VOL. 115, PARTS 1 & 2.

31 MAY, 1991

Transactions of the

Royal Society of South

Australia

Incorporated

Contents

Opresko, D. M. & Bayer, F. M. Rediscovery of the enigmatic coelenterate

Dendrobrachia, (Octocorallia: Gorgonacea) with descriptions of two

new species - - - - - - - - - -

Rowett, A. I. The dispersed cuticular floras of South Australian Tertiary coalfields,

Part 1: Sedan - - - - - - - - - -

Kemper, C. M. & Ling, J. K. Whale strandings in South Australia (1881-1989) -

Campbell, E. M. & Twidale, C. R. Relationship of residual hills and sheet fractures

in the Gawler Ranges and environs, South Australia - -

Dayies, M. Descriptions of the tadpoles of some Australian limnodynastine

leptodactylid frogs - - - - - - - - -

O”Donoghue, P. J., Phillips, P. H. & Shepherd, S. A. Perkinsus (Protozoa: apt ia

infections in abalone from South Australian waters -

Nobbs, J. M. Description of the male of Tylenchorhynchus tobari Sauer & Annells,

1981 and observations on the morphology and host range of the

female in arid South Australia - - - - - - -

Wallace, M. & Pring, A. The Mangalo meteorite, a new (L6) Olivine-Hypersthene

Chondrite from South Australia - - - - - -

Ling, J. K. Recent sightings of Killer Whales, Orcinus orca Gerace Bae

in South Australia - - - = 3 s Mi

Tyler, M. J. Crinia Tschudi (Anura: Leptodactylidae) from the Cainozoic of

Queensland, with the description of a new species - - -

Tyler, M. J. A large new species of Litoria (Anura: Hylidae) from the Tertiary of

Queensland - - - = is az 2 L 3 :

Brief Communications:

Beveridge, I. The distribution of Echinocephalus overstreeti Deardorff & Ko

(Nematoda), a parasite of elasmobranch fishes in Australian

waters - - - - - ~ - - - - -

Harris, W. K. & Twidale, C. R. Revised age for Ayers Rock and The Olgas -

PUBLISHED AND SOLD AT THE SOCIETY’S ROOMS

SOUTH AUSTRALIAN MUSEUM, NORTH TERRACE, ADELAIDE, S.A. 5000

TRANSACTIONS OF THE

ROYAL SOCIETY

OF SOUTH AUSTRALIA

INCORPORATED

VOL. 115, PART 1

TRANSACTIONS OF THE

ROYAL SOCIETY OF SOUTH AUSTRALIA INC,

CONTENTS, VOL. 115, 1991

PARTS 1 & 2, 31 May

Opresko, D. M. & Bayer, F. M. Rediscovery of the enigmatic coelenterate Dendrobrachia,

(Octocorallia: Gorgonacea) with descriptions of two new species -

Rowett, A. I. The dispersed cuticular floras of South Australian Tertiary coalfields, Part |:

Kemper, C. M. & Ling, J. K. Whale strandings in South Australia (188]-1989) - -

Campbell, E. M. & Twidale, C. R. Relationship of residual hills and sheet fractures in the

Gawler Ranges and environs, South Australia - - - -— -

Davies, M. Descriptions of the tadpoles of some Australian limnodynastine pti

Frage) o> b=" “pee ar HE we - =

O’Donoghue, P. J., Phillips, P. H. & Shepherd, S. A. Perkinsus (Protozoa: a chines ined

infections in abalone from South Australian waters - -

Nobbs, J. M. Description of the male of Tylenchorhynchus tobari Sauer & Annells, 1981 and

observations on the morphology and host range of the female in arid South

Australia ee SS ee ee ee

Wallace, M. & Pring, A. The Mangalo meteorite, a new r ALS} inhi pehne Chondrite

from South Australia - = - mie

Ling, J. K. Recent sightings of Killer Whales, Orcinus orca (Cetacea: Delphinidae), in South

Australia Ne Si tt ke ee ae

Tyler, M. J. Crinia Tschudi (Anura: Leptodactylidae) from the Cainozoic of un Ansel

with the description of a new species - - -

Tyler, M. J. A large new species of Litoria CAR: Hylidae) f from the Pees of

Queensland - - -

Brief Communications:

Beveridge, I. The distribution of Echinocephalus overstreeti Deardorff & Ko silks

a parasite of elasmobranch fishes in Australian waters -

Harris, W. K. & Twidale, C. R. Revised age for Ayers Rock and The Olgas - +

103

107

109

PARTS 3 & 4, 29 NOVEMBER, 1991

Koste, W. & Shiel, R. J. Rotifera from Australian inland waters. VII. Notommatidae (Rotifera:

Monogononta) - - - - - - + - - - Til

Cann, J. H., De Deckker, P. & Murray-Wallace, C. V. Coastal aboriginal shell middens

and their palaeoenvironmental significance, Robe Range, South Australia 161

Zeidler, W. A new genus and species of phreatic amphipod (Crustacea: Amphipoda) belonging

in the “chiltonia” generic group, from Dalhousie Springs, South Australia

- ee - Se SH + RT

Lee, D. C, & Birchby, C. M. Paraphauloppia (Acarida: Cryptostigmat: aii diaebi and

its occurrence in South Australian soils - 189

McEntee, J. C. Lake Frome (South Australia) aboriginal trails ~ - - - - 199

Southcott, R. V. A new trombellid mite (Acarina: Trombellidae) from South Australia - 207

Obendorf, D. L., Beveridge, I. & Andrews, R. H. Cryptic species in populations of

Globocephaloides trifidospicularis Kung (Nematoda: SHER GRONET ICA.

parasitic in macropodid marsupials > es - 213

Smales, L. R. A new species of Antechiniella Quentin & Beveridge, 1986 (Nematoda:

Acuariidae) from the Australian water rat, isos de nt eogreler ede

1804. - - - - 217

Insert to Transactions of the Royal Society of South Australia, Vol, 115, paris 3 & 4, 29 November, 1991

REDISCOVERY OF THE ENIGMATIC COELENTERATE

DENDROBRACHIA, (OCTOCORALLIA: GORGONACEA) WITH

DESCRIPTIONS OF TWO NEW SPECIES

BY DENNIS M. OPRESKO* & FREDERICK M, BAYER}

Summary

Since 1876, Dendrobrachia fallax Brook, an arborescent, noncalcareous anthozoan coelenterate

with a spiny, proteinaceous axis, has been assigned to the suborder Antipatharia in spite of such

equivocal features as the probable presence of eight pinnately branched tentacles and a solid axial

core. In recent years, specimens resembling D. fallax have been collected from off the southern

coast of Australia and from the Straits of Florida. These specimens represent two new species of

Dendrobrachia and are here described as D. paucispina sp. nov and D. multispina sp. nov.

Furthermore, anatomical and histological studies demonstrate that Dendrobrachia has characteristic

octocorallian features. The solid, noncalcareous axis and absence of calcareous sclerites suggest a

relationship with the holaxonian family Chrysogorgiidae, but the distinctive skeletal features

warrant retaining the genus in a separate family Dendrobrachiidae.

KEY WORDS: Gorgonacea, Octocorallia, Antipatharia, Chrysogorgiidae, Dendrobrachiidae,

Dendrobrachia.

Transwehons uy the Royal Society of & Aust. (1991), 115¢1), 19.

REDISCOVERY OF THE ENIGMATIC COELENTERATE DENDROBRACAHIA,

(OCTOCORALLIA: GORGONACEA) WITH DESCRIPTIONS OF TWO NEW SPECIES

by DENNIS M. OPRESKO* & FREDERICK M. BAYER}

Oreesku, DM. & Bayer, FM, (199)) Rediscovery of the enigmatic coelenterate Dendrobrachia (Octocorallia;

Gorgonacea), with descriptions of two new species. Trans. R. Soc. S. Aust, VIS(1), 1-19, 31 May, 1991,

Sie 1876, Dendrobrachio fallay Brook, an arborescent, noncaleareous anthozoan coelenterale with a spiny,

protei aceuus axis, has been assigned fo the suborder Antipatharia in spite of such equivocal features as the probable

presence of eight pinnately branched tentacles and a solid axial core In recent yeurs, specimens resembling 2 fallax

have bees collected from off the southern coast of Australia and from the Straits of Florida. These specimens represent

two new species of Dendrobrachia and are here described as D. paucispina sp. nov and D. mullispina sp. noy.

Furthermore, anatomical and histological studies demonstrate that Dendrobrachia has characteristic octocarallian

fearures. Uhe solid, noncalcareous axis and absence of calcayeous sclerites suggest a relationship with the holaxanian

family Chrysogorniidae, but the distinctive skeletal fealures warrant retaining (he genus in a separale family

Dendrobrachiidae.

Key Worns: Gorgonacea, Octocorallia, Antipatharia, Chrysogorgiidae, Dendrobrachiidae, Dendrobrachia,

Introduction

On 27 March 1876, at 8903'S, 14°27'W, off

Ascension Island, H.M.S. “Challenger” dredged twa

specimens of an arborescent coelenterate with spiny

proteinaceous axis at station 343 in 425 fathoms of

water. These specimens subsequently were described

as Dendrobrachia fallax by Brook (1889) and

assigned to a new family, Deridrobrachtidae, placed

in the suborder Antipaiharia, The species was taken

again in 190}, off the Cape Verdes by the Prince

of Monaco, and reported briefly by Thomson

(1910), who concurred with Brook in placing it in

a special family of Antipatharia.

In splte of their equivocal features — pinnate

tentacles probably eight in number absence of a

hollow axial core — the spiny axis and lack af

calcareous spicules overrode other considerations

and classification of Dendrebrachia among the

Aatipatharia has prevailed until the present.

Durlig 1988 and 1989, Karen Gowlett-Holmes,

while aboard trawlers in the Great Australian Bight

and farther south aff the Continental shelf of

Australia, collected a number of deep sea octocorals

and antipatharians which were deposited in the

Sourh Australian Museum. Along with specimens

of Chrvsogorgia were six colonies also tentatively

identified as chrysogorgiid gorgonians. When

examined more closely, these were found ta resemble

Brook's enigmatic Dendrahrachia fallax. This tew

material represents a new species of the genus, and

* Health and Safety Research Division, Oak Ridge

National Laboratory, FO. Box 2008, Oak Ridge,

Tennessee 37831-6051), LSA,

| Depariruent of Invertebrate Zoolegy, US. National

vt eae of Natural History, Washington, Dc. 20560,

SA,

is sufficiently well preserved to permit a more

detailed description of the soft anatomy and a

reevaluation of the placement of the family

Dendrobrachiidae. Also included in this report is

a description of a new species of Dendrobrachia

from the western Atlantic, which is based on one

specimen collected from the Straits of Florida.

Systematics

Subelass Octovorallia

Order Gorgonacea

Suborder Holaxonia

Family Dendrobrachiidae Brook, L889

Genus Dendrohrachia Brook, (889

Dendrobrachia Brook, 1889:159; Hickson, [895:40;

Thomson, 1910:142.

Type species: Dendrahrachia fallax Brook, by

monotypy.

Diagnosis: Atborescent Gorgonacea with purely

proteinaceous axis lacking hollow core; axis marked

by conspicuous ridges and grooves, with more or

less numerous and prominent spines along the

summit of the ridges, Polyps and coenosare without

calcareous sclerites.

Distribution: Eastern Atiantiic Ocean off

Ascension and Cape Verde Islands; western Atlantic

off Cay Sal, Bahamas; Great Australian Bight.

394-(089 m,

Dendrobrachia fallax Brook

FIGS 1, 2; Table |

Dendrobrachia fallax Brook, 1889:159-160, pl, 10,

figs 1-8, Thomson, 1910;142-143.

D. M. OPRESKO & F. M. BAYER

1544 toKy 7" 24Gum (1544 10K

Fig. 1. Dendrobrachia fallax Brook, axis. Top, lateral view of terminal twig; bottom, cross section below apex.

Stereoscopic pairs, SEM.

KeOISCOVERY OF THE ENIGMATIC COELENTERATE DENDROBRACHIA 3

Material Exermined: pe spevimens, South

Adlantic, off Ascension Island, 8"03°S, Jae27' W,

425 fm (777 m1), “Challenger” Expedition, Sta, 343,

27 March 1874, 2 spec. THritsh Museur (BMNH)

1890,4.9,27).

Description: Colany planar, sparsely branched,

with some averlupping of adjavent branches;

branchlets bilateral, very irregularly alternate, rarely

opposite, highest order branchlets mostly 1-3 om

long, occasionally longer, 0.5-0.8 mm in diameter,

on average about | cm apart on same side of

branches. Usually six, occasionally five or seven

axial ridges on branchlets, increasing in number on

larger branches. Spines present on ridges, one row

per ridge: spines variable in sive and shape but

increasing in size with increasing thickness of

branches, 0.2-0,25 mm long on branches about i

mm in diameter. Polyps placed bilaterally, alternate

or in nearly opposite pairs, 3-6 polyps per em.

Diseusstons Neither of the (pe specimens has a

basal holdfast and one specimen appears to have

been broken off a larger colony. Both specimens

dre abour 20 cm fall and 5-10 cm wide. The basal

“Stem” diameter is 2 # 2.5. mm in one colony and

about 3 mint in the other. Overall, the branching o!

(he colimies ts sparse, open, lo gbpul the eighth

order, and generally in one plane with some

overlapping of adjacent branches (see Hrook 1899:

p. WW, fig. }}. On the central portions of the

corallum, the branching is very weakly sympodial

i) that somie Of the higher order ramifications

became thicker and longer than the branches fram

which they arise. The branch angles (delineated by

the inner or distal side of a braneh and the lower

order branch from which it arises) are generally

breater than 45°. The smallest ramifications of the

corallum, the branchlets, occur al all levels of the

corallum. They are straight oc slightly curved

upward, unbranched, about 0.5-0.8 mm in diameter,

and usually pot more than 3 cm long. They tend

to be positioned bilaterally and ina yery irregularly

allernaling patiern along the sides of the branches,

The distance between ile bases of adjacent

branchlets and/or branches on the same side of the

lower urder ramificauion froar which they anse

ranges from about 6-9 mm, and the average

distance i¢ 9.4mm (n ~ 24). The distance between

adjacent branenlets ald branches on opposite sides

of the axis ranges [rom about 2-17 mm, and the

average distance is 6.3 mm (a = 245).

AL the tips of the smallest branchlets the skeletal

axis has six (Occasionally five or seven) longitudinal

ridges separated by grooves or channels. The ridges

do Nol radiate Ouc [rom 4 commun central port,

hut rather appear Lo develop as bifurcations at the

ends of a primary skeletal plate which is narrow and

rectangular in cross section. This primary axial

structure of branchlets can be seen at rhe core of

the thicker branchlets. when viewed in cross Section

(Fig. 1, bottom).

Extending down from the tips ef the branchlets

for a distanve of about t-1.5 cm, the ridges remain

smooth or possess only small, scattered rounded

elevations about 0,05-0,08 mm high, Further down

on the branchlets distinct spines develop along the

ridges (Fle. J, top). Although the spines are not

stricily uniform in size or shape throughout the

coraltum, they generally become tonger as the

diameter of the branchlets and branches increases..

Muny spines typically have a relatively sharp apex

and a flared base (Fig. 2); however, others are more

blunt, and some even have a swollen or knob-like

apex, On branchlets 0.6-0,8 mim in chameter, the

spires are usually 0,10-0,15 mm high; on branches

0,8-1.0.mm in diameter, they are usually 0.95 to 0,20

mim high (bur up co 0.25 mm in some places). 'They

are arranged in 5S to 6 rows (ane row per skeletal

ridge) with 4-5 spines per millimeter in each row,

Generally, the ridges remain relanyely narrow with

a single row of spines, On the thicker branches mare

longitudinal rows of spines develop. On a branch

{2 mm mi dtameter (excluding spines}, rhe spines

are about 0.3 mm high, 5-6 rows of spines can be

seen in one lateral view, and there are about 3-4

spines per millimeter in each row, The height of the

spines near tlre tase of the stern (diameter 2,0 mm)

is about 0.5 mm, there are 3-4 spines per millimeter

in cach row, and about 12 very irregular rows of

spines can be seen in one lateral view, The rows are

separated by wavy lines of coenenchyme which

intersect one another at varying inlervals, On the

second specimen (basal stem diameter about 3mm).

the spines reach a maximum size of about 0.4 mm,

and as many as 17 rows of spines can be seen from

one aspect. In the illustration given by Brook (1889:

pi. 1, Fig. 8) there appear to be about 38 or more

rows around the entire circumference of a branch

from near Uhe base of the corallum-

The polyps arise from the coenenchyme in the

grooves formed between the skeletal ridges. Their

atrangement on the coralium is not strictly regular,

although it is generally bilateral. On some

branchlets.a regular alternating pattern can be seen

with the palyps spaced about § mm apart.

Oceasionally the polyps occur in nearly opposite

pairs, In the latter case there can be as many as 6

polyps per centimeter along the branchlets. The

condition of the polyps in both specimens is poor

and allows for only 4 rough estimate of their size.

They appear to be no taller than about 2-2,5 mm

(including tentacles). The body column is about 1.0

nim in length and about 0.5 mm in diameter at the

base, As noted by Brook, the polyps do not appear

D. M. OPRESKO & F. M. BAYER

1544 1QKY

158um 1544 18KY 154um

Fig. 2. Dendrobrachia fallax Brook, axis. Stereoscopic views of axial spines, SEM.

REDISCOVERY OF THE ENIGMATIC COELENTERATE DENDROBRACHIA 5

TABLE. 1. Morphometrics for Dendrobrachia spp.

Parameter

Corallum:

Stem diam./height (%)

Branchlets:

Maximum length” (cm)

Diameter (mm)

Avy dist. apart’ (mm)

Axial Ridges:

Number on branchters"

Axial Spines:

Number of rows per ridge

Number per mm per row

Height of Spines* (mm):

Axis diam, <0.5 mm

Axis diam. 0,6-0.8 mm

Axis diam. 0.8-1.0 mm

Axis diam. 1.0-2.0 mm

Axis diam, >2,0 mm

Polyps:

Maximum height! (mm)

Number per cm

Pairs of pinnules

fallax paucispina multispina

1.2-1.5 1.1-1.7 0.7"

va, 3 ca. 3 ca, 3

0.5-0.8 0.4-0.8 04-05

9.4 9,0-9.5 13

5,6,7 6,7,8 5

1 Q-L 1-2+

34,5 34,5 45,6

0.05-0,10 - 0.06-0,12

0.10-0.15 0.00-0.07 O.10-0.25

0.15-0,25 0.07-0.12 0,26-0,32

ca. 0.3 0,12-0.18 0.25-0,40

ca. 0.5 012-025 -

2.5 2,2 ca. 2.3

3-6 3-6 3+

6-75 8-10 -

* Specimen may be part of a larger colony.

* Unbranched branchlets, rarely longer.

' On the same side of the axis,

' Mast common condition italicized.

” Height of spines very variable, commonest size ranges shown.

' Including tentacles.

“ As reported by Thomson (1910)

to stand out at right angles to the axis, but rather

are ifclined distally,

Remarks: Thomson (i910) reported that the

polyps were often in subopposite pairs, with the

pairs spaced at intervals of two lengths or more

apart. He also noted that there were six or seven,

pairs of pinnules on the tentacles.

Distribution: Known from off Ascension Island,

777 m, (“Challenger" Expedition), and from Cape

Verde Islands, 394 m,

Dendrobrachia paucispina sp. nov.

FIGS 3-8, 12-13; Table 1

Material Examined; Off the southern coast of

Western Australia: 125 nautical miles East of Cape

Arid, 34903'S, 125°31'E, 1011-1020 m, F/V

“Adelaide Peart", 31 July 1988, Coll. K. Gowlett-

Holmes, K. Olsson, M. Cameron, syntypes; South

Australian Museum SAM H715, | spec; USNM

87769, 1 spec; about 80 nautical miles SW of

Pearson Island, 35°04'S, 133°35'E, 900-960 m,

F/V “Comet”, 13 April 1989, coll, K. Gowlett-

Holmes, syntype: SAM H716, 1 spec; about 100

nautical miles due west of Whidbey Point, 34°49°S,

133°07’E, 884-859 m, F/V “Longva III”, coll. K.

Gowlett-Holmes, 11 November 1989, syntypes:

SAM H728, | spec.; USNM 87768, 1 spec.

Description: Colony planar, loosely branched, with

some overlapping of adjacent branches; branchlets

bilateral and very irregularly alternate; highest order

branches usually not more than 3 em long, 0.4-0,8

mm in diameter, on average about | cm apart on

same side of lower order branches. Six, occasionally

seven, rarely eight longitudinal axial ridges on

branchlets, increasing in number on _ thicker

branches. Spines sparse, often absent, arranged in

one row per axial ridge where present. Spines not

uniform in size and shape, but increasing in length

on larger branches; about 0,1 mm long on branches

1 mm in diameter, Polyps bilateral, in alternate or

subopposite pairs, 3-6 per cm.

Discussion; The type series consists of six

specimens, all of which are to some degree bent over

D. M. OPRESKO & F. M. BAYER

Fig. 3. Dendrobrachia paucispina sp. nov. Syntype colony SAM H715.

REDISCOVERY OF THE ENIGMATIC COBLENTERATE DENOROBKACHI A 7

and iwisied in the upper part of the corallum, a

condition which may have resulted in part from

preservation and storape in small containers, When

Ubfolded into what was probably ils natural shape,

the largest colony is about 4§ cm tall and 15 em

wide, with a basal stem diameter of 7-5 mm. The

smialles! colony is 15 cm tall and about 7 em wide,

with a basal stem diameter of about 2 mm.

A colony of medium size (SAM H715) a8

described in detail. This specimen is about 28 cm

tall and 6 cm wide, and has a basal stem diameter

of4 « 43mm (Fig. 3), The branching pattern of

the corallum is generally planar with some

overlapping of adjacent branches, The branching

is irregular to about the 9th onder. The highest order

branching consisls of small, relatively thin

branchlets arranged in a very loose bilateral and

alternating pattern. The smallest unbranched

branchlets range in length from 0.4-3,.0 cm (xX =

147 cm, n «= 23), and they are 0,4-0,6 mm in

diameter. The average distance between the bases

of adjacent branchlets and/or branches on the same

side of a lower order branch is 9.0 nim (range 4-19

nm, n 30). The average distance between

adjacent branchlets and/or branches on oppasite

sides is 4.8 mm (range 3-8 mm, a = 9). The branch

angles (delineated by the distal side of a branch and

the lower order branch from which if arises) are

generally greater than 45",

At the tips of the smallest branchlets, the axial

skeleton cousists of sik, occasionally seven, and

rarely eight narrow longitudinal ridges separated by

deep, U-shaped grooves (Fig. 4, lop). The ridges

increase in number on the thicker, alder parts of

the cortlum (Fig, 4, bottom). are present on the

srem and also faindy visible on the upper part of

the holdfast, On branches 2-3 mm in diumeter, 12

or more ridges can be seen in one lateral view,

Over most of the corallum, and particularly on

branches less thab 1 mm in diameter. the skeletal

tidges are smooth (Fig, 5, top); however, in same

places scattered spines oceur ia single row along

several of the ridges On the smaller branchlets they

take the form of small rounded elevations (Fig. 5,

bottom on the older branches they are mare spine-

lke (Pip. 6). Where present, the spines occur on only

one ora few of the ndges and never on all the ridges

around the circumference of rhe axis, Although

quite variable, the heigtic of the spines is generally

about 0.07 mm on @ branchlet 0.6-D.8 mm in

diameter, 0.07-0.12 mm on branches abour 0.8-1,0

mini it diameter, and 0.12 to 0.18 mm on branches

1. 2,0) mm in diameter. There are usually 3-5 spines

per millimeter in each row, Spines are-also found

at the base of several of the small branchlets which

arise directly from the sient or thicker branches. in

these cases the spines can be 0.25 mm or more in

height.

Polyps are present throughour the corallum, They

ure nat arranged in a strictly regular order but are

mere abundant on the lateral and front sides of the

branches (relative to the plane of the corallum). Out

many of the larger branches a narrow band of

coenenchyme without polyps extends down the

middle of the posterior side. On the branches the

polyps are spaced irregularly (Fig, 7), sometimes

unilaterally, and sometimes bilaterally with

alternating Or opposite pairs. Most polyps are about

S mm apart, and there can be 3-6 polyps per

cenlimeter depending on whether they are placed

singly or in pairs. They are usually inclined drstally,

often with their adaxial side lying against the axis,

The largest polyps are about 2.2 mm tall (including

tentacles); the body column is about t mmm in Jength

(Fig. 8). On several polyps in which the tentacles

were adequately expanded, 8-10 pairs of pinnules

could be seen on each tentacle,

The remaining specimens in the type series are

similar in most morphological features te the one

described above, All the colonies have a rypical

planar branching pattern, with the highest order

unbranched branchlets being pot more than 3 cm

fong and spaced about t cm apart in a very loose

bilateral and alternating fashion, fn the larger

colonies, there are infrequent fusions of same of

the lower-order branches and the larger branches

and stem have a distinctive golden, somewhat

iridescent sheen_

Fram specimen to specimen, and even within a

single colony — from branch to branch —

differences can be seen in the occurrence, size, and

density of ihe axial spines. tn the 1wo largest

specimens (SAM H728 and USNM 87768}, the

spines are overall slightly smaller and sparser ian

those in the described colony, Furthermore, spines

are only rarely present on branches of about 2 mm

in diameter or mare; fowever, the axial ridges

remain distinct and can be seen on both the

branches and the stem.

In another specitnen (SAM H728, height 30 cm,

basal stem diameter sbout 4 ram) spines are present

onall parts of the corallum including the stem and

basal plate; however, they are nor uniformly

distributed, are absent in some places, but present

on all the axial ridges in other places, This

variability im the occurrence of the axial spines 1s

a characteristic feature of this species:

Remarks! Although the general form ol the

corallum af D, paucispina sp. wov. is very similar

to that of D fallax; this species can be differentiated

on the basis of the very sparse number and relatively

smaller size of the axial spines (Table 1), The

D. M. OPRESKO & F. M. BAYER

LOKY 38mm | 154@ 19KY 38mm

Fig. 4. Dendrobrachia paucispina sp. nov., axis. Top, lateral view of terminal twig; bottom, lateral view of larger

branch SAM H715. Stereoscopic pairs, SEM.

REDISCOVERY OF THE ENIGMATIC COELENTERATE DENDROBRACHIA

1546 . (1546 = 10KY

238mm é i 98mm

Fig. 5. Dendrobrachia paucispina sp. nov., axis. Lateral views of terminal twigs, SAM H715. Stereoscopic pairs, SEM.

W) D, M. OPRESKO & FT. M,

BAYER

Fig. 6. Dendrobrachia paucispina sp. noy,, axis. Lateral view of larger branch, SAM H7I5, Stereoscopic pair, SEM,

complete absence of spines on many parts of the

axis is quite distinctive. Where present, and for

branches of similar thickness, the spines in this

species are only about one-half as large as those in

D. fallax. Although in both species the smallest

branchlets most commonly have six primary axial

ridges, in D. paucispinag sp.nov. there are

occasionally seven and sometimes even eight ridges

present. In contrast, in D fallax there are

occasionally five or seven ridges on the smallest

branchlets. The polyps in D. paucispina sp. nov. and

D. fallax appear to be similar in size and number;

however, there is some evidence suggesting that the

number of pairs of pinnules on the tentacles may

be different in the two species. In this species there

are eight-ten pairs of pinnules per tentacle.

According to Thomson (1910), there are only

six-seven pairs per tentacle in D. fallax.

Distribution: Known only from the Great

Australian Bight, 884-1020 m, on the continental

slope.

Dendrobrachia multispina sp. nov.

FIGS 9-11, Table 1

Material Examined: Straits of Florida, west of

Cay Sal Bank, Bahamas, 23°51.9' N, 80°42.7' W,

1080-1089 m, R/V “Gerda” Sta. 1111, 30 April 1969,

Holotype, USNM 87770.

Description: Colony planar, very sparsely branched;

branchlets bilateral, very irregularly alternate;

highest order branchlets usually not more than 3

cm long, 0.4-0.5 mm in diameter, on average 1.3

cm apart on same side of branches. Five

longitudinal axial ridges on branchlets. Spines in

rows on axial ridges, one row per ridge on smallest

branchlets, multiple rows on larger branchlets and

branches. Spines not uniform in size or shape,

becoming tall and acicular on larger branchlets and

branches; about 0.3 mm long on branches about

1 mm in diameter. Polyps mostly bilateral and

alternate, 3 per cm.

Discussion: The type specimen lacks a basal

holdfast and may have been broken off a larger

colony. The upper portion of the specimen is bent,

perhaps due to preservation (Fig. 9). With the

branches straightened and extended vertically, the

corallum is about 19 cm high and 7 cm wide. The

diameter at the basal end of the “stem” is 1.4 mm.

The colony is very sparsely and openly branched

in an irregular bilateral fashion. The average

distance between the bases of adjacent branchlets

and/or branches on the same side of the lower order

branches is 13 mm (range 8-21 mm, n = 24). The

REDISCOVERY OF THE ENIGMATIC COELENTERATE DENDROBRACHIA I

Fig. 7. Dendrobrachia paucispina sp. nov. Tip of terminal branch with polyps intact, critical-point dried, SAM H715.

Composite stereoscopic pair, SEM.

average distance between adjacent branchlets

and/or branches on opposite sides is 6 mm (range

1-16 mm, n = 22). The branch angles are mostly

60° or more. The smallest unbranched branchlets

range in length from 0.7-3.1 cm (KX = 1.5 cm, n

= 12) and they are relatively stiff and straight. They

are about 0.4-0.5 mm in diameter.

At the tips of the smallest branchlets, the axial

skeleton consists of five narrow longitudinal ridges

separated by wide v-shaped grooves. For a distance

of 0.5-2 cm from the tips of the branchlets the

ridges are smooth or have only small irregularly

rounded or triangular elevations which are generally

not more than 0.07 mm high (Fig. 10). With

increasing thickness of the branchlets these develop

into spines which eventually become relatively tall

and acicular, the ridges become wider and lobe-like,

and the grooves become relatively narrow (Fig. 11,

top). Multiple rows of spines develop on the surface

of the enlarged ridges. In some places two relatively

straight rows of spines are positioned along the

edges of the ridges adjacent to the grooves with

additional irregular rows or scattered spines

occurring between the edge rows. Overall, the axis

2 1M. OPRESKO & fF, M. BAYER

eh eaea ta

.68nn

Fig. 8, Dendrobrachia paucispina sp. noy. Single polyp, critical-point dried, SAM H715. Stereoscopic pair, SEM.

becomes quite spinose; however, remnants of the

five primary grooves, although quite narrow, remain

recognisable, even at the base of the corallum.

The spines are not uniform in size or shape from

branchlet to branchlet. They stand out at right

angles to the axis and often are slightly crooked.

The height of the spines is 0.10-0.25 mm on

branchlets 0.6-0.8 mm in diameter (Fig. 11,

bottom), 0.26-0.32 on branches about 0,8-1.0 mm

in diameter, and up to 0.4 mm on branches 1.0-1.2

mm in diameter. The largest spines are relatively

narrow, about 0,05 mm in diameter at their center,

There are usually four or five, sometimes six spines

per millimeter in each row. Occasionally the spines

are slightly more developed at the base of small

branchlets arising directly trom the stem and thicker

branches.

Polyps of this specimen are poorly preserved,

They are arranged irregularly, in a somewhat

bilateral pattern. There appear to be about 3 per

centimeter, The largest polyps are about 2,3 mm tall

(including tentacles); the tentacles are about 0.8-1.0

mm long.

Remarks; In size and general appearance D,

multispina sp. noy. is similar to D, fallax and D.

paucispina sp. nov. All have a somewhat planar

corallum with rather irregular, but generally

bilateral and alternating branching. The branching

of the corallum of D. inultispina sp, nov. is slightly

more sparse and open, but this may not be typical,

considering that the specimen may only be part of

a larger colony. The taxonomic character in which

this species differs consistently from the other two

species is the greater development of the axial

spines. In this species the spines on the larger

branchlets and branches develop in multiple rows

along the axial ridges, whereas in the other two

species there is only one row of spines per ridge,

The spines in D. multispina sp, nov, are also

relatively larger and more crowded together (han

in the other two species (Table 1). In addition, in

D, multispina sp. nov. ihe number of primary

skeletal ridges on the branchlets is consistently five

throughout the corallum, whereas in D. fallax it is

usually six, occasionally five or seven, and in D.

paucispina sp. nov. it is usually six, occasionally

seven, and rarely eight,

Distribution: Known only from the type locality.

Anatomy and Histology

Anatomy and histology have not been described

for very many species of Octocorallia, and much

of the published information deals with a few

pennatulacean species, Hickson (1895b) described

the anatomy of A/evonium digitatum Linnaeus,

REDISCOVERY OF THE ENIGMATIC COELENTERATE DENDROBRACHIA

Fig. 9. Dendrobrachia multispina sp. noy. Holotype colony, USNM 87770.

Bouillon & Houvenaghel-Crevecoeur (1970)

described the anatomy and histology of Heliopora

coerulea (Pallas), and Bayer & Muzik (1976a)

described the general structure of the solitary

Taiaroa tauhou Bayer & Muzik.

Among the gorgonaceans, the structure of

Pseudoplexaura crassa Wright & Studer was

investigated in considerable detail by Chester (1913),

and that of Plexaura homomalla (Esper) by Bayer

(1974). The anatomy and histology of those species

conform in general with those of all other

gorgonaceans the structure of which has been

described.

The anatomy of the polyps of all monomorphic

octocoral species and of the autozooids of

dimorphic species is remarkably uniform, As this

basic form has been described and illustrated

adequately in general treatises (e.g., Hyman 1940;

Bayer 1956) as well as in specialized accounts

(Hickson 1895b; Chester 1913; Bayer 1974; Bayer

et al. 1983), it need not be repeated here. The polyps

of Dendrobrachia conform to the general

gorgonacean pattern. Paraffin serial cross-sections

of a polyp of Dendrobrachia paucispina sp. nov.

clearly show that the general form corresponds to

the usual gorgonacean pattern, differing in only

D. M. OPRESKO & F. M. BAYER

i ~~

eee peepee ueunaee

8mm | 1545 10K 38mm

10kKy e@Gum 1545 ; 208um

Fig. 10. Dendrobrachia multispina sp. nov., axis. Lateral views of terminal twig USNM 87770. Stereoscopic pairs, SEM.

REDISCOVERY OF TRE ENIGMATIC COELENTERATE DENDROBRACHILA is

mio details. As the specimens were not prepared

with #natomical and histological investigation in

mind, fixation is adequate only for deiionstration

Ol gross anatomy and @ very sitperficial

invesiigation of histology,

From Fig, $2 itis clearly seen that the polyps of

Dendrobrachia have the usual four couples of

complete mesenreries and a single siphonoglyph,

The arrangement of longitudinal retractor muscles

is, as-usual, on the suleal face of the mesenteries,

bur the retractors of the asulcal and sulcal

mesenteries are relatively weak (at least at the level

of the body that was sectioned) and development

of mesogloea! plates an the asulacal side of rhe four

lateral miesenteries (Figs 12, 13) is aclear indication

that muscle fibers are present on both faces of at

least the lateral mesenteries,

The siphonoglyph is well developed and the

epithelium of the pharyax at its upper end is thrown

into abouc 10 fongitudinal folds, similar to the

condition in Aleyonium observed by Hickson

(1895b). These seem to fade oul proximad, where

they zive way lo (ransverse falding of the pharyngeal

wall probably related to shortening of the pharynx

resulting from contraction.

The epidermis (Fig, 13) is composed of the usual

tall, abeonic coyer cells (ef Chester 1913; Hayer

1974), thrawn inte conspicuous lobes and covered

by delicate cuticle, which in many places has been

torn off during processing. Nematocysts of

unknown type but probably atrichous isorhizas are

very Widely scattered in the epidermis of the body

wall, not densely clustered in batteries as 1s the case

in Antipatharia (van Pesch 1914),

The mesogloea of the body wall of the palyps

is Unusual in that if appears 10 consist oF two layers,

the inner stained pink as usual jn haematoxylin-

cus, the outer, somewhar thinner, dark purple.

However, as disruption of tissues during the

sectioning process separates the wwo layers here and

there, it is more likely that the darkly stained outer

layer is, in fact, a basement membrane of the

epidermis.

The gastrodermis of the body wall is unusually

thick, in some places nearly or quile as thick as the

epidermis, and composed of elongate, spindle-

shaped cells. In the immediate vicinity of the

mesenteries, narrow ridges of mesogloes extend into

Ihe gastradetmis, strongly suggesting the presence

ol longitudinal museles.

AFfinities within the Octocoratiia

It js or insignificant that the curator of

invertebrates af the South Australian Museum

assoviated Dendralvachia with Chrysogarsia.

Often, superficial similarity may reFlect more

fundamental relationships,

The absence of calearcous svlerites in

Dendrobrachia eliminates one af the classic

characters in gorgonacean classification, leaying

only the nature af the axial supporting skeleton as

a clue (0 relationships. Ayiong the gorgonians, only

two species of Trhichogorgia lack sclerites, one of

them having been the basis for the genus

Malacogzorgia and family Malacogorstidae

(Hickson 1904), Apart trom the complete absence

of calcareous skeleton, Melacogorzia capensis is

identical to Trichegergia flexilis and both have lone

been placed in the family Chrysogargiidae on the

basis of Lhe typical chrysegareiid sclerues of 7

Jlexilis (Kokenthal 1924; Bayer & Muzik |976b),

Tichogorgia viola Deichmann, 1936, from the Gulf

of Mexico is morphologically similar and also has

sclerites of the chrysogorgtid (ype, Colonies of

Trichogorgia lyra Bayer & Muzik, 1976, from the

southwestern Caribbean, which sre not unlike

Malacogorgia (=Thichogargia) capensis in some

respects, like it lack sclerites but have valeareaus

deposils in Lhe avis:

Gorgonacean families with an unjointed, more

ar less calcified axial skeleton without a hollow core

are Primnoidae, Ellisellidae, (Walukellidae, and

Chrysogorgiidae. Among these, only Trichogorgia

capensis in the Chrysogorgiidae has an axis devoid

of calcareous deposits,

The axis of Dendrobrachia lacks calcification as

well as a hollow core, thus resembling that of

Malacogorgia ( =Trichagorgia) capensis and, like it,

also lacks selerites. Some species of Plunizorgia

are very scantily provided with selerites of very small

size and considerably different from the prevalent

chrysogorgnd type, and Ua/ukel/a yanii Bayer has

so few as Lo be virtually sclerite-free. However, both

genera are inhabitauts of shallow, reef-associated

waters, and the axis in both develops a strong, even

massive, calcareous holdfast. In the case of

ffafiikella, the holdfast has conspicuous

longitudinal graoves and ridges thal are sharply

serrate or laciniate, but che arborescent part of the

axis is only weakly ridged, as in many. other

gorgonacean genera (Bayer 1955), The holdtast of

Dendrobrachia js not calcified und forms a

minimally spreading encrustation on solid objects.

Anatomically, the polyps of Dendrobnnhia agree

with those of Chrysogorgia, Stephanngirgia,

Trichogergia and talukelle. Unfortunately, the

anatomy and listology of those genera have not

been described, but direct comparison with serial

sections of Chrysagersia elegans (Verrill),

Stephanogertia Jaulkneri (Rayer), Tichogareia lyra

Bayer & Muzik. and I/ulukella panii Bayer reveals

Similar features,

D. M. OPRESKO & F. M. BAYER

1542 10K¥Y | —

1542 1oKy un | 1542 © 10KY iSdum

Fig. 11. Dendrobrachia multispina sp. nov., axis. Top, lateral view of larger branchlet; bottom, axial spines of same

USNM 87770. Stereoscopic pairs, SEM.

RFEGISCOVERY OF THE ENJGMATIC COLLENTERATE DENDROBRACHIA 1

epidermis

gastrodermis

lateral mesenterial

filaments

asulca| mesentery

folds of

pharyngeal wall

lateral

pharyngeal mesenteries

wall

nematocyst

siphoraglyph

nematocyst

sulcal mesentery

mesogloea

Pig. 12. Dendrobrachia puucispina sp. nav, Cross section of polyp, somewhal oblique, at level of pharynx. Scale

= 0.15 mm.

DM, OPRESKO & F. M. BAYER

mesogloea of pharynx

epidermis

mesogloea

epidermis

basement membrane

of epidermis

gastrodermis

mesogloea of pharynx

REDISCOVERY OF THE ENIGMATLO CORL ENTERAL DENDROBRACHIA 19

In spite of the general similarity of the palyps

of Dendrohrachia with those of the uncalcilied

species of Chrvosogorgiidae, the unique nature of

the axial skeleton convinees us that the Family

Dendrobrachiidae should be maintained, at least

until material becomes available for a more

comprehensive investigation and comparison of

anatomical and histoloyiceas features of all the

species mvolved.

Acknowledgments

We would like to thank Mr Shane Parker of the

South Australian Museum for providing the

specimens of Dendrebrachia paucispina, Dv Esther

Peters for preparing the histological sections, Ms

Cynthia Ahearn tor eritical-point drying of polyp

material for whole mount preparations, and Mr

Walter Brown, chiefof the SEM laboratory at the

U.S, National Museum, for taking the SEM

photographs, This work was supported by the

Smithsonian Institution, Washington, D.C, and by

Oak Ridge National Laboratory, Oak Ridge,

Tennessee, The Australian material was collected

during the South Australian Mnseum’s continuing

deep-sea trawling programme, undettaken with the

Benerous cooperation of the masters and crew of

numerous trawlers and research vessels. ‘Ihe

specimen of Dendrobrachia multispine trawled in

the Straits of Florida by R/V “Gerda” was obtained

during the DeepSea Biology Programme of the

University of Miami, supported by the U.S,

National Science Foundation and the National

Geographic Society.

References

Bavuk, PM 955) Conttiburians ro the nomenclature,

systematics, and morphology of the Octocorallia. Frac.

US. Nat. Mas 105, 207-220, pls. 1-8.

1958) Octocorallja. Pp. 163-231. Jn Moore, RK.

Coclenterata", (Geolowicer! Society of Ameriva &

University Of Kansas Press),

(1974) Studies on the anatumy and histology of

Plexaura horromatla in Florida. in Bayer, Fo M. &

Weinheimer, A, J. (Eds) “Prostaglandins from Plexaura

homonrallay ecology, utilization and conservation of a

major médical marine resource”. Stud. Trop, Oveanogr.

12 62-100, figs 1-62.

_ & Muzik, KR. M. (19768) A new solitary

vctocoral, Tiiaroa tauhois gen. et sp. nov. (Coelenterata:

Protoalcyonaria) from New Zealand, J. R. Soe, New

Zeatund 6(4), 499-515, figs. 1-10,

(19766) New genera and species of the holaxonian

Family Chrysegorgiidae (Octococorallia: Gorgonneea).

4ool. Meded. Leiden 50(5), 68-90, fies 1-10, pls |-7.

et al. (ds) (1983) “UWastrated trilingaal glossary

of morphological and anatomical terms applied to

Octocorallia”, (E, J, Brill’ Br W. Backhuys, L¢iden.)

Bouition, J. & HOUvENAGHEL-CREVECOELIR, N, (1970),

Etude monographique du genre Heliopora d¢ Blainville

(Coenothecalia-Aleyonana-Coclenterata), Mus, Ray,

Afrique Centrale. Tervuren Annmules (Serie in-k,

Sviences Zoolopiques) 17BL 7 vive 83, pes. 1-12

Brook, G. (B89) Report on the Antipathana. Kepts Sci.

Res, Vavage Challenger, Zool. 32, vi + 222.

CHESTER, W. M. (1913) The structure of the gorgonian

coral Pseudeplexaura. crassa Wrighi and Studer, Pree,

Amer Acad, Arts Sci, 48(20), 737-773, pls. 1-4,

Dickmann, EB. (1936) The Alcyonaria of the western

part of the Atlantic Ocean. Mem, Mus. Comp. Zoal.

Harvard 53, 1-317, pls, 1-37-

Hickson, S. J. (895a) Antipatharia. B40 ins Wright. B.

P. The scientific results of the Challenger” Expedition.

Natural Science 7, 7-75,

_____ (189Sb) The anatomy of Alcyonitm digitdturt

Quart, J. Microsc, Sei. (new series) 37(4), 343-388, pls.

36-39,

—____ (1904) The Alcyonarta of the Cape of Ciood Hope,

Part WH. Marine Invest, 8. Africa 3, 21\-239, pls. 7-9,

Hyman, L. H. (6940) “The invertebrates: Protozoa

through Ctenophora”. (McGraw-Hill Book Company,

New York.)

KUKENTHAL, W. (1924) Gorgonaria, Das Tierreich 47,

i-xxvii4 1-478 209 figs.

THomson, J. A. (1910) Note on Dendrobrachia fallax, a

tare and remarkable antipatharian, J, A. Microscep,

Sac, 1910, 142-143.

Van Pescn, A. J. (1914) The Antipatharia of the Siboga

Expedition. Sibega-Expeditie Monographe 17, 1-258.

(B. J, Brill, Leiden)

Fig, UO. Dendrabrichia paucispina sp. nov., A. Cross section of body wall with Mesentery sliowing longitudinal

mesoyloeal plates far insertion of retractor muscles on both faces; B. Cross section of body wall showing epwertnal

lobes and mesentery with longitudinal mesogineéal plates for retractor rouscles on beth faces; Scale al A — 0,05

mm; scale ai B = 1.1 mm.

THE DISPERSED CUTICULAR FLORAS OF SOUTH AUSTRALIAN

TERTIARY COALFIELDS, PART 1: SEDAN

BY A. I. ROWETT*

Summary

Dispersed cuticles were recovered from the three seams of the Sedan Coalfield and sixty-two

parataxa have been identified. The floras of the two younger lignite seams are distinct from the

underlying lignitic clay. The lignites are dominated by Proteaceae cuticle types with one cuticle

type, identified as the cuticle of Banksieaephyllum laeve, i.c. Banksieaephyllum aff. B. laeve in

abundance. The older lignite seam is distinguished by a large monospecific Myrtaceae component

and a cuticle type identified as aff. Agathis (Araucariaceae). Other families represented in both

seams include the Casuarinaceae, Elaeocarpaceae, Myrtaceae and Podocarpaceae. The flora of the

under-lying lignitic clay is dominated by Lauraceae cuticle types.

The presence of Banksieaephyllum aff. B. laeve in the Sedan lignites suggests either an Oligocene-

Miocene age for the lithotype or a longer stratigraphic range (extended lower limit) for the leaf-

fossil.

KEY WORDS: Palaeobotany, Tertiary, Eocene, dispersed cuticles, Sedan, South Australia

Tanners af ihe Revu) Seelery of S. duse (99, LIS, 21 30,

THE DISPERSED CUTICULAR FLORAS GF SQUTH AUSTRALIAN TERTIARY

COALFIELDS, PART 1: SEDAN

by A. |b. RowrrtT*

‘Suminary

Rowert, A. T, (1991) The dispersed cuticular floras of Sonth Australian ‘Tertiary Coalfields, Part } Sedan,

Trans. R. Sow. 8, Aust. 115(1), 21-36, 31 May, 1991,

Dispersed cuticles were reenvered tram the three seams of rhe Sedan Coulfield and sixty-two parawxa

have been identified. The floras of the two younger lignite seams are distinct from the underlying lignite

clay. The lignites are dominated by Proteaceae cuticle types with one cuticle type, identified as the cuticle

al Banksievephyllumn laeve, i.e Banksieaephyllien att, B, laeve in abundance, The older lignite seam is

distinguished by a large monospecific Myrlaceae courponent und a culicle type identilicd ws all, Avaliis

{Araucarisceae). Other families represented in both seams include the Casuarinaceac, Klacocarpaccac,

Myrtaceae und Padoearpaceae. Ve flora uf the under-lying ignite clay deniinaied by Laurgeeae cusicle

types.

he presence of Hunksiedephyliam att. B leeve in the Sedan lignites suggests either an Oligocene Miocene

age lov the lithotype or a longer strativraphic range (extended lower limit) for the tcat-fessil,

Kry Worps Palacoborany, Ternary, Eocene, dispersed citicles, Sedan, South Australia

Introduction

Chere are phumerous coalficlds found throughout

South Australia tangiog in age from Permian to

Tertiary. The Teriary deposits represent large

resourecs of Jow-rank coal (lignite) of Middle to

Lute Fogene age that are distributed throughour

sedimentary basins in the southern part of the State

(S,A,0.M.B. 1967), These deposits include the

Lochiel, Clintun, Beaufori, Bowmans and

Whitwarta deposits of the Northern St Vincent

Busin, Kingston in the south-east and rhe Anna and

Sedan deposits on the edge of the Murray Basin.

Three of these localitics, Sedan, Loehiel and

Kingston, are Lhe subject of this study,

Previous dispersed cuticl: investigations, the

majority of which have been undertaken in the

northern hemisphere (eg, Peters 1963; Schneider

196; Kovach & Dilcher 1984; Rowett 1986!) have

revealed that hgnites gencrally contain an-

abundance of cuticle fragments which can. be

assigned to cither natural or form taxa. These data

muy subsequently provide information about rhe

source vegetalion associated with the formauon af

the specific lithotype us well as the past environment

and climate. Such information is currently being

noxl by the wuthor in the correlation and dating

of coal deposits.

The Sedan coaifield is locared 75 km northeast

of Adelaide (34" 34°, 139° 18’) on the tnarein of

~ Dept of Bolany, University of Adelaide, G.PO, Box aye,

Adelaide, S. Aust, SN0L.

‘Rowell, A, 1, (1986) Megalossil and Micrafossil Flotas

of the Curlew houndation, Queensland. Ph,L) thesis,

Borany Dept. University at Adelaide. Unpubl,

the Murray Basin, between the Mount Lofty Ranges

and the River Murray (Fig. 1). Exploration in the

vicinily of (he nearby Anna deposit dates back ta

the 1920's but it was not until 1979 that the Sedan

lignite was discovered, The deposit fics inva sorth-

trending trough § km wide and 13 km long and

comprises [wo main lignite seams, The upper seam,

which is (he most economically viable, is continuous

and upto & m thick, The lower seam is lenticular

and averayes 5 im in thickness (S.A.D.M.E. 1987).

‘ertiary (oraminiferal bidstratigraphy (McGowran

1989) wnd a preliminary palynologi¢l analysis (N,F-

Alley, S.A.D.MLE, pers. comm.) have dated the

lignites as Late Focene-Oligacene,

Materials and Methods

Drill core samples from both seams and jhe

underlying lignitic clay taken from Sedan core CSR

CO74R (7.5 cm diameter), housed in the

S.A.D.M.E. core Jibrary are used in this

investigaGon. Examination of these samples

revealed considerable amounts of heavily

carbonised dispersed cuticle and wood fragments,

Cuticle fragments were extracted from 100g of

inatrix by modifying the maceration technique

ouilined by Chrisiophel ef a/, (1987) to inglude an

additional preliminary step whereby the crushed

sample is placed in warm Schulze solution for

approximately four hours prior Loa dilute (50%)

hydrogen peroxide solution step. The use of an

additional oxidizing step assures (le removal of any

organic material from the culicle and makes

Possible more aceurate ideniificutions and

deseripuoas of cuucle types. The cuticle fragments

were collected by passing the maceral througha

A. I, ROWETT

{Lachiel

«Sedan

Gatden Grave

Kingston

Atijlesea Yalloury

Cumulative Lignite Thickness

(metres)

Contours of overburden thickness

(metres) aa 45

QO 5

a ce ee NR on A

kms

SEDAN DISPERSED CUTICLES 23

single Mine pore sieve (150 jam diameter pore). The

use of a muluple sieve series Was considered

unnecessary because Of the small range in fragment

size} none of the cuficles were greater than 2 mm’,

Examination of the cuticles revealed (hat no single

culicle paralrxon was size specific.

Only cuticle types (parataxa) that displayed

stomlates were considered in the overall analysis as

the stomatal features are often diagnostiv of a

particular plant family, thereby greatly improving

the chances of determining rhe modern aff|nities

of fossil cuticles.

Five replicate samples of 200 cuticle fragments

each (Rowet! & Christophel 1990) were obtained

from core material reeovered from three depths,

54.8 im (upper seam), 67.5 m (lower seam) and

75,3 m (lignitic clay). The frequency data presented

for both individual parataxa and modern families

(ic. Table 1) isa percentage calculated fram the total

ol the five replicates of each lithorype.

The Dispersed Cuticle Mlora

The three samples thar contribute to the Sedan

cuticle flora are separated by substantial intervals

(approx. 10 m) of barren sediment (K, Wiggles-

worth, 5.A.D.M.E. pers, comm.,).

Lignilic Clay: The Vlora of (he basal lignitic clay

is dominated by the Lauraceae which forms 83.0%

of the total cuticle sumple (Table 1). This

component is particularly diverse, containing I

parataxa, of which the most frequent are parataxa

No. AA 006 (30% of total sanyple, Figs 4&5) and

No, 55 00S (19% of total sample; Figs 6 & 7), A

uuober of these Lauraceae parataxa show possible

alfinines to the extant genera Andiendra and

Cryptocarva, In Endiendra the epidermal cells are

angular in appearance with irregularly thickened

anticlinal walls, stomates are generally clongate with

narrow Bul prominent culicular scales (ex. No AA

QOU7, Fig. 2), These scales lie between (he sunken

guard cells and the overlying subsidiary cells.

Cryptocarya is generally characterised by epidermal

cells, with smooth anticlinal walls of uniform

thickness, Stomates are generally rounded with

prominent reniform cuticular scales. These scales

aré dark-staining and give a bulterfly-hke

appearance to the slomata (No, AG 005, Fig, 3).

The Myrtaceae component (3.1%) is represented

in the Mora by the cuticle types No. LC ON (Figs

8 & 9) and No. S5 00d (Figs 11 & 12), with the latter

being the most abundant, The most distinctive

cuticular feature of the Myrtaceae is the oil gland

lid cell, These cells, which may occur on either/or

both cuticular surfaces, are generally isodiametric,

divided by a curved to sinuous sinus, surrounded

by a ttumber Of concentric circles of radially

arranged epidermal cells and display some degree

of cuticular thickening.

Falcatifolium afl, £ australis Greenwood (Figs

{3 & 14) is the principal contributor to the small

Podocarpaceae component (0.8%). The parataxon

is suggested as having an affinity to Falcarifalium

australis on the basis of the circular appearance of

the stomata, a fearure considered [o be distinctive

by Greenwood (1987), who first identified the

species fram the Upper Middle Eocene fossil (laura

ofthe Anglesea locality. A very small Proteaceae

component (0.2%) is also present in this lithotype,

Lower Lignite Seany The cuticle flora of the

lower lignite seam is sipmificantly different trom (hal

al (he underlying clay, The most obvious differences

include the replacement of the Lauraceae us the

principal (loristic component hy the Proteaceae, the

TABLE 1, Thre culicte Jrequencies (0) of extant plant families represented in the three Sedan lithological unily, The

Jamilies represented are Podocarpaceae (POD), Araucuriaceae (ARAUC), Myrtaceae (MYRT), Elaeocaurpaceae

(ELAEO), Proreaceae (PROT), Lauraceae (LAUR) Casnarinucede (CAS) The OTHERS calezary represents all other

culicle paratasa whese modern fumily affinities ure unknown,

LOCALITY POD ARAUG MYRT ELAEG PHOT LAUR CAS OTHERS

SEDAN

54.68 m 24 = O.4 13 33.0 a7 a1 1,2

SEDAN

67,50 1m 3.0 2.0 i4.5 a0 40.4 os O5 25.6

SEDAN

75,30 m 08 - EDI OA 0.2 $3.0 - 12.5

Fi. 1, Map showing the extent and thickness of lignite in Lhe Sedan deposit. The awn of Seda is situated ahuve

ihe deposit. Inset: Map of castern Australia showing the location of (he Sedan, Lochiel and Kingston coal localities

relative to the Tertiary Megafossil localities of Golden Grove (Eocene), Anglesea (Eocene) and vallousy (Olwocene)

SEDAN DISPERSED CUTLCLES 25

presence of an Araucariaceae and Casuarinaceae

component and increased abundances in the

Myriaceae and Podocarpaceae components. The

Elacoearpaceae is present For the first time.

As Stated above, the flora is dominated by the

Proteaceae with parataxon No. Si O01 the major

contributor, Parataxun No, 51 O01 (Figs 17 & 18)

fas a very distinctive cuticle with slightly raised

bruchyparacytic stomates, small simple hairs

associated with a single epidermal cell which are

common ower the stomatiferous surface and large

multicellular hair bases. These features are

characteristic af the cuticle of the fossil leal genus,

Banksieaepiy lian (Proteaceae). The genus was first

used to deseribe six fossil leaf species, ie &

engustunt, B. acwminatum, B laeve, B. abavatuen,

& piinatumn and B& fastigaruim, with affinities to

the modern genera Buritsia and Drvandra

(Cookson & Dyigan 1950}. These leaves were

recovered [ram the Oligocene aged brown coal sean

at Yallourn, Victoria. At present eleven species of

Banksieaephyllum are recopyised. These species

have all been recovered from southeastern

Australian localities ranging from Early Eocene to

Barly Miocene in age and include. the Middle

Eocene Maslin Bay deposit (Blackburn 198); Hill

& Chiistopbel 1988), Early Eocene Deans Marsh,

late MiddJe Eocene Anglesea, Middle Eocene

Golden Grove, Late Bovene-Oligocene Cethana,

Miudie-Late Eocene Loch Aber, Oligovene Pioneer

Oligocene-Early Miocene Loy Yang (Hill &

Christophel 1988) und Mrocene Morwell (Blackburn

1985). OF the eleven described species, cuticle of

Banksieaenhyllunm laeve from the Yallourn Oligo-

cene brown coal flara (Cookson & Buivan 1950)

and Miocene flora of the Morwell seam at Morwell

most closely resembles parataxon No. SI 001.

‘The joereased Myrtaceae component (14.5%) is

represented in the flora by a single parstaxan No.

53 G01 (Figs 15 & 16)° This parataxon only occurs

in (his lithotype and is distinct from the other Sedan

Myrtaceae by the robust nature af the generally

larger cuticular Soarures.

The Casuarpacese component, allhough a minor

contributor to the overall Mora.is important in that

the presenee of the parataxon No. DM O07 (Figs

19 & 20), idenufied as Gwrinastome is the Sirst

dispersed cuticle record of the occurrence of this

well known ‘Tertiary genus outside of eastern

Australia, The cuticle of this parataxon is very

distinctive with stomates oriented perpendicular (o

(he longitudinal axis of the epidermal cells, The

slumates may fave one or two lateral subsidiary

cells. The genus was first reported from the

Anglesea deposit by Christophel (1980) as a

megafossil of the Casuarinaceac, Division

Gymnostomae, and has since been discovered in the

Moras of a number of Tertiary plant fossil localities

which tange from Middle Eocene to Miocene and

include Deans Marsh (Middle Eocene), the Curlew

Formation (Middle Eocene), Moranbah (Middle

Bocene; Scriven & Christophel 1990), and Yallourn

and Morwell open cut mines (Oligocene-Mincene,

Blackburn 1985), Grrinostema cone impressions

have also beer identified in the siloreres of the Eyre

Formation (Late Palaeocene-EBacene), Willilinchina

Sandstones (Eocene) and Eradunna Formation

(Oligo-Miocens) (Greenwood ef al’).

Parataxon No. AWF GO] (Figs 21 & 22), the sole

representative of the Araucariaceae component is

only found in this tithotype and may therefore prove

to be a useful local stratigraphic indicator. This

cuticle type is tentatively assigned to Agarhis on the

basis of the oblique orientation of stomata and the

curved to rounded nature of the epidermal cells,

The farge Elaeocarpaceae component [9.5%%) is

represented in this lithorype by two cuticle types

Nos.. 53 004 (Figs 23 & 24) and $3 005 (Fig, 25),

the former being the major contributor with 8.05%

of the cuticle sample. Both parataxa occur for the

first time in the Sedan deposit. Features of the

cuticle which are considered characteristic of the

family are the presence of hydathodes (Blavkburn

1985), Slaurocylic stomates and a prominent,

flarrow Outer stomatal ledge which bears small polar

extensions, termed an apiculate outer stomatal ledge

by Wilkinson (1979),

*Blackburn, D. T. (1985) Palaeobotany of the Yallourn

and Morwell coal scams, Palaeohotany Report No. 3,

State Eleciricity Commission of Victoria, Unpubl.

‘Greenwood, D. R., Callen, R. & Alley, N. F (1990)

Biostratigraphy of the Cainozoic Macroflora of the

southern Gyre Basin in South Austraha. Report, Seuth

Aibstraban Department of Mings and Energy, Unpubl,

bigs 2-7. 2. Pararaxon No, AA 007, afl) Endfandra, Lauraceag. The presence of prominent, narrow culjcular scales

and anedlar epidermal cells are features associated with the extant genus, Scale = 2D ym 3. Paraiaxon No, Ag

OOS utt, Creprocarva, Lauraceae. Whe presence of proninent reniform to crescent-shaped cuticular seales (Lees,

ariowed) whieh give the stomatal apparatiis a butterily-like appearances stamata and undulate epidermal cells are

features assbcialed with the extant genus. Seale = 24 am. 4. Paratawein No. 44 006, Lalrraceac, showing stomaniferois

surfdoe. Note the sinuous nature of the epidermal cells and poral trichome bases. Seale = 20 yin, 8 Pafataxof

No. AA 006, Lauraccac, shovviig the paraeylic stomatal arrangement and prominent cuticular scales, Scale ~ [2

wir. &, Paratanon No, AA 008 all, Crypfecarya, Lauracear, showing stomatiferous surface. Nore the smooth, woiformly

thickened anziclingl walls of the epidermal ells. Seale — 20 jun. 7. Parctaxon No. AA OOS al, Crprovorva, Lauraceae

showin pareytic stomatal arrangement dnd ereseent-shuped to rediform cutivular seules, Seale ~ 20 atin

SEDAN DISPERSED CUTICLES 7

Although showing an increased abundance

(3.0%), with the infroduction of parataxon No. LE

O09 (Fig, 20), the Podacarpaccae coniponent

remains 9 miner contributor al- this depth,

Paralaxon No. LE 009 is easily distinguished from

Falcatifolium ale F eustralis by the suwong

bultressing of anticlinal walls of the epidermal cells.

Upper Lignité Sean) The tipper seam flora, is

the most diverse with 30 paralaxa represented, the

majority of which are recognised as having alfimities

with the families, Proteaceae, Elaeovarpacear.

Myrtaceae, Podtcarpaceaer, Lauraccue and

Casuarinaceae. The Proteaceae daminates the flora

(93.0%) with two parataxa, Banksieaephvilum alt.

& laeve-and parataxon No, SI 002 (Figs 27 & 28),

in particular abundance. The only other fannilies

of any significance in the flora are the

Podocarpaccac and Elaeoearpaceae which record

frequencies of 2.1% afid 1.3%, respectively. The

Podocarpaceae component shows an increased

diversity in this seam with six parataxa represented.

The paratuxa which dominated the component of

the other floras, 18, Faleatifaliamt afl & australis

and parataxon No, LF 009, are again the major

contributors in this lithotype The small Myrtaceae

component is marked by the reappearance of

parataxon No. LC OL

Four different cuticle types comprise the

Elacocarpaceae component with parataxon No.

ALL O11 the most abundant, Cuticles taken from

mummified leaves ascribed by Christophe! &

Greenwood (1987) to Sfoenea/Eleeucarpus fro

€lolden Grove and Jeaf type “Serrate 1" described

by Christophe! ez v/ (1987) from Anglesea are

identical to those identified as parataxon No. ALL

OH ja the Sedan flora (Pig, 29),

Dispersed Caticle Descriptivns

The dispersed cuticle flora of the Sedan voalfield

is represenred by 62 cuticle paraiaxa but for ihe

purpose of this paper only the principle cuticle

paralawa, stratigraptilcally significant parataxa and

those with known modern affinities are described

here. Several of the paratawa have been previously

ideniitied by the author from other Eocene

localities, These and many more are included in the

NERDDC reference catalogue of Australian Bocene

cuticle types, All parataxon numbers are preceded

by an abbreviation of the type locality.

The terminology used in the description of Lhe

Sedan curtele parataxa has been derived from that

proposed by Stace (1965), Dilcber (1974) and

Wilkinson (J980),

Cuticle Parataxon No. AA 007

FIG. 2

Only the stomatiferous surface was observed.

Epidermal cells angular to rounded (type t-2,

Wilkinson 1979), becoming clangate over the veins.

Cells 8-24 vm in length, 6-L6 pm in width. Ann-

elinal wall irregularly thickened, smooth (uniform

thickness) to slightly beaded. Periclinal wall

irregularly thickened, smooth to finely granulate.

Stomata randomly oriented, uniform distribution.

Siomata 16-24 um in length, 616 pm in width.

Guard cells, sunken, cuticular thickening on poral

wall. Cuticular scales (i.e, labelled CS2 on Fig- 1,

Hill 1986), prominent, narrow. Stomatal Arrange-

ment paracytic, Stomatal Index (S1.) 14,

Subsidiary cells cells 2. Anticlinal wall, thin,

smooth, Periclinal wall thin, sai@arh,

Trichome bases common, uniform distribution,

poral, 4-8 radially arranged surrounding cells,

cuticular thickening around pore extending along

tadial wall of surrounding cells giving stellate

appearance,

Affinity: The parataxon is assigned to the

Lauraceae, with a probable affinity ro the extaue

genus Endiundra. The inodern Australian species

of this genus are characterised hy predominantly

angular epidermal cetls with irregularly thickened

anticlinal walls, elangate stomates with prominent,

though narrow curicular scales.

Cuticle Pararaxan No. AG 00S

FIG 3

Only the slomatiferous surtace was observed.

Epidermal cells undulate to sinugua (lype 3-5},

becoming elongate over the veins, Cells 20-40 jm

in length, (2-20 wm in width, Anticlimal wall chin,

smooth. Peticlinal wall irregularly iickened,

smooth to finely granulate (Occasionally finely

striate),

Stomata randomly oriented, uniform distmbution-

Stomuta 16-24 jam in length, 12-20 pm in wiih,

Guard vells, sunken, cuticwar rhickeving on poral

wall. Guard cell/subsidisry vell wall not always

Figs $12, 4, Parataxon No, LC Ol), Myrtaceae. The dark-staining subsidiary cells ane the sinaous nanire of the epidermal

cells charactense this parataxon Identivsl in cuticular morphology toa speeimens ron) Loclitel. Seale = 20 pm,

9, Parataxon No. LC O11, Myrtaceae, showing staurocytic stamatal arrangement, Scale «

20 pm. 10, Speemmen

of Parataxon Na LO O1, Myrtaceae from Lochiel, note the sume cuticular morphology as the Sedan spevimetis

and the presence of ab oil gland lid cell (ic. 1G arrowed). Scale — 20 pm. 11. Paraiayon No, S43 U04, Myrtacede,

showing stomatal arrangement, Note the stoongly siriate periclinal wall of the epidermal gells. Scale = 1 prt

12. Pararaxon No. $5 004, Myrtaceae, showing a heavily cutinized wil gland lid cell (ic, am owed). Seale ~ 20 wm,

SEDAN DISPERSED CUTICLES 29

evident (in which case the stomatal width dimension

is taken as a measure across the (wo Subsidiary cells,

instead of the usual guard cell width), Cuticalar

scales, prominent, reniform to crescent-shaped,

giving a butterfly-like appearance ta Ihe stomate.

