. Barker; 1 <.?, 10 km W of Penneshaw. 
Kangaroo 1. on C. preissii, 24x1.1972. S. Barker: 
1 r? & I ?, near Rocky Point, Kancaroo 1. on C. 
preissii. 21.x, 1974, S. Barker, Qtd: 7 % StaJl- 
thorpc, h, Sutton, QM 

34. Astraeus (A*traem) oberthuri van de Poll, 
18Xy: 85. I00 t 101. pt. 3. fie. 13, 13a. 
Kerremans. 1892: 102. Blackburn, 1592: 
211, Kerremans, 1902: 149. Carter, 1929- 
282. Obenberger, 1930; 36fv 
FIG. 23N 
Type. Ho]ot>pc <J\ MNHN (seen by author), 
Colour. Shiny. Head, antennae and pronotum 



iu 



$ BARKER 



black with purple ami blue neJU'-ciiuns, Elytra 
black with purple reflections, each with the 
following yellow makings: a buval spot; a 
foscih commencing at the shoulder, covering 
the humeral fold ;nul running towards lb? 
surure hut not touching it. slightly concave 
towards the base; a spot at the middle touch- 
ing the margin; a preapical spot, a spot mid- 
way between the previous two which may or 
may not touch the margin: a spot midway 
between the fascia and the preapical spot, 
near the suture but not touching ii, absent in 
.••time specimens. Undersurfacc and legs metal- 
lic purple: hairs silver. 

Shape and sculpture. Head closery anil evenly 
punctured at the base and sides, punctures 
coalescing and wrinkled at the apex, no median 
keel; hairy, Pirmoium closely and evenly punc- 
tured in the middle, punctures coalescing and 
wrinkled at the skfes; A short median longi- 
tudinal impressed line projects forwards from 
the basal crypt; parallel-sided from the base 
until after the middle, then rounded and 
narrowed to the ftpcx, front edge projecting 
slightly in Ihe middle: dorsally convex in 
lateral profile; hairy. Elytra costate, the inter- 
val* between flat, each with a row of punctures 
and faintly transversely strigose: parallel-sided 
to the middle then rounded and tapered to the 
small marginal spine; suturaf spine well 
developed: humeral fold poorly developed and 
slightly anglo-l Undcrsurface with shallow 
punctures closer at the sides than in the 
middle; hairy. 

Size Males 9.7 3: 0.10 x 3.6 ± 0.04 mm (58). 
Females 9.8 £ (1.34 x .Vfi ± 0.1.5 mm ( IK). 

Distribution. Western Australia. 

Specimen* evarnined W AnAf.: Type; \ rf- Van- 
Chcp, 7.U962, t\ H. UtUer Baker; 3 S. Gusnclte 
on Casuarma sp.. 7.1.1967, 5. Barker; \<t g & ) ?, 
X km W of Rcverfy 1 .O from Kroofcton Rd, oa 
Caiuariau sp.. l.i.1%7, S. Barker; I i^. 77 km 
along main Vork Rd, on Casuoritui sp., \ a. [<)(&> 
5. Barker; 8 J & I g, Rod Hill Rd. near Midland 
I unction on Caxuuriua sp., 4.i.l968. .V. Barker; 2. 
($1 77 km along main Vork Kd. na Casuarhta sp.. 
5J.ll>6y, .V. Barker; 6 S & 2 fc 77 km along main 
Vork Rd, on Caxuarina hucvvllaaa t 1 y,\.\kl(\, $ 
Barker, 3 j & 2 ?, 7 km E of North Bannister on 
C. hue»eUiina, 19.jm.I970, S. Barker; 8 J & 3 9, 
t% km b' nf Norrh Bannister on C luw.eUatm 
1 D.m.1 970, .V. Barker; 5 £ & 3 ?, 77 ken along 
main York Rd. on C. huegeltaaa. 1 1. xi. 1970, ( S~ 
WriiUv. \ ^ t 77 km along mam xOci Rd. on 
C. fuwxelianu, 7,\iU970. 5. Barker; 3 S & 2 ?. 
Wnnnnma! oh €■ htteveUatm, 10, xii. 1H7", S. Bar- 
ker; 2 $, 135 km along Albany Highway on (7. 
hiwxeliana, KU.I973, 5. Barker. 



J5 Astmeim <Aa»traeus) watsonl sp, nov. 

FIG. \S 
Types. 

HoIOTvpe; 2* Rorilen, W, Auai.. 16.xii.l957. 
K. P. McMillan A /. A. L. Wdtsoa, WAM, 
71-1761. 

Paratypes; 1 8 ANIC; I & Ciuomalling, W. 
Au$t, SAM; I Si W. Aust, SAM; 1 9, Bor- 
den, W Aunt., Ib.xii. 14)57. K. P. MeXfitlan 
<C V. A L. Watson, MNHN; I % fcorden, W. 
Aust., 16.xii.1957, R, P. McMillan dt /. A. I. 
Watson, WAM, 71-1760. 

Colour. Shiny. Head, pronotnm jind elytra 
black with blue and purple reflections; an) en* 
nae black with blue and purple reflections, the 
base <m6 apex of the first, segment and apes 
of the second segment brown. Klytra* each 
elytron with the following yellow markings; a 
lat'gc basal spot; a broad 1'ascia commcncinc 
at the margin and covering the humeral fold. 
running transversely towards the suture but 
broken in the middle forming a large spot near 
the suture hul not touching it. after the middle, 
a broad fascia concave backwards and not 
touching the miirgin or the suture: a small 
preapical spot; a small spot between the fascia 
near the margin but not touching it. Under- 
surface black with blue and purple reflections; 
hairs silver. Legs brown: top edges of the 
femur and top surfaces of the tarsi dark brown. 
Shapr and scttfpwre. Head nhallowly but 
evenly punctured; with a median keel; covered 
with very long hair. Pronotum with punctures 
evenly dispersed; a median longitudinal 
impressed line projects forwards from ihe 
basal crypt to the middle; sides gently rounded 
and narrowed from base to apex: hairy. Klytra 
costate, ihe intervals between convex at the 
basal half, flat at the apical half, each with a 
row of shallow punctures und slightly trans- 
versely wrinkled, parallcl-sidcd until just 
before the middle, lapered gradually to the 
marginal spine which is moderately developed, 
strongly developed sutural spine: bumera' fold 
moderately developed and angled, UndcrsUr- 
facc evenly and shallowly punctured; hairy. 
Size, Females 12,9 - 0.73 x 4.6 ± 0.30 mm 
(6) 

Gi'nemi remarks Named after Dr J. A. L. 
Watson. 
SptrUneru> ewminett Types only. 

3b. Astraeus (Astraeus) raricri sp, nov. 

FIGS 1 5. 230 

Type* 
Holutype: rf, 3S3 km ahmjj: Payne'* Find Rd* 
W. Aust, on Ca.wiatina otehiana, 17 iX. 1970. 
S. Barker. SAM, 1 201M9. 



REVISION OF BUPRES'UD GENUS AStRAEOS LAPORTO & OORV 



135 



Allotype: & Lake King. W. Aust.. 18.xii.197i)> 
£. <$ A. Cartmhy. SAM. T 20950. 

Patau-pes: 3 J, 383 km along Pavne's hind Rd, 
W Aust , on C fftetitanth n.ix.WO, S. Bar- 
ker, ANIC (1 i), BM (1 ?), MNHN t L rfjj 
3 <J A I 5, Bord-fn, W. Ausl., on Casaarina 
gtmea. 16.x,l93<>. W. I*'. JtanfH. ANIC <! 
?). WAM (3 rf)- WAM, 73-68/70. 2 ?, Luke 
Grace, W. Aust., MNHN < 1 9). WAM I I 9). 
WAM. 73-66; 3 d\ Lake Cruee, // H? 
J&Wff, WAM, 35-735/*, 73-$5; Id 1 *!?, 
Southern Cross. H If. Brow, WAM, 73-67, 
73-63; 2 eT. Tollering Slalion, Pindar on 
CWWbia %p n 22.tx.f95a. S, Barker, WAM, 
73-72/3. 

Colour. Shiny. Head black with purple reflec- 
tions; antennae black with blue-green and 
puiplc reflections. Pronotum black with blue- 
green reflections on top and purple rejections 
at the margins. Elytra black with purple reflec- 
tions with the following yellow markings: a 
spot at the baW a spot originating at the 
shoulder covering the humeral fold; a spot 
above the middle touching the outer margin; a 
spot below the middle, near the outer margin 
but not touching it: three spot* close to the 
suture but not touching it, the first Ihc largest 
above the middle, the next smaller below the 
middle and a longitudinally elongate prcapical 
spot (the first of these sometimes coalesces 
with the spot covering the humeral fold to 
farm a fascia, concave towards the base). 
t;ndersurfacc and legs dark metallic purple; 
hairs silver. 

Shape and .sculpture. Head closely punctured; 
no median keel; hairy. Pronotum evenly punc- 
tured; with a short median longitudinal 
impressed line project ing forwards from the 
basal crypt, running forwards from the 
impressed line is a glabrous line formed by 
lack of perforations, better defined in females 
where it reaches the anterior margin than in 
mattes where it runs only ro the middle, sides 
rounded and narrowed to the strong marginal 
apex; dorsally flattened in lateral profile; hairy 
at the sides but less so in the middle. Elytra 
costalc, the. intervals between flat with a row 
of shallow punctures; sides at first diverging 
slightly outwards from the base then parallel- 
sided to before the middle* then gradually 
rounded and narrowed to the strong marginal 
Spines; well developed sutural spines: humeral 
fold poorly developed but slightly angled. 
Undcrsurfacc finely and sparsely punctured in 
the middle more closely at the sides: densely 
hairy 

Site. Males 1 1.2 ± 0.16 x 4.0 ± 0.0S mm 



(13). Females J 2.0 ± 0.92 X 4.3 ±ft4rtmm 

O). 

Distribution Western Australia. 

General remarks, A. carters show* closest 

atlmity with the following group of species; A. 

inacmiliani t A. curmibyi, A. hadeni, A. jafcs&nu 

A. oberthnri and A. watsoni. Named after the 

late Mr H, J. Carter, 

Specimens examined. Types only. 

37. Astraeus (Astraeus) goerlingi sp. nov, 
FIGS 16, 23P 

Typrn 

Holotype: d\ Mar loo Sin, Wurnrga, W. AuM 
193 Ul 94 1, .-1. GoerHng, ANIC.. 

Allotype; 9. Martoo Sin. Wururau. W. AUM. 
1931-1941. A. Goerlinx, ANIC. 

Paratypcs: 4 J & 3 ?, Marloo Stn, Wurargfl, 
W. Aust., 1931-1941. A. GaeHuw, ANIC 
<1 .-* & 1 S), BM(l(?&l 5), SAM (1 o*). 
WAVf ( I 6* & 1 ?J; 3 ?. 106 Km S of Payne's 
Find, W. Aust., on Casuoritm nrutivatvh, 
I&&197& .V, Barker, SAM; 2 6* & I Z 
Wurarga, W. Aust. on Cusuartna prmcipiana 
IA.ix 1931 H W. Bton-n, 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 shoulder and covering the 
humeral fold, running towards the suture but 
not touching it, concave towards the base; a 
spot at the middle touching the margin; a pre- 
apical spot; a spot midway between the two 
previously mentioned marks, elongate and 
touching the margin; a spot near the suture, 
but not touching it; midway hetween the fascia 
and the pneapical spot Undersurface and legs 
purple, coppery and bronze; hairs silver. 
Shape and sculpture. Head closely and evenly 
punctured; slightly excavated in the middle 
between the bases of the eyes; no median keel; 
hairy, Pronotum with punctures larger at the 
sides than in the middle, median longitudinal 
glabrous line from base to apex formed by the 
absence of punctures; gently rounded at the 
sides from base to apex; hairy. Elytra costate 
at the apex, punctate-striafe at the base, the 
inteivals between flat at the apex and convex 
at the base, each with a row of punctures: 
parallel-sideU to the middle then rounded and 
narrowed to the marginal spine, which is well 
developed: sutural spine sharp but shortened 
by the sutural margin being straight and super- 
ficially appearing to be broken; humeral fold 
pooTly developed and slightly angled. UnJer* 
surface evenly punctured, the punctures 




S. BARKKR 




5 m 



Fig. (7. Asiraeus major Blackburn 



5m m 



Fig. 18. Astwus watsoni sp, nov, 



deeper at the sides than in the middle; hairy. 

In the male the lasi ahdominal slernire has p 

marginal indentation in the centre. 

Size. Males 10.8 £ 0.12 x 3.8 ± 0.05 mm 

(26). Females 11.7 =t 0.19 » 4.2 ±; 0.07 mm 

<22>. 

Pitfribution, Western Australia. 

General remarks. A. fioerlitigl shows features 

in common with the preceding group of species 

(Fig. 23P) and also with xhc following group 

of two species, and because of this I place it hy 

itself between the two groups. Named after the 

late Mr A. Goerling. 

Sp/u rlmetis cxaminrd, W. Aust.: Types; 20 : i & 14 
i\ Marloo Stn, Wurarga. 1951-1941. A. UoartiriK. 
ANIC. 

IK, Astraeus (Astraeus) eyaneus Kerremans, 

1900: 295; 1902: 148. Carter. 1929: 282. 

Obenberger, 1930: 365. 

FIG. 19 

type. HoIOlype: & Standing, N.vS.VV.. DM liseen 
by author). 



Colour. Shiny, Head and pronotum bluc-grccn: 
antennae black with blue reflections. Elytra 
black with blue-green reflections, each with the 
following yellow markings; a large elongate 
basal spot not reaching the anterior margin or 
suture; a fascia at the middle, expanded 
towards the apex near the lateral margin bui 
not touching it or the suture; a huge spot after 
the. middle, not touching the margin or the 
Miture; a. spot covering the humeral fold; a 
small elongate spot in the form of a lunette 
near the preapical margin and ending at the 
marginal spine (present in the illustrated speci- 
men-, ahseni in ihe holoiype) Undersurface 
blue-green; hairs silver. Legs metallic blue. 
Shape find sculpture, Head evenly punctured: 
deeply excavated between the eyes, mainly at 
the base; no median keel; sparsely covered 
with long, fine hair. Antennae strongly serrate. 
Pronotum deeply punctured at the sides, 
towards the middle shallow punctures with a 
central ovoid area consisting of hexagonal 



RFVISION OF BLIPRFSTID GENUS ASTRAEVS LAPORTE & GORY 



137 





5 mm ^^^— 

f 

Fig. 19, Astraeus tiffinm Kerremans 

depressions, each with a small central punc- 
ture; inflated at the 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; punctatc-striatc 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- 
lel-sided to the middle, then rounded and 
narrowed to the small marginal spine: 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 presternum and abdomi- 
nal sternitcs longitudinally grooved: sparsely 
haired. 



B rti m 



Size. Males 11.6 x 3.9 mm (1) 

x 4.8 mm CI). 



Females 13.9 



Distribution. New South Wales and Queens- 
land. 



Fig. 20. Astraeus caledonivus hativel 

Specimens examined. N.S.W.: Type. Qld: I ?, 
Acacia Creek via Killarnay, Jan. 1948, Mrs J. 
Harslet!, JH. 

39. Astraeus (Astraeus) caledonicus Fauvel, 

1904: 116. Obenberger, 1930: 365. 

FIG. 20 

Type. Holotype: 2j Baie du Sud* N. Caledonie, 

Delauney, MNHN (seen by author). 

Colour. Upper ytirface glabrous. Head and pro- 
notum black with green reflections; antennae 
purple, Elytra black with yellow reflections, 
each elytron with the following yellow mark- 
ings; a large basal spot; a small spot after the 
middle near (he suture but not touching it: a 
spot at the margin covering the humeral fold: 
a spot after the middle ai the margin but not 
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 bul 
evenly punctured; excavated between the eyes 
mainly at the apex; no median keel; without 
hairs. Pronotum shallowly but sparsely punc- 



138 



S. BARKhR 




5 mm 



Fig. 21. Astraeus robmtus sp. nov 



REVISION OF BUPRESTID GENUS ASTRAEUS LAPORTE & GORY 



139 







rw 



A B c D E F 



/l\ 



i\\ 



/1\ 



M N 



A 



/T\ 



^ 




/!\ 



/I\ 



rt\ 



(II 



r 



m 



^\ 



i_ lmm_i 

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 



/\\ 



/l\ 



rf\ 



i_1m m-i 

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; V-prothoracicus; G-elonga- 
tus; H-vittatus; \-flavopictus; l-macmil- 
lani; K-carnabyi; L-badeni; M-jansoni; 
N-oberthuri; O-carteri; V-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. 



40 



S BARKER. 




Fig. 24. Scanning electron micrograph of 
the median lobe of the pronotum 
of A. jraseriensis showing the 
basal crypt. 



Fig. 25. Scanning electron micrograph of the 
head of A. frosen'ertsis showing the 
median longitudinal keel. 



Index 

To THE Vaud St'KIF.s and Synonyms op ASTRAEUS 

the Text 

Synonyms 



With the Numbering System Adopted In 



Valid Species 

uh^mms van de Poll 4 
(uiarnsi sp. nov. 20 
httdvni van de Poll 32 
buktri sp. nov. 9 
caledomcus Fativel 39 
rarnabvi sp. nov. 3 I 
curttri sp. nov. 36 
cntssus van dc Poll 22 
cyanvus Kerremans 38 
dedariens'ts sp. nov. 8 
ddutipes van de Poll 19 
tlvngaUtii van dc Poll 27 
(lovopictus LaPorte & Gory 29 
fraxerit'tisis sp. nov. 1 1 
jraterculus van de Poll 25 
gfobosus sp. nov. 1 6 
ftoerlhigi sp. nov. 37 
iniritattis Carter 21 
irregularis van de Poll 7 
jansoni van de Poll 33 
lineatus van de Poll 5 
tntwrnillatii sp. nov. 30 
major Blackburn 23 
masters! MacKeay 17 



-— nwvrhki Blackburn 



- strandi Obenberger 



teppvri Blackburn 



- splendent van de Poll 

- simplex Blackburn 



mimttus sp. nov. 10 
mnltinotat us Vfcn dc Poll 6 
navorcliis (Thomson) 24 
oberthun' van de Poll 34 
tihxt unts sp. nov. 15 
polli sp. nov. i 
prothonicicus van de Poll 26 
p\\i>tnacus van de Poll 12 
robustus sp nov. 3 
samoueilt Saunders 18 
simitlutor van de Poll 14 
smyilti sp. nov. 1 3 
tammuu'Hsis sp. nov. 2 
vtttarns van de Poll 28 
watsoni sp. nov. 35 



Acknowledgments 

1 would like to thank the following people 
lor their assistance; Mr G. Gross, Dr E. 
Mathews and Mrs B. K. Head. South Austra- 
lian Museum; Dr E. B. Britton, C.S.J.R.O., 
Division of Entomology; Mr K. Dahms, 
Queensland Museum; Mr A. Neboiss, National 
Museum of Victoria; Dr C. N. Smilhers 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. Descarpentries, Paris 
Museum; Dr G. A. Samuelson, Bernice P. 
Bishop Museum; Dr J. Jelinck. National 
Museum of Prague; Mr K. T. Richards. 
Western Australian Department of Agriculture: 
Mr A. George, Western Australian Herbarium; 
Mr EL Slater. Canberra: Mr E. E. Adams, 
Edungalba; Mrs J Harslett, Amiens; Dr F. H. 
Uther Baker, Applecross; Mr R. P. McMillan, 
Cottcsloe; 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 BUPKESTID GENUS ASTR,1EUS LAPORTE & GORY 



141 



References 



Barker, S. (1964). — Asthraeus LaPorte and Gory, 
1837 {Insecta; Coleoptera): Proposed emen- 
dation to Astraeus. Butt. zooi. N amend. 21, 
306-307, 

Blackburn, T. ( 1890) . — Notes on Australian 
Coleoptera, with description!; ol" new species, 
pan v. Proc. Litw. Soc. N.SW, 4, 1247-1276. 

Blackburn, T. (1891). — Notes on Australian 
Coleoptera, with descriptions of new species. 
Proc. Linn. Soc. N.S.W. 5, 479-550. 

Biackbukn, T. < J892) . — Further notes on Aus- 
tralian Coleoptera with descriptions of new 
genera and species. Trans. H. Soc. S. Attst. 
15, 207-261. 

Blackburn, T. (1894). — Notes on Australian 
Coleoptera with descriptions of new species. 
Pmt: Linn. Soc. NSW. 8, 85-108. 

Buackbcrn, T. (1895). — Further notes on Aus- 
tralian Coleoptera, with descriptions of new 
genera and species. Trans, tf. Soc, S, Aust. 
19. 27-60. 

Brttton, JS R. (1970). — Coleoptera- in ''Insects 
of Australia". Ch. 30. (Melbourne University 
Press.) 

Carter* H. J. (1925). — Revision of the Austra- 
lian species of Chrysobothrls (Fam. Bupres- 
tidae), together with notes, and descriptions 
of new species of Coleoptera. Proc. Linn. 
Soc. N.S.W. 50. 225-244. 

Carter, H. J-, (1929).— A cheek list of the Aus- 
tralian Buprestidae. Aust. Zooi 5, 265 304. 

Carter, H. 1. (1931).— ChecV list of the Aus- 
tralian Buprestidae, Corrigenda. Aust, Zooi. 
6, 107-108. 

Carter, H. J. (1933).— "Gulliver in the hush." 
(Angus & Robertson: Sydney.) 

Fauvfl, A. (1904). — Fatme analylique des 
Coleopleres de la Nouvelle-Caledonie. Rev. 
d'Ent.JX 113-208. 

CJemminger, Dr. & de Harold, B. (1869). — 
"Catalogus Coleoptorum hucusque descrip- 
torum synonymicus ct systcmaticus." Vol. 5. 
(E. H. Gumnti, Monachii.) 

1.CZ.N. (1966).— ASTHRAEUS La Porte & 
Gory, 1837 (Insecta, Coleoptera): Validation 
of emendation to ASTRAEUS. Ball. Zooi. 
Nonwnc. 23, 269-270. 

Imhoff, T.. (1856). — "Dersuch einer Eiufuhrung 
in das studium der Koleoptcrn." (Basel.) 

Kr-KUEMANS, C. (1885). — Enumeration dc Bup- 
restides decrits posteneuremcni au Catalogue 
de M. M. Gemminger & de Harold IS70- 
1883. Ann, Soc. ent Bel}>, 29, 11W57. 



K.erk£M.vns, C. (1892). — Catalogue synonymique 
des Buprestides decrits de 1758 & 1890. 
Menu Soc. enl. Bela. 1. 1-304. 

Kfrrrmans. C. (1900). — Buprestides nouvcaux ct 
remarques svnonymiques. Ann. Soc. ent. Belg. 
44, 282-351! 

Kkrrkmans, C. (1902). — Genera lnsectorum 12. 
Coleoptera; Senicornia Fam. Buprestidae. 

Lacordaire, T. (1857). — "Histoire naturelle des 
Insectes." Genera des Coleopleres, Vol. 4. 
(LibraJrie Encyclopedique de Roret: Paris.) 

L^PORTr, F, L, & Gory, H. ( 1837).— 'Histoire 
Nalurcllc ct tomographic des lnseclcs 
Colcoptercs." Tome I. Monographic des spi- 
locera. euiydera, nyeieis, eunosuis. Bupies- 
lides (Chrysoehroiles et AgriUtes). tP. 
Dumenil* Pari*.) 

MacI.raY, W. (1873). — Notes on a collection of 
insects from Gayndah. Trans, ent. Soc. 
N.S.W. 2, 239-318. 

Mastkks, G, (1871). — ''Catalogue of the des- 
cribed Coleoptera of Australia." (Sydney.) 

Masters, G. (1886). — -Catalogue of the des 
eiibed Coleoptera of Australia- Part HE. 
Proc. Linn. Soc. N.S.W. II, 1-106. 

OnrNfirRriRR, J. (1928). — Opuscula B;ipresto- 
logica I. Arch. Naturxexch. 92, 113-224. 

Ouj-nihw(.;h,k, (. (1930). — In **Catalogus Coleop- 
terus." Vol. 12, Buprestidae 2, pp. 365-367. 

Oblnblroer, I. (1936). — 4 "Festschrift zum 60 
Gebudst Embrik Strand, Rica." Vol. I, pp. 
97-145. 

van de Poll, J. R. H. N. (1886).— Description of 
three new species and a synopsis of the Bup- 
restid genus Astraeus, C. & G. Notts Leviien 
Mtts. 8, 176-180. 

van nE Poli > J, R. H. N. (1889).— Monographi- 
cal essay on the Australian Buprestid genus 
Astraeus C. et G- Tjschr, v, Ent, 32, 79-1 10. 

van de Volu J. R, H. N. (1892).— Note sur quel- 

ques especies d' Astraeus, Tjschr. m. Em. Jfi, 

67-68. 
Saunders, E. (1868). A revision of tbe Aus 

tralian Buprestidae described by The Rev. 

F. W. Hope. Trans, cut. Soc. Land. 1S68. 

1-68. 
Saunders, E. (1871). — "Catalogus Buprcstidarum 

synonymicus et systematieus.'' 
Thomson. J. (1856).— Description dc dix Colcop- 

teres. Rev. Mas. Zooi. 2, 112-118. 



FACTORS AFFECTING THE DISTRIBUTION OF THE LEPTODACTYLID 
FROG GEOCRINIA LAEVIS IN THE SOUTH-EAST OF 

SOUTH AUSTRALIA 

byR. G. Beck* 



Summary 

BECK, R. G. (1975). -Factors affecting the distribution of the leptodactylid frog Geocrinia laevis in 
the south-east of South Australia. Trans. R. Soc. S. Aust. 99(3), 143-147, 30 August, 1975. 
Geocrinia laevis (Giinther), a species of leptodactylid frog previously recorded from Victoria and 
Tasmania, was first reported in South Australia in 1966. A survey has revealed that this new 
population is an extension of that known to exist in south-west Victoria. The frog is, extremely rare 
in South Australia, occupying only a fraction of the potential habitat for which it shows preference. 
Some thoughts on this are tendered as a basis for further studies. 



FACTORS AFFECTING THE DISTRIBUTION OF THE LEPTODACTYLID FROG 
GEOCR1NIA LAEV1S IN THE SOUTH-EAST OF SOUTH AUSTRALIA 

by R. G. Beck* 

Summary 

Bkck, R. G. (1975). — Factors affecting the distribution of the leptodactylid frog Geocrinia 
luevis in the south-east of South Australia, Trans- 7?. Soc. S. Artst, 99(3), 143-147, 30 
August, [975. 

Geitcruihi laevis (G anther), a species of leptodactylid frog previously recorded from 
Victoria and Tasmania, was first reported in South Australia in 1966. A survey has revealed 
that thi.s new population is an extension of that known to exist in smith-west Victoria, The 
frog i3 extremely rare in South Australia, occupying only a fraction of the potential habitat 
for which it shows preference. Some thoughts on this are tendered as a basis for further 
studies. 



Introduction 

The leptodactylid frog GeotrJnta laevis 
(Ciunther) has been described in detail by 
I jttlejohn & Martin ( 1 964 ) . The specimens 
found in the south-east of South Australia 
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 laevts was known 
to exist in four disjunct populations 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 (Woodruff & Tyler 1968) of a speci- 
men from Marsh's Swamp near Me Burr, some 
SO 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, 
Mayurra, Riddoch, Hindmarsh, Grey, Young, 
Nangwarry, Mingboof, Blanche and Gambier, 
and the adjoining area east of the state border 
to (lie Gleuelg 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 



survey, field work was concentrated around 
Marsh's Swamp (Site 3, Fig. J. Grid reference 
353362, Australian Army Survey Map, Penola, 
1:250,(100 Sheet SJ/54-6). From here the sur- 
vey extended to the north-west, following the 
general /one of influence oi 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 been found, 
detailed botanical surveys were made of the 
surrounding areas, particularly of the nearest 
probable breeding site. Following the establish- 
ment of definite ecological patterns, soil sur- 
veys of these areas were undertaken. 

Typical habitats 

Geocrinia taevis normally lays eggs in areas 
which later become flooded, Breeding sites 
may be I he 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 *i ah (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 m lype 



Lynwood Park, Mil Lei, S. Aust. 5291. 



144 



R. G. BECK 




Fig. I, Map of the lower south-east of South Australia and adjacent south-west Victoria, showing main 
collecting sites of Gcocriftia laevis in relation to the Reedy Creek-Dismal Swamp Corridor. The 
inset shows known Australian distribution prior to 1966. (After Littlejohn & Martin 1964, 
1965.) 



and consist of any of Eucalyptus baxferi, E. 
obliqua, E. hitheriana or E. ovata according to 
the soil type. However, the understorey in the 
viciulty of the swamps is remarkably con- 
stant, and four species have been found in all 
areas, viz. Acacia nielunoxylon, Leptospermum 
jurriperimmi, Melaleuca squarrosa and Heli- 
chrysitm detultoiclea. 

The soil map of Stephens er al. (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. laevis have 
now heen found in South Australia, arid a 
further 10 in Victoria between the State bor- 
der and the Glenelg River, which, prior to 
1966, represented the known western limit of 
distribution. The species is common at Dart- 
moor, where on one occasion, 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. Museum. 

Of particular interest is the specimen col- 
lected at Grid rcf. 376346 (site 7, Fig. 1 ), in 
a deep pit dug to observe pine tree root growth 
at the Forest Research Station north of Mt 
Gambier. 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. 1, 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 larvae 
washed down one of the many man-made 
drains which cross the area between the Milli- 
cenl Hills and the coast, 



GEOCRINIA LAEVIS IN THE SOUTH-EAST OF SOUTH AUSTRALIA 



145 



TABLE i 

Recorded distr'thnt'ton of Geocrinia laevis in south-east South Australia and nearby Victoria 
Refer A.A.S, Map, Penola, 1:250,000, Sheet SJ/54-6 



Grid 

Reference 


Site No. 
(Fig. 1) 


No. of 

Specimens 


Collector 


Remarks 


353362 


3 


I 


H. Minchan & 
C. Taylor 


First S. Aust. specimen, SAM, 
R8118 


353362 


3 


1 


D. Woodruff 


Melbourne University Zoology 
Dept. 220/67 


353362 


3 


6 


R. Beck 


All within 2 km of original site 
at Marsh's Swamp 


353362 


3 


3 


D. Klcm 


Marsh's Swamp. One specimen, 
SAM, R10583 


354364 


3 


1 


F, Aslin 


2 km NE of original site at 
Marsh's Swamp 


38(35! 


6 


1 


C. Taylu; 


Earl*K, 16 km N of Ml Gambier 


381353 


6 


1 


R. Beck 


Hem's scrub, 17 km N of Mt 
Gambier 


378351 


6 


1 


F. Aslin 


Telford's scrub 


378349 


6 


1 


D. Klem 


2 km S of Telford's scrub 


376346 


7 


1 


D. Klem 


Forest Research Station, Soil Pit, 
SAM, R 13974 


368355 


5 


1 


D.KJem 


Hogarth's scrub 


326360 


| 


1 


J, Aslin 


Canunda Reserve, SAM R 13975 


342366 


2 


1 


F. Aslin 


Night Road Survey, 5 km NE of 
MilliceiU 


361361 


4 


] 


A. Rowley 


Lake Leake, SAM, R 141 99 




Refer A.A.S. Map, 


Hamilton, 1 :250.000, 


Sheet SJ/54-7 


414325 


8 


6 


H. Roach 


16 km E of State Border on High- 
way No. 1, SAM, R 10780 


414325 


8 


1 


R. Beck 


Same locality 


427324 


9 


Many 


R. Beck 


Common in Glenelg River 2 km 



4283 1 i 

409313 

424310 



up and downstream from Dart- 
moor 



Refer AA.S. Map, Portland, 1:250,000, Sheet SJ/54-11 



10 

12 
II 



F. Aslin 

F. Aslin 
J . Aslin 



East of Glenelg River near Jones' 

Lookout 

Lower Glenelg River 

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 is so gradual, 
being only a few cm per km, that in extremely 
wet years such as 1S96 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-cast 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 Ardoo 
Creeks. 

In the study area, G. laevis is restricted to 
areas receiving an average annual rainfall of 
700 mm 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 pattern, it would be reasonable 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. 



|4G 



R, O BECK 



Discussion 

Three tacts emerge from ihe survey: 
J While it is certain that more specimen* will 
be discovered within and beyond the present 
known range of distribution, (7. lovvis is 
extremely rare in south-east South Australia. 
Even at Marsh's Swamp, where most speci- 
mens have been found. I have not positively 
tdentitled its calK during the Aprit-May- 
Junc breeding season, when calls are com- 
monly heard at Dartmoor, Victoria. How- 
ever, Woodruff & Tyler (!96«) have re- 
ported the recording of a mating call al 
Marsh's Swamp. 
2. According to prevent known records, the 
species occupies only a fraction of the poten- 
tial habitat for which it shows preference 
wilh respect to soils, vegetal ion um\ rain- 
fall 
i. G. Icwvi.s was always found under the shelter 
of logs, litter, or stones during the day. As 
a result, the species has not been found in 
areas cleared tor agriculture or pasture pro- 
duction 

Some suggestions for the reason tor ih'S 
restricted distribution are tendered as a banis 
for further work by someone wilb more time 
and resources than the present author. Rain- 
i.iU and associated weather patterns are 
probably the major factors influencing the 
spread of any frog species. In 1967, an extreme 
draught was experienced in the lower south- 
east of South Australia. The aveiage annual 
rainfall at Mr, Gambier is 776 mm, but in that 
year only 402 mm fell, and unofficial figures 
from the Diurnal Swamp area were as low as 
2SU and 331,1 mm respectively. In this single 
dry year, many of the local 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 a few of 
the autumn arid winter breeders actually 
spawned 

Since Crocker & Wood (1947) first presen- 
ted evidence for a recent at id period, many 
workers have commented, and they arc about 
equally divided in their acceptance or rejec- 
tion of the concept (Mulvancy & Golson 1971 : 
Littfejohn 1967). 

Gentilli i 1961 ) had already offered an 
explanation for this diversity of opinion; ^Aus- 
tralia is a targe land, spanning several maior 
climatic belts, and may have experienced dif- 
ferent climatic change*, in various parts of the 



continent at the same time.' 1 This is supported, 
on a one year basis at least, by an examina- 
tion of the Commonwealth Bureau of Mtrieoro 
logy rainfall figures from major Australian 
mainland centres for 1967, Perth. Broume. 
Darwin, Cairns. Brisbane and Sydney had 
greater than average rainfall, Geraldton and 
Alice Springs were Only slightly lower than 
average, whereas Adelaide and Melbourne 
received approximately half their average 
amount. The coastal strip from the head of 
the Great Australian Bight to c«ist of Mel- 
bourne obviously was the worst affected area. 

Cluirchilf (I968-) has shown that, in 
Western Australia, during the past 5000 years 
there have been several fluctuations in climate 
of sufficient magnitude to cause the replace- 
ment of jarrah forests with karri and vice 
ier\a. Similarly, recent wofk by Dodson 
(1974) in the study area indicates variations 
in climate, wilh relatively drv periods between 
5000 and 2000 R.P., and -.gain since 1300 
BP. 

Gentilli (1^72) also supports the concept 
of changing climate. "It must be stressed thai 
climate, being rhe result of numerous variables 
variously combined in space and time, can 
vary, fluctuate, oscillate, or just change." It 
follows that changes such as these must pro- 
duce equally dramatic changes in the local 
fauna. 

With pluvial conditions prevailing in 
southern Australia during the last glacial 
period, Bassian species extended their range 
( I.itllejohn ]'>67) and it is reasonable to 
expect that G. fumh occupied much of the 
lower south-east of South Australia- With the 
return of present-day weather conditions, il.s 
range would have decreased but not to the 
limits found today. U is therefore suggested 
that in the recent past, possibly much more 
iccently than thai postulated by Crocker ik 
Wood 1 I947K there has been a period suffi- 
ciently arid to cause the withdrawal of G. 
Ifwvis and perhaps some other anuran species 
to the more favoured puTtS of south-weM Vic- 
toria or even the Grampians. It must be 
emphasised that in this context the term "'arid" 
is relative rather than absolute. 