Stomatal Arrangement paracytic, S.I- 6.2.

Subsidiary cells two. Afiticlinal wall smooth, thicker

than for epidermal cells and raised which produces

a cuticular fold that encircles and partially overlies

the stomate, Periclinal wall thin, smooth.

Tnchome bases common, uniforra disttibution,

poral, 4-8 radially arranged surrounding cells,

cuticular thickening around pore.

Affinity: The cuticle parataxon is assigned to

Lauraceae with a probable affinity to the extant

genus Crypiocarya. The madern Australian species

of this genus are characterised by epidermal cells

with smooth anticlinal walls; shape is variable,

Stomates are generally rounded with prominent

renilorm to crescent-shaped cuticular scales which

gives a butterfly-like appearance to the stomatal

apparatus,

Cuticle Parataxon No AA O06

FIGS 4-§

Only the stomatiferous surface was observed.

Epidermal cells rounded to sinuous (lype 2-5),

becoming elongate over the veins. Cells 12-36 pm

in length, 8-20 ~m in width. Antichinal wall

irregularly thickened, smooth to ridged (Dilcher;

1974), Penclinal wall irregularly chickened, smooth

to granulate,

Stomata randomly oriented, uniform distribution.

Stomata 16-20 pm in length, B-10 um in width.

Guard cells, sunken, cuticular thickening on poral

wall. Cuticular scales, prominent, narrow. $.1. 95.

Stomatal Arrangement paracylic. Subsidiary ceils

2. Anticlinal wall irregularly thickened, smooth to

beaded. Periclinal wall irregularly thickened,

smooth to granulate

Trichome bases common, uniform distribution,

poral, 5-6 radially arranged surrounding cells,

culicdar thickening around pore,

A/finity: The cuticle is assigned ta the Lauraceae

due lo the presence of paracytic sromates and

inconspicuous, sugken guard cells.

Cuticle Paralaxon Noa. 55 005

FIGS 6-7

Only the stomatiferous surface was observed.

Epidermal cells angular, becoming elongate over the

veins, Areoles. are well-defined, Cells 16-32 m in

length, 8-20 m in width. Antichaal wall chin,

smooth (occasianally bultressed), Periclinal wall

irregularly thickened, smooth to granulate to strate

Stomata randomly oriented, uniform distribution.

Stomata (8-24 m in length, 16-25 m in width,

Guard cells, sunken, Guard cell/subsidiary cell wall

absent (in which case the stomatal width dimension

is taken as @ Measure across the two subsitliary cells,

instead of the usual guard cell width). Curiculas

scales, prominent, reniform to crescent-shaped-

Stomatal Arrangement paracytic. Sl. 8.6.

Subsidiary cells 2. Anticlinal wall thin, smooth.

Periclinal wall irregularly thickened, smooth to

granulate te finely striate. Striations generally

perpendicular to the long axis of the sromate.

Epidermal cells that immediately surround the

stomate are occasionally dark-stalning, Trichome

bases are rare, only found over veins. Poral with 4-6

radially arranged surrounding cells. Bases are

thickened around the pore.

Affinity: The cuticle parataxon is assigned to the

Laurageac with & probable affinity to the extant

genus Crvplocarve due to the preserice of epidermal

cells with predominantly smooth anticlinal walls

and reniform to oresvent-shaped cuticular scales.

Cuticle Parataxyon No, LC Oit

FIGS 8-9

Only the stomatiferous surface was observed.

Epidermal cells undulate to sinuous (type 3-5),

becoming elongate-over the veins, Cells 16-32 min

length, 10-24 m iy width. Anticlinal wall irregularly

thickened, smooth ta beaded to shghtly buttressed,

Periclinal wall thin, smooth,

Stomata randomly oriented, uniform distribution.

Stomata 16-32 m in length, 16-24 m in width.

Guard cells, sunken, T-shaped thickening and polar

rods preseht, Outer stomatal ledge, prominent,

narrow, Stomalal Arrangement stauracylic (to

cyclacytic). Si, 10.6, Subsidiary cells 3-6, darker

staining. Anticlinal wall irregularly chickened,

Figs 13-18, 13. Parataxon No ABD OO), Faleari/olium all & ausialis, Podocarpaceae, showing urmingement of stormat

in short uniseriate row. Seale = 32 pm. 14. Parataxon No. ABD OO), Fulcarifolivn aff, F australis, Pacocarpaceac.

Note the shape of the subsidiary cells and the resultant circular appearance of stomata. Scale = 26 pm. 15. Parataxon

Na, 83 00], Myrtaceae, slowing the stomarilerous surface. Note the dark-staining subsidiary cells, Seale « 30 em.

(6. Parataxon No. 53 001, Myrlacede, showing an oi! gland Jit cell (avrawed). Nate fie sinuous nature of the lid

cell sinus, Scale ~ 30 um. 17. Parataxon No. Si 001, Banksieeephyliam alt. B laeve, Proleaceac, showing ihe

stomariferous surface, Scale »

20 pm, 18, Parataxon No. S1 OO!, Banksteaephylhar all. AL faeve, Proleaveac, showing

the sramatal arrangement, Note the short, cylindrical papillae are present an mast epidermal cells. The stoma

appear sihtly raised above the cuticle surface. Seale — 17 jun.

SEDAN DISPERSED CUTLCLES 4

smooth to beaded. Periclinal wall thin, smooth.

AUfinity: Myrtaceae. Although of! gland hid cells

that are usually found on myrtaccous leaves were

not observed on fragments recovered from this

locality, the general cuticular morphology Is very

similar’ to that of specimens recovered from the

Lochiel coalfield (Fig. 10) which are unequivocally

Myrluceae,

Cuticle Parataxon No, §5 004

FIGS 13-12

Only the stomariferous surlace was observed.

Epidermal cells sinuous (type 4-6), Cells 12-36 um

in length, 10-28 pm in width, Antiehnal wall

irregularly thickened, smoath to beaded to ridged.

Periclinal wall striate.

Stomata randomly oriented, uniform distribution.

Stomata 16-26 pin in length, 12-16 wm in width,

Guard cellé slightly sunken, Tshaped thickening

amd polar pods present. Guter stomatal ledge,

prominent, marrow, Stomacal Arrangement

staurocytic, Subsidiary cells 3-3 more undilate in

outhoe Anticlinal wall beaded, radial wall often

absent or incomplete Periclinal wall ihin, striate,

Oil gland lid cells commen, prominent, constricted

al sinus, sinus straight, cark-staining, thicker cuticle

than found over epidermal cells, dimensions 32-44

am in leagth, 12-24 »m in width, Lid cell

surrounded by a number of circles of radial

arranged modified epidermal c¢ells. Strialions

radiate outwards trom lid cell,

A/fini. The presence of oi) gland lid vells and the

general stomatal morphology which includes

features like a well-defined, circular puard cell

vomplex and a prominent stomatal ledge, indicate

the cuticle js of the Myyraceae.

Cubcle Parataxan No. ABD 001

FIGS 13-14

Only the stomatiferous surlace was observed.

Epidermal cells angular (predominantly

rectangular) arranged in longitudinal rows onented

parallel (6 the long axis of the leaf, Cells 32-68 jum

in length, 20-64 wm in width, Anticlinal wall thin,

smooth, Periclinal wall granulate:

Stomuta in poorly defined spiseriate rows Which

suggests the specimen is of the adaxial surface of

the leaf, The stomata are oriented parallel to the

long axis of the leal. Slomata 32-44 jum in length,

32-40 ym in width, Guard cells, sunken, poral

thickening present. Outer stomata) ledge,

prominent, thick, broad with polar extensions over

radial anticlinal walls, Florin rings evident. Stomatal

arrangement paratetracylic, circular 14 appearance,

Subsidiary cells 4, crescent-shaped lateral cells larger

than wedge-shaped polar cells. Anticlinal wall thin,

smooth. Periclinal wall irregularly thickened,

granulate to striate.

Affiniv: The cuticle parataxon is assigned to

Fulcatifolium (Podocarpaceae), wilh a possible

affinity to the fossil species F australis, This species

was described from the Anglesea deposit by

Greenwood (1987) who noted the distinctive circular

appearance of the stomata, a feature alse exhibited

by the Sedan specimens.

Cuticle Parataxon No. SI O0T

FIGS {7-18

Hypostomatic, Adaxial epidermal cells angular

becoming elongate over yeins. Cells 20-36 um in

length, 12-28 um in width, Anticlinal wall thin,

amooth. Periclinal wall irreeularly thickened,

smocth ta finely granulate Trichonte bases

common, single to multicellular, up to 7 basal cells,

heavily cutinized, wichome scar evident,

Cutiele chinner than on adaxial surface. Abaxial

epidermal cells (angular to) rounded to undulate

(type 3), becoming elongate over veins. Cells 16-40

po in length, 8-16 um in width. Anticlinal wall

irregularly thickened, smoorh ta beaded, Periclinal

wall irregularly thickened, smooth. to papillare,

Papl\llae present on most epidermal cells, shert.

cylindrical, truncale apex, up ta 16 pm in length.

Large papillae common, arise from multicellular

bases (2-celled), basal cells heavily curinized, cuticle

of papillae also thick, up to 24 um in length, dome-

shaped. Stomata randomly oriented, ualforn)

distribution. S.1. 15.5. Stomats [8-24 jum in length,

12-22 am in width, Stomatal arrangement

brachyparacytic. Guard cells slightly raised, show

Pigs 19-25. 19. Parataxon No. DM 007, Gymnostoma, Casuarinaceae, The delicate nature of this cuticle makes recovery

difficult aod stamates are rarely preserved. Scale

40 um, 20. Paralaxon No, DM 007, Gyrinostonta, Casuarinacese,

a slightly better preserved fragmen( showing a scament of a number of stomatal rows, Note te perpendicular

orlentation of the stoma(a relative co che intermediate epidermal cells. Scale = 20 par. 21. Parataxon No. AWE

OO), all. Aguifis, Araucariaceac, Note the oblique orientation of the stamates and (he prominent Florin rings

associared with the stainates, Scale = 40 um. 22. Parataxon No, AWF 001, al. Aga/his, Araucariaceae, showing

the stomatal arrangement and the darker staining Florin fing which overlies the subsidiary cells. Scale

20 jam.

24, Parataxon No. 53 004, Blaeocarpaveae, showing stomatiferous surface, Scale = 20 am, 24 Paratavon No. 53

004, Elacocarpaceac, showing (he Anomucytic and staurocytic stamatal arrangement. Note the apiculate cater stomatal

ledge, 1.e ledge with small polar protrusions, on ihe guard ells. Seale ~ 20 um. 25. Parataxon No. 53 008

Hlavocarpaccae, showing Ihe stomanferous surface, note che staurovytic stomatal arrangemen|. Seale = 20 am.

32 A. 1. ROWETT

Figs 26-29. 26, Parataxon No. LE 009, Podocarpaceae, showing distinctive buttress thickening of the epidermal cells.

Scale = 40 um. 27. Parataxon No. SI 003, Proteaceae, showing the stomatiferous surface. Note the broad, thick

stomatal ledge and prominent T-shaped thickening on the guard cells. Scale = 40 xm. 28. Parataxon No. SI 003,

Proteaceae, showing the brachyparacytic stomatal arrangement. Note the prominent striations on the subsidiary

cells. Scale = 19 pm. 29. Parataxon No. ALL O11, aff. Sloanea/Elaeocarpus, Elaeocarpaceae. The thin guard cells

are surrounded by, up to 6, dark staining subsidiary cells. Scale = 32 ym.

slight poral thickening. Subsidiary cells 2. Anticlinal

wall thin, smooth. Periclinal wall irregularly

thickened, smooth to finely granulate, slightly

thicker cuticle, slightly darker staining. Glandular

bodies present.

Affinity: The cuticle parataxon is assigned to the

Proteaceae on the basis of a brachyparacytic

stomatal and multicellular trichome base

arrangement. The cuticular features are identical

to those of the cuticle of the Yallourn open cut fossil

Banksieaephyllum laeve. The Sedan cuticle type is

therefore identified as Banksieaephyllum aff.

B. laeve.

Cuticle Parataxon No. S3 001

FIGS 15-16

Only the stomatiferous surface was observed.

Epidermal cells undulate (type 3-4). Cells 36-64 pm

in length, 12-36 pm in width. Anticlinal wall

SEDAN DISPERSED CUTICLES w

irregularly thickened, smooth. to beaded. Periclinal.

wall thin, smooth.

Stomata randomly oriented, uniform distribution.

Stomata 20-28 yam in length, 24-28 jen in widlh,

Guard cells not sunken, granulate, small T-shaped

thickening present, Outer stomatal ledge prominent,

broad, dark-stuining. S.[. 14.6. Stomacal

arrangement staurocytic. Subsidiary cells 3-5, dark

staining, Anticlinal wall irregularly thickened,

smooth to beaded, Periclinal wall thick, smooth,

Oi) gland lid eclls rare, isodiametric, slightly

constricted 4 sinus, sinus undulate (2-3 waves), up

to 10 scarcely modified to sadially arranged

surrounding cells, dimensions 32-48 wm jin leneth,

40-48 ppm in width,

Affinity: The cuticle parataxon has been assizned

to the Myrtaceae due to the presence of oil gland

lid celle of the lype seen on modern members of

(he Farmnly,

Cuticle Parataxon No, DM O07

BIGS 19-20

Stomatiferous stem/shoot material was observed,

Epidermal cells angular (usually hexagonal),

arranged in longitudinal rows oriented parallel to

the long axis of thé shoot/stem. Cells 8-28 yum tp

fength, 16-20 wm in width. Anriclinal wall chin,

smooth, Periclinal wall irregularly thickened,

smooth to granulate.

Stomata arranged in a uhiseriate row near the

shoar/stem margin, oriented perpendicular co the

long axis of the shoot/stem. Stomata 2()-24 pm in

length, 8-12 «m in width. Guard cells not sunken,

Outer stomatal ledge, prominem, broad, raised.

Stomatal Arrangement none to paracytic.

Subsidiary cells 0-2. Antictinal wall, thin, smooth.

Periclinal wall irregularly thickened, smooth to

granulate.

Affinizy: The cuticle parataxon has been: assigned

to Gymnostoma (Casuarinaceac) on Uhe basis of

predominently angular epidermal cells and the

perpendicular orientation of stomares relative to the

long axts Of the shoot (Dilcher er ai. 1990),

Curicle Parataxon No. AWE 001

FIGS 21-22

Only che stomatiferous surface was observed.

Epidermal cells angular to rounded. Cells between

stomatal bands sare generally arranged in

longitudinal rows oriented parallel to the long axis

of the leaf, Cells 28-115 pm in length, 16-48 ~m

in width. Anticlinal wall thin, sooth. Periclinal

wall irregularly thickened, smooth to finely

grinulate.

Stomata arranged in shore discontinuous

Uniseriate rows, oriented obliquely, transverse to

parallel to the long axis of the leaf. Stomatal rows

separated by 2-5 rows of reclangular epidermal

cells, in narrow stomatal bands. Stomata 40-A4 jam

in leneth, 20-40 pm in width. Guard cells

sunken. Stumatal Jedge, prominent, narrow, Guard

cell/subsidiary cell wall heavily cutinized, raised.

Florin rings evident. Stomatal Arrangement

cyclocytic.. Subsidiary cells 4-5. Antictinal wail

irregularly thickened, smooth to beaded. Periclinal

wall irregularly thickened, granulate to striate.

Affinity: The curiele parataxon has been assigned

lo the Araticariaceae, The predominantly oblique

onentation of the stomata ald rounded epidermal

cells Suggests an affinity to Aga@ihis (Cookson &

Duigan 195]; Stockey & Taylor 1981; Hill &

Bigwood 1585, £987).

Cuticle Parataxon No. 53 004

FIGS 23-24

Only the stomabtferous surface was observed,

Epidermal cells undulate (type 3), Cells 12-32 pot

in lergth, 8-16 wm in width. Antictinal wall thin,

smooth. Periclinal wall thin, smooth.

Stomata randomly oriented, uniform distribution.

Stomata 12-20 am in length, 12-16 wm in width.

Guard cells not sunken, Outer sramatal ledge,

Prominent, narrow, apiculate, Stomatal

Arrangement staurocytic to cyclocytic, Subsidiary

cells 3-5, Anticlinal wall, thin, smooth, radial walls

may be poorly defined, Perictinal wall thin, smoot

Trichome bases common, over veins, poral, 7

radially arranged surrounding cells, cuticular

thickening around pore, Hydathodes rare, over

veins,

Affinity: The cuticle parataxon is placed in the

Flaeacarpaceac due to the presence of hydathodes,

an apiculate stomatal ledge and staurocytic stomatal

arrangement,

Cuticle Parataxon No, 33 005

FIG 25

Only the stertanterous surface was observed,

Epidermal cells angular. Cells 10-33 pmiin length,

3-23 pm in width, Anticlinal wall thin, smooth.

Periclinal wall granulate:

Sromata randoinly oriented, uniform distribution.

Stomata 12-20 un in length, 10-15 wm in width.

Guard cells, nor sunken, T-shaped thickening

present. Outer stomatal ledge, prominent, narrow,

apiculate, Stomatal Arrangement staurocytic.

Subsidiary eclls 2-5. Anticlinal wall, thin, simoath.

Periclinal wall granulate. Hydathodes rare,

Affinity: The cuticle parataxon is placed in the

Elaeocarpacrse due io |he presence of hydarhodes,

an apiculate slomatal Jedge and staurocytic stomatal

arrangement.

SA A. 1, ROWELT

Cuncle Paratoxon No. LE 009

FIG. 16

Only the stomatiferous surfyeo was observed.

Epidermal cells rectangular, arraieed in

lougitudinal rows oriented parallel lo the long axis

of the leal. Cells 8-36 jon to length, 8-18 gm in

width, Anticlinal wall buciressed. Periclinal wall

thin, smooth. Stomata arranged in uniseriate rows,

often short, occasionally merged, onented parallel

to the Jone. axis of the leat) Sromata 14-28 am in

length, 12-18 pm in width. Stomatal arrangement

paratetracyric. Giuard eclls sunken, Stomatal ledge

praminent, irregularly thickened. Subsidiary cells

4, heavily cutinized, lateral cells larger than polar

cells. Polar vells may be shared. Anticlinul wall

buttressed, Periclingl wall thick, smooth. Flona ong

evident.

Affinivvs Although very fragmentary the general

cuticular morphology of specimens suggests an

affinity to the Podocarpaceae,

Cuticle Parataxon Wa. S71 003

FICS: 27-28

Only the stomatal surface was observed.

Epidermal cells sinuous (4), 64-164 «im in Jength,

$6-80 ym in width Anticlinal wall irrcevlarly

thickened, sinoolfi to finely beaded to ridued.

Periclinal wall yranulate, Stomata randomly

onented, uniform distribution. Sramara 34-36 ~m

in Jenwth, 32-38 pn in width. Stonaatal arrangement

brachyparacytig, Guard cells sunken, broad polar

rods, T-shaped thickening and poral vhickening

presen(. Qu(er stomatal ledge promiient, ‘ery

broad, Subsidiary cells 2. Anticlinal wall beaded.

Periclinal wall striate. Trichome bases common, 1-2

basal cells, heavily cutinized, raised anriclinal wall,

BeS scarcely moditied, radially arranged

surrounding cells, Striations radiate outwards anta

other epidermal cells.

Affinity: The cuucle pardivden is assigned to the

Proteaceae due to presence of brachyparacytic

Shortie and rublicelular trichome bases.

Culicle Paratayon No, ALT. Ol

PIG, 29

Oily stomatiferous surface observed, Epidermal

cells angular io rounded, becoming elongate over

veits Cells 8-40 pam in length, 8-20 wm in width

Anticlbal wall thin, smooth. Penelinal wall thin,

smooth. Starnata randomly oriented, uniform

distribution within large areoles. Spomata }2-24 pm

in length, 12-76 pm in width, §.f. 15.6 Stematal

Urmungemen! aclinocytic to sraurocyric. Guard cells

pot lo shiwhily sunken. Subsidiary cells 3-6, darker

staining than epidermal cells. Anticlinal wall chin,

smooth. Periclinal wall irregularly thickened.

smacth to slightly granulate. Outer stamatal ledge

prominent, narrow, Polar rads and some T-shaped

\hickening present on guard cells. HEydathodes rare,

over veins, 28-40 nm in lengch, 16-20 am in width,

Affinity The cuticle paratuxotr ay placed ln che

Elaeocarpaceae due to the presence of Indathodes,

an apiculate stomatal ledge and staurocytic stomatal

arrangement.

Floristic Comparisan of Samples

The tloras of each of the three samples possess

a number ol floristic features that distinguish them

from one another. The flora of the upper seain is

characterised by the dominance of the Proteaceae

ctomporent| and of one parataxon in particular, No,

St O01 which matches the cuticle of

Banksieaephyllum laeve (Wigs U7 & 18).

The flora of the lower lignite sean has a similar

overall composition to that of the younger lignite

scam bul with a aumber of diflerenecs, These

include; 1) the presence of parataxk No, $2 001 and

No. AWF 001 both of which are absent trom the

other floras as well #5 being the sole contributors

fo (he Myrtaceae and Araucariaceae components

respectively, Both paralaxa are potentially useful as

stratigraphic Indicators, at least locally within rhe

depositional! basin; 2} an increased abundance of

the Podocarpaceay, Myrtaceac, Elacocarpaceac and

Casuarinaceae componeits which when combined,

account for more than 25% of the flora, In no other

lithowype do these components artain the same levels

of significance emher individually ot combined.

The flora of the basal clay is easily distinguished

from the ether Sedan floras by the abundance of

Lauraceae parataya, of which some ate related to

the extanr genera Exdiandra and Cryprocarva. and

the minor occurrence Of Proteaceae panitaxa,

Comparison of the floras of the three litharypes

reveals a distiner floristic difference between the (wo

hgnite seams and the basal lignitic clay seam. Both

lignite floras are dominated by the Proteaceae with

puratdxan Nu. $1 00) the mujer contributor. The

flora of the lignitic clay is very different with the

Lautaceae dominating and the Proteaceae reduced

to. an insignificant level. Irie also-evident that there

isa cortelaliun between the lithotype and dominant

floral type, ic the Proteaceae ure associated with

the liguire Whereas the Lauraceae are associated

with the clay.

Examination of modern sédimentars

enylronments has shown thal quite different leaf

floras (and by inference, different cuticle floras) are

found jn separate sedimentary envircinents

{Burnham 198%; Taggart 1988). Within the sanie

stratigraphic sequence these changes in environment

SEDAN DISPERSED CUTLICLES as

may be due to either hydrological succession Gr

climatic changes which subsequently induce

chatives 1 the plant community (Luly er af, 1980),

The correlation between the two lithotypes and the

two chispersed cuticle floras theretore indicates that

both floras are rhe product of different sedimentary

environments (Taggart 1985); ithe Lauraceae-

dominated lignitic clay and the Proteaceae-

dominated lignite, The lignitic clay is likely ta nave

heen deposited under open-waler conditions, with

mainly allochthonous, potentially extra-local

{Lautaceae-dominated flora), plant remains,

Whereas the henite represents swamp conditions

with muinly local deposition (ie. Proteaceae-

dominated flora).

Comparison with Other Australian Tertiary

Depasits

The dispersed cuticle floras of the Sedan deposit

inchide a number Of paralaxa Which are known lo

occur in other Australian Tertiary deposits. The

presefice of the very distinctive cuticle of

Banksfeaephvllum feeve in the Sedan lignites, Le

Banksteaephyllum aff, B faeve, would appear to

be of some biostratigraphic significance. A laeve

has to date only been reported from the Yallourn

(Oligocene) and Morwell (Miaceng) coal scans ul

the latrobe Valley and the Inkerman-Balaklava

lignites of the Bowman deposit, 100 kn north of

Adelaide (Blackburn 1985), The latter deposit is

located a shart distance south of the Sedan deposit

in the adjoining St Vincent Basin. Although the

stratigraphic evidence is limited, it is. consistent, with

& loeve only being reported from Oligocene-

Miocene coals and as one of these coal deposits fies

is clase vicinity to the Sedan deposit then a younger

age could be inferred.

Cunversely, based on palynological and

micropalacontological eviderice, the presence of B&.

laeve gulicle in the Sedan lignite could imply the

leal fossil has a longer stratigraphic ramge than

previously reported, i¢ Late Focene-Miocene,

The basal lignitic clay is dominated by the

Lauraceae, which is of no stratigraphic significance

as the family was well represented throughout the

Australian Eocent. A comparison af Bocene

Lauraceae culicle types from a number ot localittes,

including Anglesea (Rowett & Christaphel 1990)

and Nerriva (Hill (986) has shown that both of the

geneta identified in this lithowpe, ia Croprocerve

and Andiandra, were well represented in terms of

species and abundance throughout the period, It

should be noted that the lauraceous megalossils of

the Nerriga locality were assigned to the form genus

Laurophyllum by Hill, but affinities to Bratandre

and Cryptocurww were also Lndicated for a numiber

ot these leaf types. It must also be mentioned that

the two genera are the major contributor to whe

modem Australian Lauraceae.

However, the presence of the Lauraceae

parataxon No. AA DOG may prove to be an

important stratigraphic indiestor, bascd an the

author's unpublished information. This parataxon

appeurs to have a restricted distribution, being only

recorded in Middle Eocene depasits and reaching

inaximum abundance in the Lipper Middle Eocene.

[t has not at chis present time been identified in Late

Eocene or younger sediments,

In conclusion, it is evident from the analysis of

the dispersed cuticles of the Sedan coal sequence

thal L) (wo separate dispersed cuticle flaras are

Tevognised, ic. the Lauraceae-dominated flora of

the basal lignine clay and the Proteaceac-dominated

flora of the younger lignites; 2) the change in

lithotype and Mors recognised between the basal

clay and lignites may be climatically induced; 3) the

key cuticle type Banksiewephyltum all. B laeve

recorded at Sedan occurs in younger lignite

sequences in the Latrobe Valley (Victoria) and

Bowmans deposit (South Australia} sugyesting

either that the Sedan lienites are younger than

indicated by both the micropalacontology und

palynology or that the stratigraphic range

(Oligocene-Miocene) of the cuticle rype is longer

than suggested by the presenr records

Acknowledgments

The author iy grateful dor the support provided

by NLE.R, DDC. grant No, 1174. Thanks must go

10 David Christophel for his encouragement and

assistance throughout this project and David

Greenwood for his valuel comments on the

manuscript, Thanks also go to the Direclor General

of the South Australian Department of Mines and

Energy for permission to examine core material,

Neville Alley for assistance in core selection and

palynological stratigraphi¢ information, Brian

McGowran, for providing foraminiferal

stratigraphic information and to Anthony Fox for

his assistance with photogcapliy.

% A, 1. ROWETT

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& Briwooo, A, J, (1985) Tertiaty Araucarian

Macrofossils from Tasmania, Aust. J, Bor, 33, 645-656.

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Tasmama: Araucariaceae. Alcheringa 13, 325-335,

& CHkistorHer, D.C, (1985) Tertiary Leaves of

the tribe Banksiewe (Proteaceae) from southeastern

Australia. Bor. £ Linn. Sac. 97, 205-227.

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shudics of Tertiary brown coals: Preliminary analysis

of lithatypes within the Latrobe, Valley, Victoria.

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University, Melbourne).

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(Fds) “The Cainozoic of the Australian Region”, (Geol,

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WHALE STRANDINGS IN SOUTH AUSTRALIA (1881-1989)

BY CATHERINE M. KEMPER & JOHN K. LING*

Summary

Cetacean strandings were collated from museum records, and published and unpublished sources.

The identification of stranded animals was verified in 85% of events by examination of voucher

specimens or photographs. Trends in the stranding record were documented, but reasons for

strandings were not investigated in detail. From 1881 to 1989, 309 stranding events, involving 24

species, have been recorded in South Australia, while an additional species may have stranded in

the State. Stranding events most commonly inyolyed Tursiops truncatus, Delphinus delphis,

Physeter macrocephalus, Caperea marginata, Mesoplodon layardii, Globicephala melas, Kogia

breviceps, and Balaenoptera acutorostrata. Strandings were less frequent on the western shores of

Gulf St Vincent and Spencer Gulf, possibly because these are protected from strong southwesterly

winds. There was a trend towards more frequent events being recorded in January/February and

September/October. Some species showed seasonal trends in stranding, possibly related to their

migratory patterns or movements inshore/offshore. Eighty-nine percent of stranding events involved

single animals; 18% involved groups of two or three animals and 3% were of four or more.

Twenty-eight percent of stranded animals were juveniles. In some species (e.g. Balaenoptera

acutorostrata, Caperea marginata, Globicephala spp., Kogia spp. and other Balaenopteridae),

juveniles constituted a high proportion (> 30%). At least 15% of stranding events involved live

animals, although more accurate observations and reporting in recent times indicate that live

strandings are probably more frequent. There were live strandings of 16 species, including baleen

and toothed whales. There was a tendency for large whales to strand more often in a moribund or

decaying state than small species. On average, about 20 recorded stranding events have occurred in

South Australia each year since 1985.

KEY WORDS; cetaceans, stranding, mass stranding, trends, South Australia.

Transactions of ihe Rovel Society uf 5 Ausi (99), UASEL), 7-52.

WHALE STRANDINGS IN SOUTH AUSTKALIA (1881-1989)

by CATHEKINE M, Kemper & JGHN K- TING?

Summary

Kraipor, C. M. & Ling, JK, (1991) Whale strandings in South Australia (1881-1989) Trans, R. Soc. §.

Ast UW5(1), 37-52, 31 May, 1991.

Celacean strandings were collated fram museum records, and published and unpublished sources. The

identification of stranded wnimals was verified in BS%y of events by examination of voucher specimens

or photographs. Trends in the stranding record were Jocumented, bul reasons for strandings were nor

investigated in detail. From 1841 1a 1989, 309 stranding events, involving 24 species, have been recorded

in South Australia, while an additonal species may have stranded in the State, Stranding events most

commonly invelyed Tirsiaps trunculus, Delphinus delphis, Physeter macrocephalus, Cuperea marguviata,

Mesoplodon layardil, Giohicephuta melas, Kogia breviceps, and Balaenoptera acutorestnnd, Strandings

were less frequent on the western shores of Gulf St Vincent and Spencer Gulf, possibly because these are

protected from strong southwesterly wings, There was a trend towards more trequent events being recorded

in Jantiary February and September/October, Some species showed seasonal trends in siyanding, possibly

related lo Lieir Migratory patterns or movernents Inshore/offshore. kighry-nine percent of stranding events

involved single animals; [$¥%o inyolyed eroups of two or Lhree animals and 3% were of tour or more. IWwenty-

eight percent of stratided animals were juveniles. In some species (e.g. Balaenopiera ucuiurostrale. Capered

mareinata, Glabicephula spp. Sagia spp. and other Balaenopteridae), juveniles constituted a high praparhor

(>30%). Ai least 15% of stranding-events involved lve animals, although more acourate observahons and

reporting in recent times indicate that live strandings are prabably more frequent. There were live snandiings

of 16-species, including baleen and toothed whales. There was a tendency for large whales to strand more

often in. moribund or decaying slate than small species. On average, about 20 reenrded stranding events

have oocurred if South Austratia cach year since 1985.

Key Wopbs: cetaceans, stranding, wiass stranding, trends, South Ausrralia

(Introduction

Cetacean strandings have captured the artemlion

of humans since Aristotle’s time (see Aristutle 335

RC). but only in the last few decades have these

events been studied in detail, More recently, tretids

hawt been analysed and plausible hypotheses

proposed on the possible causes for stranding of

live animals (reviewed by Geravi & St Aubin 1979},

There are many possible causes of ectacean

sirandings, e.g. discase, injury (both natural and

anthropogenic), birth difficulties, adverse weather,

parasile infections, food supply, old age and toxic”

pollutants (Geraci & St Aubin 1979), A recent

hypothesis suggests that cetaceans may ‘make

navigational mistakes’ if Lhe earth's geomagnenic

field is disturbed (Klinowska 98), Sergeant (1962)

concluded that nearly afl animals invelyed in single

strandings were diseased or purisitised, but thosc

involved in mass strandings were not.

jn Australia, the must detailed accounts of

cetacean strandings exist for Tasmania (reviewed by

Nicol & Croume 1988). Sume published

* South Australian Museuny, North Terrace, Adelaide,

S.A. 5000 ;

Sopp, B. A. (1964) Ceracean sicandings in Sani

Australia 1881-1984, Field Study for Assuciace Diploma

in Wildlife and Park Management, South Australian

College of Advanced Eduencion,

observations are available for Vivtoria (Wakefield

1967: Warneke 1983, 1988) and the Hlawarre district

of New South Wales (Robinson 1984), Species

recorded in Queensland are discussed in Bryden

(1975), Paterson (1986) and Paterson & Van Dyck

(1990). Since 1934 all states, but nut the Northern

Territory, have been reporting strandings to the

Australian National Parks & Wildlife Service.

‘The South Australian Muscum has taken a

special interest in cetaceans since Lhe mid- to late

1800s (Hale 1956). E. R. Waite, H. M. Hale and,

more recently, PF Aitken established a stranding

reporiing and collecting network which provided

us with sufficient records ro analyse trends, Aitken

(1971) poblished a sunny of the 18 species which

he regarded as gocurring in South Australia, based

on strandings or sightings (also summarised by

Semmeanit 1982). A revised account was presemed by

Ling & Aitken (1981). Stopp (1984)' compiled a

detailed account of the locations of many South

Australian strandings up 10 1984, with some other

details for each specimen.

Here we summarise the entire stranding record

for South Australia, Trends in strandings are

analysed in relation to species composition and

abundance, geographic and seasonal distribution,

group sive of stranded animals, age, and active vs

passive stranding events. No attempt has been made

{U investigate or explain the cause of strandings,

Ax C.M, KEMPER & JK. LING

except ih very general lerms. Special attention was

paid to the possible influences of the unique

ticeaneyraphic conditions of South Austrilia on the

cetaceans of southern Australian waters.

Oceanoycaphy of the South Australian Coast

The South Australian coast measures

approximately 4000 kr of which about 1500 km

(38%) border mwo large gulls, St Vincent and

Spencer, Several laree buys occur wlong the seaward

coast, The 200-m bathnmeric contour lies 10-200

kn offshore for much of the coast but 15 Glaser

inshore (SO kim) jn the sourhense of the Sra. Hence

there is a broad vantinental shell which, with the

exception of Bass Strait. unique forthe southern

half of the continent, Deep canyons are found to

the southeast of Kanguroo Island and seuthwest of

Cedunz. Annual mean water temperatures in jen-

gull waters vary from 18° — 18°C (Radok [976).

Cold oceanic currents generally Mow in an easterly

direction across southern Australia ar cates of 14-25

kim per day. exceprin sismmict when a westerly eclely

is found in the ¢Circat Australian Bight region

(Federal Department of Transport 1987). A Fast (35

kin) per day) casterly current occurs in the Bight

during winter, The Leeuwin current, which

onginates in the Indian Qeean off Western

Ausiniia, brings warn water into the Bight in

autumea and winrer (Pearce & Cresswell 1985). A

surface temperature gradient of up te 59°C may he

noted along the southern ooast of Western Australia

at this tine Little has been documented on gulf

and inshore currents in Suuth Australia alchough

studies are in progress (PB. Petrusevies pers. comm.

1990)

In South Australia, the prevailing winds are

generally southerly or southeasterly in summer and

souchwescterly of norlherly in winter (Climatic Anas

ol Australia 1988). There may be some deviation

from this pattern in the region of the gulls, Wind-

induced upwelling events are important in the

suulheast of the State im summer jLewis LY8l;

Schihinger 1987). Other areas of upwellinw are

found off the southwest coast nf Kangaroo sland,

and between Coffin Bay and Anxious Bay Quat

south of Streaky Bay) (P, Petrusevies pers. comm.

1990).

Bye (1976) described the guifs (Spencer Gulf and

Ciull St Vincent, separated by Yorke Peninsula) and

their associated waters {Investivator Strait and

Encounter Nay) as a complex zone of fundamental

impartance to South Australia’s marine

environment, (Investigalor Straic (north of

Kangaroo Island) and Encounter Bay feast af

Kangaroe Island) are not considered part of the

gulfs, because they ure very much influenced by

oceanic patierns), Salinity al We head ol the gulfe

is auch higher than at thar mouths (far Speneer

Gulf as high as 48% in the late summer and abuul

43%, in late winter {Nunes & Lennon |4s6)), Due

to the shallowness of the water in the gulls | <50

m), lemperatures range from about 12°C in winter

to about 24° in summicr (Bye 1976; Nunes & Lennon

1986). Again, there is a north-south grathent. Whe

gulls are somewhac protected from the full effects

of the Southern Ocean by their surrounding land

nhasses.

Another urea of interest Gceauographically. bur

abour witich litleis Known, is the Head of the Great

Austrahan Bight, In winter, temperatures and

salinities are higher than the water to the south

{Pearve & Cresswell 1985).

Muterials and Methods

Information ot cetacean strandings Was cathered

Irom speeimens and photographs licld in the

collections of the South Australian Museu:

published records in Aitken (1971), Hale (L991, 1939,

1959, 1962), Ling & Aitken (19MI), Stopp (1984),

Waite (1919), and newspaper articles; and

unpublished records from various sources, No

specimens from South Australia exist in the

collections of the Western Australian Museuin,

Australian Museum, or Museura of Victoria, ‘The

identifications of all specimens in the South

Australian Museum have recently been verified;

hence stime published observations few. Aitken

1971} muy not be consistent with ours. OF the 309

stranding events reported here, 75% are verified

with voucher specimens in the South Australian

Museum and 10% are veriticd with photographs

alone.

The taxonomy used here is that of Bannister

(1988) with the exceplion of Globicephala melas (=

ieluenu) (Rice J989), the placement ol Cuperea

marginata in the family Neobalaenidae (Rarnes &

MeLeod 1984) and the use of the specific name

miacrocephatus, nor cefeden, for Physeter (see Rice

1989),

A ‘stranding’, in the present study, is any event

involving the beaching or washing up on shore of

live or dead cetaceans, a4 well as observations of

dead vetaceans floating offshore. Skeletal material

from the cous! is included in our definition ol

stranding (bur not dated), except tir the case of

Eubalaena dustralis near old whaling stations. Jt is

assumed that mose museum specimens lacking data

were collected as a resull of stranding events.

although caution was exercised in the cases ot

Phuseter mocracethalus aud Euhalaenw uustealts

WHALE SIRANDINGS IN SOLTH AUSTRALIA 39

due to whilling activities, 1} is possible that some

small cetaceans could have been caught accidentally

if fishing mets, but this information is not recorded

by the Museum, A single Caperea nmtergirate

(Encounter Bay. Septeanber 1887) was caught in a

fishing nev.

The relative age of individuals was estimated only

in those cases where the total length of (he animal

was knows or, in a lew cuses, was eslimated from

pholographs Neonates were defined as individuals

approximately the lenech of newborn animals,

based on information From other studies (Perrin ef

al. 1984; Ross 1984), Juveniles were defined as bang

less than, or cyual to, 4 certain proportion of the

species length al physical maturicy, These are: 30%

(E, australis. P wacrocephalus, Glubicephale spp,

Grampus griseus), 95% (CO mersinent), 60%

(Balaenopteridae, Tursiaps truncetus, Delphinus

delmis), 65% (Xp2ia spp.) and 70% (Aiphiidae).

These proportions were based On information fram

several sources (Dryden 1972; Ross 1984; Mcad &

Potter 194); Koss pers comm. 1990).

Where the actital date of stranding was unknown,

the month was estimated on the basis of

photographs of the animal or a description of its

slate of decomposition,

The term ‘pulfs' refers here only to Gull St

Vineettt and Spencer Gull. The (erm ‘gulls region®

includes Investigator Strait and Backstairs Passage

und the north coast ef Kangaroo fsland,

Results

Species

Twenty-four species al cetaceans were verified as

having stranded along the Sowih Australian couse

between 188) and 1989 {Table 1). Thene is same

uncertainty abaul the additional spevics,

(Balaenaplera berealis) which, although in the

Sourh Ausuatian Museum collections, may not have

stranded in the State, Reeent re-examination of

baleen plates (SAM M4829), previuusly identified

us A wedeoi, resulted in the identification being

Jetermines! as Bo berewlis {based on characters

described by Mead (1977) and Horwood (1987),

However, the plates have no avcompanying data and

therelore could have been collecred elsewhere, or

they may not have been from a stranded animal.

No species are recurdetl as stranded in the State

based solely on photographs or other reports

Jacking voucher material (Table 1).

Jn terms of individuals (bur excepting the mass

strandings of Pseudenw crassidéns), the most

commonly stranded species were the dolphins, 77

(utcatas ind 0. delphi, These are followed by six

species; JMipserer muacrocephalus, Caperea

marina, Mesopladon fayardiy, Sogia breviceps,

Globicephala melas and Baluenoprera

dcuterasteata, whose stilus yaries depending on

whether spevinven records lacking vouchers or

lgeality inlorniation ar: included. AU have greater

than 1) recorded stranding events. Belienopicra

physalus, Berardius arnuyxii and Kogia sinus have

stranded only once. There is some uncertainily about

whether fasinaceties shepherd? has. stranded once

or twice (G, Ress pers, comm, 1991),

Geographic distribution of stranding events

Stranding events having kLeovraphic Inealily

information (238) were not evenly distributed along

the South Australian coast (Fig. 1). More strands

oceurred im (or were reported trem) the Caltin

Ray/ Port Lincaln and Coorong regiuns, followed

by the castern side of Gulf St Vincent, the north

cuast of Kangaroo Islind and the Streaky

Bay/Ceduna regions. Very few strandings were

reported from the westera sides of Gull St Varteent

and Spencer Gull’, despite moderate levels of hymn

activity along these coasts, There were lew

stfandings reported in the tur west of the Stale,

much of which is lined with inaccessible cliffs and

a covky shore. Forty-six pereent of srrandings

occurred within the gulls region whieh includes

about 41% of the South Australian coast

Mass strandings (1c, more than three individuals)

have occurred at six locations (Piz. 1), Five were

in the gulls region, Three locations (north of

Adelakle and northeastern Kangaroa Island) were

where other live strandings have uccurred (Fig. 2).

Active (= five} strandings have been reported less

frequently in remote areas than around centres of

population, @g. in the gulls region (Fig, 2),

Nevertheless, it appears that some areas (ee. Port

Lincoln/Coffin Bay, Nepean Bay, eastern Cult St

Vincent, Murray River mouth) are more pron¢ te

active strandings than are others,

There are trends in the geographic disinbution

of stranding events of species or species groups (Figs

3-8), Both 7 iruncatus and 0, delphis have

stranded alone much of the coast but

conerntrations of events exists around Adelaide

Port Lincoln, Nepean Bay and the Murray River

mouth (Fig, 3 The wo records of FT truncatus

from Lake Alewandrina (north of the Coopone) were

made before che barrages were built in the T9208.

Few dolphin strandings have been recorded trom

the northern ends of the pulfs. OF the 36 sirandings

within the gulls region, Inete were of 2. rridteatis

than B delphis (P <O10, y* 2.78, 1 da,

Few strandings of Glahicephata macrorhynchus

and CG. delay were reported in the gulfs region (Vig,

4), The only two records within che gulfs proper

being G. mracrorhveckus one af these a ess

stnnoding.

40 C. M. KEMPER & J. K. LING

TABLE |, Stranded cetaceans along the South Australian coast recorded from various sources, South Australian

Museum (SAM) specimens with no locality data are in parentheses and ure tneluded in the total numbers to their

left. Events from other sources includes photographs, published and unpublished records. Total individuals includes

specimens plus individuals fram other sources, Unidentified records not listed.

SAM

Family, genus & species specimens

BALAENIDAE

ubalaena australis 735)

NEOBALAENIDAE

Cuperea marginata 27(8)

KALAENOPTERIDAE

Balaenoptera acutorostrata 11(2)

B. burealis* (W)

B. edeni 4(0)

B, musculus 5(1)

B. physalus (0)

Megaptera novaeangliae 4(0)

DELPHINIDAE

Delphinus deiphis 39(12)

Globicephala sp. 0

G. macrorhynchus 10(1)

G. melas 1)

Grampus griseus 2(0)

Orcinus orea 3(1)

Pseudorca crassidens 2(0)

Tursiops truncatus 71(21)

PHYSETERIDAE

Rogia breviceps 20(1)

XK. simus 1(0)

Physeter macrocephulus 26(4)

ZIPIIIDAE

Berardius arnuxii 1(0)

Hyperoodon planifrons 7{0)

Mesaplodon sp. 0

M. bowdoini 2(0)

M, grayé &(0)

M. layardii 22(0)

Tasmacetus shepherdt 1(0)

Ziphius cavirostris 2(0)

TOVALS 290(58)

Events from

other Total Total

sources individuals events

0 7 7

2 29 29

1 12 12

0 1 1?

1? 6 6

a # 5

0 I i]

1 5 3

16 60 54

2 2 2

0 0 6

4 15 14

2 4 4

0 3 3

0 cyn0 2

12 126 45

! 22. 13

0 2 1

g 4 34

0 i 1

0 7 T

5 6 5

0 2 2

0 8 &

I 24 19

0 1 |

O 2 2

56 WS 309

* Doubtful S.A. record

Strandings of C. marginata have been reported

from the Victorian border to Streaky Bay (Fig. 5)

although the latter location is unsubstantiated.

Sixty-five pervent of strandings have occurred in the

Port Lincoln and Nepean Bay areas. No strandings

have been recorded Irom well within the gulfs.

Balaenopterids show some interesting trends (Fig.

6). B. edeni, with the exception of a dubiously

identified specimen from Cape Banks (approx.

38°S), has always stranded well within the gulfs, B

acutorostrata, on the other hand, has almost always

stranded outside the gulfs and usually in specific

areas (e.g. Coffin Bay, Port Lincoln, Nepean Bay

and the Coorong). No trends were apparent for

other species. The single stranding of a juvenile B

physalus occurred at the northern end of Gulf St

Vincent,

The numerous strandings of Physeter

macrocephalus have occurred along almost (he

entire South Australian coast (Fig.’7). Few have been

recorded within the gulfs, The record from Adelaide

was a specimen of bones of unknown. date, buried

in sand in shallow water. No record could be found

of a large whale stranding near Adelaide in the past

50 years, 50 we assume that these bones represent

an older stranding event. Kogia breviceps has also

WHALE STRANDINGS IN SOUTH AUSTRALIA 4l

142" 1aat doa Tobe a6" yar

(348° rage y4n* yah

a ; T T4340

| i

1 4aoe

Great Australian Biaht

uae Biss

is]

200m as es

ae 4

ee <=,

aa" 7 aan

. in)

fel!

% : : 4

gael : : s a - ——— n+ y las"

hs *

rr:

=| |

>»! |

360 —hi jase

“TJhokm

Too km

a7 ingston an

; i

a4 2 i* os f f Stl ft

q?Re Ti0? Vit 32" aaa" vgae wise 45° qaire taae ya" 1409 vane

Fig. 1. Geographic distribution and number of reported stranding events in South Australia, 1881-1989, Shaded areas

indicate coastline oyer which events took place. Solid triangles denote mass stranding Jocalities.

: iar 130° asi" 152F 133° iar (a5 136° 137" 138? vag? raat vay

— ae TO - i Tyaee

| !

| Eucla

32" 5 eF

} Great Australian Bight

sae ee lel gs

[200m ec

Are =]

aw =z

ge” x To Hse”

\ , we

Se Woe

a \

‘Ry “

% 4 |

ane ml! | a5°

Xe 0

us 7 ——= | = 1388

Thakm <3

Pin ingston at

ae) |

Sareea r Pe e— -- ! te — — is 1”

tage (aoe aie laa 133" taae 135" yaar Vat jane raat 7408 vate

Fig. 2, Geographic distribution of reported active (OQ) and passive (@) stranding events in South Australia, 1881-1989.

42 Cc. M, KEMPER & J. k. LING

Igae yaar vaie ise" 139° igs AGBS 136° 137° var vase i4o"

i447

4a = E a ‘a 1 f= > me or ee 8 eh eh i

| |

lEuel | | i

ucla pa ~. !

cla |

ype deg. gece A b— + aye

i !

| | j |

Phrt August

Great Auptralvan Bight

aan * 4s

200m _ c!!

_* 4

saa ad la

352 “mt 35"

7gac00, - +

vlads a \ a >

anal

Nepoah Bay

A .

ae ma a jaan

ja0km | % | | ve

os lea

ane _ » Ja8e

* = + = = + ——— — — . ' (eos pe

van" 720! Tote be kag 133° 734° ads? 136° wre 138° 139° T40" 14a

Fig. 3. Geographic distribution of reported dolphin strandings in South Australia. 1881-1989. Each symbol represents

a Delphinus delphis (@) or Tursiops truncatus (W@) stranding event.

124" 130° ist* jae igs? gar 135° 36? are 13BF dae yaar i4are

my ~ oT - = T H “ya

| | }

{Eucla ee SUSE } | |

—— |

a2" ~ 4 = 928

x duna i

~~ a }

Greal Australian Bight |

e. ao ella

\ 7 o

i Ar OG

Po |

— 200m \ | a

SIR i

~ - a 4

e Nv |

var eae

. le

iG Va

“i

: NS 4

35th : mij 35

| o ,

| een =

* Pr.

ae <4 Jane

rookm '

ar 2 Kingston 14 ype

{

aa") f rf ‘ =| {pee

par rane yar yaa" 133° isa? 135° jae Tar" 38° yatge 1400” 1419

Fig, 4, Geographic distribution of reported pilot whale strandings in South Australia, 1881-1989. Each symbol represents

a Globicephala macrorhynchus (@), G. melas (d&&) or Globicephala sp, (Wi) stranding event.

WHALE STRANDINGS IN SOUTH AUSTRALIA 43

129* wage way 132° 133° 134" 135° 136° 1378 ida? waa" Vane vate

ait + “ts ] T T - ] T i oy | f yds

| j | H

| i | i | |

lEucta | te | | | | |

ee ™~. | 1

» Le | ;

den ' - opt - : — 2 en » age

an a Cina a

~ ir

S) 1 Port Augusta

Great Auetralian Biaht 2 H 1 |

bd |

ne Lou sua Bay J 4 Bo 339

on™ 2

200m on * , es \ e

= = te a ‘

aat es 4 & ao jae

os a «TA oy;

Ac { oF yl i i ie

are

NS eet st Af <

<a LS \ {

NM, { w

™, Le se!) Fs 4

rH ™ Zz sor

eae al > |

ante © E

NN Agaroo web. . rx 4

: uot d ‘ *

” — st slang \ hi |aee

Tookm A Ge |

— Nepean Bay ‘|

a } ’

a7} ae @ ingston bbe

— \

| A \. |

| Ny

26° *s \ |. wae

124° ~~ 7a0e vir 732" jaa" yaa jase Tsar ~OC ae vas" ——V39° me) Ta

Fig, 5. Geographic distribution of reported strandings of Caperea marginata in South Australia, 1881-1989. Numbers

in circles represent numbers of events. Streaky Bay stranding is unsubstantiated.

y2gr 130° i3i° 132° 433° ygee 195" 14a8o 137? jaar 73s* i407 jair

are Oks eet ae - - . ri T ate

lEucla i

_ SB : |

ae | !

gz + mimes tem —- — age

oe eg) Ceduna

| py: . Port Auguste

Groal Australian Bight im i |

| ‘ '

| Streaky Bay _ _-t\ et se

10% % sone bad

200m_ — ee a

— ie | i

| =

44h . — | ea 4a

' T

N oot wo } mt co a

NL. e\ ot } e

a | |)

S ° ‘4 PEUAIDES 4

‘ \. — me ele

ae Oo sts Pr

ee UR 2 {

Ngarog 5s, al

a8) x See 7 \ Se eel

d Tookm ? . \ ako

| NS Nepean Bay |

esl

ay _— a A@Kingstan aan

Po all | roa

Siete 1

mL i

NY \

se .

. oN

3a" a ; ; 4 he wa aae

i29t ye tare tart 188° yaar 15° ya6F .37° 138° 133° tac7 141°

Fig. 6. Geographic distribution of reported strandings of rorquals (Balaenopteridae) in South Australia, 1881-1989.

Each symbol represents a Balaenoptera edeni (@), B musculus (&), B. acutorostrata (Wi), B. physalus (4 ) or

Megaptera novaeangliae (QO) stranding event.

44 ©. M. KEMPER & J, K. LING

year y30* {are 4324 i3a¢ tsa" 138° Va5° say 138° yaat

vage 1416

ane Tryarr

ae * aoe

Great Australian Bighl

—= } ert

ant tp

. ciiydasnvbnines

aa°

a6"

aR’ Ret

100

a7 ww

aA" an4

= 4 = .- — = ——

rer “Vagt ate yaze 143° yaar 235% 146° yarn 136" 138° 140" Tate

Fig. 7. Geographic distribution of reported strandings of Physeteridae in South Australia, 1881-1989. Each symbol

represents a Kogia simus (&), K. breviceps (@) or Physeter macrocephalus (™@) stranding event.

ioge 130" vane raze rage 1gae qast raee 4378 3B" 139° i4ae 1442

ait —— | = : 7 | Ta"

"Eucla | ‘

ge t + 7-133"

Port Augusta

Greal Australian Bight

ay oe Be

200m c

ee = 3 4

ny ja

ss * a”

“yg , ~ i twee

\ metal

S e

SK, e

—

‘, =

3h Me ald ed

ae >

—l

» |

an" ~4} 446"

yookm

az h fare

ape f , {aah

_ $, Seger " aes. L DS Mt

year aet tat? 132" 1a3° wae \a5" 136° wT 438° iag° sao" tare

Fig. 8. Geographic distribution of reported beaked whale (Ziphiidae) strandings in South Australia 1881-1989, Each

symbol represents a Berardius arnuxii (+), Ziphius cavirostris (©), Hyperoodon planifrons (@), Mesoplodon

bowdoint (QD), M. grayi (@), M. layardii (&), Mesoplodon sp. (+) or Tastnacetus shepherdi (*) stranding event.

WHALE STRANDINGS IN SOUTH AUSTRALIA os

stranded over a wide stretch of the coast, including

the gulfs, but none has stranded on Kangaroo

Island. The single stranding event of K, sintus is

from Adelaide.

Beaked whale strandings have been recorded

along the entire coastline (Fig. 8}. Only 25% of the

beaked whale stranding events have heen within the

gulls region. There are many records from the

western side of Eyre Peninsula and along the

southeastern coast of the State.

Four species have not been illustrated on the

foregoing distribution maps. To our knowledge,

EBubalaena australis stranded three times during

1881-1990; an adult at Port MacDonnell in the far

southeast and two neonates al. the Head of the

Great Australian Bight. Grempus griseus has

stranded at four localities: Point Turton (Yorke

Peninsula), Coorong, Willson River (Kangaroo

Island) and Denial Bay (near Ceduna). The

locations of possible Q. orca strandings have not

been reported (see Ling 1991). Mass strandings of

P crassidens occurred twice and are discussed under

‘Group size of stranded animals’.

Seasonal trends

Stranding events (all species combined) were

recorded most frequently in January — April and

September — October (Fig. 9). However, this trend

does not always extend to individual species, or

species groups (Figs 10 and 11). Beaked whale and

T, truncatus strandings were recorded most often

in summer (Fig. 10). An imcrease in torqual

strandings. was evident in September (Fig, 11), with

six out of the eight being juvenile animals. We have

h bh

aoas

Number Stranding Events

5 oO N DO

J FMAM J Jd oA

Month

Fig, §. Month of occurrence of reported stranding events

in South Australia, 1881-1989,

Ziphitdae

Tursfops and Oelphinus

_ st ot

o tw

Number of Stranding Events

yn 2m OO

oS F&F DP OD

J oF M AM J J A S OWN DO

Month

Fig. 10. Month of occurrence of reported stranding everits

of Ziphiidae, (Meseplodon lavardii solid bars), and

Delphinidae (7iirsiops truncatus solid, Delphinus

delphis open, G. macrorhynchus solid, G. melas, open,

Globicephala sp, dotred) in South Australia, 1881-1989,

Numbers above bars refer ta numbers of neonatal

individuals stranded.

insufficient data on other species, or species groups,

to draw conclusions, although there did appear to

be slight increases during September - October for

Glehicephala spp., C. rharginata and T. truncatus,

Same species appeared to have stranded in

specific periods (Figs 10 and 11), Kogia breviceps

stranded from April to October, mostly between

July and October, Physeter macrocephalus stranded

between July and December, M. lavardii (January

~ July) and G. macrorhynchus (May — September)

have also stranded in somewhat restricted periods,

but sample sizes may not be sufficient to be sure

of such trends. The stranding of an adult &.

australis in February is noteworthy, since it is outside

the period of sightings of live animals in coastat

South Australia and Victoria. The nature of its

injuries ~ almost decapitated (possibly by a large

Number of Stranding Events

CM, REMPER & J. RK. LING

Balaenopteridae ~

F M A M J oJ

Kogia spp-

J oF M A M Jo od

Physeter macrocephalus

J F M A M Jd Jd A S GO N D

Month

Pig. 11, Month of pecurrence of reported stranding events

of rorqnals (Balacnoptevidae), Caperea niarginata,

Aogia spp, and Physeter macrovephalus in Sourh

Australia, 1881-1989. Numbers ubove hars refer to

numbers of neonatal individuals stranded,

ship's propeller) — suggests that it may have dicd

some distance offshore.

Strandings of T. rruncativs were most frequent in

January — March, both within and outside the gulfs

{Table 2). Events were recorded turoughout the year

in both regions, with a possible reduction in July:

September outside the gulls. The patrern for D.

delphis was similar but there were no strandings

recorded in April — June within the gulfs. This could

suggest an absence of live D. delphis from the gulls

during the autumn quarter.

Group size of stranded unimals

There: were 184 stranding events involving single

animals, Fifteen (8%) events had two or three

individuals and six (3%) had five or more (Table

3). When two stranded, most of the events were

cow/calf associations, especially Mesoplodon spp.

and K. dbreviceps. Excluded from Table 3 is a

possible association between an adult and subadult

male Ayperoodon planifrons which stranded 60 km

and one day apart.

Mass strandings (i.e. more than three individuals)

are rare in South Australia. Those that have

occurred ave described below:

St Kilda (34°45°S, 128932 F) September 1903

Ifale (931) described bricfly the suatding of five

G. wieleena (= melas) now identified as Ci.

éhacrorhwnchus, mi the mud of the St Kilda maneroves,

A female stranded first (alive) followed by four males

when the tide went our.

Port Prime (34°31 'S, 138°18°E) approximately 3

October 1944

Hale (1945, 1959) described the mass stranding of

about 250 P crassidens in Gulf St Vincent, north of

Adelaide. Approximately 200 stranded (some alive),

along the coast of tidal flats in one group, with abour

St) others stranding 2.5 kim north of the main group

and isolated individuals along a 30-km stretch of coast

berween Port Parham and Pert Gawler Very little

biological data were collected or published on the

stranded animals whitch were misidennfied as G.

mieluena (= melas) ai the time at stranding.

Merdayerrah/Eucta (31°41 'S. 129900" E) Aueust 1963

Tn August 1963, abour 49 /? erassidens stranded over

abour cizgh( miles (= 13 km) of sandy beach berween

Merdayerrah Sandpatch, S.A. and Eucla, W.A, About

12 were on the sandpateh and 47 at Eucla. All were

ive or freshly dead when found, Mr H. Gurney (pers.

comm. 1989) believes that they stranded at high tide

because there were furrows in the sand ws i! they had

been driven up the beach. He recorded males, females

and calves but did not count exch category. The males

were about 20-21 feet (6.4 m) in length and rhe calves

(of varying length), a little fess. According to Mt

Gurney a ston did not precede the strandiny.

TABLE 2. Strending events of Tirsiops truncatus (a) und Delphinus delphis fh) which necurred inside und vutside

Gulf St Vincent und Spencer Gulf during four periods of the veur,

location Jan — Mar Anr — June July — Sept Oct -— Dec

(a) Inside gulf 9 3 2

Ourside wally 9 4 1 4

(b) Inside gulls 3 0 2 4

Outside pulfs 7 3 Z 4

WHALE STRANDINGS IN SOUTH AUSTRALIA aE

TABLE 3. South Australian cetacean stranding events involving more than one animal.

Number of jndividuals

Species 3

5 14-17

Delphinus delphis

Tursiops truncatus

Mesoplodon sp.

Ad leyaeddil

Kovia lireviceps

Ky sintus

Globivephala melas

G. macrorhynchus

Pseudorca crassidens

hee Re

Memory Cove (34°58'S, 136900'B) 2 May 1977

A school of S§ 7. truncefus beached on the

afternoon of 2 May at Memory Cove, 27 km southeast

of Port Lincoln, Thirteen died and I6cal authorities

returned the remaining 42 to sea. Twelve of the dead

were collected, measured and lodged in the South

Australian Museum. Total lengths were 2.29 m — 3.10

m (6 90,6 oo). None had food tn the stomach

and one-small male showed evidence of a recent shark

attack on his side.

An unsubstantiated report of about 1X) dolphins

beaching themselves aj Louth Bay, 45 km north of

Memory Cove on the night of 3 May 1977 was found

in P. Aitken’s field notes (held in the SAM). Apparently

three of these animals diced,

American River, Kangaroo Island (35°47’S, 137°46" B)

before 22 December 1982

Tive male D, delphis, were found ona rocky beach

al. American River, Photographs of these animals

indicate that they had been dead for a few days when

found, American River is at the entrance to Pelican

Lagoon, a shallow body of water with a rather

restricted entrance: At least five other ectaccan

strandings have occurred there.

De Mole River, Kangaroo Island (35°42'40"S,

136°46‘40"E) 27 April 1985

Between 14.and 17 7. rruneatus stranded ot Snug,

Cove at the mouth of the De Mole River, on the*

northwest coastof Kangaroo Island. The sandy beach

(200 m wide) and river mouth are very smalland form

a shallow depression in the liff-lined coast. The

stranding was recorded by a local resident who

observed a group of about 80-100 Tl fruncalus

swimming offshore at about the time that the stranding

oveurred. One animal swam in the area for three days

afier the event. Mout stranded animals were aboul §-9

feet (2.4-2.7 m) tong (D. Scikman pers, comm. 1987).

Age of stranded animals

Overall, 28% of the stranded cetaceans were

juvenile, but there was: great variation between

certain species or species groups. Those having high

proportions (>30%) of juveniles were &

acutarosiraia, ©. marginaia; Globicenkala spp,

other Balaenapteridae and Kogia spp: (Table 4).

Juvenile & wextorostrate ranged from 3.23-3.96 m

(approximately the size at weaning, Jonsgard 1951).

Only. one physically mature & acutorastrata (9.13

m) and one neonate (2.3 m in October) were

recorded, Length at birth has been estimated at

about 2m in © marginuta, based on the discovery

of what was believed to have been a full-term foetus

(McManus e? a/. 1984). Two neonates (2.01 m, which

accords with this finding, and 2.10 m) have been

recorded in South Australia, both in October (Fig.

11). About 40% of the measured C. marginaia were

roughly half maximum size (6.4 m).

Ziphiids had a low proportion of juveniles (14%),

including a neonatal M. grayi (2.10 m) that stranded

in December (Fig. 10), A stranded neonatal FE.

australis (5.32 m) was found in October. Its siate

of decompasition suggests it had been dead at least

one month and could have been dead for up to four

months based on the known calving season for this

species in the area (Ling unpublished data), Only

9% af steanded T truncatus were juveniles (‘Table

4).

TAale 4. Relative age of strunded cetaceans in South

Australia, Species with insufficient numbers af individuals

not inclided, See text for definitian of juvenile length,

N = number of indiwiduals for which lenzth recorded.

Species N % Juvenile

Balaenopiera

acutorostrata 1 Bat]

Coperea targinala 6 56

Gloabicephala spp. 7 55

Other Balaenopteridae 2 33

Kogia spp. 14 tt

Delphinusdelphis 17 13

Ziphiidae 35 14

Tursiops truncaius 32 9

ts] © MM. KEMPER.& JK. LING

Achve and passive strandings

At teast 15% of the recorded stranding events

involved live animals (active strandings), In many

older stranding records it was not noted whether

atinials were dead or alive at the rime of stranding,

50 this figure must be\considered as minimal. Since

70% of the reported aetive strandings were after

1970, it is quite likely that more thorough

investigations of fatire strandings (as have been the

case Im recent years) will resul( in a higher

proportion being recorded as active. Sixteen species,

representing all four families occurring in South

Australia, are known to have stranded alive (Table

5). Only obviowsly moribund or decomposed

animals Were classed 4s passive stranders and

therefore this propartion (6%) of the total number

Of events 1s able io be an underestimate (Table 5),

Decompased carcasses found in remote areas were

not added to the passive list unless observers

mentioned the carcass having washed up at the time.

There was a disproportionate number of large

species (tg, the large mysticeres and -

rmacrocephalus) in (his category. (See also Fig. 2 tor

geographic fovation of active and passive stranding

events.)

TALL S$. Celacean stranding events involving live

factive) and obviously dead (passive) cetaceans in Santh

Australia.

Number of events

Species Acie Passive

Eubalaena australis

Caperea marginata

Balaenaplera acutorostrata

B. edeni

B. musculus

Meeaptera navaeangliag

Delphirius delphis

Clobicephala macrarhynchay

G melas

Grampus. Qriseus

Pseudorce crassideris

Tirsiops truncatus

Kogia breviveps

R. sims

Physerer macracephalus

Berardius arnuxit

Ayperaodon planifrons

Mesoplodon sp

MU wrayt

M, lavardii

Unidentitied large whale

Toritle

ee ae ok Pe

Se cel ieee ee elie ne a are ed el

-—o0c0F-cyaSsS3SS

—

Discussivn

Twenty-five cetacean Spevies have stranded in

South Australia, This includes Afesupladon hectori

recorded in February 1990 from the Courong but

not the doubtful B bareues. Onc other species,

Mesoplodaa mirus, has been reportedly sighted

(live) in South Australian waters, bur without a

specitnen or photograph to confirm (he identity of

such a difficult prowp, the beaked whales, we cannor

include tis in the State’s ceracean fauna. Front }9

(o 29 species have stranded or are known to occur

in other scates (Bryden 1978; Paterson 1986; Nicol

1987; Warneke 1988; L. Gibson pers, comm, 1990:

J. Bannister pers. conim. 1990), Western Australia

has the highest number of species, ro doubt becanse

its long coastline includes tropical and temperate

waters. South Australia has the next highest, bul

lacks The tropical dolphins found in Queensland,

Western Australian and oceasionally New South

Wales waters, Australia’s cetacean fauta bears i

resemblance to that of the southeast coast of

southern Africa (Ross 1984). In ierms ot species

composition and in some respeels relative

abundances in the stranding record, the situation

in South Austraba is most similar io Victoria and

Tasmania. The major differences are many more C:

marginaia and beaked whale (especially MZ, Jeyvardii)

events in South Australia than in Victoria; and fewer

Cr. melus, P crassidens and 2, cavirosrris; but niore

HA, planifrons than in Tasmania, The seventh mosr

frequent strander in South Australia, &. breviceps,

has not been recorded in Tasmania (Nigol 1987).