Inoccupation of the south-east of South 
Australia by G. /«em would have occurred by 
way of the Cilenelg River and the Dismal 
Swamp when wetter conditions returned. This 
may have taken place even as recently js with 
in historic times. 



GEOCHINIA LAEVIS IN THE SOUTH-EAST Ob SOUTH AUSTRALIA 






Since the settlement of the isouth-eaKl of 
South Australia by white man, the aTea 
reached its physically wettest stale in 1S96. 
This Information was obtained from S.E. 
Drainage Board records and also, some years 
ago. from old residents who remembered the 
district during the nineties. They claimed "you 
could tow 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 century to the clearing of 
natural scrub. Subsequent widespread plant- 
ing of pines and establishment of drainage 
schemes have greatly reduced surface waters 

Very recent reoccupation of the south cast 
by O v . /tft'Ws as outlined above would account 
for the limited spread of thz species. Further 



distribution has been prevented by a combina- 
tion of clearing the natural habitat, and the 
drying out of the district by drainage, pine 
plantations and the establishment of better 
pastures. It is obvious that much more study 
is required to explain satisfactorily this limited 
distribution. 

A din o wlcelgm en Is 

The author gratefully acknowledges the 
col lection of specimens by the following mem- 
bens of the South East Field Naturalists' 
Societies: Mr and Mrs F. Aslin, Messrs D. 
Klcm, P Roach, C. Taylor, and Miss A. Row- 
ley. Also thanks are due to Mr M. ). Tyler and 
Dr M. Littlejohn for advice and encourage- 
ment, and to Mr D. Lewis from the S.A. Dcp1* 
of Agriculture for his help with soil identifica- 
tion. The Royal Society of South Australia 
Irtc, ? kindly granted $50 towards petrol 
expenses. 



References 



CM rcuill, T). M. ( 19681.— The distribution 
and prehistory of Eucalyptus diver sicol or F. 
Muell., E. marxinula Donn. ex JSm,» and E, 
calophylhi R.Hr f in relation to rainfall. Ausi. 
I. Bot. 16, 125-51. 

Colville, J. S« & Holmes, .1. W (1972) Water 

table fluctuations under forest and pasture 
in a karstic region of Southern Australia. 
Hydrology 17, 61-80. 

Crocker, R. L., & Wood, I. C. (1947).— Some 
historical influences on the development of 
the South Australian vegelational communi- 
ties and their bearing on concepts and classi- 
fication in ecology, Trans, H. Sac. S, Anst. 71, 
91-136. 

Dodson, J. R. ( 1974).-- Vegetation histury and 
water level fluctuations at Lake I. cake, south- 
eastern South Australia. I. 10,000 B.JV to 
Present. Attst. '. Bot. 22. 719-741. 

Gentilll J. U96I), — Quaternary climates of the 
Australian region. Ann. N.Y. Acad. Set. 95, 
465-WI. 

CiLNiit.LL J. (IV72). — "Australian Climatic Pat- 
terns." (Nelson - Melbourne.) 

LrnLEJOHN. M. J- (1967). Patterns of Zoo- 
geography and Speciation in South-eastern 
Australian Amphibia. In A. H. Weatherlcy 
(F.d.), "Australian Inland Waters and their 
raunty Klcvcn Studies." (Australian National 
University: Canberra.) 



Littlejohn, M. J., & Martin, A. A. (1964). — 
The Crima laevis complex (Anura: Lepto- 
dactvlidae) in south eastern Australia. Autt. 
J. Zool. 12. 70-83. 

Littlejohn, M. J.. A Makiin. A. A. (1965). — 
The vertebrate htuna of the Bass Strait 
islands: 1. The Amphibia of Flinders and 
King Islands. Hoc, H, Sol. Vict, 79, 7.47-256 

Martin, A. A. (1967). — Australian Aoitran lite 
Histories: Some Evolutionary and Ecological 
Aspects, In A. H. Weuthcrfey (Ed.), "Aus- 
tralian Inland Waters and their Fauna, Eleven 
Studies." (Australian National University: 
Canberra.) 

Mulvanev, D J., & Golson, .1. (1967).— 
"Aboriginal Man and Environment in Aus- 
tralia/' (Australian National University: 
Canberra.) 

Sprjgg, R. C. (1952).— The Geology of the 
Soulh-Easl Province. South Australia, with 
Special Reference to Quaternary Coast-line 
Migrations and Modern Beach Developments. 
South Australia Department of Mines, 
Bulletin 29. 

Stephens, C. G, Crocker, R, L.. Butler, B.. £ 
Smith, R. (1941)— A Soil and Land U<e 
Survey of the Hundreds of Kiddoch, Hind- 
marsh. Grey, Young, and Nangwarry, County 
Grew, South Australia. Conn, Sci. /nd< R&i 
Aus't. Bull. 142. 

SVoodklm-, D. S.. & Tyi.kk, M. J. (t%8).— 
Additions to the Frog Hmina of South Ausi 
Ree. S. Attst, Ate. 15(4), 705-709. 



THE THALLUS AND SPORE DEVELOPMENT OF LOBOSPIRA 
BICUSPIDATA ARESCHOUG (DICTYOTALES: PHAEOPHYTA) 

BYTlKVAHEDELSTEIN*ANDH. B. S. WOMERSLEYf 



Summary 

EDELSTEIN, TIKVAH &, WOMERSLEY, H. B. S. (1975).-The thallus and spore development of 
Lobospira bicuspidata Areschoug (Dictyotales: Phaeophyta). Trans. R. Soc. S. Aust. 99(3), 
149-156, 30 August, 1975. 

The apical growth of Lobospira bicuspidata, release of tetraspores, and growth of the spores in 
culture to plants up to 1 cm across, are described. Both development of the axes and growth of the 
sporelings is from a marginal row of apical cells, and the thallus is monopodially developed. 
Lobospira is therefore placed in the Zonarieae group of the Dictyotales. 



THE THALLUS AND SPORE DEVELOPMENT OF LOBOSPIRA BICVSPJDATA 
ARESCHOL'G (DICTYOTALES: PHAEOPHYTA) 

by TiKVah Edelstein* and H. B. S. Womersleyt 



Summary 

fcDELSTFiN, Tikvar, & Womersley. H, B, S, (1975).— The lhallus and spore development of 
Lubospira hicuspidata Areschouu ( Dtctyotatcs: Phaeophvta). Trans. H, Sac. S. Aitsi. 
99(3), 149-156, 30 August, 1975. 

The apical growth of Lobospha hicuspidata, release of tetrasporcs, and growth of the 
spores in culture to plants up to I cm across, are described. Both development of the axes 
and growth of the sporelings is From a marginal row of apical cells, and the thallus is niono- 
podially developed. Lohcfspira is therefore placed in the Zonai ieae group of ihe Diclyolales. 



Introduction 

Lohospifa bicuspichsta Areschoug is a dis- 
tinctive brown alga referred to the Dictyotales. 
It occurs from Nickol Bay, Western Australia, 
around southern Australia to Eden, N.S.W.. 
and around Tasmania (Womersley 1967. p. 
2! 5) 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. 2A) is easily recognized by 
its spirally twisted axes, with a phyllotaxis of 
about 1/3, bearing laterals with bicuspid, de- 
terminate ramuli (Harvey 1858, pi. 34). and 
with lower branches bearing recurved attach- 
ment tendrils. Kjellman (1897, pp. 295, 297) 
and Oltmanns (1922, p. 185) considered thai 
ihe thallus develops from an apical cell, with 
sympodial branching, and Lobospira has thus 
been considered as a member of the Dictyo- 
tcae. The sporangia, about I00/*m in diameter, 
occur scattered over the thalitis (Fig. IB), they 
are developed from cortical cells and sunken 
in the thallus (Fig. IB). 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 occasion. 
Sexual reproductive cells alio have never been 
observed. While Lohospira has usually been 
placed in the DictyotaJes, and the Dtctyoteae 
I Womersley 1967, p. 215), its relationships 
have not been established. 

This paper reports observations on apical 
development, spore release and early growth 



of the thallus, made in 1972 while the firM 
author was on leave at the University of 
Adelaide. 

Methods 

Plants (ADU, A42264) were collected in 
drift at Aldinga reef. South Australia, on 27 
May, 1972, and transferred to the laboratoiy 
in sea water. The specimens (Fig. IB) bore 
mature sporangia, many of which released 
tetrads of spores. Fertile branches were placed 
in a glass jar with Provasoli ES medium 
(Starr 1971, p, 359) in a 15X culture room, 
and spores allowed to settle on slides durinv 
the next two days. On day 3 the slides with 
attached sporelings were transferred to pe:n 
dishes (5 cm in diameter), and germanium 
dioxide at a concentration of 5 p. p.m. added 
to the medium. Single sporelings from 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 sporelings. Afler 4 weeks 
cultures were maintained in SWM 3 medium 
(Chen. Edektein & McLachlau 1969). 

As the sporelings developed, considerable 
difficulty occurred with bacterial (and at one 
stage fungal 1 ) contamination. Addition of peni- 
cillin to the Provasoli medium bad little effect, 
but streptomycin (100-150 mg streptomycin 
sulphate/ 1 of seawaier) eliminated most of Ihe 
bacteria, and a commercial fungicide, Myco- 
statin-Dusting Powder ( 1 ,000.000 unils I 
(E. R. Squibb & Sons Ltd, Melbourne) proved 



1 Atlantic Regional Laboratory. National Research Council of Canada, Halifax, N.Sw, Canada. I?\RCC 

Mo, 14511.) 
-" Department of Botany, University of Adelaide. Adelaide, S. Aust. 5000. 



50 



TIKVAH EDfcLSTHIN & H. B. S. WOMERSLEY 




Fig- f. A. Apical development, with two -young latctals present, the older one (right) becoming bicus- 
pid. Trie axis meristem (a.m.) continues growth of the branch, and the lateral meristcms 
(I.m.) may or may noi develop further into laterals (ADU, A42264). 

B Cross section of ramulus bearing sporangia (ADU. A42264). 

C. Various sporclings I week old, with a rhizoid 3-5 cells long and early .stages of the u ceU- 
mass". 

O. A sporeling 3 weeks old, with a well developed cell-mass. 



to be effective when used as a single treatment 
of 200 mg of "Mycostutin" per 100 ml of sea 
water for 20 hours. Repeated treatments with 
streptomycin and Myeoslalin were used and 
frequent cleaning of the sporclings with glass 
needles was carried out. While this damaged 
some sporclings, the majority survived and con- 



tinued to develop to plants consisting of clus- 
ters of branches up to 1 cm across, but from 
which it was impossible to clean the epiphytes. 
In general, cultures were maintained at 15 3 C 
under a light intensity of 600-800 lux provided 
by 40 watt white fluorescent lights, and a 
regime of 14 hrs light/ 10 hrs dark. The me- 



THALLUS AND SPORE DEVELOPMENT OF LOBOSPIRA 



151 



B 




c ' 1 1 * 'H HP? 


9 ■ jPJ 1 ? 1 



m 




Fig. 2. 



152 



TIKVAH EDELSTEIN & H. B. S. WOMERSLEY 



- 

■ 

kj «... *...*? 



I 



\^>i3P 



*" ». 



\ 4 " ' 






*-* 



B 



**% 



50/UlTi 




►4 

M 



200/jm 



'»2ffin 




100/jm 






! %* 



^1 



Fig. 3. 




200/jm 



THALLUS AND SPORE DEVELOPMENT OF LOBOSPIRA 



153 



200/jm 




Fig. 4. 



154 



TIKVAH EDELSTEIN & H. B. S. WOMERSLEY 




Fig. 5, 



1KALLUS AND SPORE DEVfcLOPMFNT OF LOHQSMRA 



155 



Fijj, 7- A Thallus of Lobospira, showing basal entangled branches with tendrfJ-beaflng laterals, and 
erect branches with ultimate bicuspid ramuli. Stanley Bench, Kangaroo I„ S. Atwl. 
{f ( h'h(tin & Krujl 126, 25.iv.197Z). 
B. Surface view of a branch showing tctrasporanam (undivided) and released tcttaspurcs 
(arrow). (ADU, A42264.) 

Fi£. 3. A. A multicellular sporeiing, with rhizoid. about 2 weeks old. 

B. A plant about 3 weeks old, showing development, of a flat thatlus wkh a memtemalic 
margin. 

C Planl t? weeks old, with two lobes. 

O Plant 6 weeks old, showing development of lobes from the margin and especially from 
the rhizuidal region. 

Fig. 4. A, Plant Ik weeks old, wilh 3 main lobes. 
H. Planl 71 weeks old. becoming convolute. 
C Plant Ml weeks old. with several irregular lobes. 

D. Plain 174 weeks old, consisting of many irregular branches, each with marginal lobes of 
various sizes. 

Fig. 5 4, Plant 24* weeks old: general view of merislemalic apex, with degenerating cells in CCftt/c 
of each lobe. 
H, As in A. with two darkly-staining cells (arrows) separating in centre of margin of lobe, 

C. As in A, with degeneration of cells between two darkly-staining cells. 



<lium was changed every 3-5 days for the first 
6 weeks and thereafter at weekly intervals- 
Results 
Development of the thatiua 

Dissection pf apices (Fig. 1.4) of mature 
field plants of Lobospira shows lhal develop- 
ment is from a row of apical cells, from which 
lateral groups of apical cells separate of! alter- 
nately and differentiate rapidly to form a 
larger, abaxial, spinous or bicuspid process 
wilh a group of meristematic cells on its adaxial 
side. Growth is thus monopodial and the affi- 
nities of Lobospira arc with the Zonarieae. In 
actively growing apices, the young laterals 
deveiop rapidly and overtop the apex. The 
mcrLstematic group of cells on the lateral may 
persist indefinitely, in which case a long lateral 
develops; or it may persist to give a shon 
lateral with only a few pointed famuli; or it 
may not develop lurther. resulting in only a 
pointed or bicuspid ramulus, at the apices of 
which a single cell remains prominent (Fi*. 
\A ). Alternate series of these ramuli fringe the 
longer axes or branches. 

The indefinite axes twist spirally so that the 
branches or ramuli become arranged wilh a 
pbyllotaxis of about 1/3. 

The mature thallus consists of a cortical 
layer of cells which are arranged more or less 
in longitudinal lines and tend to radiate up- 
wards. The medulla consists of cells of similar 
size but rather irregularly arranged (Fig. IB), 
and is 2-5 cells thick; in older branches a 



slight midrib is present where the medulla js 
thicker- Older axes are ovoid to round in sec- 
tion and a central core of narrower and mure 
elongate celJs may be present. Hair groups are 
of frequent occurrence on the thallus. 

Sporofigia and spore development 

Division of the sporangia into four tetra- 
hedrally arranged, non-motile spores (Fig. 2nT) 
appears to occur only shortly before their 
release, and no divided sporangia have been 
observed in any herbarium material. Fertile 
collections have been made mainly in autumn 
(April to June), 

The spores germinated on slides within two 
days of their release, forming a short rhizoid 
which was cut off by a cross wall when slightly 
longer than the spore. The rhizoid became 3 
or 4 cells long before the sporc-rcsidual cell 
enlarged and divided, fry day 7, a variety of 
cell arrangements (Fig, \C) was present 
amongst the sporclmgs, which, during the next 
2 weeks, developed into clongatc-ovotd masses 
of cells (Figs ID. 3/4), their arrangement 
depending un the early cell divisions. 

From this cell mass, which was attached by 
a relatively long rhizoid of several cells, a Hat. 
ovatc-spathulate disc of cells developed, mostly 
two cells thick and with a distinct apical row 
of mcristcrnatic cells ( Fig. 35, plant 3 weeks 
old). During the next 4-5 weeks, further rhi- 
zoids developed ftom the basal cell mass, and 
the flat, erect, frond developed further from the 
apical metistein, usually becoming lobed (Fig. 
3C>, Smaller lobes developed both from the 



156 



TIKVAH UDELSTEIN & H. B. S. WOMERSLEY 



basal cell mass and lower parts of the erect 
fronds (Fig. 3D). 

By S 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 
weeks, numerous fronds were present (Fig. 
4C). usually branched or lohed, and thalli 17J- 
24i weeks old formed a cluster of fronds (Fig. 
4D) up to 1 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 mert- 
stematic and lateral margins, the fronds were 
mostly two cells thick, increasing to 3 or 4 
cells 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. 
5A). Plants 24i weeks old showed a further 
apical development of possible significance, in 
that centrally along the meristematic margin 
of each lobe, two cells became more promin- 
ent (Fig. 55) with denser protoplasm,, and 
breakdown of tissue occurred between them 
(Fig. 5.4, ( ). Whether this was only a break- 
down feature before death is uncertain, but the 
two cells concerned, which when first notice- 
able were densely protoplasmic and appeared 
healthy, could possibly correspond to the single 



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- 
molilc 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 Zonarieae and not the Dictyotcac. 
The distinctive morphology of Lobospira separ- 
ates it generically from all other genera of the 
Zonarieae. 

While the sporeting and juvenile stages are 
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 
occurs at this stage. 

Acknowledgments 

The second author acknowledges a grant 
from the Australian Research Grants Commit- 
tee for technical help, and the assistance of 
Mrs E. I.. Robertson and Miss C G. Anderson 
with the culture work and illustrations. Dr 
G. T. Kraft assisted in early stages of the 
study. 



References 



Chen. L. C-M- Edei.sthin, T.. & McLachlan, J. 
(I9fi9). — Bonncmuisoniii hantifcra JIariot in 
nature and in culture. /. Phyeot. 5, 21 1-220. 



Harvey. W. 11 (1858), 
Vol. I, Hates 1-6(1. 



Phycologia Australica 1 



Kjfu man, r\ R. (1897).— Fhoeophyceae. In A. 
Hnglcr and K. Prantl, "Die Naturlichcn Pflan- 
zenfamklicn". Th. 1, Abt. 2, pp. 176-297. 



Oltmanns, F. (1922). — "Morphologic una" Bio- 
logic dcr Algen/* Vol. 2. (Jena.) 

Stark, R. C. (1971).— The culture collections of 
algae at Indiana University — additions to the 
collection July 1966-Jiily I97K /. Phycol. 7, 
350-362. 

WoMERS^EV, H. B. S. (1967).— A critical survey 
of the marine algae of southern Australia. 11 
Phaeophyta. Attst. A Dot. 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 

Dolhunty, J. A. Shoreline Shingle Terraces and Prehistoric Fillings of Lake 

Eyre 183 

Late, P. K. Notes on the Relict Palm Livistona mariae F. Muell. in Central 

Australia 189 

Twidale, C. R., and Bourne, Jennifer A. Geomorphological Evolution of Part 

of the Eastern Mount Lofty Ranges, South Australia - - 197 

Haslett, P. G. Woodendinna Dolomite and Wirrapowie Limestone — Two New 

Lower Cambrian Formations, Flinders Ranges, South Australia 211 

Womersley, H. B. S., and Cartledge, Sally A. The Southern Australian Species 

of Spyridia (Ceramiaceae: Rhodophyta) - 221 

Obituary— Graham Frederic Whitten, M.Sc. 235 

Annual Report of Council, 1974-75 237 

Award of the Sir Joseph Verco Medal -238 

Balance Sheet 239 



PUBLISHED AND SOLD AT THE SOCIETY'S ROOMS 

STATE LIBRARY BUILDING 

NORTH TERRACE, ADELAIDE, SA. 5000 



HYDROIDS OF BRUNY ISLAND, SOUTHERN TASMANIA 

byJeanetteE. Watson* 

Summary 

WATSON, Jeanette, E. (1975).-Hydroids of Bruny Island, southern Tasmania. Trans. R. Sot: 
£ Aust. 99(4), 157-176, 30 November, 1975. 

A systematic collection of the sublittoral hydroids of Bruny Island, southern Tasmania, using 
SCUBA, yielded 34 species, including three newly described, three new records for Australian 
waters and 11 new records for Tasmania. 

Most Haleciidae, including two new species, are epizoic, occupying sheltered microhabitats. 
Few species of Sertulariidae are recorded, and Amphisbetia operculata is now rare in a former 
habitat. The Plumulariidae is represented mainly by small epiphytic forms, and Plumularia angusta, 
P. crateriformis and P. wilsoni are recorded for the first time from one locality. Two species newly 
described, Halecium bruniensis and H. luteum, are each closely related to endemic New Zealand 
species. The occurrence of these, and the first record of Salacia farquhari outside New Zealand 
waters (where it also occurs south of 43's) suggests active progress of speciation and dispersal 
across the Tasman Sea. 



HYDK01DS OF BRUNY ISLAND, SOUTHERN TASMANIA 

by Jeanette E. Watson* 

Summary 

Watson, Jeanette, E. (1975). — Hydroids of Bruny Island, southern Tasmania. Ttan.v. R. Soc. 

R Atmt. 99(4), 157-176, 30 November, 1975. 

A systematic collection of the sublittoral hydroids of Bruny Island, southern Tasmania, 
using SCUBA, yielded 34 species, including three newly described, three new records for 
Australian waters and 1 1 new records for Tasmania. 

Most Haleciidae, including two new species, are epizoic, occupying sbellcred micro- 
habitats. Few species of Sertulariidae are recorded, and Amphisbetia operculata is now rare 
in a former habitat. The WumuJariidae is represented mainly by small epiphytic forms, and 
Ptutmdaria an$u$ta, P r ermeriformis and P. wilso/ti are recorded for the first time from one 
locality. Two species newly described, Halecium bnmienxh and H. luteum, are each closely 
related to endemic New Zealand species. The occurrence of these, and the first record of 
Salacia farquhari outside New Zealand waters (where it also occurs south of 43 C S) suggests 
active progress of speciation and dispersal across the Tasman Sea. 



Introduction 

Bruny Island (43°25'S, l40 e 20'E) is situated 
oft the cast coast of Tasmania, 25 km south o( 
Hobart. The island is approximately 50 km 
long and is separated from Ihe Tasmanian 
mainland by the narrow waters of the D'Entre- 
castcaux Channel, The indented coastline of 
Bruny Island provides a range of environ- 
mental conditions varying from the sheltered 
bu* swift flowing tidal waters of the D'Entre- 
ensteaux Channel to the rough-water eastern 
coastline 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 t Sampling, using 
SCUBA equipment, was carried out. over the 
entire depth range (0-20 m) presented by the 
rocky sublittoral along the coastline at Satellite 
Island, Great TayJor Bay, Simpsons Bay, the 
adjacent DTIntxecasteaux Channel, and at 
Fluted Cape, Penguin Island and Adventure 
Bay 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 
or the sandy beaches of the D'Entrecasteaux 
Channel. It is unlikely that cither 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 species 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 Dcrwcnt 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 1 1 new 
records for Tasmania, including 3 new records 
for Australia, and 3 species are newly described. 
Only 19 species of Hodgson's list appear in the 
present collection. 

Holotypc and parutye mieroslides of new 
species, and other microslides and material, are 
lodged in the National Museum of Victoria, 
Melbourne (referred to as NMV). 

No athecate hydroids were recorded from 
Bruny Island. The Campamilanidae is repre- 
sented by 5 species, Lafoeidae by L Haleciidae 
6, Syntheciidae J, Sertulariidae 9, and the 
Plumulariidae by 12 species, including 2 varie- 
ties of one species. 

LIST OF SPECIES 

* Denotes a new record for Tasmania 
t Denotes a new record for Australia 



National Museum of Victoria, RussclJ Stteet, Melbourne, Vic 3000, 



I5K 



/EANRTTF Jr WATSON 



Til EC AT A 

Family CAMPANULARHDAK 

*i Campanularia ambiplica Mulder A: Trcbifcock- 

"* Campttnuluria pule rath ecu Mulder & Trcbdcocfc. 

Clytiu -\p. 

Orthopyxix caliculata Hincks. 

Silieitlaria mwv? Meyen. 

hamiiy LAFOEIDAf 

flfbt-lhi ?funu Millard. 

Family HALECUDAK 

HoU'cium dt'Jiartulum Coughtrey. 

flatccium sp. 
- Halecium heanu (JohuNlun), 

HuU'riuM brutriensis n.sp, 

Halecium htteum n.sp, 

Phylactothcca armata Stechow. 

Family SYNTHRCJIDAF 

Synihi'ciutn pat alum Busk. 

Family SERTULAiUIDAE 
E Sutu<to farqidiari (Bale). 
Stereo theca clot/gum (Lamouroux). 
Scrtularclla robusta Coughtrey 
Symplectoscyphuz pygmacus Bale. 
Sertularia acuta (Stechow) - 

* Sertalaria macraenrpa Bale. 
Amphisbt'thi minima van intr>rm?dia Bale. 
Ampinsbetia oprrculata (Linnaeus). 
Amphisbctiu avia n.sp. 

Family PLUMULARIIDAE 

UaUcortiopsis elegant (Lamarck). 
v Amenella campanuUformix (Mulder & Trcbil- 
cock). 
Pyrnotheca mirabilix var. mirabilix (AilttUtn). 
Halopleris campantda var. campanula (Husk). 
Ptumularia filicaulis Kirchenpauer 

* Plumttlurta hyalina Bale. 

* Phttmduria attRusta Stechow. 

* Plumuhuia crater if or mix Stechow. 

* Plunmlaria wilsoni Bale. 
AgUtophvma plumosa Bale. 
Thccocurpt4s t xUvaricafus var. typiea (Busk). 

* Thccocarpux dharicatus var. hrt&gsi Bale. 
Haiieornaria tanxiroxtrix (Kirchenpauer) , 

Systematic Section 
Family CAMPANULARIIDAK 
CumpsifiuJaria ambiplica Mulder <Sr Trebilcoek, 
1914a; pi, 2, tigs ^ > 4. Shepherd & Watson, 
1970; 140. 
Paraeatix ambiplica Stechow, 1925: 209, fig. H. 
Record: Penguin I., on a red alga and sponge 
in crevices nn rough-water side of island, 
16-20 ft] deep. 

Material: Colonies infertile. Hydrothecae with 
7 fairly sharp teeth, the embayments between 
wider than the teeth. 

Remarks: Although occurring on the rough- 
water side of the islnnd, this small delicate 



species occupied a microhabital in sheltered 
crevices at a depth below turbulence due K> 
surge. 

This is the first record of 0. ambiplica from 
Tasmania. Other localities: Victoria; Champion 
Bay, W, Aust. 

(Jainnumiluria pulcratheca Mulder & Trebil 
cock. 1914a: 11, pi. 2, figs 1, 2. Blackburn. 
1942; 105. 

Paracalix pulcratheca (Mulder & Trebilcoek, 
1.91 4ft). Stechow, 1923a: 3 

Record: Satellite I. (no depth recorded), on 
red alga Defixea. 

Material: Colonics infertile. Hydrorhiza tubu- 
lar. Stems 0.H3-1.33 mm long. 0.06-0,09 mm 
diam.. perisarc thick, a spherule between stem 
and hydrotheca Hydrothecae long and tuhular. 
perisarc thickening distally, a distinct dia- 
phragm near base and a flexure almost iwo- 
thirds the distance up the hydrothecal wall from 
the base, Margin with 8-10 teeth. !3iam. at 
hydrotheca at margin 0.32-0.41 mm, depth to 
diaphragm (including teeth) 0,77-0.92 mm. 

Remarkv: The Tasmanian materia! compares 
well with the holotype of C, pulcratheca (in 
NMV), although the present specimens have 
fewer marginal teeth. 

This is the first record of C. pulcratheca from 
Tasmania. Other localities: Victoria; S. Ausl, 

Clytia sp. 

FIG. 1 
Record: Adventure Bay, 10 m deep on *l«m 
of Thccocarptis divaricatus var. typt'ea. 

Material: A few infertile stems. Stems of vari- 
able length, 0.70-1,90 mm, irregularly undu- 
lated, in some places smooth. Hydrothecae 
campanulate, expanding from base Eo margin, 
perisarc fairly thick, and a well defined dia- 
phragm with a thickening of the thecal wall 
below. Depth to diaphragm 0.30-0.40 mm. A 
small spherule between hydrolhecae and pedi- 
cel. Margin 0.20-0.32 mm diam., with 12 
bluntly pointed teeth, the embayments between 
slightly wider than the teeth. 
Remarks: The hydrotheca of this relatively 
small species corresponds in some respects with 
C. hemixphaerica (Linnaeus, 1767), but the 
stem lacks the typical proximal and distal an- 
nulations of this species, In the absence of 
gonosome it is not possible to further identify 
the material. 

Ortbopyxis caliculata fHincks, 1853). ftfite, 
1914b: 74, VI 11, fig. 1, pi. 12, fig. L; 



HYDROIDS O* BRUNY ISLAND, SOUTHERN TASMANIA 



J5« 



1924: 232. Hodgson. 1950: 7, ligs 14-16. 
Shepherd & Walson, 1970: 140. 

Ctitnpanularin adiculata Htntfc* 1853: 178, pi. 

*. fig. 5. 

tucopclUi calk-utata (Hincfcs) HirohitO. 1969; 

tig. ft 
Record: Penguin l.„ 15-20 m deep, on a red 
alga and on sponge in a crevice. 
Material: A lew fertile colonies, tlydrothecae 
very shallow ;ind expanding; hydrothecal pedi- 
cels spirally annotated, a few wilh smooth 
regions. Hydratuh wkh 24 tentacles. Gwoiheca 
smooth with thick pcrisaic, containing mature 
eonophorcs. 

Remarks: The specimens from sponge possess 
longer pedicels and have a thinner perisarc 
than those epiphytic on algae. 

O. colic alata was not abundant at Bruny 
Island, 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). 

Silfciilaria rosea Meycn, 1834: 204, p]. ?5. figs 
1-11. Millard. 1968: 259. 

Silicuhria bilahiam (Coughirey, 1875 i. kaiph, 
1957; 842. 

bacvpella campatwUlrtu voa Lendenreld. 1883, 
Dale, tfigg; 751. pi 13, figs 9-J5. 
SiltcuUtria campamduriu (von Lcndenfehl, 
IWO). Hodgson. 1950: 6, figs 12. !3. 

Record; Fluted Cape (no depth recorded) on 
the brown algae Seirococcus axillaris ,,nd Sryto- 
thalia dorycarpa. 

Material: Luxuriant fertile mate and female 
colonies on the algae. Developing gonophnres 
presenl. 

Remarks: The present material conforms to 
descriptions given by Ralph (1956. 1957) for 
Silicularia bilabiata forma subtropica (demon- 
strated by Millard (1968) to be a synonym of 
S. rosea Meyen), a form occuring only north 
of the Subtropical Convergence, and typical of 
the southern Australian coastline, including 
Tasmania. 

Family LAFOEIDAE 
HcbcUa ? furax Millard. 1957: 200, % 8; 
1964; 10, fi& 2B-D. Millard & Bouillon, 
1973: 59. 

FIG. 2 
Record: Penguin I., on stem of ThectKarpus 
divarieatuit var. typita. depth 20 m. 
Material: One small colony. Hydrorh'tza a 
Uroscly winding tube. Hydrothecae large, cam- 



panulate, pcris3rc delicate, smooth. Hydrolheca 
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, perisarc thick, strongly undulated to 
smooth, widening distolly to pass into base 
of hydrotheca below diaphragm. Immature 
hydrothecac truncated with a thin cap-like 
operculum, 

Dimensions (mm.); South 

Hruny 1. Africa 

Hydrolhecit — 
depth to diaphragm 0,80-0.92 0.74-0.S4 

citato, at margin 0.50-0.f.8 0.55-06^ 

dtam. at diaphragm 0.1 4-0. 18 0,22-0.2* 

Pedicel 
length from diaplnagm 20-0.29 0.52-0.72 
minimum diam. 0,1)5- 0.08 0.08-0. 10 

Remarks: Comparison of the Bruny I. material 
wilh paxatype microslides of flebefla fura\ 
Millard from False Bay, South Africa (pro- 
vided by Dr N. A. H. Millard) shows that 
although similar in shape to the South African 
specimens, the hydrothecae of the Tasniunian 
material are generally deeper and narrower at 
the diaphragm. lack the distinct thickening of 
the lower thecal wall, and have a more pro- 
nounced eversion of the margin than the South 
African specimens. However, in the absence 
of gonophorcs it is difficult to determine the 
specific status uf hydroids of simple morpho- 
logy, hence the presenl specimens arc pro- 
visionally assigned to Ih furax, 
This is a new record for Australia. 

Family HALECIID AE 

Halecfium Uelicatufum Coughtrcv, 1#7fia; 299; 

IS76b: 26. pi. 3, figs 4, 5. Ralph, 1958: 

334, %s 1 1, 12 (synonymy >. 

Hateciitm flexile Allman. 1883: U, p|. 5, fit. 2- 
Hodfiwta, 1950: 16. figs 25-27. 

Records: Adventure Bay, Penguin I., Satellite 
I., on the kelp Macrocystix pynfera. 2-7 in 
deep; Fluted Cape, 10 m deep, on Tbecorarpus 
divarictUiis var. typica (Busk). 
Material: Luxuriant fertile colonies. Stems to 
I cm long, simple und branched. Many hydro, 
phores with marginal replications; secondary 
hydropborcs arising trom the pedicel ot 
primary hydrophorcs. Colonies dioecious with 
mature gonophores ansing from proximal parb. 
of stem and on hydmrhiza. Distal parts of 
blastostyie cap-shaped |p both sexes. Colour, 
irophosomc yellow, gonophore bright orange. 
Remarks: Hodgson (1950) described ami 
figured H. flexile Allman (= H. deficatulum) 



160 



TEANETTE B. WATSON 




Fig, 1. Chtia sp, Hydrolheca. 

Tig 2, Hebella ?jurax Millard. Hydrothcca epizoic on J hecocarpus elivaricaius. 
Figs 3.4. Halecium sp. Fig. 3.— Whole colony. Fig. 4 — Distal part of colony enlarged. 
Figs 5,6. Halecium beanii (Johnston). Fig. 5.— Whole colony. Fig. 6.— Part of stem showing hydro- 
phores. 



from Eaglehawk Neck, but did not record the 
substrate. 

The colonics of H, delicttmlum in the pre- 
sent collection, while very abundant on Macro- 
cystis and other brown algae, were, however, 
strictly epizoic, always growing on the surface 
of the crustose bryozoan Membrhwpora mem- 
brinacea, a common epiphyte on the aging 
fronds of Macrocystis. 

Halecium sp. 

FIGS 3, 4 

Record: Penguin I., on a red alga in sheltered 
water, no depth recorded. 
Material; One infertile stem. Stem 3 mm high, 
lightly fascicled at the base, branching irregu- 
larly sympodial. Stem internodes of variable 
length, 0.30-0,40 mm, narrow, perisarc fairly 



thick, with one proximal annulation and a distal 
apophysis giving rise to the succeeding inlcr- 
node. Hydrophore fairly deep, expanding to 
margin, slightly asymmetrical, adcaulinc 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 0.01 mm 
below line of attachment of hydranth, usually 
a strong thickening of the inner wall immedi- 
ately below diaphragm, best seen in older 
hydrophorcs, often absent in younger terminal 
hydrophores. Punctae not visible; if present, 
obscured by hydranths. Pedicels of hydrophorcs 
of variable length; shorter pedicels usually 
annulatcd, longer pedicels smooth. Hydro- 
phores regenerated 1-3 times, regenerated pedi- 



HVDROIDS OF BRUNY ISLAND, SOUTHERN TASMANIA 



161 



eels often with a deep proximal constriction 
Hydramh short and stubby, wfrih approximately 
14 tentacles. 

Retmrkx; The single specimen from Hnmy 1. 
resembles H. tewllum Hinks, 1861, and Hale- 
dum sp. recorded from Pearson 1. (Watson 
1973. p. 167). However, H. tenellum is muno- 
siphonic tuid the dimensions given by Millard 
(1957, p. 193) and Ralph (1958, p.~34(l) for 
H. tenelhtm arc creator than iho.se of the pre- 
sent material. The specimen from Pearson J-, 
while similar >n habit, is monosiphonk. the 
margin of the hydrophore is more everted, and 
the overall dimensions are smaller, Without 
adequate fertile material it is not possible to 
make a decision on the specific status or rela- 
tionships of the specimen from Bruny I. 