The species which occur in South Australia are

cither cosmopolitan, lropical/subtropicul,

temperate/subAniarctic or mixed-water in their

distribution patterns (Ross 1984), Only & edent und

G_ mecrarhyachus ave considered to be

Iropical/subtrapival in nature, The cool waters

(1S-18°C annual mean) in South Australia would

not be expected to support a tropical fauna,

However, Gulf St Vincent and Spencer Gull are

appreciably warmer (han the waters alt the seaward

coast in summer. Also, the iropical Leeuwin current,

which originates in autumiin the Indian Ocean and

dissipates in winter in South Austrailia, may mix

with warm Bight waters and move vastward

(Rochtord 1986),

All four reliably recorded stranding events of

eden/ occurred well within the gulfs (Fig. 6) during

April (2), Sepretiber (1) and November (1).

Balaenoptera eden occurs off Western Australie

(Chittleborough 1959), but since no sightings of live

B, edeni have been cecarded in South Austratia

(1. K. Ling unpublished data), WW is not known

whether this species is resident in the wulls or simply

an occasional visitor, Globicephala macrarhynchus

WHALL STRANDINGS IN SOUTH AUSTRALIA 49

has Stranded during the winter months inside and

ourside the gulls. We suggest that ihe presence wf

these species and the frequent strandings and

sightings of turtles (A, Edwants pers. -comum, 1989)

could be related to the Leeuwin current and the

warm oulf waters. Zeidicr (1989) has suggested that

the Lecuwin current may have been respansible for

tropical octopus oecasionally being recorded in

Sonth Australia.

Sergeant (1979) concluded char abundance in the

stranding veeord in miost cases reflects the

abundance of the free-living population in a

particular region. Mead (1979) suggested thal X,

drevicens thiwlil be an exception, Other camparable

exumples are (4 orca, which occurs in easterty and

southern Australia bur rarely strands (Bryden 1978;

Warnecke 1988; Ling 1991; present study), and &

australis, Which is sighted tréquenitly in coastal

Waters bul also parely strands, Only five verified

strandings of E ausinalis have been reported in

South Australia, Vietorit and Tasmania (Warneke

198K; presen siudy),

i the stranding record reflects abundance, species

that stranded on only one or bwa accasions (Table

Tt) mity fe designared as rare or occasional visitors

To Souil Auscralian waters. Hence R, physalis, P

erusvidens, &, simus, BR arnuxi, Mf, Hector, M.

bowdeini, T shepherd! and Z capirostris are

probably rare here, al least in shallow (<200 m)

walers between (he conlinenmtal shelf and the enase.

Deep-warer and pelagic species such as P crassidens

and 4. cavirosteis (Leatherwood & Reeves 1983:

Ross |964) are frequent stranders in Tasmania

(Nicol 1987) and New Zealand (Baker 1983), where

deep walers oocur closer lo shore than in South

Australia. The same may apply to To shepherd: a

gare common strander in New Zealand (A, Baker

pers. comm. 1990) than in Australia. Beaked whales

fend to live in deep waters ¢>200 im} beyond the

contiyedtal shelf where they feed on deep-warer

fishes and sywid (Ress (989), They may also teed

in wreits having underwarer canyons (Gi. Ross pers,

comm, 190) such ss the Murray Canyons,

southeast at’ Kanparou Psland, and the canyons

southwest af Ceduna (Griffin & MeCuskill (986).

Whether cetacean strandings happen (er are

recorded) depends wpon many factors including

animal Movements (seasonal or diurnal), human

activiry and awareness, ibe physical ¢nvironment

and the climate. Lit Jasmania, Nicol (1986) found

Wiat celavean strandings were recorded most

frequently in regions With high human activity and

a entiples coastline. Geraci & St Aubin (1979)

concluded thar most strandings oceur om gently

sloping beaches on maruval traps" in (he shoreline,

in Sowh Australia, reported stranding events do

Hol accur evenly won the couse (Fig. 1p, Phey were

frequent in regions wilh large hays fev. Coffin

Bay/Pori Lincoln. Nepean Bay, Streaky Bay) ancl

along westward and southwestwanl facing voasts

(cg. Coorong, and eastern gall shores), Observer

effurt may partly explain these concentrations of

Teports, because several areas of frequently reported

strandings are nevr cities or are popular with

‘holidaymakers, Reduced observer effort, however,

does not accounr for the smal! number of reports

from the western side of Gull! Si Vinwent and

prebably noc Spencer Gulfi We believe that

Hrevailing sourhwesterly winds in winter and spring

may contribute ta the frequency of strancings along

eastern shores (see Tig, |), Other studies have

suggusted that adverse weather is assacated wilh

suandings (Duguy 1978), bul there are few

substantialing dala (Geraci & Se Aubin 1979)

Another unporiant reason for the trends may be

Thal strandings are nore frequent where cetacean

abundance is high (Sergeant 1979). Presurnably

abundance is high because productivily of the

marine environment is high in some areas, such as

where upwelling events oveut or deep sea canyons

are part of the bottem topography. Upwellings are

common summer features in the soucheast of South

Australia (Lewis 1981; Rochford 1984) and along

the coast narth ef Coffin Ray (P, Petrusiviez pers.

vumm, 1990). Canyons occur fram Kangaroo Island

to the Victorian border and in the Great Australian

Bight, Some of the ureus of high stranding

frequency (Fig. 1) can be explained by Wiese ocean.

Ographic !ealures.

In South Australia, cetacean stranding events

were recorded rhroughoul the year (1g, 9) with ju

peaks in Frequency; summer and early auriinin

(January —Apol), and carly spring (September aid

October). Observer elfort would be expected to

increase during the summer months, when people

spend more time on the beaches and are likely to

find stranded animals. Nical (1986) found that ihe

highest! tumber of strandings in Tasmania was

recorded in January and that the winter months of

July and Augusi had fewer recorded events.

Warnecke (1988) also noted that in southeastern

Australia more strandings were recorded tn suniner

than ua winter. One would expect that in South

Australia the stranding frequency of the known

migratory species, generally the baleen whales,

would increase in winter, bul if was spring when

many stranded (Fig. 11), [eis norewarthy thar mauay

of the rorquals that stranded jn spring were &

devmnresttvate, most ol which were the size of

animals at weaning. Rorquals were absem froit the

Tasmanian swanding record duviig the summer

Manths (Nicol 1984),

The carly spring peak m te uverall record (Fig,

9) cane! be explained by observer effoic alone:

40 CoM. REMBPER & b KR. LING

Wind may he an important factor. Wind velocity and

frequency, austrated In the form of wind roses in

the Climatic Atlas of Australia (1988), show an

increase Inthe October quarter for much of coastal

Suuth Ausicalia,and coming from asourhwesterly

direction onto the coast.

Masa strandings are cvents restricted to the

Odontocetres (Sergeant 1982) ‘and are maser

frequently recorded in) Glohicephala spp, &

crassidens, Lagenorhynchus acutus and P

macrocephalus (Geracl & St Aubin 1979). ) hese are

highly social and oceanic species. Mass strancings

generally occur where a species is abundant

(Sergeant $982). U1 is therelore not surprising to find

that in Souch Austnilia, where shallow waters extend

fur wIT the coast and oceanic species would nur be

expected to occur in large numbers, there have been

only six recorded muss stranding events, A similar

picture is (rue for Victuria (seven events; Warneke

1988) and Queensland {Iwo events; Paterson 19%6),

both with wide continentalshelfs, The low munber

(six; L. Gibson pers. comm. 1990) of mass

stgandings in New South Wales where waters are

deep, may be a reflection of the relatively

uncomplicated nature ol the coast. In South

Australia the species which have niass stranded are

P cressidens, CG. mecrorhyacius, tf! wuncatus and

Le delphis.

Owerall, the percentage of juveniles which

stranded was rather high (28%), a trend also noted

by Duguy (1978) in France Sergeant (1982)

eoncluded that odontocetes Have a lower juvenile

mortality than mysticeles, probably because of Ue

wrealér parental care generally associated with

odontocete social structure, The present study

confirms (his, og. high juvenile pruportiuns of

nlorginara and &B atucoresirata, but some

odontocetes also had about a Unird juveniles (Table

4). Sergeant (1982) reported rhat newly weaned and

old B acytorostrata strand selectively. In South

Australia young R deularesirata, 3.23-3,96 m in

length, have stranded between (he months of August

aud Ocrober. This is slightly less than che estimated

length ar weaning (45 mJ reported by Lockyer

(1984). Births in & avutureasérare in the southern

hemisphere are reported to occur in Mav and June

Clackyer 1984}, bur in our records one neonagtal

animal of 2.3m stranded in South Australia in late

September ur early October, The body lenzths al

stranded 2 renga (2.01-6.20 m) are more evenly

disinbured than & acurerosrrata, but bere is also

a tendency for animals lo be about the estiniled

length (ce 3,2-3.6 m)at weaning (Ross ef af 1975).

In thy eastern United States, 17% of the ectuccan

stranding events involved five animals (Mead 1979).

All species with a high incidence of active stranding

were offshore forms, Pépseter macrocephalys, 2

deep water species, mast commonly stranded alive,

in contrast to the situation in South Australia and

other places (Rice 1989), where single animals are

usually dead or moribund. In ‘South Australia, at

least 15% of the recorded strandings were active,

but recent information suggests a much higher

perentage.

The South Australian marine environment is

unique: low nutrient levels are offser by upwelling

events and proximity to the rich warers south af the

Antarctic Convergence; the Indian Ocean influences

the occtanographic conditions through sever!

currents; and the gulf waters afford some protection

frony wind and cold wafer, al least al some times

af year.

It is hoped that additiums to the cerecan

stranding data base. coupled with a better insight

to the State’s oceanography, will permit nmiore

detailed analysis of the trends identified kere and

lead toa greater understanding of the biology of

the many species of whales that occur in sourhern

Australian waters.

Acknowledgments

We owe special thanks to National Parks and

Wildlife Service and Department of Fisheries

officers for reporting and assisting with collecting

stranded specimens, and to B. Stopp for helping to

coordinate the repurting system and allowing aecess

1o National Parks and Wildlife Service records. J.

Bannister and L. Gibson supplied information on

Western Australia and New South Wales strandings,

respectively. We also thank L, Queale, T, Sim, P.

Cockerham and J. MoNamara for technical

assistance and the many voluntecrs, especially M.

Senn, who helped jn collecting and processing

specimens, offen under the most trying and

unpleasant circumstances, Dr P. Petrusevics

provided helpful cammenis on oceanographic

features of South Australian waters and Dr G, Russ

on the manuseripr. G. White and H. Gurres drew

our attention ta the 1963 mass stranding at

Merdayerrah, J. Norman and T. Rajic laborigusly

checked many possible sources to complete our

stranding records and J. Foulkes examined the

phetographic file, J, Thurmer prepared the figures

and D. Lowery typed the manuscript,

WHAIF STRANDINGS IN SOUTIT AUSTRALIA 5

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distribution of Leeuwin current waters off southern

Australia. Aust... Mar. Freshwater Res, 37, 1-10.

Ross, G. J. B. (1984) The smaller cetaceans of the south

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___ (1989) Ziphiidae. Pp. 957-964 Jn D. W. Walton

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TRANSACTIONS OF THE

ROYAL SOCIETY

OF SOUTH AUSTRALIA

INCORPORATED

VOL. 115, PART 2

RELATIONSHIP OF RESIDUAL HILLS AND SHEET FRACTURES IN THE

GAWLER RANGES AND ENVIRONS, SOUTH AUSTRALIA

BY E. M. CAMPBELL & C. R. TWIDALE*

Summary

The domical hills developed on silicic volcanic rocks in the Gawler Ranges are typical of many,

perhaps most, of their counterparts in other parts of the world, in that they are most satisfactorily

explained as etch forms. That is, they are features that have evolved in two stages, the first

involving subsurface weathering, the second the stripping of the regolith to expose the bedrock

surface beneath. Some of the residual hills developed on granite in the western part of the Ranges,

in the Kondoolka and Hiltaba areas, not only differ in morphological detail, but also in their

possible mode of origin. Moreover, the region offers evidence relevant to a long-standing debate

concerning the origin of sheet fractures.

KEY WORDS: Gawler Ranges, silicic volcanics, granite, sheet structure, bornhardts, etch surface

Trunseceiis of the Raval Sucwty of S. Avse W989, 115(2), 53) 66.

RELATIONSHIP OF RESIDUAL HILLS AND SHEET FRACTURES IN THE

GAWLER RANGES AND ENVIRONS, SOUTH AUSTRALIA

by E.M. CAMPBELL & C,R, Twa) FF

Summary

Campuun., EM. & Twibank, C. R. (L991) Reladonship of residual hills and sheet fraciures in the Gawler

Ranges and environs, South Australia, Trams, KR. Suc: §. Aust, W5(2), 33-66, 31 May, 19ST.

The domica) hills developed on silicic volcanic rocks it the Gawler Ranges are typical of many, perhaps

most, of their counterparts in other parts of the world, in tha) they are mow! salisfactoniy eaplaines) as

erch forms, ‘That is, they are features that have evolved in two stages, the first involving subsurlgce weathering,

the second the stiipping of rhe rezohrh to-expose the bedrock surface beneath. Some of che vesidial bills

developed on granite in the western par( of the Ranges, in the Rondoolka and Hiltaba areas, non only Wither

in. morphological detail, but also in their possible mode af orivin. Moreover, the region offers evideriye

relevant to & lone-sfanding debate concerning she origin of sheer fractures.

Kry Worps Gawler Ratiges. silicic valeanics, granite. sheet structure, bornhardts, clch surface

Introduction

The Gawler Ranges ts a prominent upland

located to the north of kyre Peninsula, in the aridl-

semiarid interior of South Auatalia (Fig. 1

Campbell 1990‘). Developed mainly on the Gawler

Range Voleanies of Middle Proterozoic age (Blissctt

& Radke 1980; Blissett ef u/, 1989), Lhe massil

stands highest in the south, where it rises steeply

200-250 m from the predmiont plains ro elevations

of over 400 m, Ta the north, hill crests are tower

In the Moonarce area the maximunt height abovy

sealevel is 336 m in Bond Hill and Chitunilga Hill,

ar 312 m above sealevel, is the highes! peint in the

Kokatha region. Here the terrait ts alsa. more apen,

but the transition trom hill to plain is everywhere

abrupt, For the most part, and especially in the

suuth, the massif comprises ordered rows of

damical hills or bornhardts (see Borohardt 1901);

Willis 1994; Twidale 1982a) developed on valcanic

racks.

The Gawler Range Volcanics are crystalline and

compact. The rock is physically hard and

impermeable. Though well fractured, ftw of the

partings ate open and the rock is esseritially nassive.

On the orher hand, angular blocks up to SU cin

diameter form a discontinuous veneer on the

hillslopes.

The impermeable character of the yalcanics

cnhunees the significance of [ractures, for it is only

by Way of such parlings that water can penetrate

Dept ai Gealozy and Geophysics, University of

Adelaide, G.POe Bow 498, Adelaide, South Aust. S001,

) Campbell, LM. (990) Structure and surface in the

Gawler Ranges, South Australia, Pt D. thesis, University

of Adelaide, Unpublished,

Blisselt, AD, Parker, Al & Schelfler, ALL 1989)

Cywler Range Fxcursion, October 6-9th 1989 Pepe

Mines & f£neruy SN. Aust Rep BR RII

into the rock mass, Three lracture systems,

orthogonal, sheet and columnar, crucially influence

the morphology of the hornharuts. Each bornhardr

is developed on a fracture-defined block of square,

rectangular or, Tnore rtrely, (rianegular plan shape

(Fig. 2). Each is related to & well-developed, if in

places distorted, orthowenal fracture system the

components of which trend predominantly

northwest-northiriarthwest and north-northeast.

(In profile the domical shape of the hills is

associated with convex-upward sheet structures up

to 3 m thick (Fig, 3). In detail, banks of columns

defined by polygonal fracture systems, are

prominent components of hillslopes (Fig. 4). Some

of these columnar joints are due ta contraction on

cooling but orhers are of tecronic origin and are

related to the regional scale orthogonal systems.

The eslumnar and orthogonal systems are of

<onsiderable antiquity, both predating the Late

Proterazoic (Campbell 1990'; Campbell & ‘Iwidale

1991). The age of the sheet fractures is more difficuls

ro determine but they are probably older than the

Beck Surface of Jurassic age (Cumphell 1990').

Thus the basic morphology of the hills is

determined by gsealogical features of great antiquity,

The Gawler Range Volcanics consist

predominantly of dacite, rhyodacite and rhyolite

that originated as a layered sequence of ashilow

deposits orignimbrires. Thev were deposited abour

1592 Ma. About 1583 Mu they were miruded by the

Hiltaba Granite Sujie (Fanning et a/, 1986; Webb

ef al, 1986). The radiometric age determinations

oblained from the veleanics and the granite confirm

the observed ficld relations becwwise the former are

intruded by the latter. The oceurrenee of renimanis

of the valvanicvs at M1 Cooper, Perfection Well and

the Nuyts Archipelago, suggests that the plutunie

mass was largely eniplaced within the ignunbrite

depnsic, though much af the original volcanic

4 E.M. CAMPBELL & C_R, TWIDALE

Lane Hatt aye

A y

oe.

Chitanilga., Hi Mey

Bq Sokatha

atc SM

Yarrana Hill A) 3 = ye

et. ¥

Eromanga Bagin

‘ p 4

_r~ Eyre). Ade

Le Peninsula ra tinB a

> Major upland m4 the

Gawler Ranges.

(720 Lakes

—— Highways

» Hills

@ Towns

Homesiaads

Waulkinna iad? Alfalone

Mt Sturt

aYarwondutta Rock

SIREAKY Bay

Gpita Hin 4 Mt Couper

WUDINNA

Ueontichia Hille

a Wudinna Hill

* Ponda Rock

ae Barna Hil

4 Carappee Hill

big. 1. Location map of the Gawler Ranges, South Australia

outcrop has been eliminated by weathering and

erosion, reducing the original minimum extent from

ubout 80,000 km’ to the present 25,000 km*

(Blissett 1985):

Within the Ranges the granite is extensively

exposed only in the western region around

Kondoolka and Hiltaba homesteads and in the

nocth near Kokatha Homestead. In the southwest,

the granite was emplaced in wu northwest-southeast

trending zone extending through Catton Nob

(Fig, 2), The zone of intrusion runs parallel to the

fHacturé-related Clorrobinhie Depression (Bourne et

al, 1974; Binks & Hooper 1984) suggesting that it

too may have been developed along a major fracture

2one, The centact between the granite and the

valeanies is irregular in detail. Patches of voleanic

rocks ate capriciously distributed on what are

essentially granite hillslopes. Some contacts are

veitly inclined, but most are steep (ig, 5). A

yegelation change commonly accentuates the

contrast in rock type, for spinifex is characteristic

of the upper slopes of volcanic hills, whereas the

vezelation of the granite outerops is generally

dominated by Acacia tarculensis JM. Black

(Whibley 1980) and Eucalyptus brachyealyx —

Blakely Clessop & Toelken 1986),

Origin of the bornhardts

Borhhardts developed in Gawler Ranee Valeanics

Though all are domical, the bornhardts of the

Gawler Ranges, like their counterparts in other

areas, Varv in their precise form. Individual hills rise

to amaximin of 200 m above the adjacent valleys

or plains, but low Tises only a few metres high are

also common. [solated forms are up to 2 km in

diameter, but many burnhardts form interconnected

froups that together measure up-to 10 km in length

(Fig. 6). Typically. the bornhardts are rounded or

RELATIONSHIP OF RESIDUAL HILLS AND SHEET FRACTURES 55

Canter y: \ s

Gawler Range Volcanics

Hiltaba. Granite Suite

Sand dunes

Fractures

Drainage

Traoks

Foliations

Homesteads

Hills (elevation jn metres)

Bores (depth in metres}

Fig 2, Plan of the western Gawler Ranges showing the

fractures,

elliptical in plan, though the linked forms are

irregular.

As was recognised by Dickinson (1942) there are

prominent bevels preserved on the bedrock crests

of many of the bornhardts together forming a

summit surface which has been called the Nott

Surface (Twidale er a/. 1976). Most bornhardts are

devoid of regolith. Evidence of significant deep

weathering of the bedrock occurs only in major

valley floors, though the upper slopes of some

bornhardts carry patches of a ferruginous rind, The

only known remnants of an old regolith, with

corestones set ina Weathered matrix, are preserved

on midslope sites north of Paney Homestead and

near Notining Homestead.

\MT POLLARD, |

eo) 305

“ / {- >

wt ke

Ne aa

ae MT PYRAMID)”

*™ 380', a

ieee

BARBER HLS NY

156 we

a "74,

distribution of valeanic and granite outcrops and inferred

These bornhardts are all probably of etch or two-

stage type (Falconer 1911; Linton 1955; Biidel 1957;

Twidale 1964, 1982a, 1982b; Campbell 1990!),

During the Jurassic (and possibly earlier times) the

then outcrop of the volcanics was reduced to a

surface of low relicf beneath which a regolith

developed, largely as a result of moisture attack

(Fig. 7). The depth of weathering beneath this

surface — the Beck Surface — varied, but was

deepest along the fractures that define the

orthogonal blocks. Then, during the Early

Cretaceous, rivers were rejuvenated, The regolith

was stripped to expose the weathering front, the

Nott Surface. Because of differential weathering,

it had been shaped into a series of domical

56 E.M. CAMPBELL & C.R, TWIDALE

¥ ° es } = -— fat we _ _ ae

Fig. 3. Sheet structure coineident with hillslopes in Gawler Range Volcanics, north of Hiltaba Outstation, western

Gawler Ranges.

{ wa rte ie St sii, ae _ Bama tl

Fig. 4. Banks of columns exposed in lenticular sheet structures, Paney district, southern Gawler Ranges.

RELATIONSHIP OF RESIDUAL HILLS AND SHEET FRACTURES 5?

Fig, 5. Sketch showing various examples of the contact of Gawler Range Volcanics with the Hiltaba Granite Suite

in the western Gawler Ranges. g = vranite. v = voloanic vorks.

‘COTTON NOB

regolith

y N

s | corestone

- Sheet fractures

“ bedrock

zone ot secondary p

fluctuating orthogonal ‘ractures eck Surface

water tabla

bee Ae, 765 ove el nay ee ae

MT HILTABA JURASBIG- = = peujorial Stale orthagdnal tractures

Cadna-owie Tarmation

\ Mt Anua Sandstone (wth

Gawler Range’ Volcanics

cobbles)

EARLY CRETACEOUS > - reas

Algeoucxina Sandstone

Nott Surtac®-S Isumm\)

ay silcrete

- 3 ey sor Py

~Nort Surface -H

(hillslope) 7

EARLY TERTIARY

oe sea

Fig. 6. Contour plan of a group of bornhardts, west of PHESENT , ©

Hillaba Outstation, western Crawler Ranges (Drawn

from South Australia Department of Lands, 1:100,000 Fig, 7, Sections showing development of the Gawler

Topographic Series, Yactoo sheet). Ranges from Jurassic times to the present,

38 EM. CAMPBELL & CR TWIDALF

protuberances separared by fractuye-contrelled

lmear depressions (Wopfner 1969; Campbell 1990;

Campbell & Twidale 1991). The landseape revival

may have been a Gonsequence of uphill alang the

Cprohiuinie Faull, lo give rise ro rhe prominent

south-facing escarpment that delimics the Ranges

on their southern side; or subsidence of the present

Lake Eyre region !o produce the Fromunea Basin;

or both,

The chronology of events responsible for shaping

the volcanic bornhardis is suggested by the

relationships berween the etch surface in ihe Gawler

Ranges and the stratigraphy of the sedimentary

sequenoes preserved in thet Eromanga Basin, located

ta. the north and northwest ofthe upland, The Mt

Anna Sandstone member of the Cadna-owie

Formation, of Barly Cretaceous (Neacomisn-

Aplian) age, contains boulders of Gawler Range

Voleanics (Woplner 1969; Wopfner ef, al 1970),

These houlders ure construcd as erstwhile

corestones formed within the Beck Surface regolith,

The boulders low in the Mt Anna Sandstone are

more weathered than those above them, consistent

with their being derived front the stripping of a

regolih. Current directions and thickness

measurements also indicate a southerly provenance

fer the Mt Anna Sandstone (Woplner 1949). All this

suggests thal Che barnhardes were initiated in the

subsurlace ducing Jurassic times and were exposed

as landfornts during the Early Cretaceous, Thus the

Beck Surface beneath which the bornhardts of the

Gawler Ranges were initiated, was apparently part

of a widely developed middle Mesozoic kindseape,

temmanes Of which are found in various parts o!

central and southern Australia (Iwidale & Canipbell

1988). That rhe exposure of the residuals was

essentially complete by the Cainozdic is suggested

first by the paucity of Tertiary sediments in the

Eromanga Basin and of sediments derived fram the

valeanics in the Eocene and Pliocene formations

preserved Im che Corrobinnie Depression

pulacochannels (Bitks & Hooper 1984), Also,

silgrete is preserved in valley Moors within the

Ranges (Hulton er af. 1978) and low in the

londagraphy, well below the summit surface,

marginal tothe upland The silerew here is probably

of Esuly Tertiary age (Firman 1983) implying chat

the higher (etch) surface is older,

Suyh a two-srage mode of develapment 1s nol

unusual Most bornbardts, though by no means all.

evolve in thts fashion. Such subsurface weathering

demonstrates the primacy of fractures in

determining the subterranean penetration af

meteor'le Waters, hence in controlling the pattern of

weathering developed, and thus in shaping the

bedruek surface and jhe eventual faid' orm

assemblage developed. The Gawler Ranges

bornbardis also iustrate the antiquity af elements

of the Australian landscape (IWwidale (976; Twidale

& Campbell 1988),

Bortharits developed in granite in the southwesrern

Gawler Ranges.

In general, the granite expusures stand lower in

the landscape than da thase of the volwanics, They

are more readily weathered than the volcanics. At

a regional scale ihe granites ef north and

norchwestern Eyre Peninsula have been reduced to

extensive plains while the Gawler Ranges remain

upstanding. This may reflect the lower retative

position of (he eranile, or lectonic uplift of the

Gawler Ranges, bul the vuleanic massif survives

essentially intact whereas only small isolated

granitic residuals, such as Weortitchie Hill und

Wudinne Hill stand above the level of the plains.

In the western Gawler Ranges (Fig. 2) major valleys

have been incised in the granite. yet a summit

surface of considerable extent, and cut in granine,

is preserved argund Kondoclka Homestead (Fig. 8).

Some higher remnanis such os Waverley fill

(385 m) stand almost as high as the niost prominent

local peaks formed iw voleanic rocks.

The preservation of such high eranice surfaces

needs explanation. There is no evidenve af the

height to which the granite was emplaced, because

the highes| peaks, in the Waverley HI[l area, do nal

carry remnants of the valeanic roof thal once must

have been present, The granire emplacement must

have been exposed by the Late Jurissic, sine the

Nott summit surface is well represented on the

granite hills, hoth in the high plains around

Wavetiey Hill, cast of Biltaba Outstation and in the

fartish crests af isolated remnants snch as Corian

Nob (378 m)p. These high remnants owe their

elevation to tectonic forces, as they reflect the height

of upwelling of the granite magma during {ts

emplacement some 1583 Ma,

Cotton Nob is a nubbin that siyes same 30 m

froina platlonn or plinth ile eroded in granite and

which tn furn is hordened by a scarp abouc 21) m

high. Some of the boulders chat cover Colin Nab,

including some at the ercst, are flared, syegesting

thar they were once covered by a regolith (Twidale

1962), They, and the stepped morpholugy of the

area as a whole, can be understood in terms of

Phased or episodic exposure (Twidale & Bourne

S975, Twidale 1982c) and the occurresive of several

low hills in the general area and sruinding some 20

nt ahove the plains or valleys lends support tw this

interpretuiion, On the other hand, the gramte

exposed in the 20 m scarp is, jn phages. more

Mtensely fractured (han cs (he granite of (he nubhbin,

suggesting (hal Ihe stepped morphology could be

RELATIONSHIP OF RESIDUAL HILLS AND SHEET PRACTURES “

Fig. &. Bevelled crests of (he bornhardts together form a prominent summit surface, shown here developed on granite

east of Kondoolka Homestead, western Gawler Ranges.

a reflection of structural factors, though it is

unlikely that fracture density is arranged in

concentric patterns as the plinth and boss pattern

of topography around Cotton Nob implies.

ML Wallaby is another high residual of unusual

shape and origin. Standing 333 m above sea level

and some 130 m above the surrounding plain, Mt

Wallaby is a markedly asymmetrical east-west

trending ridge of granite (Fig. 9). Its northern slope

is gently inclined and boulder-strewn and stands in

marked contrast with the bare, precipitous, even

overhanging, southern face on which polygonal

cracking is well developed, The whole feature, but

especially the southern scarp is linear. That it may

be related to east-west fractures is also suggested,

first by the presence of many prominent latitudinal

joints, many of them with associated lineation of

the bedrock, and second by the presence, a few

hundreds of metres to the east, of two rectilinear

east-west drainage lines, In addition, what may have

originally been essentially flat-lying joints do, in

fact, dip gently northwards. This suggests that Mt

Wallaby may be a half horst upthrust at its southern

margin.

Most of the wranitic bornhardts of the western

Gawler Ranges are low domes standing aboye the

valley floors and plains, and there is here good

gencral argument to suggest that they, like their

volcanic counterparts in the Gawler Ranges, are

two-stage forms. Admittedly there are no exposures

like those at Ucontitchie Hill on Eyre Peninsula and

elsewhere (see Twidale 1982a) that demonstrate

contrasted fracture density between hill and plain.

But, whereas the domes are invariably constructed

of massive bedrock, the rocks beneath the

intervening plains were probably densely fractured

since they are altered to depths of up to at least

80 m, and commonly to depths of 40 or 50 m

(S.A.D.M.E. borelog information — see Fig. 2),

The valley floor domes are characterised by the

presence of numerous boulders, suggesting that the

outer shells or sheets of yranite have disintegrated.

Where domical forms are partially covered by the

voleanic host rock, but which can be seen in section

(Fig. 5), the outer layers of granite are bouldery,

There is no suggestion that the outer boulders

represent globules of liquid granite that have

migrated into the voleanies (cf Barbeau & Géze

1957): they are weathering forms. The presence of

boulders so close to the contact suggests that the

breakdown of sheets takes place beneath the

surface, as a result of the infiltration of meteoric

0) LM, CAMPBELL & C.R. TWIDALE

Fig, 9, Mt Wallaby, an asymmetrical graniti¢ residual in the western Gawler Ranges.

waters through the overlying volcanic rocks and into

the granite below. This suggestion is sustained by

the flared character of some of the boulders

(Twidale 1962). On the other hand, the volcanics

adjacent to the granite are not themselves notably

weathered. Indeed, where the two rock types,

voleanic and granitic, are exposed side by side there

is no major topographic break between them,

though the detailed forms contrast sufficiently for

them to be identified from afar (Fig. 5).

The similarity both in shape and in size of the

exposed and still partially covered pranite rises

suggests the possibility that some of the granite hills

located close to the volcanic-granite contact may

be stocks, bosses or protuberances developed at the

margin of the batholith and exposed by the

preferential erosion of the overlying host rock

(Twidale 1982a, 1982b), Both the exposed residuals

and those seen only or partly in section are boulder-

strewn nubbins. The exposed and the still-covered

appear to display a Similar range of size and shape

(Fig. 5). In these terms the domes are tectonic

forms, Le. they acquired their essential morphology

during intrusion, On the other hand, the voleanics

are evidently more resistant to weathering and

erosion than is the granite, making it difficult to

explain why the host rock should have been

weathered and eroded, leaving the granite

protuberances intact. Doming and stretching of the

volcanics, possibly consequent on the intrusion af

the granite, could provide a partial explanation, bul

the problem essentially remains.

Thus, whereas the bornhardts of the yaleanic

outcrops of the Gawler Ranges are sensibly uniform

in morphology and are all likely of the same origin,

the hills of the granitic areas vary in their detailed

morphology and, possibly, in their genesis. Some

are congeners of the volcanic bornhardts, but others

may not be. All however display fracture control in

plan.

Inselbergs af adjacent areas

Some voleanic bornhardts within the Gawler

Ranges, particularly those located on the margins

of the massif, stand in essential isolation and thus

can be considered inselbergs — rocky residuals

standing abruptly from the level plains around

them. Inselbergs are also well represented in the

southern piedmont of the Ranges and in the areas

west and south of the upland, Indeed, the inselbergs

of the southern piedmont like the twin peaks of Mt

Sturt, Waulkinna Hill, Peterlumbo Hill and the well

named Mt Allalone are of some historical interest

for they, together with the scattered volcanic hills

of the eastern part of the Gawler Ranges, stimulated

Eyre (1845, p. 203) to compare them with ‘so many

islands in the level waste around them’, thus

anticipating the now commonly used term

‘inselberg’,

RELATIONSHIP OF RESIDUAL. WILLS AND SHEET FRACTURES al

The lithology and morphology of these inselbergs

varies. Most are granitic in composition though

their textures aud detailed ages differ from area to

area. All, however, are members of the Hiltaba

Granite Suite, The hills developed on these materials

include such prominent peaks as Wudinna Hill and

Uconntchie Hill as well as low, large-radius dames

like Childara Hill, and Polda Rock, Carappee Hill

is a prominently stepped gneissi¢ inselberg, and (he

low domes that comprise Waddikee Rocks are also

gneissic, Most are dormical but Waulkinna Hill is

a nubbin and several others like Ucontitchie Hill

carry blocks and boulders.

Isolated elongate residuals, such as Uno Range

and Botenella Range composed of sedimentary and

metasedimentary rocks are prominent features of

the landscape of northern Eyte Peninsula. Some,

like Caralue Bluff, have bevelled crests (see also

Dickinson 1942). In addition, there are several

domical inselbergs also composed of sedimentary

material. The areal extent and relief amplitude of

these doinical forms is similar to that of the voleanic

bornhardts of the Gawler Ranges. Mt Allalone,

developed on Proterozoic conglomerate, is a low

domical hill tising to 342m above sea level and 120

m above the surrounding plain. Curtinye and Barna

hills are composed of Proterozoic quartzite.

Curtinye Hillis over 440 m above sea level and rises

110 m above the surrounding gewly undulating

plain. Approximately | km in diameter, it forms

part of an ancient fold belt, the sediments of which

were highly metamorphosed during the Proterozoic

Kimban Orogeny. As a result of this

metamorphism, the rocks in Curlinye Hill were

altered Lo quartzites and a sehistosity, striking 125°

and dipping 80° to the northeast, was developed.

The domical form of the hill is only slightly

dissected by streams which radiate from the summit,

Some have straight sectors which follow the

schistosity; others follow the trend of a series of

vertical lractures which are predominantly aligned

at 55° and 175°, Sheet fractures essentially parallel

to the surface of the hill are particularly well

exposed Of Oulcrops near the crest and on the

eastern slopes (Fig. 10).

Whatever their composition however, the

inselberg bornhardts and nubbins of northern Eyre

Peninsula are defined by fractures (hat form

orthogonal systems (Vig, 11) and thus reinforce the

suggestion (see e.g, Birot 1952: Rognon 1967) that,

through the control they cxert on the course of

weathering, fractures are of prime importance in

Fig. 10, Downslope view on Curtinye Hill, near Kinba, northern Eyre Peninsula, showing sheer fractures parallel

To the surface and cutting aeross neur-vertical foliation in the Warrow Quartzire.

62 E.M. CAMPBELL & C.R. TWIDALE

Legend

Inferred fractures

~ Sheet fractures

* ~~ Foliation

Ucontitchie Hill Yarwondutta Rocks

Sedimentary Rocks

- MY lie

Mt Allalone

Curtinye Hill

Waddikee Rock

Fig. 11. Prominent orthogonal fracture systems are of prime importance in determining the shape of the inselbergs

on Eyre Peninsula.

RELATIONSHIP OF RESIDUAL HILLS AND SHEET FRACTURES 63

landforin development. The inselbergs of northern

Eyre Peninsula are, like their volcanic congeners in

the Gawler Ranges, two-stage Forms (Twidale 1964,

19820). The crests of some of the higher forms,

residuals like Wudinna Hill, Ucontitchie Hill and

Carappee Hill have been putatively dated as bate

Mesozoic (Twidale & Bourne 1975), If so they are

equivalent to the Now Surface and were iniliated

by weathering beneath the Beck Surface.

Implications for explanations of sheet structure

Sheet fraciures in gianitic rocks

Sheer structure in the granitic rocks, bow within

the Ranges und to the south on Eyre Peninsula, very

much gantorms with chat deseribéd froni similar

lithologteal environments the world over, tis well

developed and exposed iw bornbardts. Flac tying

Tractures occur wherever granite is exposed, as for

instance af Yarwondutta and Calea quarries, In

other parts of the world sheer fractures extend to

al least 100 m (with no indication of being limited

to Chat depth), but here nu such deep exposures are

available. The sheet fraciures, which take rhe form

of either a single parting or a complex of closely

spaced fractures, are cammionly disposed convex

upwards in the hills. They ru) parallel to (he surface,

though sume converme to produce wedees. The slabs

defined by sheee structure ate up to Bot thick fonly

about 4m on Eyre Peninsula), and there is a

suypestion that thickness increases with depth, The

Fractures increase in inclination fear jwajor vertical

fracttires, and cut across lineation, foliation, Mow

structures and urihogunal ioints. Sheer fractures are

In some areas older chan surficial lamination, bur

younger than the orthogonal fracture systems.

Sheer Jraomures 17 volounie racks

In the volcanic areas, hillslopes consist of hanks

of partially overlapping slabs, that form short,

wonvex-upward bedrock slopes (Fig. 3). They are

bounded on their lower sides elther by cliffs up to

10 m high in which verical fractures defining

columns are well exposed, or by debris-covered

slopes, At and near the summit of the hills the stizer

structure 1s exposed in horizontal benches, In the

mittor valleys cut in the barnhardts, sheet fractures

are generally synformal, though in places, such as

near Eucarro Dam, southwest of Nanning Horme-

stead aad al Pelerby Hill, comvex-upward sheer

Fracruires are exposed in the valley floors. In other

vallevs, hawever, the shee factures dip towurds phe

valley anis. Che sheet fractures are inclined as nich

as 40° from the horizontal, but more commonly.

within the range 5-20° Sheet (ractures in the Gawler

Range Yolcanics, unlike those reported from many

otfier areas. tend to divide the rock mass into »

series of lenses, each lens being about 4-30 m across

and J-8 m high (Fig. 4). In places the geometry of

the sheets is itregula:, tor example in a valley incised

in Peterby Hill swells and depressions that are V-

shaped in cross-section reflect the disposition of the

shect fractures. The sheets below, though of similar

morphology, are offset with the depressions being

located below the swells of the sheet above and the

tyoughs beneath the arches.

The maximum observed thickness of sheets in

Gawler Range Voleanics is 3 m. Jn many exposures,

eg. al Spring Hill, sheets abour 1 m thick are

underlain by thinner (10 emi) sheets. However, deep

exposures are rare. Evidence (rom the quarry at Mt

Cooper suggests that sheet scructure may nor extend

to great depth; alechough sheets of one metre

thickness are present at the surface, sheet steucture

is not developed in the [0m or so of rock exposed

in the quarry! faces.

The surfaces of sheels are essentially fresh, with

file or no weathering of the adjacent rock.

Polished sur faeces, siickensides and the offsetting of

columns along shect fractures at a few sites suggest

local lateral dislocation.

Though lucally variable, sheer fractures tend to

he essentially flar-lying on the crest of the

bornhardts and to increase in declinatiun downslope

lowurds the valleys that are associated with the

regional and smaller scale orthogonal systems of

fractures, suggesting thal the sheel structure is either

contemporaneous with, or younger than, the

orthogonal systems.

Sheet [ractures and vertical eolumnar joints

impart astepped profile to many hill slopes. Some

sheet fractures terminate abruptly against the

columnar joints, although many merge with the

sheet fracture above or below. Sheet fractures

transect columnar joints, both of contraction on

cooling and of tectonic origin. The sheet fractuses

must postdste the columns. This relalive age ts

indicated by the offserting of columns lune the

sheet Tractures.

The sheet structure in the bornhardts developed

in Gawler Range Voleanics is not typical of rhat

Uescribed from ihany other areas in thar it is

associated With columnar joints due to contracrion

on coolijie. The lenticular forms are out typical ot

sheet structures in general, nor are the evident

restriction of the sheet structure to the superticial

zones and the presenve of thinner sheets beneath

thicker ones. The close association of sheet siructure

with the morphology of the borthardis is, however,

characteristic.

64 boM. CAMPBELL & C.K. TWIDALE

Sheet fraclares in sedimenmry rocks

Sheet fractures have been deseribed from

sandstone in several parts of the world. Bradley

(1963), lor instance. described thick sheet structures

develuped m the massive Navajo Sandstone of the

Colorado Plateau. Ayers Rock in central Australia

ig a well known example of an inselbere developed

in Cambrian arkose and displaying slicet structure

(Twidale M78) Similarly, Sheet lractures are well

developed in the domical inselberes cur in quarrzie

on northern Eyre Peninsula: Although on Me

Allulone che sheet fraciures are apparent on only

8 few Limited outcrops, on Curtinye Hill they are

Clearly exposed as bare ruck autereps, particularly

on the eastern side, The sheet structure cuits across

the tolation, and also dips towards the bottom ul

the miner joint-conirolled valleys that score the hill,

Tt is flat lying near the crest, but inereases i) dip

to about 4° further dows the slope.

Origins of sheet fractures

Sheet fractures have been explamied ii many ways

One widely accepted explanation links them wilh

erasional offleading or pressure release, The

geomeiry of the fractures is regarded as consequent

upon the form af the land surface. The hypothesis

was developed in relation 10 granitic (Gilbert 1904)

and other plutonic rocks that were, a was angued,

emplaced deep in the crust, at a depth of several

kilometres, but Which have been exposed as a resull

of the erosion of the original superincumbent toad.

The implied vertical unloading has given rise ta

radial gapansigon of which tungetitial liractires

sheet fractures - are an capression. Though the

hypothesis has been enticised (see eg. Dale 3923;

Coates 1964; Twidule 1964, 1973) the offloading,

unloading (see eg. Skinner & Porter 1987) a1

pressure release hypothesis ts sll (he mast luvqured

explanatian of sheet structure Indeed many

geologists refer to sheer fractures as pressure release

or witloading jos.

The ather generally accepled explanation ol sheet

fractures involves horizantal compression, There is

much evidence and argument to support this

suggestion (Merrill 1897, Ditle 1923; Twidale 1964,

1973). In particnlar, many parts ol Australia have

been shown by direct measurements to be in

Substantial conipressive stress (Moye 1958; Denham

ef at 1979), Also a suile of minor landforms

associated with the release of compressive stress has

heen described fram northwest Eyre Peniisule

(lennings & Twidale 1971; Twidale & Sved 1978;

Twidale 19846), In (lis refard it is notable that within

and around the Gawler Ranges sheer structure

geeurs an bornliandes. of varlons lniologies.

inchiding silicic valeanic rocks, which have not been

deeply buried. This poses problems for ite

offloading hypothesis. Che orthagenal fractures of

the Gawler Ranges are of Proterozoic age. The

iniiidlion of the bornhardts at thé weathering [runt

did not take place until the early or middle

Mesozoic. Lis difficult to undersrand why, wiver

the existence of orthogonal fracture systems, anv

tendency ty radial expansion consequent on

unloschng was not relieved along partings already

in existence rather than through the development

of a aew sel of tractures, Jt is alsa questionable

whether the pile of volcanic rocks ite whiell the

sranite of rhe western Gawler Ranges was introded

was of a sufficient weight w have imposed vers ical

stresses the relict of which would have produced

sheet fractures.

The occurrences of sheet fractures in sedimentary

tovks is difficule to explain in terms of the

offloading hypothesis where regional and local

evidence is suggestive of Compression. Curtinye and

Barna hills are purl of an ¢lonvate outcrop of

metasedimentary rocks that ought to give rise te

strike fidges such as indeed gre found on

Proterozoic quartzites, for example, in the Botanella

Rangc, in a varied sequence al Proterozoic

sedimentary rocks on Eyre Peninsula and in fold

mountain belts the world over. Similarly Ayers Rock

ought tobe a strike ridges instead there is a bevelled

dume. ‘The sheel stricture is surely not consequent

on the form, as is implied by (he offluading

hypothesis, fur the ‘nitural’ form is a ridge or a

range, Cross folding or shearing may be responsible

for the stresses Which gave mse to shee! structure

in Ayers Rack (Twidale (978) and shearing nay have

produced the sheet fractures of Curlinye and Barna

hills (Fig. 12).

Be that as ic may, the sheet [ractures af the

Navajo Sandstone oveur in a diflerent teetoni¢

regime, namely within or close to the Rio Grande

(extensional) Rift (Baldndge & Olsen 1984).

Convex-upward sheet structure as developed in the

Organ, Mountauts and alsa in the Ria Grande

tensional zone, southern New Mexico (e.g. Seager

1981). Thess, too, may be interpreted as cvidence

that sheeting fractures can develup in non-

compressional environments, There can be local

compression even in 4 region of extension, just as

horsts have been thruse uy within tensional pitts.

Sheet Tractures could, for example, be interpreted

is due to upthrust during emplacement of the hor

lencous rocks tof Beche 1839; Whitney 1865; Harris

185s). On atance, however, the offloading

hypothesis of sheet Fracture formation sugeested hy

Bradley (1963) muy. be valid in the Nuvajo

Sammdbsturie, though Wseess untikely i the oorlexs

oY the Ausiraliun areas discussed,

RELATIONSHIP Ol RESIDUAL FILLS AND SHEEL WRACTURES 65

| |e ae

Lesa! meats ~~) a

Hie. 12, Cross-lolding and shearing, suggested

explanations for the stresses which give rise to sheet

structure, and hence the damical foo of the inselberys.

A — compression, B - shearing.

Conclusion

The bornhardts developed on volcanic rovks in

the Gawler Ranges are features. Of considerable

antiquily that have evolved in two stages, the first

involving subsurface differential weathering, the

second the sreipping of the regolith to expose the

bedrock surface beneath, The plan form of the

bornhardts is related to complex systems of

orthogonal fractures which developed during the

Middle Proterozoic. Their domical profile is a result

of the development of sheet fractures, which were

apparently present prior 16 a Jurassic period of deep

subsurface weathering. The detailed slope

morphology of the bornhardts reflects the presence

of orthogonal fractures.at various scales, of sheel

fractures and also of columnar joints due to

contraction on cooling of the extrusive mass, The

bevelled crests of the bornhardts are part of a,

sumroit surlace which was expased by stripping of

the regolith in Early Cretaceous times.

In Ube western Gawler Ranges barnhardts of

varied morphology are developed on granite which

was intruded into the Gawler Range Volcanics in

Middle Proterozoic times, Orthogonal fracture

systems, atid shee| fractures. are of fundamental

importance in determining the shape of these

features. The development of’ some of these granitic

bornhardis may be due to tectonic forces or to

strucioral factors; some may be stocks exposed by

erosion of the host rock; most, like their

counterparts in the volcanic rocks, are two-stage

farms,

The shape of isolated domi¢al inselbergs

developed on resistant rocks of varied composition

on porthwestern Byre Peninsula is also fracture-

controlled, Orthogonal and sheet fractures

influenced iheir morphology. These aay algo be

etch forms.

The residuals of different lithologies present

various problems tor the offloading hypothesis of

sheet fraclure formation. Although there is no direct

evidence that the sheer structure is due to lateral

compression, the confinement of the domes with

sheet structures to blocks defined by orthogonal

systems, the forms and measurements indicative of

crustal comipression, and the development of dames

in quartzites can all be construed as corroborating

this suggestion.

Acknowledgments

Thanks are due to the Royal Society of Sourh

Australia Inc. for two grants from the Saciety’s.

Endowment Fund.and to the Australian Research

Commission and the Commonwealth Postgraduate

Scholarship Scheme for financial assistance.

Techical ussistance was provided by S. Proferes and

R. Barrett. The authors wish to.thank J. A. Bourne

for constructive suggestions on an early draft of the

paper, and Drs V. A. Gostin and J. E. Mueller for

helptul comments. during the review process.

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DESCRIPTIONS OF THE TADPOLES OF SOME AUSTRALIAN

LIMNODYNASTINE LEPTODACTYLID FROGS

BY MARGARET DAVIES*

Summary

Early development and late stage larvae of the limnodynastine frog species Heleioporus

albopunctatus and larvae of Neobatrachus centralis, N. wilsrnorei, N. kunapalari and Mixophyes

schevilli are described and illustrated.

KEY WORDS: larvae, embryos, Heleioporus albopunctatus, Neobatrachus centralis, Neobatrachus

wilsrnorei, Neobatrachus kunapalari, Mixophyes schevilli, frogs

fhomsections of ike Koval Sec ar § dust. ePOOl). U1SC2), 67-76.

DESCRIPTIONS OF THE TADPOLES OF SOME AUSTRALIAN LIMNODYNASTINE

LEPTODACTYLID FROGS

by MARGARET DaVies*

Summary

Davins, M. (199t) Deseriptions of the tadpoles of some Australian limmodynastine leptodactylid: frogs.

Trans, R. Sov, 8. Aust (t8(2), 67-76, 3L May, 1991.

Karly development and lute stage larvae of the limnodynastine frog species Heleioporus albopunctalys

and larvae of Weohareachus.centratiy, N. wilsmarei, N. kunapatar/ and Mixupiiyes schevilli are described

und ilhrstrared,

Rey Worn larvae, embryos, Heleioporus albopunctatus, Neobatrachus centralis, Neobutrachus wilkmorei,

Neubatrachus Kunapalari, Mixaphves sehevilli, trogs

Introduction

Despite considerable advances over the last

twenty years, larval forms of many Australian frogs

remain unknown, Tyler (1989) documented those

for which descriptions have been published, but the

breadth of qaformation available for the listed

species varies considerably, For instance, same

papers provide a composite description of larvae

but lack any early developmental data (e.g, Waison

& Martin 1973) or illustrations (eg. Lee 1967):

authors rarcly have access to complete lile history

lala.

In addition, larvae attributed to. species in the

carly 1960's may be wrongly identified (c.g, larvae

of Neobatrachus ptetus and N. centralis, Martin

1965; Watson & Martin 1973). For this reason

additional data are hecessary to ensure association

of larvae with adults of species as currently

recognised.

During studies of the ontogeny of bane of

lininodynustine leptodactylid frogs, tadpoles of

various species have been reared. Some of these have

not been deseribed, or have been unreliably

associated with adults. Here | provide information

on the early development.and larvae of Heleroporus

albopunctatus and on the larvae of Neobatrachus

centralis, N. wilsmorei, N. kuaapalari and

Mixophyes schevilli,

Materials and Methods

Material reported here is housed in the Dept of

Zoology, University of Adelaide. Larvae were staged

according to Gosner (1960). Measurements were

tnade using dial calipers measuring to 0.05 mm or

with the aid of an eyepiece micrometer.

Measurements taken (in. mm) were; total length

(TL), body length (BL.),

* Depr of Zoology, University of Adelaide, GPO. Bow

498, Adelaide, S, Aust. 5000

Line drawings were made with the aid of a

camera lucida attached to a Wild M& sterca

dissecting microscope. The format of the larval

descriptions follows Anstis (1976).

Results

Heleioporus albopunclatus Gray

FIGS 1-5

The earliest stage available was an embryo within

the egg membranes at stage 20 (Fig. 1) which is

perched on 2 large yolk sae around which the jail

is wrapped Isterally, In five of six embryos the tail

is Wrapped sinistrally and in the sixth, cextrally. The

cornea is Not yet transparent but the eye is partially

plgmented, External gills are not apparent and

ventral adhesive organs cannot be located. The

stomodaca! pit has differentiated into a mouth

consisting of an upper lip overhanging an

unperforated mouth cavity, The-olfactory pit also

is unperforated, A well-developed orifice (anal

opening) is located ut the junction of che tail with

the body (Fig, 1), The embryo is lightly pigmented

with a brilliant yellow yolk sac.

By stage 22 the ollactory pit has. become

perforated to form the nares. Two upper and two

lower tooth rows have begun io keratinize, The tail

remains Wrapped around the yolk and curved up

over the face covering, one nostril and the corner

of the cye (Fig. 2). The yolk sac shows the

beginnings of coiling and the anal opening remains

prominent. Blood vessels are present on the tail fins.

A yellow spot occurs on the top of the head at the

level of the anterior extremities of the eyes, The

embryo remains in the egz membrane at this stage.

External gills are absent but an aperture is

detectable on the left hand side in the usual position

Gf caternal gills. Ventral adhesive organs also are

absent. The embryo is covered with a Nive dusting

of fine brown pigment.

68 M.. DAVIES

Fig, |, Embryo of Helcioporus albopunctatus at stage 20: a, dorsal and b, lateral views. Scale bar = 1 mm,

ee |

Fiz. 2, Embryo of Heleiaporus albopunciatius at stage 22: a, lateral and b, dorsal views. Scale bar = [ mm.

By stage 24 the embryos have hatched. The tai!

is flexed at its tip and the mouth is situated

anteroventrally (Fig. 3). Later al this stage the tail

loses some of its flexure, The anus 1s dextral and

open, The opercuium has closed but the spiracle

has not formed.

By stage 25, the mouth ts directed anteroventrally,

The tail is still very slightly flexed and the spiracle

is forming. Coiling of the gut is detectable, but it

remains yolk-filled.

At stage 26, the gut appears yolk-filled but is fully

coiled. Keratinization is incomplete on the tooth

rows but is apparent in some state on all the

presumptive mouth structures,

By stage 28 the tadpoles are feeding and the

mouth parts are fully formed,

A larya al stage 33 is illustrated in Fig. 4,

The body is elongate and widest at the eyes. I'he

snout is evenly rounded in dorsal view and slopes

gently in lateral view. The nares are dorsolateral and

sessile. The moderately large eyes are dorsolaieral.

The spiracle is sinistral, ventrolateral and visible

(ram above. It opens posterodorsally and has a

constant diameter along its length. The anal tube

is dextral, long and opens about hallway up the

ventral fin. The tail fin is not arched and is gently

rounded terminally with the dorsal fin extending

along the posterior 1/10 of the body and deepest

TADPOLES OF LIMNODYNASTINE LEPTODACTYLID FROGS 69

Fig. 3, a, Dorsal, b, ventral and c, lateral views of a larva

of Heleioporus albopunctatus al stage 24- Scale bar =

1 mm.

approximately halfway along its length. The yentral

fin is deepest about 1/3 of the way along its length.

Tail musculature is moderately thick and tapers to

a fine point posteriorly. The mouth is anteroventral.

There are six upper and three lower rows of labial

teeth. The second - sixth upper and first lower rows

are divided (Fig. 5). The horny beak is of moderate

proportions. Tail musculature is pigmented and

Pigmentation also occurs on the dorsal and ventral

fins (Fig. 4).

Meristic data of tadpoles are provided in Table 1.

Commeni: Lee (1976 p,388) reported that the tooth

row formula as recorded here is the “maximal

observed” and that variability occurs: “commonly

1 or 2 complete, up to 5 incomplete upper labial

tooth rows; 3 lower labial rows, 2 may be

incomplete”.

The mouth illustrated in Fig. 5 is essentially the

same as that illustrated by Watson & Martin (1973)

for H, australiacus. However the absence of external

gills in developing 4. a/bopunciatus is not mirrored

in H. australiacus. Watson & Martin (1973) reported

external gills in this species. No comment was made

about absence of adhesive organs. Adhesive organs

(oral suckers) rapidly disappear after stage 21

_—— |

Fig. 4.-a, Lateral and b, dorsal views of a larva of Heletoporus albopunctatys at stage 33. Scale bar = 10 mm.

70 M. DAVIES

TABLE 1. Measurements (in mm) of tadpoles.

Species Stage Total length

x (range)

Neobatrachus 27 23.8

centralis

29 36.7

32 35.5

(35.5)

34 37.9

(37.6-38.2)

35 37.2

36 43.31

(33.7-50.7)

37 50.0

(48,2—51.8)

38 49,99

(46.0-57.1)

39 50.93

(49.3-52,3)

41 49,77

(45.6-54.5)

42 39.47

(35.7-41.7)

43 31.38

(22.5-40.7)

44 19.43

(15.8-21.6)

N. wilsmorei 34 49.6

40 59.8

41 62.9

(61.4-64,4)

N. kunapalari 36 40.7

Mixophyes 24 98.03

schevilli (86-116,4)

25 87.53

(75.4-107.2)

31 58.4

32 59,2

33 63.4

33/34 66,7

34 64.0

35 68.25

(66.7—70.6)

Body length

x (range)

9.8

14.9

15.7

(15.6-15.8)

15.9

(15.3-16.5)

17,4

18.97

(16.8-23.0)

22.7

(22.4-23.0)

21.46

(19.8—25.0)

23.19

(21.6-23.6)

19.58

(16.9-21.1)

17,23

(16.1+18.0)

17.66

(16.9-19.3)

16.95

(15.6-20.1)

19.1

26.1

(25.7—26,5)

17.1

34.6

(31.1-39.5)

30,27

(23.3-36.0)

21.5

22.9

25.23

(24.6-26.6)

TADPOLES OF LIMNODYNASTINE LEPTODACTYLID FROGS 71

TABLE |. Measurements (in mim) of tadpoles. (continued)

Species Stage Total length Body length n

= (range) ® (ange)

Mixophyes (continued) 37 70.13 23.63 3

schevillt (67.2-75.0) (23,1-24.2)

38 77 23.4 1

39 74.45 21,95 2

(72.7-76.2) (21.2-22.7)

Heleioporus 26 15.3 6.0 ]

whopunciatus

27 16.4 6.05 2

(16.3-16.5) (6.0-6.1)

28 22.6 8.6 2

(20,8-24,4) (8,0-9,2)

29 27.0 9.2 i]

33 40.9 14.1 1

35 45.7 15.43 3

(42.5-47.4) (12,5-17,.D)

a7 47.1 15.1 i

38 47.25 17.7 2

(46.6-47.9) (17.5-17.9)

41 55,9 21.2 1

42 44.6 18.1 I

43 28.98 16,55 2

(23.6-34.40) (16.0-17.1)

44 18,5 18.3 !

(Gosner 1960). The earliest stage examined here is

slage 20 and it may be that the structures disappear

at an earlier stage in A. albopunctatus (the form

or oral suckers varies both systematically and

ontogenetically (Gosner 1960)).

sefoncaseainl It

OK Creel

Uy gba

ag ain

uel ‘

5 Minin

wai men

4 so Git Le i

- “ee Mixophyes schevilli Loveridge

FIGS 6-7

A tadpole al stage 36 is illustrated im Fig. 6. The

body is ovoid and widest behind the eyes. The snout

is evenly rounded in dorsal and lateral views. The

nares are dorsal and sessile, opening laterally. The

relatively-large eyes ure dorsolateral, The spiracle

is sinistral, short and ventrolateral with a large

Fig, 5. Oral disc of a larva of Heleianarus albopunctatus orifice directed posterodorsally, and is not visible

at stage 33. Scale bar = 1 mm, from above. The diameter of the spiracular tube is

re M. DAVIES

Fig. 6. a, Lateral and. b, dorsal views of a larva of MixopAves schevilli at stage 3G. Seale bar =

relatively constant along its length. The anal tube

is dextral and hidden in a membranous sac which

also surrounds the developing hind limbs. At this

Slage, the feet protrude from the sac. The anus

opens abour halfway up the ventral fin. The tail fins

are arched, the dorsal fin commenting in ihe

posterior 1/10 of the body, being deepest

approximately half way along its length. The tail

fin is slightly rounded at its terminus. Tail

musculature is thick, tapering 10 # point posteriorly,

Tadpoles are heavily pigmented, pigmentation

extending over the tail musculature and fins. The

mouth is ventral with the oral disc surrounded by

a papillary border of niodeérately fine papillae (Fiz.

7). There are six upper and three lower rows.of sabial

ieelh and five or six rows occur laterally on cach

side near the angle of the jaw. All the upper rows

except [he most anterior are divided. The second

and thicd lower rows aré undivided, The horny beak

is of relatively fine propartions,

Meristie data on tadpoles of M. schevillt are

provided in Table I.

Camment: M. schevilli can overwinter as very large

tadpoles at stage 25 (Trenerry 1988'). This

phenomenon is reflected in the mensuration data.

10 rim-

is en

‘ Cy

™%

or aware

om “0 wg

“0 {wasabi rma

ne xe Whe esau sgoeraeti iaaatanad i MH Gy

(

Poa a 8

Ub iy"! A iehTa ONL orange siya qavneenouet! MaHivand vue {

eh ery tog tee

Sinden

\ 4

yan Wahi Nb

Fig. 7. Oral dise of a larva of MixopAves schevillr at stage

36, Seale bar = 1 mm.

‘Trenerry, M. (1988) The ecology of tadpoles in a

tropical rainforest stream. B.Sc. (Hons) Thesis, Dept of

Zoology, James Cook University of North Queensland,

Unpubl.

IADPOLES OF LIMNODYNASTINE LEPTODACTYLID FROGS 73

Some of the material measured was reared to

metamorphosis in the laboratory whilst the

remainder was collected and preserved in the

overwitlering condition at stage 25.

Tyler ([965) deseribed the oral dise of Mt, schevilli

as Litoriu nannotis (Liem 1974), He recorded six

upper and threé lower tooth rows, but only three

lateral rows of teeth,

Martin. (1967) reported six upper, three lower

lateral and three lower labial tooth raws with a

complete papillary border in M, fasciolatus whilst

Watson & Martin (1973) recorded six upper, live to

six lower lateral and three lower labial tooth rows

with a complete papillary border in Af. halbus. The

mouth of this species, illustrated by Watson &

Martin (1973, Fig. 4B), is very similar to that of AZ,

schevillt reported here. These authors also reported

development of the hind limbs in a membranous

sac in the early stages. This latter phenomenon

presiumably is a lotic adaptation shared with other

sympatric stream-dwelling species such as

Nyctimystes dayi and Literia nannatis (Davies &

Richards 1990), However, joule adaplahons in M.

schevilli are not as prominent as those of M,

Jasctolaius (Martin pers, com).

Neobatrachus wilsmmorei (Parker)

FIGS 8-9

A tadpole at stage 34 is illustrated in Fig. 8. The

body is elongately avoid and widest posterior to the

eyes. The snout is slightly truncated in dorsal view

and gently sloping in lateral view, The nares are

dorsal, sessile and poorly-separated. The relatively-

large eyes are dorsolateral. The spiracle, with an

uttached inner edge, is sinistral, ventrolateral and

not visible from above, The spiracular opening is

wide; the diameter of the spiracular tube decreases

slightly from its origin to its opening. The anal tube

is dextral, relatively long and opens about 34 of the

way down the ventral fin. The tall fins are arched

and rounded terminally, the dorsal fin. commencing

in the posterior 's of the body, being deepest

Vig. 8. a, Lateral aud b, dorsal views of a larva of Neohalrachus wilsmorei ac slaps 34. Scale bar = 1) mim.

4 M. DAVIES

a wee

vate r

With esas cower

ON Se a alee!

Pe cae Mi

PAP CHa Mminyn rina jess

Et, eine

Fig. 9. Oral disc af a larva of Neobatrachus wilsmorei

at sliage 34, Scale bar = | mim.

approximately 24 along its Jength. The ventral fin

is deepest abour halfway along its length. Tail

musculature is thick, narrowing to a Tine point

posteriorly.

The mouth is antereventral, Labial papillae

extend anterolaterally, laterally and posteriorly

around the mouth dise with an anterior median gup

(Fig. 9). There are three upper and three lower labial

tooth rows. The first upper and the second and third

lower rows Temain undivided. The horny beak is

robust.

Tail musculature is finely pigmented as is the

dorsal fin, The ventral fin is pigmented only in its

posterior third. The head and body are dusted with

fine pigment.

Keratio appears on the distal edge of the inner

metatarsal tubercle at stage 41.

Measurements of rhis tadpole are shown in Table

1. A metamorphosing tadpole at stage 45 had a

snout-vent length of 20.6 mm.

Neobatrachus centralis (Parker)

FIGS 1-11

A tadpole at stage 36 is illustrated in Fig, 10, The

body is ovoid and widest across its mid region. The

snout ts evenly rounded in dorsal view and lateral

view. The nares are dorsal, sessilc and poorty-

separated, Opening anterolaterally. The relatively

large eyes are dorsolateral. The spiracle is sinistral,

ventrolateral, not visible fram above and with an

attached inner edge, The wide orifice opens dorsally

and the spiracular tube increases in diameter very

Slightly from its origin to its opening, The anal tube

Fig. 10. a, Lateral and b, dorsat views of q tarva of Newhatruchus centralis at stage 36, Scale bar = 10 mm,

TADPOLES OF LIMNODYNASIINE LEPTODAC) ¥YLID FROGS a3

is median and relatively long. The tail fins are

arched and rounded terminally, the dorsal fin not

extending over the body and decpest approximately

halfway along its length. The ventraltin is deepest

approximately halfway along its length, Tail

musculature is moderately thick narrowing to

point posteriorly. The mouth is anternventral-

er ayy redige,

7” on Sek

Fig. ll, Oral dise of a larva of Nedbatrachus centralis

al stage 36. Seale bar = | mm.

Labial papillae extend anterolaterally, laterally and

posteriorly around the mouth dise, being

interrupted anteriorly. There are four upper und

three lower rows of Jabial teeth (Fig. 1). The first

upper and second and third lower rows are

undivided,

A dark pigment crescent is located around the

nares, The remainder of the body is dusted with

Pigment granules. A very faint dusting of pigment

granules covers the tail musculature and the dorsal

fin but is absent from the ventral fin.

Earlier stages of N. centralis have little or no

pigment. Keratin is formed on the distal edge of

the inner metatarsal jubercle at stage 41. A

metamorphosing individual at stage 45 had a snoutr-

vent length of 14.5 mm. Measurements of ™.

centralis tadpoles are shown in Table 1,

Neofatrachus kKunapatart Mahony & Roberts

FIGS (2-13

A tadpole at stage 36 is illustrated in Fig, 12, The

body is.ovoid and widest posterior to the eyes. The

snout is evenly rounded in dorsal view and gently

sloping in lateral view. The nares are dorsal,

sedentary and Opening anterolaterally. The

relatively-large eyes are dorsolateral. he spiracle

if sinistral, ventrolateral, not visible (rom ubove and

with an attached inner edge. {t opens dorsally and

the tube diameter decreases very slightly along its

Fig, 12 a, Lateral and bh, dorsal views ol a larva of Neodatrachas Avrtitpalurt al stage 36, Scale bar = 10 nine

76 M, DAVIES

length from the origin to the orifice, The anal tube

is long, dextral and opens about 5 of the way up

rhe ventral fin. ‘The tail Tings are poorly-arched and

rouniled terminally, The dorsal. fin does not extend

along the body and ts deepest about halfway along

Uslength, The ventral Gin is deepest.about halfway

along its length. Tail musculaiure is moderately

thick, narrowing, terminally,

The wiouth 1 anteroventral. Labial pupillac

extend anterolaterally, laterally and posteriorly

being interrupted anteromedially (Fig. 13). There

are Ihree upper and three lower rows of labial teeth,

The third upper and first lower rows are divided,

The horny beak is of moderate proportion

The head and boily are heavily dusted with

figment, as are pail fins and tail nisculature, A

Pie 13, Oral dise of a larva of Neobarrachas Aunapalari

al sliwee JG Seale bar = 1 mm,

circle of very clark pigment surrounds the nares. The

ventral surface is unpigmented.

Caomunent; More species of Neahatrarhus have been

described or redefined in recent times than of any

other limbodynastine genus (Tyler & Ledo 1973;

Rober{s 1978; Tyler ec e/., 1981; Mahony & Roberts

1986), For this reason, the data supplied by Martin

(1965) for N. pictus and by Watson & Martin (1973)

far N. ceatrafis may have been based on

misidentified maternal, Tyler (1989) reidentified te

N. pietus of Martin (1965) as N. swdelli and a

redescription of MN. centralis is provided here.

Neobatrachus tadpoles are known to grow lo #

very large size in (heir natural state: Martin (1947)

reported tadpoles of 73 mim teal length.

N. centralis and N. wilsmareé are laser to the

short body with a strongly arched tail recorded for

N, sudefli and N. pletis by Martin (1965) and

Warson & Martin (1973) than ts N, kunapatari

which has a more streamlined body. None of the

Species cxamined here has the spiracle free along

its inner edge as reported in N. svee//i by Martin

(1965)-

Acknowledgments

This study was supported by an Austrahan

Reseatch Council grant to the author and M. J,

Tyler, Leanne Seller and Rachel Norris assisted

with tadpole rearing whilst Garry Werren, Michael

Mahony and John Read provided tadpoles, Their

assistance 15 really appreciaied, Michael J. Tier,

Angus Marin and Graeme Watson provided

constructive criticism of che manuserip! which was

typed by Lorna Lucas.

References

Amsyis, M, ({976) Breeding biology and larval

developuien of Ligne verreque (Amira Hylidae),

Truns, Re Soc. S. ase WO, 103-202

Davis M. & Rietiarps, §. 1 (1900) Developrrenttl

biolowy of the Australian Hylid {rey Neyevinutey day

(Anuta: Myliduey Jia. pt, 207 21.

Gossek, Keb, (1960) A almplified table for staging

AnUraN anbryos and hirvwe with notes on ident estion,

Herpetolougica 16, 183-190,

Lee, A. K. (1967) Studies in. Australian amphihia U,

Tasonomy evelugy and evolutien of the vents

Holoropords Gray (Anum Leptodyetyliduct Aust.

Zoul, 15, 367-439.

Jem, DS. (1974) A review Of the fitoria Hannes

species group and 4 descuiption of a new species of

Liloria trom nerthern Queensland, Australia. Morr Old

Mus, (70), 151-168.

Manony, M. & Rogerts, | BD, (PYRG) Two new species

of desert burrowing frogs of the gens Neoburrachyes

(Amwrar Myobatrachidae) trom Wester Australia. Ree.

Hest Avi. Afey, 13, P85-170.

Mannn, A. A. (1905) ladpoles of the Melbourne area

bicrn Nal. B2¢4), 139-149

(1967) Australian anurin Jife histories: some

evolutionary and gcological aspects. fA, H, Wearherly

(Ed.) “Australian inland waters and their fauna"

(A.S.U., Canberra)

Roserts, J. D. (1978) Redefinitian of the Ausualiam

leptodietylid frog Neobatrichus pictus Peters. Trans.

R. Soe, 8. Aust, 102, 97-105.

Tyrer, M, J. (1965) Taxonomic studies of some bylid

frogs of Ansrralia and New Chunca, Prac, Lael, Soe,

London 51), 91-106,

(1989) “Australian Frogs”, (Viking O'Neil,

Melbourne),

= , Davies, Mo & MaArtIN, A.A, (981) New and

rediscovered frags front the Dyrby-Broume area Western

Australia. Rec, Wi lust Mus. 9, 147-172.

,& Lebo, W, W, (1973) The type locality ol the

leptodactylid. Troe Neabatrechus centratiy Parker. 5

Aust. Nai, 47, 75-76.

Watson, Gi T & Manrin, Ay A, (1973) Life history

larval morpholosy and relatianohiips of Austratian

teptodiactylid frogs. Truvts. R. Sve. S hush OT 13-48,

PERKINSUS (PROTOZOA: APICOMPLEXA) INFECTIONS IN ABALONE

FROM SOUTH AUSTRALIAN WATERS

BY P. J. O'DONOGHUE, P. H. PHILLIPS* & S. A. SHEPERDT

Summary

A total of 234 abalone were collected from nine sites in South Australian coastal waters and

examined for infections with the protozoan parasite Perkinsus sp. Infections were detected in ten

Haliotis laevigata from one location in Gulf St Vincent and in nine H. rubra from another location

in Spencer Gulf. All infections were characterized by the presence of macroscopic necrotic nodules

(0.5-8.0 mm in diameter) in the adductor muscles and mantle. Microscopic examination revealed

the nodules to contain variable numbers of host amoebocytes and numerous developmental stages

of the parasite, including single ovoid trophozoites (10.0-17.5 um) and larger rounded schizonts

(12.5-35.0 Um) containing vacuolated merozoites. The morphological and_ ultrastructural

characteristics of the parasites were similar to those previously described for P. olseni. A total of

240 Pacific oysters (Crassostrea gigas) were also examined from four commercial farms in

neighbouring coastal waters but no Perkinsus infections were detected.

KEY WORDS: Apicomplexa, Perkinsus, abalone, Haliotis, morphology.

Vransitciions of the Roval Suviery of S. Aust. (99. 1152), TI-B2

PERKINSUS (PROTOZOA: APICOMPLEXA) INFECTIONS IN ABALONE FROM

SOUTH AUSTRALIAN WATERS

by P. J. O"DONOGIIVE*, P. H. PHitLips* & S. A. SHEPHERIIT

Summary

O'DomocHue, P. J, PHowips, PH, & SHerHerp, 8. A, (1991) Perkinsus (Protozoa, Apicamplens)

infections in abalone from South Austrahan waters. Jrans, R: Svc, 8, Ausf. 115(2), 77-82, 31 May, 1991.

& total of 234 abalone were collected from nine sites in South Australiiin coastal waters and examined

for infections with ihe protozoan parasite Perkinsws sp. Mntcerions were detected in ten Haliatis daevivata

from one location in Gulf St Vincent add in nine A. rubra [rom another location in Spencer Cull, All

infections were characterized by the présence of mucroscupic necrotic nodules (0.5.8.0 iim tn diameter)

in the adductor muscles and mantle, Microscopic examination revealed the nodules to centinn variable

numbers of host amoehocytes ang numerous developmental stages of the parasite, including single ovoid

trophozottes (10,0175 pm) and larger rounded sehizonts (12.5-35.0 wm) contiining vacualated merosoites.

The morphological and ultrastructural characteristics of the parasites were similar to those previously

desetihed for PR alseni. A toral of 240 Pacific oysters (Crassestrea gigas) were also examined trom four

commercial farms in neighbouring coastal waters but no Perkinsus infections were detected.

Rey Worps: Anivomploxa, Perkiasus, abalone, Heliatis, morphology.

Introduction

Two abalone species are fished commercially

trom South Australian coastal walers; blackhp

abalone, @faliotis rubra Leach, and greenlip

abalone, H. Jaevigeta Donovan. Since 1972, licensed

divers have reported the occurrence of yellowish

pustules in the flesh of H. rubra collected near

Neptune Island in Spencer Ciulf, The pustules

render the flesh of the abalone unacceptable for

processing and marketing, Microscapic examination

revealed the pustules to be caused by a protozoan

parasite, Perkinsus olsenr Lester & Davis, 1981. Only

two other Perkinsus spp. have been described; P

marinus from the American oyster Crassostrea

wrefnica (Mackin ef af 1950; Perkins 1969) and 2

atlanticus from the clam Ruelifapes decussutus

(Azevedo 1989), In recent years, local divers have

become increasingly concerned with dwindling

stocks of H. /aeviyata along the western shore of

Gulf St Vincent. Perkirsus infections were detected

in H laevigata collected from reefs south of

Edithburgh (Lester 1986). The present investigation

with carried out to determine the geographic extent

of Perkinsus infections in greenlip and blacklip

abalone from South Australian coastal waters, and

whether Perkitsus infections occur in commercially-

farmed Pacific oysters (Crassostrea gigas) from

neighbouring waters.

* Central Velerinary Laboratories, Dept Agriculture,

Frome Road, Adelaide, S, Anst, 5000),

1 Dept Pisheries, 145 Picie Street, Adelaide, $. Aust. S000,

Materials and Methuds

Nine sampling sites were selected from the three

abalone fishery management zones of S.A. Pig. 1,

Table 1), Licensed divers collected approx. 30

abalone at random from each site between April

and October 1986, A Fisheries Officer also collected

approx. 60 oysters from cach of four commercial

oyster farms. The abalone and oysters were fixed

by immersion in Davidson’s fluid immediately after

collection and the species, sex and shell length were

Therny

Pansage |

Weslern done

| SAMPLES ait -4

t aes Gani haeuine

| @ Abainne -

_ Sotinein aim

| aoe

Fingrucd @&

Heel

3's 140E

Fig. |. Locations of sampling sites in South Ausiralian

coastal waters fram Which abalone and oysters were

collected.

18 P, |. O(DONOGHUE, P, H. PHILLIPS & S. A. SHEPHERD

recorded in the laboratory following shell removal.

The adductor muscles and mantle were examined

for macroscopic lesions on all superficial aspects

and internally through a series of longitudinal

incisions | cm apart. Suspicious lesions were excised

together with surrounding tssue. Small blocks of

mantle, adductor muscle and gonad tissue were also

sampled from cach abalone and oyster. Tissues were

embedded im paraffin wax, sectioned at 5 pm

thickness, stained with haematoxylin and eosin and

examined by light microscopy at 100-400 «

magnification. Tissue blocks found to contain

parasites were then processed for electron

microscopy by de-paraffinization in xylo]

containing 2% osmium tetroxide, clearing in

propylene oxide and embedding in epoxy resin

(TAAB Laboratories). Ultra-thin sections were cut

at 75 nim thickness, stained with 69% uranyl acctate

and 0.5% lead citrate and examined in a

transmission electron microscope (JEM 1() CX,

JEOL, Tokyo). Voucher specimens of fixed tissues

containing parasites.were deposited with the South

Australian Museum, Adelaide (SAM E2180 1).

Results

A total of 125 Haliotis daevigaia and 109 A, rubra

were collected from the ning sampling sites.

Perkinsus infections were detected in ten H.

laevigata (6 Go, 4 9 2) collected from Troubridge

Shoal in Gulf St Vincent and in nine H. rubra (6

ocr, 3 9 9) from Thorny Passage in Spencer Gulf

(Table 1). Infections were not restricted to any

particular size (henve age) group of abalone.

Infected /7. /aevigata ranged in size from 8.5-16.0

em in shell length and infected H. rufra from

10,0-16.5 em, No parasitic infections were detected

in any of (he 240 Pacific oysters. examined,

Infections in abalone were characterized by

macroscopic hemispherical blister-like nodules on

the superficial aspects of the adductor muscles and

mantle (Fig. 2), The nodules were soft and slightly

darker in appearance than the surrounding tissue.

Larger nodules were found io contain creamy

viscous fluid when incised. Ovoid nodules were

occasionally detected deeper within the tissues when

sectioned (Fig. 3). Intection levels ranged from 1-14

TABLE. 1. Prevalence of Perkinsus infections in abalone and avsters from South Australia.

Haliotis laevigata H_ rubra Crassostrea gigas

Lovafiten Depth (greenlip abalone) (blacklip abalone) (Pacific oyster)

(m) No. No- No, No. No. No.

examined infected examined intected examined = infected

Baird Bay 10 ‘2h Q 23 0 - -

(33°08'S, 134°16' EB)

Vhorny Passage § ps it} 16 9 - =

(34°SR'S, 136°04'L)

Hardwicke Bay 6 16 0 - - -

(34°50'S, (37922°F)

Poul Bay 20 8 i} t 0) - -

(35°13°S, 13715 'D)

Troubridge Shoal 3 30 10 - - - -

(33°08'S, 137-56'F)

Emu Bay 1S 24 0 \ 0

(3533'S, 137°34"B)

Capé Gantheaume i4 - 28 () - =

(36°07°S, 137930")

West Island 10 20 0 Is 0) -

(35'°37'S, 138°35' EB)

Ringwood Reef 6 - 24 0 S =

(37°38'S, 140°07"E)

Coffm Bay L - = - 31 tf]

(34°30°5, 13518 °D)

Bird Island ) - - S§2 0

(33°S9°S, 137¢33E)

Pori Price 1 ~ - - - 66 0

(34°15'S, 138"04'T)

Oyster Bay | - - 7 ()

(34°92'S, 17°48" B)

Total 125 10 109 9 240 0

(8.04%) (8.34%)

PERKINSUS INFECTIONS IN ABALONE

Pigs 2-5, 2. Nodules on surface of adductor muscle of Hations rubra. Scale bar = S$ mm. 3. Necrotic lesion in adductor

muscle of H. rubra. Scale bar = 5 mm. 4. Histological section through nodule in adductor muscle of H. /aevigata.

H&E. Scale bar = 0,5 mm, 5. Trophozoites (T) and schizonts (S) of Perkinsus within lesion in adductor muscle

of H. laevigata. H&E, Scale bar = 50 pm.

nodules per abalone and the nodules ranged in size

from 0.5-8.0 mm in diameter. Those detected in H..

/aevigata and H. rubra were similar in location, size,

shape and appearance. They were not encapsulated

but bound by normal tissues which sometimes

contained mild infiltrations of mononuclear

inflammatory cells (amoebocytes). The nodules

were necrotic and contained variable numbers of

amoebocytes together with other host cells, The

majority of cells appeared degenerative containing

pyknotic nuclei. Connective tissue fibres and

occasionally the remnants of muscle fibres were

found throughout the lesions forming a loose

supporting network, All lesions contained

fumerous clusters of extracellular basophilic bodies

which were identified as various developmental

stages of a protozoan parasite (Figs 4, 5), The

majority of parasite stages were vacuolated in

appearance but some were homogeneous and

stained uniformly throughout, Most stages

appeared degenerative and morphological integrity

was not well preserved within lesions, Nonetheless,

two types of parasite developmental stages were

evident by light and electron microscopy; unicellular

and multicellular forms (Fig. 6).

Unicellular stages (termed trophozoites) were

found scattered throughout the lesions as individual

cells sometimes grouped together in small clusters,

They were ovoid in shape measuring from 10,0-17.5

um in diameter (mean 15.0 nm) and were bounded

by a dense wall varying in thickness from 1,5-2.5

pm (Fig. 7). The trophozoites contained a single

nucleus with a prominent nucleolus and a highly

vacuolated cytoplasm usually containing a large

central yacuole measuring from 5-10 pm in

diameter. A small dense yacuoplast consisting of

eosinophilic granular material was occasionally

detected within the central vacuole,

Multicellular stages (termed schizonts) were

detected throughout the lesions in distinct clusters

(Fig. 8). They were larger in size than the

trophozoites measuring from 12.5-35.0 pm in

80 P. J. O’DONOGHUE, P. H. PHILLIPS & 8S. A. SHEPHERD

diameter (mean 27.5 wm). They were surrounded

by a dense wall (2.0-3.5 wm thick) with an irregular

outer margin. The schizonts contained 2-24

rounded cells (termed merozoites) ranging from

5-10 wm in diameter. Each merozoite contained a

highly vacuolated cytoplasm and a single nucleus

with a prominent nucleolus. Large central vacuoles

were not detected in the merozoites nor were

vacuoplasts. The majority of schizonts appeared

degenerate particularly towards the centre of the

lesion,

Discussion

Identical lesions and parasites detected in H.

laevigata and H. rubra suggests that both abalone

species were infected by the same parasite species.

This species is similar to P olseni previously

reported in blacklip abalone (Lester & Davis 1981).

All three parasite developmental stages

(trophozoites, schizonts and merozoites) were

similar in structure to those previously described

although some schizonts appeared larger (mean

Figs 6-8. 6. Electron micrograph of trophozoite (T) and schizonts (S) of Perkinsus in adductor muscle of Haliotis

laevigata, Scale bar = 5 wm. 7, Electron micrograph of Perkinsus trophozoite bounded by thick wall. Scale bar

= 5pm. 8. Electron micrograph of Perkinsus schizont bounded by thick wall (W) and containing several merozoites

(M). Scale bar = § pm.

PERNINSUS INFECTIONS IN ARMALONE a1

diameter of 275 um compared to 15,0 ym) and

more marure containing greater pumbers of

IMeTozgites. Ilowever, developing or immature

prezvosporangia were not detected and lesions were

not sutrounded by n loose wall of connective nssue.

Despite these dilferences, the morphological and

tltrastructural characteristics of the parasites were

considered to be consistent with those of 2 afseni

lester & Davis, (981.

Similar developmental stages have been described

previously for two other Pertinsus spp. Parasites

found in the American oyster €. virginica were

originally described as Dermacystidium marinurt

hy Mackin ez af. (1950) and luter as Labyrintionpa

imavinag by Machin & Ray (1966), Levine (1978)

subsequently renamed ihe species Perkinsns

murinvs and creeted the class Perkinsasida in che

phylum Apicumpleas on the basis of the electron

microscopic studies of Perkins (1976). This species

differs from F olsen’ in having much smaller

trophozoites (3-10 yun in diameter), membranous

rather than thickered walls and basaphilic rather

than cosinophilic vacuoplasts. More recently, thick-

walled Pertinsus-like trophozoites were found in the

sill filaments of the clan #. decussasius in Portugal

by Comps& Chagat (1987) and Chagot et ef, (1987).

These parasites were cultured in chioglycolate

medium cto form mature sporangia containing

biflagctlaied zoospores by Avevecio (989) and were

rramed F arfangicus on the basis of hose identity,

pathology and zoospore ullrastructure. The

dimensions, shaprund Hagellar organization of the

zoospures Were more regular than those of P

inarinns bil comparisons with # olsen could nol

be made because their Zoospore ullrastructute has

not yer heen delermined, Several other undescribed

Perkinsws sp. have been reporied in 57 species of

molluses [rom North America, the Mediterranean

and Australia (Andrews 1954; Ray 1954'; da Rov

& Canzonicr 1985; Goggin & Lester 1987) but

compinsons could nor be made because the only

developmental stages reported were large ovord ells

presumed to be prezanspoyaigia,

Early cross transmission studies suggested that

Perkingsus spp, may be specific for particular groups

of molluscs; P maritus for oysters (lamellibranchs)

and A asen] for abalone [gasinipods) {Ray 1954):

Lester & Dayis 981}, Flawever, revenc stiidics have

noc supperted any rigid host specificity for these

parasites, /* olseni \solated trom AL krevieafa was

juccessfully transmitted ta two laimellibranch

species (Pinctada Sugtlita and Anadara trapezi«)

and #erkinsus spp, isolated from five lamellibranchs

"Ray, SM. (1954) Biological studies of Mermuevstidium

Masten, at Vin@us. parasite of oysters. Rice Institute

Paiiphilet, Special Tssue, Linpubl,

(Anadara trapezia, Chama pucificuy, Tridavna

algas, T. erocea and T, wraxira) were suctesstully

transinitted to H, faeyivade (Goggin eral, 1989), 2

fRar ines has also been |ransmiltedl [rom the oyster

C, virginica to the pymmidellid gastropod Boonen

iapressa (White ev al. 1987). These results suggest

that Perkinsus Wlections may be transmitted

between different mollusc species inhabiting the

same walters No infections were detecied in oysters

sampled from neighbouring areas in this study bul

other molluse species remain 10 be examined.

Infected 4, rabra and FV. laevigeta were detected

at iwo different sites located 140 km upare in

adjacent Spencer Gulf and Gulf St Vincent.

Infections have previously been found in abalone

from the same general areas (Lester & Davis 1981:

Lester IYSA). The reasons for this patchy

Uistriburion of infvetions are nat known. The two

sites are separated by Vorke Peninsula but both are

situated neur the mouths of the Gulfs where the

same Ocean Current passes in an easterly direction,

However, no infections were detected in abalone

sampled from intermediate sites por front sites

locared further away in the same current Mow, There

are also no records of abalone stocks being moved

between the two sites of infection. These sites must

be regarded as potential point sources af infection

and local mollusc populations should be monitored

regularly for the spread of infections.

Significant mortalities of H. deevizata were first

reported along the western coast of Gulf Si Vincent

in 1980 and further deaths were reported eact

summer from [982-1985 (Lewis ef gf 1987),

Abalone had been abundant in this area As far Worttt

a5 Black Point but stocks have now practically

disappeared (K.L_ Branden pers. comin). Claiins

imade hy divers that mortaliues were due to

pollution were not substantiated by laboratory

investigations for heavy metals, oreanochlorines,

olganophusphates and hydtocarhons (Shepherd

1485), Subsequent studies revealed that many

abalone in this area were infected with & olsen

(Lester 1986) bur itis not known whether infections

caused the mortalities. The parasite is certainly

pathogenic antl causes necrotic lesions within hast

issues, Mortalities have been ubserved in

experimentally infected #7. rubra maintained in the

laboratory at 20°C wiiereas thase mainraiiied at

15°C recenered from infection (Lester 1986). The

continued detection of Perkirsivs infections in

abalone from dicback areas highlights (fe need for

further studies on parasite pathogentcity,

transmission and eunirni,

82 P. J, DONOGHUE, P. 11. PHILLIPS & S. A. SHEPHERD

Acknowledgments

‘The authors wish to thank the Director of

Fisheries, R. K. Lewis, for supporting the survey

with facilities and resources; A. Geering, J. Godden,

W. Guidera, J. Kroezen, J. McGovern, R. Spruyt

and A. Vermeulen for collecting the abalone; R.

Grove-Jones for collecting the oysters and B, Dixon

for conducting the electron microscopic studies. The

first author was also supported by a grant from the

Australian Biological Resources Study.

References

ANpreEws, J. D. (1954) Notes on the fungus parasites of

bivalve molluses in Chesapeake Bay. Proc. nail, Shellfish

Ass, 45, 157-163.

Azevepo, C. (1989) Fine structure of Perkinsus

atlanticus n. sp. (Apicomplexa, Perkinsea) parasite of

the clam Ruditapes decussatus from Portugal, J.

Parasitol, 75, 627-635.

CHacor, D., Comes, M., Bouto, V., RUANG, FL &

GeizeL, H. (1987) Histological study of a cellular

reaction in Ruditapes decussatus infected by a

protozoan. Aquaculture 67, 260-26).

Comes, M. & CHAGoT, D, (1987) Une parasitoye nouvelle

chez la Palourde Ruditapes decussatuy L. CR. Acad.

Se. Paris 304, 41.44,

Da Ros, L. & CANZONIER, W. J. (1985) Perkinsus, a

protistan threat to bivalve culture in the Mediterranean

basin. Bull, Kur. Ass. Fish Path, 5, 23-25.

Gocain, C. L. & Lester, R. J. G, (1987) Occurrence of

Perkinsus species (Protozoa, Apicomplexa) in bivalves

{rom the Great Barrier Reef. Dis. aquar. Org. 3, 113-117.

, SEWELL, K. B. & Lesrer, R. J. G. (1989) Crass-

infection experiments with Australian Perkinsus species,

Ihia. 7, 55-59,

Lester, R. 3. G. (1986) Abalone die-back caused by

protozoan infection? Aust. Fish. 45, 26-27.

_ & Davis, G, H. G. (1981) A new Perkinsus species

(Apicomplexa, Perkinsea) from the abalone Haliotis

ruber. J. avert. Pathol. 37, 381-187.

LEVINE, N. D. (L978) Perkinsus gen. n. and other new

a in the protozoan phylum Apicomplexa. J. Parasitol.

64, 549.

Louwis, R. K., SHEPHERD, S. A., O7DONOGHUF, P. J, &

Puivips, P. A, (1987) Protozoan parasite (Perkinsus)

infection in abalone: a progress report. SAF/SH 11, 7-8.

Mackin, J. G. & Ray, 8 M. (1966) The taxonomic

relationships of Dermoeystidiam marinum Mackin,

Owen and Collier, J, Jnvert. Pathol. 8, 544-545,

, Owen, H. M, & Coicier, A. (1950) Preliminary

note on the occurrence of a hew protistan parasite,

Dermocystidium marinum n, sp. in Crassostrea

virginica (Gmelin). Science 111, 328-329,

PERKINS, F, O. (1976) Zoospores of the oyster pathogen,

Dermocystidium marinum, |, Fine structure of the

conoid and other sporozoan-like orvanelles. J Parasitol.

62, 959-974,

SHEPHERD, 8. A. (1985) No evidence of contamination of

abalone stocks in western St Vincent Gulf. S4 FISH 9, 5.

Waite, M. E., Powrr., E. N. Ray, S. M. & WILSON,

E. A. (1987) Host-to-host transmission of Perkinsus

marinus in oyster (Crassostrea virginica) populations

by the ectoparasitic snail Boonea impressa

(Pyramidellidae). J. Shellfish Res, 6, 1-8.

DESCRIPTION OF THE MALE OF TYLENCHORHYNCAUS TOBARI

SAUER & ANNELLS, 1981 AND OBSERVATIONS ON THE

MORPHOLOGY AND HOST RANGE OF THE FEMALE IN ARID SOUTH

AUSTRALIA

BY J. M. NOBBS*

Summary

In a survey of the arid region of South Australia, over 300 sites were found to have

Tyienchorhynchus tobari Sauer & Annells, 1981. Previously undescribed males of T. tobari were

identified from only nine sites and are described here. From field observations, plant species of the

family Chenopodiaceae were most likely to have 7. tobari present. This was tested by culturing the

nematode on different host plants in the glass-house. It was found that environment affected the

morphometrics of different field populations of T tobari but not general morphology.

KEY WORDS: Tyienchorhynchus tobari, arid South Australia, males, host plant, Nematoda

Transactions ef (he Royal Society af S Aust (1991), 2S), BIRR.

DESCRIPTION OF THE MALE OF TYLENCHORHYNCHUS TOBARI SAUER &

ANNELLS, 1981] AND OBSERVATIONS ON THE MORPHOLOGY AND HOST RANGE

OF THE FEMALE IN ARID SOUTH AUSTRALIA

by J. AL Nowsrs*

Summary

Neuss, 1 M. (1991) Description of the male of Te/entchorhynchus fobori Sauer & Annetls, 198) and

observations of the morphology and hast runge of the female in arid South Ausiralia. Trams, R. Sec. 3.

Aust, 115(2), 44-88, 31 May, 199],

[na survey of the arid region of South Ausintlia, over 300 sites were found lo have 7plencharkvechua

sebari Sauer & Annells, WSL. Previously undescribed males of 7 cabaret were idenntied from only fine

sues and are described here, Prom field observations, plant species of the family thenapodiageae were

most fikely to have 7 tobari present, This was tested by cultiiring the nematode on different host plants

in the glass-house. Lt was found that environment a/feeted the morphometrics of different field populations

of 7) tehari but not general morphology.

Kiy Wokoe Tifenchorhynckus debari, arid South Australia, males, lost plant, Nematoda

Introduction

The and region of South Australia consists of

diverse vegetalion and landfarms, There is little

information on the occurrence and diversity of rhe

plant parasitic nematode fauna within this region.

During a survey of the area (Nobbs 1989), one of

the most widely distributed plant parasitic

nemalodes was Telencharhynchus tabari Sauer &

Annetls, L98l, The wide distribution of rhe

nematode over a range of environments. offered the

opportunity to examine the effect of environmental

variation on the nematode. This paper examines the

effects of environment on férnale morphometrics

and possible hosts among the diverse plant species

sampled, Males are described for the first rime.

Methods

Extraction of nematodes

Soil was collected from undisturbed native.

vegetation which occurred close to the main tracks

that cun throughout the arid region, Over 300 sites

were sampled and the sampled plant species noted.

The nematodes were extracted from $0 tm) of ¢ach

soil sample using a modilied Baenpana funnel

(Schindler 1961),

Morphology and measurements af Tylenchorhyn-

chus iobari

‘To examine the effect of different environments

On variation in morphometrics, ten sites were

selecied from different areas, From each site, ten

females were processed through an alcohol series

and mounted in eiycerol by the wax method

= CABI Institute of Parasitology. 395A Hartfield Rd. St

Albans, England, AL4 OU.

(Hooper 1986), Measurements (in mm) of body

length, body width, oesophageal length, position

of the vulya, tail length, tail width and stylet length

were them made under high magnification and the

de Man rarios (a, b, ¢ and c’) were calculated.

Analysis af variance (ANOVA) was used to

determing if there were significant ditferences in

Measprements between the ten different

populations,

OQvvurrence in the field and in pots

To determine the most likely host plant of T

tobari the number of sites on which a particular

plant species occurred was sampled and compared

with the actual (or observed) number of sites where

that particular plant was sampled and found to have

T. fobari present. The number of sites where a

particular host plant was sampled was used as a

percentage of Lhe total sites sampled (expected sites),

Using Chi-square analysis (Bailey 1976) the

observed number of sites was then compared with

the expected number of’ sites. to determine most

likely host species. Due to the diversity of the

veretation sites, grouping of the host species was

necessary (e.g. Chenopods = plant species of the

family Chenopodiaceae),

This information allowed investigation of

possible hosts of T fobari, Sceds of native and

introduced spevies including Atriplex spongiosa, A.

lindleyi, Chenopodium quinoa, Lycopersicum

esculentum and Hordewn vulgare (ev. Clipper) were

surface sterilised (3 mio, in 1% bleach),

pregerminuted ina Petri-dish, planted into a 1:4

parts soil to sand mix and inoculated with 50 female

T. tohari. After two and a half months, the shoots

were. removed und the roots and soil washed Lhrough

asetof sieves (500 um, 250 pm and 40 pm aperture).

hd IM, NOBBS

The sediment on the 250 pm and 40 ym sieves was.

collected and placed in a modified Buerman’s

funnel for three days. The nematode extract was

then counted for 7 tebert There were three

replicates from each plant species.

Results

Morphometrics of male and female

Tylenchorbynehus tobari i rhe arid region af South

Ausiratict

Males of 7. todmrd were identified from nine

different sites within the arid region of South

Australia (Fig. 1) andi mean values + standard

deviarions of morphometric measurements for all

sites (n—20 specimens) are presented below. In

addition, the same data for a single site (n=9) near

Kingoonya (grid reference 299 |80, map

KINGOONYA SHS3 11 (1: 250,000) edition 1,

series RSQ2, Royal Australian Survey Corps) are

provided, The original measurements of Sauer &

Annells (1981) for temales as well as the grad

means of the LO sites selected are also presenti.

fiemales: original descriprion (Sauer & Annellx 1981

(n 19); Body length — 690 nim (610 - 770).a = 34 (0

~4k) b = SOUS -F2ye = IDO lye’ - DBL

\ “y “ti

i l

\ \, : =

te=,p Oa |

‘4 ” |

| tye!

ee ee | tie 7

>. . ‘ 3 ;

7 “i rey

\ ) of

ae IN

\y ‘a |

“" !

ude \, |

Fin. b. The distribution of fitedchorhvaches /oburt Sane

& Annells, S98) wairhin the arid region of South

Ausitrahin, Closed rales ace sites From which f. reAuri

wasidenniticd: open vires are sires ar which nudes were

idenriticd Sites | tO were sires trom wiiel ten Lemualis

were Mmeastered,

~ 4.4), V = 54 (5) - 54) styler = 17 - 19 um. Survey 1983

1985 (i » 10D): Bolly length = 724 ~ 62 am (595 -

YOO), a - 30.3 +4 FLSA -d256 S82 = 05 O40

~7.6),¢ > 14.0 + 3.0 006-25. 6° — 3.0 = OR UL7

4.3), V¥ — 34.4 © 22 (49- 59) stylet = 17.4 5 14m

th4 - 2).

Males (Fig, 2) (n=20): Body length - 672 1 18 ym (S86

~ 792), a = 39.9 4 LS (25.8 BRI 952 ) O2 eho

SO WO8 + 0.6 (8.8 - IRM = 3M 4 OF EY

47), spicule lengih - 26.5 = 1.3 pm (ry - 30)

pubginacdlum = 113 2 2bp~m (8 17); stylet length =

16.7 = 0.7 pm (14 - 20).

Site near Kingoonya (n - 9): Bady length © 676 + 26

piu (619-727), 4 = 29.9 + 0.9 (258-324) b - 5.2

1 U2 (49 -5.8)¢ 9 U7 = 04004 L220 47 =

0.2 (2.9- 4.3) spiculelength = 25.1 + 11 pum (22 28)

gubernaculum = 12 + (3pm (8 - 17); stylet length |

17.0 + 0.7 im 4 - 18).

Description af the male

(Fig. 2) Similac to female in anterior region. Lip

region offset, 6 - 8 aunnules, stylet of mediuns

development, with backwardly sloping stvlet knobs.

‘lesris single, not reflexed. Tail enveloped by a larue,

simple, crenate bursa. Spicules distally flanged,

terminus narrow, guberauculum well developed,

generally rod-like, protruding. Phasmid easily seen,

just anterior to mid-point of tail.

Occurrence in the field und in pois

Chi-square analysis showed that 7) lebari was

jound in sienificantly mare sites than expected only

where plant species of the family Chenopodiaceae

were the most curmmon species (Table {), Therefore,

the most likely preferred host plant is a member of

the family Chenopodigcede. With the pot tests there

Was some multiplication of TL tahart with all the

hve plant species tested, but Aarriplex spongiosa

had the preutest multiplication rate (lable 2).

dnalysis of papulations

Although only a small number of females per

populalion were measured, significant differences

in morphometries were observed. OF the churucters

measured only position of the vulva (V), de Man

fulfo's a. by and c’ were not significantly different

between populations (Table 4), Body length, body

Width, tail length, tail width, ne¢xophageal length,

stylet length and de Mun oc ratio were all

significantly different between naturally occurring

populations.

Io One population (Y), almost all ofthe observed

values were ereater than the standard deviation of

the grand mean. Few of the other popalalons had

any or more than one value beyond the range of

plus or minus the standard deviation. ‘There were

no obvious. differences in general morphology

between specimens collected from the ten sites, so

the differences in measurements between the

Populations-are mest likely due to environmental

elects such as recent rainfall, host species present

and soil (ype rather than species differences.

TYLENCHORHYNCHUS TOBARI IN ARID SOUTH AUSTRALIA 85

Fig. 2. Morphology of the male of Tylenchorhynchus tobari Sauer & Annells, 1981. A = oesophageal region; B

= head region; C ~ shape of tail (internal); D — shape of tail (external). Scale in microns (um).

86 J. M. NOBBS

TABLE 1. The host plant/eroups and number of sites where Tylenchorhynchus tobari Sawer & Annells, 1981 was

collected.

Species/groups Number of Sites Chi-square % Total Sites

Ohserved Expected value Sampled in

Survey 1983-85

Chenopods 140 108.5 914 44 33,5

Ephemerals 14 14.6 0.02 4.5

Eucalyptus spp. 27 36.6 2.52 W3

Acacia spp. 71 76.8 0.44 23.7

Grasses 9 15.6 2.79 4.8

Shrubs: (Eremophila, Dodonea, Cassia sp.) 16 25.6 3.60 7.9

Trees: (Myoporum, Pittosporum, Cailitris sp.) 24 18.8 1.44 48

Salicornia spp. 8 7.8 0.01 24

Reeds ) 2.9 2.90 0.9

Zvgocloea paradoxa 15 16.8 0.19 $2

Total 324 324.0 23.05 100.0

** _ significantly different, df = 9, P = 0.01, Chi-square analysis.

#4 = significantly different, df 1, P = 0.01, Chi-square analysis.

The null hypothesis that there is no difference between the expected numbers of sites from which certain plant

species/groups were sampled and the presence of 7ylenchorhynchus tobari in the soil sample is rejected.

The % total sites indicate the number of samples from which soil was sampled in the period 1983 to 1985 and were

used to calculate expected number of sites with T. tobari.

TABLE 2. Pinal number and multiplication rate of Tylenchorhynchus tobari from an initial inoculation of fifty females

and sampled after two and a half months. (mean + standard deviation).

Plant species Mean number Multiplication rate

Atriplex lindleyi 212.7 4.2

+55,9 $1.12

A. spongiosa 1238.3 24,8

4224.6 14.50

Hordeum vulgare 56.0 1.1

{var. Clipper) =/I7.4 +035

Lycopersicum 209.7 4.2

esculentum +297 + 0.96

Chenopodium quinoa 499.0 10.0

TYLENCHORHYNCHUS TOBARI IN ARID SOUTH AUSTRALIA a7

TABLE 3, Measurements of different populations af Tylenchorhynchus tobari frard fhe arid région af South Australia,

Population Body Body Tail Tail Length of Length © Talo

fength width length width oesophagus stylet

1 699.3 23.6 52.2 17.6 130.0- 17.0 13.9

2 699.5 23.6 56.1 17.6 135.7 17,2 12.7

3 716.9 23,3 49.4 16.9 145.2 16.8 14.8

4 725.6 23.4 47.4- 17.3 (46,27 1R.7" 15.4

5 TOLD 24.0 55.5 18.8 > 133.9 18.0 13.5

6 724.2 24.1 54,5 18.6 137.0 16.2- 13.7

7 # 734.0 23,8 $8.27 18.3 133.3 174 12.7

8 # 7OL.6 23.8 56.8 18.7 130.6 16.2 12.6

9 795.9 ' 27,2" 49.6 18,3 152.1 ° 18.3° 16:6!

1 N34 22,8 49,9 16.87 140.7 17.6 14,3

Grand Mean 720.7 239 52.9 17.8 138.5 17.3 14.0

+ 3D, 61.6 24 ad 18 ILS Lda 3:0

F-value 2.40 2.79 2.28' 2.06 3.57 441 242

+ ++ +4 ‘*e es ere ¥¥

Significant at P = 0.01% level indicated by **; significamt at P = 0.001% level indicated by ***; df. = 9, 86.

Grand mean is calculated from all 100 nematodes measured and includes the standard deviation /S.D.) in italics,

# = populations where males were identified.

indicates value less than lowest value of the standard deviation of the grand mean. | indicates value greater than

highest value of the standard deviation of the grand mean.

Measurements are in microns (gm).

Discussion

Males of Ty/enchorhynchus tobari were found in

only a small number of sites and in low numbers

indicating that 72° tnbori may reproduce

parthenogenetically. Populations of 7) fobari trom

different natural habitats differ significantly in

certain, morphometric characters, However, the

description of a new species is nol necessary as the

populations are still identifiable morphologically

as T. tobart. Many workers (Davide 1980; Fortuner

1984a; Fortuner & Queneherve 1980; Kline 1976;

Roggen & Asselberg 1971; Townsend & Blakith

1975; Saha & Khan 1988; Singh ef @/, 1985) have

looked at the influence of host on morphometrics

of different species of nematode, They found that

many characters were highly variable between

populations and that ratios were of little overall

value (except V) in determining species. Fortuner

(1984b) suggested Ihat observations of several

populations were important in estimating the mean

and range of measurements. When identifying

species, morphology should always be used with

priority aver morphometrics as differences in

measurements can often be attributed tu

environmental effects.

T. tobari is a migratory ectoparasite and so has

a wide host range. In the field the most common

planis sampled with 7 tobari present were of the

family Chenopodiaceae. In pot cultures Atriplex

spongiosa allowed the greatest multiplication. In

using a hast plant that allows rapid multiplication

of 7. tobari, the host/parasite relationship can be

investigated.

Acknowledgments

I wish to thank Prof. H, R, Wallace and Dr

J. M. Fisher for their advice during the survey and

Dr D. J. Hunt for reading and commenting on the

manuscript, T would also like ta thank the

Australian Bureau of Fauna and Flora for the travel

grant to conduct the field trials in the arid region

of South Australia,

88 J. M, NOBBS

References

Bary, N. T. J. (1976) “Statistical methods in Biology”.

(Hodder & Stoughton).

Davinu, R. G. (1980) Influence of different crops on the

dimensions of Meloidogyne arenaria isolated from fig.

Proc, Helm. Soc. Wash, 47, 80-84.

FortuNer, R. (1984a) Morphometrical variability in

Helicotylenchus Steiner, 1945 5: On the validity of

ratios. Rev. de Nentatol, 7, 137-146.

(1984b) Statistics in taxonomic descriptions,

Nemutologica 30, 187-192.

& QuENEHERVE, P. (1980) Morphometrical

variability in Melicotylenchus Steiner, 1945 2: Influence

of the host on H. dikystera (Cobb, 1893) Sher, 1961,

Rev. de Nemaiol. 3, 291-296.

Hooper, D. J. (1986) Handling, fixing, staining and

mounting nematodes, pp 59-80. Jn Southey (Ed)

“Laboratory methods for work with plant and soil

nematodes”. (H.M. Stationary Office, London,

England).

Kiine, J. P. (1976) Morphometric variation in

Aphelenchus avenae with varied nutrition and time.

Nematologica 22, 94-102.

Noess, J. M. (1989) The occurrence of plant parasitic

nematodes in the arid region of South Australia. Trans.

R. Soe. §. Aust, 113, 117.

RoaccEN, D. R. & ASSELBERG, R. (1971) The use of ratios

in nematology. Nematologica, 17, 187-189.

SAHA, M. & Kuan, BE. (1988) Effect of host on the

morphometrics of Pratylenchus zeae Graham, 1951.

Indian J. Nematol. 18, 55-60.

SAUER, M, R, & ANNELLS, C. M. (1981) Three new

Tylenchs (Nematoda) from Australia. Nematologica. 27,

422-43}.

SCHINDLER, A. F. (1961) A simple substitute for a

Baermann funnel. Plant Dis. Rep. 45, 747-748.

SINGH, V., SINGH, S. P., Yabav, R. & SAXENA, S, K.

(1985) Effect of different plants on the morphometrics

of females of the root-knot nematode Meloidogyne

incognita. Nematol. Medit. 13, 81-85.

TOWNSEND, J, L, & BLAkITH, R. E. (1975) Fungal diet

and the morphometric relationships in Aphelenchus

avenae. Nematol. 21, 19-25.

THE MANGALO METEORITE, A NEW (L6) OLIVINE-HYPERSTHENE

CHONDRITE FROM SOUTH AUSTRALIA

BY MARGARET WALLACE & ALLAN PRING*

Summary

The Mangalo meteorite is a single stone, which originally weighed 1050 g, and was found near

Mangalo (33°34'S 136°39'E) South Australia in 1987. It has been classified as an L6 chondrite,

shock facies ‘e', and contains olivine (Fazs52264) orthopyroxene (Fsz1 4-223) clinopyroxene

(Wo44.7eN46.8fSg.5), apatite, nickel-iron, troilite and maskelynite. Mineral textures and compositions

indicate that Mangalo was a metamorphosed part of the L-planetoid and was heavily shocked before

reaching Earth.

KEY WORDS: Mangalo, chondrite, meteorite

Transactions of the Royal Society of S. Aust. (1991), 115(2), 89-93.

THE MANGALO METEORITE, A NEW (L6) OLIVINE-HYPERSTHENE CHONDRITE

FROM SOUTH AUSTRALIA

by MARGARET WALLACE & ALLAN PRING*

Summary

WALLACE, M. & PRING, A. (1991) The Mangalo Meteorite, a new (L6) olivine-hypersthene chondrite from

South Australia. Trans. R. Soc. S. Aust. 115(2), 89-93, 31 May, 1991.

The Mangalo meteorite is a single stone, which originally weighed 1050 g, and was found near Mangalo

(33°34’S, 136°39’E), South Australia in 1987, It has been classified as an L6 chondrite, shock facies ‘e’,

and contains olivine (Fas 2-264), orthopyroxene (Fs) 4-22.g), clinopyroxene (Wog4.7Eng¢,gFsg.5), apatite,

nickel-iron, troilite and maskelynite. Mineral textures and compositions indicate that Mangalo was a

metamorphosed part of the L-planetoid and was heavily shocked before reaching Earth.

Key Worps: Mangalo, chondrite, meteorite

Introduction

The Mangalo meteorite was found by Mr Neil

Smith while ploughing on his property at Mangalo

near Cowell on the Eyre Peninsula, South Australia

(33°34’S, 136°39’E) in June 1987. Mr Smith

brought the stone to the attention of two geologists

from the South Australian Department of Mines

and Energy, who in turn sent it to AMDEL in

Adelaide where it was positively identified as a

chondritic meteorite. After identification Mr Smith

brought the meteorite to the South Australian

Museum for detailed examination. No additional

material has yet been recovered. In accordance with

the guidelines on meteorite nomenclature, the

meteorite has been named Mangalo, after the

geographical locality closest to its site of discovery.

Mangalo is the forty-fifth meteorite to be found

in South Australia and the ninth to be recovered

from Eyre Peninsula (see Fig. 1). It does not appear

to be related to any of the Eyre Peninsula meteorites

(Corbett 1968; Fitzgerald 1979'). Mangalo was

found quite close to the site where the Cowell

meteorite was found in 1932. The Cowell meteorite

is now considered to be part of the Kyancutta fall

(Graham ef al. 1985). It is however an iron and not

a stony meteorite (Spencer 1933; Buchwald 1975).

South Australia, like Western Australia and the

south-west of the United States is a particularly

good area for finding meteorites due to its arid

conditions and the great age of the land surface

(Bevan & Binns 1986, 1989),

* Dept of Mineralogy, South Australian Museum, North

Terrace, Adelaide, S. Aust., 5000.

! FITZGERALD, M. J. (1979) The Chemistry and

Mineralogy of the Meteorites of South Australia and

Adjacent Regions. Ph.D. thesis, University of Adelaide.

Unpubl,

The Mangalo meteorite and the Streaky Bay

meteorite (Wallace & Pring 1991) are the first new

meteorites to be recovered since legislation was

enacted in 1980 to protect meteorites found in South

Australia. This legislation, in the form of an

amendment to the Museum Act, made all

meteorites found in South Australia the property

of the Museum. Provisions were included in the Act

for rewarding the finders of meteorites. In the case

of the Mangalo meteorite, the finder, Mr Smith has

been presented with a polished piece of the

meteorite and a bronze medallion commemorating

the meteorite’s discovery.

Physical Description

The meteorite is a five sided sub-rounded stone

(11 x 8 x 8cm) weighing 1050 grams (Fig. 2). The

stone is covered in all but one corner with a 1 to

AS ; °

Kaldoonera Hill @ — Yardea « Buckleboo

ss ° * Kyancutt

Streaky Bay Cocunda eure

Cy * M

e u

Kappakoola angalo

0 50 100 km

Fig. 1. Map of Eyre Peninsula showing the location of

Mangalo and the sites of other meteorite finds on the

Peninsula.

90 M. WALLACE & A, PRING

Fig. 2. View of polished slab of the Mangalo meteorite.

2 mm deep well-preserved shiny brownish-black

fusion crust, portions of which are gritty with sand

grains which may have become attached during

Weathering, The external appearance of the stone

is typical of a chondrilic meteorite which has been

exposed to the weather for a number of years. The

imerior of the meteorite is dark green-grey in colour

and fine to medium grained, Chondrules (quenched

spherical groups of olivine, orthapyroxene and

glass) are readily visible especially on polished

surfaces. Metals, both kamacite and taenite, occur

as finely disseminated grains and stringers, many

of which rim chondrules. Silicate minerals are

stained brown, probably by lerrestrial weathering,

and the sample contains several iron oxide filled

fractures, The meteorite has been cut and two of

the internal surfaces polished, Two polished thin-

sections have been made. These were used both in

the petrographic examination and in electron

microprobe analyses.

Mineralogy

Compositions of the silicate phases were analyzed

with a JEOL electron microprobe ai the University

of Adelaide. Analyses were made using an

accelerating voltage of 15 kV, a sample current of

5 nA, and a beam width of 5 pm, Representative

crystal analyses are presented in Table 1,

The meteorite is composed predominantly of

olivine and orthopyroxene with minor amounts of

clinopyroxene and maskelynite. Microprobe

analyses show that the olivine in Mangalo is

equilibrated with a mean fayalite content of Fa,,,

(10 analyses, range 25,2 to 26.4). The orthopyroxene

shows a small variation in chemical composition

with a mean ferrosilite content of Fs... (10

analyses, range 21.4 ta 22.8) and wollastonite

content which varies from 1.2 to 1.7 mol!” (average

= 1,3). The orthopyroxene has a particularly high

calcium content which suggests that it equilibrated

at high temperatures. Clinopyroxene, which is only

abundant and coarse enough for reliable analyses

in type 6 chondrites, is homogeneous with an

average composition of Wo,,,Eny, Fs, ..

Plagioclase crystals normally found in equilibrated

chondrites are not present in Mangalo; they have

been transformed to the shock produced glass,

maskelynite. Accessory minerals include l(roiliie,

iron-nickel metal and chlorapatile.

Chemical Group

Mangalo is an ordinary chondrite, an

agglomerate meteorile. This type of meteorite is

considered to be compositionally similar to the bulk

solar system, less (he volatile hydrogen and helium

components (Keil 1969), The ordinary chondritic

MANGALO METEORITE

TABLE 1. Representative electron microprobe analyses (wt. %) of selected minerals in the Mangalo meteorile

Olivine Orthopyroxene Clinopyroxene Maskelynite Chlorapatite

6 6 6

P205 - = - — 39,7]

SiO2 37.13 54,75 53,78 66.79 =

TiO2 = = 0,32 - -

Al203 - 0.16 0.63 21.03 -

Cr203 - - 0.87 - -

FeO 23.22 14,27 §.30 0.69 2,02

MnO 0.40 0,42 - - =

MeO 38,26 28.66 16.26 - -

CaO = 0.69 21,60 2.06 50.72

Na20 = = 0.65 5.49 0,28

K20 - - - inet -

cl - - = = 6.94

Total 99.00 98.94 99.43 97.18 99.68

P - = = = 0.958

Si 0,9827 1.9801 1.9838 = -

Ti - = 0.0090 - =

Al - 0.0066 0.0276 - -

Cr = = 0.0254 - -

Fe 0.5140 0.4328 0.1636 - 0.0482

Mn 0.0090 0.0130 - - -

Mg 1.5091 1.5498 0.8941 - -

Ca - 0.0270 0.8538 - 1.5488

Na ~_ - 0.0467 - 0.0157

K = = - -

c - - - - 0.3330

Total 3.0149 4.0092 4,0041 - 2.9058

Pe# 25.4 21.8 15.5 - -

- Ca=13 Ca=44,7 Ca=15.5 -

- Me=77.1 Mg=46.8 Na=74.6 —

- Fe=21.5 Fe=8.6 K=9.9 =

meteorites are divided on the basis of the iron

content of olivine and orthopyroxene into three 25 - Eyre Peninsula

chemical groups (H, L, and LL). All the meteorites

in any group are believed to have come from the

same planetoid from within the asteroid belt (Keil

& Fredriksson 1964). The iron content of the olivine

and orthopyroxene in Mangalo indicate that it

belonged to the L-group of ordinary chondrites (see

Fig. 3).

Petrologic Type

Ordinary chondrites of the same chemical group

differ widely in texture and physical history. After

the accretion of the planetoid, its components were

heated and metamorphosed: chondrules

recryslallized, silicate minerals became homogenous

and show a narrower variation in composition, and

Slony Meteorntes

Th, Gronp

r L Group

. .

Mangal (6) 7 ™ Suneaky Hay Ud)

L _

Couuiida yf A)

PW Groupe

Kaliannera HULA) Kayipakoola (6)

Fe mol% in Gribopyrosene (Fs content)

_— Kielpa (HR

[Buckleboo 1Hé:

15 ee ee ee ee ee

15 20 25 30

él

Peale Olvine (Pa uitient

Fig, 3. Iron vontents, expressed as mola Fe in olivine (Fa)

and orthopyroxene (Fs) of Eyre Peninsula slony

meteorites. Classification groups are fram Keil &

Fredriksson (1964), Data are from Mason (1974); and

Wallace & Pring (1991).

92 M. WALLACE & A, PRING

Fig. do Thin-seetion microphotograph af

meteoric showing a radial psrosece chondrule (R) and

# porphyrine olivine chrandrule (P) in a reorystallized

MATES

plasoclase leldspar erystallized out of the elassy

areas, The Mangile meteorite is a highly

equilibrated chondrite and belongs to petrologic

type 6 of a six increment scale ol the Van Schmus

& Wood (1967) classification. In thin-seetion, the

chonrules are easily recognizable, up to. 6 mm in

diameter, but are weli-tregrystallized and show

poorly delineated’ boundaries (Fig. 4). The matrix

has been reerystallived and now consists of olivine,

hyroxenes, and miaskelynite (a shock produced plass

af plagioglase composition). Silicate mineral

chemistry 7s homogeneous. Temperature

calculations using the Wells (1977) and Lindsley

(1983) geothermomelers. based on the calcium

content of co-existing pyroxenes indicate that the

mictcorii¢ has been heated tu between 700 and

ROO” within its parent body (Pig, 5),

Shock Effects

Meteofiles offen record shock features produced

ly high velocity collisions in space. Shock effects

1 qvett ‘ eee

bwin ¢

LEC Ro BD

)) eH Eure eee

tee f

; jet Soo Here dite

Vert Aly a ay ; i) IN

Hastie wn Hoon

rk ears

li, 4, Siheate mineral cheaviseey in Maniyale mutearite,

Isarberms show temoerniure calculaiowy based ag ihe

calcium vontent of orihopyraxenes and chinopyrevenes

(Lindsley 1%85).

range from fracturing apd brecelation to le

formation of high pressure ninerals and melting,

Shock related fractuniig is pervasive throngl

Manaulo; the olivine grains show muosuic extinction,

and plagioclase has heen cotiverted inte

maskelynite, Mesaic extinction of ulivine is

characteristic of meteorites which have been

shocked to pressures of 150 10 400 kilobars and the

conversion of plagioclase to maskclynite supgests

shock presstires of about 300 kilobars (Van Schmus

& Ribbe 1968; Staftler ef ai 1988). Mangalo as

therefore & severely shovked meteorite which. tas

suffered from some lorm of high velocity collision.

Classification

The average compositions of the ferro-maunesian

silicates, olivine (Pa,,,) and orthopyroxene

(Fs,,,Wo,,), show that Mangalo belongs to the L

group of ordinary chondrites (Kell & Fredrikssen

1964). The microstructure of Munvala, which shows

recognizable chondrules with inclstiner rims in x

crystalline Taatrix, indicates that the meteorite

belongs to a high metamorphic grade, petrologic

type 6 (Van Schmus & Wood 1967), The wollastonite

content of orthopyroxene tates from 1,2 10 1.7

wiM, similar to the range found in most L6

chondrites (Wo, .-Wo),, Sort ef af 1986), There

have been major alterations to the mineralogy of

the meteorite due to preferrestrial shock, Sitcale

grains are fractured, olivine crystals show masaic

extinction under cross-polarized light, and

Magioclase has heen converted to maskelynite. ‘This

is consisrent with a classification of shock facies

“e' (Dodd & Jarasewich 1979).

These observations indicate that Mangala tormed

deep wethin the L-group parent body, which is

believed to have been part of the asteroid belt. 1

was metamorphosed and reerystallized at

Lempctratures berween 7(Khand SPC (1 to 2 kilobars

of pressure) probably within the L-group paren|

hody and was heavily shocked before reaching

Eartit. The Mangalu meteorite is classified as an

Lée chondrite

Acknowledgments

We thank Mr Smith for bringing ihe metearie

ro-our atrention at ihe Sourh Australian Museum.

Gerr Horr is thanked for making thin-sections and

Jan Forrest for photography, The University af

Adelaide is acknowledged for providing access to

them clectron microprobe faciliues housed in its

bleciron Optical Centre, Phe project was landed

by the Mark Mitchell Foondauorne:

MANGALO METEORITE 93

References

BEvAN, A, W. R. & Binns, R, A, (1986) A preliminary

sorting out of new meteorite recoveries from the

Nullarbor Plain, Western Australia. Meleoritics 21,

335-336,

&_ ss (1989) Meteorites from the Nullarbor

Region, Western Australia [: A review of past recoveries

and a procedure for naming new finds. bid. 24,

127-133.

BUCHWALD, V., F, (1975) “Handbook of Iron Meteorites”.

(University of California Press, Berkely, California.)

Coreett, D, W. P. (1968) Catalogue of meteorites in the

South Australian Museum (excluding tektites). Rec, 8.

Aust. Mus, 15, 767-790,

Doon, R, T, & Jarosewicn, E. (1979) Incipient melting

in and shock metamorphism of L-group chondrites.

Earth Planet. Sci. Lett. 44, 335-340.

GRAHAM, A. L., BEVAN, A. W. R. & Hutcuison, R-

(1985) “Catalogue of Meteorites”, fourth edition,

(University of Arizona Press, Tucson.)

Kei_, K. (1969) Meteorite composition. /n K. H.

Wedepohl (Ed) “Handbook of Geochemistry” 1, 78-115.

(Springer-Verlag, Berlin.)

______ & FREDRIKSSON, K. (1964) The iron, magnesium

and calcium distributions in coexisting olivines and

rhombic pyroxenes of chondrites. 2 Geophys. Res. 69,

3487-3515,

LinpsLey, D. H. (1983) Pyroxene thermometry. Am.

Mineral. 68, 477-493.

Mason, B. (1974) Notes on Australian meteorites, Rec.

Aust. Mus. 29, 169-186.

Scott, E. R. D, Tayior, G, J. & Kem, K. (1986)

Accretion, metamorphism, and brecciation of ordinary

chondrites: Evidence from petrologic studies of

meteorites fram Roosevelt County, New Mexica, Proc.

17th Lunar Planet, Sci. Conf. E\1S—E123.

Spencer, L. J. (1933) A new meteoritic iron found near

Kyancutta. S.A. Mineral. Mag. 23, 329-333.

STOFFLER, D., BucHWALD, V. & RuBIN, A. E, (1988)

Shock effects in meteorites. Jn J. F. Kerridge & M. S.

Matthews (Eds) “Meteorites and the Early Solar

System”. (University of Arizona Press, Tucson.)

Van Scumus, W, R. & Rispe, P. H. (1967) The

composition and structural state of feldspar fram

chondritic meteorites. /bid. 31, 747-765.

& Woop, J. A. (1967) A chemical-petrologic

classification for the chondritic meteorites. Geochim.

Cosmochim. Acta 31, 747-765.

WALLACE, M. E. & Princ, A. (1991) Geological Note:

The Streaky Bay meteorite, a new (L4) chondrite from

South Australia. Aust. Jour Earth Science.

We its, P. R. A. (1977) Pyroxene thermometry in simple

and complex systems. Contrib. Mineral. Petrol. 62,

129-139.

RECENT SIGHTINGS OF KILLER WHALES, ORCINUS ORCA

(CETACEA:DELPHINIDAE), IN SOUTH AUSTRALIA

BY JOHN K. LING*

Summary

Twenty-six sightings of what are believed beyond reasonable doubt to have been killer whales

(Orcinus orca) were reported between August 1982 and March 1990 in most South Australian

coastal waters. The number of whales ranged from one to 52, with singletons and groups of two,

three, five and eight whales being the most common social aggregation. Most of the sightings were

made between December and June; this may reflect observer effort as much as any seasonal pattern.

KEY WORDS: Killer whale, Orcinus orca, Cetacea, sightings, South Australia, social groupings,

seasonal occurrences

Trmisactons ar the Raval Socyrey of S Aust (199%), 11S¢2), 9S-98_

RECENT SIGHTINGS OF KILLER WHALES, ORCINUS ORCA (CETACEA:

DELPHINIDAE), IN SOUTH AUSTRALIA

by Joun K. Lina*

Summary

Ling, J, K. (1991) Recent sightings of killer whales, Orcinus orca (Cetacea: Delpliinidae), in Sourh Australia.

Trans. R. Sow. 5. Aust. 115(2), 95-98, 31 May, 1991,

‘Twenty-six sightings of what are believed beyond reasonable doubt to have beem killer whales (Oretrus

ofva) were reported between August 1982 and March 1990 in most South Australian coastal waters. The

number of whales ranged from one to 52, with sinpletous and groups of two, three, tive and eight whilles

heing the mast cammon social ageregation. Most of the sightings were niade between December and June;

this muy peflect observer effort as much as any seusona) pattern.

KEY Woros: Killer whale, Greivuy orva, Ceracea, sightings, South Ausiralia. social groupings, scasonal

occurrences

Introduction

The only published record of the Killer Whale

(Orcinus orca) in South Australia refers to an

encounter between 1] Killer Whales and a large

whale and calf (apparently Blue Whales,

Balaenoptera musculus) in the South Bast of the

State (Cotton $943) (see Table 1), The Killer Whales

were furiously attacking the head of the larger

whale. This is typical O orca behaviour (Marejohn

1968; Baldridge 1972; Hoyt 1984). Aitken (1971)

oveclooked this record, although he belyeved that

Killer Whales probably did occur off the South

Australian coast.

Since 1982, the South Australian Museum fas

coordinated a public whale sighting and reporting

pregranime known us “Whale Watch - South

Australia’. Orcinus ovrea is one of the most

distinctive and easily identifiable cetaceans because

of its body shape, markings and behaviour. Only

one or cwo diagnostic characters are sufficient for

its positive identification. his paper records and

analyses sightings of killer whales in South

Australian waters resulting from the “Whale Watch”

programme.

Methods

Through extensive media coverage, the public

have been encouraged to report sightings ta the

South Australian Museum of any whales, alive or

dead, fram ground the coast. National Parks and

Wildlife Service rangers and Fisheries Department

officers have been supplied with pro formas

designed to enable identification of cetaceans and

record pertinent inforntarion. Where possible, a

follow-up thterview is conducted by Museum staff

with the observer,

* Soutn Austvatian Musetim, North Terrace, Advluide. S.

Aust, SOOO,

All reports are filed, but only positive

identifications have been used here. An

identification brochure (Anon 1980) is usually sent

io éach person following the interview to help

maintain interest in the scheme, and to allow further

vhecking of characters useful for identification and

follow-up of sightings,

Results and Discussion

Twenty-six reports of Killer Whales were received

Deiween Atigust 1982 and May 19h) (Table tL} th

two instances, photographic confirmation was alsa

received (Fig. 1), Killer Whales have been reported

for all months except January and October, with

most reported sightings occurring between

December and June, The preponderance of

sightings in the summer to early winter months,

may. reflect increased numbers of observers, better

visibility and calmer weather rather than any actual

increase in numbers. of Killer Whales in South

Australian waters during this period, According to

Baker (1983), Killer Whales may calve in autumn

or early winter in New Zealand waters. It is known

that Killer Wheles in other parts of the world moye

about in response to the movement of food species,

eg, herring in Norway (Christensen 1988). It is

possible that their presence in South Australian

waters during the summer. and autumn months ts

due to the availability of favoured food items such

as fish, squid and seals,

A recent reported sighting involved an encounter

with six to eight killer Whales near Point Brown 1n

the Far West, by District Fisheries Officer, Bob

Sprigas of Ceduna, on 2 March,, 1990 (sighting 26).

Mr Spriggs observed the whales from a small boat

for about two hours, There was one large (ca 10 m)

male, four to five whales 4-5 m long and one or

Iwo less than 4 im long. They were first seen

approximalely | km north of Point Brown towards

which they then headed rhrough foul ground. The

Mt J. K. LING

TABLE |. Killer whale sightings in South Australia, 1942 and 1982 - 1990).

Sighting Date

Na,

Locality

1 10 December 1942 4 km SE Port MacDonnell

= Aupust 1982

3 April 1983

Penneshaw, Kangaroo.

Istand

South Coast, Kangaroo

Island

Foot of Gulf St, Vincent

Investigator Strair

Cape Jervis

Goolwa Beach

Waitpinga Beach

Off Glenelg

4 13: December 1983

5 3) May 1986

6 ! June 1986

7 1 June 1986

4 15 June 1986

4) 1 July 1986

io 13 February 1987) Cape Dutton, Kangaroo

Island

IL 19 February 1987) Althorpe Islands

2 2 Novernber 1987

13 14 February i988

Outer Harbor

Brighton North

14 9 Mareh 1988 Smoky Bay

15 3 May 1988 5-7 km ENE Tronbridge

Shoal

16 12 May 1988

17 13 May 1988

Port Noarlunga

Port Noarhinga

18 17 May 3988 Port Elliot

19 19 May 1988 55 km south of Cape du

Couedic

20 25 May 1988 Port Nourlunga jelry

2 8 September 19R8 Arne Bay

Latitude,’

Longitude:

No. af Remarks

whales.

22 13 March 1988

33.0 -2K May 1989

44 8 December 1989

15 km west of Rapid Head

4 km south of Cape Servis

25 27 February 1990 $15 km NNE of Snug Cove,

2 km s6uth of Howlers Bay

Kangaroo Fsland

26 2 March 1990 Point Brown

27 31 May 1990 Greenly [sland

38°05°S, 140°44°E 1) Attacking other whales

(Cotten 1943)

359435, 137°S6'E 8 Photographs

37°00'S, 137° 00'E § {= #2)

34°33'S, 138710°E 3 ) adult (2); .2 young (?)

35°27 'S, 136°50'E i F(T) (= #6Y)

35°36'S, 1398°06’E 52 Herding salmon

35°30'S, 138°47'F, 2 Moa FO) Ce +O)

35° 38'S, 138°29'E ! | NM offshore (— #671

34°59'S, 138°352'b. l 1O km offshore Mi? )—

HO?)