Ifahciiim bcatiifi (Johnston, 1 838). Millard, 
1957' IKK; 1966; 464; ]96S; 256, fa, 
9A-F. Ralph. 1958: 3.12, fig. 10a, b, c-k. 

FIGS 5, 6 
Records: Adventure Bay (no depth recorded) 
on encrusting sponge on underside of the red 
alga Sotiilerofthycux auitrafiv: Penguin 1„ 15 m 
deep, on sponge in crevice. 

Material: Busby infertile colonics to 15 mm 
long. 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 terminul hydrophore. 
Stem internodes of variable length, 0.28-0.52 
mm, narrowest at node, 008-0.12 mm, widen- 
ing distally to 0. 13-0.23 mm to accommodate 
hydrophore. Nodes distinct, with a slightly 
oblique slope alternately right and left, occa- 
sionally straight, H spirophores alternate, shal- 
low, saucer-shaped, adnale to internode, diani. 
at margin 0.12-0.14 mm. Diaphragm distinct, 
0.02-004 below margin, tilted towards node, 
marked by a thickening of perisarc, p ring 
of punctac (frequently not well seen) above! 
Secondary hyttrophores given off on a short 
pedicel from diaphragm of primary hydro- 
phore. adcauJine wall convex, abcauline wall 
straight or very slightly bulged. Body of 
hydranth delicate, wilh approximately 14—1 6 
long filiform tentacles borne on a long 
peduncle, a deep constriction between 
peduncle and hypostome, 
Rrmarkx; Since the Bruny I. specimens con- 
form lo descriptions, and fall well wilhtn the 
range of dimensions given for U, beonii by 



Millard (1957) and Ralph (195S), the speci- 
mens, although infertile, are assigned to ibis 
species. 

An unusual microhabilat is the epizore 
growth on crustose sponge on the underside of 
the thick, plaie-like thallus of Sonderophycus. 

H. heanii is a cosmopolitan species, not 
previously recorded from Australian waters. It 
is rare at Bruny L 

Hakiium bruniemis a. sp. 

FJGS 7-1 5 
Type material and records: Holotype, NMV. 
C2494-microslidc; G2495-prcserve"d material, 
remainder of holotype cofony; Penguin f. t 20 m 
deep, on sponge and bryozoa in crevice. 
Description from hohtype; Etea stem 3 cm 
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 sympodia! growth. Stem inter- 
nodes of ultimate branches of fairly constant 
length, 0.48-0.64 mm, narrower proximally, 
widening distatly to 0.1 0-0.] 2 mm at node. 
Nodes oblique, well defined, sloping alternately 
left and right, often 1 or 2 annulalions above 
node. Pedicef of hydrophore given off distally 
from a well defined apophysis 0.04-0,14 mm 
long, at the same level as the node. Pedicel 
tubular. 0.13-0.20 mm long (node to dia- 
phragm), perisarc of younger parts smooth, in 
older regenerated parts heavily internally 
ridged, a deep fold just ahnvc apophysis giving 
pedicel an offset appearance. Hydrophore fairly 
deep, 0.06-O.Ofc mm margin to diaphragm, 
slightly expanding with an everted margin, 
0.15-0 18 mm in diam,, and distinct rim. Dia- 
phragm well defined, thin, with a ring of 
punctac just above and a pseudodiaphragm 
below, frequently only marked by a thickening 
of the adenine wall. Secondary and tertiary 
regenerations of the pedicel from the orifice of 
the preceding hydrophore common, each suc- 
ceeding pedicel uutally shorter than the last; in 
some instances regeneration is reduced to a 
mere replication of the hydrophore. Secondary 
branching of pedicels rare. Hydranth elongated, 
slender, with 10-16 long filiform tentacles. 
Female gonorheca very large, flattened, lenti- 
cular, but somewhat variable in shape, usually 
slightly longer than wide: greatest width G,9&- 
1.33 mm; length (excluding pedicel) 1.20-1,43 
mm, tapering proximally into a short pedicel 
arising at base of hydrophore, usually ol junc- 
tion with main stem. Perisarc very delicate, 



162 



JEANETTE E. WATSON 




Figs 7-15. 



HYDROTDS OF BRUNY ISLAND. SOUTHERN TASMANIA 



m 



distal extremity puckered, a small circular 
Otificc sealed by an operculum situated in the 
distal third of Ihc abcauline wall Male gono- 
theca small, sausage-shaped, 0,25 mm long, 
0.10 mm wide, arising from a short pedicel at 
base of hydrophore. Buth sexes arising only 
on younger, monosiphonic pans nf colony, the 
males more distal than females Colour, $tems 
straw coloured, female gonophores pink. 
Remarks: Hatecium bninienM\- is closely allied 
to //. lentiatlure Trcbilcock, 1928. in sym- 
podial habit, shape of female gonotheca and 
hydrorheca However, //. knilculure as known 
at present (Ralph 1958, p. 331) is n mono- 
siphonic species with smaller stems I less than 
I cm high ) , a considerably smaller female 
gonotheca, and a male gonotheca of somewhat 
different shape. 

The type material of H. hnmiensix shows 
growth stages of the female gonophorc from 
earliest development to maturity (Figs 12-15). 
Development of J he uonophorc begins with 
formation of a hook-shaped hlasiostyltr sur- 
rounding a central body (Fig. 12). Further 
growth and diilcrcntation into 6-41 large bodies, 
posstbly larvae (but material insufficiently well 
preserved for positive identification), then oc- 
curs, filling gonotheca (Figs 13. 14), A small 
circular aperture with slightly thickened rim 
then develops in the distal third of the ah- 
caul'mc wall through which the reproductive 
products escape (Fig. 15). The orifice of ihe 
now empty gonotheca then becomes rcsealed 
hv a very Ihrn operculum. 

Only one group of colonies of H. bnudensh 
wat lound These were cpuoic on sponge and 
encrusting bryozoa in a sheltered crevice. 

I lutecium hiteum n. sp. 

FIGS 16-lS 
type material mid Records: Holotype. NMV. 
G24%-microslide <K.OH cleared preparation); 
G2497-prcscrved material,, remainder of holo- 
type colony; paratype G24W rmcroslide; 
Penguin L. 15m deep on sponge and rock. 
Description front holotype and paratype: 
Colonies, to 2.5 cm high, growth arborescent, 
mam stem strongly fascicled, woody and stir? 



Branching more or less in one plane, the ulti- 
mate branches in any one part of the colony 
all directed anteriorly: in other parls they may 
face in other directions. Stein tnrentotlex nf 
variable length, 0.40-0.65 mm, expanding 
dismally, with 1, occasionally 2, extra nodes. 
Nodes distinct, oblique, parallel in each inter- 
uode. sloping alternately left and right in ad- 
jaccat uitcrnodcs; perisarc indistinctly internally 
ridged. Width at proximal node, measured 
parallel to node, 0.12-0,14 mm. Hydrophore 
seated on distal third of distal segment of inter- 
node, well below node, Hydrophore sessile, 
very tlat and shallow, wails thin and .UTOngly 
constricted just above diaphragm, abcauline 
wall more concave than adcauline. Margin or. 
cular, 0.14—0.16 mm in dianv. with outrolled 
rim. Depth from margin to puncjae, 0,015- 
0.02 mm; depth from margin to diaphragm. 
0.025-O.O35 mm. Diaphragm veiy strong, with 
a distinct riug of punctae above, and a wedge- 
shaped thickening oi the perisarc ol the inter- 
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, Hydnatth large with an annular 
hyposlomc and 25-28 tentacles. Colour, bright 
yellow, Gonotheca. absent. 

Remark*: Ntdectum htieum is superficially 
similar to //. corrugatissimum Trebilcock. 1V2S. 
(rom New Zealand, as il has strongly ridged 
intcrnodes and shallow hydrophore* character- 
istic of this species. In the latter species, bow- 
ever, the ridges of the stem are merely strung 
annular constrictions, not definite oblique 
nodes as in fi. furcunh Furehe/mure, Ihe hydm- 
phores, while shallow, axe somewhat deeper 
than those of the new species. Also, H. eorru- 
s}athsimum is a monosiphonic spectes, while 
H. tutemn has a strongly fascicled hahit. 

H. luteum displays several unusual morpho- 
logical features. One is Ihe presence of supple- 
mentary nodes, so well defined that the stems 
could almost he described as being alternately 
athccale and thecate; another feature is Ihc 
extraordinarily shallow hydrophore. which 
seems to offer negligible support lo ihe very 



Figs 7-15. Hatecium brtwiensis n.sp. He. 7.— Holotype colony, natural size. Fig *.— Distal part of 

colony, Shwtofc fasciculation oi stem and empty female fiOOOthevae. Figs 9, 10. Hydro- 

phurc-v enlarged, showing offset pedicel and teplications of the hydrophore Fig. II.— Male 
gonotbec*. Figs 12 15.— Development ot the female gonopbore. Fig. 12. — Parly stage 
of development showing booX-shaped bbstosrvie with developing central mass Fig. 13. — 
tatei stage ot development. Fig. 14.— Mature gonophore. Fig. 15.— Gonotheca after dis- 
charge of reproductive products. Note residual mass remaining at site of the circular orifice- 
through which contents have been discharged. 



164 



JEANETTE E. WATSON 





1 cm 





Kgs 16-18. Halevium luteum n.sp. Fig. 16, — Paralype colony, twice natural size. Fig. 17. — Distal part 

of branch. Fig. 18.- — Stem internodes with hydrophores enlarged. 
Fig. 19. Phylactotheca armata Stechow. Gonotheca with developing male gonophorc. 



bulky hydranth. The extreme thinning of ihc 
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- 
sarc 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. 
Ophiodixw fragHfr Blackburn, 1937: 365, fig. I. 

FIG 19 

Records: Penguin 1. and Adventure Bay: epi- 
zoic on solitary ascidians, bryozoa and sponge; 
epiphytic on crustose coralline algae and on 



holdfasts of Phyihsporn comoca, 10-22 m 
deep. 

Material: Luxuriant colonies, some fertile. 
St^ms of variable length, 4-15 mm, usually 
simple, occasionally branched. Colonies 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 lop, 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 blastastyle, female 
packed with mature ova. Hydnmths with a 
single row of large lenticular nematocysts 
(probably stenoteles) alternate with the ten- 
tacles surrounding the hypostome. These also 
occur in the nematocyst batteries in the capi- 



HVDROTDS OF BRLNV ISLAND, SOUTHERN TASMANIA 



165 



tulum of the retracted daciytozooids, and are 
scattered throughout the hydrocaulus in some 
stems No discharged neniatocysts were seen. 
Remarks; Blackburn ( 1937) described the 
gonophore of "Ophiodissa fragilis" (■*= P ar- 
mata) as being "subspherical, arising at the 
junction of stem and peduncles, as well as stem 
and hydrorhiza". Blackburn's type mieroslide 
of "O. fragilLv" t'NMV collection) shows three 
extremely delicate structures which appear to 
be either directly attached to, or enveloping 
the hydrocaulus. Although two of these contain 
a central mass which could possibly be a 
developing gonophore. they resemble neither 
in shape or structure 1he immature and mature 
gonophores of P. armaia as seen in the present 
material. They dp, in fact, closely resemble 
&gg capsules of certain minute gastropods. 

P, artnata shows a wide range of substrate. 
The epiphytic colonies, particularly those from 
the rough-water sites among Phylfaspora hold- 
fasts, were usually short and robust, with 
heavily ridged cauline perisarc. Eptzoic 
colonies, particularly those from more shel- 
tered situations under ledges in deeper water. 
were lax, branched, with a more delicate peri- 
sarc and fewer intranodal ridges. All stems, 
however, show a tendency towards thickening 
oE the perisarc and increase in cauline ridges 
with age. 

This is the most abundant occurrence of P. 
armaia so far recorded In Australian waters, 
and demonstrates a greater variability in stem 
characteristics than formerly known. 

Family SYNTHECHDAE 
Synlhectum patulum (Busk, 1X52). Hodgson, 
1950: IS, figs 32, 32. 
Sertutasia paiula Busk, 1852.* 390. 
Records: Satellite I., 14 m deep, tinder ledge; 
Simpsons Bay, 2 1 m deep, on scallop Equl- 
chlamys hifrvHSi Penguin l M 20 m deep, on 
sponge. 

Materiaf: A few infertile colonies. Stems to 
25 mm long, some stems immature Proximal 
internodes of sterns short, with I pair of oppo- 
site hydrothecae, followed by an Joternode 
with 2 pairs of opposite hydrothecae and 
ilislal hydrociadia; distal lnternodes with either 
I pair Or 2 pairs of hydrothecae. Hydrothecae 
on older stems with thick perisarc and replica- 
tions, of the margin. Colour, purple and white. 
Remarks: The present material conforms to 
descriptions of Synthec'mm patulum given by 
Bale (1914a, p. 5) and Hodgson (1950) How- 
ever, it is very difficult I Watsou 1973. p. 167) 



to distinguish between Synthecium etegans f. 
suhventrieosum and S. patulum on character) 
of the hydrothecae alone. The present material 
is thus provisionally assigned to S. pawtum. 
Further work may eventually prove that the 
two are conspecific. 

No morphological differences could be 
detected between stems growing in the environ- 
mental extremes of exposure to current (Simp- 
sons Bay), sheltered situations under ledges 
(Satellite I.) or exposure to surge I Penguin J.>. 

Family SERTLLARflDAE 

Salaefo farquhari (Bale, 1924). Ralph. 196!a: 

760. fig. 7, 

FIGS 20-22 

Thuiartu farquhari Bale. 192*1: 2^4, fig, 10 
TreWlcock, 1928: 19, pj, 8, fig. 4. 

Records: Penguin I., 15 m deep; Satellite I., 
3 m deep, on sponge under ledge. 
Materia}: Two infertile colonies. Stems mono- 
siphonic, to 18 mm long, simple and branched, 
the simple stems shortest. Stem Jnternodes 
tapering proximally and distally, with one pair 
of opposite hydrothecae adnate in front, widely 
separated behind. Branching regularly alternate 
from aii apophysis of the inter node 0.25-0.4 
mm long, usually 2 pairs of hydrothecae 
between branches. Branches given off at an 
upward angle with a distinct proximal genieu- 
lation at the apophvsis and a V-shaped distal 
joint. Some secondary branching, but where 
developed, these branches carry few hydro- 
thecae. 

Remarks: The specimens compare well with 
microslides of Thuiarfo farquhari Bale (NMV 
collection) and with the redescription of S, 
farquhari given by Ralph f 1961a) The present 
material does, 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 
rnternode. as well as the tendency towards 
thickening and loss of the stem lnternodes 
in older parts of the colonies, features recog- 
nized by Ralph as being more characteristic of 
Solatia bicalycuhi (Coughtrey, 1R76a) than 5. 
farquhari. Since the present material agrees in 
most respects with the latter species, particu- 
larly in the size of the colonics., it is assigned 
to S, farquhari. 

This is the first record of S, farquhari for 
Australia, and the first record of the species 
outside New Zealand watery where it occurs 
only south of 43°S, the same latitude z% 
Brnny I. 



i«6 



jFANrTTT T WATSON 



SirrcoUuM-ii elongata (Lamouroux, 18it>). 
Ralph, 196 la: 7*2. fig. 4. Waison, 1973: 
170. 
Set tula flu clotizuta Lamouroux, 1816: 189. pf 
5. Bale 1884: 75, pi, <S. fiflfi 7, 8, pi- 19. fig. 7, 
1915: 277. Hodgson. 1950: 23, figs 38, 30 
Record; Penguin I., 15 m Jeep, on rough-water 
side of island, on a red alga. 
M&rr'tai; Several fertile colonics. Stems short. 
2.5 cm. GonotJtecae long and narrow, horned 
processes very much elongated. 
Remarks: Hodgson's Oyster Bay material from 
storm-drifted weed is Ihe •long-stemmed" form 
of S. elongata. The Bruny £ material is the 
"short-stemmed" ocean form frequently asso- 
ciated with red algae. 

Scritilarolla robusta Coughtrcy, IS76h: 300, fig. 

2. Hodgson, 1950: 33, fig. 58. Ralph. 

1961a: 824, fig. 22a-d. Watson, 1973: 

171, fig. 21. 

F[GS 23, 24 
Records-: Satellite I.. 3 m deep, on sponge- 
under ledges, and 6 m deep on the red alga 
Sonderophyctts aastralis; D'Enlrecasteaux 
Channel, 1 1 m deep on dead seawhip Prim- 
noeita australasiae, and on old scallop shells. 
Equichlumys hijrons; Adventure Bay, 5 m deep 
on sponge under ledges: Fluted Oupe (no depth 
recorded) on stem of a brown alga. 
Material: Stems 3-4- mm long, the longer stems 
llexuoiJs. with long inlernodes, occasionally 
branched Shorter stems robust, with short 
iniefnodex. each stem type occurring in 
separate colonies. Hydrothccae of the "long 
iniernodc" form large with smooth, very faintly 
undulated walls* hyilrothecac of the "short 
internode" form distinctly smaller (see dimen- 
sions), with thinner perisarc. heavily ridged 
wirh 3-4 annulations and a Strong submarginal 
constriction of the thecal neck on the abcauline 
side, All material infertile except for one stem 
of the '"short intermule" form. 

Dimensions (mm) : tons short 

internode internode 
form form 

Length, abcauline wall 0.55 fl.fiO 0.33-0.45 

Length, free ndcaulinc wait 0.35-0 4? 0,28-0.38 

Remarks: Specimens of S. robust a from Bruny 
I. show a complete range of variability between 
the extremes of the long and short internode 
forms, and large and small hydrothecac. Some 
correlation appears to exist between stem type, 
environmental conditions <md habitat pre- 
ferences, the long flexuous stems growing epi- 



zoically on dead shell and other material in 
the lyEntrecasteaux Channel in situations of 
.good current How, those of intermediate stem 
length being from cryptic habitats beneath 
ledges and on the underside of Sonderoph\ctts, 
while the more robust stems occurred on the 
lower parts of brown algae in moderately tur- 
bulent open water. Dimensions given by Hodg- 
son (1950) for his material correspond to the 
"short internode" form, although the stems of 
his material were 15 mm long, and were epi- 
phytic, 

Except for the greater thickness of stem 
and very faini thecal undulations, the "long 
internode" form of the rtruny I materia! cor- 
responds very closely with Sertalarella simplex 
(Hutton, 1873) described from Pearson I- 
(Watson 1973). A specific distinction based on 
thickness of perisarc and faintness of undula- 
tions of the thecal wall as defined by Ralph 
( 1961a, p. 820) seems to be somewhat arti- 
ficial, Thus, if further material with smooth 
hydrothecae and thin perisarc is found, 3. 
wbitsta must be referred lo the synonymy of 
S. simplex. 

Synipkctoscyphus pyjjmneus (Bale, 18JU). 

Walson 1973: 176. 

Sertuhrella pv.unuwa Bale. IS81: 25. pi. 12, fitt- 
Sfj 1KS4: H>8, pi, 3, fig. 8. pi. 19, fig. 19. 
Hodgson. 1950: 36. figs 63. 64. 

Eectjrd; Penguin I.. 16 m deep on bryo/oa in 
sheltered situations. 

A'fiirrnV//* Sparse infertile colonies. Stems to 
7 mm long, simple, or with one branch. A 
delicate 3-flapped operculum visible in most 
hydrothecae. 

Remarks: The line of small dots rri the thecal 
wall diagnostic of 5. pyxmueas is obscured by 
the hydranths. However, Ihe material corres- 
ponds closely with specimens of S, pygmnens 
in the Bale collection (NMV) and for this 
reason is Assigned to this species. 

Sertularia acuta (Stechow, 1921). Millard, 195*: 
192, fig. 8. Shepherd A Watson. 1970: 140. 

Tridentata acuta Stcchow, 1921: 231. 
Sertularia favulo.w Bale, 1884: 91. pi. 4. fiji 
5, 6: 1913: 121 pi. 12. figs 7, 8; 1915: 272. 
Hodgson. 1950: 25. figs 43, 44. 

Record: Adventure Ray, 10-22 m deep, on red 
algae. 

Material: Luxuriant fertile colonics. Sietm to 
7 mm long. Cottothecae with 4—6 strong annu- 
lations. arising from stem internode below 
proximal hydrotheca. Gonophores mature. 



HYDK01DS OF BKUNY ISLAND. SOUTHERN TASMANIA 



H,T 



Remarks: Hodgson',*; infertile specimens came 
from storm-drifted Microcystis. S. acuta was 
r\o\ associated with this kelp at Bruny 1 

Serfularin macrocarpa Baie. 1&S4: ftO. pi. 5, 
fig. 2. pi. 19. fi&. 11: !9I4a: 14; 1915; 
277. Mulder & Trebilcock, 191 4ht 42. 
Hodgson, 1950; 27, rig. 47. Shepherd & 
Watson, 1970: 140. Watson, 1973; 177. 

Record: Penguin J., sheltered side, 10-22 rn 

deep, among algal holdfasts. 

S'iawriai Rare colonies, comprising a few 
stems each Stems to R cm long, infertile- 

Remarks: The internal suhmarginaJ loolh is not 
as well developed in these specimens as in (hose 
ffom the Australian mainland. 

This dark brown species wjlh distinctive 
white-tipped hydrocladia has previously been 
recorded among the hold I a si fauna of red algae 
at Pearson I. (Watson 19731. 

This is Ihe first record of S- macrocarpa from 
Tasmanian waters. 

Amphisbctia minima var. intermedia Bale. 

1915. Watson, 1973; 179, fig 29 

FIG. 25 

Sertutaria min ima Thompson T 1 879 : 1 04, pi . 
17. fig. 3. Bale, 1881: 21, 45, pi. 12 fig, 2; 
1884, 89, pi. 4. fifiS V t 10, pi. ft, fits 12, 13; 
Mulder & Trebilcock, 1 9 14b: 39, Hodgsnn, 
1950; 23, figs 41, 42. 

Records: Adventure Bay, 4-6 m deep, on iod 
alga Rhodymenia and on sponge under ledges; 
Penguin 1., 16 m deep, on red algae and in 
crevices in rough water; Satellite 1 , 1 m deep> 
on Laurencia. 

Material: Luxuriant fertile colonies, mainly on 
red algae, some on sponge. Stems to 5 mm 
long, internode length 0.30-0,38 mm; diam. at 
node 0.03-0.06 mnu HydrOlheca 0.19-0.28 mm 
long. Tubular nematothecae present in the base 
of proximal intrathecal chamber. 
Remarks: Although the hydrothecae are larger 
than those recognized as var. intermedia (Wal- 
son 1973, p, 181), the present material is 
referable to this variety on the ba\ts of the 
shape of the hydrothecae and the presence of 
the characteristic tubular nematothecae. How- 
cvei, several of the stems have rather robust 
hydrothecae and the wedge of perisarc hetween 
hydrocaulus and hydntfheca considered typical 
of (he var. pumiloides. Furthermore, one stern 
(Fig. 26) from sponge., shows distinct transi- 
tinnal features between the pumiloidcK and 
intermedia types. The stem of this specimen 
hits four hroad* robust, proximal hydrothecae 



of "pumiloides" type, and a distal rcgrowth of 
thiee hydrothecae of unmistakably "intermedia" 
type following stem breakage. The basal pair 
of "intermedia" hydrothecae bave well 
developed tubular nematothecae. Dimensions 
of the proximal and distal groups of hydro- 
thecae on this stem are given for comparison: 

Dimensions (mm) j ptter* 

pumiioides media 

type type 

Internode — 

lenpih 0.33-0. *5 30-0.34 

widrh at node 0.06 0.04 

Hydro Jheca — 

length 0,20 0.21 

width across ba^e 0.24 0.25 0.18-0.20 

widrh across margin 0.43-0-52 0.32 0.37 

The finding of iwo varieties of A. minima on 
one stem lends support to evidence (Watson 
1973) that these varieties are in reality eco« 
tnorphs, the development of which may be 
dependent upon the type of substrate available, 
or environmental conditions prevailing during 
growth. 

Hodgson (19501 did not differentiate be- 
tween the varieties of A. minima in his collec- 
tions; however, according lo his measurements, 
and the ahsence of nematothecae. as well as 
the material having come from Maerocystis 
and "drift" (brown?) algae, the material may 
be ascribed to the "var. pumiioides 1 '. 

Amphisbctta operculafa I Linnaeus. T 75K). 
Ralph, 1961a: 775. fig. 8. 
Scrtidaria opercuhta Linnaeus, 1758: 808. Bale, 
1884: 67. pi. *. fig. l, pi. 19. tm. 3. Hodgson, 
1950; 22, figs 36. 37. 
Record: Simpsons Bay. 11 m deep on stem of 
dead seawhtp Primimella austral iastac. 

Material: One small infertile colony of a few 
short stems. 

Remarks. Although Hodgson remarks that 
"this species is very abundant in the D'EoUe- 
casteaux Channel, being constantly taken in Ihe 
form of large tangled masses in scallop 
dredges v . only one colony was found in the 
present survey, This apparent rarity may be 
due to seasonal growth (Hodgson's material 
was collected in August) or to permanent 
changes in the environmental equilibrium 
caused by scallop dredging. The substrate of 
old shells preferred by A, operculata (pers. 
observ.) ts no longer available as the seafloei 
of the estuary has now heen invaded by enor- 
mous numbers of the New Zealand gastropod 
Maoricoiptts roseus. Since no colonies of /I, 
operculata were found associated with M 
roseus> it may be concluded that the smooth 



168 



JEANETTE E. WATSON 




Figs 20-27. 



HVDROTDS OF BRUNY ISLAND, SOUTHERN TASMAMIA 



169 



shell of this jtaunopod is unsuitable for settle- 
ment of larvae of A. opvrculata, 

Amphisbetia aria o. sp. 

FIGS 26, 27 
Type Material and Records: Holotype, NMV, 
G2499-micros!ide; G2500-preserved material, 
remainder of holotype colony: Adventure Bay: 
paratypes. G2501, G2502-rjrncrosltdes; Satellite 
I., all colonies on the brown alga Carpoglossum 
conflttens, 3 m deep, 

Description from holotype and paftttypex; 
RydrorhhM tubular, 0.09 mm diam., reticulate, 
very loosely wound on algal surface. Stems 
arising at slolonic junctions, simple, un- 
brartched, to 5 mm long, beginning with I or 2 
twists, then a V-shaped proximal pint, followed 
by first thecate intemode, PcnsaTC of stem and 
hydrotbecae Ihick. and very brittle. Stem inter- 
nodcx O.3&-0.46 mm long, with one pair of 
hydrotbecae, nodes slender. 0.06^0.03 mm 
wide, distal node collaT-shapcd. proximal node 
V-shapeJ and sockettcd into (he collar of pre- 
ceding infernodc; if node absent, it is replaced 
by a nanowirtjr of the intcrnodc. Hydro/necac 
opposite, on distal half of intcrnodc, tubular. 
narrowing to margin, actuate for one third of 
length, proximal adcauline wall more or less 
parallel to axis of intemode, 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 ba^e of hydro- 
theca, and a corresponding, hut not so deep 
inflexion (sometimes missing altogether) of the 
adcauline wall, opposite, but more distally 
situated, just behind margin. Width of inicr- 
node just below hydrotheca, 0.24-0.29 mm; 
length of fixed adcauline thecal wall (measured 
diagonally) 0. 15-0.1 y mm, length of free 
adcauline wall (to end of tooth) 0.12-0. J 8 mm; 
lenoih of abcauline wall (measured diagonally 
from bade to end of tooth) 0.26-0.31 mm. 
Margin horizontal, facing upwards, with two 
long, shaip, laterally placed teeth with a deep 
hortontal cmbayment between, connected lo 
the internode by a thick wedge of perisatc. 



Width across margin between teeth 0.07-0,09 
mm; width across paired hvdrothecae (outer- 
most teeth) 0.52-0.69 mm. A Plicate internal 
sheath often present within margin Gotto* 
tftetwe large, obovate, 1.15-1.33 mm long (in- 
cluding pedicel) expanding from base to sum- 
mil; max width 0.75-0.83 mm. tapering evenly 
into a narrow pedicel arising from the inf*a- 
ihecal chamber of hydrotheca on lower stem. 
Apenurc circular, 0.35 mm in diam., centrally 
situated at distal end, with a slightly raised 
collar, a ring of minute denticles within, and a 
Oat operculum. Gonorhecae identical in both 
saxes, only one borne on each stem: male and 
female gonophnre on same colony. Female 
gonophorc narrowly elliptical, not filling gono- 
thecal cavity, with 16-20 eggs; male gonopho:e 
of same shape and size as female, spermato- 
geaic mass surrounded by a thin blastostyle. 
Remarks: Amphisbetia avia is closely related 
lo Ihe Amphisbetia minima group in size, 
colour, habit, and preference for algal sub- 
strate, and is not easily distinguished from the 
varieties of A, minima in the field. However. 
the deep retroflexion of rhc abcauline wall, the 
horizontally directed distal part of the hydro- 
theca, and the Jong marginal teeth immediately 
distinguish A. avia from A. minima. 

A. avia was associated only with one species 
of alga. Curpogfossum confluens, growing in 
shallow water. 

FamiJy PLUMULARMIME 
Halicornopsis elegant tLamarck, 1816). Bale, 
1914a: 56: 7915; 303; Briggs. 1914: 296; 
1915: 309. Blackburn. 1942: 107. Hodg- 
son, 1950: 48, fig, 79. Watson, 1973. 195 
Pltwwtaria at^ans Lamarck, f816: 129. 
Record Adventure Bay. 15 m deep, epiliibic 
un vertical face. 

Material: One fertile colony. SterftX short. :o 
7 cm. Gonothecae borne prolifically on main 
stems and occasionally at base of branches on 
the apophysis of the hydrocfadium. Gona* 
shecae mature, irregularly ovate, flattened dis- 
tally or slightly flattened on one side, aperture 
closed by a thin membrane. Only female gono- 



Figs 20-22. iWwm farauhari (Bale), Fig. 20.— Part of stem. Fip. 21.— Stem mternodes, enlarged. Fig. 

22. — Single hydrotheca. anterior view. 
Figs 23.24. St'rwfan-Ua robusta Coughlrcy. Fip. 23.— "Long internoUc" farm with large hvdroThecae 

and -smooth thecal walls. Fig. 24— "Short intemode" form, showine smaller hydrolhecae 

with undulated wajls. 
Fig- "25. Amphisbetia minima var. intermedia Bale. Aberrant stem with both ^pumiio'nhs" (proxt- 

5«J P*rt of ylem) and "inter/nvtifa' (dista?) hvdrothecac. 
Hp 26.27. Amphisbetia avia n.sp. Holotype Fig. 26.— Stem with femnle gonophore. Fig. 27 —Stem 

mternodes, enlarged 



170 



JEANKITE L. WATSON 



phorcs present, spherical, half filling gnno- 
theeal cavity, packed with niaiure ova sur- 
rounded by a ihin granular blastostyle. 
Remarks: Only one small colony of H. clematis 
WAS found growing in a relatively exposed 
situation. 

MfiUire gonophorcs of It. elegrwy have not 
previously been described, 

Antcnnella cainpaiiiiliformis fMuldei Si I rehil- 
eock. 1909). Watson. 1973: 182. figs 43. 
44. 

Plumttfaria campanula or wis Mulder & Trebil- 

cack, 1909; 31. pi. K figs 6, 9. 10; 1910: 115. 
Heron!: Satellite I„ on t.enormandia marpjnatu 
and other red algae; no depth recorded. 
Material: Luxuriant infertile colonies. Erect 
stems to 8 mm high. Colour, trnphrwome 
pinkish-yellow, stolons dark brown. 
Remarks: This material conforms reasonably 
well with the description bt Mulder & Trebil- 
cock (1909). However, the hydrothecac are 
slightly more campanulate and delicate, with 
no thickening of the ahcauline thecal wall as in 
specimens described from Pearson I. (Watson 
1973). 

The hydroid was found on one side only of 
the -algal' fronds. 

This is the first record of A. compart uli/ormis 
from Tasmania. Other localities; Victoria; S. 
Aust. 

Pvcnotheca mirabilis (Allrnan, 1883) var, mira- 
bilfe Stechow, 1925; 241. Ralph, 1961b: 
50. tig. 7a, b. 

FIG, 28 

Pvenotlteca mirabilis (Allrnan, 1883). Hodgson, 

1950; 50, fiss 81. 82. 

Diplocluilas mintbilis Allrnan, 1883: 49, pt. 8, 

fig* 4_7. 

Kirr}vni> t m*tla mirabilis (All man. 1883). Bale, 

1894: 109; pi. 6. flgfi 4-7. Briggs, 1915; 308, 

Blackburn. 1942: 106. 
Records: Satellite I., 9 m deep; Adventure Bay, 
10-22 m deep, on red algae 
Material: Fertile stems to 5 cm high. Stems 
monosiphonic, arising singly from bydrorhiza. 
lower stems devoid of hydrocladia. Gonothecae 
large, adnate to lower 9tem. hydrorhiza, of 
jlgac. Perisarc very thick, strongly ridged with 
up to 9 ridges, more prominent on ahcauline 
side. Colour of gonolheca, deep red brown, 
lionophore.s, female. 

Remarks: It is difficult to distinguish between 
infertile stems of P. mirabilis and P. products 
Bale. 1KR1. Hxamrnation of fertile material of 
both species in the Bale collection (NMV). 



shows that the Bruny I. specimens correspond 
closely to a micros! ide of P. mirabilis from 
Buss Strait. Gonotbccac of P. producta col- 
lected at Port Jackson, in 1886, arc smaller 
<I,5<>-1.86 mm long. 0.75-0,84 mm wide), 
have a thinner perisarc, and arc only faintly 
undulating. These are similar lo Hodgson's 
( 1950) figure and dimensions of "P. mirabilis"- 
Ralph (1%la) noted the discrepancy between 
Hodgson's description, figures, and the actual 
dimensions of the gonotheca of P mirabilts 
and suggested that it may represent a new 
varietal form of P, prodttcta. It seems more 
likely, however, that the numbers of the two 
figures of the gonothecae of P. producta and 
P. mirabilis have been confused in Hodgson's 
paper. 

Halopteris campanula var. campanula (Busk. 

IK52). Ralph, 1961b: 47. Watson. t973: 

i»4. 

PlumuUtria cumpuru*la Rusk, 1852: 401. Bale, 
1884; 124, pi. 10, fig. 5; 1913; 133. Hodgson, 
1450: 40. 
Record: Satellite L, 14 m deep, under ledge. 
Material: One infertile colony. Stems to 2.5 cm 
long, sparingly branched, lax. Colour, yellow 
Remarks: H. campanula var. campanula is 
represented in this collection by only one sparse 
cpilithic colony, which was growing in reduced 
light in <\ cievice. 

Plumuluria hlicaulK Kirchenpaner. 1876: 28. 
p\. 5, fig. 6. Bale, I8S4: 134, pi. II. figs 
6, 7. pi. 19, figs 41, 42. Icloup, 1934; 41 
Hodgson. 1950; 42. fig. 72. Millard. 1958: 
209,^ fig. 13D f E. Shepherd & Watson. 
1970: 140 
Record: Adventure Bay, 10-22 m deep, on red 
algae. 

Material: Luvuriam infertile colonies. Usdro- 
rhiza pegged, forming a reticular network on 
the algal frond, Stems short, simple and pin* 
naLe on the one colony. Simple stems to 2 mm, 
pinnate stems io 4 mm high. 
Remarks: P. filicaulh is a common epiphyte on 
several species of red algae 1 Shepherd & 
Watson 1970). 

Plimtularia hyalina Bale. 1SK1: 41. pi. 15, fig, 9: 
1884: 14T.pl. 12. figs 4, 5 Ralph, 1961 h: 

41 

FIG. 29 
Record: Fluted Cnpe, 16 m deep, on a red alga 
Material: Abundant, sparingly fertile colonies. 
Seems io 3 mm long, stems and hydrocladia! 



HYDROIDS OF BRUNY ISLAND. SOUTHERN I ASMANIA 



171 







Fig_ 28. Pycrwthecu mirahUts vjir, mirabilis (Allman). Gonoiheca with heavily ridieed perisarc and 

female gonophore. 
Fig. 29, Plumularia hyalina Bale Stem with ripe female gonophore. 

Figs 30,31. Plumularia angu.sta Stechow. Fig. 30,— Part of hydrociadium. Fig. 31.— Gonoiheca 
Fig. 32. Piumuhma crateriformis Siechow Part of hydrociadium. 
Figs 33, 34. Ptumularia wilsoni Bale. Fig. 33.^Hart of hydrociadium. Fig. 34.— Gonoiheca showing 

downwardly directed growth habit. 



internodcs strongly ridged, hydrocladia with 
one terminal hydrotheca. Gonothecae large, 
top-shaped, greatest widlh distally, 0.7 mm 
wide. 0.9 mm long; gonophores mature, female 
only, completely filling gonothecal cavity. 
Colour, stems white to yellow; gonophores 
bright yellow. 

Remarks: This is the first record of mature 
P. hyalina in Australian waters. Ralph (1961b) 
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 gonothecae, it seems that an as yet undocu- 
mented range of geographic variants of P, 
hyalina may exist. 



Some stems of the material from Bruny I. 
have a well developed "stolonic plate" identical 
with that described for Plurmdaria epihtacteo- 
hsa Watson, from Pearson I. 

This is a new record for Tasmania. Other 
localities: Vic.; S. Aust.: New Zealand. 

Plumularia angusta Stechow, 1923b: 226. 
Blackburn, 1942: 108. 

FIGS 30, 31 
Plumularia setaceoides vars a, b, d, Mulder & 
Trebilcoek, 1910; 117, pi. 2* fig. 9, pi. 3, figs 

3, 6. 

Records: Penguin I., 15 m deep, on Macro- 
cystis holdfast and sponge; Adventure Bay, on 
Macrocystis holdfasts and the brown alga 
\4yriodesma quercijolium. 



IT 



JP.ANETTE U. WATSON 



Malarial: Abundant fertile stems to I cm long. 
Hydrothecae pitcher-shaped, but variable, some 
with a pronounced concavity of the adcauline 
wall and a constriction behind margin, others 
with a thickening of the abcaulinc wall. Colour, 
^traw-coloured, gonotbecae bright orange. 
Remark*. The hydrothecae arc very variable 
in shape even on the one hydrocladium. the 
thecal walls ranging from pitcher-shaped to 
almost straight, often closely approaching the 
shape ui the hydrotheca of P. sefaceoides. 

Pluinularia eratcrifnrmis Siechow, 1923b: 227. 
FIG 32 

PhtmulnHa sctaceoutes var. crater ijormis 

Mulder & Tiebltcoek, 1V10: 1 18, pi. 3. figs 8, 

Ra; 1VI5: 51. pi. 7. figs 3, 3a. 
Record: Adventure Buy, 4-6 m T on the brown 
alga Xlphophora gladiata. 
Material: Infertile stems to 1 cm long. 
Remarks on P. angusta and P, crateriformis: 
Sicchow (1923b, pp, 226, 227) raised Mulder 
k Trebilcock's varieties of P. setaceoides (the 
unnamed vars a, b. d. and var. t:ritterijormh\ 
to specific rank, rm distinction between Hie 
two species resting largely on the shape of the 
hydroiheca, a character of considerable varia- 
bility in this group and thus of doubtful diag- 
nostic value- The material from Bruny U 
although showing some intergnidaiion. can. 
however, be fairly readily assigned to one or 
other of these two species. 

This, together with the (act that the two 
species occur together in the one locality, 
although at different depths and on different 
olgal substrates (and one. P, angusia, was fer- 
tile, while P. crater if or mix was not) indicates 
thai these are valid, although closely related 
species radiating From ihe central P. setueeoid?$ 
stock. 

Neither P, tmgnsta and P. crateriformis have 
previously been recorded from Tasmania. They 
may have been confused with P. setaceotdts 
P. an gusto is. known from Victoria and S. Aust.; 
P. crateriformis from Victoria. 

PlumutariA wilswii Bale. 1926: 21. Ralph, 
1961b: 31. fig* 2, 3, 

FIGS 33, *4 

Ptumularia drlfrotufa Bale. ISftl: 28. pi 15. 

fig. 2; 1884: 137, pi 11, fig. !*, Midder & 

Tiebitcoek, IV 10: 1 15. pi. 2. fig. 2 
Records; Penguin I.; Adventure Bay: Satellite 
I,, 3 m deep, on sponge. 
Mit/eritU; Sparse fertile stems to 1 cm long. 
(ionothecae u>p-^hapcd, 1 or 2 arising on a 



very short pedicel from proximal part of stem, 
distal end directed downwards towards s»b- 
smiie- Petisare delicate, very faintly undulated. 
no operculum. Gonophorcs — mule, 
Remarks: The U'Ophosome of the Bruny I. 
specimens corresponds very closely with Bale's 
material of Pi wihoni from Griffiths Point. 
Victoria. The submarginal constriction behind 
the hydrothecae of the present material is 
rather variable, being more pronounced in some 
hydrothecae tiuin In othets even on the same 
hydrocladium. The hydrothecae with a shallow 
constriction closely approach those forms of 
P. setaceoides with more recumbent hydro- 
thecae. 

The gonothceac of the Bruny I, specimens 
arc identical with those of Mulder & Trebil- 
cock's (1910) microsHde from which they 
described the gonorheca of P. wihorii. 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 localities: 
Victoria; New Zealand. 

Agfaopbenia plumosa Bale, 1881' 25, pi- 14, 

fig. 6. 1884; 153, pi. 14. tig. 5. pi. 17. 

fig. 12; 1924: 257. Blackburn. 1942; 110. 

Hodgson. 1950; ?& fig 87. Shepherd & 

Watson, 1970: 140. 
Record. Fluted Cape, 20 m deep, on sponge. 
Material; Rare infertile colonics. Stems to 
1 cm long. 

Remark*. This is the robust form of A. pfo- 
PWW vvith short hydrocladial and stem inter* 
nodes. Hodgson notes that his material col- 
lected from Macroeywtis and drift had shorter 
stems (L.c. the short internode foi'iVU than those 
steins collected from the scagrass /.o.uera, 
which growls in sheltered waters. 

Thccocarpu* divaricatub var. typica iBusk, 
1852). Shepherd & Watson, 1970; 140. 
Aftlaophenta divuricutu Busk, 1852; 398. 
Records: Adventure Bay, 1-10 m deep, on 
horizontal faces among holdfasts of brown alga 
Phytlositora eomosa; 10-22 in deep, cptlithic 
in sheltered situations. 

Material: Stents infertile, to 10 cm in height, 
branched. Cawline nematothecac similar in 
shape to those of 7\ divaricatus var. cystifera, 
but much smaller, ilydrocladia close-set, hydro- 
thecae crowded on hydrocladium, no oblique 
intrunikU' sepia on ihecatc hydrocladial inter- 
lude. Colour, dark brown. 



JIYDROIDS OF BRUNY ISLAND. SOUTHERN TASMANIA 



m 



lltecocarpns divancaru* (Busk) var. briggtJ 
Bale, 1926: 22. fig. 5. Watson. 1973; 194. 
Record- Satellite I,, on the red alga Thamno- 
ilnm\tw dichototnum. No depth recorded 
Material; Straggling, irregularly branched, 
stems to 15 cm long. Colonies infertile, Colour, 
hlacfc. 

This is the first record of var. briggsi from 
Tasmanian waters. Other localities, Port 
Jackson, N.S.W. (Bale); Pearson L S. Aust 
(Watson). 

Remarks on the varieties oj T. divaricafus at 
Bruny Island The two varieties of J\ divari- 
cates recorded at Bruny I. are easily distinguish- 
able by the presence of an intranodal ridge in 
the hydrocladium of var. btiggsi, as well as in 
differences in habit and substrate preferences 
of each. 

The var, brfa&si was found only in sheltered 
rvefs near Satellite I T in the D'Entrccasteaux 
Channel, where it was a common epiphyte on 
Thamuoctonium, The more robust var rypka 
occurred on rock faces and as a common epi- 
phyte on PhyUoxpiua holdfasts in situations of 
moderate exposure to surge. 

Although showing affinities with van cysri- 
fera, the latter form docs not display the dis- 
tinctrve planar growth habit recorded for this 
variety (Watson 1973). nor the enlarged 
cauJinc nematothecac. In microstrudures it 
most closely resembles a fragment of Busk's 
type of A g'.aophenia divaricate from Bass Strait 
(NMV collection) and U thus recognized here 
as var typica. 

T, divaricates is a common and variable 
species of the southern Australian coast. 
Further study is necessary to elucidate the sys- 
tematic status and ecological relationships of 
the varieties of this species. 

Hatfcornaria longlrostris (Kirchenpauer, I872>. 

Bale. I8S4: |SJ. p (. 13, fig. 7, pj. !6. 

fig. 3, pi, 19, tie. 30. Hodgson. 1950: 51 

fig, 83. Watson, 1973: 197. 
Records; Penguin l. f 10-20 m deep, on Iheco- 
carptts divaricarus: Adventure Bay, 10-20 m 
deep, on red algae and epilithtc on vertical 
faces: Satellite I. 12m deep r on T. divaricates 
var. wtggth 

Material: Luxuriant colonies, stems 3-8 cm 
long, unhranched, infertile 
Remarks: The material from Bruny I. displays 
the same range in substrate as already noted for 
ff. hagirostris at Pearson I. (Watson 1973), 
Epilithic colonics from faces* exposed to surge 
have the longest jud most robust stems, while 



cpizoic colonies on T, divarkaius var. briggsi 
from both Penguin I. and Satellite J. H.e, from 
protected and relatively rough-water situations) 
had somewhat shorter, lax stems, givea 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- 
clatUa, the hydrocladia themselves having long 
interludes, and the. mesial ncmatoiheea extends 
well over the mouth of ihe hydrotheca. Rare 
epiphytic colonies on red algae at Adventure 
Bay had the shorted stems. There were no 
discernible differences in microstruetures be- 
tween these three ecomorphs. Briggs (1915) 
mentions that his specimens from Storm Bay 
were "4*6 mm in height (and) were found asso- 
ciated with Agiaophetu'u divaricate", Hodgson 
(1950) did not record the substrate of his 
material from Black man's Bay in the Derwenl 
Estuary. 

Discussion 

Ecology 

Campanulariidae were recorded from all 
depths, being epiphytic on red algae growing 
mostly in sheltered places. One exception, Sili- 
ctdaria rosea, was associated with the brown 
algae Scytothaita dorycarpa and Seirococcu, 
axillaris in situations of moderate water move- 
ment. 

The only representative of the Lafoeidac. 
Flebella furax, provisionally recorded for the 
first time in Australian waters, is a small form 
cpizoic on the stems of Thecacarpat divarica- 
tes var. hrigpxi Bale. The majority of species 
of the Lafoeidac known from Tasmaman and 
mainland Australian waters arc 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 Phylactorhectt armata 
and Halccium sp., ..II haleciid species in the 
collection are epizoic forms. Hulecium delicatu- 
lam Cotightrey [H. flexile of Hodgson (1950)) 
is one of i he most abundant bydroids at Bruny 
J„ the luxuriant orange-yellow colonies growing 
on the crusiose bryozoan Mcmbrinopora mem- 
brinacca epiphytic on old stipes of the large 
kelp Macmcystn pyrifem. The two species 
newly described, Hafeciutn hrunieruis and 
H. luteuni, as well as H beanii. are of cryptic 
habit, the two new species growing on sponge. 
bryozoa, and rock in crevices, white H. heanii 
occurs on the underside of the large plate-like 
alga Sondvrophycus australii, While fertile 
colonics of Phyiactotheca armata were recorded 
abundantly at all rough-water sites, rhey in fact 



174 



JEANETTE ft W A' I SON 



also occupied a relatively sheltered niierohahitat 
among the holdfasts of die brown alga rhyilo* 
xpora comosu. 

Remarkably lew species of Seuulariidae were 
recorded- Of greatest interest is ihe first record 
of Sixluctu farquhm (Rule) outside New Zea- 
land waters (sefe further discussion beluw). S. 
iarquitari was. however, rate at Bruny I. Sertn- 
lart'a acuta Stechow. while one ol the most 
abundant epiphytes in the collection, was asso- 
ciated only with red algae, and not with 
Mucracysiis as were Hodgson's specimens 
Hodgson (1950) also recorded very abundant 
colonies of AmpfvsbcJia operculum on old 
scallop shells from the DT.ntrecasleaux Chan- 
nel. Careful search in Ihe area of the DT.ntre- 
casteatix Channel covered in this survey pro- 
duced only one attenuated colony. Although 
A. npercuiata is known to display strong sea- 
sonal growth, some of the rootstock and pat is 
Of rhc colonics usually persist from one season 
ro another (per*, obscrv.l. The virtual absence 
of this species from a former habitat may be 
explained by permanent changes in the eco- 
system brought about by invasion of the gastro- 
pod Maoriculpux rosetts, following the collapse 
of the scallop dredging industry. Probably the 
shell of M. roxeux does not offer an attractive 
substrate to the larvae of A. operadata* 

Of particular interest is the occurrence in 
unc locality of Ihree species, Plumtdaria an- 
gUSlfo P cmterijormis and P wilxoru, all of 
which are known from Victoria. P. <m^uxto is 
also recorded from South Australia (Blackburn 
1942) and P. wifsonl ftom New Zealand 
I Ralph l%Jb), when: it is. however*, rare. 
These species arc closely related to one nnother 
and to PlunmUiriii setaceoides Bale, endemic to 
southern Australian and New Zealand waters, 
Irom which central stock they may have 
radiated. 

None of the larger Plumularians were of 
common occurrence. Haiiroinopxix dedans and 
Tltvt-ocarpits dhwicatux var. typ'tea were found 
in sheltered situations. The association of 7 
itivaHcatua var. hriggxi with a bluish coloured 
sponge investing the warty surface of the alga 
fhanmoclonium dichotoiimm has not pre- 
viously been observed. Jlalkornaria lon&'rostm 
was moderately abundant at all sampling sites - 
the three ecomoiphs of thi* specie* displaying 
an identical choice uf substrate with that 
reported from Pearson I. (Watson 197J). 

Zoogeography 

The Brum K material although collected 
from a restricted locality, yielded U new 



Tecords for Tasmania, including 3 new records 
for Australian waters. Seven of these specie* 
are already known Irom ihe Vieto/ian coast- 
line of lias* Strait, so would be expected to 
occur among the Tusmanian fauna. Theeocar- 
pux ttfpatitatus var. &ff{gpjf 4 now known from 
N.S.W. (Bale 1926). Tasmania and -South Aus- 
tralia (Watson 1973) has not yet been recorded 
from Victoria. 

Ol" Ihe 3 new records for Australia, flcde- 
chtm beanii is cosmopolitan, Hebella juntx is 
known only from South Africa, white Salutki 
fa/Qithtnl is a New Zealand .species recorded 
only from the South Island, where it does no:. 
however, occur north of 43 *S. The absence of 
S. farquhari from mainland Australian waters 
may thus be attributable to this southern distri- 
bution, as Bruny I. also lies close to 43 S. 

The profound influence ol the East Australia 
Current And the West Wind Drift in the dis- 
persal of species and the biological and zoo- 
geographic relationships of the trans-Tnsman 
hydroid fauoa have been discussed at length 
by Ralph (1961c). In this repaid, three species 
recorded from Bruny I. are of considerable 
interest. These are Ihe two halceiids newly 
described, and the occurrence of S. jurquhari. 
Hidccium bnmwndK bears a strong resemblance 
to H. knlk-idun\ a species known from the 
South Island of New Zealand and Cook Sir air. 
while H. htleum displays close aflmiiies with 
H. vors'tiMtussUmm* a rare species recorded 
from both North and South Islands. The strik- 
ing similarities in stem morphology between 
the members of these two pairs, a.s well as i.hc 
gross differences m habii <hoth Australian 
species mte polysiphonic, and the colonies arc 
larger*, strongly suggests active progress ol 
special ion from a common stock in addition 
to dispersal across the Tasman Sea. 



Acknowledgment* 

I am indebted to Professor: H. B. S- 
Womerslcy, Botany Department, University of 
Adelaide, for identification of algae; to my 
diving companions, S. A. Shepherd, R. Balduck 
and N. Coleman, tor assistance in the field 
and ro Mis* Rhyl I is Plant for some ot the 
drawings. I gratefully acknowledge financial 
support in Ihe form of a research grant from 
the Trustees of the C.S.T.R.O. Science and 
Industry Endowment Fund and a field research 
grant from the Royal Society of South Aus- 
tralia which partly met the costs of this study. 



HVDROIDS OF BRtTNY ISLAND, SOUTHERN TASMANIA 



175 



References 



Allman, G. J. (1872).— A Monogiaph or the 
Cymnobtasric or Tubiilarian Hydrnids, R(JJ, 
Site Lond. Part I. 1871; Purl 11 f !S72> 

Allnun. G. J. ilg&a i.— Report on Ihe Hydroida. 
Part J. Plumulariidae. Rpi. Set. Res. HM.S. 
Challenger. Zoology 7(2), 1-55, Plates 1-20. 

AUMAhr, G. J. (1888),— Kepoit on ihe Hydroida 
dredged by H.M.S. "Challenger" during the 
years 1873-1876. Part 1J. Ihe Tubularmae, 
Corymorphinae, Campanulannjje, Scrtu- 
larinae and Thalamorpha. Rpt. Vow Challen- 
ger 1873-1876, 23(70). 1-90. 

Bale, \V. M, (1881).— On the HyOroida or south- 
eastern Australia, witb descriptions of sup- 
posed new species, and notes, on the genus 
Axtaophvnia. J. ntfer, Soc. Vkx. 2(1), 15^47, 
Pis XH-XV. 

Bale, W. M. ( 1884 1.— "Catalogue of the Aus- 
tralian Hydroid Zoophytes." (Auit. Museum; 
Sydney.) 

Bale, W. M. 11888).— Un some new and rare 
hydroid.s in the Australian Museum collec- 
tion. Proc Linn. S'oc N.S.W. 3, 145-199. 
Pis XH-XXt. 

Bale, W, M. (1894).— Furfhcr notes on Aus- 
tralian Hydroida whfi descriptions oi some 
new ssecfe& Proc. R. Soc Met. (n.s.) 6, 93- 
117. i>ls Ul-VJ. 

Balk, W, M, I JV|3). — Further notes on Austra- 
lian hydroids. Jl. Proc. H. Sot'. I'fct. \t\.s/\ 
26, 114-147, Ph Xll XIII. 

Kaie, W. \f. (19l4 t O.— -KcpOil On the Hvdroida 
collected In the Great Australian Bight and 
other localities, Biol. Rrs. FJ,S vt fc.'ilde^vour*' 
2(1), 1-62, Pis I-Vri. 

Bale. W. M. ( 19 Ub).— Further notes on Aus- 
tralian hydroids. Ill, Proc: R. Soc. Vict. 
(n.s.) 27, 72-93, Pis XI-XH1. 

Baif, W\ Mi (1915). — Report on the Hydroida 
collected in the Great Auslialinn Bight and 
other localities. Part Til. Biol. Res F.t.S. 
"Endeavour" 3(5), 241-336. PH XLVI- 
XL VII. 

Baie. W. IVf. (1924).— Renort on some hydroid*. 
from the New Zealand coast with noles on 
New Zealand Hydroida generally, supple- 
menting Parquhat's list. Trans. N Z. Inst. 55. 
22<-268. 

Hale. W. M ( 1 926).— Further notes on Austra- 
lian hydroida. V. Proc. R Sot: Via. (n.s.) 
**> 13-23. 

BucKbUkn. M. (1937).— Report of the expedi^ 
rion oi the McCoy Society for held investiga- 
tion and research II. Cuelenterata, Proc. it. 
tine. Vki (rtAJ 49. 364-371 

Blackburn, M. (1942).— A systematic list of The 
Hydroida of South Australia with a summary 
of their distribution in other seas. Trans H 
Soc. 5. Aust. 66(1 ) ( 104-118. 

Bhiuos, E A. (1914),— Hydrozoa from one 
hundred lathoma, seven miles enst of Cane 
Pillar. Tasmania. Rec. Aust Mut. 10(10), 
285-30!. Pis 25. 26, 

BUM IL A. (I?|J), Notes on Tasmania* 
Hydrozoa. J. Proc. R, Soc. N.SW. 4H, 1\)2- 
318, Pis X, XI. 



Bvuk. G. 11852).— An account of the Polyzoa. 

and Scriularian zoophytes. In J. MacGillivrav, 

"The Narrative of the Voyage of the H,M.$ 

^Rattlesnake' l \ Appendix 1V. 385-402. 

(Bonne- London) 
Coughtrev, M (1875).— Nous on the New Zea- 
land Hydroideae. Tratif. Ptot: A'.Z. Inst. 7, 

281-293, PI. 20 
Couohirey, M. ( 1876a)— Critical Holes on Ihe 

New Zealand Hydroida. front. Proc. NZ 

iml S, 298-302. 
Co-lghtrev. M. i l£T76b>. — Critical notes on the 

New Zealand Hydroida. Ann. Mug. MoL ftteL 

scr, 4, 17. 22-32. PI. 3. 
Hincks, T. 1 1853) — Huther notes on Bnush 

Zoophytes, with description* oi' new species. 

Ann. Max. Nnt, Mist, ser 2, 2. 178-185, Pis 

5, 6. 
Hiroi inr>, £mt>efor of h.pan ( 1 y69 ) —Some 

hydroidN of the Amakusa Islands. Mai. Lob. 

Imperial Household, Tokxo 9, l->2. 
Hodgson, M. (1950),— A revision of the Tav 

mnnian Hydroida. Pap. Proc. R. Soc. 7 asm 

1949. 1 65. 
HciniN. F. W. (1873).— On Ihe New Zealand 

seilufarianv. Trans. Proc. vV.Z. fn.u 5, 25(*- 

259. 
1ohnston> G. (1R3«).— -A historv of Ihe British 

Zoophyl-s." 1st ed. (W. H. Uzars: Edin- 
burgh. I 

KtRCHtNPAUEa, G. H. ( 1 872 ) .— C'ber dk* 
Hydroidcnfamille Plumulandae. cinzelnr 
Gruppen derselben [inc\ ihre Fnichtehtiltcr. 1. 
Aglaaplwruu, Abh. naiurw- Vet, Hambun* 5. 
1-52, Pis IVIU. 

Lamahck, J. B. 1 1816). — "Hisioire naturalk dss 
Animaux sans Vertehres, II, Polypes." 1 

Lamourolix, J. V. F. <; 1816).— M His|aire dea 
Polypiers Coralligenes f*c\ib]e<, vulgniremeni 
nommes Zoophytes" (Caen) 

LnooK. F. (1934).— Tmis hydmpolvpes de \u 
bale de la Tabic. Ainqne, Anstrale. Bull 
Mhs, Hist. nat.Bvts. 10(19). 1-7. 

LcNDENPRi.o, R. von. fl883).— Oher CoeJen 
teratcn del Sud^ec. IV. Miuheduna Eueopelhi 
cumpanukiria nov gen. Zvhsvhr. f.iHss Zoot. 
LtipzwWA), 497-582. Pis 27-32. 

TjNNACUfi, C. (1758).— 'Svstema Namrae." lOlh 
ed. (Lipsjac.1 

MtVEN, K, J. F. (1834),— Beitrage ztir Zoolouie 
V'esammelt >ad* einvr Reise urn die Erde.^V 
Ucbei das l.eueh1en de^ Mtere« und Bcschrci- 
bun^ einigci Polypen und anderer niederer 
Thicrc. Nova Acta AteiL Caes. Leopold 
Caret, siippl. 16, 125-216. 

MniAfltv N. A. H. (1957) —The Hvdro^oa of 

False Bay, South Africa. Ann. S. Aft: Xfus. 

4% 173-243. 
Mrlartj, N. A H (1958). -Hydrozoa from Ihe 

co39t5 of Natal and Portuguese E-A*f Africa. 

Part r. Cahptotdtntea. Ann. S. 4h\ Mux. 

44, 165-226. 

Millard. N. A. H. (1964) .^The Hydrozoa of 
ihe <outh and west coasts of South Africa. 
Part II. The Tafoeidae, Syiithvciidae and Set- 
ruiariidae. Ann, S. Afrf. Mtts. 48. 1-56. 



176 



JEANETTE E. WATSON 



Millard, n, A. H. {1966).— The Hydrozoa of 
the south and west coasts of South Africa. 
Part III. The Gyinnoblastca and small 
families of the Calvptoblastea, Ann. S, Afr. 
Mas. 48, 427-487. 

Millard, N. A. H. (1968.).— South African 
hydroids from Dr. Th. Moi'tensen's Java- 
South Africa expedition. 1929-1930. Vidensk. 
Meddr. dansk nuttnh. t'oren. 131, 251-288. 

Millard, N. A. H., & BoutLLON, J. (1973).— 
Hydroids from the Seychelles (Coelentcrata). 
Artnls Mm. r. Aft. cent. ser. 8, 206, 1-106. 

Mulder, J. F. f & Trebilcock. R, E- (1909).— 
Notes on Victorian hydroida with descrip- 
tions of new species. Geclong Nat. 4, 2nd 
ser., 29-35, PL I. 

Mulder, J. F., & Trebilcock, R. E. (1910).— 
Notes on Victorian hydroida with descrip- 
tions of new species. Gedona Nat. 4 ( 2nd 
ser., 115-124, Pis 11, III. 

Mulder, J. F.. & Trebilcock, R. E. (19 14a). — 
Victorian hydroida with descriptions of new 
species. Part III. Geetonx Nat. 6(1), 6-15, 
Pis MIL 

Mulder. J. F., & Trebilcock, R. E. (1914b).— 
Victorian hvdroida with descriptions of new 
species Part IV. Gccionx Nat. 6(2), 38-47, 
Pis 1V-VL 

Mulder. J F n &. Trebilcock, R. E. (1915). — 
Victorian hydroida with description of new 
species. Part V. Geeiong Nat. 6(31, 51-59, 

pis vn-ix. 

Ralph, Patricia M. (1956). — Variation in OMm 
geniculate (Linnaeus, 1758) and Sifkularia 
hilabiata (Ooughtrcy, 1875) (Hydroida, F. 
Campanulariidae). Trans. R. Sac. N.Z. 
84(2), 279-296. 

Ralph. Patricia M. ( 1957).— New Zealand 
thecate hydroids. Part I. Campanulariidae and 
Carnpanulinidae. Trans. R. Sot. N.Z, 84(4). 
811-854. 

Ralph, Patricia M. ( 1958).— New Zealand 
thecate hydroids. Part 11. Families Lafccidae, 
Lineolariidac, Haleciidae and Syntheciidae. 
Trans. R. Soc, N.Z. 85(2), 301-356. 



Ralph, Patricia M . ( 1961a). — New Zealand 
Lhecate hydroids. Part III. Family Serin- 
lariidae. Trans. R. Soc. N.Z 88(4), 749- 
838. 
Ralph, Patricia M. (1961b).— New Zealand 
thecate hydroids, Part IV. Plurnularitdae. 
Trans. R. Sac. N.Z. (n,s.) 1(3), 19-74, 
Ralph, Patricia M. ( 1961c). -New Zealand 
thecate hydroids. Part V. The distribution of 
the New Zealand thecate hydroids. Trans. R. 
Soc. N.Z. f Zool. 1(7)> 103-1 1 L 
Shepherd, S. A., & Watson, Jeanette E. (1970).— 
The sublittoral ecology of West Island, South 
Australia. 2. The association between 
hydroids and algal substrate. Trans. R. Soc. 
S. Aust. 94, 139 146. 
Stkchow, E. (1921). — Uebcr Hydroiden der 
Dcutschen Tiefsee — Expedition nebst Bemcr- 
kungen liber cinigc andere Formen. Zool. 
Anz. 53, 223-236. 
Siechow, E. (1923a). — Neue Hydroiden der 
Deutschen Tiefsee — Expedition nebst Bcmer- 
kuttficn liber einige andere Formen. Zool, 
Anz, 56, 1-20. 
Stechow, E. (1923) — Zur Kenntnis der 
Hydroidenfauna des M ittelmeeres. Amerikas 
und anderer Gebiete. Zool. Jb. (Svst.) 47, 
29-270. 
Stechow, E. (1924).— Diagnosen neuer 
Hvdroiden aus Auslralien. ZooL Anz, 59, 
57-69, 
Stechow, E. (1925).— Hydroiden von West- und 
Sudwestaustrulien nach den Sammlungen von 
Prof. Or Michaelsen und Prof. Dr Hart- 
meyer. ZooL Jahrb. Abt. f. Syst. 50, 191-269. 
Thompson, D'A. W. (1879). — On some new and 
rare Hydroid Zoophytes (Sertulariidae and 
Thuiariidae) from Australia and New 
Zealand. Ann. Mag. Nat, Hist. (s*r. 5)» 3, 
97-114, Pis 16-19. 
Trebllcock. R. E. (1928).— Notes on New Zea- 
land Hydroida. Proc. R. Soc. Vict, (n.s.) 
41(1), 1-31, Pis 1-7. 
Watson, Jeanette E. (1973). — Pearson Island 
Expedition 1969.— 9 Hydroids Trans. R. Soc. 
S. Ausl 97(3), 153-200. 



SHOOT AND FOLIAGE PRODUCTION OF FIVE SHRUB SPECIES OF 
ACACIA AND HAKEA IN A DRY SCLEROPHYLL FOREST 

BY J. R. MACONOCHIE* 



Summary 

MACONOCHIE, J. R. (1975) .-Shoot and foliage production of five shrub species of Acacia and 
Hakea in a dry sclerophyll forest. Trans. R. Soc. S. Aust. 99(4), 177-181, 30 November, 1975. 
An examination has shown that shoot growth of Acacia niyrtifolia, A. pycnantha, Hakea rostrata, 
H. rugosa and H. ulicina is distinctly seasonal. Growth commenced in spring, finished by mid- 
summer and was preceded by flowering. 

The maximum rates of loss of foliage occurred towards the cessation of active shoot growth and 
both mature and juvenile foliage was lost. Measurements of size of shoots of the three species of 
Hakea over a series of years suggested that the available soil moisture during the growth period was 
the controlling factor for shoot size. It is further suggested that the growth habit of these three 
species is reflected in the pattern and size of new shoots along a parent shoot. H. ulicina, the species 
which showed a tendency toward apical dominance is an erect, several stemmed shrub, whereas H. 
ro strata and H. rugosa are rounded spreading shrubs. 



SHOOT AND FOLIAGE PRODUCTION OF FIVE SHRUB SPECIES OF ACACIA 
AND HAKE A IN A DRY SCLEROFHYLL FOREST 

by J. R. Maconochie* 

Summary 

Maoonochh:-, J. R, (1975). — Shoot and foliage production of five shrub species of Acacia and 
Hakea in a dry sclcrophvll forest. Trans. R. Sac. 8, Attst, 99(4), 177-181, 30 November, 

J 975. 

An examination has shown thai shoot growth of Acacia myrtifolia, A. pyenantha, Hakea 
rostratu, H. rugosa and //. ulicina is distinctly seasonal, Growth commenced in spring, finished 
by mid-suromer and was preceded by flowering. 

The maximum rates of Joss of foliage occurred towards the cessation of active shoot 
growth and both mature and juvenile foliage was lost Measurements of size of shoots of the 
three species of. Hakea over a series of years suggested that the available soil moisture during 
the growth period was the controlling factor for shoot size, It is further suggested that the 
growth habit of these three species is reflected in the pattern and size of new shoots along 
a parent shoot. /?. ulicina, the species which showed a tendency toward apical dominance is 
tin erect, several stemmed shrub, whereas //. rostwta and H. rugosa are rounded spreading 
shrubs. 



Introduction 

This study on shoot production of the five 
temperate shrubs. A cacia myrtifolia (Sm. ) 
Willd., A. pyenantha Beruh., Hakea rostraia 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 1 970, 
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 Ml Lofty 
Ranges, South Australia from April 1965 until 
January (967. The occurrence and size of new 
shoots produced on about 120 tagged shoots 
of each species of Hakea and A, pyenantha^ 
and of 25 tagged shoots on two bushes of A. 
myrtifolia, were recorded monthly. The tech- 
nique has been 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 Austra- 
lian Bureau of Meteorology, 

Results 

Qualitative Aspects 

The percentage cumulative gains and losses 
of foliage for A. pyenantha and A. myrtifolia 
are presented in Fig. 1, .4. myrtifolia did not 
produce any new shoots in the first year of the 
study but the burst of growth during the 
second year was in phase with that of A- 
pyauintha. The "stepped" effect displayed by 
A. pycnaiuha was also produced by H. 
rostratu, H. rugosu and //. ulicina. New shoot 
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; soil moisture values were 
taken from Martin & Specht (1962). 

Shoot growth reached a peak during Octo- 
ber when daily temperatures were increasing 
and soil moisture was available (Figs 2 and 
3). In both 1965 and 1966, flowering, which 
commenced in early August and rinshed by the 



* Animal Industry and AgricuHure Branch, Arid Zone Research Institute. Dept. of the Northern Terri- 
tory, Alice Springs, N.T. 5750. 



178 



L R. MACONOCHIE 




Fig. 1. The percentage cumulative leaf gain and loss of A. vycnantha (□, total new positions; A 
leaves; Q, scars), and A T my rti folia (■, new positions; A, leaves; •, scars). 



end of September, preceded shoot growth. 
Immature flower buds on the Acacia species 
were observed to 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 season 
may be predetermined by conditions at the 
time of bud development. 

Although the A. my rti folia bushes did not 
produce new shoots in 1965, flowering did 
take place. 

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, nticina and H, roslrata 
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 arc 
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 
each species and for the years 1965 to 1971 
inclusive are presented in Table h Analysis 
of these data showed that for some years 



SHOOT AND FOLIAGE PRODUCTION IN ACACIA AND HAKEA 



U? 




4 



ML 




i * 9 C 



Fig. 



The relative rates of gain (block) and loss 
of foliage of (a) A. myrtijal'to, (b) A. 
pyrnarttha, (c) i/. ulicina. (d) //. rostrata. 
and (e) //. rugasa, for the years 1965 and 
1966. 



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 set of correlation coefficients (r) and 
regression equations were computed between 
total rainfall (x) for the period September 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. 
H. M//Wna— axillary shoots 

r^ + 0.74 0.10>P>.05 
y - = 5.0 4- 0.014x 



//. rostrata — terminal and axillary shoots 

r=-f-0.52 P>0.10 

y = 5.1 + 0.008* 
H. rtigosa — terminal and axillary shoots 

r= + 0.83 .05>P>.G2 

y « 4.1 + 0.029x 

H, rugosa was the only species showing an 
acceptably significant correlation between 
mean shoot size and rainfall during the grow- 
ing season. 

Calculation of the correlation coefficient 
between the mean shoot size and rainfall 
period September-November of the preceding 
year gave the following: H. ulicirtu — 0.65, H, 
rostram — 0.07, and H. rugosa — 0.55. 

Analysis of the data for 1966 on the rela- 
tive positions of shoots showed that there were 
no significant differences in shoot sizes 
between pairs of positions distal to the apex 
(Table 2) except for H. ulcina. In this species 
there was a significant difference (P < 0.05 1 
between the mean of the terminal and that of 
the first axillary position. Further analysis 
showed significant differences (P < 0.05) 
between terminal and axillary positions two 
and three. It would appear, therefore, that H. 
idkina is the only one of these three specie* 
which has a tendency towards apical 
dominance. 