35°38'S, 137°90°L 5 adults, 2 young

35°22'S, 136°52'E 4 3 adults, b young

(= #107)

34°45", [RRO29' bh 3 heuding north; 25-30

35°01'S, 19893 'E 1 F (2)

32°18°S, 133°50'°L 5 2Ms + 2 Fs

(photographs)

35°08 'S, 137748°F 2 Attacking other whales

36°09'S, 138°29'b 5 1M, 4 Fs

25°09'S, 138°29°F, . Fs(?) Heading north

(= A167)

35°32'S, 138°41'E 2 (~ 167)

36"94‘'S, 136°42'E 24 (- #6?)

35°09'S, 138"29°L 2? (= rhe?)

33°36'S, 138°35'°E 5 OM, 40 find © 20 fh

Fs(?)

35°31'S, 138" 10'E 3

32°00'S, 132"27°E 2

35°5R'S, 138'06"E 8-9

8 10m Jong

M, & int others, 5.6 m

2999

#227)

2 Ms, 1 FC?) (=

#227)

1M, 25-30 Tt Ps

1S-20iT

(photographs: Fig. 1)

1 M, 4-5 V's, 1-2 young

(= #267)

35°94°S, 130°52'E 3

32°94"S, 193°SUE 8

34°30°S, 134°45'E 6-8

whales were also seen ut One stage to he working

ina kind of pack and harassing a Common Dolphin

(Deiphinus delphis). The latter appeared to be quite

frantic in trying to escape from the Killer Whales

and was leaping out of the water and somersdulting

and cartwheeling in the air. Several of the smaller

Killer Whales were also seen leaping into the air and

re-entering the water head-first. [ is rather

remarkahle that five Killer Whales were seen (and

photographed) in almost the same area on 9 March,

1988 (sighting 14). Although it was not possible to

match any photographs to the same whales scen in

the two sightings, it is not inconceivable that the

same whales were involved on both occasions, and

avain on 3) May, 1990 ar Greenly Island (sighting

27).

Killer Whales are known to form close-knit,

highly stable, and probably permanent groups in

British Columbia (Ford & Ford 1981). Given that

Killer Whales are seen infrequently in South

Australian waters, it seems extremely likely that

several of the 26 sightings could have been of the

same animals or of the same herd, For example,

sightings 5, 6 and 7 and possibly sightings 5, 6, 7.

8 and 9 sightings 10 and 11; and sightings LS to

20 probably ineluded all or some of the same

animals. Furthermore, some of the sightings seen

in different months or even years could have been

of the same whales, e&.g. 2 and 3; and 22, 24 and

25, as well as 14 and 26 referred to above: Excluding

possible repeat sightings, social graupings were as

follows: | whale (3); 2 whales (3); 3 whales (4): 5

KILLER WHALES IN SOUTH AUSTRALIA 97

Fig. 1. Killer Whales, Orcinus orcu, near Point Brown,

South Australia, 2 March 1990. Top, male; centre, male

and female; bottom, female. Photographs R. Spriggs,

whales (4); 8 whales (3); 24 whales (1); and $2

whales (1).

Recent Killer Whale sightings in South Australia

ure not confined to particular areas, but are spread

from Goolwa Beach on the south coast to Fowlers

Bay in the Far West, and into both gulfs. Indeed,

at this stage, it is not possible to nominate any one

locality where regular observations might be carried

out. Taking Cotton's (1943) report from the South

East into account, also means that Orcinus orca has

been observed over almost the entire South

Australian coast. Only sightings from land or boats

close to land have been reported to date.

There have been 46 strandings in New Zealand

(Baker 1983) and two in Tasmania involving at least

three animals (Nicol 1986), Nicol believes that (he

Killer Whale’s predatory lifestyle in and familiarity

with inshore waters should include an ability to

navigate safely away from hazards that might affect

other (particularly oceanic) species and cause them

10 strand. However, many strandings occur on

Vancouver Island, Canada, where large resident and

smaller transient populations frequent the waters

between Vancouver Island and mainland British

Columbia, Canada, and Washington State, USA;

despite the Killer Whales’ presumed familiarity with

the area (Ford & Ford 1981).

There is very little Killer Whale material in the

South Australian Museum collections, and

associated data are imperfect (Table 2), We have no

documented evidence that Orcinus orca has ever

stranded in South Australia, although the five teeth

referred to in Table 2 cannot be discounted entirely

as such evidence.

The results presented here emphasise the value

of encouraging people to look out for whales and

report sightings to the authorities and, in particular,

to try and obtain good clear photographs of dorsal

fins and other markings by which whales may be

identified. Such data will lead to a_ better

understanding of the biology of this spectacular

species in South Australian waters.

Acknowledgments

The interest and cooperation of the many people,

particularly Mr Bob Spriggs, who have reported

sightings, provided information during interviews,

and supplied confirmatory phatographs which have

made this article possible are gratefully

acknowledged, Thanks are also due to Dr Catherine

Kemper and Dr Graham Ross for commenting on

earlier drafts of the manuscript. Trevor Peters

Prepared the photographs for publication and

Debbie Lowery typed the paper.

98 J. K. LING

TABLE 2. Orcinus orca specimens registered in the South Australian Museum

Reg. No. Date of registration Locality Material/Remarks

or collection

M 1590 11/9/1922 “near Mount Lofty Range” Part skull (specimen lost)

M 3224 2 1932 ‘South Australian beach” Five teeth

M 5345 ? 1945 Portland, Victoria Mandible ramus; no data*

M 5649 2? 1945 unknown Part skull (no mandibles); no data*

*These could be of one and the same animal

References

AITKEN, P. F. (1971) Whales from the coast of South

Australia. Trans. R. Soc. S. Aust. 95, 95-103.

ANON. (1980) A coast-watching scheme for marine

mammals. Wildlife in Australia 17(September), 66-71.

Baker, A. N. (1983) “Whales and Dolphins of New

Zealand and Australia: an Identification Guide”.

(Victoria University Press, Wellington).

BALDRIDGE, A. (1972) Killer whales attack and eat a grey

whale. J. Mamm. 53, 898-900.

CHRISTENSEN, I. (1988) Distribution, movement and

abundance of killer whales (Orcinus orca) in Norwegian

coastal waters, 1982-1987, based on questionnaire

surveys. Rit Fiskideildar 11, 79-88.

CorTTon, B. C. (1943) Killer whales in South Australia.

South Aust. Nat. 22, 2-3.

Forp, J., & Forp, D. (1981) The killer whales of B.C.

Waters 5, 3-32.

Hoyt, E. (1984) “Orca the Whale Called Killer” (Camden

House, Camden East, Ontario).

MorEJOHN, G. V. (1968) A killer whale - gray whale

encounter. J. Mamm. 49, 327-328.

NICOL, D. (1986) A review and update of the Tasmanian

cetacean stranding record to the end of February 1986.

Univ. Tas. Envir. Studies Working Paper 21, 1-97.

CRINIA TSCHUDI (ANURA: LEPTODACTYLIDAE) FROM THE

CAINOZOIC OF QUEENSLAND, WITH THE DESCRIPTION OF A NEW

SPECIES

BY MICHAEL J. TYLER

Summary

The leptodactylid frog Crinia presigngera sp. nov. is described from a series of Oligo-Miocene sites

at Riversleigh Statioin northwest Queensland. This finding represents the first record of the genus

Crinia from the Tertiary and the first record of fossil material of Crinia from Queensland. Crinia

remota Tyler & Parker is reported from a Quaternary cave deposit at Riversleigh Station.

KEY WORDS: Crinia, ilia, Leptodactylidae, Cainozoic, Queensland, Australia

Trans tivas uf the Rew) Syeiers of S. Aust 99)), 11512), 99-101.

CRINIA TSCHUDI (ANURA: LEPTODACTYLIDAE) FROM THE CAINOZOIC OF

QUEENSLAND, WITH THE DESCRIPTION OF A NEW SPECIES

by MICHAEL J. Tri ER*

Summary

Tyiex, M. J. (1991) Crinta Tschudi (Anura; Leprodactylidac) trom the Cainozoic of Queensland, with the

description of a new specics. Trans, R, Soe. 8S. qust, UUS(2), 99-101, 31 May 1997,

The leptodaciylid frog Crima presignifera sp. nov, is described tram a series of Oligo-Miucene sites at

Riversleigh,Starion im northwest Queensland, This tinding represents the first record ef ihe genus Crinia

from whe Tertiary and the first record of fossil material of Crinia from Queensland. C'rinia remota Tyler

& Parker is reported from a Quaternary cave deposit at Riversleigh Station

Key Worps Crinia, ills, Leprodactylidag, Cainozoic, Queensland, Australie

Introduction

The genus Criniv Tschudi, including frogs

relerred lo Ramidella Girard (according to Heyer et

a/, 1982), is a group of 12 small, ground-dwelling

species that live close to water. It is represented in

all bul the arid, central portion of Australia, and

{he central coastal area of Western Australia. One

species (C. remota (Tyler & Parker}) occurs in

worthem Ausuralia and southern: New Guinea.

The genus Crinia bas been the subject of diverse

studies, particularly in the fields of polymorphism

and of pre-mating isolaling mechanisms.

Consequently, published data on this genus are

more Substantial than those available for any other

genus in Australia,

The phylogenetic relationships and the origin of

Crinia are unclear, Merphological evidence {Heyer

& Liem 1976; Davies 1989) suggests a close

relatiouship with Psexdophryne Fitzinger and

Upernleia Gray.

The current fossil record of Crinia-consisis of the

ealanl species of C. signifera Girard, trom

Pleistocene deposits at

Victoria Cave, in the southeast of South Australia

(Tyler 1977), and the extant species C georeiana

Tschudi from Pleistocene deposits at Skull Cave and

Devil's Lair in the extreme squthwest of Western

Australia (Tyler 1985),

Here I report the first Tertiary record of Cristia,

and the first Quaternary record of Cringe from

Queensland. I'he genus Crinia aceurs at several

Caozoty sites at Riverslcigh Station in northwest

Queensland, Previously, two other feplodactylid

genera have been. reported) from that area!

Leefiriodus Boulenger (Tyler 1989) and

Lintiodynastes Fitzinger (Wier 1990).

* Dept of Zoology, University of Adelaide, Box 498,

€..P.0., Adelaide, 3. Aust. S001

Henschke’s Cave and-

Maierial and Methods

The material is deposited in the Queensland

Museum, Brisbane (QM) and the South Australian

Museum, Adelaide (SAM), Letters following, the

abbreviations are departmental identifications.

Comparative studies were based on the

esteological collections ut the Department of

Zoology, University of Adclaide.

Osteological nomenclature and methods of

measurement follow Tyler (1976, 1989),

‘Syslematics

Family; Leptoadactylidae Werner, 1896

Sub-fannly; Myobatrachinae Schlegel, 1850

Genus; Crinia Tschudi, 1838

At the time of the preparation of a description

of the ihal eharacteristigs of Australian frogs. by

Tyler (1976), Crinia was considered a monotypic

genus, and Renidella distinct from it. The principal

morphological features distinguishing C. georgiana

from the species of Ranidella examined /R.

parinsignifera aad R. signifera) were considered to

be the extent of the dorsal protuberance, aid the

presence of a very slight longitudinal indentation

upon ihe lateral surface of the ilial shaft of Crinia

that was absent from the Ranidella species (‘Tyler

1976).

Examination of thes¢ particular features in

additional species formerly referred io Ranidedla:

bilingua, desetiteolu, glauerti, invxignifera,

pseudinsignifera, remota and riparia indicates that

the generic differences proposed by ‘Iwler (1976)

cannot be sustamed. Nevertheless, and perhaps

morte sizmificantly, despite its larger adult size, it

is evident that C, georgiana has a more robust ilium

than the species of Renidella so far examined, in

which the ilial shaft proportionately is deeper and

more substantial than in those species {hat are now

its congeners.

LOU MJ TYthe

Diugnostie generic features of Crinja are the

curved and medio-laterally flattened shafl, lacking

a dorsal crest and possessing a very slight medial

indentation; large acetabular fossa with a broad

peripheral rimy slight development ot the ventral

acetabular expansion anda subacetabular vone that

dogs not protrude anteriorly, The dorsal acetabular

expahsion is poorly developed. The dorsal

prominence is low and the dorsal prolubyrance

slichily developed.

Crinia presigaifera sp. nov.

PIG. 1A

Holotype: QM FA7630. A left lium collected at

Wayne's Wok Site, Riversleigh Stalion, northern

Queensland,

Deseription of holorype: Uial shaft slender and

slightly cucved, Lacks dorsal crest but with

moderately deep, slight Tatcral voncavity along

proximal one-third of shalt. Distal end of ilial sbatt

incomplete inferiorly.

Acetabular fossa Jaree and deep, with prominent,

clevated rim. Dorsal margin of acetabular foxsa

superior to utferior margin of ilial shall, Pre-

icetabular zone evenly rounded, expanding

inferiorly into protruding rounded Pare of sub:

iwetabular expansion. Inferior margin of sub-

acetabular expansion lacking.

Dorsal acetabular expansion raised slightly,

Dorsal prominence poorly developed. Dorsal

proluberance narrowly oval, prominent, projecting

laterally,

Length of ilium: 6.3 mm.

Paratypes: ‘There are 18 paratypes — Outasite Site:

OM F17634-36, 18155, SAM P31230-33: Quentin's

Quarry Site; AM F17631; Neville’s Crarden Site: (QM

F48156-58, SAM P31234-35; Two Trees Site: OM

F17632, SAM P31228; Camel Sputum Site! QM

117633, SAM P31229.

The largest of the specimens in which the ilial

shall ts complete measures 7.) min. A paratype is

shawn in Pig, 1,

Throughout the series the dorsal prominence and

Jorsal protuberance are conspicuous, atid the dorsal

acetabular expansion ts clevared only slighuy, The

acetabular fossa is consistently large, but the

breadth of ihe adjacent pre-aceiahular zone varies

from extremely narrow (at its closest proximity to

the fossa) 10 moderately wide. The ventral

acetabular expansion is incomplete in mast

specimens,

Somperison with ather species: The overall

similarity in external marpholagy of species af

Crinia (excluding © geargiang) is aceoarpanied by

an extreme conservansim in the form of the ium,

Most of the specific characters are slight, when

7 waar!

Le nr. ial

hate —_— -

| co —.,

be fe ; en : =

| a a =

I { 45 _f

big. loo. Crnnad presipiifferd sp. nay paratype: QM

FIT630, B, Cinta rerio SANT PITIIG,

compared with those distinguishing members of

other genera e.g. Limnodynastes and Litorte.

Nevertheless one feature distinguishing

presigaifera from congeners is the narrow pre-

ucefubular zone clearly demonsirated by com-

parison with C. remota in Fig. 1. Amonese extant

species the one with the narrowest pre-acctabular

zone is C signifera but even in that species it js far

more substantial than in the new species.

Stratigraphy and lithology In the Riversleigh

Station area Archer, ef af, (1989) recognined u

minimum of five types of Oligo-Miocene

carbonates that are rich in bones. The sites bearing

©. presignifera comprise two. sequences of lacustrine

carbonates that contain principally non-aquatic

local faunas, These collectively are referred to by

them as “Sysiem B” and “System C”

Etymology: In adding pre- (1, prae) as a pretix to

signifera 1am alhiding to the ancestral nature of

the fossil specics relative lo cxtant species,

Crinta remota (Tyler & Parker)

rig. 1B

Material; A single right iium, SAM P31236 from

Carrington Cave, Riversleigh Station, Queensland,

Descripiive notes: The tlial shatt is cylindnecal and

the terminal portion 1s missing. Lixisting length 4.6

mn, The superior portion of the dorsal acetabular

expansion is missing. The ventral wecetubular

eXpansion is broadly expanded,

Comonents: Crinia remota (Fyler & Parker 1974) was

described from southern Papua and is now known

to occur also in northern Queensland and the

Northero Territory including Melville island andl

Groote Fylandt (Tyler ef af, 1985, in press). The

present specimen does not differ from specimens

examined and is (he lirst fossil record of ihe species.

CRINIA TSCHUDI FROM THE CAINOZOIC OF QUEENSLAND wi

Carrington Cave is situated in a hill adjacent to

the Gregory River. The specimien was found near

the surface of a vast mound of fragmented bones

derived from the excreta of the ghost bat

Macroderma sigas, and is located approximately

100 m trony the entrance to the cave,

The age of the deposit is unknown but it is

presumed to be Holocene or Late Pleistocene.

Acknowledgments

Tl am extremely grateful to Professor Michael

Archer, Dr Suzanne Hand and Mr Henk Godthelp

of the University of New South Wales, for their

continued aid and support throughout my study of

the fossil frox fauna at Riversleigh Station.

Laboratory studies have been funded by a grant

from the Australian Research Council, Miss Leanne

Seller sorted and documented the material, and

prepared the illustration. Research facilities were

provided by the University of Adelaide. ‘The

assistance of Parke Davis Pty Ltd is gratefully

acknowledged.

The materials upon which this srudy was based

were obtained through the support of the following

funding bodies and organisations to M. Archer, S.

Hand and H. Godthelp; Australian Research Grants

Scheme; Department of Arts, Sport, the

Environment, Tourism and Territories; National

Estate Programme Grant Scheme; Wang Computers

Pry Lid; Australian Geographic Pty Ltd; Mount Isa

Mines Pty Ltd; the Queensland Museum; the

Australian Museum; the Royal Zoological Society

of N-SW.; the Linnean Society of N.S.W.;

Ansett/Wridgways Pty Ltd; Mount Isa Shire

Council; the Riversleigh Society and the Friends of

Riversleigh.

References

ARCHER, M., GoptTugtp, H., Hann, S. & MEcikiam, D,

(1989) Fossil mammals of Riversleigh, northwest

Queensland: preliminary overview of biostrativraphy,

correlation and environmental change. Aus. Zool.

25(2), 29-65.

Davies, M. (1989) Ontogeny of bone and the role of

heterochrony in the myobatachine penera L’peroleiu,

Crinja and Pseudophryne (Anura: Leptodactylidae:

Myobatrachinae). J. Morphal. 200, 269-300.

Heyer, W, R. & Liem, D. S§, (1976) Analysis of the

intergeneric relationships of the Australian frog family

Myobatrachidae, Siithson. Contrib. Zaal. 233, 1-29.

, DAUGHERTY, C, H. & Maxson, L, R. (1982)

Systematic resolution of the genera of the Crinia

complex (Amphibia; Anura: Myobatrachidae). Prac.

hial. Soc. Wash, 95, 423-427.

Tyier, M. J. (1976) Comparative osteolopy of the pelvic

girdle of Australian frogs and description of a new fossil

genus. Trans. R. Soe. 8. Aust, WO, 3-14,

(1977) Pleistacene trogs from caves at Naracoorte,

South Australia, /bid. 101, 85-89.

_. ___. (1985) Quaternary fossil frogs.trom Skull Cave and

Devil’s Lair in the extreme south-west of Western

Australia. Rec. Wi Aust, Mus. 12, 233-240.

(1989) A new species of Lechriodus (Atura:

Leptodactylidac) trom the Tertiary of Queensland, with

a redetinition of the ilial characteristics of the genus,

Trans. R. Soc, S. Aust. 113, 15-21.

(1990) Limnodynastes Fitzinger (Anura:

Leptodactylidae) From the Cainozoic of Queensland,

Mer. Qld Mus, 28(2), 779-784.

____, Davies, M, & Watson, G. F (1986) The frog

fauna of Groore Eylandt, Northern Territory, Australia.

Zool. J. Linn. Soc. 88, 91-101.

(in press) The frog fauna of

Melville Island, Northern Territory. Beagle.

& Parker, F (1974) New species of hylid and

leptodactylid frogs from southern New Guinea, Trams.

R. Soc. S. Aust. 98, 71-78,

A LARGE NEW SPECIES OF LITORIA (ANURA: HYLIDAE) FROM THE

TERTIARY OF QUEENSLAND

BY MICHAEL J. TYLER*

Summary

A new species of large hylid frog of the genus Litoria Tschudi is described from a small series of

disarticulated, and fragmentary ilia from Tertiary freshwater limestone deposits on Riversleigh

Station, Queensland. The species exhibits unique characters in the form of a pair of depressions

situated on the ilium, superior to the acetabular fossa, and a horizontal flange projecting mediad

from the ilial shaft. Litoria magna sp. noy. is the first record of the genus from the Tertiary of

Queensland, but several congeners await further study and description.

KEY WORDS: Litoria, new species, Tertiary, Riversleigh, Queensland

Transactions of the Royal Society of S. Aust, (1991), 115(2), 103-105.

A LARGE NEW SPECIES OF LITORIA (ANURA: HYLIDAE) FROM THE TERTIARY

OF QUEENSLAND

by MICHAEL J. TYLER*

Summary

TyLer, M. J. (1991) A large new species of Litoria (Anura: Hylidae) from the Tertiary of Queensland.

Trans. R. Soc. S. Aust, 115(2), 103-105, 31 May, 1991,

A new species of large hylid frog of the genus Litoria Tschudi is described from a small series of

disarticulated, and fragmentary ilia from Tertiary freshwater limestone deposits on Riversleigh Station,

Queensland. The species exhibits unique characters in the form of a pair of depressions situated on the

ilium, superior to the acetabular fossa, and a horizontal flange projecting mediad from the ilial shaft.

Litoria magna sp. nov, is the first record of the genus from the Tertiary of Queensland, but several congeners

await further study and description.

Key Woros: Litoria, new species, Tertiary, Riversleigh, Queensland

Introduction

Documentation of the rich Tertiary frog fauna

from freshwater limestones on Riversleigh Station

in northwest Queensland, commenced with the

description of a new species of the leptodactylid

genus Lechriodus Boulenger (Tyler 1989a). Tyler,

et al. (1990) discuss the significance of the numerical

abundance and dominance of that genus amongst

the frog fossils recovered.

Two Tertiary species of Limnodynastes Fitzinger

have been reported from Riversleigh Station (Tyler

1990), but as yet the hylid fauna is unknown.

Here [ describe the first of several species of the

hylid genus Litoria Tschudi. It is represented by a

smail series of ilia, is distinguished by a suite of

unique characters, and is noteworthy for its

particularly large size.

Material and Methods

The material is deposited in museums abbreviated

in the text as follows: Queensland Museum,

Brisbane - QM; South Australian Museum,

Adelaide - SAM. Letters following the abbreviations

are departmental identifications.

Comparative studies were based on the

osteological collections of the Department of

Zoology, University of Adelaide.

Osteological nomenclature follows Tyler (1976),

methods of measurement and orientation of

specimens follow Tyler (1989a), and stratigraphic

interpretation is after Archer ef al. (1989).

Systematics

Family: Hylidae Gray, 1825.

Sub-family: Pelodryadinae Giinther, 1859.

Genus: Litoria Tschudi, 1838.

* Department of Zoology, University of Adelaide, Box

498 G.P.O., Adelaide, S. Aust. S001

The diversity in external morphology in this

genus is paralleled by the form of the ilium;

comparative osteological data for 21

Australopapuan species was presented in tabular

form by Tyler (1976). Diagnostic features are the

absence of a dorsal crest upon the ilial shaft, and

the presence of a shallow, longitudinal groove upon

the medial surface of the shaft. The dorsal

protuberance and dorsal prominence are not raised

much above the superior border of the ilial shaft,

but usually are well differentiated.

Litoria magna sp. nov.

FIG, 1

Holotype: QM F17627. The proximal two-thirds of

aright ilium collected at Camel Sputum (C.S.) Site,

Riversleigh Station, northwest Queensland.

Description of holotype: lial shaft curved, deep,

robust, cylindrical in section proximally and lacking

a dorsal crest (Fig. la). Medial surface of shaft with

horizontal, superior flange becoming progressively

more prominent and rendered more conspicuous by

accompanying inferior indentation. At its distal

extremity, flange extends from shaft for distance

equivalent to depth of shaft (Fig. 1b).

Acetabular fossa large, with narrow but

prominent rim. Pre-acetabular zone evenly rounded,

with narrow separation from acetabular fossa. Sub-

acetabular zone incomplete. Medial surface of

acetabular region bears shallow, central cavity ca.

1.0 mm in diameter and 0.7 mm deep.

Dorsal acetabular expansion slightly truncated

but apparently poorly developed, being elevated

only slightly above level of ilial shaft.

Dorsal prominence and dorsal protuberance

poorly developed and replaced by a pair of distinct

depressions: one located superior to anterior rim

of acetabular fossa, the other superior to the centre

of the fossa.

104 M. J. TYLER

Fig. 1. Holotype ilium of Litoria magna sp. noy. A. Lateral surface; B. medial surface.

Length of ilium 18.7 mm; estimated length of

reconstructed complete ilium 25-27 mm.

Paratypes: Four incomplete ilia: Camel Sputum

Site: QM F17628, SAM P31220-21; Gag Site QM

F17629.

Variation: Each of the paratypes is an incomplete

and variously abraded proximal head of an ilium.

Although the ilial shafts terminate at or proximal

to the development of the horizontal flange, the

conspecificity of the material with the holotype is

demonstrated by the presence of a pair of

depressions superior to the acetabular fossa. In both

SAM P31220 and QM FI17628, the anterior of the

depressions leads anteriorly to a groove connecting

to the superior margin of the shaft.

Paratype QM F17629 is highly silicified, and the

elaboration of bone bordering the depressions is

suggestive of localised, secondary exostosis.

The paratypes appear to be from individuals that

would have been smaller than the holotype and

further differ in having a more elevated dorsal

acetabular expansion.

The pit reported on the medial surface of the

acetabular area of the holotype corresponds to the

position of a foramen in QM F17629 and is

presumably the effect of artificial enlargement of

such a feature.

Paratype SAM P3122] consisted originally of the

lateral face of the acetabular region and proximal

ilial shaft. This fragile specimen disintegrated into

three smaller fragments after it had been drawn. It

probably was the corresponding portion of the more

complete SAM P31220 which was extracted from

the matrix at the same time.

Comparison with other species: The presence of a

pair of depressions superior to the acetabular fossa,

and the horizontal flange upon the ilial shaft

distinguish this species from all congeners.

The depth of the ilial shaft of the holotype is

indicative of a robust-bodied species, whilst

examination of ilia of large species of Litoria,

including L. /esueuri (Dumeril & Bibron), L.

caerulea (White) and L. infrafrenata (Gtinther)

suggest that L. magna could have been bigger than

any extant species, certainly having a snout to vent

length of more than 120 mm.

Stratigraphy and lithology: Archer, Godthelp, Hand

& Megirian (1989) provided a _ preliminary

assessment of the stratigraphy of the major fossil-

bearing sedimentary deposits at Riversleigh. The

NEW TERTIARY LITORIA SPECIES 105

Iwo sites from which L, meena has been recovered

occur within a series of Oligo-Miocene lacustrine

carbonates that range in age from approximately

15 to 25 million years BP. Camel Sputum Site occurs

in limestones interpreted to be of early to middle

Miocene age (within the Archer e7 af. “System BY”),

whereas Gag Site occurs in slightly younger middle

(o early late Miocene limestones (i.e. “System C”

deposits):

Etymology: Latin magna: large, alluding to the size

of the fossil specics.

Discussion

Liforia is the most spectose venus in Australia

and New Guinea, and osteological data have

supported the sub-division of the unit into assumed

natural groupings. of species (Tyler & Davies 1978)..

The Tertiary record of Litoria is poor. Isolated

specimens (disfed in Tyler 1989b) have nor been

identified to species. Thus J. magaa is. the first

Tertiary member of Litoria to be described.

The unique horizontal flange upon the ilial shaf,

and the pair of supra-acetabular pits set L. magna

apart From all extant species, and hence does nor

appear tu be ancestral to any modern species graup,

Other species of Litoria from Tertiary deposits

on Riversleigh Station await description, bul none

approaches the size of L. magna, The large size of

i. mugaa is uausual amongst the Tertiary frogs

known from Riversleigh Station, for the species

there are predominancly small, a feature

charavteristi¢, amongst extant frogs, of geographic

areas that experience a high raiifall (Tyler 1989b,

Fig. 8) and are pot exposed to seasonal aridity,

Acknowledgments

Progress on the elucidation of the Tertiary frog

fauna of Riversleigh Station has been supported

generously by Professor Michael Archer, Dr

Suzanne Hand and Mr Henk Godthelp.

Laboratory studies in Adelaide have been funded

by grants from the Australian Researeh Council. L

thank Veronica Ward and Leanne Seller for their

invaluable assistance in the sorting and

documentation af the material, Research facilitics

were provided by the Liniversity of Adelaide. The

provision of gelatin capsules for hone storage by

Parke Davis Piy Ld is gratefully acknowledged.

The materials upon which this study was based

were obtained through the support of the following

funding bodies and organisations to M, Archer, 5,

Hand and H. Godthelp: Australian Research Grants

Scheme; Department of Arts, Sport, the

Environment, Tourism and ‘Territories; National

Estate Programme Grant Scheme! University of

New South Wales; Wang Computers Piy Ltd;

Austrahan Geographic Ply Ltd; Mount Isa Mines

Pty Ltd; the Queensland Museum; the Australian

Museum; the Royal Zoological Saciety of NSW;

the Linnean Society af N.SW,; Ansett/Wridgways

Pty Ltd; Mount Isa Shire Council; (he Riversleigh

Society and thy Friends of Riversleigh,

References

ARcHUR, M,, GobiHELY, ML, BANU. S. 1d MEGIROAN,

D, (1989) Fossil mammals of Riversleigh, northwestero

Queensland: prelirinary overview of biostraripraphy,

correlation and ¢nvironmental chanee. dist, Zeel,

25(2), 29-65,

Triek, M. J, (1976) Comniparative ostcology of the pelvic

girdle of Aystralian frogs and description of a new fossil

genus. Trans. R. Soe. S$. Aust. 100, 3-14.

(19890) A new species of Lechriveus (Anuray

Leptodactylidas) trony the Tertiary of Queensland, with

a redefinition af thy ilia) characteristics of the gents

Vhict V3, 15-21.

——_ (19896) "Ausiratian Froes." (Viking 'Neil,

Melbourne.)

(1990) Limnodynastes Fitginuer (Anna!

Leptodactylidae) from the Cuinozvie of Queensland,

Mem. ld Mus, 28, 779-784.

—.— . & Daviks, M. (1978) Species-sioups within the

Australopapuan hylic trow genus Literia Tschudi, Aus,

J. Zool. Suppl. Ser. 63, 1-47.

_.> Hawn, 'S, & Wakn, V1. (1990) Analysis of

the frequency of Lechriadus invrergerivvus Tyler (Anura:

Leptodactylidae) in Olivo-eMiovene loval faunas of

Riversivigh Station, Queenslund, Prec. Linn, Soc

NLS.HE 112(2), 105-109,

THE DISTRIBUTION OF ECHINOCEPHALUS OVERSTREETI DERDORFF

& KO (NEMATODA), A PARASITE OF ELASMOBRANCH FISHES IN

AUSTRALIAN WATERS

BY IAN BEVERIDGE

Summary

Transavtions af (he Royal Soctery af S Aust. (1991), 115021, 107.

BRIEF COMMUNICATION

THE DISTRIBUTION OF ECHINOCEPHALUS OVERSTRERTI DEARDOREF & KO

(NEMATODA), A PARASITE OF ELASMOBRANCH FISHES IN AUSTRALIAN WATERS

A recent study of ihe cchinocephalid nematode parasiles

of elasmobranchs trom Suuth Australian coastal waters!

identiticd the species present as Evhirocephalus overstreeti

Deardorit & Ko, (983, previously known only froma ray

Jaeniura melanaspilas Weeker trom the Marquesas.

Islands je 10°S, 40°F) in the southern Pacitio Ocean’,

and sinee reported from another species of ray,

Urogyminus asperrimus (Bloch & Schneider), trom

Emiwetok Atoll (2°N, 164°E) (also spelled Enewetak

Atollp in the Marshall Islands of the castern Pacific’.

A number of records east ol Ech inovephatis sp. from

regions of Ausrratie other than South Australia, fram

Molluscs, lleasts, dolphins and turcles', hur in phese

causes Lhe species involved could ner be determined with

certainty, since (he reports were based on larval specimens.

In addition, adults uf a second. species, &. sinensis Ko,

1975, have been found in a may, Daspetis Jlevieruim Ogilby

off the southern Queensland coast, and there are no

known morphologieal criteria which reliably distiiguish

the larvae of the two species!

Since alw study of & overseer i South Australian

Waters was completed, 9 further 770 clasmobranchs,

coreriiig an additional 72 species, have been examined far

Parasites, comprising specimens irom north-eastern

Queensland, the Northern Territory, the north-west and

southavest coastal arcys of Western Australia and tram

lasmania, Detailed records of all elasmobranch loses

cxamined for parasives have been lodged in the

helminthologica) collections of the ‘South Australian

Mhiscum (SAM).

Adult specimens of &\ aversrreed’ were recovered from

(numbers in parentheses are specimen registration numbers

Wi SAM) 2of 2 (1000) Dasvatis sephen (Forsakal) from

Fog Buy, N.T. (HC 17194) and 13-01 15 (87%) at Nickol

Bay, W.A, (NHC 17200-17205), trom t of 12 (89h)

Myhobutis austratis Macleay fram Bunbury, WA. (HO

17208) and 2 of 4 (S005) Mererodouiis portusjackseni

(Meyer) from Bunbury, WA, (HO 17252) and tor |

(100%) at Stantey, Tats. (HO 17231).

Larval sluges Of Echiaacephelus which could not be

identified to species were recovered from one of 10 (aut)

Carcharhinus plurrtbeus (Nacdo) fron, Bunbury, WA,

CHE 17216), two of 10 (204) RAvachahaons diiddonsis

(Porsskal) from Flac Top ts,, Qkt (HE (6041, (6042) bar

innone af Wat the same species examined in the Northen

"Beveridge, 1. (1987) Trans. R. Sov. S. Avat, TH, 79: 92.

“Deardortf, T. L. & Ko, Ro. (1483) Proc. Helminthol,

Soc. Wash. S@, 285-293.

Territory and Western Australia, fram one of 14 (7%)

Armentura varnak (Porsskal) from Fou Bay, NuT. (HC

17219), but from none of tive H, warneck from Western

Australia and Queenstand ancl 1 of 1 (00%) Aerobarus

narinary (Evphrasen) trom Fog Bay, NT, (HC 17214),

Adult specimens of E. overstreeti were thus recovered

from elasmobranchs in-coastal waters of three of the four

shites from which fish were examined, inferring an almost

circum-continental distribution. Taken together wilh pre-

existing records’ ', the data suggest that £. overstreeti

is widespread inthe Pacific region. &. sinensis, described

originally from Metobutus narinari Ruphyasen trom Hong

Kong* was tor encountered in this survey, so the sole

record of the species [rom (he Australian region vemains

That trom 2D fluvierum trom Queenstand!,

To spite of the large numbers and wide range of species

af elasmobranchs exammed, Echinucephalus adults and

farvac-aceurred al a muh lower overall prevalence (2.9% j

than reporled earlier for South Australian coastal waters:

(47.3%), The difference does not reflect the very few

hererodonnforins vallected outside South Australia, siree:

the respective prevalences when these hosts. ave removed

are 2.5% and 39.89, mut may réfect biases in the other

host species surapled.,

Th South Austrtlian waters, gravid specimens of £

beerstreeli were tound only in Helerodontus

portusjacksant (Meyer, 1973). None of the specimetis

elected from hosts other than //, portayjuckseny,

included in this report, were gravid, contirming our edcher

observations, However, the type specimens of #.

aersireet, collected Tom the ray 7 melaraspilas were

Sravild sugecsting that heterodonritorm sharks are not the

only suitable hogs. The srate al maturity of the specimens

fram Lt asperriptus ton Bniwerok Atoll was tor

reported, and no echinocephalids were founsl ina single

U. usperrimus examined by us Strom northern Australian

COAT,

Thanks are due ta Mr B G, Robertson who collected

the specimens of AeAintucephaius, and ta the Australian

Biological Reswurces Study who tinaneially supported the

colleching-

‘Brooks, 1, R. & Deardortt, 1, 1. (1988) J. Parasitol, 74,

459. pS

"Ko, ROC (1975) 4cay J, Zool, 53, 490-50),

IAN BEVERIDGE, The University of Melbourne, Verennary Clinical Centre, Princes Hiehway, Werribee, Vig, 30311

REVISED AGE FOR AYERS ROCK AND THE OLGAS

BY W. K. HARRIS & C. R. TWIDALE

Summary

tratisdetiany if the Ravel Sariery uf S&S Aet (M91, TES 18.

URILE COMMUNICATION

REVISED AGE FOR AYERS KOCK AND THE OLGAS

In an sunlier paper , vas dediiced chat Ayers Rock

and rhe Otpas were already uplagds 60-65 My ago, in

mittle Palucoeene limes It wis not suggested, «as

Oller’ has claimed with respecr ro Ayers Rock, that ue:

residual looked “very muell as it does today in Palaeocene

rimes mor Mo Route residuals are eroded in

Cambruursediments, The steep bounding slopes. that vive

them their dramatic appearance were formed during the

later Cainozoig, as u result of searpfoot subsurface

Moisture attack", That there were topographic rises on

the present upland sites during or by the Barly Tes cary

was inferred from palacantolagical evidenve discovered

during 4 programme of drilling instigated during a search

for wiler in the nineteen sixties.

A broad shallow valley cul inthe Cambrian sediments

has Deve Cilled by a maximum of jar over 100 ny of

Cainovoic sediments. The basal sequence mw paludal but

the sediments include riverine as well as surlicial aeolian

Materials, Che wpresent plain surface of low relief is

essentially constructional and gives no hint of the bedrock

relict below, The suggestion that the ancestral Ayers Rock

and the Olgas are of considerable antiquity derives first

fram the aruiment that if there were a valley there must

have oeen higher ground on cirher side, aud that the

precursors of the Modera residuals occupied sume of this

higher ground; and second [rom the occurrence near the

base of the fill sequenge and at depths of 81-84 pot

lignites that contin ab assemblage of plant comains that

was conydered (Oo be of iniddle Palacocene age. The

uplands probably look the form of a Kail-covered low

domy en the case of Ayers Rock - the present bevelled crest

plus a regolithie veneer ~ and ola vamplex af low rock

domes in the ease of the Olvas,

This carhier assignment of a Pulueocene ape ro the fossil

assemblaze (from Sample No, $4065) was busecon ihe

ringes of severd! species, in particular, Merkosporites

elliortil Stover, Prutvacidites angulotus Srevet,

Beaupreaidiles elevansiformis Cookson aad B vernicusus

Cookson. Flements which are naw cansidered (a have a

lnited. Late Cretaceous er Early Tertiary age tnelude

Quadraplanus brassius Stover and Rrrace/parires

verrucasus Stover, More recently A.W, Partricle has

eolified frawmenty of eirapae/ipore evunsii Stover &

‘Twidale. . KR. & Hargis, W. KR, (1977) Tray, Re Soe

S -lwst D1, 45-50,

(Mller, €S DL 11977) pp. S§ 98 iA DON. Jeain (Rdg

“Austral A Geography.” (Sydney Lintversity ‘Press,

Sydney),

‘Ollier, CD. (1986) pp. 97-116 fe DN. Jeans (da

“Austolias A Cicowraphy Volume 1, The Natta!

Favioament” (Syvdovy University Press, Sydieyt

rides C. R978) A. Geomorph, Suppl-Banel 3,

177 206,

Partridge (pers. comm, to W.K,H,) the avarciial Pls

species has a middie to late Maastichtian-age in Australia

and Now Zeahand*,

The material was previously equated wirh che

Gambierina edwards zone (Whe Lysiirmepatlenttes

balmei zone), However vn alternative assignment is (0 the

Tricalpites longys zone (Maasirichtian (o basal Danian’)

based on the presence of GO) Arossius and G Pvansié The

younger Palaeocene clement may reflect downo-hole

contaminarior.

Since the earlier paper was published, fay ther material

from ihe region has come to hand and fias been examined.

Bore RN 17577 (lat. 25° 21° 30° S, Jong. 191° 03° OO"L)

yielded a-cathonaceous sequence between 67 and 84 in,

Palynomorph assemblages recovered trom this interval are

of Late boven age (Upper Nothofagidites asperis 7o0e

equivalent) indicuted by the presence of

Malvaccarumpollix si. Quintinia sp. GYraslenon sp,

an ubundanee of conifer pollen und a low diversity of

“proteueeous™ pollen.

This evidence, together with thar of dhe oriwitad samples,

indicates a vomples Late Cretaceous and Tertiary

depositional history for che valley between Ayers Rock and

Lhe-Oluas. A similar history occurs in tlre adjacem Lake

Eyre Basin’ and it is tetipling lo relate these: seemingly

sporadic Uuepusitiodal episodes within the craton om

regional warping, such as resulted in the disruption of

drainage anid inthe formation ot the Amadeus and otter

basins of imernal drainage in Central Australia.

Thus we conclude that there were ap leasr thee

depositional phases in the area: vie oy Late Cretaceous,

a possible mid ta lare Palaeucence and a fate Eocene

sequence, The earlier phase implies a sliettty older aue

~ by seme 5-10 My for the ancestral Ayers Rock and

the Olgas than was previously deduced. Ewen this slightly

older age isa Minin age, The bevelled upper suurfices

af Ayers Rock ind of the varidus domes of the Olgas

complex mitv well nurn our to be part of the even older

Larly Cretaceous or even Late Jurassic surface of which

remnanes remain we the MacDonnell, Flinders and Crawler

Fafaes, did ppan which impacted the neteweie responsible

for Gosses Bluth’.

bale C.F & ortridge, AL 1K (1984) Molinelann 8.

139-144,

"Helby, R., Morgan, R, & Partridge, AL 1) (1984) afew,

Jlysoe. otestralas, Poldeant, 4, 179.

“Woplner, Ho Callen. KR. AL & Parris, WOK. INT)

thee! Suc. Avs. 2), 17 52,

“Milton. D. 3., Barlow, HL C2, Kroji, BR, Brown. A, K.,

Glikson, A. ¥., Manwaring, B A., Mass, 8, 4, Sedmik,

Eo E., Van Son, 1, Young. G. A, (19%) Sener 179,

1149-1307,

WOR, HARRIS, Selinal ol Apphed Scinee, Uiriversity College of Southern Queensland, laowyormbu 4340, and

CR TWIDALE, Deparitvent of Geology and Geophysics, University of Adelaide, Adelaide 5006,

VOL. 115, PARTS 3 & 4

29 NOVEMBER, 1991

Transactions of the

Royal Society of South

Australia

Incorporated

Contents

Koste, W. & Shiel, R. J. Rotifera from Australian inland waters. VII. Notommatidae (Rotifera:

Monogononta) - - - - = - - - - - -

Cann, J. H., De Deckker, P. & Murray-Wallace, C. V. Coastal aboriginal shell middens

and their palaeoenvironmental significance, Robe Range, South Australia

Zeidler, W. A new genus and species of phreatic amphipod (Crustacea: Amphipoda) belonging

in the “chiltonia” generic group, from Dalhousie Springs, South Australia

Lee, D. C. & Birchby, C. M. Paraphauloppia (Acarida: Cryptostigmata: Cabaimhaes) and

its occurrence in South Australian soils — - - - -

McEntee, J. C. Lake Frome (South Australia) aboriginal trails - - - - -

Southcott, R. V. A new trombellid mite (Acarina: Trombellidae) from South Australia-

Obendorf, D. L., Beveridge, I. & Andrews, R. H. Cryptic species in populations of

Globocephaloides trifidospicularis Kung (Nematoda:

Trichostrongyloidea), parasitic in macropodid marsupials - - -

Smales, L. R. A new species of Antechiniella Quentin & Beveridge, 1986 (Nematoda:

Acuariidae) from the Australian water rat, Hydromys chrysogaster

Geoffroy, 1804. - - - - - - - - - - -

PUBLISHED AND SOLD AT THE SOCIETY’S ROOMS

SOUTH AUSTRALIAN MUSEUM, NORTH TERRACE, ADELAIDE, S.A. 5000

lll

161

177

189

199

207

213

217

TRANSACTIONS OF THE

ROYAL SOCIETY

OF SOUTH AUSTRALIA

INCORPORATED

VOL. 115. PART 3

ROTIFERA FROM AUSTRALIAN INLAND WATERS. VII.

NOTOMMATIDAE (ROTIFERA: MONOGONONTA)

BY W. KOSTE* & R. J. SHIELT

Summary

Keys are given to 14 genera and 70 species of the Rotifera: Monogononta in the family

Notommatidae presently recorded from Australian inland waters. Available distribution data and

ecological information are given for Cephalodella (26 species), Dorystoma (1), Eosphora (5),

Eothinia (1), Itura (3), Monommata (11), Notommata (14). Pleurotrocha (1), Resticula (3),

Scaridium (1), Taphrocampa (2) and Tetrasiphon (1). Drilophaga is recorded here for the first time.

A record of Rousseletia is incertae sedis. The genera Metadiaschiza, Pseudoharringia,

Pleurotrochopsis, Sphyrias and Tylotrocha are not recorded from the continent.

KEY WORDS: Rotifera, Australia, taxonomic revision, Notommatidae, Cephalodella, Dorystoma,

Drilophaga, Eosphora, Eothinia, Itura, Monommata, Notommata, Pleurotrocha, Resticula,

Rousseletia, Scandium, Taphrocampa, Tetrasiphon

Transaenons of the Reval Noelety OF S. Aust, (1991), TESCR), T1150.

ROTIFERA FROM AUSTRALIAN INLAND WATERS. VII, NOTOMMATIDAE

(ROTIFERA: MONOGONONTA)

by W. Kosre* & R, J. SHIELT

Summary

Kusre, W. & Stile, R, J. (1991) Rotifera from Australian infand waters Vil, Norommatidae (Rotifera:

Monogononta) Trans. R. Sac. 5. Aust, TESG), 11-159, 29 November, 1991,

Keys are given to 14 genera and 70 species of the Rotifera-Monogononta in the family Notonmmardae

presently recorded from Australian inland waters. Available distribution data and ecological information

are piven for Cephilodella (26 species), Darystoma (1), Eosphara (5), Eothinia (1), ftura (3), Manammata

(11), Netommala (14), Pleurorrocha (\), Restioula (3), Scaridium (1), Taphrocampa (2) and Terrasiphon

(1), Drilophaga js recorded here for ihe first time, A record of Rousseletia is incertae sedis. The genera

Metadiaschiza, Pseudoharringia, Pleurotrochopsis, Sphyrias and Tylofrocha are not recorded from the

continent.

; Koy Worps: Rotifera, Australia, taxonomic revision, Notommatidae, Cephulodella, Barystania,

Drilophaga, Eosphara, Eorhinia, Vura, Monommata, Notommata, Pleuroitocka, Resticula, Rousselelia,

Seurtdivit, Taphrovampa, Terrasiphan

Introduction

The purpase of our series of papers has been to

document the rotifers recorded from Australia,

primarily to provide usable keys ta them. We have

demonstrated the existence af a diverse Australian

rotifer fauna distinct from that of Europe or North

America, with >S50% of recorded species in some

genera endemic. Also, there is marked latitudinal

and longitudinal variation in species assemblages,

Taxonomic resolution of this component of aquatic

systems permits more informed understanding of

community ecology in inland waters and, thereby,

betler ifterpretation of environmental impacts on

water quality,

The taxonomy of the rotifer families we have

reviewed to date (see Koste & Shic! 1990b) has been

reasonably straightforward, The Notommatidae, in

contrasl, appears to be the repository of everything

not readily placed elsewhere. In their revision of the

Wisconsin notommands, Harring & Myers (1924)

noted of the family that “there has been a steady

accretion of new species and a periodic shifting of

the old ones, until it has become a veritable

Serbonian bog, carefully avoided by everybody or,

al least, trespassed upon only under compulsion”.

li seems that time has only compounded the

confusion. In the ensuing years same efforts were

made to resolve loosely affiliated taxa by erectian

of new families, eg, Linditdae, Dicranophoridae,

particularly by Remane (1933). Notwithstanding,

the Notonimatidae still contains 21 genera In two

* Ludwie-Brill-Srrasse 5, Quakenbriick D-4570, Federal

Republic af Germany,

{ Murray-Darling Freshwater Res. Centre, P.O. Bax 921,

Albury, N.S.\W. 2040.

subfamilies (Koste 1978), with one-third of the

genera monospecific, and an extraordinary

profusion of ‘species’ in others [>200 in

Cephalodella (Rutiner-Kolisko 1974)]. A systematic

revision of the family is needed urgently. Given

recent technological advances, particularly scanning

electron microscapy (SEM) (cf. Koste & Shiel

19904), resolution of the group js feasible.

Natommiatid rotifers are predominantly littoral

(epiphytic or epibenthic) in habit, collected in and

around vegetation in shallow. waters oF lake or river

margins, in billabongs, and in the roots or scales

of floating macrophyte mats. Most are grazers,

feeding on detritus, bacteria and epiphytic algae,

particularly diatoms. Some taxa are omnivores,

taking protozoans. and other rotifers, and several

species. are parasitic an worms or leeches (Pourriol

1965; Koste 1978),

General problems associated with rotifer

systematics are more pronounced in the

Notommatidae, especially those arising from study

af preserved material, sometimes long alter

collection (cf, Berzins 1982). Animals may be

strongly contracted in preservatives, artefacts are

produced by distortion, colours of organelles may

fade, etc, leading to erroneous observations,

incorrect measurements and misidenti fications, For

these reasons we consider that some of the

notommatid taxa reported from Australia probably

do not occur here. They may be good endemic

species named for the European taxa they most

closely resemble, or a known species wrongly

identified.

To minimise erroneous identifications, we suggest

thal live material be examined wherever possible,

followed by critical examination of miustax

t2 W. KOSTE & R. J. SHIEL

morphology, particularly the species-specific

sclerotised /rophi (Fig, 2), Extensive use of trophi

morphology has been made in Ruropean revisions.

General trophi structure was reviewed in Koste &

Shiel (1987a), but in view of the difficulty of

working with these small structures (some less (han

20 jam), we attempted in earlier parls to produce

keys to species based on morphometry, ¢.z. body

y. toe length, claw:toe ratios, etc. Unlike most

metazoans, rotifers are eutelic (cell numbers

constant between generations) with little

intraspecific morphological variation, hence

comparalive measurements of body parts can be

used diagnostically. In the Notommatidae, close

examination of the trophi is necessary to prevent

confusion of closely allied taxa or juveniles of large

species with adults of smaller forms.

In this review the format of earlier parts is

followed; for convenience, genera and species are

treated alphabetically. Keys to roufer families are

included in Koste & Shiel (1987a), which also

contains brief descriptions of general morphology,

Known distribution and ecological information are

given for the species we have encountered, Available

type locality and holotype information is included.

Where type locality is not known, probable place

of origin is given in parentheses; some early authors

did not specify origin of material, however we

consider if likely that in the late 18th-early 19th

century their collections derived from proximal

localities.

Methods

Live animals can be collected with floating or

submerged macrophytes from most standing waters,

Stems of Fallisneria, Eleocharis, Myriophyllum,

and other submergents provide rich rotifer

assemblages. The floating liverwort, Rieciocarpus,

or the fern, Azolla, usually have diverse rotifer

faunas associated with their submerged parts,

Fig. |. Low vacuum system for mounting rotifers and

clearing trophi onto a Nuclepore filter

Whole plants or segments can be examined under

LP microscopy and resident rotifers removed by fine

pipette onto glass slides for HP microscopy.

Animals can be restrained by light pressure of a

coverslip (supported on plasticine “feet” or coverslip

fragments), or In a purpose-built compression

chamber (Martin 1986); all measurements of body

morphology can be made on uncontracted

individuals, If live material is not available, recently-

preserved is preferable to long-preserved,

When all body measurements are taken,

preparation of trophi for light microscopy should

be made by clearing the animal(s) in sodium

hypochlorite; at least several preparations should

be made to permit interpretation of the orientation

of the minute components. A drop of bleach

solution placed beside the coverslip is drawn

underneath it by carefully touching lens tissue to

the opposite side, The clearing animal should be

in view during the process because rapid flow of

the bleach may move it or the trophi, and the

preparalion will be lost. A microscope-mounted

video camera is a useful accessory for recording

both whole-animal and trophi morphology during

this procedure,

Fig. 2. Trophi of Notommata copeus from R. Murray

waters, Barmah Forest. F = fulerum, M. =

manubrium, R = Ramus, U ~ uncus, Cambridge S600

Stereascan. Seale line 20 um.

ROTIFERA FROM AUSTRALIAN INLAND WATERS Ils

Treatment of trophi for SEM will depend on their

robustness; some larger trophi can simply be

extracted from cleared animals by micropipette,

rinsed through a graded ethanol series and pipetted

from the final 100% ethanol or acetone onto a SEM

stub (cf, Fig, 1). For more delicate trophi, the system

shown in Fig. 2 was adapted from Markevitch &

Koreneva (1981). Rotifers removed from field

collections are rinsed through filtered water,

Fig. 3. Jétrasiphon hydrocora Ehrenberu: (a) dorsal, swimming (g = glands); (b) lateral, swimming; (c) trophi; td)

vesting egg. Scale lines: a, b, d 100 pm; c 20 wm. After Koste (1978).

pipetted in a small drop of water onto a Nuclepore

membrane and treated with sodium hypochlorite

for 5-10 min. Low vacuum is then applied to remove

the hypochlorite solution, the cleared trophi are

rinsed gently with distilled water, and the filter is

removed and dried over silica gel, Critical point

drying is not necessary, If sufficient numbers of

animals are used, standard sputter coating with

gold/palladium and examination under SEM

It W. KOSTE & R. J. SHIEL

enables detailed resolution of trophi in different

orientations. An alternative method detailed by

Kleinow ef a/. (1990) permitted high-resolution

micrographs of undistorted trophi preparations of

a brachionid species, Brachionus plicatilis, but has

not yet been used for rotifers with more complex

trophi, Further details of trophi ultrastructure are

given by Markevitch & Kutikova (1989),

SYSTEMATICS

Family Notommatidae Remane

The charactensties of (he family were described

by, infer alia, Harring & Myers (1924), Remane

(1933) and Koste (1978), It is a diverse assemblage

of illoricate or partly loricate taxa comprising 1wo

subfamilies: Tetrasiphoninae (two genera) and

Notommatinae (19 genera) separated on the basis

ol presence (Tetrasiphoninae) or absence

(Nolammatinae) of a whorl of bulbous glands

between the stomach and intestine.

Subfamily Tetrasiphoninae

Ol two described genera, Repaulina and

Tetrasiphon, only Tetrasiphon is known from

Australia.

Genus Terrasiphon Ehrenbers

Tetrasiphion Ehrenberg, 1840, p. 219. Monolypic genus.

Type: Tetrasiphon hydrecora Ehrenberg, 1840

p. 219.

Tetrasiphon hydrocora Ehrenberg

FIGs 3, 4

Syn. ?Repaulina dicerea Berzins, 1960, pp, I-3.

Type locality; Berlin,

Holotype: Not designated.

Description: Body illoricate, cylindrical 10 fusiform;

no annular ring separating head and trunk, cuticle

somewhat stiff; abdomen ends in short stumpy

projection over cloaca; foot short, bi-segmented,

with lwo acutely pointed, elongaled toes; corona

oblique, extended ventrally to. elongated ‘chin’

tentaculate dorsal paired lateral antennae; lateral

antennae towards posterior end of abdomen

similarly elongate, with exceptionally long sensory

setae; single cerebral eye; trophi with single toothed

unci; manubria with complex projections (Fig. 3c);

rami long, lyrate, curved dorsally, with pointed

alulae; large hypopharynx muscle inserted in mastax

wall; adult animal commonly in yellowish

gelatinous sheath.

Length 450-1000 pm, toes 60-80 ym; subitancous

Fig. 4. Tétrasiphon hydrocora grazing on Pleurotaenium.

Tallandoon billabong, Mitta Mitta River, Vic. Kodak

T-max, 1/30 seu,

egg 140-154x110-L15 pm, with curved spinules 45-65

pn long; resting egg 200 x 155 um; male to 300 pm;

male egg 102-126% 88-92 pm.

Ecology: In Sphagnum pools, acid waters in Europe,

N, and §, America; billabongs of upper Murray

tributaries, N.S.W., Vie.) dune lakes in Tasmania,

Specialist grazer on large desmids, e, Cosmarium,

Micrasterias, Pleurotaenium (Fig. 4), Staurastrum.

During filmed feeding experiments, an individual

from a billabong on the Mitta Mitta River at

Tallandoon, Vic, ingested 10 Staurastrum in 30 min.

As the cells passed along the gut they gradually lost

colour, the semicell branches were fractured by

muscular action, and the fragments were egested-

Literature: Pourriot 1965; Koste 1968, 1978.

Subfamily Notommatinae

The subfamily has 19 named genera, 13 of which

are known from Australia. Metudiaschiga Fadcev

(Fig. S:1), Pleurotrachopsyis Berzins (Fig, 5:2),

Pseudoharringia Fadeev (Pig. 5:3), Sphyrias

Harring (Fig. 5:4) and Z/otrecha Harring & Myers

(Fig. 5:5) are not presently recorded here. For

further information on them, see Koste (1978).

Drilophaga is a new record, reported here for the

first time. A single report af Rousseletia is

considered incertae sedis.

ROTIPFERA FROM ALSTRALIAN INLAND WATERS hs

’, T ~

Ty)

ede

aM) .

“20° 4

~ oq ‘Qe

Sor =)

a id ay

an and

z 4

a Ay

ae:

Peal

es ete

5a \

Fiz. 5, 1, Metadiaschiza Fadeev: (a) lateral; (b) dorsal; (c) trophi ventral; (d) trophi, lateral. 2, Plewrotrochopsis Berzins:

(a) lateral: (b) posterior abdomen and toes, dorsal; (c) trophi, ventral; (d) distal end of fulerum, (e) ventral cuticular

‘jamella’ with hooks. 3. Pseudoharringia Fadeevy: (a) lateral; (b) trophi, ventral; (c) fulcrum and rami. 4, SpAyrias:

Harring: (2) dorsal; (b) trophi, lateral; (c) trophi ventral; (d) trophi, apical; (e) unci, lateral: (f) resting cee. 5, Dlotrocha

Harring & Myers: (a) lateral; (b) dorsal; (c) trophi, ventral; (4) trophi, Jateral, 1 after Wulfert (1937), 2 after Berzins

(1973), 3 after Kutikova (1970), 4 after Koste (1978), 5 after Harring & Myers (1922). Seale lines: adults 50 pry

trophi 10 zm;

Key ta genera of the subfamily Notommatinae

1. Corona o1cylindrical extrusion/evagination, with

cirtumapical ciliation (Fig. 14a); annular adhesive

organ present; no lateral ciliary auricles; mouth

deeply invaginated.,... Prilophaga Vejdovsky

(Fig. 14)

Corona not extruded, may he frontal, oblique or

extending ventrally; no adhesive organ; ciliary

auricles may be present; mouth not deeply

invaginated ..,,.....4-. tedteeeelaiet ss garts me

20). ViteNanum band or ribbon-shaped........, 3

Vitellarium aval or kidney-shaped..... 5.55.5

(2). Nuclei arranged linearly, 22.0... .5 0 -2e eve 4

Nuclei irregularly distributed, .....Fnteropleu

Ehrenberg (Fig. 15:2)

4(3), Eyeless; foot two to three-segmented.........

Lb gate weyb are Psevdohartingia Fadeev (Fig. 15:3)

Two frontal eyes on papillae; foot one-segmented

with annuli......-Sphyrias Harring (Fig. 5:4)

5(2).

(5).

10(8).

Foot and toes Jonger than body... .--. 6

Foot and toes shorter than body... .----- 7

Toes of dissimilar length; toot shart, mostly 2-

rarely three-segmented, .,Moneommiata Bartsch

(Figs 19, 20)

Toes of similar length; foot 3-segmented and very

long, »...4.-- Scaridiven Etrenberg (Fig. 28:1)

Poot with single log... . Tletroche Harring &

’ Myers (hig. 5:5)

Foor with two toes. 2.0.0... 0. 2. eee eee 8

Rump or last foot segment wilh spine,...,.. 9

Rumip or last fool segment without spine... 1)

Rump with curved spine............-...---

voc. Dorvsromm Warring & Myers (Fig. 1571)

Foot-end with short spiné.... .... Rowsseletia

Harring (Pig. 29)

Trunk lorivate with 3-5 species-specific vuticulay

plateye.. .-.o--s essen eraemidaatie

Trunk illoricate,,.----. 2-2-2, eras Y

Il W, KROSTE & R. 1. SHIEL

1i(10), 2) dorsolateral plates; 4

playin

praniianad vetilral

( ..... Cephetadella

Bory de St Vincent (Figs 7.03)

2 dorsoventral, 2 (er 5) venwrolateal plates, alsu

J smuoth trunk plates present... 20.

--- 4), Metudiasehiza Fadeey (Fig. 4: l)

Cutiele with rows of tiny hooks...

by opens ca Pleuratrochopsts Berains Kix, $:2)

1210),

Chilivle lackiie HOOKS... eee 3

(4012), Trunk with conspicuous ‘annul. : Tapnroaritpe

Oosse (Fig, 28:2, 3)

AAU AOBERL. be a eked eee eee sa.l4

1443). One cerebral eye und two widely-separated frontal

VCR ep is

Cerebral eye absent (or if present, no fromial. eves

WH SHOVE)... cee eee . ame 17

[Sqld), Stomach with blind sacs. 5 frura

Harring & Myers (Fig. 18)

Stomach within blind saes —, = 16

L6(15) Masrax with single salivary bland, .. -Enthinia

Harring & Myers (Fig, 17:2)

Muastas with paired salivary elands, Sosphera

Ehrenberg (Pigs 16, 13:1)

Wud). Coruna displaced ventrally; aillaey aurieles

generally presen hi cpt § Netomputa

Ehrenberg (higs 2!-25)

Corona wulerior, TO auricles presene . 8, 18

Salivary glands syiminetrical.,, ..Plewratrocha

herzins (Fig, 26:1)

Salivury glands asvymmenical or rudimentary. .

» Resticula Warring & Myers (Pigs 27:24)

inhdth

Genhs Cephalpdelle Bory de St Vincent

Crphalodella Bory de St Vilwent, 1826, p. 43,

Tipe: Cercaria cutellina Miller, 1786, p. 130. =

Cephalodetla cetetling (Miiller).

Type levality; Copenhagen,

Fusiform notomniaud rotifers of varicus shapes,

from clongate to short and stumpy; occasionally

iNaricate, but mostly with one or more lorica plates,

position of whicl) varies according ta species; shehi

constriction between fread and trunk, none between

trunk and short foot, which bears two toes; in

loricate taxa, dorsal and fiateral sulci distiner

between plates; corona frontal, oblique, with long

marginal cilia and two Jateral tufts of long

swimming cilla; buccal fleld lightly ciliated; mastax

virgale, wilh long straight fulcrum, poorly

developed rami; retrocerebral organ rudimentary or

absent; eyespol cervicul, single or paired frontal, or

absent.

Of > 2) Cephalodella species worldwide, Koste

(1978) described 132 taxa from Europe, Twenty-Four

Ol these, and two endemies, are known trom

Australia, Othet taxa (especially numing dubia) are

giver by Harring & Myers (1924), who also discuss

the confused generic nometelaruce.

Trop) marphelagy: In {he descriptive section

below, we recognize the six traphi types deseribed

by Wulfert (1937). In descriptions of trophl

Structures, we use ‘Proximal’ to refer to the head or

anterior end and ‘distal’ to the tail oF posterior end,

‘Basal’ as used by Wullert implies proximal.

Type A (Fig, 601): fulerum spatulate distally; rami

single, without teeth on inner margin; manubria

slender, rodlike, curving inwards in top view, from

straight shaft, no basal lametlae or dist! dilation

of manubria, which form characteristic creseentic

shape when closed.

Type B (Fig. 6:2): falcrum as Type A; inner margin

of rami toothed or striated, at times with alulac;

manubna with single or bilateral basal lamellae,

distally Teshaped (termed ‘double-crooked’ by

Wulfert, referring to a curved shepherd’s ‘crook’,

This implement, and the term, no longer seem 10

be in common usage). Several species (a2. C. eva)

have 4 spatulate dilation of the manubria ends

rather than a free-standing T, but in all other

features conform to Type B trophi,

Type C (Fig. 6:3): features distinctive ringlike

fenestrations at distal ends of manubria, considered

by Wullert to be derived From double-cruoked ‘Type

B trophi.

Type D (Fig. 6:4): is most complex, with trophi parts

not found in other types. Fulcrum short, dilared

distally talsu in lateral view), narrower in niiddle;

rami from above widely separated, with comblike

teeth on forcipate tips (absent in srenroosi);

manubria proximally with wide bilateral lamellae,

distally with single abrupt inward curve or crook;

behind basal expansions, branehed structures

(subunci) occur; unci single, offen with dorsal plate:

In some species (orficula, gigantea, tenuiseta), a

large delicate l'rontal plate with denticulate margin

oceurs above rami,

Type (Fig. 6:5); known only in C! mezalocephuler,

Fulcrum not dilated; rami right-angled dorsally

(visible in lateral view), cloned distally (at fulerum)

and separating proximally; basal Jamellac of thin,

S-shaped manubria apparently separate.

fype V (Mig, 6:6): recorded only in-€. ynira, whieh

is mot known trom Australia, This tophus is

comprised entirely of delicate rods.

Key to species of Cephilodetla revorded from

Australian inland waters

1. Ratio rotal length/me length <4. ..,, 0, \,

Ratio tora! lengih/loe Jength >3

With single or double cerebral eye, -

Fycless.. eee

32). Toes curved. a doa, distinctly segmented, ...

>) tantillvides Hauer (Fig. 13:1)

- '

ROTIFERA FROM AUSTRALIAN INLAND WATERS 0

6b 5a

Fig. 6. Trophi types recognized in Cephalodella species. 1, Type A? (a) ventral; (b) lateral; (¢, d) vartant. 2, Type B:

(a) ventral; (b) lateral; (c, d) variant, 3, Type C: (a) ventral; (b) lateral, 4, Type D: (a) ventral; (b) lateral, (c) uncis,

lateral. 5, Type E: (a) ventral; (b) lateral. 6, Type F: (a) ventral; (b) lateral. After Wulfert (1937). Scale lines 10 ym.

4(2).

Toes sigmoid in lateral view, not segmented...

Oe eee C. nana Myers (Fig. 12:4)

Abdomen with hooked caudal projection.....

tet thw, ae C. mucronata Myers (Fig. 11:4)

Without hooked caudal projection..........,

wtpbt dL i oteae lat C. biungulata Wulfert (Fig, 7:3)

Ratio total length/toe length 3-5.........., 6

Ratio total length/toe length >5...---...- 21

. Toe tips with longitudinal denticle row (2-4) on

functionally ventral side........ C. lindamava

Koste & Shiel (Fig. 11:1).

Toe tips without ventral denticles, bul may haye

median dorsal denticles.......)-.00-+-51-- 7

7(6).