Discussion 

Cambagc (1918, 1927) measured the height 
growth of A. pyenantha at Sydney Botanic 
Gardens and his study showed the normal 
pattern of rapid growth during the juvenile 
stage followed by a decreasing rate as the 
plant matured. The plants studied at Para 
Wirra were mature and the rates of growth as 
retlected in both Figs 1 and 2 were of a 
"steady state" nature, as occurs at tbc plateau 
of the sigmoid growth curve. 

Martin & Specht (1962) measured the 
moisture relationships in a dry sclerophyll 
forest, of which these species arc shrub com- 
ponents. Their studies showed that the more 
mesie community of this vegetation type had 
a higher index of evapotranspi ration and could 
be subjected to a drought period during the 
mid-summer. The three species of Hakea 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 




Fig 3 Climatic data of Parra Wirra study site; soil moisture ex Martin & Speehl (1962), other data 
means for 1965-1970. , soil moisture; O, mean temperature; 9 rainfall; Q, poten- 
tial evapotranspi ration (P/H 0.75); , florescence and duration of active growth. 

TABLE 1 

Mean number of leaves per shoot for seasons 1965 to 1971 



Species 


Shoot 
Posn. 


1965 


1966 


1967 


Year 
1968 


1969 


1970 


1971 


Hakea ulicina 


Total 

Terminal 

Axillary 


7.4* 

9.6 

5.9t 


8.1* 
10.2 
7.1' 


8.2 
9.4* 
4.9 % 


8.7 
9.4 

8.1* 


9.2 
10,9 
6.7t 


9.4 

10.6 
8.9 


9,8 

11.2 
8.4 


Hakea rostraia 


Total 
Terminal 

Axillary 


Mf 

6.1 
6.5$ 


5.8t 
6.1 

5.7 


5,2 

5.7 
5.2 


5.4$ 
5.4$ 
5.4$ 


6,5 

7.0 

6.4 


6,7$ 
6.5$ 
6.8* 


9.0 
10.6 

7.5 


Hakea ru^osa 


Total 

Terminal 
Axillary 


1.1% 
7.3* 
6.9$ 


9.1$ 

9.4 
9.0^ 


4.6$ 
8.1 
4.4 % 


8.7 

8.8 
8.8 


8.4r 

8.8 

8.3* 


9.2$ 
9.5$ 
8.9$ 


13.7 
15.9 

12.7 



Significant differences between mean and mean of succeeding year are indicated. 
* P < 0.05; fP< 0.01; t P < -001. 

TABLE 2 

Mean number of leaves per shoot for terminal and axillary positions distal from shoot tip for year 1966 



Relative 
Position 

H. ulicina 



Term. 



AX 



AX. 



AX ;! 



AX 4 



AX r , 



109 



7.8 



7.5 



?.: 



7.6 



7.4 



AX< ; 
6.6 



AX T 



AX* 



7.6 



5.6 



H. rosrraia 



6.2 



6.0 



5.3 



5,6 



5.3 



5.3 



H. rugosa 



9.3 



8,6 



9.6 



10.2 



10. 



8.2 



::; Indicates significant difference (.05 > P > .01) between mean and that of succeeding position, 



SHOOT AND FOLIAGE PRODUCTION (N ACACIA AND HAKE A 



181 



rainfall) during the growing period of the 
previous year suggests ibat shoot size in the 
following year is not necessarily determined 
at the bud 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 
{ 1 f J 65 ) . who recorded that the shoot growth 
commenced in December when soil moisture 
was decreasing Specht (1957) showed that 
drought conditions occurred in both December 
and January. 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 A> sowdeniu /i. Uxukifu and 
A, tnunayarWt and possihle non-seasonal and 
seasonal shoot growth responses on A. aneura 
and A. kempeana, Weiherell (1966) recorded 
Mushes of growth on A. harpophylla m Queens- 
land during spring and summer, and during 
one period of early winter. A. pycrtanthtt, by 



contrast, actively grew only during the spring 
period at the study site, and although shoot 
growth on A. myrti folia was only recorded 
during the second year, it appears that from 
this slender evidence that the shoot growl h 
follows a seasonal pattern also. 

Both H. rout ram and H. ru^osa have a more 
spreading bushy habit than H. ulkhia which 
has a tendency to be more erect. The habit 
of these plants is reflected in the sizes of new 
shoots on a parent shoot. Both H. roxtmra and 
H. rugosa did not produce significantly smaller 
3xiilary shoots in comparison to the. terminal, 
but the terminal shoots of //. 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. 

A cknowledgments 

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 



Camdace, R- H. (1918).— The vertical growth of 
Irees. I. /. Proc R. Soc N.S.W. 52, 377-384. 

Cambagh. R. H. (1927).— The vertical growth of 
trees. IJ. Proc, /?. Soc. N.SW. 61. 279-2K4. 



Grovhs, R. H. < 1965). — Growth of Heath Vege- 
tation, II. The seasonal growth of a heath on 
ground-water podzol at Wilson's Promontory, 
Victoria. Aust. A Bot. XX 281-289. 

MicONOCHfE, J. R., & Lange, R. T. (1970), — 
Canopy dynamics of trees and shrubs with 
particular reference to the arid- zone topfecd 
species. Trans. R- Soc. S. Ai4st 94, 243-244. 

Maconochiii, J,. R. (1973). — Leaf and shoot 
growth on Ac arid kempcana T\ Muell. and 
selected other arid-zone species, Trnpiuil 
Grasslands 7(1), 49-55. 



Martin, Helerte A. ? & Specht, R. L. (1962).— 
Are mesic communities less drought-resistant? 
A study on moisture relationships in dry 
sclerophyll forest at Inglewood, South Aus- 
tralia. Aim. J. Bot. 10(2). 106-118. 

Spechi. U. L.. & Rayson, Patricia. (1957).— DaTk 
Island Heath (Ninety-Mile Plain, South Aus- 
tralia). T. Definition of the ecosystem. Auss 
J. Bot. 5, 52-85, 

Swht, R. L. (1957).— Dark Island Henlh 
(Ninety-Mite Plain, South Australia"). V. The 
watei relationships in heath vegetation and 
pastures on the Makin Sand. Aitst. /. Bot. 5, 
151-17^. 

Wr.TirEHAU., A. J. ( 1966).— Leaf growth of Briga- 
low (Acacia harpophylla) suckers in relation 
to seasonal conditions. QUI J, (fttrip, /iw/w. 
Set, 23. 453-456. 



SHORELINE SHINGLE TERRACES AND PREHISTORIC FILLINGS OF 

LAKE EYRE 

BY J. A. DULHUNTY* 



Summary 

DULHUNTY, J., A. (1975). -Shoreline shingle terraces and prehistoric filling of Lake Eyre. 
Trans. R. Soc. S.Aust 99(4), 183-188, 30 November, 1975. 

Investigations were carried out of (i) development of contemporary wave-built shingle terraces 
during the 1974 record historic filling of Lake Eyre, and (ii) old shoreline shingle terraces built by 
prehistoric fillings of the Recent salina to levels above that of 1974. 

Evidence indicates at least three prehistoric fillings to levels of 280, 160 and 70 cm above that of 
the 1974 filling. No precise evidence of the ages of the prehistoric fillings was obtained, but 
estimates inferred from field studies suggested dates of the order of 3,000, 1,500 and 500 years 
before present. The "3,000" year old filling was probably the deepest during the Recent salina 
episode of the history of Lake Eyre. The possibility of more permanent, deeper and widespread pre- 
salina filling of the lake, and late Recent subsidence of its southwestern areas, was also inferred 
from field observations. 



SHORELINE SHINGLE TERRACES AND PREHISTORIC FILLINGS OF 

LAKE EYRE 

by J A. Dulhunty* 

Summary 

Dclhuntv, J. A. (1975). — Shoreline shingle terraces and prehistoric filling of Lake Eyre. 
Trans. R. Soc S, Ausl, 99(4'),, 183-188, 30 November, 1975 

Investigations were carried out of (i) development of contemporary wave-built shingle 
terraces during the 1974 record historic filling of L<>kc. Byre, and (ii) old shoreline shingle 
terraces built by prehistoric fillings of the Recent salina to levels above that of 1974. 

Evidence indicates at least three prehistoric fillings to levels of 2S0, 160 and 7t> cm 
above that of the 1974 filling. "No precise evidence of the ages of the prehistoric iillings was 
obtained, but estimates inferred from field studies suggested dates of the order of 3,000, 1,500 
and 500 years before present. The "3,000"' year old filling was probably the deepest during 
the Recent salina episode of the history of Lake Eyre. The possibility of more permanent, 
deeper and widespread pre-salina filling of the lake, and late Recent subsidence of its south- 
western areas, was also inferred from field observations. 



Introduction 

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 internal 
drainage system (Bonython 1955; Mason 
1 Q 55; Wopfncr & 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 
occasions 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 salt crusts 
(Oulhunty 1974; Bonytlum 1956). 

When the lake contains water, its shores arc 
actively eroded by wave action, producing 
well-defined shorelines with shelving strands 
or low clitfs 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 j.s 
10 m above the lake bed. 

Around Babbage and Hunt Peninsulas and 
in Belt Bay ( Fig. 1 A) , shores are beinij 
eroded in hard dolomite, silcrete and ferru- 
giniscd siltstonc (Williams 1973 1 ) Wave 
action, during periods of filling, has produced 
quantities of pebbles or shingle consisting 
largely of slightly flattened and rounded 
pebbles from 2-10 cm in diameter. Single 
strands or beaches have developed, and break- 
ing waves have washed up across the beaches 
long parallel deposits of shingle. These 
deposits, formed during the present salina 
episode of the lake's geological history, as dis- 
cussed later, are Recent to contemporary 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 were observed and recorded 
by Williams (1973 1 , 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. 

1 Williams, A. F. (1973).— Explanatory Notes for the LAKE EYRE 1:250,000 Sheet S. Aust. Dept. 

Mines. R.B 72/93 (unpubl.j. 



1X4 



J. A. DULHUNTY 



include the Recent shoreline shingle terraces 
desert bed in this paper, a nd other gra vel 
deposits, possibly at higher Levels, and more 
remote from the present shoreline, which may 
be older and related to prc-Recent history of 
the Lake. Williams fpcrs. coram- J 975) also 
noted shoreline shingle terraces 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 examined 
more remote terrat.es up to 20 km from %he 
present lake shore 

The tilling &l 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 to lake levels and wave action. 
This was undertaken to gam a better under- 
standing of the nature and origin of the old 
shoreline Jerraces and enable interpretation of 
their significance regarding the maximum 
deplh to which the lake had been filled during 
its role as the salina of the present internal 
drainage sysfcm* and the time since it was 
filled tu. or above, the level of the 1074 filling- 
Such investigations and results, recorded in 
this paper, are confined to the significance only 
of terraces along present shorelines, arid do 
not represent a cornprehensvic study of all 
shingle deposits and their regional distribution 
around l-ikc Eyre, 

Investigation of shingle terraces 

Contemporary mem\ Joke iVvW 

During the killer half of 1?>74* water level 
around the shores of the lake provided a hori- 
zontal datum, at any one lime within certain 
limits of variation, which was used extensively 
;ii determining levels of shoreline shingle ter- 
races tn relation to each other and to mean 
lake level. Short term variations in wafer level 
were caused by wind tides, currents and 
possibly other factors. Wind tides were most 
pronounced on windward shores where water 
level rose and fell, within a maximum range of 
about 45 cm, very quickly with rise and fall 
of wind velocity and wave height. When wind 
dropped, wave action usually ceased and water 
♦evel fell lo "nnrmar wllhfn 5 to 6 hours. 
"Normal" watci* level or the level free of wind 
effects during periods of calm, varied as a 
result of other factors within a range of about 
20 cm. Tn view of this n was assumed, for 
the purpose of Ihe present investigation, that 



water level at any point along the .shore* 
measured after a calm period 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 reached its maximum during May 
It rose by only about 20 cm during ApTil. 
remaining steady during May and fell by 
about 20 cm during June and July Therefore, 
rhe shingle terraces studied in August had been 
built over a period of four months during 
which mean lake level rose and fell through 
only about W 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 lei races 
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 mean lake level 
period, and finally remained stranded when 
water fell, providing evidence of the highest 
level reached, 

The 1974 contemporary shingle terraces 
varied in width up to 500 cm. and in depth 
or thickness up to 151) cm above pre-exislmg 
beach material. The height of their lops above 
maximum mean lake level <Mt<uned during 
their development, reached a general maxi- 
mum of ^0 em f=t 5 cm) on exposed shores. 
On sheltered shores, terrace lops were lower. 
All the terraces formed at any one time, 3t 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 level of 
the tops of ihe 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 
lake levd at which they had formed, There- 
fore, it was concluded that the maximum mean 
lake level of any past filling would be very 
clove to 90 cm below the general level of the 
original tops of the highest shingle terraces 
produced dining that filling. 

Greater depths of water during prehistoric 
fillings are not likely to hnve produced waves 



SHINGLE TERRACES AND FILLINGS OF LAKH EYRE 




UCALfTV BUtft SOUTH.V^- -^lOM <rf j AKE EYRf tfCRlH 



ggPgj PL^fi TsH '.3LE fUWACESmWILLCW 5A 



prehistorc iAflAce 

»-i'.-E:. I 
dWUB MIUCLE 



t TERGAL t 



LAKE 

COO' Tfi fcF 




"I'lrii"" i'fi 

* f-Ou" rff &p 

'5W&Y 5M1WGLE 
LQMITE _| J 1 - f '■'fr 

OflY.LAKE GEO BHffiEUNE 7hVtL-'-~- " _"_"_.'" |^>£j 
I - — !--■—! 1 



D* 



SECTION F- ri£.E?CftrjS10NI ACWOSH SHINQts TEKKACES 




Fig. L Wave-built shoreline shin 

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 "to 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 lake 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 to 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 



gle terraces in Lake Eyre North. 

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 Silcrcte Island off 
Bonython Head (Fig. 1A). Although eroded, 
the old prehistoric shmgle terraces all bear 
close resemblance in general form and nature 
to the 1974 terraces, and follow contours con- 
forming to the present shoreline. They were 
formed in a 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- 



1*6 



S. A. DULHUNXY 



lOtic terraces would seem to have been similar 
to that of the [VTA filling, as already discussed. 
Cower rates of evaporation in prehistoric mid 
10 laic Rec«nl rime could have resulted in 
greater duration of peak lake- level periods and 
Singer limes of terrace format iom 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, 
equilibrium terrace height appeared to have 
bee* reached after 4 months in 1974. So the 
terrace top height of 90 em above mean lake 
level, -attained in 1<>74, is assumed to have 
heen the equilibrium height attained in the for- 
mation of the prehistoric, terraces, Whilst 
longer time of terrace formation is not likely 
to have produced higbei terraces, it cou^d well 
have buitt wider terraces in association with 
wave cut benches However, after allowing 
for erosion, ihc old terraces do not seem to 
have been wider than the contemporary ones, 
and there is no general evidence of associated 
vwve cut benches. 

Measurements of the relative levels of old 
shingle terrace tups at many places indicated 
the occurence of three groups at successively 
higher levels above the 1974 terraces and 
nifKirntrm mean lake level, They are described 
as the lowest, middle and highest groups for 
the purpose of this paper (Pig. IB, D). Each 
is. considerably more eroded and obviously 
older than the owe beneath ft, They represent 
three prehistoric tilling* of the lake to levels 
higher than that of 1974. No higher terraces 
were found on the present shoreline slopes of 
Hunt and Bahhage Peninsulas, or on Silerete 
Island. Strand gravels recorded by Williams 
H973 1 ) at distances up to 8 km from the 
present b;ke shore iyinji ouiside the field of 
this investigation, contain Coxiella which Is 
absent in the recent shoreline terraces. They 
may have been disturbed by IcetODism, fcnrt 
could teprescnt older, possibly Pleistocene, 
water levels of an earlier phase of lake history 
as discussed later. 

It was concluded that the highest, group of 
old shingle terraces on the present shnrelme 
slope was built bv the deepest lilHng of Lake 
Pyre during the Recent geological history of 
the snlina as we know it to-day. Terraces Of 
this group were studied up greatest detail to 
ascertain as precisely as possible the height 
and age of the deepest prehistoric filling of 
the presen! valina. 



Mostly the old shingle terraces of the 
highest group have heen greatly eroded or 
completely removed. However, in protected 
places, where less erovion has occurred, their 
tops reach uniform maximum levels which 
vary in different areas from 280 cm to 350 
cm above the 1974 maximum mean lake level. 
' They reach levels of 1 541 cm along both 
eastern and western sides of Hunt Peninsula. 
320 cm on both sides of ftabbage Peninsula, 
and 280 cm on Silerete Island. Comparison 
with the shape and general nature of the 1974 
terraces <uggesis thai the least eroded of the 
old terraces, whieh were those selected for 
measurement, have Inst unly about 20 cm from 
their tops. This means that the original maxi- 
mum level of the old terraces was 370 cm on 
Hunt Peninsula, 340 cm on Babbage Penin- 
sula and 300 cm on Silerete Island. It h 
evident that the highest group of old terraces, 
in each of the three areas, is of the same age, 
having been formed during the same prehis- 
toric filling. Therefore, the present differences 
between the terrace-top 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 
wider areas of the lake must be carried out 
before any general conclusions can he reached 
about Recent tectonk movements. The pur- 
pose of the present investigation was to gam 
some knowledge about prehistoric fillings. 
Therefore, Hunt Peninsula was selected as the 
most centrally placed area of old shingle ter- 
races, and likely to have heen least affected by 
tectonic movements known to have been 
prevalent alone the western side of Lake Eyre 
(Williams I973 1 ; Wopfner ■& Twidale 196*7) 

I'i-l ! liiU no Fillings 
Aa discussed above, the terrace-lop level of 
the highest group of old shore-line shingle 
terraces on Hunt Peninsula wus 370 em above 
the peak mean lake level of the 1974 idling, 
and the contemporary 1974 shingle icnacc-top 
level was 90 cm above the mean lake level at 
which the terraces were formed (Fig ID). 
Therefore, the peak mean lake level of the 
prehistoric Jilting which built the highest group 
of old shingle terraces must have been very 
close to ISO 'cm above that of the 1974 tilling- 
This means that the peak mean lake level of 
the highest prehistoric filling of Lake Byre 
salina, in Recent lime, would have reached a 
depth of about 640 cm above the lake bed 
along, the southern shores of Madman Gulf. 



SHINGLE TFRKAChS AND KILLINGS OF LAKE EYRE 



m 



about 850 cm in the deepest pari* or" the gulf, 
and a level of approximately 280 cm above the 
1974 water level in Lake Byre South and along 
I he Centra? Australian Railway line. 

The fevefs of i he terrace tops in the middle 
and lowest or the three groups of terraces on 
Hunt Peninsula measured 230 cm and 140 cm, 
respectively, above the 1974 peak mean lake 
level. Alluwing for 20 cm of erosion, and 90 
cm for (he height of the terrace top levels 
above the mean lake levels at which they 
were built, the two groups would represent 
fillings ot the lake tn depths of 160 cm and 
70 cm above the 1974 peak mean take level 
iPig. ID). 

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 buik terraces, a: inter- 
mediate and lower levels, which were probably 
destroyed or modified by redistribution of 
shingle during subsequent deeper fillings. Any 
terraces built by a Tilling 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 thai of the 1974 filling, includ- 
ing terraces of the 1950 filling, have been 
modified or removed by the 1974 filling. 
Antitftiifv of fitting 

The ages of the three prehistoric fillings of 
Lake Eyre, to depths greater than that of the 
J 974 filling, arc ve-Ty difficult to ascertain pre- 
cisely. The only available evidence is very 
vatfuc and indefinite, and the only conclusions 
possible are of th^ order ot* magnitude of age 
rather than exact periods of time. 

Extensive searches were made for 
carbonaceous materia) within the old shoreline 
shingle terraces, suits hie for carbon dating. No 
shell material or animal bones were found. No 
plant matcrrat such as driftwood was found, 
although sticks and fragmentary pieces of 
wood were plentiful in the contemporary 1974 
terraces, Plant roots Were found in varying 
degrees of decay, but rt was evident that they 
belonged ro plants which grew on the terraces 
after rhev were built. The environment of 
burial in the old shrngie terraces was evidently 
one of maximum aeration, oxidation and pene- 
tration of rain water, lending to complete des- 
truction ^nd loss of all plant material 
originally Included in the terraces. 



Shell material was absent from both con- 
temporary and old terraces, suggesting that the 
kilter were built hy intermittent fillings 
between which aridity and dryness of the 
salina prevented establishment of shell fish 
The more remote and higher gravel deposits 
described by Williams all contain small gastro- 
pod shells (Williams 1973', and per?;, comm. 
1973) supporting the suggestion that they were 
deposited during an earlier pre-salma stage 
when the lake was permanently filled wuh 
water carrying a shell fish community, Studies 
by King fl°5r\ I960) suggest that gypseous 
lacustrine clays with Coxiella occurring along 
the southern shores of Lake Eyre are nf 
Pleistocene ago. and that younger shoreline 
deposits snd sand ridges originated in associa- 
tion with recent delation 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 middle and lower groups have each suf- 
fered successively less erosion. Rainfall is very 
low, bin this is offset by high runoff from arid 
surfaces. FJuvial erosion under these condi- 
tions could be comparable with that of higher 
rainfall area*, where runoff Is icdured by 
vegetation. Wind erosion is not effective as the 
shingle is too large te be removed. 

Where shingle terraces of the highest group 
have not been completely removed, they have 
been breached in places by flowing! water dur- 
ing rare occasions of heavy rain Terraces 
built across pre-existing or antecedent creeks, 
trend upstream in L'-bends as would contours. 
Where hreached, their eroded ends still turn 
upstream (Fig. 1B). Terraces built across 
gently doping surfaces with no pre-existing 
creeks, have been hreached in places by water 
held back on the sloping surface, leaving the 
eroded ends of the lerraee "in-line" and not 
turned upstream (Ftg. 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 weatheiing dolomite to depths of 
130 cm below the bases of the terraces. Tribu- 
tary consequent creeks carrying water along 
the top sides of the terraces towards the 
breaches have eroded channels in dolomite to 
50 cm deep (Fig. 1C). Whilst erosion oF (his 
kind is very difficult to refale |p time, it is 
evident thai the terraces concerned are of con 
siderahle antiquity. 



!** 



J. A. DUL1IUNTY 



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 early to mid Kecent sediments. 
Cjypcrete, silcrete, ealcrcte and ferruginite have 
been formed (Williams 1973 1 ; Wopfncr & Twi- 
ddle 1967). and the processes still appear to 
he in operation. No cementation has been 
found ih the old shingle terraces or the more 
sandy and earthy material forming their basal 
layer*. Suggesting limited antiquity. 

The evidence outlined above is not at all 
precise or satisfactory in attempting to estab- 
lish ages in years, 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 
certamly of mid to late Recent, and most 
probably late Recent, age. For the purposes 
ol physical geography and hydrology of Lake 
hyre, estimates are required for (i) the age 
in years of the deepest prehistoric fining 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. 



After making extensive field studies, taking 
all factors into account, and balancing the 
xnmewhai 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 befoie 
present, but. less than 5,000 years, Further 
narrowing of these, limits must involve infetred 
speculation, but in the author's opinion the 
age may be of the order of 3,000 years, which 
provides an estimate of the age of the deepest 
prehistoric filling of the present salina. The 
middle and lowest groups of old shingle 
terraces arc certainly younger, and could well 
he of the order of 1,000 and 500 years, respec- 
tively. 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 lake 
had previously been filled to the level of the 
J374 iUling. 

Acknowledgments 

it is wished to- acknowledge (■) valuable 
assistance and co-operation of Muloorinp Sta- 
tion which made field investigations possible, 
(ii) valuable discussion with Mr A. F. 
Williams of the Geological Survey of South 
Australia, and (iii) research facilities provided 
by Ihe Department of Geology and Geo- 
physics, University of Sydney. 



References 



Bonytmon, C. W. (1955).— in u U#e F.yre, South 
Australia. The Great Flooding Ot !94SM95(ft 
The report of ihe Lake Eyre Committee, K. 
eeogr. Soc Aust. (S. Aust. Branch), pp. 7-9, 
27-56. 63-70. (Griffin Press: Adelaide.) 

BoNYtftON, C W. (1956). The Salt of Lake 
fcvrc — »t.s occurrence in Madigan Gulf and 
Its possible origin. Trans. H. Snc. S. Aust. 
79, 66-92. 

Dirt monty, .1. A. (IV7*1)< — Salt crnst distribution 
and lake bed conditions in southern areas of 
Take F.vre North. Trans. K- Soc. S. Aust, 
i»8(3), 125-I33. 

Kin<7, D. ( 1956), — The Quaternary stratigraphie 
record at Take Eyre North and the. evolution 
of existing topographic forms. Trans. R. Sov. 
S. Aust. 79. 93-103. 



ICiNC, D. (I960)-— The sand ridge desert* of 

South Australia and related Aeolian land- 
forms of the Quaternary arid cvclc. Irons, H- 
Soc. S. Aust. 83, 93-108. 

Mason. B. (1 955). — In "Lake Byre, South Aus- 
tralia. The Great Flooding of 1949-1950* 
The Report of the Lake Eyre Committee, R 
grogr Soc. Aust. (S. Aunt. Brunch), pp. II- 
2&. I'CitttTin Press: Adelaide.) 

Wuuams, A. V. (1975).— LAKH FYRF map 
sheet. Geological Atlas of South Australia 
1:250.000 scries. Geoi $W\ Si AtiSJL 

Wupfner, H., & Twumi.k. C. R. (1967), — Geo 
morphological history of the Lake Eyre 
Basin, in J. N. Jennings, & J, A, Mahhutt 
Fd.s.. "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 RFXICTPALM UVISTONA MARIAE F. MUELL. 

CE3STRAL AUSTRALIA 



IN 



by P. K. Latz* 

Summary 

Lata P. K. < 1975). — Notes on the relict palm Lhistona mariae F; Muell. in central Australia. 

Trans, R.Sac, S,Au&t. 99(4,), 189-195. 30 November, 1975. 

Comparison of photographs taken after intervals of up lo 56 years indicate thai the relict 
palm Uvistotia mariae may reach an age of up to 300 years. Aspects of the ecology of the 
palm are presented. 



Introduction 

Palm Valley, situated in the Finke Gorge 
National Park in the MacDonneli Range sys- 
tem, Northern Territory (Fig. 1) is of con- 
siderable interest to tourists and scientists alike 
largely because of the presence of a stand of 
the relict cabbage palm Livistona mariae F. 
Muell. This species of Livistona is restricted 
W "«n 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 Livistona 
to the north. The closest relative to L. mariae 
occurs on the Fortescue River in the Hamcrsley 
Ranges, W. Atist., a stand in many ways simi- 
lar lo that at Palm Valley. The relict nature of 
the palms is discussed by Keast (1959), Bur- 
bidgc (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 
VaMey itself, and its discovery is attributed to 
a Lutheran missionary from Hermannsburg 
Mission some time later. The valley was in- 
vestigated by members of the Horn expedition 
during 1S94 (Tate 1896). After a visit to the 
Valley, Lothian (1959) discussed aspects of 
the palm's reproductive behaviour. 

Chippendale (1959) listed 200 plants found 
to occur in Palm Valley. Since this time a total 
of 333 plant species have been recorded from 
the valley (about a 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 Ln 
the valley fed by permanent water seepage. 

Recently a large gas field has been dtv 
covered in the area 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 a 
suitable habitat for its continued survival must 
have a permanent shallow water supply over 
an area large enough to support a viable breed- 
ing population The area must also be pro- 
tected from severe erosion forces or have a 
stable soil environment, as the palm will not 
survive scouring of surrounding soil. 

PaJm Valley appears to be one of the very 
few areas in the central ranges which meets ail 
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 from a large catchment 
area, down lo where the valley dissects these 
strata. The position of this dissection ensUTes 
a continuous permanent seepage area 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 area. 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 Springs, N.T. 5750. 



190 



P. K. IATZ 




Fig. 1. The distribution and numbers of mature individuals of the cabbage palm Livistorm marwr in the 
Finke- Gorge National Park, Northern Territory. 



shallow water table for much of its length, but 
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 Rales and Ages 

The author was fortunate in obtaining a 
number of accurately dated photographs of 
certain 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 pholographie vantage points, 

Of the 13 photograph sites relocated, three 
are presented in Figs 2-4. 'I he 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 
taken) are now still present. The present 
heights of the living palms were estimated with 
a Suunio clinometer. By using the cliff back- 
ground ns a scale, palm heights at the time of 
the earlier photograph were estimated. Distor- 
tion due to angle of viewing and to lens aberra- 
tion was estimated to be less than 10 percent. 
A mean annual growth rate of 10 cm was esti- 
mated for this group of palms. 

Palms in Fig. 3, how r ever, have grown 
approximately Hi m in 38 years, a mean 
annual crowth rate of 30 cm. Palms in Fig. 4, 



which apparently were absent in the earlier 
photographs, taken in 1918, are now up to 12 
m high, a minimum mean annual growth rate 
of 22 cm. 

Palms grown from 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). <\ palm in the cooler climate 
of Melbourne Botanic Gardens is reputed to 
have grown to only 3.75 m in 30 years r a 
growth rate of 12.5 cm per year (Australasian 
Post, 31 May, 1973). 

In its natural habitat, the rate of growth of 
the palm appears to be mainly dependent on 
the water supply. The palms showing the fastest 
growth rate (Fig. 3) are at Palm Bend on The 
bank of the Finke River about 6 km east of 
the Valley. This area has a greater depth of 
fertile alluvial soil and a better water supply 
than most areas in Palm Valley. Palms in Fig 
4 arc 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 UVISTONA MARIAE 



191 




Hg, 2. The upper photograph was taken in 19 17 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 



192 



P. K. LATZ 




Fig. 3. The upper photograph was taken in 1935 and the lower in 1973. The white bed 
of the Finke River can be seen in the foreground. 



range. However, statistical tests showed no sig- 
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 UVISTONA MARIAE 



193 




Fig. 4. The upper photograph was taken in 1918 and the lower in 1973. This area was 
mi?E tw usl 7S.the large rock right of centre in the earlier photograph as a 
marker. This rock is hidden by palms in the later photograph 



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 



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 



I'M 



i». k i \\y 



1973 indicated that about 750 palms of repro- 
ductive age occur 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 to 
show thai rough estimations give a lower 
number than is actually present and may 
account for the previous low 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 i 1896) reported that the white basal 
parts of the inner leaves of the young palms 
were eaten by aborigines in the area. Thi& 
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 rcgrowth ol some edible plants 
and signal their presence to other members of 
(he tribe. Only two tires of restricted extent 
are known to have occurred in Palm Valley 
during this century. Before this time the 
aborigines may have burnt the area mote fre- 
quently to stimulate growth of young palms 
which could then be used tor food. 

Contrary to Tate's observations, there are 
as many palms outside Palm Valley as in the 
valley itself fRg. D. The second largest 



iuiliuiv is in 



little Palm Creek where 550 



mature individuals have been counted. Except 
for about 6 palms further downstream in the 
Finite River, the location of all individuals of 
Lhhtotw mariav known to occur naturally is 
indicated in Fig. I. All ot the palms of repro- 
ductive age have been counted and total 1 500 
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 or established individuals 
of Livhrona mariae (the total naturally occur- 
ring world population) is estimated to be up- 
wards of 3000. 

Alter a good season, each mature palm pro- 
duces an abundance VA seeds with a high 
natural rate of geimination. Although few of 
these seedlings arc able to get their roots into 



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- 
minate and quickly lose their viability unless 
kept in humid conditions (dc Leon 19581, 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 pou-niial 
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 ftotu the lowcrinr til 
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 m other areas of Australia to ensure 
that replacements arc available in case of 
natural disaster is recommended. 
Conclusion 
Palm Valley is situated on the edge of a 
targe gits iield. 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 ihe overall diversity ol the flora, 
A recluse area such as this, is by nature fragile, 
and therefore any development in this area 
will require great care and foresight. 

Acknowledgments 

I acknowledge the help in interpreting and 
dating the photographs lent to 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. 



BuRHRirxih, N. T. (19601.— The r>hytoieo?.raphy 

ol the Australian region. Au.\t. J. Bat. 8. 7?- 

211. 
CHii'PtNDAi.t, O. M ( 1 9?V>— Plants o( Palm 

Valley. Central Australia. Kto Nor, 75, 192- 

19V. 



References 

Chippendale G. M ( 1963 J —The rehe nature of 
some Centra) Australian plants Trans. R. Sc*\ 
\. Aust. 86. 31-34 



r>E LnoK *N. J. ( 1958).— Viability of Palm Seedi 

/Vi>fc//^v2(3>.9h- l >N 



THE RELICT PALM LIVISTONA MARIAE 



195 



Giles, 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. A ust. Mus. Mae. 
13(3), 81-86. 

Lothian, T. R. N. (1959).— Further notes con- 
cerning the Central Australian Cabbage 
Palm — Livistona mariae. Principes 3(2), 53- 
63. 



Tate, R. (1896).— Botany. In W. B. Spencer 
(Ed.) "Report of the Horn Scientific Expedi- 
tion to Central Australia Pt. Ill — 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 

byC. R. Twidale* and Jennifer A. Bourne* 



Summary 

TWIDALE, C R., & BOURNE, Jennifer A. (1975).-Geomorphological evolution of part of the 

eastern Mount Lofty Ranges, South Australia. Trans. R. Soc. S. Aust. 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. 



CEOMORPHOLOGICAL EVOLUTION OF PART OF THE EASTERN MOUNT 
LOFTY RANGES, SOUTH AUSTRALIA 

by C. R. Twidale* and Jennifer A. Bourne* 

Summary 

Twidalh, C. R., & Bourne:, Jennifer A. (1975). — Geomorphcilogical evolution of part of the 

eastern Mount Lofty Ranges, South Australia. Trans, R. Soc. S. AusL 99(4), 197-209, 

30 November, 1975. 

Scattered relicts of a I a ten tic erosion surface of probable early Mesozoic age occur as 
(be Wnalley Surface high in t\\t relief in the eastern Mount Lofty Ranges. However the greater 
pari of the prominent summit surface of the uplands, the Tungkillo Surface, is an etch plain 
due to the stripping of the weathered bedrock and exposure of the irregular weathering front 

The surface of the adjacent Murray Plains is of Pliocene age and was graded to a shallow 
estuary which then occupied the present lower Murray valley. It is cut across Miocene strata, 
though there arc relicts of embayments eroded in Cambrian rocks where major rivers debouch 
from the uplands. It carries a veneer of ferruginous grit, Quaternary calcretc, dunes and 
alluvia. 

The scarps associated wilh the Milendella and Palmer faults are primarily of late Mcso- 
zoic-carJy Tertiary Age. It has been claimed that there has been 60-90 m of dislocation on the 
Milendella Fault since the Miocene, but this is questioned partly because the evidence is 
equivocal, and partly because in some areas the lower part of the scarp is erosional, being 
due to the exploitation by streams of intensely weathered rocks in the fault and scarpfoot 
zones in Quaternary times. 



Introduction 



as the riverine features of the Finnis valley (de 
Mooy 1959), the granite forms of the Palmer 

The Mount Lofty Ranges have received and Caloote areas (Twidale 1968, p. 95 et seq.; 

considerable attention from geomorphologists, 1971. pp. 5-44), and the Permian glacial 

but even in studies which are ostensibly con- features of the Strathalbyn area (Maud 

cerned with the uplands as a whole it is the 1972) have been investigated- Although there 

western areas that have been investigated in has been significant geological mapping of the 

detail (sec Benson 1911: Fcnncr 1930, 1931, region (Kleemun & White 1956; White & 

1938; Sprigg 1946), There has been passing Thatcher 1957: Mills 1965 5 : Thomson 1969) 

reference to particular landforms which occur there has been no consideration of the evolu- 

in the eastern Mount Lofty Ranges (e.g. Fen- tion of the eastern Mount Lofty Ranges as a 

ner 1931. p, 23); there have been reconnais- whole, no attempt to correlate its development 

sanec studies of small parts of the area (Saies with that of the western Murray Basin, and nc 

1968 1 ; Breucr I96S-'; Forrest 1969 s ; Frahn investigation of the interplay of tectonism and 

1971 4 ); and specific features or problems such land form development in the area. 

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

1 Saies, Janet U968).- ''Geomorphology of the Marmum Falls Area". (Unpubl, B.A, Hons thesis, Univ. 
Adelaide.) 

- Breuer, Margaret (1968). — "The Geomorphotogy of Harrisons Creck"\ (Unpubl. B.A. Hons thesis, 
Univ T Adelaide.) 

* 4 Forrest, G. L (1969). — "Geomorphoiogical Evolution of the Bremer Valley" (Unpubl. BA. Hon* 
thesis, Univ. Adelaide.) 

' Frahn. D. (1971). — "Geomorphology of the Milendella Area of South Australia*. (Unpubl. B.A. Honi 
thesis. Univ. Adelaide.) 

: - Mills, K. J. (1965). — "The Structural Petrology of an Atcr Hast of Springton, South Australia. (Un- 
publ, Ph.D. thesis, Univ. Adelaide.) 



w 



C. K TWTOALE &: )£NN1?£K A BOUKNF. 



5*be.?, 1 --"-* . j y- 




Fig. 



Physiographic reeioilS- of part of lhe eastern Mount Lofty Ranges, and (inset)- location map. 
The area nrouncf Rueljcns Hill and depicted in Fig. 5b is indicated. A-R ind CD are lhe tides 
of seciion shown in Fig- ^a- and X-Y lhe sec-lion represented in Tig. 7. 



Genera! Description 

The area under consideration lies between 
lhe River Marne in the north and Gorge Creek 
in the souih. Il falls naturally into three physio- 
j^uphic regions; the high plain, lhe Murray 
PUtttfi and the intervening escarpment (Fig. 

the Summit Hi$h Plain 

Standing at elevations of 200-300 m nhove 
wa level, the summit high plain or Ttiugktllo 
Surface* lies 320-170 m higher than the nearby 
Murray Plains It slopes down to the south 
•'rom some 430 m elevation near lhe Marne to 
330 m south of Harrisons Creek. 

The Surface is eroded in gneiss, schist and 
granite (Fig. 2) which are essentially un- 
weathered and which give rise to distinctive 
iandform assemblages. Boulders known locally, 
and inadvisedly (Twidale 1971, pp. 14-17), as 
'tors\ btivc developed on the northern pari of 
ihc Palmer granite outcrop where the rock is 
wcO-joinicd h*rt not shattered and where split 



rocks, perched rocks or loganstoncs (Fig. 3a) 
together with lhe many clusters of boulders 
iorm a distinctive landform assemblage, The 
foliation of the gneisses has been exploited by 
weathering agents to produce pen item ( Acker- 
man n 1962) or tombstone rocks <Fig. 3b) but 
in other plates, as for instance o-K km cast 
of Mount Pleasant, the gneiss is more massive 
and forms low (up to 10 m) angular hlocky 
residuals (Fig, 3c) or miniature ensile koppics. 

Also standing above the level of the Tung- 
killo Surface are several scattered low (ngam 
up ro 10 m) mesas or conical hills which arc 
cither underlain by laleritic weathering profiles 
carrying a ferruginous encrustation, or are 
capped by alluvial gravels and boulder* of pre- 
dominantly qustrlwjic composition, The.se 
residual hilts coJlccliveiv form what has been 
called the Whatley Surfaced. 

The Tungkillo Surface is deeply incised by 
such Intermittent streams as ihc River Manse, 
and Saunders, Milendclla. Harrisons and 



GKOMOKPHOLCXilCAL EVOLUTION OF Tttfl EASTERN MI LOFTY RANGES 1*>9 




paiivozolQ socinTientt 



Fig. 2. Generalised geological map of part of the Mount Lofty Ranges (after S. Aust. Cieo]. Surv, mop 
sheet SI/54-9, Adelaide). 



Gorge creeks. Their valleys are however sur- 
prisingly narrow in proportion to their depths, 
and in many places the streams run 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 
M comprises two north-south trending scarps 
which arc 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 Raerjens Hill and south through Palmer 
(Fig.2). 

The scarps are intensely dissected. The 
Milendella Scarp is 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 latter 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. 
In some places the upland scarp displays a basal 
steepening and scarpfoot depression, as do 
some of the ridges within the upland (Fig. 4a). 
In some few sites, prominent flats occur at 
levels intermediate between the Tungkillo Sur- 
face above and the Murray Plains below. One, 



the Millcndella Bench, is greatly dissected by 
the present Milendella Creek and its tributaries, 
but there are nevertheless broad 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 deseend by way of a 
steep slope to the Murray Plains (see Tepko 
1 .50.000 topographic series (6728-111) 
bftween grid lines 320 and 347 ? and 440 and 
493; see also Fig 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 
ot alluvium. It was associated with an in- 
trenched meander simiiai to those found on the 
present Milendella Creek but of roughly 5—6 
times greater radius. The meander loop and 
core developed when the precursor of Milen- 
della Creek flowed at a level some 80 m 
higher than at present. 

A second bench occurs where Harrisons 
Creek leaves the uplands to llow across the 
Murray Plains, where it is known as Reedy 
Creek (see Tepko 1 : 50.000 topographic 
series (67284II) around MR3I3374; see Fig. 
4b). Less extensive than the Milcndelld 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. ^Ijjcarpfoot depression developed at the base of a gneiss ridge at Raetjens Gap. (C. R. 

b. The Palmer Scarp west of Tepko, showing the Tungkillo Surface, the low scarp (a) the 

scarpfoot equivalent of the Palmer Bench (b). the break of slope on the hogback scarp (c) 

ft? \v£ P n 0Ot e Val ! ey (d) " and ,he lower stce P« "»rp face (e). (C. R. Twidale) 

L P e ,T^ nL f 6 (a> and - Tl ! n ^ i110 S,,rface (b) west of Mount Pleasant. Outcrops of the 
gneiss country rock are seen in the foreground (c). (C. R. Twidale.) 



202 



C R- TWIDALF & JENNIFER A. BOURNE 



» 



.'JM 



3 






run i. i"'.m'! i 



KILOMETRES 



Fig. 5. a. 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. 




Okm 

i 



contour interval 20 metres 



Fie 5 b Generalised topographic map of the Milendella Bench showing the old meander loop and the 
g ' iSSSoM?S^hm 339). Drawn from S.A. Lands Dept. 1:50,000 topographic sheet 

6728-1 UTcpko. 



GfcOMORPHOLOGIOAI. EVOLUTION OF THE EASTERN MT LOFTY RANGES 2£l% 



Ifrfit, 50 m above the adjacent plains and 130 
m below the Tungkillo Surface and in detail 
Includes several levels which represent stages 
in downcutting. This Calmer Bench (so called 
because the site of the original Palmer town- 
ship was only a short distance away) carries a 
mantle of quartzkic blocks, some of which are 
iron-stained It has been dissected by Reedy 
Creek which in this sector displays wclE- 
developed intrenched meanders. 

Other, similar, perched benches have been 
noted near the mouth of Pine Valley, south oF 
Tepko, and particularly west of Strathalhyn 
where there is an extensive dissected bench, 
but as these arc outside the study area they are 
not considered further. 

Murray PUthts 

The western Murray Plains comprise a roll- 
ing lowland which stands 200-150 m high 
near the scarp foot, to 70-50 m at the cliff top 
overlooking the River Murray. In several sec- 
tors there is a scarpfoot depression up to 50 
rtl deep fronting the Mount Lofty Ranges fFig. 
I). 

These western Murray Plains are underlain 
by Caino/oic strata 80 m or mare thick. In the 
west. Quaternary fnnglomeTates predominate 
hul these thin to the east where Miocene 
marine slrata occur in bores, in Uibutaiy 
valleys, and in tfie high precipitous bluffs bor- 
dering the Murray trench (Fig. 2). Borelog 
data indicates thai the Miocene strata rest on 
an irregular surface cut in acid and basic crys- 
talline rnckv Basic crystalline rocks ale 
exposed at Black Hill, where Ihc so-called 
'norile' ha* been extensively quarried; similar 
rocks have been located in bores near Sedan 
and Walkers Flat. Granitic rocks and schists 
arc exposed extensively in the valley of Reedy 
Creek, but also in many other minor outcrops. 



Hmv did this landscape evolve? Of what age 
and origin are the ntains ,? What is the nature 
nf ihc scarp which separates the high from the 
low plains? 

The lineai ity of these scarps and their 
association with faults exposed in Ihc gorges of 
the River Myrnc and Saunders Creek or in- 
ferred from the displacement of strata, suggests 
that the topographic forms are fault generated 
and probably fault scarps, i.e. due directly and 
wholly to tectonic displacement This is ccr- 
laittJy the view of Mills 1 who with resneci 
lo the Milendclla Fault, which fe of reverse 



type, argued ihat several stages could be 
detected in its development: 

1. Initiation of The fault in the Palaeozoic 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 2fi0 m. 

4. Miocene marine incursion with strata over- 
lapping the eroded fault-scarp. 

5. Recurrent movement on the fault wince the 
Miocene, the east side being lowered by 60- 
90 in. 

Sevetal aspects of this chronology can be 
challenged, and in particular the date and ex- 
tent of faulting arc open to question. The proh- 
Icm involves a consideration of the age and 
character of erosion surfaces identified in Che 
Mount Lofty Ranges and on the adjacent 
Murray Plains. 

Age and Nature of mW Summit Surface 

The summit surface is a complex feature. Tin; 
iatcritic remnants nf the Whallcy Surface arc 
crucial to the interpretation of the whole up- 
land. They arc sufficiently scattered to suggest 
that they arc relicts of a once-contiguous 
weathered surface of low relief, over which 
lluwed streams carrying, a gravelly and 
bouldcry quatuose bcdload, Remnant deposits 
of these boulder beds occur in places, and a 
discontinuous mantle of angular quartz frag- 
ments forms a veneer on Lhe Inw divides. The 
weaihering extended only some 10 m beneath 
the surface but it is notable that the base of 
significant weathering — the 'weathering front* 
of Mabbuft (1961) — is everywhere coincides 
with the local lever of the tungkillo Surface. 
underlain by intrinsically fresh rock. Thus it 
is reasonable to suggest that the Tungkillo Stu- 
face is an etch plain (Wayland 1934), formed 
as a result of the stripping of the Interitic deep 
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 frntil 
and exposed as a result of the stripping of 
weathered material. 

The age of the WhaMey Surface and of the 
deep weathering is Meso/oic and probablv 
early Mesozoic (Triassic). The latente rem- 
nants can be traced to ihc southern Mount 



204 



C rt TWTDALE & JENNIFER A BOURN F 




Fig. jS. Diagrammatic section Through the eastern 
Mount I ofty Range** showing the rela- 
tionship between the laleritiscd Whaliey 
■Surface muferjalll by weathered bedrock 
and the etch plain, coincident with the 
weathering front , uf the TungkLllo Sur- 
face, 



Lofty Ranges where the laterite surface stands 
high above valleys pnitially iilled by Miocene 
marine sediments, and where the fault angle 
depressions resulting from the fault-dislocation 
of the summit surface contain basal marine 
sedimeni of early Tertiary age. and with in- 
clusions of laterite blocks I'Glacssner 1953; 
Glacssner & Wade 1958; Campana 1958). 
Thus the summit .surface and the laterite it 
carries must be of earliest Tertiary or late 
Mesozoic age However, evidence from Kan- 
garoo Island, just, across Backstairs Passage 
from Fleurieu Peninsula, suggests a much older 
age. There the laterite plateau formed on Cam- 
brian and Penman strata was partially dissected 
before the extrusion of Middle Jurassic basalts 
near and just west of Lhe present site of Kings- 
eoie. Pataa »climatic consideration suggest the 
Tiiassic as the most likely period fur laterite 
development (Daily, Twidale & Millies 1974). 
Thus by short-distance correlation the 
Whaliey Surface and the associated deep 
weathering are considered to be of early Ivfeso- 
aoic age. The development of the Tungkillc 
Surface followed the disruption of the upland, 
probably by faulting. Streams were rejuvenated 
and they exploited the contrast in cohesiveness 
and resistance between Ibe rcgolilh and the 
fresh Tocks beneath, wirh the result that an 
etch surface of low relief, the Tungkillo Sur 
face, was initialed. It i* still extending. 

Disruption nf (he Summit Surface 

In view of the character of the Milendella 
and Palmer scarps it is obvious that the reason 
for the disruption of the summit surface could 
have been a renewal of movement on tbe two 
recognised faults, If this were so, rocks similar 
1o those of the nearby upland should be found 
beneath the Cainozoic strata of the western 
Murray Basin GTamtes and gneisses arc 
exposed widely in the valley of Reedy Creek, 



in minot outcrops throughout the western 
Mutray Basin, and they are commonly en- 
countered at shallow depths west of tbe River 
Murray. But there is one feature missing, 
namely buried laterile. though weathered 
t.kaoltnised) Cambrian bedrock is recorded in 
bores beneaih the fresh crystallines and tbe 
overlying Cainozoic strata. There are also 
superficial ferruginous encrustations on high 
plain remnants and at higber elevations than 
the Miocene strata and therefore of lalei 
Cainozoic age, in the Murray Plains, for 
example just south of the Mannum Falls. 
Further reference is made to Ihese occurrences 
below, but no laterite profiles have been found 
cither in outcrop or in bores. 

There are a number ol' possible explanations, 
For some reason or reasons at present un- 
known, laterite may not bavc developed on 
Ihose rocks of what was- to become the down- 
thrown block; this is rejected as being 
irrational. Second, The laterite could have been 
eroded before burial by Miocene and later 
sediments, but then some remnants should 
surely have been preserved. Third, the laicrtie 
may indeed be preserved beneath the Cainozoic 
cover, but has not yet been discovered. This is 
unlikely because many bores have been sunk 
anil though there is some reference to 
weathered crystallines, there is none lo laterite. 
Fourth, tbe weathering ot the Cambrian rocks 
could have taken place beneath the Cainozoic 
cover, but it is unlikely thai the oxidation i\^- 
rcsetired by the weatheted bedrock could have 
developed in such conditions. Fifth, the latenre 
could have "been buried and iron oxide duiicrust 
deslroyed by groundwater attack in the sub- 
surface, Such dissolution of iron oxide by water 
charged with organic acids bas been shown to 
be possible (Bluomrield 1957*. Coulson, Davjs 
& Lewis I9b0; Hingston 1963), 

This last explanation *eems to be the tnosi 
likely. All other indications arc that the 
Whafley Surface wav dislocated by faulting, 
with the cast side subsiding by a matter of 260 
m if] the north. The concentration of ground- 
waters in the scarp foot zone (see Ruxtun 
1958; Twidale 1962, 1%7) would account for 
the subsurface elimination of the ferruginous 

crust, 

Thus if it is accepted that faulting occurred 
after the development of the lateritised sumnm 
surface but prior to the Miocene marine trans- 
gD&slrJt], a numbei of geomorphological 
changes eau bo related to this period; 



GrOMORHHOLOOlCAL EVOLUTION 01- THE EASTFRN MT LOFTY RANGTS MS 



1 Pie rejuvenated streams began the stripping 
of the lafeWte regolith and the development 
of the Tungkillu Surface. 

2. The rejuvenated streams incised into the nn- 
weaihered bedrock creating quite deep, nar 
row, and in places meandering gorges, 

3. As they emerged from (he uplands, lhe 
streams eroded quite broad ewhayments in 
the crystalline basement rocks of U>e uplands 
contiguous widi the then surface of the 
.Murray Plains. The Milendella Bench stands 
considerably higher than either the Palmer 
Bench or bench remnants at the mouths ol 
the Saunders and Marne gttt^cji. This may 
reflect its development at a stage before 
faulting hud ceased, and its grading to the 
Miocene shore, or if may indicate toca* late 
Cainuzoic faulting, 

When did faulting begin? There is. no direct 
evidence, but the stratigraphy of the Murray 
Bnwn vO'Driscoll I960; l.udbrook 1969) sug- 
gests that part of (he region became a jeMrictcd 
marine emhayrnent during the early Crcla- 
eeuus. The next proven marine transgression 
took place during the Late Eocene ami marine 
influence continued through to the Middle Mio- 
cene. However, though strata of Late Creta- 
ceous age have not been located, they may 
have been deposited and .subsequently been 
removed by erosion (l.udbrook 1969. p. 159). 
The fail It dislocation was probably gradual and 
possibly occupied the whole of this late Mesu- 
zoic-middle Tertiary period 

Age of tlie Plains Surface 
The broadly /oiling plains surface of the 
western Murray Basin is bfkhg dissected by the 
few through-flowing streams such as the Marnc 
Milendella and the Harrisons-Reedy systems, 
which emerge Horn the Mount Lofty Ranges 
and persist to rcaef> the Murray. The River 
Murray trench is a Quaternary feature, though 
there wa-s an earlier shallow estuary coincident 
with the present Murray valley as tar north as 
Morgan. In this long narrow and shallow estu- 
ary, marine sediments, the Norwesl Bend For- 
mation, of Late Pliocene age were deposited 
(l.udbrook 19931}, Remnants can be 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 Pliocene 
stiata, it must be younger and there is much 
general .i-gumem as well <is specific evidence 
(von der Boreb 3968) to suggest that it is or 



Pleistocene age, being related to low glacial 
stands of the sea. 

However, in Pliocene times the regional baie 
level on the western Murray Plains stood 50- 
70 m higher than it does now, and it is to this 
higher ba.selevel that the streams responsible 
for the erosion of the plains were graded. 
These streams transported to the then Murray 
estuary the detritus which in part comprises The 
Norwest Bend Formation. Thus the Murray 
Surface Js in the study area of Pliocene age. 
There is some corroboration from the thin 
ferruginous duricrusts found on some parts of 
the surface, as for instance just south and 
south-west of Mannum Fatls. for surfaces of 
comparable age on northern Yorke Peninsula 
(Hbrwftz & Daily 19581 and Eyre Peninsula 
(Twidale, Bourne & Smith 1976) also carry & 
ferruginous encrustation, as indeed do posl- 
Miocene valley floors in the southern Mount 
Lofty Ranges (Horwiu 196CM, and the 
Karoonda Surface of the central Murray Plain* 
(Firman 1973, p. 23). 

lhe occurrence ol iron staining, as well as 
their elevation, suggest that lhe several perched 
benches found at the mouths of major upland 
gorge* were also formed at this time, even 
though they are now separated from the plains 
by scarpfoot valleys up to 50 m deep. This is 
corroborated by profiles relating the benches to 
the main plains surface (Fig. 5a). 

Although little is known of the geometric 
rclationship between river discharge, channel 
width md the radius of intrenched meanders 
tcf. Bales 1939 and Dury 1954, 1964; with 
respect to flood plain meanders, also Gey I 
196$) tf> e abandoned meander loop preserved 
o*i the Milendella Bench suggests rivers of 
much greater I? occasional t discharge than 
those active at present. 

Thus, during the Pliocene, |hc scarp foul 
stood 80 m higher than present at the mouth 
of Milendella Creek and 50 m above present 
Ivucleve! where Harrisons Creek debouches oi: 
to the plain. Is the post Pliocene scarp tectonic 
or erosional, or are there element oi" both 
origins present? 

The Question of Post I ate Cainozoie Faulting 

The lower part of the Milendella, and by 
implication the Palmer, fault scarps are seen 
by Mills (1965, pp £36-442)* as due to post 
Miocene faulting. He discovered several out 
liers of Miocene sediments perched on the 
scarp north ot Milendella. correlated them with 
Miocene strata found In bores in the Carahrai 



473 metres 




C. R. TWIDALE & JENNIFER A. BOURNE 



perched 

Miocet<« Inoostone 
162 _ 
QuarernaTyalluviiim 

upper Miocene sand 

Miocene* limestone 

76 . 

Eocene lipniticsands 

40 

Sea levei 




Ckm 
\ 



fig. 7 Section through the Milcndella Scarp near Suundm Creek, I After Mills 1965) 



and Milcndella areas, and took the eievational 
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 ihe Pleistocene faoglomcrates 
abutting the fault plane display orientation sug- 
gestive of recent movement along the plane. 

This suggestion of recent and continuing dis- 
location is borne out by the seismic record 
(Sutton & White 1968) which shows that the 
eastern border of the Mount Lofty Ranges is 
subject to earth tremors. The distinctness of 
the escarpments both here and to the south 
suggests geological youthi'ulness (see e.g. .leune 
Si Chittleborough J974). The higher level of 
ihe Milcndella Bench (see earlier) can also be 
interpreted as indicating local late Cainozoie 
dislocation. 

On the other hand, the scarps have been 
deeply and intricately dissected, the Milendetla 
scarp in particular being greatly cut about yet 
retaining its overall linear 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 strati- 
graphically lower or older Mannum Formation 
and the higher and younger Morgan Formation 
(Ludbrook 1969). The two are very difficult to 
distinguish in the field and can only be dilTeren 
riated palaeontologieally. The faunas of the 
perched outliers are unfortunately not diagnos- 



tic, Mr. J. M. Lindsay reported (pers. comm.) 
in 1975: Ci A sample of recrystailised quartzosc 
fine-grained limestone, from a perched oulliet 
of mid-Tertiary rocks 3.5 km north of Sander- 
ston> contained small benthonic forammifcra 
bur no diagnostic forms and no lithological 
features which would distinguish between Mor- 
gan Limestone and Mannum Formation." As 
the stratigraphic position of the Miocene en- 
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 cufre- 
lation is unwarranted. It can be argued that the 
topographically higher outliers are Morgan 
rormatioo left behind as remnants of cireum- 
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 For- 
mation in situ, and thus that no post~Mit»ccne 
dislocation of any significance is implied by the 
evidence (Fig, 8). This crosional hypothesis 
receives support from the character of the 
Milcndella and Palmer scarps. If the scarps 
were wholly of tectonic origin, then the scarps 
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 



UEOMORPHOLOGICAL EVOLUTION OF THE EASTERN MT LOFTY RANGES 



207 




a fatting 
impMSd 



Morgan 




b no posl Miocene 
faulting implied 



Fig. 8. Two possible modes of evolution for the 
eastern scarp of the Mount Lofty Ranges. 

or 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 (see Figs. 1, 4b and 
5a), frequently with intensely weathered rock 
exposed in the valley floors. In between these 
sectors arc others where (here are no scarpfoot 
streams, where the escarpment is of lesser 
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 die upper part of the scarp 
is of fault or tectonic origin, though of con- 
siderable antiquity (late Mesozoic or early Ter- 
tiary), but that the lower part, where present, 
is due to erosion by streams eroding the 
■weathered strata of the scarpfoot zone (Twi- 
dalc 1967): i.e. that the lower scarp is of fault- 
line type and that no significant recent disloca- 



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 erosionai 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 Milendclla 
and Palmer scarps. It is not a matter of con- 
trast in rock type or volume of run-off, for Hie 
lithological units of the eastern Ranges run 
north-south and are essentially similar behind 
both scarps. It cannot be a question of climatic 
differences. It may be due to the Milendella and 
Marne draining a higher scarp, and eroding 
regressivcly more quickly to capture the head- 
waters of the Torrens drainage. More likely 
however, the dissection of the southern part ot 
the study area has been retarded by the base- 
level control exerted on the Harrisons-Reedy 
creek system by the large mass of crystallines 
exposed west of Caloote. The lower elevation 
of the Palmer as compared to the Milendella 
scarp may be a contributary factor. 

Conclusion 

The development of the eastern Mount Lofty 
Ranges and the parts of the Murray Plains is 
summarised below. 



Cambrian 
Early Palaeozoic 

l-ale Palaeozoic 

Early Mesozoic 



Late Mesozoic- 
Middle Tertiary 

Pliocene 



Quaternary 



Deposition in Adelaide Geosyncline 

Orogenesis, granite emplacement and mctamor- 
phism 

Permian glacialion (not evidenced in 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 rejuvenation 
and stripping of regolith 

Essential withdrawal of sea, only estuarine sedi- 
mentation. Planation of Miocene sediments and 
of emhayments in crystalling rocks on face of 
scarp 

Lowering of sea level. Murray Trench formed, 
tributaries rejuvenated, valleys and scarpfoot 
zone excavated 



Whalley Surface 



Tungkillo Surface. 
Scarp formed. Gorges 
developed 

Perched benches and 
Murray Surface 



Scarp foot valleys, 
abandonment of scarp 

benches 



20K 



C. R. IW1DALH fr JHNNIFKK A BOWKNk 



Acknowledgment* 
rhe writers wish id thank Dr Brian Daily, 
licology Department, University or Adelaide, 
lor a critical reading or" the paper, and Dr 



Kiugsley Mills, Department of Geology and 
Geophysics, University of Sydney, for helpful 
discussion and continuing interest in ihe evolu- 
tion of the area under investigation. 



References 



Ackhrmann, E. (1962). — Biisscrsteinc-Zjcugen var- 
zeitlicher Grundwassetscb.Wank\irjgen. '/.. Geo- 
morph. (N.S.) <i. I4&-1S2. 
Bai'ES, R, h\ ( 1939 i -- -Geomorphic history or the 
Kickapoo Region, Wisconsin. Ball. %eoL Sw 
Am. 5», 819-879 
Ulnson, W. N. (1911). — Note de&eriplive of a 
steieogium ot' the Mt J-ofly Ranges. South 
Australia. Trans: R. Soc. S, AitSt, 35 } 108-11 1, 
Hloommei d, C (1957). — The possible signifi- 
cance of polyphenols in soil formation J. Sci, 
FJ A^ric, 8, 389. 
Cam('/us% B. 1, 1958).— The Mt f_ofty-OJary regiun 
and kangaroo Island, hi M. F. Glaessner & 
L. W. Parkin (eds) 'The Geology of South 
Australia", pp. 3-27, (Meib. U.P ) 
COI'ISON, C, R. Davis, R. 1., & Lrwis. D. A. 
I I960). — Polyphenols in plant, humus and 
soil. I Polyphenols of leaves, Vnter. and super- 
ficial humus from mull and mor sites. II Re- 
duction and transport by polyphenols of iron 
in model soil columns. J, St/iL Sci. M, 20. 
ruiiv, B, Twiiivth, C. R., & Milnes. A R 
(1974). — The age of the lntenu>ed summit 
surface on Kangaroo Island and adjacent 
areas of South Australia. J, Gtol. Soc. Attst. 
21.3517-392. 
Dt/RY, G. H. 1 1954) . — Contribution to a general 
theory nf meandering valleys Am J Set. 
251. 193-224. 
Duhv, G. H. (1^64 /.-Principles of underfii 
Streams Prof. Pop. U.S. geol. SktV, 452-A. 
J -67. 
FrNNER. C (I9"HI). — The major Mruetut'ul and 
physiographic features of South Australia. 
Trans. R. Soc. S. Awit. 54, J-36. 
Fi.NNbR, C. (1931), — "South Australia: A Geo- 
graphical Study." (Wbiieornbc and Tombs - 
Melbourne.) 
t-tNNFR, C. (1939). — The significance of the topo- 
graphy of Ansiey Hill, South Australia. Trans: 
R.Soc.S.Austtt, 79-K7. 
Fkman, J B. (1973).— Regional stratigraphy of 
surfkinl deposits in the Murray Basin ami 
Gambicr Embayment. Rep!. Invest. w»l. 
Surv. S. A ust. 39. 
Geyl, W, F. (19GH).— Tidal stream action rtnd 
sea level change as one cause of valley mean- 
ders and under fit streams. Aust, Gent*, Sou!. 
6, 24-42, 
Glaes.sner, M. F. (l-WI— Soqje problems of 
Tertiary geology in southern Australia. A R, 
Sac. JVJ5.lV. 87(2), 31-45, 
G»ahssner, M. F., & WAUh. Mary (1958).— The 
St. Vincent Basin in M. F. GlaessneT & 1 . W. 
Parkin teds). "The Geology of South Aus- 
tralia", pp 115-126. (Meib.U.P.) 
Hinumun, F. J. (1963), — Activity of polypheno- 
ls constituents of leaves of Etu:alvptus and 
other spevtes in completing ;md dissolving 
iron oxide. Auxt. /. Soil. Rr.w HI), 63-73 



Horwitz. R. C. ( 1960). — Geologic de la region 
de Mt Compass (feuille M'huig), Anslrulte 
meridionjle. fcW/v,i>. Gtol <teh 53, 2M-2fr\ 

HoRwtiA R- Cm & DmiX B. tl95B). -Yorke 
Peninsula, in M. F. Glaessner & L. W. Par- 
kin <eds). "The Geology of South Australia". 
pp, 46-r,0- (Melb. U.P.) 
Jeone, R. P., & CHtTTtnuoBoiioH, D. F (1974). - 
City of Monarto. Preliminary Geotechnical 
Investigations. A'. Aum, Drpt, lijincs Rent. Bk 
74/105'. 

KiruNHN, A. W, ik White, A. I. K. (t95r>).— 
The stiuctural petrology ot portion of tr»e 
eastern Mount Loftv Ranges. /. Gcol, Hoc, 
Auxt. y 17-3J. 

LtmbftOOK. N. H. ( 195S). — The Murray Rasiu in 
South Australia fa M F' Glacssner & 1.. W, 
Parkin (edsl "The Geology of South Aus 
1rafia H 7 pn. 102-114. tMelh U.P.) 

I iT.i-ipnrtn. N. II. 1 1969). — ^Tertiary Petiod. ht 
L W. Parkin {cd.\. Handbook of South 
Australian Geology", pp. 172-203. (S. MJL, 
Geol. Surv.: Adelaide ) 

MAiruun, J. A. (I9(>1)! — "Bufiui surface' or 
''weathering front". Proc. Geol. Ax\: London. 
72. 357-358. 

NUl'o, R. R. (I972). — Geology, gcomorphology, 
and soiln ot* central County llindmarsh i.Mt 
Compass-MUang), South Australia. C.SJ.R.O. 
Soils PabL 29. 

im Mooy, C. .1. (1959).— Notes on the- isomor- 
phic history of the area surrounding lakes 
Alcxandrina und Albert, South Ati*.Uulia 
Trans, R. Soc. S. Attst. 82, 99-1 18. 

O'DRificoi.t., E. P. (I960) — The hydrology of the 
Murray BaKifi Province in South Australia. 
Butt. veo/. Snrv. S Ausj. 35. 

RovrON, B. P. 11958!'.— Weathering nnrl subsur- 
face erosion in granite at the piedmont angle, 
Balos, Sudan. Gcol A/ffj?. 45. 353-377. 

Svkiug. R. C 11946). — Reconnaissance geological 
survey of portion of the western escarpment 
ol" the Mount Lofty Kafigca imns, R. Hoc, 
S.Auxt.lO, 313-347 

StmoN, D. f„ & Wnmt. R, H. I 1968)— The 
^eismicity trf South AtMialia, / Gtol. Sot: 
Aw*. 15. 25-32. 

Thomson. B tv (196^).— AUUn'th SA Gcologi- 
cnl Atlas Senes_ Scale 1 :250«00U Sheet St.- 
?4-9, zones 5 & 6. (S, Aum. geot. Surv.: Ade- 
laide.) 

Tuorvton, R. C. N. r, 1974 ) . — Hydrocarbon 
potential ol' western Murtuy Babio and jnfiA 
basins. S, Aust. Geo}. Surv, kept. Invest. 41. 

TwtDAifc. C. R, (1962).— Steepened margins of 
inselbergs from northwestern F.yie Peninsula 
Soudi Australia Z. Gcomorptr (N.S.) 6. 51- 
69. 

'IwiDALt, C K. (1967). — Origin of the piedmont 
angle as evidenced in South Australia. J- 
OVaf, 75,393-411. 



GEOMORPHOLOGICAL EVOLUTION OF THE EASTERN MT LOFTY RANGES 209 

Twidale, C. R. (1968). — Geomorphology with von der Borch, C. C. (1968). — Southern Aus- 

special reference to Australia. (Nelson: Mel- tralian submarine canyons: their distribution 

bourne.) and ages. Mar. Geol. 6, 267-279. 

Twidale, C. R. (1971).— Structural Landforms. Wayland, E. J. (1934).— Peneplains and some 

(A.N.U. Press: Canberra.) erosional platforms. Geol. Surv. Uganda. Ann 

Twidale, C. R., Bourne, J. A., & Smith, D. M. Rept. Bull, 77-79. 

(1976).— Age and origin of palaeosurfaces on White, A. J. R., & Thatcher, D. (1957).— Man- 
Eyre Peninsula and the southern Gawler num Geological Atlas 1 mile series, No. 811 
Ranges. Z. Geomorph. (in press). Zone 6. (S. Aust. geol. Surv.: Adelaide.) 



THE WOODENDINNA DOLOMITE AND WIRRAPOWIE LIMESTONE - 
TWO NEW LOWER CAMBRIAN FORMATIONS, FLINDERS RANGES, 

SOUTH AUSTRALIA 

BY P. G. HASLETT* 



Summary 

HASLETT, P. G. (1975) .-The Woodendinna Dolomite and Wirrapowie Limestone-two new Lower 

Cambrian formations, Flinders Ranges, South Australia. Trans. R. Soc. S. Aust 99(4), 

211-219, 30 November, 1975. 

Two new formations, the Woodendinna Dolomite and the Wirrapowie Limestone, are proposed for 

two distinctive units of the Lower Cambrian carbonate sequences of the Flinders Ranges, which 

have not, hitherto, been differentiated from the presently defined formations. 

Both formations are widespread in their occurrence, but are best developed in the eastern Flinders 

Ranges, north of the line of the Blinman, Wirrealpa and Frome Diapirs. The Woodendinna 

Dolomite and Wirrapowie Limestone record a significant period of supratidal and shallow, 

sheltered, intertidal deposition respectively within the Cambrian of the Flinders Ranges. 



THE WOODENDINNA DOLOMITE AND WIRRAPOW1E LIMESTONE — TWO 
NEW LOWER CAMBRIAN FORMATIONS* FLINDERS RANGES, 

SOUTH AUSTRALIA 

by P. G. Haslett* 

Summary 

HytSLfixr, P- G. (1975). — The Woodendinna Dolomite and Witrapowie Limestooe — two new 

Lower Cumbrian formations. Flinders Ranges, South Australia. Trans. /?. $<n:. S. Aust. 

WHJ, 211-219, 30 November, 1975. 

Two new formations, the Woodendinna Dolomite and ihe Wirrapowie Linieslune, are 
proposed for two distinctive units of the Lower Cambrian carbonate sequences of the Hinders 
Ranges, which have not, hitherto, been differentiated from the presently defined formations. 

Roth formations are widespread in their occurrence, but are best developed m the eastern 
Hinders Ranges, north of the line of the Blinman. Wirrealpa and Frome D tapirs. The Wooden- 
dinna Dolomite and Wirrapowie Limestone record a significant period of supratidal and shallow. 
sheltered, interiidal deposition respectively within the Cambrian of the Flinders Ranges. 

Introduction the WUkawiliina Limestone, the Parara Lime- 
Formations within the Hawker Group (Dal- stone and the Ajax Limestone (Table 1), several 
garno 1964) of the Lower Cambrian of the distinctive lithofacies exist. Not only do the 
Adelaide Geosyncline characteristically show present formations contain separate and InJi- 
frequent changes in both thickness and facies, vidually mappable Ethologies of significantly 
This results in complex intertonguing of litho- different depositions! origin but in the past the 
logical units. The Ten Mile Creek type section formation names have in some instances been 
(Daily 1956), measured within a syndeposi- loosely applied (Dalgarno 1964, pp. 134-1 35"L 
tional graben structure northeast of the Ora- table i 

parinna Diaplr, represents one of the thickest F.xixrinz jotmatinn names ami their sources for the Lower 
««.j «*»*•*. .,~~,.,i .*~ *. A „ «.««„.. «£ *u~ xj ...1,^ Cambrian carbonate sequences of the Flinders Rang*;* 

and most complete sequences or the Hawker : : — 

Group in the Flinders Ranges. Mapping by the western CE ea1stern NI> 

S.A. Department of Mines (Dalgarno & lake torrens flinders flinders 

i,i~ laii, r-w* i tanri\ „a u. t» AREA RANGES RANGES 

Johnson 1966; Coats et al. 1973) and by B. 