8(7).

9(8).

10(9).

Body >300 pm...... ccc eee eee eee eee 18

Body <300 pine...) 0... cee eee 8

Body >90 pm, toes >20 pm. ...........--9

Body <90 ym, toes $20 pm,......C. gisleni

Berzins (Fig, 10:2)

Toes (straight or curved) taper evenly from base

segmentation. - 2.20 022s eee eee 10

Tips of toes sickle-shaped, recurved..........

Ly salto get imets C, apacolea Myers (Fig. 7:1)

[Ik

11e9),

QUT),

1311).

1413).

15(13).

16(15).

741.

18(7).

19(18).

2018).

212},

22(21).

23(22).

25424).

26(25),

W KOSTE & RB. J. SHIEL

Distal 'a of toes demarcated hy transverse Seprun

into segmented tip. ee Co terute,

Myers (Fig 10:5)

locs >60 pm. pee ae Se 12

Toes <A) pine... eee, ca 13

Trophi > 70 pms cvves ea ella

(Bhrenbere) (Fig. 9:6)

Trophi ea, 30 pot... 2. eee {~ tinea

Wullert nal 13: ‘ye

THOS FD GO AT eee

Toes 230 pm... eee : "4

Body <127 pm; toes 20-26 win trupli <wW an

-C_ exigua (Close) (Fig. 9:3)

Body >125 ym; tees 25 28 pm; trophi 30-34 pm

$A Ben C. venitipes Dixon-Nuttall (Fig. 13:4)

Trophi <30 pm... o eee cexicicik as

Trophi >30 jum... 6... ee - 7

Paired eyespots with -cryssalling lens: toes > 143

hody length, .C. miszuraus Wulfert (Fig. 14:3)

No evespots; toes < 1/3 body lengrh —_.

iti hpetettee C, farficate (Ehrenberg) (Fig. 9:4)

Single evespot al posterior end of ganvlion; corona

with prominent tips; manubria nor crooked. . .

we aphtctt¢ Pa Sutee o5 C. hoodi (Gosse) (Fig, W4)

Paired frontal eyespots in single capsule; corona

without prominent lips; manubria crooked. .

estat wetgye tts C, sterea hanes ore 12 4)

Toes > 100 win, , f Sits, SY

Toes <100 mim... 0... eee 20

Toes ca. 1/3 body length; trophi >70 pm.

prea C, gibba (Ehrenbery) (Tig, 10: 1)

Toes L/6 body length; trophi <70 pm.......,

Spatittione stutt ge C. panarista Myers Wig. 12:2)

Distinet ‘cycapat: wes 1/5 body length

i © forfienty (Fhrenberet [Fig 9:5)

No Evespot, lacs “body lengifh......) 0.2...

AC. fenuisera (Burn) (Fig. 13:2)

ote stele WIT y wm elelels fyi) de

.C. culellinu

“(Muiller) (Figs 7:4, 8)

Body SIM pM cece eee eee ee 23

Body <I) panics iia teceveee cei es 24

(NB: Occasionally individuals of C. parasitica may

exceed 190 pm: see species determinanon.)

Toes >50 pin (at least 1/3 body lengih),.....

AAC Eee oe C. ever (Gosse) (ig. 9:2)

Toes <50 nm, 1/0 body length.......

Eyespot(s) visible, coloured or colourless. ..25

No evespol, , .C. parasineu (Jeunings)

(Fig. 12:3)

‘Two verebral cyespats; lorica keeled in dorsal 1/3

‘ .C, euderby/ Walfert (Fig, 9:1)

Single. evespor, Coroufless or coloured; no dorsal

keel on posterior lorica......,....--..--- 24

Lyespot reddish; traphi >30 pm.

- 11 & auriculata (Miller) (Tig. 7: 2)

Eyespot ‘colourless: trophi <30 pm..........

san sihih at CG gracilis (Ehrenbera) (Fig, 10y3)

Toes ‘>20 am

Toes <20 yt,

Cephalodella apecelea Harring & Myers

FIG, #1

Cephaladella anocolea Harring & Myers, 1924, p. 509,

bigs 33:1, 2.

Dype locality. Not specified,“ . commen in weedy

ponds and bogs.” Vilas and Oneida Counties,

Wisconsin, are first localities mentioned by Harring

and Myers.

Holotype: Not designated, ? Myers Collection,

American Museum of Natural History (AMNH),

New York,

Pescription: Body wransparent, clongate, slightly

compressed laterally; plates obvious, with distingr

lateral sulci; foot large, robust, with small

protruding tail; foot glands large, pyciform) tues set

wide apart, straight, ending in recurved, siokle-

shaped tips (toes may be crossed in swimming

animal); corona oblique, convex, without

protruding lips; trophi modified type A, slender,

with delicate; curved manubria; fulcrum with slight

terminal bend; eyespor absent,

Total Jength 125-185 pm; twophi 29-33 am

(fulcrum 12-20 pm, manubria 20 pm); toes 32-58

pln.

Ecology: In acid-neutral waters on submerged

plants, in periphyton, where it feeds on diatoms and

unicellular algae. Europe, N, America, New

Zealand, pH tolerant. Single record: Magels Creek,

NT, (billabong not named, possibly Mine Valley).

Literature: Koste 1981,

Cephalodella auriculata (Miiller)

FIGj, 7:2

Farricella avriculata Miitler, 1773, ys. LLL,

- ceri uel auriculata: Warring. & Myers, 1924, p. 479,

hss

For extensive synonymy see Rutikovy (1970: 240), Koste

(hO7R: 366),

Tepe locality: Copenhagen,

Holotype: Not designated.

Description: Body short, stout; head conspicuously

wider than trunk, with small rostrum; lorics rigid,

plates distinct; foot very short, toes shart (<1/4

lenigth); caudal antennae setae long; mastiax large

with two round, clear salivary plands; trophi type

A, with small, thin mallei, slender recurved

manubria, fulcrum long, expanded distally.

Resembles C. venrripes, but has single red cervical

eyespot,

Total length 120-160 pm; trophi 36 jan: toes 22-28

am; male 95 jum.

Ecology: Cosmopolitan in beach sand, in littoral

of stilland flowing waters, where its main food is

phytotlagellates and detritus. Qld, NUT, Tas, Vic.

common in 'R, Murray billabongs. 16.0-22C, pH

6,4-7,3, 37.3-274 2S cm.

Literature: Colledge 1914; Koste 198]; Berzins 1982;

Koste e/ al. 1988,

ROTIFERA FROM AUSTRALIAN INLAND WATERS We

Fig. 7. 1, Cephalodella apoealea Myers: (a) lateral; (b) dorsal; (c) trophi, lateral; (d) toe tips, lateral; (¢) fulcrum,

lateral, 2, C auriciilaia (Muller): (a) dorsal; (b) lateral; for trophi see Fig, 3:Ja, b. 3, C biunagulata Wulfert: (a)

lateral; (b) toe tip, lateral; (ey trophi, ventral; (d) trophi, lateral; (¢) uncus; (f-h) views of ramus; (i) toes, ventral,

4..C. catellina {Miiller): (a) lateral; (b) trophi, ventral; (c) trophi lateral; (d) trophi of C. armata Rudescu. 1, 2

after Wullert (1940); 3 after Wulfert (1937); 4a-c aller Harring & Myers (1924); 44 after Rodewald-Rudescu (1960).

Seale lines: adults 50 am; traphi 10 pm.

120 W. KOSTE & BR. |. SHIFI

Cephalodella biungulata Wylfert

FIG, 7:3

Peg a eae binegulaia Wulfert, 137, p. G17 48, Fie,

Tipe /ocatity. (Germany).

Holotype: Not designated,

Pescription: Body hyaline, oval, dorsally arched;

head short, slightly oblique; abdomen projects over

short foor, both covered by dorsal plates; toes long,

flexible, >1/3 total length, with distinctive bifurcate

lips due to presence of spinule at distal end; mastax

lacks salivary glands; trophi of C. gibba type (B),

with symmetrical rami denticulate on inner margin;

manubria doublecrooked, with bilateral proximal

lamellac (Fig. 7:3c); shaft of uncus with semicircular

lamella. Eycx absent. Close to € gibha,

Distinguished from it by the lack of eyes,

characteristic longer bifurcate toes and iophi

differences,

Toral length 250-313 jum; toes 88-112 pm; ophr

40-34 pin.

Exology: Rare in littoral/moss of pools and sireams

Pig. 8. Ceptaladella catelling (Muller), (a) lateral, ih)

seennu individual, lateral; (wv) mule, laterals Gt) trophi,

lateral; (¢)trophi, ventral: (f) distal end of manubrium.

Koste, orig. Scale linesy adul $0 pm, trophi 1 yn

in Europe. Several individuals in a collection from

the filling Dartmouth reservoir in (978, probably

incursions. from a.submerged littoral habitat, Not

collected subsequently.

Cephaledetle eateltinag (Miilter)

FIGS 7:4, §

Cercaria catelling Muller, 1786, p, 130, Fig. 20;12)13,

Cephalodelta cateltina: Bary de St Vincent, 1826, p. 43,

See Harring & Myers (1924, p. L83-184) and Koste

(1978, p. 371) for extensive synonymy.

Wpre locality; Caopenhapen.

Holotype: Not designated.

Description: Body short, stout, bulbous posteriorly;

abdomen laterally compressed, with wide lateral

sulci separating indistinet lorica plates; foot short,

ventral, beneath overhanging ‘tail’: toes short,

approx, 1/10 body length, thin to conical; mastax

with ventral salivary glands (may be absent in small

specimens); trophi type C, with long. filerum

slightly expanded distally; manubria rod-shaped,

decurved, ending in semicireular dorsal expansion

(Fig. 81, Hooked manubria ends may result {rom

hypochlorite digestion (cf, Fig, 7:dd) (Kaste 1978);

twa separate red frontal ¢yespots,

Total length 80-160 pm; toes 9-14 pm (18-20 ~m

in Harring & Myers 1924); trophi 27 am (45 pn iin

Harring & Myers); male 140 xm, toes 17 um.

Ecology: Cosmopolitan in fresh to brackish water.

i occasionally reported in coastal (marine) waters.

| N.SMW.,, Tas., Vic. 12,5-23,5"C, pH 4.4-6.8, 69,5-600

ws cm, 2,9-300 NTU. A parasite of bolvex

colonies (Europe and North America) is referred

to ©. eatellina, C. catellina volvacicole

(Zawadowsky). It is not recorded from Australia,

Literrture: Bvans 1951; Shiel & Kaste 1979: Kaste

& Shiel 1987b,

Cephalodella euderbyi Wulfert

FIG, 9;

Cephalodella euderbyi Wullert, 1940, p. 564, Fig, 4,

Wpe locality: Birkhorster Moor (eastern Germany}.

Holotype: Not designated.

Description: Body stout, lightly arched dorsally;

head approx. 1/3 body length; corona with slightly

protruding lips; posterior dorsal lorica compressed,

resembles keel; foot short, almost covered by

pointed ‘tail’; toes short, approx. 1/5 total length,

thin, straight, occasionally slightly recurved; mastax

large, with distisict salivary glands; trophi of type

A - fulcrum spatulate distally; manubria thin,

rodlike, with single crook distally; rami single, with

very small alula teethy rwo small ruby-red cerebral

CyeSpors.

ROTIFERA FROM AUSTRALIAN INLAND WATERS I2I

Fig. 9, |, Cephalodella euderhyi Wulfert: (a) lateral; (b) trophi, ventral; (c) trophi, lateral. 2, CG eva {Gosse): (a) lateral:

(b} trophi, ventral; {c) trophi, lateral. 3, C. exigwa (Gosse): (a) lateral; (b) trophi, ventral; (c) trophi, lateral. 4, C

Jorficata (Ehrenberg): {a) lateral; (b) trophi, ventral; (c) manubrium, lateral; (d) fulcrum, lateral; (e) uncus; (f) ramus.

5, C. _forfieula (Ebrenberg): (a) lateral; (b) trophi, ventral; (c) trophi, lateral, 6, @. gibba Ehrenberg: (a) lateral;

(b) trophi, ventral; {c) trophi, Iateral, 1, 2b, c, 3c afrer Wulfert (1940); 2a, 3a, 4a, 5 afler Harriny & Myers (1924);

db, 4c-f after Donner (1949); 46 after Wiszniewski (1936); 6 after Wulfert (1937). Scale lines: adults 50 zm; trophi

10 ym.

Total length 150-168 pm; trophi 32 »m; toes 28

pm; male 110 pm; resting ege 53 pm.

Ecology: Described from a shallow moor in

Germany, where it was noted to graze green algae.

Single unverified record from Boar's Lagoon, Vic.

Not seen in our material,

Literature: Berzins 1982.

Cephalodella eva (Gosse)

FIG, 9:2

Furcularia eva Gosse, 1887, p, 864, Fig, 14:9.

Cephalodella eva: Harring & Myers, 1924, p. 507.

Type locality: (England), “Lacustrine,”

Holotype: Not designated,

Description: Body slender, laterally compressed,

dorsally gibbous; head short, broad, neck clearly

marked; plates distinct; corona oblique without

projecting lips; foot large, with large pyriform foot

glands; toes long, approx. 1/2 total length, very

slender and generally curved ventrally; trophi type

B, with long, strongly crooked manubria; no eyespot

or retrocerebral organ.

Total length 190-285 pm; trophi 23-30 pm

(IZ W ROSTE & BR, J SHIFE

(fuleriim 19 zm, manubnum 23 am, right uncus 1

pm}; toes 50-85 pm.

Ecology: Cosmopolitat i periphylod, diss beach

sand, moors. Grazes diatoms. Rare, from a

billabony, near Bildon, Vie; Tas. 17.9-19.7 °C, pH

7.2-7.3, 009.2, 87 pS aml, 5 NTU,

Cephalodella exigua (Hudson & Gosse)

FIG, 9:3

aac thiag exlgva Hudson & Gosse, 1886, 2, p. 78, Fig.

22:15,

Cephalodeélig exigtiy Harring & Myers, 1924, p. 481,

Type Jocaliiy; Chellenham, England. “., , windnw

lank."

Holotype: Nor designated.

Description; Body short, stout, appears truncated

at posterior erid; lateral sulci between Jorica plates

disuaer neck clearly demarcated; foot small,

tubular; toes short, <1/5 body length, tapering

eradually to acute tips; salivary glands not

described, may be absent; trophi modified type B,

fulcrum slightly expanded distally, rami denticulate,

manubria strongly curved distally, but two ends do

not meet to form the fenestration typical of type

C trophi; two red cerebral eyespors; caudal antenna

very distinct; male known; resting egg with smooth

dark brown shell.

Total length 90-125 pm, toes 21)-26 pm; trophi

3) pm; male 74 ym; toes 19 pm,

Feolagy; Cosmopolitan between water plants,

Sphagnum. Common in R Murray (Vic.) billabongs

in Spring, Tas. 11,0-13.6°C, pH 6.2-7.4, 220-1900

eS cm, 2.7-120 NTU,

Literature: Berzins 1982; Kaste & Shiel 1987b,

Cephalodella forficata (Ehrenberg)

FIG. 3:4

Notwmmaia forficata Ehrenberg, 1842, p. 134.

Cephalodetla forficata; Harring & Myers, 1924, p. 499,

Fig. 33:7.

Type localirv: Bertin.

Holoivpe; Not designated.

Description; Body elongate, slender slightly

compressed laterally; meck well-marked; loriea

plates distinct; sulci narrow, parallel-sided; toes

approx 1/4 body lengih, widely spaced ar base,

short, stout, taper to acute apices; foot glands large,

pyriforms gastric glands large. red-pigmemted in

oldet individuals; trophi type B, manubria crooked,

eyespot absent.

Toral length 144-265 zm) toes 36-56 pin, Crophil

16-26 pm,

Eeology: Cosmopolitan in vegetation of standing

and flowing waters, Rare; N.SW., Qld, Tas.

16.5-I8.5°C, pH 4.8-6.3, 25-100 pS em. |,

Literature: Shiel & Koste 1970

Cephalodella forficuia (Ehrenberg)

FIG. 45

Disremma furfieula Ehrenberg, 1832, p. 199.

Cephulodetla forficula: Harring, 113, p. 12-

Tipe locality: Berlin,

Holotype: Not designated.

Description: Elongate, spindle-shaped body, slight

constriction at neck; integument flexible, without

lorica plates; abdomen tapers to ill-defined foot

toes short, stout, recurved, about 1/5 total length:

loes have distinctive transverse spicule row (2-4) on

dorsal median surface which terminates in a larger

spine; mastax with salivary glands; jrophi rype D,

manubria dilated distally, bur nor crooked, with

distinctive oval basal plate; single (rantal eyespot;

no retrocerebral organ.

&cology; Pancontinental. 12.0-25.0°C, pH 4.8-6,8,

25-440 pS em-', TDS 19.7 mel '. 73-25 NTU.

Literature: Shiel & Koste t979; Koste & Shie! 1987h.

Cepkalodelta gibhe (Ehrenberg)

PIG. 9:6

Furcutaria gibba Ehrenberg, 1832, p. 130, Fig. 4:16,

Cephalodetla gibba: Harring & Myers, 1924, p, 472.

Tipe locality: Berlin.

Holotype: Nat designated,

Description: Body stightly elongated, compressed

laterally; gibbous rump; lorica firm with disline

plates; sulci widen slightly posteriorly; toes tong,

straight or recurved, slender (ca. 1/3 body length);

masiax very large; trophi type B; rami with

denticulate lamellar combs on inner ventral

margins; manubria strongly crooked, shaft with

both sides lamellate; no retrocerebral organ; single

frontal eyespot,

Total length 250-450 yum; toes 67-150 ms traphi

70-9) pm.

Ecology: Cosmopolitan in littoral of fresh-brackish

waters, also in branchial chambers of Crustacea.

Eats unicellular alpae, flagellares, also carnivorous,

particularly on ciliates, Euryiopic, pancontinentat

in Australia, most common representative of the

genus. Abundant in acid waters in Tasmania,

9.5-23.0°C, pH 4.7-7.8, 92-700 pS em |, 17-110

NTU,

Comment: Several forms (ecolypes or a species

complex?) are listed in Koste (1978), Harring &

Myers (1924) noted that C. gibha is “somewhat

variable”. A distinctive ssp., C. gibba nricrudactyla

Koch-Althaus, 1963 (Wig. 10:1) was recorded from

a roadside pool near Scorts Peak Dam, Tas, 18,0°C,

PH 64, 122 wS cm |. This appears to be a good

apecies, however more detailed comparison of the

Tasmanian matenal with the notminare species is

required.

ROTIFERA FROM AUSLRALIAN INLAND WATERS 123

Fig. 10. 1, Cephalodella gibba microdactyla Koch-Althaus: (a) darsal; (b) lateral; (c) trophi, ventral, right manubrium

omitied; (c) trophi, lateral. 2, C. gis/feni Berzins: (a) dorsal; (b} lateral; (c) trophi,. ventral: (d) trophi, lateral. 3,

C, gracilis Ehrenberg; (a) lateral; (b) trophi, ventral; (c) trophi, lateral; (d) manubrium, lateral, 4, C. Aoodi (Gasse):

(a) lateral; (b} trophi, ventral; (c) forcipate rami; (d) trophi, lateral; (e) posterior end and toes. 5, C. intuta Myers:

(a) lateral; (b) trophi, ventral; (c) trophi, lateral. 1 after Koch-Althaus (1963); 2 after Berzins (1953); 3 after Wulfert

(1937); 4a, 5 afler Harring & Myers (1924); 46-e after Donner (1950), Scale lines: adults 50 pm; trophi 10 wm,

ne! W. KOSTE KR

Lirerature: Pourriot 1965; Koste 1978; Kose & Shiel

ISG, LOBTb:

Cephalodella gisten’ Berzins

FIG, 10:2

Cephalodella gislent Bermns, 1953, p. 4-6, Figs 1-4.

Tipe locality: Porongotups, W.A.

Holotype: Not designated,

Description: Frotn Berzins’ deseriptiot of a single

contracted individual ,.. “Body squat, rounded,

with small bulge in dorsal posterior part; ventral

margin lightly convex; head very large, almost 1/2

body length, somewhat narrower than body;

dorsally a distinct longitudinal sulcus visible; lateral

sulci indistinct; cuticle somewhat stiff, enabling

head to be seen; dorsal and lateral boundaries of

plates obscured, not distinct; animal very hyaline;

foot stout, distal, extending slightly over base of

Loes; toes proximally thickened, compressed dorso-

ventrally, tapering abruptly in distal 1/3 to sharp

points; in dorsoventral view [there is] a very peculiar

semicircular deflection of toes: in thickened part.

nf toes.is relatively large ducit mastax Jarge, with

two salivary glands; rropli symmetric, intermediate

hetween type A (manubria) and type B (rami);

fulcrum slender [unusually short for genus), slightly

dilated distally; manubria slender, terminally

[slightly] crooked; rami wide, of simple

construction, withoul leeth . . . eyes or lenses not

observed".

Total length (contracted) 85 pm, body

(contracted) 44 pm; head width 40 um; trophi 20

um; tocs 20 wm; proximal toe thickness 5 am.

Ecology: No ecological information other than that

collections came from small alkaline waters or

“Brackwasser". A single individual closely

resembling Berzins' description was recorded from

Salt Creek, near Berri, 5. Aust. (19.0°C, pH 7.8,

220 gS em, DO 9.2 mgl-', TDS 132 mal +, 45

NTU) (Shiel & Koste 1979) and another in Ryan’s

| Billabong at Albury (Shiel unpublished}.

Conmnent: We regard C. gisleni as a vahd species

on the basis of the distiitctive trophi, toes, and other

characters described above, despite its descrimion

from a single specimen. This specics may be mare

widely distributed across southern Australia than

the sparse records indicate.

Cephalodella gracilis (Ehrenberg)

FIG, 10:3

Furcularia gracilis Ehpenbers, 1834, j. 130: 1838, p_42t,

Fig. 48:6-

Cephaladella practi: Harting & Myers, (924, p. 474.

Tipe locality: Berlin.

Halatppe: Not designytedt,

I SHIFT

Description: Short, laterally compressed body,

round posteriorly; lorica thin, flexible, plates

distinct; sulci narrow, small tail: toes short, ga, 1/5

body length, slender, recurved slightly to acuie.tips;

foot glands moderately large, pyriform; mastax

large, trophi (type A) variable (Rost: 1978); fuleruns

curved, manubria distally cruuked or fanlike,

denticles may be developed on inside of rami,

pseudoallulae symmetric of asymmetric:

occasionally a reduced number of nuclei in

Vitellarium (4-6); single eyespot may be colourless.

Total length 125-150 sim; toes 22-30 pm; trophi

22-27 um; male 65-75 ppm,

Exology: Cosmopolitan in frésh, also in athallassic

saline waters. Berzins (1982) noted if was

“widespread” jn Victoria. We bave found C. grucilis

only twice a 1987 collection in ‘Tasmania (shallow

vegetated pool, Miena-Deloraine road near Golden

Valley), and Sept. 1990 in Ryajis 2 Billabong,

Wodonga, Vic. 16.0-22,0°C, pH 6.4-6.85, 114-292

pSom |,

Literature: Kosie er af $988.

Cephaiodelta hood! (Gusse)

FIG. Wh4

qa iuschiza hoodi Uudson & Gosse, 886, p, 79, Fie.

Cephalodelia hoadi: Harring & Myers, 1924, p. 482

Cephalodella remanei Wisniewski, 1934, p 353, Fig.

597-2).

Type locality: Loch near Dundee, Scotland.

Holotype: Nor designated,

Description; Fusiform body, glbbous dorsally;

abdomen unusually elongate, with dorsal plates;

foot small, conical; tail prominent; toes shore ¢f/'4

total length}, stout, decurved, tapering to acute tips;

foot glands small, pyriform; corona oblique, with

prominent beak-like lips {rostrum}; trophi type A,

with slender, short, rodlike manubria, curved at end

but not crooked; two pleural rods present; rami

sotrictimes with ‘pseudnalulae’, toothed inner

tiargin; caudal aniennae setae longs retrocercbral

Organ present; eyespot lange, at posterior end of

cerebral ganglion.

Total length 110-195 ym; toes 32-47 jum; trophi

30-38 um (fulcrum 16-25 yin, rami 14 pm,

manubria 14-20 wm, unci 8 um); male 10-LL5 jem.

Ecology, Cosmopolitan in fresh and inland saline

waters, in beach sand, submerged muss, also in

flowing waters. Rare, Gippsland, Vie, and Mr Field

Nal, Park, Tas, 16,0%C, pH 7.4 (Shiel & Tan

unpublished),

Literature: Berzins (982.

Cephalodeila intite Harring & Myers

FIG. 10:5

Cephatudetla intact Harring & Myers, 1924, p SOU-Stb,

Fig. 35:2-5,

ROTIFERY "ROM AUSTRALIAN INLAND WATERS 25

Tipe focality: Loon Lake, Vilas County, Wisconsin.

“collected among submerged Spdagrturn.”

Holotype: Not designated. IMyers colleytion,

AMNH, NY. Na, 566 in AMNH is a co-type,

Description: Body moderately elongated; head

longer dorsally than ventrally, corona markedly

oblique; lorica rigid, plates distinct; toes long and

slender (1/4 body length), rapering to acute Lips with

transverse basal septa; mastax large with salivary.

glands; trophi type By fulcrum expanded distally;

manubria crooked; rami ends strongly toothed; rami

with small alulae (easily lost in caustic or bleaching

solution used to clear trophi); gastric glands red to

red-brown, ducted. retrocerebral sac present; no

eyespat. May be confused with C forficuta, but has

Telatively longer toes (toe:total length ratio <4 in

G intuta, >4 in C Jorficata).

Total length t1$-225 um; toes 30-60 ym; trophi

30-40 jum.

Ecology: Cosmopolitan in standing and flowing

fresh waters, in Moss and periphyton of submerged

vegelation, Rare, NT, Tas,, Vic, 13.0-18.0, pH

4,7-1.8, 42-213 uS cm ?.

Literature: Koste 1981; Koste & Shiel 1936.

Cephalodelia tindamaya Koste & Shie

FIG. Us)

Be ppionetia lindamave Koote & Shiel, 1986, p. 95-6,

ig. J-

Tipe levality: Stock dam J km south of Copping,

Tasmania.

Holorype: South Australian Museum (SAM) V4019.

Coll. R. J. Shiel, OL.x11.1985,

Description: Body shert, stout; head broad,

defiexed; lonca flexible, plates indistinct; toes

relatively long (>1/4 body length), basally

thickened; terminal claws curved with acute tips,

four distine! spinules in row inside claw; mastax

large; trephi type B, fulcrum long, narrowest in the

centre, flaring at distal end; manubna unusual,

terminally crooked, leaf-shaped; rami with

denticulate inner margin behind tips, uncus with

one tooth and basal lamella; foot glands large, club

shaped; eyespol not recorded.

Total length (contracted) 245 yim; toes 68 «rm

(spinules 4-6 ym); trophi 43 wm (manubria 38 um,

fulcrum 24 pin, unei 17 wm, rami 14 am,

Ecnlogy: Endemic. Known only from acid stock

dam at Copping, eastern Tas. 21,7°C, pH <4.0, 80

oS cm.

Cephalodella megalocephala (Glasscott)

FIG, lhe?

Sear wires menolocephale Glasscart, 99, 9, $6, Fiz,

4:3,

Caphatodetta megalocephata: Harting & Myers 1924,

p. 494,

Tpe locality: ({reland),

Holalype; Not designated

Description: Body stout, dorsally gibbous; head

very large, ciliary field extremely oblique; apical

ficld with two large cirri; lorica thin, flexible, plates

indistinct; dorsal median sulcus may have convex

rather than concaye. connecting integument (Hauer

1921); foot 2-scgmented; toes short (1/6 total

length), decurved, sharply pointed; trophi of

peculiar (ype (E): simple rod-shaped fulcrum;

manubria two: sigmoid curved slender rods; rami

appear semicircular trom above: unci multi-toottied,

rake-like. Sometimes triangular, lamellar, thin epi-

pharynx distinguishable; no eyespot; retrocerebral

organ transparent.

Total length 195-210 am, toes 34-38 pm; (rophi

30 pm. Larget forms to 325 pm (Donner 1949) may

be ecolypic variants or species complex.

Ecology: Cosmopolitan; mud flats, beaches, sand,

penphyton of fresh water, margins of flowing water,

Feeds on diatoms and Chlorophyceae. Wentworth

Falls, N.S\W., St Marys, Tas., Ryans 2 billabong,

Wodonga, Vie, 14-21,0°C, pH 6.2-6.8, 73-351 uS

cm |,

Literature; Berzins 1982; Koste & Stiel |986,

Cephalodella nusgurnas Wulfert

FIG. Lh3

Cephaledelia misgurmus Wulfert, 1937, p 420, Fig, 29.

Type tocolity; Single locality ool specified.

“[ ., . bottom of muddy streams like the Saale and

Unscrut ... }." (Germany).

Holotype; Not designated.

Description: Body elongate, widest in posterior

third; head and trunk loricate, three larze and rwo

smaller plates; neck clearly defined; toes long, ca,

1/3 total length, slightly wider at base, parallel for

much of their length, terminating in acute tips:

mastax small with two small salivary glands; trophi

type C, syminetiical: fulcrum straight, flaring

distally, manubria with straight shalts (no lamellae),

ending distally in a distinctive open cing; unci robusr

with quadratic plate of) outer half; subcerebral

gland present; two frontal eyespots With crystalline

lens in common capsule,

Total length 165-190 pm; toes 49-61 pm; traphi

22 pnt.

&celegy: Burope, revorded in mud, flowing water,

Single recard, Mavela Ck, N.T.

Literature; Koste T98l_

Cephelodelia mucronata Harring & Myers

FIG. th:4

Cephalodella mucronsna Harring & Myers, 1924, (), 510,

Fig. 36:2-4.

W. KOSTE & R. J. SHIBL

126

ROTIFERA FROM AUSTRALIAN INLAND WATERS mw

Type flocalizy: Vilas and Oneida Counties,

Wisconsin, “in weedy, soft-water ponds.”

Holetype: Not designated. 7Mycrs collection,

AMNH, N¥. Ne, 252 in AMNH is a co-type.

Description: Elongate slender body, with rigid torica

extending, beyond end of foot; longitudinal sulci

deep; foot sheath has triangular ventral point and

dorsal spine, separated by deep, rounded sinus; toes

exceptionally long (ca. body length) slender,

recurved and pointed; mastax typical for genus:

trophi type B; fulcrum long and straight, manubria

slender, distally crooked, rami denticulate on inner

margin; retrocerebral sac present; no eyespot.

Total length 265-275 pint; toes 120-140 4m; trophi

36 urn,

Ecology: Paniropical-pansubtropical warm

stenotherm (20-32°C) in shallow vegetated ponds,

also New Zealand. Ears diatoms, unicellular algae:

Koste (1978) noted thet C ¢rucraneta also was

predatory on bdelloid rotifers. Isolated records from

N.SW., NUT., W.A. Widespread int shallow pools in

Tas, where it appears to occupy a Breater thermal

rage than elsewhere: 94)-29.0°C, pH 3.1-7.4,

LL8-98,3 pS cm-!.

Literature: Koste & Shiel 1986.

Cephialodella nang Myers

FIG. 12:1

ri > haan cura Harring & Myers, 1924, p. 491-2,

ig. |,

Type locality: Corduroy Creek, Absecon, New

Jersey. “collection in Sphegaurnt."

Holotype: Nor designated, ?Myers Collection,

AMNH, NY.

Desvriplion: Body short, vonical, tapering gradually

from corona to base of toes; head large, ca, 1/2

Jengih of body, and wider than abdomen; lorica

moderately flexible, plates distinct; (oes ca, 1/3 body

fength, long slender, set wide apart at base with

gentle sigmoid curve, tapering 1¢ bristle-like apices;

foot glands small, pyriform; corona oblique with

prominent beaklike lips; mastax very large; trophi

type A; fulcrum slightly expanded distally;

manubria slender, siightly clubbed and recurvet

ends but not crooked; salivary glands small:

eyespots ac posterior end of ganglion, uo

retrocerebral organ.

Total length 105-14) wm; toes 35-32 amt; crophi

30-34 yan,

Ecology: in submerged Sphagnuyn Europe, N,

America; recorded from Clunes, Vic, and Little Pine

Lagoon, Tas. 80°C, pH <$.0, 33 pS emo.

Literature: Berzins 1982; Koste & Shiel 1987b.

Cephalodelig panarist# Harring & Myers

FIG. 12:2

Cephalodelia pavariste Harrine & Myers,

p. 478-9, Fig. 5-7.

Tppe lucality: Four Mile Run, Washingtan, Duc.

Holotype; Not designated. ?Myers collection,

AMNH, NY.

Description; Body large, elangate, slender; dorsal

margin curves downwards posteriorly us base of

foot; integument very Mexible, plates indistinct! foes

very long (ca. 1/3 total length), stout and recurved,

tapering to acute tips; occasionally a dorsal

toothlike spine 1/3 of length along tocs; foot glands

extremely long, clubbed; mastax large, trophi robust

(Type D); fulcrum long, siraight slightly expanded

posteriorly; manubria short, recurved posteriorly

burt not crooked: with Large basal plate; unci have

typical single tooth; eyespot frontal with front part

of capsule colourless resembling “lens”,

Total Jengil 360-375 pm; roes 102-105 um; tropn

6Sum.

teolagy: Rare. N. America, S. E. Europe. Bills-

bongs, Magela Ck NT, R. Mutray N.SW.

Literatures Koste 1978.

1924)

Cephalodetfa parasitica (Jennings)

FIG, 12:3

Pleurotrocha parasitica Jennings, t900, p. 84, Fig, 16:13,

14,

Cephalodelle perasitica: Harring & Myers, (924, p. $12.

Type lacelity; Small pool near Lake St. Clair

(US.A.).

Holotype: Not designated,

Leseription: Body tusitorm, curved and gibbous

dorsally; head unusually long, tapers from neck to

corona: integument flexible, no sign of fissured

lorica; four short and conical; toes ea. 1/6 body

length, slightly decurved io. slightly sigmoid,

tapering to acute tips; mastax large with two large

salivary glands; Lrophi type A, with sharply pointed

unci and rounded, curved rami which have curved

alulae on their outer margins; gastric glands

brownish to black; no eyespot.

Total length 110-200 jum; toes 28-35 jam; trophi

32 wm.

Fig. U1 1, Cephaladella lindaraya Koste & Shiel: (a) lateral; (b} ventral; (c) trophi; (d) manubyium, 2, C megalecephala

(Glasscott): (a) lateral; (b) trophi, ventral; (c) tropha, lareral, 3, masgurvues Wulfert: (a) dorsal; (b) ventral; (c)

manubrium; (d) fulcrum, lateral; (2) wo views of uncus; (fy 2) Two Views of ramus; (h) trophi, ventrs], 4, Co mucronata

Myers: (a) lateral; (b) trophi, ventral; (c) trophi, lateral. 1 after Koste & Shiel (1986); 2, 4 after Harring & Myers

{1924}; 3 after Wulfert (1937). Scale lines: adults 50 pin; trophi 10 ani.

128 W. KOSTE & R. J. SHIEL.

Fig. 12. 1, Cephalodeila nana Myers: (a, b) lateral; (c) dorsal; (a) trophi, ventral; (c) trophi, lateral. 2, C. penarista

Myers: (a) lateral; (b) toes, lateral; (c) trophi, ventral; (d) trophi, lateral, 3, C. parasitica Jennings: (a) lateral; (b}

dorsal; (c) trophi, ventral. 4, C. sterea (Gosse): (a) lateral; (b) trophi, ventral; (c) views of manubria; (d} trophi,

lateral; (e) uncus, la, 2, 3a, 4a after Harring & Myers (1924); Ib-d after Wulfert (1940); 3b Koste orig.; 4b-e after

Wulfert (1937), Scale lines: adult 50 am; trophi 10 um.

ROTIFERA FROM AUSTRALIAN INLAND WATERS (29

Ecology: Europe, N. America, occasionally free.

swimming, but more commonly parasitic on

oligochaetes (Chaetogaster, Nois, Stvlaria, etc).

Single report from Diggers Ck, Mt Kosciusko. Not

seen in our collections.

Literature; Koste 1978; Berzins 1982,

Cephalodella sterea (Gosse)

FIG, 12:4

Furcularia sterea Gosse, 1887, p. 864, Fig. 14:8

Cephalodella sterea; Harring & Myers, (924. p. 474.

Tipe focality: Rockery pond in the grounds of

Watcombe Park near Torquay, England.

Holotype; Not designated,

Descriplion: Body fusiform; head large; lorica firm,

plates well marked; foot large, robust; tail extends

beyond distal end of toot; toes short, stout slightly

recurved posteriorly, may have slightly undulate

margins, generally <1/4 body length; foot glands

large and pyritorm; mastax large with four small

salivary glands; trophi asymmetric type B;

manubria strongly crooked; unei with variable inner

margin denticulation; rami with three pleural rods;

retrocerebral sac present; two red [rontal eyesports

in single capsule.

Total length 140-250 pm; toes 26-56 wm: trophi

37-39 gm.

Ecology: Cosmopolitan, in moss, in mud margins

of standing and flowing waters. Rare: NT, Tas,

Yie., 12.0-22.0°C, pH 5,3-6,9, 73-351 eS cm-!,

Literature: Kose 1981p Koste ef af, 1988.

Cephalodella tantilloides Hauer

FIG. 13:)

Cephalodelia tantilloides Hauer, 1935, p. 69, Fig. 9.

Type fecality; High Moor pond, Black Forest,

Germany:

Holotype: Not designated.

Description: Body squat, vaulted dorsally, abdomen

falls sharply to clearly demarcated foot; plates and

sulci distinct; dorsal sulcus relatively deep; lateral

sulci margin parallel almost to base of toes; toes

long (1/3 body length), parallel sided for 3/4 of

their length, to a distinctly segmented tip, slightly

recurved; toe tips acute; foor glands smalls lips

Project from mouth area; trophi type A; fulcrum

dilated distally into‘a broad plate; rodlike manubria

curve upwards, rami with inner denticles: cerebral

eye present; retrocerebral sac not recorded.

Total length 104-175 wm; tocs 45-56 jam; toe

points 14 wm; trophi 34 um.

&eology: Europe, in Sphagnum. Single recond,

Bromfield Swamp, Qld. No ecological data given,

Literature: Green 1981; Koste 1978.

Cephulodelia tenulsete (Burn)

FIG. (3:2

Furcularia leruixeta Bom, 18990, p, 34, rext fig.

Diaschiza tenniserw Dkxon-Nuttall & Freeman 1903,

p. 138, Fig. 1:2.

Cephalodella renuiseta: Hatring & Myers, 1924, p. 508.

Type locality: (England).

Holotype: Not desighated.

Description: Body elongate, laterally compressed:

head large, short: abdomen unusually long, gibbous

posteriorly; lorica flexible, plates. indistinct; foot

short, conical: toes very long (1/2 body length),

slender, slightly recurved; mastax large, trophi type

D; fulcrum slightly expanded posteriorly, manubria

rodlike not crooked; no eyespot.

Total length 205-314 am (Koste) 380-390 (H&M);

toes 59-96 jum (Koste) 120-125 (H&M); trophi 35-39

am.

Ecology: N. America, Europe, Rare: Vic, WA,

16,0°C, pH 7.1, 264 pS cm’', Eats unicellular

green algae and diatoms.

Literature: Berzins 1953; Koste 1978.

Cephalodella tinea Wulfert

FIG, 13:3

Cephatodelte fince Wulferi, 1937, p. 622, Fig. 31

Type focality> Dram outtlow (Bad Lauchstadt)

Germany,

Holotype: Not designated.

DHescription: Body elongate, laterally compressed;

abdomen slightly bulbous prior to short tail

extending beyond foot; plates and sulci distinct; toes

relatively short, ca. 1/3 total body length, thickened

at base, with slight medial swellings; mastax with

two large salivary glands; trophi type D; fulcrum

dilated distally, broad-bladed proximally (seen

Jarerally), rodlike viewed apically; manubria with

blunt, slightly enlarged tips; paired eyespots in single

capsule,

Total length 260-280 xm; trophi 29-3) yn; toes

52-70 um; male 160 pm; subitaneous egg 60. pn.

Ecology: Europe, in drains, decomposing

vegetation, manure pits and piggery outflows, Eats

diatoms, Rare: Vic. (billagong), Tas. (stock dam)

15,0-19.0°C, pH 4.9-7,1, 264-273 uSem-', DO 10.3

me t-',

Literature: Koste ef il, 1988.

Cephalodella vertripes Dixon-Nuttall

FIG. (3:4

” peathist ventripes Daxon-Nuriall, 1901, p. 25, Fie.

" Caphalodette venirives: Harring & Myers, 1924, p. 484,

Type locality: Knowsley Park, Lancashire, England,

Holotype: Not designated,

130 W. KOSTE & R. J, SHIEL

ROTIPERA FROM AUSTRALIAN INLANIZ WATERS lat

Fig. 14, Drilophaga bucephalus Vejdovsky; (a) darsal; (b)

trophi,. ventral: (c) tropli, lateral. Alter Koste (1978),

Seale lines: adult 50 pm; trophi 10 pm.

Description: Bady short, stocky, bulbous dorsally;

abdomen may extend beyond distal end of the foot;

plates and sulci distinct; dorsal sulcus a distinet V-

shaped groove; foot ventral, small; tocs short and

stout, decurved, ca. 1/§ total length; corona with

prominent lips; mastax large; trophi type A;

fulcrum dilated distally; distal ends of manubria

sickle-shaped, not crooked; double cerebral eye.

Total length 135-140 ym; toes 25-28 pm; trophi

30-34 wm

Ecolagy: in fitroral periphyton of mosi standing

fresh waters: N. America, Europe. Uncommon in

billabongs of River Murray, N.SW., Goulburn

River, Vic, also dams in northeast Tasmania

13.0-21.0°C, pH 4.8-7,1, 18-351 pS em).

Literature: Berzins 1982; Koste & Shiel 1987b,

Genus Dorystema Harring & Myers

Dorystoma Harting & Myers, 1922, p. 5$5. Manatypic

genus.

Type: Proales caudata Bilfinger, 1994.

Dorystoma caudata (Bilfinger)

FIG, 153]

Proales caudata Bilfinger, 1894, p. 46, Fig. 2;3-4.

Dorystoma caudata: Harring & Myers, 1922, p. 555.

Type locality: Wurttemberg, Germany.

Holotype, Not designated.

Description: Stout, iloricate notommatid rotifer:

uansparent body, gul may be coloured; corona

oblique, with lateral ciliary tufts (‘auricles’) for

swimming; constriction separates head and

abdomen; abdomen bulbous, with longitudinal

striations; foot short, apparently two-jointed; toes

long, pointed, short; at base of foot, bulb above

anus carries short spine; gut yellowish, often filled

with yellow-gold balls; mastax specialized virgate,

With long pharyngeal tube; trophi modified to

support mastax walls; specialized piercing

epipharynx present; manuhbria with wide crook:

unci absent; single bright red cerebral eyespor,

(sometimes absent); dorsal and lateral antennae in

pits in cuticle, sensillae distinct; subitaneous egg

smooth-shelled.

Total length 130-260 wm: toes 16-22 pm; spine

10-22 pm; trophi LS am; pharyngeal tube 22 um;

subitaneous egg 56 = 44 ~m,

Ecology: Isolated records from periphyton of

submerged plarits, esp, Polamogeton, Nuphar, also

in Sphagnum, Europe, N. and S. America. Eats

algae. Single Australian record: Yarnup Swamp,

W.A. 17°C, 1600 wS em-'.

Literature’ Koste 1978; Koste ef ai, 1983.

Genus Drilophaga Vejdovsky

Drilophage Vejdovsky, 1883, p. 390,

Tepe: Drilophaga bucephalus Vejdovsky, 1883,

p, 390, Fig. 171-8.

Body slender fusiform; cuticle soft, flexible, with

indistinct annuli; head cylindrical, elongate, with

simple circumapical ciliation; small jail projects over

foot; toes minute, conical, ca, 1/20 body length;

foot glands with reservoirs; mastax with two lateral

and one posterior salivary glands; trophi with

Fig. 13. 1, Cephalodella tantilloides Hauer: (a) lateral; (b) teophi, ventral; (c) traphi, lateral; 2, C, tenuiseta {Burn):

{a} lateral; (b) trophi, ventral; (c) trophi, lateral; 3, C. rinca Wulfert: (a) lateral; (b) traphi, ventrat; (c) Iranhi, taleral;

4, ventripes Wullert: (a) lateral; (b) wrophi,. ventral; (c) trophi, lateral. | after Hauer (1935), 2a, 4a after Harring

& Myers (1924); 2b-c, 3b-c, 4b-c alter Wulfert (1937), Scale lines: adult 50 yam; trophi 1) pm,

32 W, KOSTE & R. J, SHEL

Tig. $5. 1, Dorystoma caudaza (Bilfinger); (a) lareval; (b) dorsal; (c) rophi, ventral; (d) trophi, lateral, 2, Bnleroplea

faviestris Ehrenherg: (i) lateral; (b) dorsal; (c) trophi,. ventral; (d) trophi, tateral; (e) iacus, oblique frontal view.

1 after Wulfert (1960); 2 after Harring & Myers (1924), Scale lines! adult 50 am; trophi 10 pm.

anchor-shaped incus; rami curved inwards; unci

short, thick, bidentate; manubria distally crooked;

fulcrum long, straight to lightly curved distally;

pleural rods present; gastric glands spherical;

stomach and intestine separate; vitellarium large;

large retracerebral sac dorsal to ganglion; evespats

absent; dorsal and lateral antennae present, latter

projecting from small tubular extensions of

integument.

Two other described species were synanymised

with D. bucephalus (Fig. 14) Koste (1978), BD.

bucephalus is parasitic on the integument of

otigochaetes and leeches (Lumbriculus, Rvachelnis,

Stiylodrillus, Nais, Herpobdella, Hiruda) (Koste

1978).

Total length LO-355 zm; toes 6-11 jam; trophi

20-32 pm (unci 8 pm, manubrium 16 zm, fulcrum

25 um); subitaneous ege 50-62.« 35-39 um.

Comment: The synonymy of D. bucephalus, D.

delagei Beauchamp and D. judayi Harring & Myers

requires re-examination. Although Koste (1978)

attributed interspecific differences as described and

figured to observational errors or preservation

artefacts, habit differences were noted by the

different aulhors, i.c., parasitic v, free-swimming

(judayi), also differences in morphology,

particularly wophi.

A single free-swimming, animal identified 2s

Drilophaga was collected by WK from Ryan’s 2

billabong at Wodonga on Sept. 27, 1990, the first

record of the genus from the continent, No

oligochaetes or Jeeches occurred in the sample. The

living rotifer was filmed on videotape, but the

mastax was lost during clearing. preventing specific

determination. Until further material becomes

available, we can nole only that Drilophaga occurs

in Australia,

Llierature: Beauchamp 1904; Harring & Myers

1922.

Genus Enteroplea Ehrenberg

ELateropfea Ehrenberg, 1830, p. 46. Monotypic genus

Thoe: Enteroplea lacustris Ehrenberg, 1830

Enieroplea lacustris Ehrenberg

Rica, (5:2

Enteroplea lacusiris Ehrenberg. 1830, p. 46.

Tepe locality: Berlin.

Holotype: Not designated,

Description: Body with wade head, saccate

abdomen; foot directed ventrally, three-segmented,

offset from body; toes short, laterally barrel-shaped,

frontally claw-like; corona an oblique disc circled

by ciliary whorl (circumapical bard and. ventral part

of buccal field); dorsa) margin of buccal field with

type of pseudotrochus made of four closely-situated

membranelles; ventrolaicrally, beside mouth wide

row of menibranelles stand on ‘pedestal’; inner part

ROTIFERA FROM AUSTRALIAN INLAND WATERS 133

of buccal field unciliated; rwe frontal eyespots on

papillae in ‘forehead’ region; mastax resembles

fasphora, with more pronounced seizing function:

pincer-like angled rami can be protruded through

the mouth opening; inner rami margin with single

Jarge tooth, margin finely denticulate before and

behind; unei with one main- and one ancillary

tooth; no basal apophysis, however; small processes

present at insertion point of rami adductor muscle:

fulcrum boardlike; two small ventral salivary

glands; oesophagus long; stomach rounded,

cellular; intestine thin, ropelike, For additional

details of internal morphology, see Koste (1978).

Total length 500-600 pm; toes 30-35 jum; trophi

70 pm (fulcrum 21 jon; rami 56 pm; unci 35 ym;

incus width 46 pm, length 70 pm) subitaneous egg

155-160 «110-130 um; male 306 pm.

Ecolvgy; In shallow pools, ephemersl waters,

Europe, E, Asia, N. and S. America, Reported to

be carnivorous on other rotifers (Rhinoglena) in

culture (Pourriot 1965), Recornled by Colledge (1914)

from Qld, not found again until Oct, 15-18, 19%),

when individuals were identified from submerged

scales of Ricciovarpus natans, Ryan's 2 billabong,

Wodonga (Manuel & Shiel in prep.).

Literature: Colledge 1914.

Genus Eosphore Ehrenberg, 1830

EBetphora Ehrenberg 18340, p. 47,

Type locality: Toboisk, Siberia.

Type; Eosphora najas Ehrenberg, 1830, pp. 47, 84,

Fig. 7:3.

Body plump; head and neck distinguished! by

transverse sutures; abdomen sac-like with rounded

or weakly trilobed tail; foot lwo, three- or

unsegmented; two toes with long foot glands;

corona frontal; circumapical band interrupted

dorsally; two ciliary bundles laterally; buccal fleld

lightly or non-ciliated; cerebral eye at posterior end

of brain (absent in & anthadis); retrocerebral and

subcerebral glands present, size and shape variable;

mastax three-lobed; unci single toothed, may have:

small ancillary teeth; rami with symmetric braces,

occasionally with strong basal apophyses; at bend

of rami teeth on inner margin vary from 1-2 strong

to 4-§ smaller teeth in different species; fulerum

wide plank or handle-like; salivary glands differ

between species; five of six species are known from

Austratia.

Key to species of Eosphoru recorded from

Australian inland waters

I, Foot seginented, , , tel abe toca

Poot umsegmented.- ....--..... -...- 7

2(1), Papilla at baseof wer. 2.2200... Bayes

Ehrenbere (Fig. 1:3)

No papilla at base of toes.....-., E. ekrenbergi

Weber (Fig- 16:2)

3(1), Obvious cerebral eye Present, - 4

Cerebral eye absent. -....,.....-.- E. anthadis

Harring & Myers (Mig, 16:1)

4(3), Trophi leneth <40 yin, oon E. thofdes

Wolfert (Fig. 17:

Trophy length 50 pm.... 222-54. as &. thea

Harring & Myers (Fig. 16:4)

Sosphora anthadis Hamug & Myets

FIG. 16:1

Lasphora anthadi=s Harring & Myers, 1422, p, 641, Fig.

58:9-13,

Type locality: Not specified * .

widely distributed,”

Holotype: Not designated. 7Myers Coll, AMNH,

NY,

Description: Body broad and robust, oa, three times

longer than wide; integument soft, body

transparent; stout abdomen tapers from median lime

to base of broad foot; foot wrinkled but not

segmented; toes short, stout (1/20 fength), seen

dorsally margin almost forms hemisphere; seen

laterally, dorso-ventrally flattened, appear as normal

conical toes; mastax modified virgate; rami

symmetrical with four or five small teeth in median

section on each ramus margin; unci with one tooth,

small subsquare striated plate at base vestiges of

accessory teeth; fulcrum of two. plates joined

longitudinally to form V; manubrium a straight rod

slightly expanded at each end; salivary glands not

seen; gastric glands large, elongate, cylindrical,

terminating in mucus reservoir at base of toe,

retroterebral sac and two subcerebral glands

present; no eyespot.

350-410 wm; toes 16-22 pm; trophi 33-35 am

(fulcrum 20 pm, rami 18 ~m, manubria 22 pm),

Evology: ln acid waters or mildly saline waters

(Utricularia) in Europe, N, Anierica, New Zealand,

Japan.

Single Record; Crackers Swamp, off Brand Hwy,

W.A. 20.0°C, B00 wS cm-!.

Liferalune Koste 1978; Koste et af 197

.. appears to be

Eosphara ehrenbergi (Ehrenberg)

FIG, 16:2

Notommata najas Ehrenberg, 1832, p_ 142.

Eosphora ehrenbergt: Weber & Montel 14218, 0, 123.

Type locality; Berlin.

Holotype: Not designared

Description: Body broad, robust, coloured light

brown in fresh specimens; integument firm;

indistinct transverse folds between head/neck and

14 W. KOSTE & R. J, SHIEL

My tii hy

Fig. 16. 1, Easphora anthadis Harring & Myers: (a) dorsal; (b) lateral; (c) trophi, ventral (cl) (rophi, lateral. 2, £

ehrenbergl Weber: (a) dorsal; (b) lateral; (c) trophi, ventral; (d) trophi, lateral, 3, A. najus Ehrenberg; (a) dorsi;

(b) Tateral; (c) Lrophi, ventral; (d) trophi, lateral. 4, £. thea Harring & Myers; (a) dorsal; (b) lateral; (c) trophi,

ventral; (d) rrophi, lateral, 1, 2 after Harring & Myers (1922); 3, 4 after Harring & Myers (1924). Scale lines: adult

50 am; traphi 10 jin,

neck/abdomen; abdonien routided posteriorly with

short tail (two small lateral lobes); foot long,

cylindrical, two-segmented; toes short, acute,

conical; virgate trophi adapted for seizing prey; rami

approximately triangular, two teeth on each ramus;

fulcrum a broad plate with diagonal ventral edge

where abductor muscles attach; unci with large

subsquare basal plate with strong Ventral tooth;

manubrium with straight central section, expanded

anteriorly into: broad triangular plate, two large

salivary glands present: foot glands long, without

mucus reservoir; retrocerebral sac and 1wo

subcerebral glands; eyespot large, dark red, at

posterior end of ganghon; hypopharyny muscle

rudimentary. Male known,

Total length 350-450 yin; toe length 24-30 um:

trophi 65 xm; male 212 pm.

Ecology: Probably cosmopolitan between water

plants in fresh to slightly saline water, Single

unverified record from Victoria.

Literature: Anderson & Shephard 1892; Koste 1978.

Easphora najas Ehrenberg

FIG, 16:3

Eosphora nojas Ehrenberg, 1830, pp. 47, 84, Fig. 7:3,

Type locality: Tobotsk, Siberia,

Holotype: Not designated.

Description: Body robust, integument firm, bady

in fresh material light orange in colour, Head and

ROTIFERA FROM AUSTRALIAN INLAND WATERS 135

neck sutures distinct; abdomen broad and oval; tail

with targer median lobe, two small lateral lobes; foot

indistinctly three-segmented; toes long, straight,

slender; mastax modified virgate (for seizing prey);

yami triangular in ventral view; left ramus with

single large tooth, right with two teeth; rami with

finely denticulate dorsal extension; unci small,

subsquare basal plate with single clubbed ventral

tooth; manubrium broad, lamellate, anteriorly.

Lapering to knobbed posterior end: ventral salivary

glands distinct, right Jonger than left; retrocerebral

sac and subcerebral gland as in other species;

eyespot at anterior end of ganglion, two

(accasionally four) lateral eyespots in small

projections of corona,

Length 260-610 pm; toes 26-48 zm; trophi to 80

pin; male to 300 gm; subitaneous egg

140-150 120-130 ym; resting egg 130%170 xm; male

egg 100-110 %« 120-130: pm.

Ecalogy: Cosmopolitan in littoral between water

plants, preys on rotifers including Colurella,

Lepadella, Lecane, Monostyla and bdeltoids. Early

records from Vic. and Qld. In our material, Gwydir

R. at Moree, N.SW. (24.v.78), and recently (30,~.90)

in MyriopAvliam in a flooded roadside marsh,

Ryans property, Wodonga, Vic, 13,0-22,5°C, pH

3,97-8.0, DO 8&4 mg {-', 60-400 pS cm-!, 160

NTU.

Literature; Colledge 1914; Eyans 1951; Koste 1978,

Eosphera ifioa Harring & Myers

FIG. 16:4

unaeerts thoa Hurting & Myers, 1924, 9. 523, Fir.

Type locality: Cemetery Pond, near Eagle River;

Vilas County, Wisconsin.

Hototype: Not designated. ?Myers collection,

AMNH, NY,

Description; Body robust, integiiment flexible,

hyaline; head and neck fixed bui suture visible

between neck and abdomen; abdomen tapers to

broad tail; unsegmented conical foot; toes heart

shaped in dorsal view; mastax modified virgate;

fulcrum short.and broad; rami elongate with single

blunt tooth on inner edges and posteriorly with ca.

20 denticles; unci with robust clubbed ventral tooth;

small retrocerebral sac and two subcerebral glands;

large eyespot at posterior end of brain.

Fig. 17.1, Easphorg thoides Wulfert: (a) dorsal; (b) lateral; (c) rophi, ventral; (d) trophi, lateral. 2, Zothinia elongata

(Ehrenberg): (a) dorsal; (b) trophi, ventral; (c) trophi, lateral. | after Wulfert (1935); Scale Jines: adull 40 jm; trophi

1 pin,

Ro W. KOSTE & R_ 5. SHIEL

Length 300-500 um; tocs 20-35 am; trophi 50 am

long, 60 pm wide.

Erolagy:; Soft water, N. America. Two records:

Forestdale Lagoon, near Perth, WA. (Coll. J. van

Alphen, Murdoch University) arid Ryans 2

billabong at Wodonga, Vic. (Coll. J, De Manuel,

University of Barcelona). 16.0-24.0°C, pH 65-67,

263-310 oS em~'.

Enasphera thoides Wulfert

FLG, '7;1

EBusphora thoides Wulfert, 1935, p fi00, Fig. (Sa-d.

Thpe lacality: Saale River, eastern Germany.

Halotype: Not designated.

Description: Body resembles &. thee as above; lung

cilia from fateral margin of head but not auricles

per se; neck suture distinct; abdomen tapers to wide

font, its greatest width in anterior third; foot 3-4

segmented by light transverse lines; toes appear

triangular in dorsal view, conical in lateral view;

mastax with lateral salivary glands extends slightly

past neck suture; trophi with long, distally widened

fulcrum; rami triangular in dorsal view; in lateral

view curve downwards at right angle to acute tips;

unc with double-looped framework at free end (Fig.

17:\¢); manubria slightly asymmetric, with fanlike

lamellae at base, bifurcate epipharynx above trophil;

foot glands large, right larger than left; mucus

reservoir as large as toe present; retrogerebral sac,

subcerebral glands and eye as in & (how.

Body length 460-510 gm; (oes 26-35 pm; Inophi

37 pm,

Ecology: Europe. Two records: billabong of Magela

Creek, N-T. (Koste 1981), and margin of L. Mulwala,

Vic, (Shiel, unpublished).

Literature Wulfert 1935; Kosle 1981,

Cemment; Wulfert noted the resemblance of &

thoiges to E. thea, the former is distinguished by

the toe morphology, more elongate vilellarium; and

above all, differences in trophi structure ag

described. The animals found in our samples

resemble E. thoides, however minor differences in

lrophi structure were nated. Further matezial is

necessary for detailed examination,

Genus Eothinig Harring & Myers

Sothinia Haring & Myers, 1922, p. 555.

Tipe: Easphorr elongata Ehrenberg, 1832 =

Eetkinia elongata (Ehrenberg, 1832),

Type fecality: Berlin-

Kothinia was erected by Harring & Myers to

accommodate Easphora elongata Ehrenberg, 1832,

the mastax of which differed from Eosphora but

which could pol be included in the related genus

Sphyrias because of other morphological

differences,

Body elongate, slender; head and neck clearly

marked by transverse sutures; trunk with

longitudinal lines tapering to rai! of variable form;

cuticle very transparent; foot short, 2-3 segmented;

two toes and foot gland; corona slightly oblique,

with ciliated buccal field and marginal wreath of

cilia (reduced dorsally) with two lateral auricle-like

curves of strongly developed cilia; mastax virgate;

trophi with compact, fine denticles on inner margin

of triangular rami; unci generally single-toothed:

no preuncial teeth; fulcrum clongate, distally

dilated; manubria rod-shaped with triangular-

section al proximal end; large retrozerebral and

subcerebral glands: cerebral eye and two widely

separated frontal eyes. Eight taxa were referred [a

the genus by Koste (1978); one is known from

Australia.

Eothinia elongata (Bhrenberg}

FIG. 17:2

Losphorn elongata Ehrenberg, 1832, p. 145.

Suthinia elongata: Harring & Myers 1922, pp. 595.

646-648, Fig. 6151-3.

Type locality: Saale R., eastern Germany.

Holotype: Not designated,

Description: Transverse folds indistinct: foot jonger,

2-segmented; toes straucht will; conical tips, about

1/10 total length; corona frontal; trophi with

triangular rami; symmetrical; inner edges. armed

with numerous compact denticles; fulcrum of long

straight plates fused in a V-shape; distal end of

fulcrum finely subdivided; unci single toothed;

manubria straight rod-like; two pleural rods pair

transyersely across mastax for support during

PLUNping actionvembedded in mastax walls at dorsal

ends of rami (Fig. 17:2c); eyespots at posterior end

of brain; two accessory cyespots. on corona,

Length 350-310 wm; toes 32-45 pm; trophi 56-69

em; unci 14 pm; male 150-215 ym; toes 13 emt spiny

subitaneous egg 92% 115 ym; spines to 38 xm long.

Ecology: Widespread in littoral between water

plants. Europe, Asia, N, America. Carnivore of

other rutifers, particularly bdelloids. Known only

from Ryans billabongs at Wodonga, Vie,

14,0-22,0°C, pH 6.2-7,1, DO 4.1 mg L-!, 73-374

es cm, 5 NTU,

Literature: Koste 1978; Koste & Shiel 1980,

Genus Jew Harring & Myers

ftira Harring & Myers, 1928, p. 684.

Type: Diglena avrita Ehrenberg, 1830 = fined aurita

(Ehrenberg).

Tbpe facality: Bertin.

ROTIFERA TROM AUSTRALIAN INLAND WATERS

maT

Fig. (8.1, Léara aurita (Ehrenberg): (a) dorsal; (b) trophi, ventral; (¢) trophi, lateral. 2, 4. srrersd Walfert: (a) dorsal;

(b) trophi, ventral. 3. 2. viridis (Stenrons): (a) dorsal; (hb) trophi, ventral. 1, 3 after Harring & Myers (1928); 2 after

Wulfert (1935). Scale lines: adult SO pm; trophi 40 2m,

Body elongate, fusiform, gibbous posteriorly;

cuticle thin, flexible, body may be green due to

symbiotic zoochlorellae; two dorsal transverse

sutures separate head and neck; corona with

stumpy, nom-retractile lateral processes; foot and

toes short; one cerebral eye at end of brain; twa

frontal eyes on apical field, the latter sometimes

with speckled pigment flecks; single dorsal antenna;

lateral antennae small; tetrocerebral sac large;

mastax resembles forcipate type of Dicrenophorus

spp., but rami cannot be extended through mouth;

unci long, with bifureate tips and knoblike median

swelling: manubria long rods, proximally with small

lamellae (dorsal and ventral), distally hooked,

crooked or dilated; rami lyrate or forcipate, widely

separated, with dorsally curving tips; tips dilated

distally, toothed; inner rami margins (one or both

sides} with finely striate or smooth lamellae,

sontetitnes also on outer margin; alulae, sometimes

asymmetrical, may be present; fulcrum in lateral

view hooked or boardlike, frontally rodlike;

tudimentary epipharynx and oral plate may be

present. Female 180-500 «xm. Mate known but

undescribed. Three species are known from

Australia.

Key to species of Itura recorded from Australian

inland waters

1, Rami with ssvmmetric lamellae on outer border.

dbs GoM ole Sach! 1, aurita (Ehrenberg) (Fig. 18:1)

Rami without Jamellary ribs on outer border_.2

2(1), Subcerebral glands very long....-.-.. J. myersi

Wulfert (Fig. 18:2)

Subcerebral glands missing or pootly developed.

oe, EAA a fetys !, viridis (Stenroos) (Fig. 18:3)

Ttura aurcita (Ehrenberg)

FIG, 18:1

Diglena aurita Ehrenberg, 1830, p14.

Tura aurita: Harring & Myers 1928, p. 685.

Tepe locality: Berlin.

Holotype: Not designated,

Description: Body elongate, fusiform, may be green

due to symbiotic zoochlorellae; toes short, conical

with obtuse tips; mastax virgate; trophi asymmetric,

robust; rami lyrate, knobbed at tip and bearing 5-6

long teeth; inside left ramus has narrow lamella,

right ramus has broad denticulate Iametla; alulae

large, acutely pointed: manubria broadly expanded

proximally, abruptly curved distally; gastric glands

W. KOSTE & R. J. SHTEL

138

ROTIFERA FROM AUSTRALIAN INLAND WATERS yy

abseut; retrocerebral sac present; cerebral eye and

two Frontal eyespors (with lenses) present.

Length 180-220 pm; toes 9-18 ym; trophi 38-45

xin (eg. fulcrum 20 zm, manubria 30-33 pm, uncus

18 nm).

Ecology: In littoral of still and slowly Mowing

waters; Europe, N. America. Eats euglenoids and

other unicellular algae, Early reeord from Qld. In

Hur material, rare, unly in Vie, (hillabongs) and Tas,

(stock dams), 13,0-18,0°C, pH $.7-7.7, 42-3330 uS

em.

Luénature: Colledge 1914; Koste & Shief 1986,

finns myersi Wulfert

FIG. 18:2

Jira myersi Wulfert, 1925, p. 589, Fig. 6a-c

Iype locality; Near Halle, eastern Germany,

Holotype: Not designated.

Description: Vanable form, may be broader or

narrower than figured; toes short, tapered; frontal

eyes with lenses; gut sac ghassy preert, whereas in

other species tood balls are brawn; Jang asymmetric

subectebral glands; trophi robust; rami elliptical in

outline with inwardly curving tips which earry 5-6

teeth; right ramus with broad denticulate lamella

on inner margin, left ramus with narrower finely

denticulate lamella; alulae winglike expansion,

without sharp points; fulerum distinetive for

species, very short and wide,

Total length 270-406 pm; toes 21-27 am

(Fulcrum 13-16 pms rami 27-34 pm; manubra to

37 am: unci 22 am,

#eology: Europe, Asia. Bats Eugienu, Scenedesmus,

Pediasirum, Trackelomonas. Single record from art

Eleacharis bed, Snowdon’s Billabong, Wodonga,

Vie, 14.7°C, pH 7.1, DO 4.1 mgt", 240 pSem-',

5 NTU.

Literature: Kosie 1978; Koste & Shiel 1980.

Itura viridis (Stenroos)

FIG. 18:3

Eosphora virtais Sienroos, 1898, 136; Fig. 130-32.