Daily and a number of his students (Gehling 1 ; limesi'ONE 

Haslett-: Hatcher 1 : Mi>unl :i ; Hull"; Daily andamooka. ajax (Duit> iSSfi) 

Irt-fc-, , . ... It. -,- c LTMESTONE. LIMESTONE 

|y/2a) has resulted m the recognition of (Johns 196S) (Daily 1956) wilkawillina 

several significant litbological sequences not ^e^JwTSgF 



present within the Ten Mile Creek type section. 

The thick basal Cambrian carbonates in par- Whilst it is extremely impractical and un- 

lietilar exhibit complex and varied facies inter- desirable to introduce an excessive amount of 

relations and it has become obvious that within formal stratigraphic terminology for the Lower 

<hc presently established broad divisions, viz. Cambrian carbonate rocks, it is necessary that 

4 Department of Geology and Mineralogy, University of Adelaide, Adelaide. S. Aust. 5000. 

1 defiling, J G. (1971). — The Geology of the Reaphook Hill Area. Honours Thesis, Univ. Adelaide 

(unpublished). 
" Haslett, P. G. (1969). — The Cambrian Geology North of the Wirrealpa Dinpir, Flinders Ranges, South 

Australia. Honours Thesis, Univ. Adelaide ("unpublished), 
s Mount. T. J. (1970). — Geology of the Mt Chambers Gorge Region. Honours Thesis, Univ. Adelaide 

(unpublished). 

5 Hatcher, M. I. (1970*),— The Geology of the Mt Chambers Mine Region, Northern Flinders Ranees. 
S. V Honours Thesis. Univ. Adelaide (unpublished). 

^Htdl, K. G. (1973). — The Lower Cambrian, Puttapa Synclinc, FlfudetK Ranges. S.A. Honours Thesis. 
Univ. Adelaide (unpublished). 



212 



?. O. HASTKTT 



sufficient nomenclature be available Ic map and 
describe, on a regional scale, the major ruck 
relationships preseni within the depositions 
basin, Careful recognition of mappable units 
fil uniform or repciitive lithulogical nature 
should facilitate rather than cucumber further 
work, Such accurate lithologieal subdivision, 
coupled with thorough palaeontologieal 
evaluation, will result in a more lucid inter- 
pretation of the deposiiional history within the 
ba&in. Jt is with this intention, that the new 
formation names Woodendinna Dolomite and 
Wirrapowie LimeMone 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, discotiformabiy overlies the 
Bonncv Samisione Member of the Pound 
Quartzitc, and ii extensively dolomitizcd ai the 
1>3SC In general the unit is massive, pure and 
pale cream to pink coloured. II consists pre- 
dominantly of ooid and skeletal grainstnnes. 
which may he patchily silicified and lecrystal- 
Jurd. An abundant fauna js characteristic of 
the upper parts, some beds consisting almost 
entirely ot broken skeletal remains of arcbeo- 
eyathids. and associated braeh'topods. hyo- 
liihids. and trilobites. Algal Temains arc 
common throughout and although some 
cryplalgalaminatcs and oncolitc horizons are 
present at the very base, stromatolites arc vir- 
tually ahsem. Suhaerial desiccation features arc 
not represented in the type section Daily 
(1956, 1972o) considers the Wilkawillina Lime- 
stone 1o have hecu deposited on a stable shelf 
under shallow marine conditions. 
t'antra Limestone 

Flaggy to tubbly dark to black impure lime- 
stones with intcrbedded dark calcareous shales 
form the lower Member of the Parara Lime- 
stone m i he Wilkawillina graben. The Upper 
Member consists of uaggy and thin bedded 
dark jrrey to black aphanitic limestones and 
(bin calcareous .shales. Although scantily 
Ussilifcrous throughout, the Parana Limestone 
contains trilobites, hyolufiids, brachiopods, 
sponge spicules, evnehostracans and rare 
uichcocyathids (Daily 1 956). Stromatolites, 
ITarpebblc conglomerates and desiccation 
features are all completely absent. 
AjdX Limestone 

Daily (J 95b) proposed that the term Ajax 
Ltmcstont- should be used 'or all carbonates 



above the Pound Quan/ite W\ beneath the red 
elastics of the Billy Creek Formation in the 
Mt Scort Range area. Thrs comparatively thin 
carbonate sequence incorporates a number of 
different lithological types. Daily (1972b) has 
subsequently distinguished a 1 ower and Upper 
Member, and In view of the thickness of this 
unil, any further subdivision of the Aja.x Lime- 
stone may prove unpractical on k regional 
scale. It is very imponanc to realize, however, 
that a large number ol lithotypes present in the 
Ajax Limestone are present within the Cam- 
brian carbonate formations mapped elsewhere 
in the Flinders Ranges (Daily, pers. eomm.) 
and that the change in stratigraphic ter- 
minology in the Mt Scott Range area is in large 
part historical rather than geological. 

Proposed New Formations 
A significant propoition of the basal Cam- 
brian carbonate sequences, particularly those hi 
the north-east Flinders Ranges, are not readily 
assignable to any of the above Mrali graphic 
units. Although mapped c>n the COPLEY 
1:250.000 and parts of the PARACHILNA 
1 :250,tl00 geological maps as Wilkawillina 
Limestone, the carbonates show only scant 
similarity to Wilkawillina Limestone as defined 
in the type section. In part, these carhnnates 
show greater similarity with the Parara Lime- 
stone of the type section, hut crucial differences 
in Ikhology and depositionai environment are 
apparent on careful examination. It is for these 
distinctive and widespread basal Cambrian 
units that the new names WtwlenJittna Dolo- 
mite and Wirrapowie Limestone are proposed. 

Type Section 

The type section is located in Witrapowic 
Creek and one of its tributaries, about 14 km 
due east of Point Well Station homestead in Hie 
Central Flinders Ranges (Fig. 1). Unfortunately 
the section presents some difficulties with 
respect to access, which involves \S km of 
travel by four-wheel drive vehicle from a point 
on the Point Well-Narrina road to Wooden 
din mi Bore, and a further 2 km on foot alon.a 
a creek to the section. 'Ibis disadvantage is tar 
outweighed by the completeness of the 
sequence and the good exposure of both the 
Upper and lower boundaries of both forma- 
tions, 

The combined Woodcndinna Dolomite find 
Wirrapowie Limestone in the type section con- 
sist of 392 m of carbonate rocks which over- 
lie, with apparent amformtty, Diplorratcrion 



LOWER CAMBRIAN FORMATIONS, FLINDERS RANGES 



213 



MAP A 



WOODENDINNA DOLOMITE 
WIRRAPOWlE LIMESTONE 
TYPE SECTION LOCATION 



31 00 — 




PARARA LIMLSTONE 
WIRRAPOWIE LIMESTONk 
WOODENDINNA DOLOMITE 
HARACHILNA FORMATION 
POUND OUARTZlTL 
All facks shown are passable to fouf-wheel drive vehicles onlj 



MAP B 



— 31 00 




SOUTH AUSTRALIA 



■\ 1 A 



£ MAP B 



h\ 



J' 
^ { ) \ 

—I ! i i \ i 



PGM 75 



Fig. 1. Location of type section. 



214 



P. G. HASLETT 



TYPE SECTION woodendinna dolomite and wirrapowie limestone 



PARARA 
LIMESTONE 



392in 



-r-fWv-w^ 



^^Xk 



300m 



WIRRAPOWIE 
LIMESTONE 



I i 



'*^ VVyy Wv > 



200rn 




100m 



WOOOEN0INNA 
DOLOMITE 



m 



PARACHILNA 
FORMATION 




Very r«fti,fiiiy indued, .5tratiq"ly EiuttrappHg finC grtlifcd 
"ill iter tori as ^Ufi 1 itjnler coloured inalcy ir-tei-beJs . 
'jparsaly fossil i fet l ous and lacking dcsT^catiOi foafUfM 
and tldt-gelljlt: cciitjloniei aiOE,. 



Tina.- ijraincd riffij t« tii*c1v ?iity 1 iivestones .-ind 
LdTrfi^fftaua siit'jiu.iifMi, w.vrti very vtstjit far 3-'ij wr 

ii't'tdi"^ riapk fiiifiy lamim*tPi} a]qnl Uruno.Uj.l Tie 
£ed$ (5) 4ty I PP1T1 ru ! di' tld.-btfbble LUtinloiuei'titfs .ire 

cairj-ior,. I'li-io'r- cross-aedded utk ^nnstone beds (.3) 
PBJ.W* l.iii-oughnul. Sui iwn-i; ai'teofititti -letri tus is 
w—uriil," dbsi-nL j m the type section. Tire upperiros-t 
m jmiii-v -^ this ufi~ is spJicwfeS tjradati'crtal in*& Eli* 
i-'ARMA LIMESTONE iiftt3 r - Btyecd at Uw tawicst: al' : i*i 

stftuM-JftHM bed. 



Tellcw-We£^icrinc|, flatly to n,i>.sivn 41q1ut,;-qs 
4fi; loTonitie l iiaestotrES'i Wf? ujiiniiurw tloiSni cress- 
bedded qaafM SriivisMirti-. j;,), in!."j1 ut nntional 
conglomerate and ile^c^uLiail iflJilrtWfcS-. Auundan: 
•jtrouia*Ql1te beds .( = ), 0C]it.k 1'imafituncs (VI 
o:uir sda< ttw Up. 



MdSbive outuiiflpiHT] dull (pey dulaml L'7t."d 1 fl>itfii±EH*&. 
U^ally cry^UHine! prm..vy n«r>o-si ticrial ;eKti*T5 hw*«j 
difficult ta toUmnr.e nr tiic liieid. jHj4ra1 I1n«it«tifv 
wure u'^ticmtr>-av>t I _y cm'.:.-.- bedded air.d uralruunes 
■juiitiittn-mi mattered qujrcz s^rd ^r^nii (foSiiVf 
I a. Litr'ei "ly*'l inf.ed stroma to! T t&s are cuwui'- 



HlSSiVtt LU fl-l'liv yd, low-WEMtH(>rn".g JulOI'ilitflS 3-nd rffi Idi-IIMi.: 
limr^trn as. Tit WS^ h4 Ww 'aIKMITWIE UKC STONE tS 
rtp-jd^nily confonr^hla v, i Lh Lire itnrievtying pARAClltLflA 
HHMAilUN and is plat«»t: it tht* v-cla !.i vclj' dbnpt Kf&nttWtfiiiB 
<jf wpamitts wJira^'iU. flia :tfrbwwM i>-p rwruirGy 

■■:»',' kic o'Ui r.i v,| i L it, FUt-petiblS CWlgfte^ttt*, 
nu;-:racks <uH •.Ln>u,Lu' ftcs \i) »te a'laa -resent . 
Trm,. ■Gffiis-teddeu qyarU ^njattjnps (3; art imer'bedded 
will dcteiU-^ a-ncl aalitU i iituj-ptonsn in upusr parts. 



»it« is In biologically icieit-irra! »<-J\ U-nUurl .y'lrlo POUNt) 
[<'llAM71TE., but contains atsjnd,= nt Bj^gf|ipricn buri'ows. 
"Hiy uti'L Lucones red atttr^r.cras^intjn u ay -rich toward r.ha 
UiSi P' £l". ; rdluf n:n bur-jv-s bp^n^ present thf'f un^'i**- 
T-'u, uonskTs '.if rjroMi-fjrpy, poarly r;jtc.rop';1n'] Sflnie^a. 



Fig. 2. Type section. 



LOWER CAMBRIAN FORMATIONS, FLINDERS RANGES 



215 



ROCK RELATION DIAGRAM Woodendinna Dolomite and Wirrapowie Limestone 

(Not to scale ) 



Wirrealpa 




WEST 

Mt Scott 

Range 




WOODENDINNA DOLOMITE 



ehe 
€hd 
Ghr 
Ghw 
Ch, 
Ghp 
Gu 



Ewp 
Disconformity 



CAMBRIAN 
Nepabunna Sittstone 
Midwerta Shafe 
Parara Limestone 
Wilkawillina Limestone 
Ajax Limestone 
Parachilna Formation 
Uratanna Formation 

PROTEROZOIC 

Pound Quartzite 



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 




.- 

ft.; :: *^Miiln iiint^Plf** 7 "! 1 



'^*^to 









Figs 4-11, 



LOWER CAMBRMN FORMATIONS. FLINDERS RANGES 



2i: 



Stones (Fig, 6), Ripple maTks and chevron-type 
cross bedding are pieseni in (he ooid grain- 
stones and quartz sandstones (Figs $ and 9), At 
135 rn from the base, a thick cross-bedded 
quartz sandstone With abundant creamy yeJIow 
dolornitic mudsione cla^ts is developed. Toward 
the top of the Woodendinna Doiomite> ooid 
grainstone beds commonly show a primary 
wbbly or cobbled upper surface, reminiscent of 
that found in modern environments on area< 
of stihaemdly exposed, partially lithificd car- 
bonate sands (Fig, 7). 

Winapowie Limestone 

The hase of die Wirrapowie Limestone coin- 
cides with the relatively abrupt disappearance 
of dolomite in the sequence Flaggy fine- 
grained, dark grey, limestone and green grey 
calcareous ailtstones dominate the sequence. 
Desiccation features arc less ahundant but stro- 
matolites, intraformational conglomerates and 
minor cross-bedded ooid grainstones arc 
pieseni throughout (Fig. 11). Quartz sandstones 
are virtually absent, As in the Woodendinna 
Dolomite, stromatolites arc finely laminated. 
low, laterally linked domes which, on bedding 
plane exposures, show little if any preferred 
orientation fFig. 10). 

The Wirrapowie Limestone in the type sec- 
tion is overlain by Parara-type limestones. The 
topmost limit of the Wirrapowie Limestone is 
selected at the top of the uppermost algal 
stromatolite bed. Although a superficial simi- 
larity with the Wirrapowie Limestone remains 
above this limit, the sequence lacks ooid grain- 
stones, flat-pebble conglomerates and evidence 
of subacrial desiccation. The dark limestone* 
are far more strongly outcropping, purer and 
more crystalline above the contact. They also 
tend to he sparsely fossil if erous. with archeo- 
cyathids, hyolithids and various hraehiopods 
present. This fa in contrast to the underlying 
Wirrapowie Limestone and Woodendinna 
Dolomite which are notably depleted in fossils. 
Apart from alga! stromatolites, tbe only evi- 
dence of organic activity found to date are very 
minor irregular hurrows in some dolomitic 



rnudstones of the Woodeodmna Dolomite, and 
minor trails and apparent btotuibarjcm of bed- 
ding which may be found in impure limestones 
of the Wirrapowie Limestone. 

Dwribution 
The Woodendinna Dolomite and the Wirra- 
powie Limestone form the basal Cambrian car- 
bonate succession over much of the Flinder.v 
Ranges. They are best developed in the eastern 
part of the ranges, north of the line of outcrop 
of the Blinman, Wirrcolpa and Chambers db- 
pirx (sec Fig. 3). In this region their combined 
thickness ranges from 300 m to 500 m and they 
appear to thin gradually to the wesi, where 
comparable Jithologics are thinly developed in 
iht AJax Limestone at Mt Scott fDaily, pers, 
comm.) and the Andamooka Limestone in the 
vicinity of Yarrawurta Cliff (Wopfner 1 969; 
Daily 1972a). To the south, along the above- 
mentioned line of diapirs. the Wirrapowie 
Limestone thins very abruptly and is overlain 
by, and intcrlingers with, Wilkawillina Lime- 
stone. Syn- and post-deposttional tectonism 
along this line of diapirs. and the consequent 
development of complex local facies variations, 
has complicated the regional facies relation- 
ships along ihi.s line. Lateral intcrfingering of 
the Wilkawillina Limestone and the Wirrapowie 
Limestone- h strikingly demonstrated, however, 
in outcrops just north of the Wineatpa Diapir 
(Haslctt, in press). Abrupt lateral change south- 
ward from Wirrapowie Limestone may also be 
seen in tbe vicinity of Fregunda Creek. Iu addi- 
lion, Hatcher* indicates a similar southward 
thinning of a wedge of Wirrapowie-type lime- 
stones within the northern part of an area ol 
thickly developed Wilkawillina Limestone near 
M t Frome. 

In tbe north-eastern Flinders Ranges, the 
Wirrapowie Limestone is in all cases overlain 
by Pararn-type limestones. To tbe best, of the 
author's knowledge, Wilkawillina Limestone is 
absent from the north-eastern Flinders. Those 
rocks which are mapped as Wilkawillina Lime- 
stone in this area on tbe COPLEY 1:250.000 
Geological Map are assignable to the Wood- 



Fig, 4. Bedding plane exposure of stromatolites, Woodendinna Dolomite. 

£l g ' *' 2 7 oI i? n ? ,,e i c,ast ? m inlraform ational conglomerale. Woodendinna Dolomite, 

hig. 6. Well developed rmidcracks on the underside of a bedding plane, Woodendinna Dolomite. 

Ng. 7, Cobbled upper bedding plane surface of ooid erarnstone, Woodendinna Dolomite, 

hig. 8. Chevron-type cross beds in quartz sandstones, Woodendinna Dolomite. 

Fig. 9, Ripple marks cm bedding surface of a thin quartz sandstone, Woodendinna Dolomite. 

Mg. 0. Squat stromatolite bioherms on bedding plane, Wirrapowie Limestone. 

frig. It Impure flaggy-bedded limestones of the Wirrapowie Limestone. Note the lenticular agereealcs 
of uitra-formatKMial conglomerate. 



2IK 



P. O. HASIJtTT 



cndinna Dolomite and Wirrapowie Limestone 
in Ibi! lower pari and obviously hitherto undif- 
ferentiated, and in the upper part can be much 
more justifiably related to Parara Limestone 
Itthologies than those of the Wilkawillina LiuKV 
stone. 

Thin but well developed Woodendinna Dolo- 
mite does oceiir south of the line of the Klin- 
man-Wirrcalpa-Chamhers diapirs along the 
southern limb o( the westward plunging ami- 
dine 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 
Ranges, the Woodendinna Dolomite rapidly 
thins in response to erosion, or nondeposition 
or both. At the Ten Mile Creek type section 
itself, Wilkawillina Limestone disconformably 
overlies the Bonncy Snndstone Member of the 
Pound Quart2ile (Dalgarno 1964; Pierce' 1 ). 

I he very highly dolomitired basal Wilka- 
willina Limestone in this section, however, 
superficially resembles parts of the Wonden- 
tfinns* Dolomite, but due to the degree of dia- 
genesiv, the original nature of these basal car- 
bonates has no! been resolved. For convenience 
they are considered at this siage to represent 
dolomitized Wilkawillina Limestone litho- 
facics. 

Tn the east-central Flinders Ranges Gcnling* 
describes approximately 25 m of carbonates, 
which are tithologieally similar to Ihuse of the 
Woodendinna Dolomite, from the base of the 
Cambrian carbonate sequence at Reap hook 
Hill. 

Along the entire western outcrops of the 
Cambrian sequence* from I'arachilna Gorge 
south, a thin sequence of flaggy grey lime- 
stones, dolomitie mudstones, stromatolites, nat 
pebble conglomerates, and imcrbedded sand- 
stones, occur beneath typically fossiliferous 
Wilkawillina Limestones'. These beets are 
assignable to the Woodendinna Dolomite and 
the Wirrapowie Limestone and roughly cor- 
respond to what Dalgarno called the Lower 
Member of the Wilkawillina Limestone (DaJ- 
garno 1964). although as staled above, these 
lilhologics are not present within the defined 
type Wilkawillina Limestone, Dalgarno & 
Johnson (1963) report significant thicknesses 
of Wirrapowie-type limestones from jus! north 



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 ha* confirmed the presence of both 
Woodendinna- and Wirrapowie-type carbonate 
sequences in these southern and south-western 
parrs of the Flinders Ranges (Daiiy, pers. 
comm.). 

Depnsitinnal F.nvironraenl 

From the above descriptions, it is apparent 
that although the Woodendinna 1>olomiie and 
Wirrapowie 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 
anuthcr both temporally and spacially and 
clearly have formed in closely related deposi- 
tional environments. 

The Woodendinna Dolomite xhowi- 
numerous features characteristic of deposition 
on emergent high intertidal and suprMidal 
mudflats. Prevalent desiccation muilcracks 
record repeated periods of suhaeriaJ exposure, 
ami the highly restricted nature of the environ- 
ment is suggested by the paucity of fauna and 
dominance of primary dolomhic mudstones or 
pene-eontemporanoously dolomitized car- 
bonate mudstones, The only organisms capable 
of tolerating the extremes of restriction and 
desiccation of the environment were algae, as 
evidenced by the ahundance of stromatolites 
in the sequence. 

Cross-bedded ootd grainstones and quartz 
sandstones record brief episodes of high energy 
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 .sheets 
of coarser elastics from more open marine 
areas, over the exposed mudflats. Quart/ sand- 
siones Sn the Woodendinna Dolomite show con- 
siderable variation in iheir lateral development, 
reflecting the local availability of w qnariz sand 
source, which in many cases probably cor- 
responded to areas of exposed, unlithif*ed 
Pound Quartzitc. 

The Wirrapowie Limestone has developed in 
a sheltered but less restricted upper intertidal 
to lagoonal environment. The absence of dolo- 
mite and the rarity of desiccation mudcracks 
suggest that only limited exposure and restric- 
tion prevailed. The sequence lacks coarse detri- 



B Pierce. P. R. 11*691. — "the Cumbrian ecology south of thr Wirreatpa Piapir. Flinders Ranges South 
Aitttralin. Honours Thesis, Univ. Adelaide ^unpublished). 



LOWER CAMBRIAN FORMATIONS, FLINDERS RANGES 



219 



tus, current bedding and any preferred elonga- 
tion of stromatolite bioherms indicating that 
energy conditions were in general still very low. 
As described above for the Woodendinna Dolo- 
mite, however, thin grainstone sheets attest to 
the periodic penetration, probably during 
storms, of high energy conditions into the shel- 
tered tidal mudflat environment. 



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. Dalgarno who 
read a draft of this paper and offered valuable 
criticisms. 



References 



Coats. R. P., Callen, R. A., & Williams, A. F. 
(1973).- COPLEY map sheet, Geological 
Atlas of South Australia, 1:250,000 Series. 
(Geol. Surv. S. Aust: Adelaide.) 

Daily, B. ( 1956) — The Cambrian of South Aus- 
tralia. In HI Sistema Cambrico, su Paleoeeo- 
grafia y el Problem de su Base 2: 91-147. 
(xx Congresso GeoL Tnternacional, Mexico.) 

Daily, B. (1972a).— Aspects of carbonate sedi- 
mentation in the Cambrian of South Aus- 
tralia. Abstracts, Joint Specialists Groups 
Meetings, Canberra, GeoL Soc. Aust. cl0-cl4. 

Daily. B. (1972b).— The base of the Cambrian 
and the first Cambrian faunas. Centre for Pre- 
cambrian Research, Univ. Adelaide, Spec. 
Hap. L 13-41- 

Daily. B. (1973).— Discovery and significance of 
basal Cambrian TJratanna Formation, Mt 
Scott Range, Flinders Ranges, South Austra- 
lia. Search 4(6), 202-205. 

Dalgarno. C. R. ( 1964).— Lower Cambrian 
stratigraphy of the Flinders Ranges. Trans. R. 
Soc. S, Aust. 88, 129-144. 



Dalgarno, C. R. ? & Johnson, J. E. (1962).— 
Cambrian sequence of the Western Flinders 
Ranges. Quart, geol. Notes, geoL Surv. S. 
Aust, 4. 

Dalgarno, C. R., & Johnson, J. E. (1963). 

Lower Cambrian of the Eastern Flank of the 
Flinders Ranges. Quart, ^eol. Notes, geol. 
Surv- S. AusKl. 

Dai.garno, C. R m & Johnson. J. E. (1966).— 
PARACHTLNA map sheet, Geological Atlas 
of South Australia, 1:250,000 series. (Geol. 
Surv. S. Aust.: Adelaide.) 

Haslett, P. G. (1975, in press).— Lower Cam- 
brian stromatolites from open and sheltered 
intertidal environments, Wirrealpa, South Aus- 
tralia. In M. R. Walter (ed.) "Stromatolites". 
(Elsevier: Amsterdam.) 

Johns, R. K. (1968).— Geology and mineral 
resources of the Andamooka-Lake Torrens 
area. Hull. geol. Surv. S, AusL 41, 1-103. 

Wopfner, H. (1970).— Early Cambrian Paleogeo- 
graphy, Fromc Embayment. South Australia. 
Bull. Am. Ass. Petrol. GeoL 54(12). 2,39*- 
2.409. 



THE SOUTHERN AUSTRALIAN SPECIES OF SPYRIDIA 
(CERAMIACEAE: RHODOPHYTA) 

byH. B. S. Womersley* & Sally A. Cartledge* 



Summary 

WOMERSLEY, H. B. S., & CARTLEDGE, Sally A. (1975).- The southern Australian species of 
Spyridia (Ceramiaceae: Rhodophyta), Trans. R. SOC. S. Aust. 99(4), 221-233, 30 November, 1975. 
Four species of Spyridia are recognised on southern Australian coasts. S. filamentosa (Wulfen) 
Harvey (including S. biannulata J. Agardh, S. breviarticulata J. Agardh, and S. spinella Sonder) is 
common in sheltered to moderately rough water. S dasyoides Sonder (including S. opposita Harvey 
and S. prolifera Harvey) is fairly common on rough-water reefs and in deeper water. S. squalida 
J. Agardh (including S. wilsonis J. Agardh) is a less common, usually deep-water, species. 
S. tasmanica (Kuetzing) J. Agardh occurs in relatively calm localities but where there is often a 
strong current. The Australian species differ in vegetative aspects such as arrangement and diameter 
of the ramelli, but agree well in the development of the thallus and reproductive structures, and 
emphasize the generic uniformity of the species ascribed to Spyridia. 



THE SOUTHERN AUSTRALIAN SPECIES OF SPYRIDIA 
(CERAMIACEAE: RHODOPHYTA) 

by H B. S. Womersley* & Sally A. Cartledge* 

Summary 

Womersley, H. B. S. t & CumtooK, Sally A. (1975). — The southern Australian species of 
Spyridia (Ceramiacejie: Rhodophvtn). Trans. R. Sox:. S, Attst. 99(4), 221-233, 30 Novem- 
ber, 1975. 

Four species of Spyridia are- recognised on southern Australian coasts. $, fiiantwtosa 
tWuifen) Harvey (including Si bianmdata J. Agardh, S. breviarticulata J. Agardh, and S. 
spinefla Sender) is common in sheltered to moderately rough water. S dasyoides Sonder 
(including S. opposita Harvey and S. proVtfera Harvey) is fairly common on rough-water reefs 
and in deeper water. S. squalida J. Agardh (including S. wilsonis L Agardh) is a less common, 
usually deep-water, species. S. tastnanica (Kuetzing) J. Agardh occurs in relatively calm loca- 
lities but where there is often a strong current. The Australian species differ in vegetative 
aspects such as arrangement and diameter of the rarnelli, but agree well in the development 
Of the thallus and reproductive structures,, and emphasize the generic uniformity of the species 
ascribed to Spyridia. 



Introduction 

While Spyridia is a distinctive and easily 
recognised genus of the Ceramiaceae, the taxo- 
nomy of the southern Australian species has 
been confused. Some 1 species have been 
credited 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 or 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 its 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 general 
morphology, presence of nodal and internodat 
cell bands, branches of limited growth (ramclli 
or * H brachyblasts") and unlimited growth, and 
in reproductive features. However, the features 
which Hommersand (1963, p. 177) used to 
distinguish 5. fdamentosa from other species of 
Spyridia do not apply satisfactorily to the Aus- 
tralian species. Cortication by tiers of nodal 



and internodal cells is found in all species (the 
cell shape varies considerably m S, fdamen- 
tofin), and all species have radially disposed 
rarnelli (opposite in S. dasyoides) which (ex. 
cept 5. 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 
rarnelli 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 a band around the 
node between two axial cells. Each of these 
periaxial (nodal) cells cuts off two cells from 
its lower end. and these elongate and become 
attached by pit-connections to th$ nodal cells 
of the next lower segment. Thus the thaUun 
shows bands of shorter and broader nodal cells 
alternating with the bands of iutemodal cells. 



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

t This term is used for cells cut off from, but of different form to, an axial cell, Cells of similar form 

to the axial cell, as found in the pofysiphonous families of the Ceramiales. are still referred to a* 

per [central cells. 



222 



H- R. S. WOMERSLEY & SALLY A. CARTLEDGE 



which arc longer narrower, and approximately 
twice as many as the nodal cells, Purine! cor- 
ucation occurs some distance from the branch 
apices, from descending rhizoidal cells deve- 
loped from the nodal cells, anil this ohscurcs 
the regular pattern of nodal and internodal cell 
hands, especially in certain species (e.g. S 
•qaallda). 

The cells uf (he ramelli each cut off a ring of 
6-8 celts from their upper end. and these deve- 
lop into a nodal band 1-3 cells broad in S. 
(ihmwntosa. Spyridhi is readily recognised by 
the alternating nodal and internodul bands, and 
ramelli with corticated nodes. 

Ueproductively Spyhdia is also distinctive, 
especially in (but the carposporophyte becomes 
surrounded, by periearpic filaments developed 
from the segments above and helow the one 
bearing the procaip These filaments can eive 
the appearance of a eystocarp with a well deve- 
loped pericarp wall heint; held together by a 
mucilaginous sheath and some lateral pit-con- 
nections, but ihey disintegrate fairly readily in 
preserved material. 

Procarps (V-6) are produced on small 
lateral branchlets of rcslricied growth, Accord- 
ing lo Hommersatul (1%3. p 191 .), three peri- 
central (periaxial) cells are normally formed 
in each fertile segment, one of which (the sup- 
porting cell) bears the earpugonial branch, and 
each pericentral forms an auxiliary cell Hom- 
mcrsand reported that the carpogoniurn fuses 
with the third cell of the curpogonial 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 carpogoniurn divided into two 
cells, each of which fused with an auxiliary 
cell. 

Spermalaiigia cover several cells in the lower 
pan of the Tamelli, usually excluding the basal 
cell. Thev are derived from filaments originat- 
ing from' the "odal cells, which grow over the 
two adjacent cells, then cut oif spermatangial 
mother ecu's before forming the continuous sur- 
face layer of spermatangia. 



Tetraspornngia occur on the lower ceils of 
the nunelli. being sessile and mostly on the 
upper (adaxtal) side. Hommersand (196$, p. 
196) considers that they arise directly as pro- 
trusions from the axial cell, while Krishn;,- 
munhy i IU6S, p. 47) considers thai they arc 
formed from periaxial cells- 
Some 10 species of Spyndia have been 
recorded from southern Australia. This study 
recognises only four species, including S. fikt- 
mentoxa. The thallus development ajid mor- 
phology, and the structure of reproductive 
organs, are very similar to those of S. filtt- 
mvntosd* and the Australian species differ 
mainly in vegetative features. 'I he descriptions 
below are therefore confined largely to recog- 
nition of the species, with brief notes only on 
reproductive details 

Key to southern Austrian species 

]. Ramelli robust, opposite and decussate, usually 
100-150 /mi thick with i»H(Ji;»metTic cells and 

nodal bands 3 5 cells broad 

& tkuyoidt'.s (p, -31 ) 
V Ramelli slender, -single or whorled but not oppo- 
site, less than 70aiu thick, usually with cells 
longer than broad, imd nodal band* 1-3 oclls 

broad • , ,. ( 2 

2. Ultimate bmnchleis stout (t-l mm thick), 
markedly basatly constricted, hi?avily corti- 
cated tn their apices, bearing slender, irregu- 
larly branched ramelli . S. squub'da <P- 225) 
2. Ultimate branch let* slender < under '• aim 
thick), not or only sliehily hastily constricted, 
corticatinn only on older branches, with 
ramelli eirher one per segment or veuualluic 

J; Ramelli one per segment, 3."5-t>5 /urt thick, nodal 
hands 2-t cells broad S. glshieitt&m <P- 222) 

3. Ramelli becoming Veflfcllfetfi <!-/•), 2»M0 Mn 
thick, nodal bands t cell broad 

S. rusmmiiva (P 2*7 1 

Spyridia fijarmnlasa (Wuli'cn) Harvey 1833: 
336: 1844: 449; WW*, pi- -*6: W3f= **?! 

1859: 329; 1863. svnop.: 42. J. Agardh 
IK52: 340; 187b: 2b«: t897: 13. Boerge- 
scn 1917: 223, figs 222-226. Pddmann- 
M;.7.oycr 1940: 348. Guiler J9S2: 98. 
Hommersand 1963: 177. fig$ 4-10. 
Hooker & Harvey 1K47: 409. Krishoa- 
muTthv 196K: 42. Newton 1931: 394. fig. 
1^6. Okamura 1932; 130. ReinboUl 1X97: 
60. Sondcr 1853: 680; 1880: 16. Tate 



FiE I &9ttfamm**te&. 4. Pt Sranvae. S. Atist. (Uwh. MaMfc^ 

B * plant with vo.ing cystoearp . ft Denison. W. A*tf. Kraft, l4:o..W1: ADO. Mil flW.« . MuJc 
plant with "sperLtangial ramelli f&lfai, S. Ausl. CWlMft 30 -" 1 W2; ADU - **»»! °- 

Tetrasporangia! plant (A41KI5V 



THE SOUTHERN AUSTRALIAN SPECIES OF SPYRWIA 



223 






■ B-D , 



# 300/um 









* ft 



f 














Fig. 1 



224 



H. B. S. WOMERSLEY & SALLY A. CARTLEDGE 




THE SOUTHERN AUSTRALIAN SPECKS Oh SPYRIfUA 



225 



1 832: J 8. Tisdall 1898: 505. Wilson 
1692; iSl Wornersley 1958: 157, 
Ftwuf fxUovctitaws Wulfcn 1803: 64. 
Jf, fftvmtMtim VflPi urhtixcuta Sender 1855; 518. 
£ btamHtfote J. Agardb Jg76: 267; |gfe: 13. 
De Toni 1903: 1426. Guiler 1932; 98. Lucas 
190$: 52: 1929a: 25; 1929b: 53 Lucas & Perrin 
1947: 363. May 1965: 369. Oltamura 1932: 
130. Retnhotd 1897: 60; IS99; 50. Sender I8S0; 
16. Tadall 189ft stXV Wiison 1892: I8L 
Wornersley J950: 180. 

.V. hrrviarUttttaUi J. Agardh 1876: 268; 1897 
13. Dc Toni 1903: 1427. Guile* 1952: 98. 
Lucas. I9U9: 52; 1929a: 25; I9&&! 53. Lucax & 
Pen in 1947; 363 May 1965: 369. Osamura 
1932: 130. Reinbold 1897: 60; 1890: 50. Sonder 
1**0: 16. 

i. stnneUa Sonde r 1845: 53: 1846 168; ISS0- 
J6. J. Agardb 1852- 342; 1876: 269; IH97 13 
He Tooi 1903; 1430. Harvey |B63, SVflOB.! 42. 
KuctzJng 1*49; 668; 1862: 16, pi. Sled. Lucas 
1909: 52. May 1965: 369. Mzzzu 1925. no. 824. 
Okamum 1932: 130. 

FIGS I, M.B 
Thatlui (Fig. M) usually 7-18 cm high, 
epilithic or epiphytic on various larger algae 
and scagrasses. lax and soft, irregularly much 
branched on all sides with longer and shorter 
branches intermixed, with one to several axes 
(often poorly defined) from an originally dis- 
coid holdfast, soon becoming fibrous or stoloni- 
tcrous and entangled, commonly grey to grey- 
red, sometimes red-hrown, in colour. Axes and 
larger branches corticated, terete; axes i-1 
t— If) mm thick, tapering to branches 300-500 
/ntO thick and hranchlets 100-300 ,/m thick; 
laterals arising from periaxial cells or advents 
liously from cortical Cells. Segment* usually 
clearly defined on branchlcts (Fig. J/?), vari- 
able in length and proportions but usually 
(i— H-l times a.< long as broad, with bands of 
shorter nod^l cells and longer internodal cells 
alternating; nodes with 11-14 periaxial cells, 
each corresponding to two internodal cells ex- 
cept for the (.usually) larger periaxial eel) bear- 
ing the famellus. Cortication usually commenc- 
ing ■ few mm from the apices but very 
variable, consisting of rhi/oidal cells lying 
between the internodal cells and gradually 
forming a continuous cortex l(-2) cells thrck\ 
Raoidh (Figs }B t 3A, B) single per segment, 
irregularly spirally arranged, j-j* mfn ~ | on g 
with 12-201-27) cells, linear or gently taper- 
jng apart from the terminal 2-3 very short 



cells I Fig. 3 A) which taper abruptly to a 
mucronate cell, f 35-) 4f>-55<-65 > pn\ thick 
with cells (I J-) N-241-3) times as long as 
broad; mucronatc end cell often Inst from 
older ramelli. ramelli with about 9 nodal cells, 
each usually cutting off 1 (-2> cells anleriortv, 
giving a nodal band 2-3 cell* broad. 

Cys/ocarps (Fig. \B) short-stalked, usually 
bilobed, lobes globular, 300-700 pip across. 

Spermutangia covering the lower (except 
basal) several segments of ramelli (Fig. IC). 
forming maie organs 75^120 jam in diameter. 
letrasporanfjia (Fig. ID) sessile, 1-3 per 
cell on lower cells of ramelli, mostly on the 
upper (adaxial) side, spherical, 50-75 ,»m in 
diameter, fctrahedrally divided, 
Type locality; Adriatic Sea. 
Type: 1 

Distribution; All around the Australian coast 
(including. Tasmania) in conditions of mod«r- 
atc to slight water movement. 

Spyndia fifonwitosa is recognised as a 
widely distributed species, having been 
recorded from most seas, and manv authors 
(e.g. Harvey 1846, pf. 46; Felduiiinn-Mazoyer 
1°40, p. 348) refer to h as a very variable 
species J. Agardb (1876, p, 268) in segregat- 
ing two Australian species [S. hUmwtota and 
S. hreviar/jculata) from 5. filamentosa, referred 
to their similarity in habit with Si fitamentoxa 
and the large number of forms classed as this 
species. J. Agardb apparently regarded his two 
segregate species with some doubt, and a de- 
tailed study of extensive collections of Ausira- 
lian material does not provide any satisfactory 
way of segregating S. binnnukita and 5". brevi- 
art'wulQia from S. filamentosa. 

The type of 5. bimmulata is from Tasmani;. 
(Georgetown, Tav. Gtwn. LD, 51300, selected 
as lecto(ype) and was distinguished by J, 
Agardh in having more conspicuous bands of 
nodal and internodal cells and being less cor- 
ticated above. Si breviarticttlaiu is based on 
specimens from Whitsunday T., Queensland 
(lectotype in LD, 51311). and was distin- 
guished by having the nodal and internodal 
bands shou and of almost equal length. The 
variation in S, filammtosa encompasses the 
above features of both S. buttmutata and S 
hwviaeticHfaUt. 



Fl * % SESfettSP** A ForuuHngtott, Vic. (Woilastrm, I7.vlli.1956; AM), A20S67) B Femak- 

SEK h%^f^A^^ri^^ ".melll (Taplcy Shoal. % A„s, , 15 m % W 
rtwot —n.ivnv, ajju, Aj>>38). D. Tetrasporangial plant (A4fQ$gt, 



226 



H. B. S. WOMHRSLEY & SATXY A. CARTLEDGE 




Pig 3 4, B. C Wumentout. Apex and mid region oC ramellus, with periaxial cells in face view 

(A41815). , fli , n ^ 

C\ O. 5. wsmanica. Ditto (A41066). 

jSjf£' 7 885 ADUT A35287>. c.br.-carpofipnial branch; p.a.-pcnax.al cell; 8 .c.-suppw t- 

flf f\ snualida. Apex and mid region of lamellus, with periaxial ceils in face view 

(A26375). 

/, K. S, tla^oides. Ditto (A20I70) 



THE SOUTHERN AUSTRALIAN SPECIES OF SPYRID1A 



227 



S, filamentosa var. arhuscula 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, 15.1.1951; ADU, A 15142) agree well with 
the type in having densely aggregated ramelli 
400-800 yxm long, composed of 15-18 cells, 
50-65(-90) fim thick and 1-H 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.viii.1973; ADU, A43938). 
However, material from Brest, France 
(Cabioch, Dec. 1972; ADU, A43050) has dis- 
tinctly more robust ramelli (about 100 ^m 
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.iii.l972; ADU, A4188I) 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 



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, pi. 42 c, d. 

Sonder 1853: 680. 

S. filamentosa var. tasmanica Kuetzing 1849: 
666. 

S. filamentosa var. verticil lata Harvey 1 844: 
449. 

Wrangelia setigera Harvey 1859: 309. pi. I9M; 
1863, synop.: 27. J. Agardh 1876: 622; 1879: 
pi. 32, fig. 3. De Tom" 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, AA-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 
l-li(-2) mm thick, branches about £ mm 
thick, lesser branchlets 200-400 ^m thick. Seg- 
ments (i-)J-H 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) l(-3) per node near apices, becoming 
whorled (3-6(-8) per whorl with the addition 
of adventitous ramelli), arising from enlarged 
periaxial cells, £-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 
( 1 2-) 1 5-20 cells long, greatest diameter 
30-40(~45) ^m and l£-2i 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, i-£ mm in 
diameter. 

Spermatangia cover 2-6 cells (Figs 2C, AB) 
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. 



22X 



H. B. S. WOMERSLEY & SALLY A. CARTLEDGE 




Fig. 4. 



THE SOUTHERN AUSTRALIAN SPECIES OF SPYRIDIA 



229 



Tetruxporanxia (Figs 2D. 4C) on 1-4 cells 
near the base of the ramelli, borne mostly on 
the upper (adaxial) side, 60-100(-120) M m in 
diameter when mature, tetrahedraliy divided. 

Type hiatity; Tasmania (probably Cmnth ex 
Hooker I 

Type: l (941, 3ti...37n. 

Distribution; From Elfiston. S. Aust. lo 
Western Port, Vic. and aiound Tasmania. Gen- 
erally in relatively calm localities, often wilb 
cuiiNtdcrablc current. 2-35 m deep, occa- 
sionally in partly sheltered habitats and shaded 
tuck pools on tough-water coasts. 

Development of reproductive structures in 
S. itisniattica appears to be very similar to that 
in 5. ftlamentaw. The female axes (Fig. 3G) 
arise as short laterals and bear up to 5 pro- 
carps, separated by one or two sterile segments 
each with eight periaxial cells, one of which 
hears 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 
not been clearly followed, but usually two 
groups of gonimohlast cells develop, probably 
from fusion cells originating from auxiliary 
cells cut off from the supporting cell and one 
of the other ferule periaxial celts. Sterile peri- 
carp filaments arise from the periaxial cells of 
segments above and below the proearp-bearing 
segment, forming a cysfocarp similar to that in 
S fiiamettfoxft. 

Development of spermatanvia and tetraspo- 
rangia is similar to that in S. fifamentoxa. 

S, fasmanica is a distinctive species with its 
v* hoi led ramclli and single row of nodal cells. 
The ramelli are typically slender (20-35 /A m 
thick), with matuie cells 3-5 times as long as 
hroad. However, some plants from rougher- 
water localities |e.g. Elliston, S. Aust., 7 m 
deep (Shepherd, 20.x. 1970; ADU, A37622) 
and Cape Lannes, S. Aust., in shaded pool 
I Krttft, 12.ii. 1972; ADU, A41S09 ) | have 
more robust ramelli, 30-40f-45) yjix thick 
CFifiS 3£. F, 45, C) and mature cells 1 t-2i 
times as long as broad. Otherwise the latter 
specimens are similar to the majority of plants, 
and the type, of & tasnumha, and the more 
robust* shorter-celled rametli are regarded as 



more characteristic of plants found under 
rougher-water conditions. Flutter studies on 
this form are. however, desirable. 

Spyridia squalida ), A*»ardh J 876: 270; 1897; 
16. De Toni 1903; 1436. Lucas 1909; 52; 
1929b: 53. Lucas & Perrin 1^47; 364 
May 1965: 369. Okamura 1932: 130. 
Reinbold 1897; 60. Sotukr IKKO; 16. 
Tate 1882: 18. 

$ wiJsortisJ. Agardh 18^7: 16, De Toni (90S: 
M35. Lucas 190V; _S2. lue;is ft Perrin 1941 
3c»4. May 1965: 396". Okamuia 1932: 130. 

•V. KttlkUi Sontlrr 18X0* 16 (uomen nudum). 

FIGS 3//. /, 4 A /. 

ThaUus (Fig. 40) usually 10-30 cm high, 
robust, erect, irregularly and proliferouvly 
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 in colour and when 
dried Often appearing somewhat farinaceous, 
All branches terete and corticated to thcij 
apices, axes and main branches linear, branch- 
lets (Fig. 4E) basally constricted and bearing 
densely arranged ramelli, especially on their 
upper parts, sometimes denuded below. Axes 
li-2i mm thick, denuded below or with short, 
proliferous branchlcts, tapering slightly in 
branches 1-1 f mm thick and lesser branchlcts 
4-1 mm thick. Segment* largely obscured by 
eorticatiou, \b**& as long as broad, with 16 
periaxial cells and about twice as many inter- 
nodal cells; eortication commencing within a 
few axial cells of apices, pseudo-parenchyma- 
tous, 2-3 cells thick on branchlcts, several cells 
thick on axes. Ramelli (Figs 3//, /, 4E) one 
per segment close to apices and derived from 
periaxial cells, but scattered adventitious 
ramelli (usually) densely cover the hnmehlcts. 
sometimes persisting onto larger branches; 
ramelli *-1(-li) mm long with (IO-)I4-20 
(-24) cells of greatest diameter (20-) 30 40 
(-45) pzh and (1-j I J-2(-2J ) limes as tone 
as broad. Ramelli with a single row of small 
nodal cells (Fig. 31) derived from 5-6 peri- 
axial cells each of which cuts off 2-3 outer 
cells in the same transverse plane. 

Cystocarps short-stalked, globular-bilobed, 
\-i mm in diameter 



1% 4. Spyrun tasmanca (robust form . A. Female plant with quashed mature cvstocarp (EUislon 
S. Aust., J m deep in bay. Shepherd, 20.X.I970; ADU, A?7ft2?). B. Male plant wiUi sperm 
langud ramelli (A37622). C. Telrasporangbl plant (A3TO2) P 

Spyndta sxjualidti D. Victor iLirhor, S. Aust. iWotnerslev, iK.iii iVfcft; ADU UOtmi K 
Branchlcts and ramelli (Pt Elltol. S. AnM- Dodd, J2.iii.I463; ADUi A26575J 



230 



H. B. S. WOMERSLEY & SALLY A. CARTLEDGE 




THE SOUTHERN AUSTRALIAN SPECIES OP SPYJUDlA 



23 1 



Sperrnatattgia cover the lower 2-6 cells 
(except basal cell) of ramelli, forming a male 
organ 50-1*0 plti in diameter, 

Tetraspvrangia borne on the iower several 
ceffs of tbe ramellK largely on the apper 
(adaxial) side. 1 (— 2) per cell, sessile, spherical 
to slightly ovoid, 40-60 pm in diameter when 
mature., tetrahedralfy to sub-cruciately divided. 

Type locality: "Nov. Holland, australem". 
Type: Herb. Agardh. LD, 51533. 
Distribution; From Geographe Bay, W, 
AusL to Waoatafi Bay, S. Gippsland, Vic., 
usually in deep water i 2-24 m deep) . 

Female axes of 5. squckluto develop as short, 
adventitious branchlets which are more heavily 
-a r f , ;:.it. d than in other species but less so titan 
in vegetative branchlets of this species, Alter- 
nating segments each bear a procarp, with the 
sterile segments be^Ting ramellr. Usually three 
periaxial cells occur in fertile segments, one 
line supporting cell) producing a 4-oeUed car- 
pogontal branch. Two, or probably ofien 3. 
auxiliary cells are formed, leading to >i carpo- 
sporophyte with two or three lobes. The pen- 
carp develops similarly to that in other species 

S. squaUda is a distinctive and robust species 
of Spyridia, having cortication to the apices 
and thus forming swollen, basally-constrided, 
branchfets, bearing ramelli often densely scat- 
tered hut usually soon denuded. The farina- 
ceous appearance is also a common feature of 
older, dried plants 

S. wiltot\ii J. Agardh is typical S. sqttaJida, 
The type of the former is from Pt Phillip Hds, 
Vic {J. B. Wilson, 1857; LD 51532), and tbe 
thallus is not compressed ns stated by J. 
Agardh ( 1-897) and May (1965V 

5. vul'ulu Sonder (1880: 16) is a nomen 
nudum, based on a specimen in MEL (45195) 
from Geographe Bay. W. Aust. (Bunbury. 
IH75), accompanied by Sonder* drawings. It 
is typical S. sauaiida. 

«Y. xtjuulUh was recorded from Port Alfred 
(Kowie), Souih Africa by Barton (16%, p. 
?96>, and the record repeated bv De Toni 
(1903, p. 1436) and Lucas & Porria (1947, 
p. 364), This record almost certainly applies 
to some other species, probably to S. pfamosa 
Schmitz ex J. Agardh (G. F. Papenfuss. pers. 
comm.1. 



Spyridia dasyoldes Sonder 1853: 680; 1880: 
16. J. Agardh 1876; 272. De Toni 1903: 
1437. Harvey 1863, synop.: 42. Lucas 
1904: 52. Lucas & Pcrrin 1947; 364. Mav 
1965; 369 Okamura l ( >32: 130. Tate 
I8S2: l«. Tisdal! 1S98: 505. 

S, oppoxha Harvey 1855b. 256; I860: pi, 15*. 
Adi.ms. 1972: S3. J. Agardh 1876; 270; 1897: 
H De Toni 1903; 1431; 1924: 502. Guile* 
1952: 9fc. Lucas 1909: 52; 1929a: 25; 1929b- 
53 Lucas * JVrrin 1947* 363. fig, 1*2. Mav 
1965: 369 Mama 1912, no. 421. Okamura 
1932: 130. Reinboid 1897: 60. Shepherd & 
Womersley 1970: 135. Sonder ISSOr 16. Taie- 
IS82: 18. Tiwlall 1X98 505. Wilson 1892: 1*1. 
WnnwNley 1950: 180; 1966; »51. 
5. pralifera Hnrvcy 1863: (A. 274. J. Agardh 
1*76: 269; 1597: 14. De Toni 1903: 1431. 
Lucas 1909- 52. Lucas & Perrin 1947: 362. fie. 
Ifil. Mav 1965. 369. Mazza 1912. no. 422. 
Sonder 1880: 16. 

FIGS 3i, A', 5 

Thallus (Fig. SA-C) usually 10-20 cm high. 
eroc(, eprlithic or epiphytic, much branched 
with one to several axes from an originally div 
coid holdfast which soon becomes fibrous and 
stoionrferous, dark red to red-brown in colour. 
Axe* and larger branches heavily corticated, 
terete to angular and becoming four-Mdcd with 
thickened cortical flanges in line with the 4 
ranlcs of ramelli, densely branched. Axes 1-2 
(— 2i) mm thick, often denuded but sometimes 
with numerous, short, proliferous branchlets, 
tapering to branches }-£ mm thick and lesser 
branchlets k-\ mm thick; branching usually 
subdistichous (Fig. $C) with laterals arising 
from nodal cells. Segments largely obscured by 
cortication, Kl times as long as broad, with $ 
perineal cells producing 16 intemodal cells 
and the 8 cells soon with interposed rhteoidal 
cells giving both nodal and mternndal rings of 
16 cells (Fig. S/)i; cortication commencing 
within a few segments of apices, of clongalc 
cells later appearing pscudo-parenchymatoui, a 
few cells thick on branchlets, many (especially 
on flanges) ccih thick on axes. Ramelli (Figs 
3/, K* 5£>, E) arising from an enlarged peri- 
axial cell, in opposite and more or less decus- 
sate pairs (Fig. SD) on successive segments 
(often displaced to two rows on each side in 
the plane of branching), n-}|*-2(-2*) mm 
long with (16-HX-22 cells, relatively uniform 



Fig. 5 Spyridic desyoides. A Holotype (MEL, 45128). B. Robe. S. Aim. <CartIcdge t i4.v,J^72; ADU, 
A42I74)— an irregularly branched form C_ TrivesUcalor Strait, S. Atist., 43 m deep \Watson\ 
27.U97I. ADU, A38143)— distichously branched form, D, Branch showing arrangement of 
ramelli i Pt Oemson, W. Ausi. Kraft, I4.xii.1971; ADU, A4I7301 E Ramelli with tetra- 
sporangia (Vtvonne Buy, Kangaroo I-, S. Au»i. Wamtrstey, 30.1.1956: ADU, A20170) 



232 



H. B. S. WOMERST.EY & SALLY A. CARTLLDOE 



in diameter and tapering fairly abruptly to a 
point, (70-) 100-1 50 ^m thick, cells about 
l( H) limes as long as broad. Ramclb with 
16-20 nodal cells, each cutting off 1-2 cells 
(which otren divide again 1 on both skies (an- 
teriorly first), producing a nodal hand (2-) 3-5 
(-6) cells broad (Fig. 3A K). 

Cystocarps short-stalked, irregularly globular 
to bilobed, 400-600 ^.tn in diameter. 

Spermutansict cover the lower (except ha.sal) 
several segments of young ramclli of adventi- 
nous hranchlets lying between older ramelli. 
forming male organs 120-200 /*ni in diameter. 
Terras f'Oruttgia (Fig. 5E) sessile, 1-3 per 
cell, mostly on the upper (adaxiul) side of the 
ramelli, Mibspherieal, 50-90 /j,m in diameter, 
tcUahcdrally divided 

Type totality: Holdfast Bay, S. Ausl. (F.v. 
Mueller). 

ry^.MHL, 45128- 

Mxfribittion: From Port Dcnison, W. Ausl. 
u> Gabo L, Vic. S. dasyoides usually occurs on 
rough-water coasts from low tide level to 
depths of 33 ra. Deeper growing plants are 
usually more delicate than those growing in 
turbulent conditions. 

The reproductive cells develop very sunilarly 
tn those in S. filametitosa or S, 7asmank-<t. Fe- 
male axes correspond well with that illustrated 
(Fig. $G) for S. iusmanivu, having alternating 
sterile and proearpic segments, with the latter 
comprising a 4-celled caipogonial branch on 
the supporting cell and usually iwo other peri- 
axial cells. The mature eystocarp encloses two 
or three discrete carposporophyle lobes and ihe 
pericarp is relatively lirrn at its periphery. 

S. tUisyoides h characterised hy its robust, 
opposite and more or less decussate ramelli. 
with cells ahout as long as broad and nodal 
bands 3-5 cells broad., logcther with the largely 
distichous branching. Eight periaxial cells are 
formed in branches *nd soon become separated 



by rhizoidal cells, thus forming both noda) and 
intcmodul rings of usually lf» cells, the bands 
being about equal in length. 

The type of S dosynidts (Fig. 5 A) in MEL 
is typical of this f-peeies, previously known 
mainly as S. opposita, Sender's hand-wriucn 
label with the holotype gives the locality as 
"Adelaide'', inland from Holdfast Bay. The 
type of S. opposha (in TCD) is from Preserva- 
tion Harbour, on the souih-west coast of the 
south island of New Zealand (LyalL Jan 
1851), and agrees very well with the southern 
Australian plants. Adams f 1 972, p. 83 ) 
records S. vppoxtta from several localises near 
Wellington, New Zealand, and although it is 
apparently not widely known in New Zealand, 
the specimens (e.g. CHR, 55775, from Patu- 
rau, Nelson) agree well with Australian mate- 
rial. The type of S. protifera Harvey, in TCD, 
is from Fremantle. Western Australia (Clifton), 
and is a plant with denuded branches hearing 
proliferous hranchlets. S. prolijtta represent *> 
older plants of S. dasyoides. where Ihe ihick 
axes and branches arc probably remnants of 
the previous yeats' growth, from which num- 
erous short branchlets have arisen prolifcr- 
nusly. The structure of the ramelli. is idenlicnl 
with that of 5. da$y*>Ui?s. Plants referred to 5. 
prvJifera are apparently not infrequent on ihe 
Western Australian coast, where younger aud 
typicol plants of S. dasyoides also occur. 

Acknowledgments 

The first author gratefully acknowledges a 
gram trnm the Australian Research Grants 
Committee and the technical assistance pro- 
vided by Mrs Hnid Robertson and Miss Cheryl 
Andcisoo- Dr G. Sartoni. instrluto Botanico, 
Fircnze, Iialy. and Or J. Cabioch, Station Biu- 
logtquc de Rosccff. France, kindly made avail- 
able material of .Y. fflamentoM from I'm/ope 
The Director of the National Herbarium, Mel- 
bourne, is thanked for the loan of specimens. 



References 



Adams, Nancy M. (1972). — The marine Algae of 

the Wellington area. A list of species. Hec- 

Dom. Mus. 8(5), 43-*>8 
Aoaruh, J. Ci. ( 1852). -"Species, 0«nern et 

OidJncs Algnrum M . Vol. 2, Pi. 2, pp. 337-720. 

(Lund.) 
Acabow, I, G. (1 876 ) . —"Species, Genera et 

Ordines Algatum". Vol- X Pt. I, pp. 1-724, 

Fpicrisis systematise Hoiidcarum. (I.und.) 
Agardh, J. G t!879). — Florideernes inurphologi. 

K. Veten&k Akad. Forschumll, 15(6), 1-199, 

Plates 1-33. 
AitAftDH, J. O. (1W7). — Aiwdeeta Algoluftiea . 

Cxm. IV. Acta Unh>. tutttl, 8, l-LOfi, Plates 

1-2, 



Bauton. Hihel S. (181*6), —Tape Afepe J. Hvl 
M, 193-1^8. 

BOBRQtsaN, F <J9I7>. — Th^ marine ulfiae OX the 
Danish West Indies. Vol. 2. Rhodophveeac. 
Fmt H. D*m.<k. box. Ark. 3. 145-240. 

DE ToNt, G, B, (1*97)— "Syllope Alsarum om- 
nium hucusuue Goguitafhim « Vol. 4. Tloh 
dicae-. Sect J, pp- 1-388- (Padua.) 

D»: Tonj, G. B. (1903). — "SyJIogc Algurum <>rn- 
llJOBfl hticusqne CucniUiiuui". Vol 4. Mori- 
deae. Sect. 3» pp, 77^-1525. (Padua.) 

Dq Tuni, G. R. (1924).— "SyllOgc Alganim utn- 
niiim bucusque Cngmiarium , " ( Vol b, l-lo*i- 
deae. i Padua.) 



THE SOUTHERN AUSTRALIAN SPfcCifS OF SPYRWIA 



23» 



hFLDMANN-M^OYEft, G, (1940).— "Reche<xbes 
s*tr les Ccramiacees de la Meditcrranee occi- 
denialc". (Alger) 
Gordon, Elizabeth M. ( (972). — Comparative 
morphology and taxonomy of the Wtange- 
Iicae, Sphondylothamnieae,' and Speraiotham- 
nieae fCeianiiaoeae. Rhodophvtrtl AUAJ. J, 
Dot. Suppl. 4, 1-180. 
Gru.r.R, B R (19521.— The Marine AUae of Tas- 
mania. Check list wnh localities, "Pop. Proc 
R. Sot:. ftuw, 86. 7Md6, 
Harvey, W. H. (I83V» —Algae. In W. J Hooker 
"The British Flota". Vol, "2, Pnn f, pp. 248- 
401. [Also hi W I. Hooker. l Thc Fnelish 
Flora Of James Edward Smith", Vol. S, >arl 
I. pp. 252-405.1 
Harvey, W H. ( 1844).— Alfiae of Tasmania. 

Lund. /. Hot, 3, 428-454. 
Harvey, W. H. ( 1846).— -Phycologio Briutmica", 

Vol. 1, Plates 1-72. (London.)" 
Marvpy. w. H, (1855a).— Some accoutw of ihe 
marine botany of Ihe colony of Weston Aus- 
tralia. Trans, R. In'sh Acad. 22, ^2^-566 
Harvey, W- H. ( 1855b).— Algae, //i ?. D. 
Hooker. "The Botany of (he Antarctic Voy- 
age , Part JI, Flora Novae-iiciandiac. Vol 2 
pp. 211-266, Plates 107-121. 
H.ARVtv, W. H. H859)— Algae. /« 1 D, Hooker. 
"The Botany of the Antarctic Voyage." Part 
lit Flora Tasmanrne. Vol. 2, pp. 282-343 
Plates J 85- 19ft. ' 

Harvf.y. W. H mwh— "PhycolocJa Australia". 

Vol. 3. Plates 121-180. 
Harvey W. H, ( 1863).— "Phycologia Australia" 

Vol. 5, Plates 241-300, synop.; 1-7^. 
Hommprsand, M H. (1963).— The morphology 
and clarification of worne Ceramiaceue and 
Rhodomelaceae. Univ. Catif. Pubis Hot 3<i 
(2), 165-366. 
Hooker. J, D-, k Harvey, W. K. j IS47) — Algae 

Tasmamcae. Lond. A Bot. u, 397-417 
Kmshnamisithy, V. 1 1968).— The morphology of 
gfSWw fiiamentosu <Wulf.) Harvey. Phykos 

KVBWWi?, E T. i 1849) —-Species. Alearum' 

(Leipzig.* 
Kultztnu F. T. ( 1862).— "Tabulae Phycolo- 

Ricae. Vol. 12. <Nordhausen.) 
Lucas. A. H. S. N 909). —Revised list of the 

rucoidcae and Floritfeae of Austrnlr^. Proc 

Linn. Soc. NSM 34, 9-60 
LCCAA A. H. S. flSWK— The marine algae of 

Tasmania. Pap. Ptot\ R. Soc. fa&rL 1928 

6-27, 
Lucas, A. H. S. < 1929b).— A census of the marine 

3lgae of South Australia. TMni, R. Soc. S 

Attn. 53. 45-5 V 
Locas. A. H. S., 4 Perrin, F. (1947).— "The Sea 

weeds ot South Australia". Part 2: The red 

seaweeds, pp 109-458. (Govt. Printer, Ade- 
laide.) 
May, V (1965).— A census and key to the species 

Df Rhodophyccae (red alg.ie) recorded from 

U94? COhtf ' NSW - miH ' Htrh ' 5<6 >- 



Mxzzty A. (1912). — Saggio di algologm oceanica. 
Nnova Notarts'ia 23. Nos, 415-447. 

Mazza. A. 1 1919),— Saggio di algologia oceanica 
Nno\'tt NotctritUi 30. Nos. 661-683. 

MAZZA, A. < 1925).— Saggio dl algologia oceanica. 
Nos, 8UI-927 (m 1692-2096). (Publ. pri- 
vately?) 

Newton, T (193 1).— "A Handbook of the Bri- 
tish Seaweeds" fBntish Museum: London.) 

OkamUra, K. ( J 932). —The distribution of mat in* 
algae id Pacific waters. Kcc, Oveanogr. Wkx 
Japan 3, 30-150. 

Reinhold, T. 11897)— J>ie algen der Laccpede 
Uttd Ouichen Ray. Nuo\a Notorlsia 8, 41-62. 

RriNBOLD, T. (1899). -Meeresalgcn von Investi- 
gator StraeL (Sad AustralieriK geaammelt von 
Miss Nellie Davey fWalTham. Horn ton I 
H?4wigta 38, 39-51. 

SHtPMERD, S. A- & WoMfRSLE^, H. B. S. (1970), 
— The &ublittoral ecology of West Island. 
South Australia, I. Environmental featured 
and the elga] ecology. Trans. R. Soc, & t Ami. 

Sonoish, O. G*. (1845).— Nova aiMarum genera ei 
species, mux in itinere ad oras occidental 
Novae Hollambae, collegit L. Preiss, Ph.D^, 
Bot. 2fg 3, 49-57. 

So>Ji>r.R, O. G. (1846).— Algae, hi C. Lelimann, 
"Plantae Prcissianae", Vol 2, pp I4S-295 

Sonoer. O. G. (1853).— Plantae Muellerhaae. 
Algae. Linnaea 25, 657-709, 

Sondfr 0_ G. (1855).— Algae annis IN52 et 185J 
collectac. LhmtSto 26, 506-528 

Sower, O. G. fl880).— In R von Muellci. 
"Fragmenta Phytographiae Anstrahae" Sup- 
plementum ad volumen undectmirn; Algae 
Australianae hacienns cognitae. pp. |-42, IJJ5- 

Tai>. R. (1882).— A list of (he charas, mosses, 
liverwort*, lichens, fungs. and algal* of extr.i- 
tropjcal South Australia. Tram. H. Soc $ 
Aust. 4, 5-24, 

Tlsdalu ft X (1*98).— The algae ot Victoria. 
Rep. 7th Meet Aust. Ass Adv. Sci., Sydney, 
1898, pp. 493-516, 

Wilson, J. B. ( 1892).— Catalogue o( algae col- 
lected at or near Port Phillip Heads and 
Western Port. Proc, R. Soc. Vicr. 4. 157-190 

Wollaston, Elise M. (1968).— Morphology and 
taxonomy of southern Australian ucnc'ra of 
Crouameae Schmitz (CcramiaccaeZ Rhodo- 
phyta). Aust. /. Boh i*\ 217-417. Platea 1-10, 

WoMBRSLhY. H. B. S. (1950).— The maiine afeae 
Ot Kangaroo Island. OI Lisi of species' J 
Tmn R- Soc. S. AUtt* 73, 137-197 

WoMrRstnv, H. R. S. I1958j.— Matinc algae from 
Arnhern Land, North Ausrralia. Rec. Amcr. 
Aust. Sc. Exp. Amheo> Land. Vol. 3, 139- 

WomdrslgV, H. B. S. tl966).— Port PhilUp ^m% 
yey, 3957-1963- Algae. Mem nam. Mus, 
Vic!. No. 27. 133-156. 

Wulfen, X. f 1803). — Cryptogamu aquatica. 
Koemers Archiw Bol (Leipzig) 3, J -64 



OBITUARY: GRAHAM FREDERIC WHITTEN, M.SC 

by P. B. Webb 

Summary 



2>5 



OBITUARY 

GRAHAM FREDERIC WRITTEN, M.Sc. 
16TV.I924-16.I1U975 



Graham Whitten, former Deputy Director of 
Mines in South Australia died on 16 March, 
1975, after a long illness, With his passing the 
State lost an outstanding geologist 

Whitten graduated as B.Sc. from the Univer- 
sity of Queensland in 1946 and subsequently 
jnined the Electrolytic Zinc Company where he 
worked as geologist and later Senior Geologist 
at the Rosehery Mines in Tasmania, After 
several years at Rosehery. he became respon- 
sible for exploration for 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 m 1965 to Super- 
vising GeoJogtai and during the next five years 
was responsible for all activities of the Explora- 
tion Services Division, and in 1970 was 
appointed Chief Geologist, 

With the untimely death of the then Acting 
Director of Mines, Dr K. R. Miles, ui March 
1972, Whitten was left in administrative con- 
trol of the entire Department for a period of 
eight 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 m 
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 
Society 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 (U.S.A.). 

His energy and drive will be remembered by 
all who weie privileged to know and to work 
with him. His many official reports and 
published papers are a fitting record of his 
career. 

To his wife Beuy and to his three daughters*, 
we offer our deepest sympathy. He wiD be sadly 
missed by his professional associates and bv 
his many personal friends, 

B. P. WEBB 



I9tf 

|yM 

J 965 
1965 

1965 

1065 
1966 
1*66 



Investigation of Aeromagnetic Anomalies 
hundreds of Carina, Chandada and Ripnn— 
Western Eyre Peninsula. Kept, fnxesi,, %rol. 
Sitrv, S, Ausi, 23. 

Ironstone deposits. Radium Hill district 
Min. Rev. Adelaide, 117, 80-87. 
Metallurgical testing of Pceralilla H||I 
latcn're. Min. Rev Adelaide, 117, 89-90. 
Investigation of Kopi acromagnelic ano- 
malies. Min. Rev, Adelaide, 118, 116-123. 
The Uhc of AeronrtHgndii Maps ill Geo- 
logical Regional Mapping. Quart. Geo!. 
Notts. $eo!. Sttrv. S. A MM. 15. 
Iron Ore deposits in South Australia out- 
side the Middleback Kanges. Ptor, Eighth 
Comm, Min. Metull. Congress. Aust. AVir 
Zealand, J 965. 1 , 309-3 11. 
Leached Outcrops, Prov. Eighth Ccmm. 
Min, MetalL Conpress. Aust, Nwot Zealand 
1965. % 193-204. 

The Geology of some South Australian Iron 
Deposits. M.Sc. Thesis, University of Ade- 
laide (unpublished). 

Suggested Correlation of Iron Deports 
within South Australia. Quart. Geo}. Notev 
■zeal. Snrv S A us/. 18. 



Bibliography 

iyf>7 



1968 



1968 



1 5*8 



3 969 



19711 



1971 



1972 



Investigation of Carrow and Neill aero- 
magnetic anomalies. Min. Rev , Adelaide, 
121, 40-46. 

Type Section of Iron Formations, Tarcoola 
District. Quart, Geo!. Notes, geof. Sarv, S 
Aust. 26. 

Atlas of Technical Data. Quart, Gent 
Notes, geot. Surv. S. Ami. 26. 
With Kisefy. B. G. A ground magnetic and 
gravity survey of the Wanamboo Aeromag- 
netic Anomaly, Central Ryre Peninsula. 
Repl. Invest, gcal. Sur\'. S. Aust. 32, 

Regional .gcochemical investigations in the 
Ctare one-mile sheet aTca. Mineral ftesour 
Rev.,S. Aust. 127,34-45. 

The Investigation and Exploration of ;he 
Rtuorbacfc Ridge Iron Deposit Rrp. Invest., 
f?eol, Surv. S. Aust, X$, 

The Investigation of the Almanda mine 
area. Mineral Resour. Re\\, S. Aust, 131 
25-37. 

Annual Report of the Director of Mines 
and Government Geologist — for year ended 
30th June, 1972. South Australia. 



ROYAL SOCIETY OF SOUTH AUSTRALIA INCORPORATED 



Patron: 

HIS EXCELLENCY THE GOVERNOR OF SOUTH AUSTRALIA 
SIR MARK OLIPHANT, K.B.E. 



OFFICERS FOR 1975-76 



President: 
C. R. TWIDALE, Ph.D. 



Vice-Presidents: 



B. R WEBB, M.Sc. 



N. H. LUDBROOK, M.A., PLD. 
D.I.C., F.G.S. 



Secretary: 
C. J. M. GLOVER, M.Sc. 



Treasurer: 
S. A. SHEPHERD, B.A., LL.B. 



Editor: 
H. B. S. WOMERSLEY, PLD., D.Sc. 



Assistant Editor: 
W. K. HARRIS, M.Sc. 



Librarian: 
I. M. THOMAS, M.Sc., MX, Biol. 



Program Secretary: 
W. V. PREISS, Ph.D. 



Minute Secretary: 
R. H. FISHER, A.U.A. 



Members of Council; 



P. J. M. GREENSLADE, M.A., Ph.D., 

Did 
J. K. KING, Ph.D. 



R. W. R. RUTLAND, PhJX 
J. H. J. SZENT-IVANY, Ph.D. f 
F.R.E.S. 



Auditors: 
Messrs MILNE, STEVENS, SEARCY k CO. 



Registered for posting as a publication Category 'C* 
Printed by Graphic Services Pty Ltd, 86a PIrle Street, Adelaide 6000