{tere viridis: Harring & Myers 1928, p. 692, Fig. 241-2

Thpe locality: Lake Nurmijarvi, Finland.

Hatatype: Not designated.

Descripfion: Variable in form, readily confused with

congeners; duct of retfocerebral sac much shorter

ian congeners; rami armed with 12 teeth; right

ramus has broad striate, denticulare lamella, Jeft

ramus not lamellate; alulae large, broad, pointed

posteriorly; fulcrum nearly as long as rami; frontal

eyes with large spherical lenses and generally with

accessory pigmentation; characteristic sttaight,

narrow, sharply pointed toes; zoochlorellae in gut,

Total length 260-400 um: toes 16-26 wm; trophi

45-48 um.

Ecology: Littoral, possibly cosmopolitan, Europe,

Asia, N. and 8, America. Recorded from Trentham,

Vic. (Coll. 1. 3, Powling, Melbourne) and Rapseys

2 stock dam, Wodonga (Coll, F. Duan, MDFRC).

22°C, pH 6.0, 98.1 pS em-!.

Literature: Kaste 1978.

Genus Monomriaia Bartsch

Moromimara Bartsch, 1870, -p. 344.

Type: Vorticella longiseta Miiller,

Monommata longiseta (Miller, 1786).

Type locality: Copenhagen.

Body cylindrical or fusiform; suture between

head and abdomen; cuticle thin, firm, laterally and

dorsally with longitudinal striae; foot indistinctly

two-jointed: toes extremely long, almost {wice body

length, right longer than left (with exception of M.

aequails), corona slightly oblique, with marginal

whorLof cilia and lateral auricle-like tufts of longer

cilia for swimming; apical field utciliajed, buccal

freld ciliated; mastax variable, from simple virgate

to intermediate between virgate and forcipate type;

in farmer type (Fiz. 19:1b), rami lyrate or triangular

without inner teeth, manubria simple rods, unci

with one weak tooth or reduced to thin lamellar

plates (Myers 1930): in intermediate type (Fig,

19:7b), rami lyrate with one or more teeth on inner

1786 =

‘margin, manubria broad and lamellar at base, unci

with Three unequal long, slender clubbed teeth;

dorsal antennae single or paired on papillae in some

species; lateral antennae normal; cerebral eye at

posterior end of brain {absent in M. caecu).

Variations from generic characters are detailed by

Koste (1978) and summarised in the species

diagnoses below, Eleven species have been tecorded

from Australia.

Key to species of Monommats tecorded from

Australian inland waters

1. Toes of sitmitar length. ..-.._, vee MM, aequatis

Bhrenherg (Fig. 19:2)

Fig. 14. 1, Monommmata oclices Myers; {a) lateral; (b) trophi, ventral; (¢) bophi, lateral. 2, Mf. ceqvotis Ehrenberg:

(a) lateral; (b) trophi, ventral; (c) trophi, lateral, 3, Mf. aeschyna Myers: (a) lateral; (b) trophi, ventral; (¢) traphi,

lateral. 4, M4. arndti Remane: (a) lateral; (b} trophi, ventral; (c) trophi, lateral. 5, ML dentata Wulfert: (a) lateral;

bh) trophi, ventral; (c) trophi, lateral. 6, Mf. diaphora Myers: (a) lateral; (by trophi_ ventral; ¢c) tropa, lateral. 7,

J, grandis Tessiit, (a) lateral) (>) frophi, oblique (rootal, () rophi, tater; (0) Lophi, dorsal, 8.4%, deneiseta (Miller):

(a) Jateraly (b) (rophi, ventral; tc} Grophi, Jateral, |, 3, 6 8 after Myers (1930); 2 after Myers (1937); 4 afler Koste

(1972); 5 aller Wulfert (1940). Scale lines: adult 50 jm; traphi 10 um.

halt W. KOSTE & R, J, SHIEL

Toes of dissimilar lenath....., ’ woah

Stomach with sacs, M. viridis

"Myers (Fis. 208)

Stomach without savs_-.....2, 0.0... 0-2... 2

Dorsal antenna single, .

Dorsal antennae paired,.,.,., weet

Body +200 jan; rightleft toe ratio “<42. Ne a he

a wvoetititreets. Af, arndti Remane (Fig, 19:4)

Body < 200 poi rightileft toe ratio >1.2....--

San M. actices Myers {Fig. 19: (4

Rami with teeth on inner margin...,..... ;

Rami without teeth on inner margin

Toes <200 pm: rophi 25-35 am. a

Toes >200 pm; Gophi <25 yan... .M. diaphara

Myers (Fig. 19:4)

Right toe >168 pms rami lyrate; unei with 5-6

linear leeth..,,....46 phoxa Myers (Fig. 20:2)

Right ine <165 pm; rami triangular; unci single-

toothed... .. M, aeschyna Myers (Fiz 19:3)

Raini with [~S inner margin tooth pairs. ...- 9

Rami with | or 2 occasionally unpaired inner

murgin teeth, MM dentara Wulfert (Fig. 19:5)

Sh, Unej with plate-like (eeth terminally with finger-

ike extensions....M. grandis Tessin (Fig. 19:7)

Unci only with Jong tlagger-like teeth, . . 10

10(9), Cnei teeth paired... .- eptt fas Mh meoutara

Harring & Myers (Fig. 20:1)

Unei teeth single...-.2...22. 2... M. longiseta

(Miiller) ¢Fiz. 19:8)

2).

3(2).

43)

5(3).

6(5),

Tih).

(5).

Monammata actices Myers

FIG. 19:L

Maonomimata actices Myers, 1930, pp. 394-5, Figs 4-7.

DBipe locality: Permanent bodies of acid-waters in

Atlantic County, New Jersey,

Hojotype: Not designated. ?7Myers Coll., AMNH,

NY.

Description: Body slender, cylindrical, tapering to

foot; head clearly separated trom trunk by dorsal

projections and folds of integument, which is

striate; single dorsal antenna tubular, retractile,

when extended has papillate base: lateral antennae

with two associated round reddish or clear areas in

lumbar region, corona oblique, typical; mastay

minute, simple virgate; fulcrum long, todtike; rami

slender, lyraie, crooked downward medially; alulae

todtike; unci with two shotr teeth; pair of thin

lamellar plates bounded by dorsal pair of unc teeth:

oesophagus short; gastric glands round: no

constriction between stomach and intestine: foot

glands, bladder and retrocerebral sac small; eyespot

at posterior end of ganglion; egg spinulate,

Body length 150-195 zm; trophi 22-24 pm; right

toe 2000-210 am; lett toc 150-170 ym; subitaneous

ege dR« 58 um, spinules 15 pm.

Ecology; Europe, N. America, in moor pools and

lightly acid waters. Rare, billabong ai Jabiluka, NIT;

Goultiirn R. billabong at Seymour Vic; Cradiz ML

Nat. Park, Tas. 23,0-25,0°C, pH 5.7-6.2, 48 »S

m ', DO 3.7 me] '.

Literature: Donner 1978; Koste & Shiels; Koste et ¢i/,

1988.

Comment) The single dorsal antenna was

considered by Wulfert (1960) co bé dowbi ful; Koste

(1978) suggested that paired antennae were

overlooked by Myers, and thar the appearance of

M. actices is identical with M. arndli Remane. The

dimensions given in Koste (1978) are contradictory,

and there are differences in the trophi as figured

by the two authors. We retain both taxa here until

finer resolution of them can be made,

Monommiata aegualis Ehrenberg

FIG, 19;2

Nolommata longiseta aequelis Ehrenberg, (R42, px 134,

Monommata aequalis, Eyferth, 1878, p. 84.

Type focality; Berlin.

Holotype: Not designated.

Description: Long, slender, hyaline body indistinctly

demarcated from head; single dorsal antenna: rami

lacks inter margin teelh; rami, seen laterally, at righe

angles to fulcrum; unci uniformly lamettate, (luted

or with marginal denticles; fulcrum rodlike;

manubria bilaterally lamellate; subcerebral glands

absent; toes of equal or similar length; red or yellow-

orange vesicles beside: intestine,

Total length 200-227 ym: toes 110-120 pm.

Ecology: Rare in vegetated waters, Europe, 5S,

Anverica, Early record fram Qld. In our Tasmanian

cOlleclions from Great Lake, Central Plateau

S.0-1L.0°C, pH 7.6-7.8, 9,0-13.4 pS em-'.

Literature: Koste 1978; Koste ef al, 1988,

Monommata aeschyna Myers

FIG. 19:3

Monommata aeschyne Myers, 930, p. 387, Fig. 24:4-6

Type locality: Corday Creek, Atlantic County, New

Jersey.

Holotype: Not designated, ?Myers Coll., AMNH,

NY,

Description: Body elongate, fusiform; head

demarcated by slight constriction; body tapers

Bradually to 3-segmented foot from midline: single

dorsal antenna on low papilla; mastux virgate,

stall, simple; fulcrum Jong, straight, slightly dilated

distally; rami triangular, without teeth or denticles;

alulae prominent; manubria simple rods, distally

curved, with medial blunt tooth-like process

dorsally; unci single-toothed; gastric glands,

tetrocerebral sac small; eyespot on ventral side of

ganglion.

Body length !30-150 jum; right toe 150-165 jum;

left toe 120-145 pm; trophi 25-35 xm.

ROTIFERA FROM ALISTRALIAN INLAND WATERS tdi

Distribution: In Spliagnuin, melt-water pools,

Europe, N, America. Rare, billabong at Seymour,

Ryans 2 Billabong, Wodonga, Vic., and shallow

pools, western Tas. 13.0-27.0°C, pH $.4-6.9, 25-292

wScm ', TDS 16.2-21.b mg !~, 1.1-7.3 NTU.

Literature: Koste (978,

Monommate arndt) Remane

FIG, 19:4

Monommiata arndit Remane, 1933, p. 567-68,

Type locality; Moor pool near Kiel, Germany.

Holotype: Not designated,

Description: Resembles M. actices. Paired dorsal

antennae alse on retractile papillae, lateral antennae

tubular, mastax with two paired and one single

salivary gland; retrocerebral sac with distinct paired

excretory ducts present; two tubular subcerebral

glands; gut contents generally yellow-gold; Koste

(1978) notes Unat paired red lumbar bodies develop

in response to food intake, and possibly are

excretory deposits; ratio of body/toc Icngth changes

during development; subitaneous egz spiny, male

egg smooth-shelled, Male undescribed,

Length 210-S00 pum; right toes to 250 gm; left toc

ta 210 pm; subilaneaus egg 35x75 om; male ceg

48%40 um.

Ecology; Burope. Eats phytoflageliaies, e.g. Syrwna,

by holding colonies in the corona and sucking cell

contents. Single record: billabong of Goulburn R.

at Alexandra, Vic. 20,0°C, pH 7.2, DO 9.1 me I!

Literatiire: Koste 1972, Koste & Shiel 1980,

Mornommata dentate Wulfert

FIG, 19:5

Moanommaia dentata Wulfert, 1940, p. $78, Fig 22

Type locatity: Sphagnum poal, Birkhorster Moor,

beiween Svharfeabriick and Neuendorf.

Halotype; Not designated,

Descriptian Body squat, glassy; stomach and

intestine yellow-grey; right toe shorter than budy;

coloured vesicles absent, although mastax

sometimes contains colourless balls/spheres; single

dorsal antenna; trophi structure variable; rami with

paired of single (or missing) teeth an inner margin;

pleural rod present; unci two-toothed; fulerum

spatulate distally, basal apophysis (medial pointed

process on Fulcrum) present; manubria bilaterally

lamellate. not crooked terminally.

‘Total length to 400 wm; right foe 115-200 py left

toc 89-160 pm; trophi length 16-22 pm.

Ecology; Previously known only from Europe,

Described by Koste (1978) as pH tolerant,

¢urytherm, otigo-euryhaline. A population closely

resembling M. dentata was collected from Tasmania

(Golden Valley) in 1947, with a second record from

Ryans 2 Billabong, Wodonga in Oer 1990.

15=23.0°C, PH 6,2-6.7, 140-310 pS ¢m-', 4.0

NTU,

Literature Koste & Shiet 19876,

Mortomrmate diaphore Myers

FIG. 19:6

Monommata diaphora Myers, 1930, p. 388-3, Figs 7-9,

Tipe locality; Acid water of the littoral region of

ponds and lakes in Atlantic County, New Jersey,

Holotype: Not designated. ?7Myers Coll., AMNH,

NY.

Description! Body very long, cylindrical;

characteristic sWelling, above anterior part of

stomach; right toe shorter than body; corona

typical; single dorsal antenna; mastax virgate;

fulcrum without basal apophysis; ramt triangular

in ventral view, laterally right-angled, without teeth

or dentictes; alulae prominent; manubria reduced

to simple rods, crooked terminally, attached to rami

by thin lamellar plates; unci single toothed; mastax

with two large confluent salivary glands; icyespot

with lens.on ventral side of brain at posterlor end;

retrocerebral sac with rudimentary excretory duct,

Body length 225 um; right tue 260 pm; left toe

225 am; trophi 25 «am.

Ecology: Littoral of weakly acid waters (pH

6,2-6.4), southeast Europe, N. America. Single

record from Rushy Billabong, &. Murray fear

Barnawartha, Vic. (Shiel unpubl),

Mononvnata grandis Tessin

FIG. 19:7

Mondrian prardis lesain, 1390, p, 151, Fig, prt-12

Tipe locality: Rostock, eastern Germany.

Holotype: Not designated.

Description: Body elongate, fusiform; single dorsal

antenna on raised prominence; lateral antennae

ysual; characteristic red pigment spots beneath

faleral antennae; foot indistinctly two-jointed,

mastax of intermediale type; fulcrum similar length

to tami; rami with thin lamellae medially; large

basal apophysis; large alulac; inner margins of ram

with 25+ comb-like ventral denticles and two pairs

of four Jong, slender opposing oral teeth; each

uncus with platé-shaped ventral tooth, ending in

five tooth-like projections at tip, and distal rod-like

tooth; manubria crooked distally, lamellate

proximally; retrocerebral sac small, clearly ducted

to corona surface; no subcezebial glands; mastax

has. conflyent salivary glands; eyespot ventral at

posterior end of ganglion.

Total length 350-680 ym; body 190-240 wm; right

toe 210-470 nm; left toe 180-336 wm-

Ecology: Cosmopolitan, rare, generally single finds

I42 W. ROSTE & R. J. SHIEL

in littoral of standing waters in Europe. Rare, Qld,

Tas,, Vic, 16.7-27.0°C; pH 4,52-7.2; '25.4-60.0 1S

cm'; TDS 16.2-24,9 mg 1-1; 1-9.4 NTU,

Literature: Koste & Shiel 1980, 1983, 1987b.

Menommata longiseta (Miller)

FIG; 19:8

borticella longiseta Miller, 1786, p. 295, Fig. 42: 9-10.

Monommata loagiseta; Bartsch 1870, p. 344.

Type locality; Copenhagen.

Holotype: Not designated.

Description: Body slender, elongate, with

transparent integument marked with closely spaced

‘striae; resembles M. dentata, Can be distinguished

by trophi differences: ram bent at right angle near

mid length, with long slender tooth at angle on each

tamus; right uncus has three long slender teeth, left

uncus two; fulcrum lacks basal apophysis;

manubria broad lamellar proximally, distally

rodlike, outward curving; length of toes variable.

Total length 200-250 «m:. body length 86-115 gm;

right toe 155 ym; left toe 120 um; trophi 15-16 um.

Ecology: Casmepolitan in vegetated waters. Earlier

records from N.SW., Qid, Vic. In our material a

single record from Scottsdale, Tas. 15.0°C, pH 7.1,

105 «S om !.

Comment: This taxon may not be as widely

distributed in Australia (or globally) as the records

suggest, because of confusion with other species by

earlier authors.

Literature; Shie! & Koste 1979; Koste ef al. 1988.

Fig. 20, 1, Monomtata riaculata Harring & Myers; (a) lateral; (b) trophi, oblique frontal view: (¢) trophi, ventral;

(d) trophi, tateral. 2, 47, phoxa Myers: (a) lateral; (b) trophi, ventral; (c) tophi, lateral. 3. AZ, viridis Myers: (a)

lateral; (b) trophi, ventral; (c) trophi, lateral. 1, 2, after Harring & Myers (1924); 3 after Koste (1972), Scale lines:

adult 50 ym; trophi 10 pm.

ROTIFERA PROM AUSTRALIAN INLAND WATERS 34

Monommata maculata Myers

FIG. 20ht

Monommata grandis Harring & Myers, 1924, p, 538,

Fig. 43; 6-10,

Monommaia maculata Myers, 1930, p. 385.

Type locality: Not specified. “~~, common in

weedy ponds..."

Holotype Not designated. 7Myers Coll, AMNH,

NY,

Description: Body slender. elongate; integument

striated; foul two-segmented; toes variable, but

always unequal; mastax intermediate between

virgate and forcipate; fulcrum lrontally short,

dagger-like, laterally planklike; abnormally large

basal apophysis; inner margin of rami with unique,

complex denticulation - three groups of \eeth:

ventral group with 12-14 comb-tike teeth; middle

oral group with four large curved, pointed

interlocking teeth; dorsal group with three long

needle-like teeth; unci with three long clubbed teeth:

manubrium terminally crooked with ventral strong

lamella; retrocerebral sac present.

Total length to 680 pin; right toe 340-470 um; left

toe 270-410 jun; trophi 26-34 pm,

Ecology: Europe, Asia, N. and 5. America in

vegetated waters. NT, Old, Tas,, W.A. 19.0-27,0°C,

pH $.4-6.9; 25,4-33.5 48 em ', TDS 162-214 mg

I-', 1.1-2.2 NTU.

Literatiire; Koste 1978; Kaste ef al, 1988,

Monommata phoxa Myers

FIG, 20:2

geil phaxa Myers, 1930, p. 295-6, Fig, 26;

Tepe fecality: Acid pond tear Brglish Creek,

Atlantic County, New Jersey,

Holotype: Not designated. ?7Myers Coll. AMNH,

NY.

Description: Body long, cylindrical; singh: minute

dorsal antenna; trophi virgate, with long, distally

spatulate fulcrum; rami lyrate in ventral views unci

with 5-5 linear teeth} two rods run from bases. of

teeth to tips of tami, marking limits of two thin

lamellar plates resting on sides of rami; manubria

with median lobulate projection, distally with slight

expansion and outward curve; retrocerebral sac

small,

Body length 150 jun; right toe 190 um; lett oe

140 um; trophi 35 pam,

Ecology: Europe, Asia, N. America, ?Africa, iti acid

moor pools, Tas., Vic. 24.0-27,0°C, pH 5,4-5,8,

25-119 pS cm', TOS 16.2-76.3 me ]-', 2.0-2.2

NTU

Literature Koste ef al. 1988.

Monormmara viridis Myers

FIG, 20:3

% Monavie viridis Myers, 1937, p. U-1, Fig. 13, 19,

Type locatity: Atlantic County, New Jersey. * — . . int

the littoral region amang Sphegeurn in bodies of

acid waters,”

Holotype; Not designated. ?Myers Coll,, AMNH,

NY. Paratype; Cat No, 604, AMNH,

Description: Body slender, ne constriction behind

head; dorsal antenna double papillose projection;

stomach ends in four blind sacs; put contents

yellow-green spheres; gastric glands present; mastax

modified virgate; fulcrum short, in lateral view

curved, tapering; rami slender, lyrate, bent dorsally

near mid-length at approximate right angle; dorsal

portion has small tooth on inner margin; unci

reduced to thin lamellar plates, posterior edges

thickened to resemble slightly clubbed teeth:

manubria curve dorsally, have small digitifarm

process near mid length; retrocerebral sac round,

clear, ductless,

Total length 382 um; body length 166 ~m: right

toe 216 pm; feft toe 150 pm; subitaneous ceg

63«$2 um,

Ecalogy; Common in Sphegniamn, pH 4.5-6,5, N.

America, Europe, Single record from dune lakes

area, western Tas. 17,0°C. pH 3.1, 20.6 pS om |,

Literature: Koste 1978; Koste et af, 1988,

Genus Noidmmata Ehrenberg

Notommata Ehrenberg. 1830_ p: 46.

Type: Vorticella aurita Miller, 1786 = Notommoerta

aurita (Miller)

Type locality: Copenhagen.

Body cylindrical, spindle-shaped, sac-like,

conical, or with lateral alae; neck suture evident;

caudal tail usually present; foot short, often

2-seginented, occasionally indistinctly segzinented;

also rudimentary; with and without caudal sensillac.

always with two toes; cuticle generally soft, often

with longitudinal striae; carona broadly triangular,

displaced ventrally ("Netommata type”) (Fig. 21:1b),

in some species extending into pronounced “chin”;

auricles generally large, narrowly to widely

separated, retractable; apical field small; mastax

Virgate; (rophj asymmetric in most species; unci

often with accessory teeth, occasionally also with

rudimentary fulcrum, long and plank-like, straight

or curved; pleural rods, epipharynx and oral plates

may be present; hypopharynx muscle attached to

fulcrum; stomach and intestine separated by weak

coistriction; retroacerebral organ generally wedl-

developed; protoncphridia visible in head- 4-6 pairs

of flame cells; cerebral eyespats present or absent;

Wt W_KOSTE & R

Male fot khown for all species. Fifty-five

Nolommata specics are listed or described in Koste

(1978); 12 of these are known from Australia, with

two endemic species described subsequently, These

are keyed below, however we caution that in view

of gross morphological similarities between sume

taxa, and ready confusion of juveniles of large

species with adulcs of smaller forms, the key can

be used reliably only with living adults. For

preseryed material, and doubtful live material,

trophi dilfferénces.as described and figured enable

accurate specific identification.

Key to species of Notommata recarded from

Australian inland waters

1. Cylindrical, fusiform ar conical body, wirhour

lateral extensions (alaey.... 226.02 02008 22

Body with pronounced alae,....---.... 0.0...

_ N. Spine nov. noo. TFig. 24:4)

Rump with projecting digitiform process (Pig.

7A0 Le a SR Ae ee 3

Rump withouc obvious appendage... . 4

Body >500 yan; trophi +80 pm; caudal process

with articulated tip, does not reach base of tocs

N. capers (Ehrenberg) (Fig. 23:1)

Body <200 um; trophi <30 pm; caudal process

not articulated, extends past base of toes,

oN, deipus Elueuberg (Fiz. 25:1)

Caudal ail’ more or less covering base of foat

Rump rounded, with no obviaus iil -._- 12

Tacs <0 yin; body vermiform; auricles indistingt

aa oh. S88 S N. cantorta (Stakes) (hig, 223

‘foes >10 pmy body fusiform or glbbous; auricles

GISINED. + tu msses Son tt Petey fi

Toes >3S pm..... 2. 22. eee ee - 7

Toes <35 wn

Toes 40-42 pm; body < 310 ui trophi 40. 45 am

oN. doneta Harring & Myers (Big, 23:3)

‘Toes 35-75 ium; body 300-800 um; trophi > 75 wm

coy N. puchyura (Gosse) (Fig. 24:1)

Trophi “<40 yen toes 16 20m; body <350 zm

. AN, auring (Muller) (Fig, 21)

Trophi >40) utd; toes 15-35 jum; body 300-750 yan

a(S).

Tif).

B16].

beter

Wish Mastax large. frophi 100 pm; body 500-750 yom

ON. collaris {Bbresibers) (Fig. 21:4)

Nlastux sirnaller, trophi <70 pm; body 300-680 jan

MTT ere , 1a

Tail with median lobe and two tateral lobes; traphi

45-60 jim wed fete i

Tail rounded; trophi 60-70 tt

.-N. glyphura Walfert (Fig. 23:4)

Toes 30-35 yin; traphl symmetric, unei singte-

HOOMINED, uuu cee eens N. pseudocerberns

De Beauchamp (Tig. 24:2)

Toes (5-35 am; trophi asymmetric, uncl

J-5-toathed ...N, cerberus (Gosse) (Fig, 21:2)

Toes 24-30 pmy trophi 30-35 pm, WN. eyrrepus

Goase (Pig. 23:2)

foes <2) pin, Lroplil <25 pm -.---- 4.3

19).

119),

(24).

J. SHIEV.

43(/2). Body 169-200 pm; toes 8-10 win, trophi 24 pam

..... oN, silpha (Gosse) (Pig. 24:3)

Body ‘<140 pin; toes 12-16 pm, trophi 20 pm,

ei pleg 4 JN, tylert Kosio et al, (Pig. 25:2)

Nolommata aurita (Miller)

FIG, 2h:

VorticeHa aurita Muller, 1786, p, 288, Fig, 41:1-3

Notonmmnata aunt’ Ebrenbete 1830, p 46,

Dpe locality: Copenhagen.

Holotype: Not designated.

Descriplion; Body short, stocky, integument

transparent, with Jongitudinal striae; loot

2-segmented, first segment nearly covered by

tongue-shaped caudal projection (“tail”); small

“pee” between toes; auricles. short; corona with well-

developed “chin" region; mastux virgate; trophi in

ventral view with left ramus more strongly

developed; fulcrum long and stout, distal end Y-

shaped for attachment of hypopharynx muscle;

tami subsquare in ventral view, bent at right angles

lo anterior point; unci plates with single main tooth,

basal plate with 1-2 rudimentary teeth; small

preuncial teeth at tip of right uncus; manubria with

broad angular anterior plate; triangular oral plate

and thin pleural rods presetit; retrocerebral sac large,

spherical, generally opaque; cerebral glands

rudimentary; eyespot large, beneath retrocerebral

sac, only visible laterally.

Total Jength 250-350 pnt; toes 16-200 4m; trophi

34-36 pm; male tS80 jam.

Ecology: Cosmopolitan i fresh to brackish water,

abundant in spring and autumn in Palacarctic

waters; preys on bdelloid and other rorfers as well

as algae. Early records from N.SW., Qld, Vic,

recently collected (Sept-Oct. 1990) in Ryans

Billabongs, Wodorga: 13.0-17.0°C, pH 6.85-6.92,

274-292 xS cm !.

Literature; Koste 1978; Shicl & Koste 1979.

Notommiare cerbertus (Hudson & Ciosse)

FIG. 21:2, 3

1a Conens cerberus Hudson & Gosse, 1886, p, 34, Fig,

" Notorimat cerberus; Beauchamp tY68, py, 401. Fie.

Type locality: Sandhurst Wood, Herkshire.

Holotype: Not designated.

Description; Body slender, integument flexible;

slight transverse folds between head ahd neck: tar!

rounded, with median lobe and rudimentary lateral

lobes; foot with two joints; small setose pil in centre

of dorsal side of second joint: toes slender pointed,

1/15 total length; corona extends ca. 44 length of

body ventrally; auricles short, with robust cilia tufts}

mastax virgate; Y-shaped Jamellac at base of

ROTIPERA FROM AUSTRALIAN INLAND WATERS ids

big. 2h. 1, Neroniniata aurita (Miller): (a) dorsal; (b) lateral; (cl) Grophi, ventral; (d)} trophi, lateral, 2, No cerberus

{Gosse): (a) dorsal; (b) patterning of integument, dorsal; (c) trophi, ventral, manubria omitted; (d) fulcrum, lateral;

{e) unci; (1) foot and toes, lateral. 3, NM. cerherus longinus Wulfert: (a) dorsal; (b) trophi, ventral; (c) fulcrum, lateral;

(d) unei; (e) abdomen, foot and toes, dorsal: (f) manubria. 4, N. collgris Ehrenberg: (a) dorsal; (b) trophi, ventral;

lines adult 50 pn trophi 10 ym,

fulcrum to which hypopharyngeal muscles attach;

rami asymmetric, inner edges striate but not

denticulate, outer margin with asymmetric

apophyses and short toothed alulac; right more

developed than lett; pleural rods present; unci plates

with 3-5 teeth; ventral main tooth has small

accessory teeth: four foot glands, one pair large,

one pir small; retrocerebral sac pyriform teaches

almost to end of mastax; eyespor large, lenticular.

‘Total length 300-600 pm; toes 15-35 rn; trophi

46-60 pin (46 am trophi has 42 pm rami; 32 ~m

fulcrum; 234m unci).

Eeology: Sphagnum, periphyton, waters in northern

hemisphere. Omnivore, cats rotifers, desimids,

diatoms, flagellates. Early records from N.SW., Old,

Vic, Recently collected (Sept, 1990) from a billabong

of the Mitta Mitta River at Tallandoon, Vic.

Liferqture: Evans 1951; Shicl & Koste 1979.

Ma W. KOSTE & R. 5. SHIEL

Comment: A variant described by Wulfert (1961)

as N. cerberus vat. longinus occurs in eastern

Tasmania. Comparison of the morphological

differences shown in Figs 21:2 and 21:3, particularly

trophi, suggests that specific status may be

warranted. 19.0°C, pH 6.8, 215 pS em-!.

Notommata collars Ehrenberg

FIG, 214

Notommata collaris Ehrenberg, 1832, p, 134, Pig. 4:01.

Type locality; Berlin,

Holotype: Not designated,

Description: Body short, stoul, integument

‘eathery’, transparent; neck long to accommodate

large mastax; anterior sutures well marked;

abdomen bulges to twice anterior width to rounded

posterior with short broad tail which covers ca, 4

of first of two foot joints; toes short, conical about

1/30 total length; auricle ciliation continuous with

corona; corona elongate ro form prominent post-

oral chin; mastax vitgate, trophi asymmetric, the

largest (in proportion to body) of all Notommata

spp; fulcrum with V-shaped lamellae for muscle

attachment; right ramus with broad lamellar tooth

with denticulate lamellar margin; left ramus

excavated; unci with three teeth, only ventral tooth

developed; inner unci margin striated; manubria

Jong and broad with wide thin lamella extending

nearly to posterior end on dorsal margin; pleural

Tods present; fool glands slender; retrocerebral sac

long, pyriform;: eyespot ac posterior end of brain.

length 500-750 pm; toes 25-32 wm; trophi 100

am; male 240 jams vesting egg 170%«215 (415)am.

£eolovy, Palaéarctic, Nearetic, abundant in

dystrophic waters in periphyton, Eats Closterizim

and filamentous algae, N.SW., NT.

Literature: Shiel & Koste 1979; Koste 1981.

Notormmata conlarte (Stokes)

FIG. 22

Diglena contorta Stokes, 1897, p. 630, Fig. 1425.

Notommaia coatorta, Harring 1913, p. 78,

Tope locality: “. . - shallow clear-water pool ina

rocky wood near Trenton, New Jersey”,

Holotype; Not designated,

Description; Body elongate-subcylindrical,

vermmiform, gibbous posteriorly, integument notably

flexible; head rounded, convex, with small hook-

like proboscis beneath which frontal border is

conspicuously emarginate; rump depressed into

cylindrical tail overhanging and almost completely

surrounding short papillate foot; two small conical

divergent toes; buccal field elongate, almost

horizontal, excending ventrally ca. 1/3 body length;

‘chin’ absent; lateral ciliated auricles small, rarely

big, 22. Notommata cantaria (Stokes); (a) dorsal; (by

Jateral; (c) trophi, ventral; (cd) traphi, lateral. After

Harring & Myers (1922), Seale lines: adult 50 jum} trophi

em.

extended; two dorsal and two lateral antennac; small

yellowish-orange cerebral eye; retrocerebyal sac with

bacteroids, subcerebral glands short: trophi simple

forcipate; lulcrum jong, dagger-like, with basal

apophysis and terminal hook-like dorsal curve; uncl

single-toothed; rami simple, with strong right-

angled bend.

Total length 206-300 um; toes 7-10 pm; buccal

field 83-86 ym; trophi 17-20 pm (folerum 12 pm,

manubria 13 pm}.

Ecolagy: In acid waters, in periphyton of submerged

plants in pools and lakes. Previously known from

Palearctic and Nearctic (Koste 1978), First record

from Australia 19 Sept. 1990, Ryans 1 Billabong,

Wodonga, in Azol/a. Subsequently also found in

nearby Ryans 2 In Rieciocarpus, 20.0-22°C, pH

§.39-6.57, 114-274 pS cm |.

Notominata copeus Ehrenberg

FIGS 2, 23:1

Notornmutus copeuy Ehrenberg, 1834, p, 213,

Type locality: Berlin,

Holotype: Not designated.

Description: Large species, elongate body, truncate

posteriorly; amterior sutures distinct; tail a lone

round, conical projection with articulated tip

ROTIFERA PROM AUSTRALIAN INLAND WATERS 447

Fig. 23, 1, Nofommiata copeus Bhrenberg: (a) dorsal; (b) trophi, ventral; {c) unica; (d) fool and toes, dorsal, 2, .N.

eyriopus Gosse: (a) dorsal; (b) lateral; (c) trophi, ventral; (d) trophi, lateral. 3, NM. donera Harring & Myers: (a)

dorsal; (b) lateral; (c) trophi, ventral..4, WV. glyphura Wulfert; (a) dorsal; (b) trophi, ventral; (c) uni, 1a, o d after

Donner (1954); Ib, 2 after Harring & Myers (1922); 3 after Harcing & Myers (1924); 4 after Wulfert (1935). Scale

lines: adult 50 xm; trophi 10 am.

segment; foot with two broad segments, with setose

papilla at distal end of second joint between toes;

toes ca, 1/16 total length, long and slender, slightly

decurved; very long ciliary auricles (seldom

extended) and exceptionally elongated chin; lateral

antennae setae long; mastax virgate, asymmetric,

left side more developed; rami tnangular with broad

lamellar tooth on right ramus, socket on left; left

uncus with large ventral tooth and three smaller

teeth; right ramus with tour undeveloped teeth;

manubria and pleural rods as N. collaris;

retrocerebral sac and subcerebral glands present;

eyespot large, at posterior end of brain

Total Jength 500-1100 um; toes 40-65 pm; trophi

80-100 pm; male 280-350 pm; toes 35 pm.

Ecology: Cosmopolitan in algal-tich standing

waters, pH 4-6.5, temp. 15-30°C, Eats filamentous

algae (Spyrogira, Zygnema, Mougeoria) swallows

stnaller Filaments in short pieces. N-SW., N-T., Old,

Vie

Literature: Shiel & Koste 1979; Koste L981.

Lik W. KOSTE & R, J, SENEL

Nolommata eyriopus Hudson & Gusse

FIG, 23:2

Noatommara cyrionus Hudson & Gosse, 1886, 2: 22. Paz.

17:7.

Tipe locality: Wideombe Pond, Bath.

Holotype; Not designated.

Description: Body stout, integument flexible,

transparent; single obvious suture between

neck/abdomen; abdomen rounded posteriorly

without tail; foot 2-segmenred; toes long (1/7 total

length), curved and tapering; auricles short, cillation

continuous with corona; mastax virgate; tropht

slightly asymmetric; Fulcrum stout; rami

approximately triangular in ventral view; inner cdgc

of right ramus concave, Icft convex, both

denticulate; alulae well-developed; right ramus has

four short curved preuncinal teeth at tips, left has

five; unci with single: main tooth and rudimentary

second tooth (two thin slightly curved pleural rods);

retrocerebral sac, subcerebral glands and eyespot (at

posterior end of brain) present; male known.

Total length 175-250 ym; toes. 22-28 pm; trophi

30-35 um (matiubria 18-23 zm, fulcrum 18-21 pm;

rami (2-14 wm) male (otal 139 ym: toes 20. em.

Ecology: Cosmopolitan in periphyton in littoral of

flowing and standing waters. Single early record

from N.SW., and a Sept. 1990 record from Ryans

I Billabong at Wodonga, Vie,

Liferature; Koste 1978; Shiel & Koste 1979.

Norommata dereta Harring & Myers

FIG, 23:3

Notomautte doneta tlarring & Myers, 1924, p 448-St,

Fig. 22: 1-4.

Type focality: Starvation Lake, Vilas County,

Wisconsin.

Holotype: Nol designated. YMyers Coll., AMNH,

NY.

Deseripuion: Body transparent; prominent tail with

rounded median lobe; foot with two short joints;

toes ca. 1/6 total length; corona extends to small!

chin; mastax virgate, Lrophi nearly symmetric; rami

broadly triangular in ventral view, armed on inner

margins with ca. 12 minute teeth; fulcrum long,

slender, slightly curved distally; unci with well

developed main tooth and rudimentary second

tooth; left uncus with seven smal! accessory teeth,

right uncus with six; manubria long, slender, with

small basal plale; pleural rods and two slender

epipharyngeal rods present; tetrocerebral and

subeercbral glands large. Distinguished fram close

relatives. (yurita, cyriopus) by long, peculiar tocs,

Ecoiogy; In Sphagnum, periphyton on submerged

plants, Europe, N. America, A notommatid

resembling WV. donela was collected from a humic

pool adjacent to L, Strahan, W. Tas. 14.0°C, pRB

3,35,

Naomi glythura Waller

FIG, 23:4

Notommata glypaura Wulfert, 1935, p. 590, Fig. Ja e¢.

Type tocality; pouls near Mersehurg, Germany,

Holotype: Not designated.

Deseripiion: Body with wide, plump abdomen; tail

rounded; head and neck clearly defined; foot

covered by tail (except distal end of terminal

segment); toes straight, tapered to blunt tips; auricle

of medium size; corona extends venirally to form

chin: retrocerebral sac brown to black; subcerebral

glands long, wide; mastax large; trophi asymmetric;

rami with hook-like alulae exceptionally long with

inner margin teeth; unc plates with main and three

secondary leeth on right, uncus fused to rectangular

plate; three preuncial teeth in front of main tooth;

ynanubria dilated distally.

Length 325-S00 pm; toes 20-24 nm; trophi 60-70

pm; male to 280 pm} resting epg (spiny)

115%150. pm,

Ecology: Europe, in fresh to brackish water, in

littoral standing and flowing waters. Eats algae,

rotifers, scavenges dead microcrustacea, Recorded

front billabongs of the Goulbuin and Murray, Vic.

13.0-17,8°C, pH 6,2-7.2, DO 8.9 mg I-|, 74 pS

om’, 8 NTU, also from Nankeen Billabong,

Mayela Ck, NT.

Literature; Koste 1978, 1981,

Nolammata pechyura (Hudson & Gosse)

FIG, 24:1

Copeus pachyures Hudson & Gosse, ARG: 2, p. 31, Fie

16:4.

Notorunatu pachyura: Warrine 13, p, 79.

Type lacality, pools, Sandhursi, England.

Holaiyne: Not designated,

Description; Body fusiform, transparent; anterior

sutures distinct; lobulare tail projects over first fuot

joint; fool 2-segmented with small papilla between

tocs; toes long (tu. 1/12 total length), conical;

auricles Jarge, stout; post-oral chim prominent:

mastax virgale, strongly asymmetric; fulcrum long,

stour, posterior cross section V-shaped; right ramus

with broad lamellar tooth, left with socket, alulae

large, also asymmictric; unci with one main and four

secondary tevth on left, one main and three

secondary on right; two sigmoid pleural rods:

somewhat quadrangular oral plate present; foot

glands long; retrocercbral sac extremely tone, clear

subcerebral glands small; large cyespot a! pesterice

end of brain.

ROTIFERA FROM AUSTRALIAN INLAND WATERS 9

Fig. 24. 1, Netommata pachyura (Gosse):; (a) dorsal; (b) trophi, ventral; (c) unci; (d) foot and toes, lateral. 2, N.

pseudocerberus De Beauchamp: (a) dorsal (b) trophi, ventral; (c) fulcrum & rami, lateral; (d) foot and toes, lateral;

(e) toes, dorsal, (fF) manubriwm, 3... silpha (Gosse): (a) dorsal; (b) lateral; (c) trophi, ventral; (d) unci. 4, N. spinata

nom. noy. (a) dorsal; (b) trophi, ventral. 1, 2a, b, d, 3 after Harring & Myers (1922); 2c, e, f after Donner (1954);

4 after Koste (1981). Scale lines: adult 50 pm; trophi 10 pm.

Total length 300-800 jm; toes 35-75 um; trophi

75-102 pm; male 300 zm.

Ecology; Cosmopolitan in aquatic plants of littoral

of pools, floodplain waters etc.. in Sphagnum. Eats

desmids, occasionally nematodes and rotifers. Early

records from N.SW., Qld, Vic. [In our samples, NvT,

and Tas. 14.7°C, pH 6.5, 23.2 «S em-!.

Literature: Koste 1978; Koste & Shiel 1986.

Notommatia pseudocerberus Beauchamp

FIG, 24;2

Notommata pseudocerberus Beauchamp, 1908, p. 400.

Type locality; ? France.

Holotype: Not designated.

Description: Body slender, integument ‘leathery’,

striated; tail has small tonguelike rounded median

150 W. KOSTG & R41 SHIFT

lobe, two lateral lobes; foot two-segmented, toes

conical, slender, ca. 1/12 body length; corona

extends down ventral side for ca. 1/4 body length;

rostrum above mouth, slight chin below; auricles

long, robust; mastax virgate, simple; fulcrum long

slender; rami broad, lamellate, without

denticulation on inner edges; unci with single

slender tooth, 4-5 accessory denticles on either side;

pleural rods present; salivary glands vestigial;

retrocerebral and subcerebral glands large,

Total length 400-680 pm; toes 30-35 pm; trophi

45-56 wm (fulcrum 24-27 ym).

Ecology: Cosmopolitan, rare finds in periphyton of

littoral, standing and flowing waters. Preys on sessile

ciliates, Single record from a stream near Bicheno,

Tas. 19.0°C, pH 6.8, 215 wS em-', 11 NTU.

Literature: Koste ef al. 1988.

Notommata silpha (Gosse)

FIG. 24:3

Diglena silpha Gosse, 1887, p, 2, Fig. 1:2.

Notommata forcipata In Hudson & Gosse, (886: 2, p.

23, Fig.. 18:1. [non-N. /orcipata Ehrenberg, 1838]-

Notommata silpha: Harring 1913, p, 79.

Type loculity: *. . . the middie of lreland”.

Holotype: Not designated.

Description; Body elongate, fusiform distally with

annular striae; foot rudimentary; toe apices curved

lightly inwards and ventrally elevated; evertile

ciliated auricles and chin absent; trophi prehensile

without sucking function; fulerum short, ventrally

stick-like and laterally plank-like; rami bifurcate,

asymmetric (Fig. 24:3c); unci with three large teeth,

nearly symmetrical, clubbed at tips; ventral basal

jamella of manubria with hoad-like process;

retrocerebral sac rarely with bacteroids; subcerebral

gland absent.

Total length 160-200 wm: toes 8-10 pm; trophi

24 yum.

Ecology; Isolated records from Nearctic and

Palaearctic in periphyton and in dystrophic waters,

Single unconfirmed early record from N.SW,, Not

seen in our material.

Literuture: Koste 1978; Shiel & Koste 1979.

Notammata spinata nom. nov.

FIGS 24:4; 25

Notommata pachyura ¢. trianealata: Koste (981, p42),

Fig. 16a-e, 18.

BT i a a fridneulatus Kirkman, 1906, p, 264, Fig,

Type locality: Magela Creek, N.T., Australia.

Holotype: Not designated,

Description: Body laterally expanded inte two

distinct alae so that total width is only slightly less

than total length (in preserved material alae may

om ol

Pig. 25. Netomsnata spinata nom. nov., photomicrograph,

exceed body length due to contraction of head and

foot); lateral tips of each ala armed with stout

triangular cusp (possibly retractible; partly

contracted individuals may have alary lips

withdrawn (Fig, 25); no dorsal “hump” (ef,

pachyura), triangular tail commences at level of

lateralantennae, distal end a blunt point at

beginning of first foot joint; head and neck sutures

distinct; two foot segment; toes conical, long, tapet

to pomt; corona ventral, “chin™ present; lateral

auricles with long swimming cilia; mastax modified

Virgate; trophi asymmetric; fulcrum expanded inte

fan at distal end, manubria curved posteriorly,

dilated distally; retrocerebral sac long, extending

past neck/abdomen suture; pigmented, subcerebral

glands large; eyespot hemispherical at posterior of

brain.

Total length 600-720 pm; toes 58-60 xm; trophi

140 am (fulerum 100 um; rami 80/60 wm; unci 60/48

gm; manubria 116/100 jm).

Ecology; Recorded from billabong of Magela Ck,

NT, A similar form also occurs in WA. (C.H.

Fernando pers. comm.)

Literature: Koste 1981.

Comment’ Koste’s (1981) record of this taxon

invluded figures and measurements but no

ROTIFLERA FROM AUSTRALIAN INLAND WATERS 1S

Fig, 26. 1, Nofemmata tripus Ehrenberg: (a) dorsal; (b) lateral; (c) trophi, ventral; (d) unci. 2, N. fv/ert Koste & Shiel:

{a) dorsal; (b) fateral: (c) trophi, ventral; (d) incus; (e) incus, lateral; (f) manubrium, lateral. | after Harring &

Myers (1922); 2 after Koste & Shiel (1986), Scale lines: adulc 50 pm; trophi 10 um.

description. The above description must be

considered provisional until more material is

available, however its status as a distinct species

rather than a form of N. pachyura is justified on

the basis of the characteristic alae morphology,

particularly the distal tooth, shorter toes, much

larger trophi and a differences in trophi

morphology. It differs from the winged taxon

figured by Harring & Myers 1922: Fig. 43: [-4

(Copeus triangulatus = Notommata triangtilata

(Kirkman, 1906).

Notommata tripus Ehrenberg

FIG. 26:]

Notommata tripus Ehrenberg, 1838, p. 434, Fig. 1:4.

Thpe locality: Berlin.

Holotype: Not designated.

Description: Body short, broad; integument rigid,

transparent; head and neck form a single segment,

dorsally humped; single transverse suture distinct;

abdomen rounded posteriorly; projection carries tail

and toes, no true foot present; dorsum has keeled

appearance. due to tapering of top half of abdomen

from about 1/3 its lengih towards tail; rounded tail

has spur-like projection with knobbed base, which

projects about 1/3-1/2 the length of toes; toes

relatively long (1/10 total length), straight, acutely

pointed; auricles short, stout, ciliation continuous

with corona; mastax virgate, irophi highly

asymmetric; rami roughly triangular; right ramus

152 W. KOSTE & RB. J. SUEBL

minutely denticulate, left ramus with four blunt

teeth; right alula of normal size; left alula hugely

dewloped; unci teeth 3/3, one main tooth:

manubria broad triangular lametlate proximally,

distally slender curved rods; foot glands large,

porilerm; retrocerebral sac almost spherical; eyespot

at posterior end of brain.

Total length 150-200 wm; toes 16-20 pm; trophi

26-H) am; caudal process 16-22 pm.

Ecology: Grazer of algae and periphyton, littoral

vegetation. Europe, N, America, Rare: N.SW., NT,

Tas. Vic. 22.0-24.5°C, pH 4.7-6.4, 45,5-114 pS

em-!.

Literature: Koxte 1978; Koste & Shie] 1986.

Notommata rpleri Koste, Shiel & Tan

FIG. 26:2

eg elaneneete tylori Koste, Shiel & Tan, 1988, p. 125-6,

ig, 2S.

Type locality: Arthur's Lake, Tasmania, Australia.

Holotype: SAM V4IL0, Collected by R. J. Shiel

22.1x.87.

Description: Very small, stout species; bouy dilated

distally to rounded rump; indistinctly seemented

lobulate foot with two short, acutely pointed toes;

mastax modified virgate; rami strongly convex on

outer margins, no inner denticulation; fulcrum

straight, spatulate disially; mranubria slender,

curved, with distinctive median ringlike

fenestration; foot glands elongate, club-shaped.

Total length 120-139 jam; toes [2-16 um; imcus 19

am; subituneous ege 30-45 x 50-65 pm.

Ecology: Endemic. Free-living in Arthurs Lake,

Tasmania. 8.0°C, pH 7.7, 17.4 pS em-!,

Genus Pleurotrocne Bhrénberg, 1830

Pleurotrocha Uhrenberg 1830, p 46,

Type: Pleurotrocha peiromyzon Ehrenberg-

Thpe locality: Berlin.

Notommatid rotifers. with short, stout iloricate

body; head and neck clearly demurcated; foot long,

cylindrical; toes short, conical; corona slightly

oblique with ciliated circumapical band and lateral

auricle-like tufts of long cilia for swimming; buccal

field finely ciliate; mastax virgate, trophi simple;

fulctum long, rod-hke; rami curved, triangular, not

denticulate; manubria long with reduced basal

plate; unct with only single tooth; hypopharyax

Jarge; mo retrocerebral organ; eyespot single,

Fourteen species are referral to this genus (Koste

1978). Only 2 pertromiyzon is known [rom Australia.

Pleurotrocka petromyzen Ehrenberg

FIG, 27:1

Pleurotracha petromyzon Ehrenbera, 1830. p 44,

Type locatity; Berlin.

Holotype: Not designated.

Description, Body short, stout, abdomen wider

than head; integument soft, transparent;

constriction berween neck and abdomen; abdomen

pyriform, lapers posteriorly to hase of foot; no tail;

foot 2-segmented, basal square, distal twice basal

segment length; toes short, conical; foot glands long

with reservoirs; cirgumapical band of cilia and two

lateral auricle-like tufts; mastax virgate, trophi

simple; rami triangular with large rounded alulae;

fulerum very long slender rod, slightly expanded

distally; unci triangular plates with one weak vertral

and second rudimentary tooth; manubria slightly

sigmoid with characteristic lobe projecting ventrally

from 1/3 along its length; pleural rods present; no

retrocerebral organ; eyespot minute at posterior end

of ganglion,

Total length 220-480 pm) toes 20-26 pm; trophi

30-37 am.

Feofozgy: Mass development after cladocerat

“pulses” or im bacteria or protozoa blooms, in ia

wide range of fresh to brackish waters,

Cosmopolitan. Necrophage - cleans out carapaces

of dead mites and microcnistacea, also eats sessile

ciliates, Rare, N.SW., Tas. Vic. 12.0-26.0°C, DO

10.4, pH 6.4-7.7, 114-3330 nS emo"

Liferatire: Evans 1951; Shiel & Koste 1979; Kaste

& Shiel 1986,

Genus Resticuia Harring & Myers

Resticusa Harring & Myers, 1924, p. 518.

Type: Furcularia mielandocus CGosse, 1887 =

Resticula mefandacus (Gosse)

Tipe focaliz): Woolston, England.

Very slender, Tusiform iloricate body, tapcring

gradually from mid-length to base of tocs: 1wo toes

short, with bulbous enlargement at base containing

mucus reservoir; corona frontal or oblique with

marginal wreath of short cilia and two lateral

auricle-like tufts of long swimming cilia} buccal

Field ciated; mastax virgate; fulcrum long, slender;

rami Iriangular, symmetrical with right-angled

median bend; unci with single well-developed tooth;

epipharynx. rudimentary or absent; two salivary

glands; retrocerebral organ a small round ductless

sac} eycspol a loose aggregation of réd pigment

granules in vacuoles of sac,

Seven species are recognized by Kosre (1978);

three of them ate knewn from Australia.

Key to spevies of Resticula recorded from

Australian inland waters

|, Foot fram end of abdomen to base of toes long (ca_

Iya Cotal bermpttth, occ ce cee K. gelida

(Harring & Myers) Mig, 27:2)

ROTIFERA FROM AUSTRALIAN INLAND WATERS 153

Fig. 27. 1, Pleurotrocha petromyzon Ehrenberg: (a) dorsal; (b) lateral; (c} trophi, ventral, 2, Resticula gelida Harring

& Myers: (a) dorsal; (b) lateral; (c) trophi, ventral, right manubrium omitted; (d) fulcrum, lateral. 3,.R. melandocus

(Gosse):; (a) dorsal; (6b) trophi, lateral; (c) trophi, ventral; (d) differenr views of ramus; (€) uneus. 4, R. ryssa Harring

& Myers: (a) dorsal; (bh) trophi, ventral; (c) manubrium; (d)fulerum, } after Harring & Myers (1924); 2 after Wulfert

{1935); 3 after Wulfert (1939); 4 after Koch-Althaus (1962). Scale lines: adult 50 um; trophi 10 ym.

154 W, KOSTE & R. J. SHIEL

Foor short and fidistinet, less than Jength of toes

£2600 Se, -— #2

TO Eyes a mass of red pigment granules.

.R, nyssa

Harring & Myers (Fig. 27:4)

No evespol discernible....., 0... RB, melanducesy

(Gosse, 1887) (Fig 27:3)

Resticula gelida (rlarting & Myers)

FIG, 272

P Eosphura gelida Harting & Myers, 1922, p, 642, Fig,

1-6.

Resticula pelida: Harting & Myers 1924, p, 519,

Type locality! Washington DC,

Holotype: Nol designated, ?Myers Coll., AMNH,

N.Y,

Descriplion: Body fusiform; transparent cuticle

with variable folding; abdomen occasionally

conspicuously wide; head clearly defined; tail

rudimentary; foot long (>3»% toes), without

segments; toes short (1/20 total length) tapering to

obtuse tips; corona frontal, circumapicul band

interrupted dorsally; mastax modified virgate (for

seizing prey); rami broad triangular; intricate

supporting fenestraled framework of slender, round

rods (Fig. 27:2c); fuleruny long and straight,

spatulate at distal end for attachrnent hypopharynx

muscle; unci reduced to single slender tooth with

three small preunvial tecth on left and two on right;

manubrium slender, curving slightly posteriorly

with ventral lobe anteriorly and triangular dorsal

Jobe; retrocerebral organ small, has pigment

granules, clustered af posterior end of ganglion

where eyespot would be,

Total length 350-600 jum; toes [8-30 wm; trophi

45-75 wm (in 45 pm trophi, fulcrum 26 wm, unci

17 om; manubria 32 pm); male 150-200 pra,

Ecology: Isolated occurrence in periphyion, still and

slowly flowing Water, Europe and Asia. N. America.

Single record, from Yarnup. Swamp, W,A. L7,0°C.

1800 pS cm |.

Lilerature: Koste et al, i983.

Resticula melandocus (Gosse)

FIG, 27:3

Purcularia melandocus Gosee, 1887, p. 2, Fig. bd.

Resticula melandocus; Harring & Myers 1924, p. 518.

Type locality: Woolston, England (lacustrine).

Hololype; Not designated.

Description: Body transparent, variable shape;

cuticle thin, with longitudinal striac; foor 3-4

segniented; toes variable, bulbed at base; toc points

curve inwards; single salivary gland to right of

mastax; unci with single main teoth, left uncus with

bev) preuncial teeth, right uncus with one; rami

triangular in ventral view, right angle bend at

midpoint toward dorsal surface; two large teeth in

each ramus; manubria nearly straight, dorsal and

ventral extensions are thin lamellae; pleural rods

which support mastax during pumping action

embedded in wall of mastax on each sidc, under

upper ends of manubria.

Total length 210-320 ym; toes 25-37 pin; trophi

34-42 um.

Ecology: Cosmopolitan in detritus-rich periphyton

Carniverous on rotlters (particularly bdeloids) in

littoral. Rare, N.T., Qld, Tas, Vic, 22,0-30.9°C, pH

3.4-5,4, DO 4.15 mg 1-', 29,0-59.3 pS em-'.

Literature: Shiel & Kaste 1979: Koste & Shiel 1983,

1986.

Resticula nyssa Haring & Myers

FIG, 27:

Resticula nyssa Harring & Myers, 1924, p. 521, Fig. 38:

-4.

Type locality: Atlantic City, New Jersey.

Holotype: Not designated,

Descriptian, Vermiform with longitudinal and

transverse siriae, head protrixion anteriorly

corresponds to rostrum in other taxa; foot two

jointed, very short, broad; toes short, with bulbous

basal enlargement and claw-like tip; mastax

modified virgale; fulcrum long, spatulate distally;

rami triangular with large asymmetric alulae: basal

apophysis prominent; three or four small teeth at

apex of rami; unci with two teeth each, five

accessory teeth on Jeft uncus only; manubria long

and slender; pleural rods slender, fibrillate at ends

and fused to rami; pedal glands small pyriform;

retrocerebral sac but no subcerebral glands; no true

eyespot, but mass of red pigment granules occupy

portion of sac.

Total length 300-630 pms toes 14--21 am; trophi

50-60 pm; male 160 um_

Ecology; Petiphyton, Gurope, Asia, N. Ametics,

Acidophile. Single record: Lake Garcia, western Tas.

0.2. «.87. 17,0°C, pH 4.3, 98.3.8 om-!, 0.5 NEU,

dark, humic water.

Literature, Koste 1978; Koste er al 1983.

Genus Searidium Ehrenberg

Searidium Elwenberg 1830, p. 47.

Type: Trichoda longicauda Mijller =

Jongicaudum (Miiller).

Type locality: Copenhagen,

Body: cylindrical or fusiform, partially lorivaw

both dorsally and ventrally; neck behind dorsal

antennae also with stiffened cuticle; foot very long,

three-segmented, tot fetractable, with short

basal., long distal-segment; complex internal foor

musculature to move very long, straight (oes; corona

Scartaium

ROTIFERA FROM AUSTRALIAN INLAND WATERS 5

oblique, with perioral buccal field and narrow

dorsally-interrupted circutmapica) ciliation; Uorsal-

and lateral antennae small; eyespot and retrocerebral

organ absent; mastax rounded, trophi yirzate; unel

toothed plates, points offser; manubria widened

proximally, hooked distally; fulcrum long, strong,

distally spatulate; rami somewhat triangular with

alulae, long pointed ends of which curve ventrally;

each ramus with a long pointed tooth arising

ventrally; epipharynx large, complex, reddish

colouration resembles an eyespot, gastiic glands

large; separate stomach and intestine; male

significantly smaller, with much shorter foot and

toes, cerebral cye present; subitaneous egy hairy,

resting ege spinulate.

Of the two described species, only 8. /ongicalidunt

is known from Australia,

Scaridium longicaudum (Miller)

FIG, 28:1

Trishoda longleauda Mijjlet, 1786, p. 216, Fig. 31:8-10.

Scaridium fongicaudum: Ehrenberg 1830, p.47,

Type Jacality; Copenhagen,

Holotype: Not designated.

Descriptions As lor genus; toes almost as long as

foot; unci lamellar plates with acute offset teeth;

manubria proximally “fish-hook' shape, distally

almost right angled,

Total length 360-450 pmy foot 118-150 jum; toes

122-145 pm; trophi 48 jum (fulcrum 30) zm; rami 15

am; manubrium 22 pm; ramus 16 pm); male 150

um; subitaneous egg 5!) pm,

Evofogy: Cosmopolitan warm stenotherm, pH

tolerant, swims slowly between macrophytes itr

shallow waters. Pancontinental, common in wide

range of habitats, 8.0-22°C, pH 7.2-7.4, DO 9.2-9.6

mg [-?, 60-160) nS em-') 8 NTU,

Comment; Distinguished from Scaridium bostjani

Daems & Dumont (from Nepal) by larger size

(>3) pm vs <288 ym) and wide tongue-shaped

articulations of the manubria with the unei vs a

pointed elongation in the Nepalese species. Detailed

analysis of variants reported in the literature is

lacking (Koste 1978), however it is likely that these

represent mbore than ecolypic variations, ie, 2

complex of species is involved,

Genus Taphrocampa Guss¢e

Yapnrocamipa Gosse, [BS1, p, 199.

pe: Tuphracampa annulosa Gosse.

Tepe focaliry: Leamington, England.

Notomimatid rotifers, cylindrical or Fusiform body

wilh more or less distinct transverse plicae; corona

oblique on anterior surface of head, with lateral

ciliated auricle; mastax yirgate with asymmetric

trophi; fulcrum, long slender; rami approximately

hemispherical; manubria long, slender with

eudimentary basal plate; unci with 2-3 narrow teeth;

well developed hypopharynx muscle; rami with large

alulae, Two of the four species listed in Koste (1978)

are known from Australia.

Key do species of Taphrocaimpa recorded from

Australian inland waters

1, Body generally <200 ym; toes <IS pm. .,,, 2.

we ie Ree, 2 Oe ...T. ananlosa Gosse (Fig. 28:2)

Body >220 um; joes >25 wm...,.....7) selenuna

GSosse (Fig. 28:3)

Taphrocampa ankulesa Gosse

FIG, 28;2

faphrocarnpa anrtulasa Gosse, 1851, p. b99-

Type focalicy: Leamington, England.

Holotype: Not designated,

Description: Movement shigzish, creeping, gliding;

body stocky may be thin when extended; three head

“segments”; dorsum of same appearance as T

selenuray seen faterally more or less coarsely

serrated; ciliary. auricles. rarely extended; short,

indistinc(, smell tooth on inner margin of left ramus

may be present; no basal apophyses,

Total length 130-230 jan; tues 11-15 ym; trophi

25-26 pm,

Ecology: Cosmopolitan in detritus tich periphyton

(littoral), N.SMW., NT, Qld, Vie.

Literature; Koste 1978, 1981; Shicl & Koste 1979.

Taphrevampa selenura Gosse

FIG. 28:3

Taphrocampa selenura Gosse, 1887, p. 1, Fig. UL

Thpe locality: England, not specified. Lacustrine.

Holotype; Not designated.

Description: Body elongate, cylindrical, tapering to

foot; integument “leathery”. Dorsally with

transverse plications usually 10-12; ventral surface

less distinctly plicate; tail separated from abdomen

by transverse groove; toes long, tapering. decurved,

form semicircle (viewed dorsally); lel! side of trophi

more developed than right; piston muscle attached

to end of fulcrum; left ramus with blunt tooth on

inner edge, lamellar plate behind this, then two large

teeth (Fig. 22:3c), right ramus with eight

rudimentary teeth, left uncus with large ventral

tooth and smaller second tooth; night uncus one

large and two small; manubria have subsquare basal

plates with straight posterior branch and terminal

expansions.

Toll leneth 220-290 jorg toes 25-33 yam trophi

J& pm,

156 W. SOSTE & R. J. SHIEL

Fig. 28. 1, Scaridium fongicaudum (Muiller): (a) lateral; (b) trophi, ventral; (c) trophi, lateral; (d) trophi, dorsal; (e)

two views of uncus, 2, Tuphrocampa unnitlosa Gosse: (a) dorsal; (b) lateral; (c) traphi, ventral; (d) unei, 3, T.selentre

Gosse: (a) dorsal; (b) lateral; (c) trophi, ventral; (d) unci. | after Donner (1943); 2, 3 after Harring & Myers (1924),

Seale lines: adult 50 mm: trophi 10 pm,

ROTIFERA FROM AUSTRALIAN INLAND WATERS iS?

Fxology; Cosmopolitan in detritus rich periphyton

and eutrophic decomposition areas. Rare, N.5S.W.,

NT.

Literature: Shiel & Koste 1979, Koste 1981-

Genus. Rousseletia Harring

Roussefetia Harring, 1914, p, #93.

Type: Rousseletia corniciata Harring, 1914 (Fig.

2),

Type focality: Kenilworth, DC, USA,

Small stout illoricate body; slight constriction

between head and abdomen; tail large, collar-like,

projects over Jong foot; foot has dorsal seta, may

be broken off; two short conical toes on ventral side

of foot tip; corona terminal with circumapical

ciliation; two papillae project from unciliated apical

plate; buccal field semicircular, ciliated, with mouth

near ventral edge; mastax disproportionately large

{ca. \4 body length), of specialized virgate type;

fulerum spatulate distally; rami large, domed,

without marginal denticulation; manubria simple

curved rods with ventral spur, unci absent) rod

shaped epipharyny present; eyespot large, cervical;

retrocerebral sac large, filled with highly refractive

granules; large stomach extending to blind sacs on

either side of mastax; no intestine; gastric glands

small; foot glands club-shaped.

Rousseletia if not positively identified from

Australia, Sudzwki & Timms (1977) listed a rotifer

identified as Russelletia [sic] parroti Russell from

Myall Lake, N.S\W. No description or figures were

provided, If this is the rotifer described by Russell

(1947), it was réferred to the genus Lindia

(Lindiidae) (see Koste & Shiel 19906), We regard

the record as iceriae sedis, but include the generic

descripuion should the Myall Lake rotifer be

encountered again.

Acknowledgments

Collectors. acknowledgec in earlier parts also

contributed marerial included here. The Deutschen

Fig. 2%. Rousseletia cornlentuia Harring: (a) dorsal; (b)

ateral; (c) trophi, ventral; (d) trophi, lateral. After

Harring & Myers (1924). ‘Scale lines: adult $0 um; trophi

10 jam.

Forschungsgemeinschaft continues to support WK

with microscope and photographic facilities. The

support of the Murray-Darling Freshwater Research

Centre, particularly the Director, Dr David Mitchell,

in funditig.a six-week visit to the Centre by WK and

assistant C. Leutbecher is gratefully acknowledged,

The patience of Lor-Wai Tan in converting. a part-

pencilled/part-bilingual draft manuseript to final

form on a word-processor, and her continued field

assistance, has been invaluable. The systematic

expertise of Dr Alice Wells, similarly patient, made

our task somewhat easier for this paper and the

TeMaining papers of the series. For technical and

field assistance respectively, Garth Watson and

Fiona Dunww are thanked.

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Srenroos, K. E. (1898) Das Tierleben im Nurmijarvi-See.

Eine faunistischebiologische Studie. Acta Soc. faun,

Siar, Fenn. 17, 1-259,

SToKes, A. C. (1897) Some new forms of American

Rotifera. I. Ann, Mag: Nat. Hist. Ser. 6, 19, 628-633.

Note added in proof

The status of Lindia parrotti Russell (sce Koste

& Shiel 1990b: 141) = AKusselletia parrotti (Russell)

in Sudzuki & Timms (1977) was resolved after this

paper was completed. Mr Phil Parr of Levin, N,Z.,

sent us the photograph of Russell’s /,india parrotti

missing from our copies of the original description.

Mr Jordi De Manuel, University of Barcelona, sent

a copy of the description of Russelletia Sudzuki

(1959), erected to accommodate L£. parrotti

Mrs: La-orsri Sanoamuang, Department of

Zoology, University of Canterbury, Christchurch,

collected at the type locality (Lake Victoria) a

population resembling garroifi as figured by

Russell, and checked the type collection of the

Supzuki, M. & Timms, B, V, (1977) A new species of

Brachionas (Rotitera) from the Myall Lakes, New South

Wales. Proc, Linn. Soc, NSW. WL, 162-166.

TESSIN, G, (1890) Rotatorien der Umgegend von Rostock,

nee renee Natur, Mecklenburg, Gustrow 43,

VEsnowsky, F. (1883) Uber Drilophaga bucephalus a. By

Tl. Sp. ein parasitisches Raderthier. Sitzungsber.

Bohmischen Ges, Wiss, Prague (for 1882), pp. 390-397.

Weer, E. F & Montet, G. (1918) Rotateurs, Cul,

Invert. Suisse 2, 1-339.

Wisniewsk!, J. (1934) Wrotki psamimotiowe, Avin, mius,

zool, polon. Warsaw, 1, 339-399,

———_ (1936) Uber die Nomenklatur der Cepkalodella

calellina ahnlichen Rotatorienarten, Zool, Polon Lwow

1, 171-179.

WULFERT, K. (1935) Beitrdge zur Kenninis der

Radertierfauna Deutschlands. 1. Arch, Hydrobial, 8,

$83-602,

(1937) Beitrdge zur Kenntnis der Radertierfauna

Deutschlands, Til. fbid, 31, 592-635.

(1939) Beitrage zur Kenntnis der Radertierfaung

Deutschlands. 1V. /bid. 35, 563-624,

(1940) Rotatorien einiger ostdeulscher Torfmoore,

Ibid. 36, 552-587.

(1960) Die Radertier saurer Gewasser der Diibener

Heide. {1. Die Rotatorien des Krebsscherentiimpels bei

Winkelmuhle. fhid. 56, 311-333.

—_____ (L961) Die Radertier saurer Gewasser der Diibener

Heide, (1. Die Rotatorien des Presseler und des

Winkelmthler Teiches. Zhid. 58, 311-333.

Canterbury Museum for Russell's material, We

(hank these people for their efforts.

The type material could not be located, but from

trophi analysis and comparative photographs of the

Lake Victoria rotifers, we consider Lindia parrotti

a junior synonym of Proalides tenfaculatus De

Beauchamp, 1907.

References

BracHamp, P, De (1907) Description de trois Rotiféres

nouveaux de la faune Francaise. Bull. Soc. gaol. Fr. 32,

148-157,

Sup2ZUKI, M. (1959) A new proalid rotifer, Liliferotrechka

urawensis n.g.n.sp., from ma marsh in Urawa, Japat.

Seat. Rept Tokyo Kyotku Daigaku Sec, BY, 9-33.

TRANSACTIONS OF THE

ROYAL SOCIETY

OF SOUTH AUSTRALIA

INCORPORATED

VOL. 115, PART 4

COASTAL ABORIGINAL SHELL MIDDENS AND THEIR

PALAEOENVIRONMENTAL SIGNIFICANCE, ROBE RANGE, SOUTH

AUSTRALIA

BY JOHN H, CANN*, PATRICK DE DEKKER** & COLIN V, MURRAY-WALLACET

Summary

Two superposed and stratigraphically distinct shell deposits, located at the seaward edge of Robe

Range within Little Dip Conservation Park, southeastern South Australia, are identified as

Aboriginal middens. The lower midden consists of Katelysia shells and megascopic charcoal,

within a terra rossa soil, developed on calcarenite of the Pleistocene Bridgewater Formation.

Radiocarbon dates of 82704 80yr cal B.P. and 7910+ 140yr cal B.P. were obtained for charcoal and

shell respectively. Amino acid racemisation values confirm an early Holocene age for the Kaielysia

shells and also suggest relatively recent exhumation of the midden materials. The upper midden

consists of Turbo shells, flint fragments and finely comminuted charcoal within a Holocene sand

dune. A radiocarbon date of 470+160yr cal B.P. was obtained for these shells. Amino acid

racemisation values confirm that the Turbo shells are only slightly older than modern. The midden

features, and their established timeframes, together conform to the constraints of the time-cultural

archaeostratigraphic Early Horizon and Late Horizon of Aboriginal sites in southeastern South

Australia. Accordingly, the middens site is here proposed as a type archaeological locality and type

archaeostratigraphic section for the Luebbers Early Horizon and Late Horizon time-cultural units.

Shells of the older midden were probably derived from an intertidal marine lagoon that occupied the

low lying corridor between the Robe and Woakwine ranges at the culmination of the Holocene

transgression. Alternatively, near the peak of Holocene sea level, a similarly protected sandy

environment may have hosted Katelysia seawards of Robe Range. Shells of the younger midden are

equivalent to those extant on the rocky shoreface of Robe Range. The established time difference

between the two episodes of human occupation of the site provides a valuable timeframe for

Holocene geomorphic changes within the study area.

KEY WORDS: amino acid racemisation, Australian Aborigine, archaeostratigraphy, Early Horizon,

Late Horizon, Holocene, Pleistocene, Bridgewater Formation, St Kilda Formation, Glanville

Formation, radiocarbon, Mollusca, midden, Robe, Woakwine, South Australia

Vransactions of the Koyal Sacvery af 8, Aust. (1991), 11801. 161 14,

COASTAL ABORIGINAL SHELL MIDDENS AND THEIR PALAEOENVIRONMENTAL

SIGNIFICANCE, ROBE RANGE, SOUTH AUSTRALIA

by JOHN H. CANN™, PATRICK DE DECKKER** & COLIN V. MURRAY-WAT 1 ACE

Suoumary

Casn, J. H., De Drckker, P. & MURRAY IW ar ace, CV (1991) Coastal Aborwinal shell gmddens and

their palaewenvironmental significance, Robe Range, South Australia. Trans, R. Sot. S Aust, 1N5(4), 161-175,

2% November, 1991,

Two siiperposed and stratigraphically distinicr shell depasirs, located at the seaward edet of Rohe Runge

within Lie Dip Conservation Park, southeastern South Australia, are identified as Aboriginal middena

the lower midden consisty of Aavelysia shells and megascapic charcoal, within a terra rossa soil, developed

on calearenite of the Pleistocene Bridgewater Formation. Radiocarbon dates of 4270) 80yr cal BLP. and

7910 +140yr cal BP, were obtained for charcoal and shel) respectively. Amino acid racemisation values confirm

anearly Holocene age for the Ayrelpsia shells and also suggese relatively recent exhumation of the midden

materials, The tipper midden consists of Turhe shells, Mint fraymenta and finely comminuted charcoal

within a Holovene sand dune, A radiocarbon date of 470: 160yr cal BLP? Was obtained for thest shells

Aming acid tacemisation values confirm that the Turbo shells are only slightly older Uvan modern, The

hiddlen) features, and their established timeframes, together conform {o the constraints of the time-Cultunal

Arehavostradgraphic Early Horizon and Late Horizon of Aboriginal sites in southeastes) Souch Australia,

Accordingty, the middens site is here proposed as a type archaeolagical locality and lLype archacostrativraphic

seelion for the Luebbers Early Horizon and Late Horizon time-cultural units. Shells of the older miditen

were probably derived from un intertidal marine lagoon that vecupied the low lying corvidor herween the

Robe and Woakwine ranges at the culmination of the Holocene transgression. Alternatively, near [he peak

of Holocene sea level, a similarly protected sandy environment may have hosted Karelvsia seawards af Robe

Range. Shells of the younger midden dre equivalent to those extanton the rocky shoreface af Robe Range.

(he established rime difference berween the twer episodes of human occupation of rhe site provides u valuable

limefrune for Holdcene geomorpltic changes within the study area,

Key Wort: amino acid racemisation, Australian Aborigine, archacostrilgniphy, Early Hotizon, Care

Horizon, Holocenég, Pleistocene, Bridgewater Formation, St Kilda Formation, Glanville Farmauon,

radigcarbon, Mollusca, midden, Robe. Woak wine, South Australia

Introduction

This paper is prumanily concerned with sediments

and landforms that resulied frum the Holocene

transgression in southeastern South Australia, and

wilh the ampact of thab transgression on

populations of coaslal Aboriginal people. The

investigation is centred on areas close to the town

of Robe and includes the Robe and Woakwine

Tanecs and the low lying inter-dune corndor (Fig.

1), This area is part of the Robe-Naracoorte coastal

plain, which provides a unique tecord of

Quatemary sea level changes. Successive high stands

of sea level resulted in the consifuction of

beach/diine barrier complexes. on a steadily

uplifting coastal plain, The result is aseries of low

allilude ranges, approximately parallel to each

other, and lo the present coastline, and generally

increasing in age landwards.

* School of Pure and Applied Sciences, University of

South Australia (Salisbury Campus). Smith Road.

Salisbury East. S Aust. 3108.

“* Depatrment of Geolovy, Australian National

Universiy, GPO Bos 4, Canberra, ACT, 2601,

| Department of Geology, University of Newcustle,

Rankin Drive, Neweastle, N.SAV. 2308.

The Holocene sea transgressed the continental

shelf and reached present sea level about 7Xyr BP,

Uncunsolidated, mosily bioclastic carbonate sands

were driver sShorewards by the persistent high energy

waves of the southern ocean, and redistributed by

the wind to form a blanket of transgressive dunes.

Sea water flooded the Jow lying corridor between

the late Pleistocene Robe and Woakwine ranges,

thus forming a sheltered coastal lagoon. Subsequent

coastal sedimentation, upwards shoaling of lazoon

sedinjents and continued uplilt of the coasial plain

has transformed the lagoon to a series of shallow

lakes.

Late Pleistocene populations of Aboriginal

people presumably lived on tracts of coastal land

that are now inundated by the Holocene sea, Such

populations, adapted to a gatherer economy in a

coastal regime, would have moved landwards with

the Holocene transpression. Thus, the oldest sites

of Aboriginal occupation along the coast are

‘contemporary with the peak of Holocene sea level

and contain the remains of mollusean fauna

harvested from the inter-range lagoon, Younger sites

vontain shells of molluses which favour a tacky

open ocean shoreface.

The palavoenvironmental significance of these

162 J. H. CANN, P, DeDECKKER & C, V. MURRAY-WALLACE

A > vy

HIFTOR DIP (CONSERVATION PARK

WLITTLE DIF

\ eo" >

s8 ¥

R ‘ oN YF Ax LOCATION OF ABORIGINAL MIODENS

Spe By EXPOSED SECTION OF HOLOCENE SEDIMENTS

GERBAN POND Ts re ROAD *-\ TRACK =“. PARK BOUNDARY

+ SPOT HEIGHT CONTOUR INTERVAL (10 METRES}

37" 20'

sent RIVOLI BAY

BRADLEY

HILL

AT? trig POINT (METRES)

EZ AITTLE DIP CONSERVATION PARK

CGASTAL ABORIGINAL S1iZLL MIBDDENS 64

middens is evaluated ere within a tlinefrane of

radiocarbon and amino acd racemisation (AAR)

Wales obtained from shells and charcual,

Previous investigations

COASTAL. ABORIGINAL OCCUPATION IN

SOUTHEASTERN SOUTH AUSTRALIA

A mounting body of archaeological evidence

suiwpests that aboriginal populations were

established in southern Australia between 40000

and 30,000yr BP. (e.g. Allen [989; Cosgrove 1989).

From approximately 45,000 and 30,000yr BLP. in

southern Australia sea levels fluctuated between -30)

to -22m (Cann e/ al. 1988, in press}, During the last

glaciation sea levels were some 130.10 150m below

present mean sea level (Chappell 1983; Chappell &

Shackleton 1986), while the ensuing transgression

reached present sea level 7000 to 6000yr BLP. Ic has

been calculated thar Holocene sea levels rose as

rapidly as 1.5 to 2.4 cea/ye (Woodroffe ef al. L988;

Belperio i) preys). This rise in sea level and the

associated environmental changes are likely to have

significantly affected chose Aboriginal populations,

with hunter/gatherer economies, reliant on coastal

resources. It has. been argued by some thar this ios!

recen! and catreme rise in sea level would have

forced all popnlations on the now submerged

continental shelf ta retreat inland (Ross 1985), IL

is. clear that the earliest dates that can be expected

for emergent sites of coastal aboriginal occupation

in southeastern South Australia will approximate

to the peak of the Holocene Leansgtession,

The most comprehensive work detailing carly

Aboriginal sites in the southeast is that of Luebbers

(1978)'. The significance of this largely

unpublished stucy can be gauged by the extent to

Which it has been cited by other researchers (eg.

Preily ef a/, 1983; Ross (985; Head 1986; Godfrey

1988, 1989: Eeloff era, 1989; Bourman & Murray-

Wallace 1990), Luebbers (1978)- established a

chronology for Aboriginal occupation ijn the

southeast, entilying two discrete episodes of

BHecupalion Which he termed Early Horizon and

Late Horizon. The terim ‘llarizon’, in this context,

is used in a lime-cultural sense rather than in

reference lo (he physical materinis of ihe sites.

Aboriginal middens of the Karly Horizon oocut

in lerra rossa soils developed on exposed surlaces

of late Pleistocene dunes, such as the Robe Range.

' Luebbers, R.A, (1978) Meals and menus: a study in

prehistoric coastal settlements in South Australia. PhD

thesis, ANU. Canberra, unpublished.

Luebbers (1978)! describtd material from two such

middens, one from Cape Marlin and another from

Bevilaqua Cliffs, about Skin southeast of Cape

Buffon (Fig. tj. The Cape Martin site contained

shells of Karelysie and Mytilus, charcoal and fllint

tools. Charcoal yielded a radiocarbon date which

Liiebbers considered questionable, The other site

cuntained ‘a small number of nundeseript tools’

together with shells of Mlehidones and charcoal,

Dares of 6250160 and 63504)0yr BP. were

reported for charcoal and shell respectively.

Tor these sites and another inland, Lyebbers

(1978)! remained unsatisfied with the stratigraphic

control and believed it was possible that younger

overlying material, perhaps from a more recent

occupation, had been incorporated into the lower

terra Tossa soil, Thus, no specilic site was. designated

as an archacologleal type locality for tis Early

Horizor.

Late Horizon Sites occur in unconsolidated sand

and in places, such as at Bevilaqua Cliffs, may

stratig¢aphically overlie an Early Horizon site,

Luebbers (1978)! subdivided his Late Horizon inro

an Early Phase aad w Late Phase, Middens of the

Farly Phase range in age from S800 to 1300vr BLT,

contain small numbers of tools ani monospecific

deposits of Plehidanay or Brachidontes? Middens

of the Late Phase are younger than 1300yr BLP,

contain numerous Aint implements and shells of

Turbo (= Svbainella) and other gastropods extant

on southern Australian rocky Forestrores,

Recent work by Egloff er wf (1989) in

southeastern South Australta has revealed ahundant

fiurbe shells in middens, dated by radiocarbon ol

charcoal, as old as 2560-120 and 3060+230yr B.P,

These dates call into question the Luchbers (197)!

subdivision of the Late Horizon time-cullural unit

on the basis of the types of shells preserved in

voustal middens.

GBOLOGIC AND GEOMORPHIC

FRAMEWORK

The landscape of southeastern South Australia

is characterised by a series of low altitude ranges,

sub-parallel to. each other and (o the present

coastline. Between Naracoorte and Robe thirteen

géeomorphically distingt ranges can be identified on

the otherwise gently seawards sloping coastal plain

(Sprigg 1952; Schwebel 1983). The region has

undergone steady regional uplift of about

1.07mm/yr throughout the late Pleistocene

(Schwebel 1983; Belperio m press). In general terms,

the ranges are geomorphic features associated with

palaeoshorelines and they wncrease in age away from

Fig, |. Map ol study area and surrounding region indicating locations of sites ang@ features referred up in deni

Robe towards Naracoorte. The geological origin of

these features has becn attributed to Quaternary sca

level changes (e.g. Tindale 1933; Sprigg 1952; Cook

é al 1977; Schwebel 19785, 1983; Belperio in

press), The ranges were termed ‘stranded coastal

dunes" by Sprigg (1952), though he recognised that

at least some ranges. had heen constructed during

several episodes of marine transgression and that

they incorporated sediments of beach, dune and

lagoonal palacoenvironments,

SedimentsS composing the ranges are

predominantly aeolian bioclastic calcarenites with

some seaward horizons of shelly limestones in which

the Fossil molluses cun be assoviated with rocky

foreshore sedimentation. This complex of

Pleistocene sediments has been termed the

Bndgewater Formation (Boutakoff 1963). Between

the ranges closest lo the present coastline, sediments

are lagoonal and lacustrine bioclastic to muddy

limestones. Fossil] molluscs in the lagoonal

sediments indicate clearly a variety of shallow

subtidal and intertidal palacocnvironments,

In the work reported here, the coastal Robe

Range, the adjacent Woakwine Range, and the

lagoonal sediments confined by these two features,

provide an important geomorphic and

palacoenvironmental framework (Fig. 1).

* Sefwebel, D. A. (1978) Quaternary stratigraphy of the

youth east of South Australia, Ph.D. thesis. Flinders

University, Adelaide, unpublished,

KATELYSIA IN HOLOCENE

ST KILDA FORMATION

ROBE RANGE

[Roce

DUNE SANDS

FACIES OF PLEISTOCENE

BRIDGEWATER FOAMATION

TEATIARY GAMBIER LI

Woukwine Range

The internal structure of Woakwine Range has

been exposed in the Drain Ll. and Woakwine

cuttings, excavations through the range to effect

drainage of wet Jands (o the northeast. These

cuttings reveal a complex Quaternary stratigraphy

which resulted from perhaps as many as five

separate stands of high sea level. The deposits af

each high sea level are separated by conglomerates,

soils, caleretes and strong carbonate cementation

(Sthwebel 1983), Basal transgressive sediments in

the Woakwiie cutting consist of shelly and pebbly

horizons that include flirt cobbles, up to 10crn

diameter, derived from the underlying Tertiary

Gambier Limestone: These are overlain in turn by

seaward dipping subtidal sands, beach sediments

and Jandward dipping sands of the transeressive

dune facies (Belperio in press),

in the Drain L cutting Sprigg (1952) observed (hut

several exposed planes of marine erosion were

immediately overlain by sediments containing ‘a

typicul recf fauna’ fossil assemblage (Fig. 2). Blocks

of excavated shelly limestone, corresponding to this

facies, can be observed today on the roadside

overlooking this cutting. Significant faunal

components are fossils of abalone and Turbo (Fig.

3), species which are characteristic of modern rock

foreshores. Sprigg (1952) deduced that the dune

sediments had been substantially lithified during

subaerial exposure. hus they had remained as a

coherent geomorphic feature and provided a rocky

AEOLIAN FACIES OF

BRIGGEWATER FORMATION

ANADARA IN PLEISTOCENE

GLANVILLE FORMATION

—_,

ESTONE'

Big. 2. Diagrammatic section through Woukwine Range (afler Sprigg 1952) illustrating stratigraphw distribution at

distinctive fossil molluscs. Both Anadara and Aatelpsta are characteristic of lagoon facies sedimencs,

COASTAL ABORIGINAL SITELL MIDDENS 165

Fig, 3. Fossil gastropods within rocky shoreface tacies of

the Bridgewater Formation, Drain L cutting, Woakwine

Runge: A. urbe shell B. Abalone shell.

substrate follawing marine inundation. On this

basis he was able to recognise three episodes of

Marine regression followed by transgression in the

construction of the Waakwine Range.

tossils of the bivalve Anadera (rapezia occur in

some of the lagoonal sediments onlapping the

landward side of Woakwine Range (Pig, 2). This

species is characteristic of the late Pleistocene

Glanville Formation (Cann 1978; Murray-Wallace

etal, 1988a) and similiar marine sediments of earlier

Pleistocene age (Murtay-Wallace ef al. 1988b).

Rabe Range

Robe Range is the youngest of the emergent

stranded coastal dunes, Within the study area it

outcrops as an erosional rocky shoreline with

numerous irregular siacks and islands in which

acolian bedding structures are clearly evident (Fig,

4). Basal sediments of this range rest

unconformably on ‘lértiary Gambier Limestone

10-15m below present sea level and there are mo

horizons of fossiliferous rocky foreshore lacies such

as were recognised in the Woakwine complex

(Sprige 1952).

Schwebel (1983) identified three stages of

development for Robe Range, The late Pleistocene

construchonal stages were equated with oxygen

isotope sub-slages 5c and Sa, Equivalent marine

sediments were deposited in Spencer Gulf at sea

level maxinva of -8m and -l4m respectively (Hails

ef al, 1984), The most recent deposition has resulted

from the Holocene transgression, during which

there was extensive build up of dune sands over the

older components of the Robe Range. These

Holocene sands remain esseorially untlichified.

/nter-rangé sedimentation

Despite the steady regional uplift, maximum sea

level of the Holocene transgression was sufficient

co flood the low lying corridor between the Robe

und Woakwine ranges. Numerous road cuttings and

other shallow excavations reveal a wealth of

Holocene fossil molluses characteristic of relatively

protected (lagoonal) shallow subtidal and intertidal

marine environments (Fig, 5). The Moor of Lake

Robe (Fig, 1), for example, is littered with the shells

of oysters, scallops and cockles, particularly the

intertidal Katelysia, These richly tossiliferous

Holocene sediments belong to the St Kilda

Formation, In the sense oF Cann & Gostin (1985),

Present Investigation

The work reported here centres on the Little Dip

Conservation Park southeast of Robe (Fig. 1). The

area includes coastal exposures of the late

Pleistocene Robe Range (Fig. 4) and Holocene shell

beds deposited in the low lying areas between the

Robe and Woakwine ranges. These features are to

a large extent covered by trarisgressive Holocene

sand dunes, some of which are fixed by modern

vegetation, while others are little vegetated and

subject to present day erosion. The gastropod Turbo

is extant on the rocky foreshore and its shells are

easily collected at the Waters edge (Fig. 6).

The Aims of the investigation are three fold:

|. to evaluate critically the cultural-chrono-

stratigraphi¢ convept adopted by Luebbers

(1978)' within a framework of

chronologic, palaeontologic and

geomorphic investigation;

2. to propose a type section for the Early

Horizon and Lave Horizon cutural sites;

3. to document the palaenenvironmental

significance of the type area,

166 J. H: CANN, P. DEDECKKER & C, ¥, MURRAY-WALLACE

Fig. 4. Erosional rocky shoreface of Robe Range at Little Dip (Location A, Fig. 1).

Observations and Methods

ABORIGINAL MIDDENS

At the study site (Fig. 1, location A) a poorly

vegetated coastal dune immediately overlooks the

foreshore. The seaward side of this dune has been

subjected to wind deflation and a lag deposit of

abundant shells and opercula of Tirbo, together

with numerous fragments of flint, litters and surface

(Figs 7A, B). Some opercula are chipped or

fractured and are more numerous in some areas

than others, as if selectively sorted. Above the lag

deposit there are numerous conspicuous Tirbo

shells in a greyish, poorly consolidated horizon of

the dune (Figs 7A, C). The shell and flint appear

to have been derived from this layer which, on field

evidence, is interpreted as an Aboriginal midden

belonging to the Late Phase of the Late Horizon

as defined by Luebbers (1978)'. Shell from this

midden was taken for radiocarbon and amino acid

racemisation dating.

On the landward side of (he dune the Holocene

sand sharply overlies a well consolidated red-brown

terra tossa soil developed on the Bridgewater

Formation of the Robe Range (Fig. 8A). Embedded

within this palaeosol are numerous shells of the

bivalve Kate/ysia and fragments of charcoal (Figs

8B, C; 9A, B). Although no flint fragments were

observed, a human origin is also proposed for this

material. This assertion is based on the following

observations.

a. The shells are disarticulated, lack any preferred

orientation and many are severely broken. It is

difficult to imagine a natural sedimentary

environment that would cause such fracturing of

shells, but had they been naturally transported

under conditions of high wave or current energy,

the shells would have been deposited predominantly

convex upwards and tightly imbricated and also

show. signs of attrition. Such a fabric can be seen

in some of the Holocene shell beds between Robe

and Woakwine ranges (Fig. 5B). Where shells have

not been actively transported their valves usually

remain. more or less paired (Fig. 5C).

b, Characteristic ‘drill’ holes, inflicted by predatory

gastropods, were not observed in any of the exposed

Fig. 5. Fossil molluses of Holocene St Kilda Formation between Robe and Woakwine Ranges: A. Oyster shells on

the floor of Lake Robe. B. Predominantly shells of Kufe/ysia, disarticulated and convex upwards, signifying moderate

energy trausportation; small roadside excayation, Princes Highway. C, Predominantly shells of Katelysia, mostly

articulated, signifying litle or no transportation. An intertidal environment of deposition is inferred. (Location

B, Fig. 1). D. Katelysia showing the characteristic “drill” hole inflicted by predatory gastropods. (Location B, Fig. 1).

COASTAL ABORIGINAL SHELL MIDDENS

168 J. H. CANN, P. DeDECKKER & C. V. MURRAY-WALLACE

Fig. 6. Shells of Turbo, extant on the Rocky foreshore of

Robe Range at Little Dip. (Location A, Fig. 1).

Katelysia valves, yet within the nearby Holocene

shelly sediments such valves with ‘drill’ holes are

numerous (Fig, 5D). Clearly some form of selective

process has operated to eliminate bivalves affected

by this type of predation. Selection also seems to

have favoured larger individuals.

c. Sand enclosed by paired Katelysia valves from

the Holocene lagoonal sediments was

microscopically examined and found to contain

species of Foraminifera also known from intertidal

sandflats of Gulf St Vincent (Cann & Gostin 1985).

Species included Elphidium crispum, E.

macelliforme and Miliolinella labiosa. Gill et al.

Fig. 7. A. Lag deposit of 7urbo shells and flint fragments apparently derived from stratum indicated by arrow. This

deposit is interpreted as a Late Horizon Aboriginal midden, (Location A, Fig. 1). B. Flint fragments from the lag

deposit. C. Turbo shells within the stratum indicated by the arrow in A.

COASIAL ABORIGINAL SHELL MIDDENS Ino

(1991), used the abundance of Foraminifera ina shell

deposit at Warrnambool to show thal it was a

natural estuarine deposit aod nat of human origin.

Mictoseopic examination of the terra rogsa matrix

enclosing the Kate/ysia shells revealed no

Foraminifera within the deposit under present

discussion,

d. Stratigraphic elevation of the Na/elysia deposit

at the crest of Robe Range makes natural

sedimentation implausible 1f these shells are to be

correlated with those undisputedly deposited in the

Holocene lagoon between Robe and Woakwine

ranges. Iris alsa possible thal Kotelysia could fave

occupied relatively sheltered intertidal seaward

environments that probably existed immediately

prior ta the culmination of the Holocene

transgression, a suggestion lavoured by Luebbers

(pers. comm, 1991).

(Although there is abundant charcoal, both

within and surrounding the shell deposit, and

unambiguously embedded within the terra rossa

soil, there are no clear signs of localisation that

tight be easily interpreted as camp fires. Thus, for

this site, the presence of charcoal does not

necessarily, in itself, constitute evidence of human

occupation).

Thus the deposit is interpreted as an Aboriginal

midden belonging to the Early Horizon as defined

by Luehbers (1978)!, Shell and charcoal trom this

midden Were taken for radiocarbon assessment.

Additional shell was taken for amino acid

racemisalion dating. Paired Aate/ysia valves. from

the nearest accessible deposi! of Holocene laznonal

sediments (Fig. 1, location B), were taken for

comparajive AAR dating,

DATING

Charcoal and Karelysia shell were carefully

removed from thie teria fossa matrix of the Late

Horizon midden and packed in clean plastic bags,

Similarly, Yurha shells. were taken from the Early

Horizon midden (Fig. 1, location A), These

matenals were forwarded ta the radiocarbon

laboratories of (he Australian National University

and the University of Sydney for radiocarbon

dating, Conventional radiocarbon dating followed

the methods of Gupta & Polach (1985).

METHODS: RADIOCARBON

DATING METHODS: AMINO

RACEMISATION ANALYSES

The following materials were collected for AAR

analysis for the purpose of age determination:

a disarticulated shells of Karelysia rhyipkere and

X, wealarina from the Barly Horizon midden (Fig.

1, location A),

ACID

b. articulated shells of A. sca/arina trom Holocene

Jagoon sediments (= St Kilda Formation) within

the Robe-Woaakwine corridor (Fig, 1, location B);

c. articulated shells of K. sealarina from fate

Pleistocene lagoon sediments (= Glanville

Formation) on the landward side of the Woakwine

Range, exposed in a small quarry adjacent ta

Princes Highway;

d- shells of Turbo sp. from the Late Horizen

midden, and from the immediately adjacent

modern shoreface sediments (Fig, lL, location A).

AAR analyses were undertaken on all the

collected shell materials. Data obtained from the

Kavelysia shells of the Early Horizon midden were

compared and contrasted with the extent of

Tac¢emisation in specimens obtalnaed tron the

Holocetie and late Pleistocene lagaon sediments.

As the nature of racernisation kinetics in Jinbo spp.

shells is not well documented, the significance of

AAR analysis of shells from the Late Horizon

midden was assessed with reference to data derived

from radiocarbon dating of an adjacent midden

shell and AAR analysis of modern specimens.

Amino acid analyses were for the ‘total acid

hydrolysate’. complex peptide mixture of varying

molecular weights, and follawed the methods

outlined by Kimber & Griffien (1987). Analyses of

the N-pentailuoropropionyl D, L-amino acid

2-propylesters were undertaken using a 25m Sused

silica Chirasil-L-Val capillary column and Hewlett

Packard model 58904 gas chromatograph with a

flame ionisation detector and hellurn earner gas,

The integrity of the analytical procedures

undertaken using the AAR technique was evaluated

by analysing international interlaboratory

comparison samples of Wehmuiller (J984a). Results

Werte within two standard deviations of the arand

mean of the international comparison.

The hinge region of each Kolelvsia shell was

analysed as the highest concentration of residual

Protein oerurs within this region. The volumes was

analysed in the Tiirbo specimens. In excess of 205

by weight for each shell was analysed by AAR to

reduce yarlabjlity which may potentially arise when

analysing small fragments (Wehmiller 1984b).

Amino acid D/L ratios were compared with

calendric radiocarbon aves (cal BLP), converted

from ovarine reservoir corrected radigearbon years

(BP: Libby half-life) according to the methods

outlined by Gillespie & Polach (1979) and Stuiver

ef al, (4986) (Table 1).

Results

RADIOCARBON DATES

Charcoal from the Barly Horizon midden (lat.

code ANU-7448) yielded # tadiocarbon age of

8270+ 80yr cal BP, Awfelvsia shell, also from the

170 J. H. CANN, P. DeDECKKER & C. V. MURRAY-WALLACE

COASTAL ABORIGINAL SITELL MIDDENS hi

Fig. 9. Selected yalve from (he Early Horizon midden

showing (A) internal and (B) external features of

Katelysia.

older midden (lab, code SUA-2613), was determined

to have a marine reservoir corrected age of

7910 +140yr cal B.P, Jiurbo shell trom the overlying

Late Horizon midden (lab. code ANU-7447) had

a tadiocarbon age, again corrected for the marine

reservoir effect, of 470+ 160yr cal BP.

AMINO ACID RACEMISATION ANALYSES

The relative extent and degree of racemisation for

the different amino acides in Aarelysia spp. and

Turbo spp. is in aecord with other radiocarbon

dated Holocene fossils in southern Australia (for

example, Murray-Wallace er a@/. 1988c,; Murray-

Wallace & Bourman 1990) and the theoretically

predicted dilferential rates of amino acid

racemisation (Table 1),

The similarity in extent of racemisation of amino

aads ln Aavelysia scelarina obtained from lagoonal

sediments and molluses from the Early Horizon

midden points tO &@ common age given the

assumption that they have experienced equivalent

diagenelic temperature histories. The validity of this

assumption is strengthened by the close proximity

of these two sites. The location of the lagoonal

facies close to the feather edge of the Holocene

transgressive sediments, which have elsewhere been

dated at approximately 7000yr B.P,, also points to

a common age for ihese two sites.

A calibrated radiocarbon age of 79104 Id0yr cal

B.P. was obtained on Aavelvsia from the Early

Horizon midden (Table 1), These data are

significant, for they document Aboriginal coastal

occupation during an early transgressive phase af

(he post glacial Holocene marine tratisgression. ‘To

evaluate the amino acid data of the Early Horizon

midden, these results are compared with a sequence

of similar age at Smoky Bay, Eyre Peninsula

(Table 1), A z-score, according to Lhe methods of

Gupta & Polach (1985), indicates that these Iwo

calibrated radiocarbon dates are not significantly

different at the 5% level (z-score of 1,55), Although

comleTnporary mean annual temperatures (CMAT)

between these two sites differ by approximately

2.2°C, itis unhkely that the diagenctic temperature

differences will be detected for the Holocene record.

In contrast, such temperature diflerenves are

significant for last interglacial fossils (for example,

Murray-Wallace ef al, (991).

The similarity in extent of racemisation for the

Tadiocarbon calibrated Holocene Karelysia from

Smoky Bay indicates that the Karel/vsitt shells of the

Early Horizon midden were butied tor a significant

portion of their diagenetic temperature history, Had

the shells been subaerially exposed for much of their

diagenetic history, the extent of racemisation of

amino acids would be significantly higher (compare

Murray-Wallace e¢ al. J988c), The suggestion,

independently based an AAR data, (hat the Early

Horizon midden had heen buried, then exhumed,

is consistent with the geomorphic evidence of the

site, that is, recent dune deflation, This clearly

represents 4 novel application of AAR in the

recognition af exhumed sequences.

The lower extent of AAR in Tirrbo sp. from the

Late Horizon midden, than in Aare/isia spp, trom

the Early Horizon midden, is consistent with a

younger age, as independently determined by

radiocarbon dating. However; as this is the (irst

eee

Fig, 8. A. Early Horizon Aboriginal tmidden within (erra rossa palaeosol vo Bridgewater Formation, Robe Range.

(Location A, Fig. 1). a. Karstified rocky oulcrop of Bridgewater Formation. b. Xatelysia shells of (he Early Horizon

midden. c. Terra rosga palacosol. d, Holocene sand dune which includes the Late Horizon Turbo midden (Pig

6). Nore the sharp contact between this and the underlying terra réssa palacosal. B, Detail of Kulelysia shells wuhin

the midden, Note that shells are disarticulated, lack preferred orientation and pane shows signs of gastropod predation,

C, Detail of part of the middenin which charcoal, indicated by arrow, is embedded in the terra rossa palacasal.

172

J. H. CANN, P. DeDECKKER & C. V. MURRAY-WALLACE

TABLE 1. Extent of amino acid racemisation (‘total acid hydrolysate’) in late Quaternary Mollusca from the south

east of South Australia.

Location/ Depth of Species CMAT+

description burial (m) (°C)

Robe Beach surface Turbo spp. 14.7

Robe midden surface Turbo spp. 14,7

(Late Horizon) (exhumed)

Robe midden surface Katelysia spp. 14,7

(Early Horizon) (exhumed)

Robe/Holocene 1 Katelysia 14.7

lagoon facies scalarina

Woakwine Range/ >1 Katelysia 14.7

Back barrier scalarina

lagoon facies

Smokey Bay, Eyre 1.82 Katelysia 16.9

Peninsula/coastal 1.88— rhytiphora

sediments!

Age PB* Amino acid D/L ratio#

& [lab code] VAL LEU ASP PHE GLU

modern 4 0.02 n 0.02 — 0.03 0.04

*0.001 *0.004 *0.005 *0.01

(840 +80) 0.04 0.07 018 0.12, 0.09

470+160 * 0.002 *0.01 *0.002 *0.01

[ANU-7447]

(7480+70) 0.05 + 0.09 0.28 (0.22 | 0.11

79104140 *0.01 *0.03 *0.01 *0.02 ~0.01

[SUA-2613]

= 0.06 0.08 0.32 016 0.12

0.02 +002 *0.01 *0.04 +0.01

last interglacial | 0.20 0.35 0.54 co 0.31

125 000 0.02 +0.01 *0.06 * 0.03

(6940 +170) 0.08 0.16 023 — 0.10

7410+290 +002 *0.02 * 0.004

[CS-450]

C.M.AT. — Contemporary mean annual temperature (atmospheric).

Conventional radiocarbon age indicated in parentheses with associated error term (16). Marine reservoir corrected

sidereal ages without parentheses with 26 error term. See text for discussion on marine reservoir correction and

calibration of radiocarbon ages to sidereal years,

# amino acids: VAL — valine; LEU — leucine; ASP — aspartic acid; PHE — phenylalanine and GLU — glutamic

acid. Error terms indicate analytical precision and intershell amino acid D/L ratio variation (16).

! data of Murray-Wallace ef al. (1988c).

AAR analysis on Turbo spp. from southern

Australia, it is not possible to evaluate critically the

relation of the kinetics of racemisation in Turbo to

the moderate racemisation rates that are

characteristic of Katelysia. It is likely that

racemisation rates vary between bivalves and

gastropods, as noted by Miller & Brigham-Grette

(1989). However, the small difference in extent of

racemisation of amino acids between Turbo, of the

Late Horizon midden, and Kafelysia, of the Early

Horizon midden, may also point to a history of

subaerial exposure for the former.

The extent of amino acid racemisation in

Katelysia scalarina from the back barrier lagoon

facies of the Woakwine Range barrier complex is

consistent with other last interglacial Mollusca

(Murray-Wallace ef a/, 1988a) and is in accord with

a temperature-geographic latitude kinetic model for

last interglacial shell taxa in southern Australia

(Murray-Wallace et al. 1991).

These data assist in constraining the time

framework for the Aboriginal coastal occupation

of southern Australia.

Discussion and Conclusions

THE ARCHAEOLOGICAL SITE: A TYPE

LOCALITY

The sharp stratigraphic boundary between the

Katelysia bearing terra rossa palaeosol of the

Bridgewater Formation and the overlying Turbo

bearing Holocene dune sand is clearly shown in Fig.

7. the palaeosol is well consolidated and has not

been contaminated by younger overlying sediment,

shells or artifacts.

The two sets of midden materials, as described

in this paper, undoubtedly conform to those

specified by Luebbers (1978)! for his Early

Horizon and Late Horizon of aboriginal

occupation of southeastern South Australia.

Equally, the age determinations of both shells and

charcoal, particularly the close agreement of ''C

and AAR dates, meet the constraints of time

applied to this time-cultural classification.

Given that the site meets these tight stratigraphic,

archaeological and time constraints, and given that

it is located within the boundaries of a National

Conservation Park, it is here confidently proposed

COASTAL ABORIGINAL SHELL MIDDENS Wa

asa type locality and type section for the time-

cultural Barly Horizon and Late Honzoy of

Luebaers (1978)!

OTHER ARCHAEOLOGICAL SITES

POSSIBLE REFERENCE LOCALITIES

It is likely thar Future investigations will reveal

Other sites that will equally illustrate, or futher

clarify, the Luebbers (1978) chronulogy, If

appropriate, such sites should be desienated as

reference lovalities and referenve sections.

In this context, middeus within Discovery Bay

Coastal Park near Cape Bridgewater in

southwestern Victoria seem worthy of further study,

Godfrey (1989) differentiated middens in this area

into two episodes of occupation. though did nor

use the Luebbers (L978)! terminohogy-

The older middens, 8490-3860y7 BLP, ane in terra

rossa souls al the Bridgewater Formation and

contain mussel shells of a species no longer extant

along the present shore. The younger middens are

in unconsolidated sand and contain shells of

species, such as Turbo, which inhabit the present

shoreline, together with numerous (lint fragments,

Dates of 1050-360yr BP, were reported by Ciodfrey

(1989) for these younger sites.

AND

Environmental history of the study area

The following is.an account of the interaction of

physical and biological processes, from c.125.4)00yr

BP. to lare Holocene, leading co the evolution and

early human exploitation of rhe study environment,

At L35,000yr B.P, oxygen isotope sub-stage Se,

southern Australian sea level was Slightly higher

than aj present. Various estimates place vlobal sea

levels at +4 to =4m, bul distribution of late

Pleistocene Mambray Formation (-Glanville

Formation) in northern Spencer Gull suggests that

the Se sea level was only «Im (Hails ef af 1984).

At this times, the seaward side of Woakwine Range

formed a rocky coastline and the shoreface was

inhabited by molluscs favouring such a subsrrare

ina high cnergy wave regime. Abalone, limpets and

Trio were significunt faunal elements. Seawater

flooded areas landwards of Woakwine Range

forming a coastal lagoon, the sheltered walers of

which Were extensively colonised by molluscs such

as Kajelysia and Anadara.

Following marine regression during uxygen

isorope sub-stage $d, at 105,000vr BLP, sub-stage

Sc, marine transgression brought palaco sea level

to -8n.{Halls ef af. 1984; Belperio in press), Robe

Runge stage [El sediments accumulated as

unvonsolidated beach and dune sands (Schwebel

1983).

Duting isotope sub-stage 5b, the sea receded

allowing subaerial diagenesis and at Teast partial

lithifcation Of the carbunate rich sands.of (he stave

Jl) seditients. Trotective calerctes developed on

exposed surfaces

Ar BO,000yc BEF, Isotope sub-sii@e Sa, marine

Ininsgression brought sea level to ~14in (Hails ef

al 1984; Belperio in press}. Robe Range siage II

sediments were deposited al this time. Sea level was

not sufficiently Ligh to erede the earlier formed

Stage IL sediments, which were mantled by the dunce

Facies of the stage UL transgression.

Following this peak of sea level, the voean again

tegressed acruss (he continental shelf and for the

remainder af Pleistacene time the shoreline

remained seawards uf Robc Range. ‘The carbonate

sands thus underwent futher extensive diagenesis

and consolidation,

Beiween 45,000 and 20,WXWr BAP., oxygen isotope

slage 3, here were Muctuations of sea level between

-30m and —22m (Cann ef al 1988, in press}, but

these were insufficien! (o influence Robe Range,

Also, by 30,000vr AUP, the base of the range had

undergone about Sm of tectonic uplift, futher

compounding the impact of the regression. Early

humans may have firsc appeared in southeastern

Australia at this time.

From 30,000 to (8,000yr Bue, duting Oxygen

isotope stage 2, the last glacial meeression lowered

sea level fo — 130m (Chappell & Shackleton 1986).

Aboriginal populations occupied the emergent

continental shelf and in coastal ateas probably

exploiled a variety of sew food resources.

Ac 18,O(lyr BP, sea level begat to rise, sumetimes

as rapidly as 2.4dem/yr, totally submerging the

continental shelf hy 7,000yr BLP, (Belperic tn press).

Unconsolidated sands were driven shorewards by

The rising sews, ntantling seaward outeraps of Rabe

Range stage If and, where exposed, stage IT), Sea

water flooded the low lying corridur between Robe

and Woakwine ranges, providing sheltcred shallow

subtidal and intertidal environments in which

mollusc populations thrived, Aboriginal people

accupied Robe Range, open ocean fo ane side and

sheltered lagoon to the other At the study site they

exploited the intertidal cockle Kuefvsia, Elsewhere,

for exarople at Revilaqua Cliffs to the seurheast,

ocean beach cockles, Plehiduriax, were the prime

food source.

Bioclastic sedimentation within rhe shallow Robe-

Woakwine marine lazoun was tapid, Sedimentary

secions reveal upward shaoline sequences of

subrigal oysters overlain by intertidal Nufedysia and

Amapella. Shoaling was further facilitated hy

tectonic uplitt of about 0.4m from the ime of

stabilisation of Hologene sea level 1o present,

Meanwhile, on the seaward side of Robe Range,

sands continued to accymulare, In the absenve of

apy preferred direction of longshore transport

ha jd. . CANN, P. DeDECKKER & C. Vo MURRAYAWALLACE

(Sprigg 1952. Bourman & Mutray-Wallace 1990), (te

strong persistent ocean swell (Gostin et al. ISR)

moved sands onshore from where they were

distributed to form a blankel of dunes, Sonie sands

were alsa redistributed, both up and down the coast,

lo the protected areas of Gmichen Bay, ta the

northivest, and Rival Bay, (a the southeast.

Sedimentation |i these areas effectively.isolated the

Rebe-Woakwine Jagoon from JSuther marine

influence

At the study Site, continued onshore und

alongshore migration of the Holocene sands once

again exposed (he older lithificd sediments of Robe

Range. Their long subienal emergence thiougli the

late Pleistocene bad resulted in a high degree of

carbonate cementation and they oulcropped as a

rugged Irregular erosional coastline, A marine

mollusc launa, dominated by the large gastropod

Trrbo, became established along this rocky, bigh

wave encrpy cnvironment. Thus was repeated, an

the Holocene shureface of Robe Range, ecoloxical

events that are recorded by the Pleistocene horizons

ot typical reef fauns® (Sprigg 1952) in the

Woukwine Ranges,

For a second time, Aboriginal people occupied

the study site and exploited this newly established

(nod resource.

The two middens at the study site therefore reflect

profound changes in landscape, The elapsed rime

herween the two periods of human oceupation is

a viliable clue tp the tate of environmental change

Conclusions

|. Radioearbon and AAR dating together consirm

an early Holocene age for the Early Horizon

Rohe Kange Aboriginal midden. which is

contemporanceus with the culminstian of the

Holocene lransgression. Application of AAR as

a palacothermomeler indicates iat this early

human site has recently beet subaerially exposed,

This work has demonstrated the importance of

aul integrated approach ww archacostratigraphic

studies through the combined efforts of dating,

geomorphology, sediimentology and

paleontology. Such inlegrated studies can

reconcile otherwise seemingly conftieting

evidenee

3, The netion of archaeastratigraphic type and

reference sections provides a useful approach in

the study of Australian prehisiory and his

Potential for wider application.

Acknowledgments

Field work and preparation of this paper was

financially supported by the Lniversity af South

Australia Professional Leave Programme. The

authors thank Mr David Hritan for drafting Pig.

1, Mrs Chris Moore for typing, and Mr Neale

Draper, Dt Antonio Belpena, Dr Brian McGown

and Dr Roger Luebbers for their critical reading of

the manuseripl, We are indebted tg Neale Draper

for aceess to the Luebbers thesis and other

documents held by the Aboriginal Heritage Branch

uf the Dept of Environment and Planning. Same

of the work teported here eonsitutes part of a

brawler Invesugulion of fate Pleistocene and

Holocene s2a levels by John Cann within a Ph.D,

proeramime atthe University af Adclaite under the

supervision of Dr Bian McGowran. We also wish

ta thank Ms Gillian Taylor of the NW.G, Macintosh

Centre fur Quaternary Dating, University of

Sydney, and Mr John Head of the Radiocarbon

Laboratory of the Australian National University

for ussistance with radioearbon dating; and Dr

Kelvin Picker of the Dept of Organic Chemistry.

University of Sydney, for ascistance witltamino avid

Tdvemisation dating. br Mike Barbetti contributed

valuable comments on radiocarbon calibration,

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{Eds) “Natural History of the South East”, (R. Soc. S.

Aust., Adelaide).

Spricc, R. C. (1952) The geology of the south est

province South Australia, with special reference to

Quaternary coast-lne migrations and modern beach

developments. Geol, Suri. S. Aust, Bull. 29.

StTurver, M., PEakson, G. W. & BRAZIUNAS, T, (1986)

Radiocarbon age calibration of marine samples back

to 9000 cal yr BP. Radiocarbon 28, 980-1021,

TINDALE, N. B. (1933) Tantanoola Caves, south cast of

South Australia: geological and physiographical notes.

Trans. R. Soc. 8, Aust. 57, 130: 142,

WEnMILLER, J. 1. (1984a) Interlaboratory comparison of

aming avid enantiomeric ratios in fossil Pleistocene

mollusks. Quan Res. 22, 109-120,

____ (1984b) Relative and absolute dating of

Quaternary mollusks with amino acid racemization:

Evaluation, applications and questions. Pp, 171-193. in

W, C. Mahaney (Gd) “Quaternary Dating Methods.”

(Elsevier, Amsterdam)

Wooprorre, C, D, CHAPPELL, J. & Tuo, B. G- (1988)

Shell middens in the context of estnarine development,

South Alligator River, Northern lerritory. Archeot,

Oceania 23, 1-16.

A NEW GENUS AND SPECIES OF PHREATIC AMPHIPOD

(CRUSTACEA: AMPHIPODA) BELONGING IN THE "CHILTONIA"

GENERIC GROUP, FROM DALHOUSIE SPRINGS, SOUTH AUSTRALIA

BY W. ZEILDER*

Summary

A freshwater amphipod Phreatochiltonia anophthalma gen. et sp. noy. is described and illustrated.

It is endemic to a few small, cold, artesian springs amongst the Dalhousie Springs complex in the

north of South Australia. Unlike species of the closely related genus Austrochiltonia the new

species seems to be subterranean in habit, is completely white, lacks eyes, and coxa 4 is without a

clearly defined posterior excavation. "Chiltonias"generally are considered to be more closely allied

to the Hyalellidae than the Ceinidae, and thus transference of chiltonid genera (subfamily

Chiltoniinae) to Hyalellidae is recommended.

KEY WORDS: Phreatochiltonia anophthalma gen. et sp. noy., new species, amphipod, artesian

springs, Australia, taxonomy.

Vrunsuctiaons of the Royal Su¢iery of S. Awst. (199), (15{4), 177-187.

A NEW GENUS AND SPECIES OF PHREATIC AMPHIPOD

(CRUSTACEA: AMPHIPODA) BELONGING IN THE “CHILTONIA” GENERIC

GROUP, FROM DALHOUSIE SPRINGS, SOUTH AUSTRALIA

by W., ZEIDLER*

Summary

ZeIn.ER, W. (1991) A new genus and species of phreatic amphipad (Crustacea: Amphipoda) belonging

in the “chiltonia" generic group, fromm Dalhousie Springs, Sourh Australia, Trans. R. Soc. 8. Aust, 1W5(4),

177-187, 29 November, 1991.

A freshwater amphipod Phreatochiltonia anophrhalma gen. et sp. noy. is described and illustrated. It

is endemic to a few small, cold, artesian springs amongst the Dalhousic Springs complex in the north of

South Australia, Untike species of the closely related genus Austrochiltumia the new species seems Lo be

subterrancan in habit, is completely white, Jacks eves, and coxa 4 is without a clearly defined posterior

excavation.

“Chilionias” generally are considered to be more closely allied to the Hvalcllidae than the Cvinidae, and

thus transference of chillonid genera (subfamily Chiltoniinae) to Hyalellidae is recommended.

Key Worps: Prkreatochilzonia enopkthalma gen. el sp. nov., new species, amphipod, artesian springs,

Australia, Taxonomy;

Introduction

Recent studies of the mound spring fauna of the

Great Artesian Basin in South Australia have

revealed a diverse endemic fauna (Ponder 1985,

1986; Ponder et al, 1989; Zeidler & Ponder 1989).

This fauna, components of which are endemic to

certain springs or spring groups, is dominated by

hydrobiid molluscs (Ponder ef qa/, 1989) and

crustuccans including amphipods of the genus

Austrachiltonia Hurley, 1959 (not Afrochiltoma

K. H. Barnard, 1955 - see Zeidler 1988).

During a recent survey of the fauna of Dalhousie

Springs in the extreme north of South Australia

(Zeidler & Ponder 1989), we discovered several

populations of an undescribed amphipod similar

to the Aus(rochilionia species commonly found in

olher mound springs, but without eyes. A detailed

examination of this species has tevealed several

characters whivh preclude it [rom Austrochiltonia

or any other allied genus. [therefore propose a new

genus TO accommodate this new species.

Chiltonias (subfamily Chiltoniinae) also occur in

New Zealand. (Chiltonia Stebbing, 1899) and South

Alrica (Afrochiltonia), but until now no phreati¢

species. have been described, although Williams

(1986) reports the probable occurrence of some in

Western Australia, from springs and caves near

Perth.

The familial placement of chiltonias is briefly

véassessed and they are considered to be more

closely allied to the Hyalellidae than ihe Ceinidae,

and thus transference of chiltonid genera to

Hyalellidae is recommended.

* South Australian Museum, Nosth Terrace, Adelaide,

S. Aust. 5000

fearon Airing

| \

' 5

\ Worthern \__ Tervitory

i <7

— a

Dalhousls Springs» \

Austratia

Fig. 1. Loeation of Dalhousie Springs, South Australia.

Materials and Methods

Specimens were collected from five rather small,

Telatively cool, seeps amongst the Dalhousie Springs

complex (Fig. 1), The springs are coded following

Zeidler & Ponder (1989) (see Fig, 2), Animals were

Wh W. ZELDLER

Laele tl, Temperature measurements, al tive of collection, for habtiats of Phreatochiltonia auuphthalma sp. nov

and physteochemical data (from Smith 1989) for nearby springs (meusured us close as possible to source).

Temp. Temp. Nearest Celi Chemistry’ ee _ bab Chem,

Habirat Ait Water Spring Temp, cond. TDS PH DO, TDS PH

°C is 0G 25°C mg/L. ppm me/T

Siemens

{'b) 3 20 None - - = - — - -

Dbda 225 22.5 Db3 a5 1500 830 7! 0.6 oO. 7.6

Nih of be - - kz 46 1600 890 74 19 - -

8 2 2 El 23 1480 820 VT 7.5 930 73

(iad Zz 7 Gal AZ 2100 1180 T1 10 1220 wo

‘ Gas 44 212 1190 TI ZI 1275 77

collected with a small hand sieve ov picked off plant Systematics

debris with forceps. and once, when no surface

water was present (at Chi). two specimens were

eollected using. a houschald corron mop placed in

4 pil dug to water level (Approx. Im). A toval of 676

specimens (3669 9, 26340, 47 juveniles) were

collected and exammed,

Physicochemical daia for the sites sampled are

unavailable because too little surface water was

present for meaningful analysis. However, air and

waler temperatures were measured at the time oF

collection, and these and physicovhernival dita for

nearby springs are given in Table J.

Material reported here is deposited in the South

Australian Museum, Adelaide (SAM), and the

Australian Museum, Sydney (AM). All specimens

are preserved im 75% aleohol or 2%

forimaldehyde/propylene glycol solution, OF the

types, only the holotype, allotype and one paratype

o have been dissected (partially), with appendages

removed from the Jeft hand side of the animal

unless indicated otherwise. Dissected appendages

are preserved with the carcass ov, in the case of the

holotype, the mouthparts, pleopods 1-3, and the

uropods and telsun are mounted in poly-vinyl

lactophenol on a microscope slide.

Specimen lengih is measured alony a tareral

purabalic line drawn from the anterior extremity of

the head through the middle of the body ro the

pesterior limit of the telsau.

The (horacic limbs are relerred to as gnathopod

| and 2 followed by pereapods 3-7 to avoid

confusion. Size comparisons of gnathopods exelude

the coxa und daetyl, and of the pereopads, the coaka,

with articles being measured alona the middle.

‘The following abbreviations are used in the text,

AL = first antenna; Gl = gnathopod I (first

pereopod); G2 = gnathopod 2 (second pereopod);

P3-7 = pereopods 3-7; U1-3 = uropods 1-3.

Phrestochiftonia gen, et sp. siete

Type species: Phreatochiliania anaphtkalma

5p. nov.

Diagnostic definition: Males about 44 size of

females, Fyes absent. Antenna | longer than

antenna 2, about 's body length; ventral aesthetase

al base of distal (hree or four (rarely five) flagellar

articles. Antenna 2, length almost 0.7% antenna 1;

gland cone pressed against head, not readily visible

laterally, Maxilla L without palp, notched at palp’s

normal position, Coxaé 1-4 deep, Coxa 4 withour

well-defined posterior excavation; maximum width

greater than length (up to 1.25% in femtale).

Gnathopods | and 2 subchelate in both sexes; article

6 of gnathopod 2 enlarged in male. Pereopods

slender; pereopod 5 distinctly shorter than pereopud

4; pereopod 6 distinctly shorter than per2opod 7.

Pleopods unmodified in both sexes. Uroped 3 with

Single cumus or ramus uwbsent. Telsan entire,

subrectangular with slightly concave distal margin.

Coxal gills on gnathopod 2 to peregpod 6.

Oostegiles on gnathopod 2 to. pereopod 5,

Elymoplogy: The generic name, Phreatochiltonia,

refers to the phreatio habit of the type species, and

acknowledges its similarity to. Austrochiltonig,

Systematic status of Phreatochiftonia gen. nov.

and other chiltonid genera.

The new genus clearly belongs to the “chiltania™

generic group (subfamily Chiltoniinge). tc is must

similar 10 Austrachil/tonia, bul is readily

distinguished from this and the other two chiltonid

venera (Chilfonia and A/rochilionia) by the

tollowing characters; 1) lhe phreatic habit; 2) the

complete lack of eyes; 3) coxa 4 is without well-

Tis. 2. Dalhousie Springs complex showing sites (coded) from which Phreatochilionta anephthaima gen, evsp. nov,,

has been collected and adjacen) springs (referred ro in Table tl). Seale =

springs ancl creek becls ave stippled,)

12100,000, (Qther springs, swamps from

—_

NEW SPECIES OF PHREATIC AMPHIPOD 179

bat %,

“Dalhousie. pes

180) W, ZEIDLER

defined posterior excavation, and 4) pereopad 5 is

distinctly shorter than pereopod 4,

The current familial placement of the chiltanid

genera is uhcertain. Barnard (1972a) originally

placed the chiltanias in his new lamily, Cemidac

(Barnard 1972b). Bousfield (!982) however, includes

the chiltonias in the Hyalellidac, in a brief synopsis

of the family, but gives no reasons for this

Tearrangement; it has chus been ignored or not been

aveepted (eg. Zeidler 1988; Barnard & Barnard

byt).

‘The welusion of chiftoniss in the family Ceinidae

by Barnard (1972ai as not altogether satisfactory,

They differ from Ceina Della Valle, 893, the pe

genus, in scveral key characters; Ceine has ait

aberrant mandibulat molar consisting of a Jong,

conical, non-triturative protuberance, and a flat,

distinctly cleft telson, whereas chillonias have a

relatively large, triturative mandibular molar, and

the telson js curved over the anus and is not Cleft

except in Chiltan rivhiwake (Chiltan, 1898) (for

distal 4), Chiltonias appear to have more in

common with Ayalella Smith, 1874 and

Allorchestes Dana, 1849, particularly with respect

tothe mandible, uropod 3 and the telson. | therefore

reccommend that the chiltonias, Afrochiltonia,

Axstrochiltonia, Chdtenia and Phreatachiltenia

gen, nov, he placed in the family Hyalellidae

wogether with Allorchestes, Ayalelfa and Partiyeleltla

Kunkel, 910, subject (oa thorough revision of the

hyvalellids,

Pheeatochiltonia anophinalma sp. noy.

FIGS 3-35

Aasvochiltonia sp. Zedle®, 1989, pp, 83 84, fig. LIA.

Australia, Dalhousie Springs, from small seep uf

sprine Db4 (Db4a), 26°26'49"S 19592915 "B, W,

Zeidicr & KL. Gowleti-Holmes, 9.vi.1985_

holotype.

Paratypes; All with same collection data as

holotype, except AM specimens collected by W. F.

Ponder & D. Winn. AM P40445. 30 9, foc.

SAM C4230, Lo 3.1 mm. SAM C4231, 1369 9 (19

oviverous), 2c oc, 39 juveniles.

Other material examinca; Alt from Dalhousie Springs arva

(hig 2) AM P4446, 19, spring Cbl, 26°25 16"S

125°29°52."F,, from shallew pool at top oF niaund, W. F.

Ponder, 30.v.1983, SAM 4232, 2-2 9, spring Col, tram

mop tmp in pit dug to water level lapprox. liny, W. Zeidler,

l4.vi I9RS, SAM C4233, bor, from small seep of spring

Db4 (Db4b), 26°26'49"S 1RS°29 1S "h, W. Zeidler & K, £-

Gowlcte Toles, 997 98S. SAM Cd7I4, 294 (both

ovigerous), Hom <enrall seep ameangst trees (Melaleuca

glomerata) just north of spring &2, 26°284'30°S

98°29 DS"E, W, Zeidler & KO | Geawlett-Hotnies,

vi 1985. AM Tadad7, 239 9 lone ovigerous), Wer cr,

spring E& 26°28'21"S 135'38'52"E, from small seep on

north side of (mound, W, F. Ponder & D, Wink, TS.) I9RS.

fromm smut! seep on nomh side of mound, W. Zeidler &

kK. L. Gowlett-Holmes, 10vi1985, AM P40448, 277 2

fone ovigerous), 60°, spring Gu4, 26°29'23°S

135529°10"R, from small seep, W. Ponder & 2. Winn,

41.1985, SAM C4236, 759 9 (20 oviverous), 20.7 o, 8

juveniles, spring Gud, from small seep, W. Zeidler & K-

1. Gowlert-Holmes, Svi1985.

Definitions As for genus with the following

udditions. Specimens entirely white, relatively small,

body length of femules up to 4.4 mm, males up to

3.2 min; relatively robust. Coxal plates 1-3 slightly

wider proximally than distally, Ooslegites relatively.

large, that on coxal plate 2 almost as long as

enathopod 2. Urapod 3 4 single article

Description of holotype Male 2.9 mm (Fig, 3),

Coxal gills sausage shaped, present from G2 to P&é.

Head abuul as lung as deep, length equivalent

io first 1-S pereonites,

Antenna | (Fig. 4) abour 3= head lengthy article

Lof pedunele, width 0.4= length and slightly more

than 1.3.< length article 2, article 3 slightly shorter

than article 2; flagellum length almost 15+

peduncle, of nine articles with one ventral aesthetasc

at base of each of last four articles.

Antenna 2 (Fig. 5), article 1 of peduncle as long

as wide, article 2 almost twice as lony as wide, twice

length article | and ahour 2% Jength article 3;

flagellum only slightly longer than peduncle, of

seven articles.

Upper lip (Fig. 11%; slightly wider tham lone,

apically rounded. bearing numerous short setae

distally,

Lower lip (Fig. 12): inner lobes vestigial, outer

lobes subovale with setose distal and inner margins,

Mandibles without palp: left (Fig, 9) with incisor

of seven tecth plus one tiny protuberance, Jacina

mobilis of five teeth, spine row of three feat here!

spines, molar trifirative; right (Fig, 10) with incisar

of seven teeth, lacina mobilis of four teeth, spine

row of two feathered spines, molar triturative with

one long Ceathyered sett,

Maxilla | (Fig. 7) outer plate with gine cours

like spines apically; inner plate very narraw with

two feathered spines apically.

Mazxilla 2 (Fiz. 8}: outer plate slightly longer thin

inner, selal row restricted ta apex; inner plate with

one large seta on inner margin about 0.6 from base.

setal row apically and along inner inarein almost

to large seta.

Maxilliped (Fig, 6): inner plate reaching extremity

of article 1 of palp, rectangular, width aboul 3x

lenech, with three stout leeth apically, the inner ane

NEW SPECIES OF PHREATIC AMPHIPOD 181

Figs 3-12, Phreatochiltonia anophthalma gen. et sp. nov., holotype o*. 3, lateral view; 4, antenna 1; 5, antenna 2;

6, maxilliped; 7, maxilla 1; 8, maxilla 2; 9 & 10, mandible, left & right; 11, upper lip; 12, lower lip. Scale bars -

0.5 & 0.1 mm respectively.

NEW SPECIES OF PHREATIC AMPHIPOD 1s

smaller, few feathered setae apically and along inner

Mazpin, Ould plate ovate, reaching 0.74 along article

2 of palp, shightly wider than inner plate, bearing

several setae apically and aleme about distal “2 mner

margin: palp article | with oblique distal margin,

length outer margin about 2.5 « anner; palp article

2 slightly broader than Jong, slightly expanded

distally, as long as outer margin article tL, bearing

group of sctae on inner distal corner and along

distal *% inner margin; palp article 3 about a3 long

as broad, slightly expanded distally, as long as

article 2, bearing close-set sctac on inner distal

corner and for about distal !s inner margin, several

setae On outer distal corner and outer 4 distal

margin, palp article 4+ small, conical, width %%

length, slightly less than 4 arti¢le 3, four setae

terminally; dacty! sharp, slightly longer than article

Grathopoad | (Fig. 13 coxa width 0.8% length,

slightly longer than article 2, anterodorsal and

posteradorsal comers slightly produced, distinctly

wider proximally, anterior margin little longer than

posterior margin, distal margin evenly rounded with

several evenly spiced setae; article 3 length about

1.2» maximum width, posterodorsal lobe with

close-set row of eleht stout, pectinate spines; article

6 irapezoid, half as long again as article 5, slightly

wider distally, width abour 4 length. posterodistal

comer wilt two sldul spines on either side of dacty!,

several long sctac on anterodistal corner and on

posterodistal Jobe, several srnall setae on distal

margin: dacty! slightly shorter than widih article

6, lilting neatly against palm.

Gnathopod 2 (Tig, 14): leneth about L4e Gh

coxal gill tength more chan 3 width, little shorter

Than cox; coxa slivbuly longer thao wide, about

OB» article 2, Wistingtly wider proximally, distal

margin evenly rounded with several evenly spaced

selae; article 4 with right-angled bend; article 5

small, withour pectinate spines; article 6 little

shorter Than article 2, length anterior margin |,2

maxiinum width, postcroproximal corner forming

rounded lobe, palm oblique with numerous spines

of yaryme lengths on either side of cutting edge

(allowed by shallow eroove for tip of dactyh; dactyl

claw-like, as long as anterior maryin article 6.

Pereupod 3 (Fig- (5): slightly Jonger than G2;

coval vill length 2,5 width, little shorter than coxa;

coxa like that of G2 but slightly larger; article 4

length 2% width, about 0,5» article 2, anteradistal

comet produced; article 3 length 44 article 4; article

f about as long as arncle 4; dactyl length 0.5»

ariiele 6; ull articles sparsely setuse as illustrated,

Pereopod 4 (Fig, 16): slightly shorrer than P3,

‘et

otherwise identical except for coxa; coxa distinctly

wider than tong, wider distally, anterior margin

straigh|, distal margin evenly rounded extending to

small posterodistal lobe, posterior margin with

Slight indentation but without proximal excavation,

Perepod 5 (Fig, 17); length about 0.9% P4; conal

pill nearly twice as long as wide, as long as article

2; coxa width about 1.5% width article 2, length

anicrior lobe almost !4 width coxa, length posteciot

lohe 0.6% width coxa or O.8« length arucle 2;

article 2 slightly longer than wide with {ypical

expanded posteriar margin and posterodistal lobe

reaching to about '4 article 3; article 4 length 13x

width, 44 Jength article 2, with posteradistal corner

produced; article 5 shghtly longer than article 4,

width about ¥4 length; article 6 length 1.3» article

5, width 4 length: dactyl length about !4 article

6; all articles sparsely spinase as illustrated,

Pereopod 6 (Fig, 18): length 14x PS; like PS

except for coxa} coxa almost as wide as article 2,

length anterior lobe \% article 2, length posterior

lobe 34 article 2 or 0.8 width voxa.

Pereopod 7 (Fig. 19): longest pereopod, leneth

about 1.2% P6; like P6 except coxa is semi-circular

and lacks coxal sill, width 1.4« length, and article

2 posterior margin 14 disinctly serrated above

Wisertion of setae with acute proximal shoulder.

Pleopods (Fiz. 203: all unmedtfied (cf.

Chiltonia).

Uropod 1 (Fig, 2i): length about 1&« U2;

peduncle with small spine o4 inner and outer distal

curners, and three Jaree spies on dorsal outer

margin; rami equal in length, about 45 peduncle,

outer ranius with two large and one small spine al

lip and one large spine medially on dorsal marein;

inner ranius with ewo large and rwo small spines

#t lip and one large spine medially on dorsal

margin,

Uropod 2 (Fig. 22): peduncle with two large

spines on dorsal margin, one medially, the other

near distal comer: rami subequal, as long as

peduncle, outer ramus with one large spine

terminally, one small spine subterminally and three

large spines on dorsal margin; inner rainus like outer

bul inaddinon a row of five small spines, adjacent

to larger spines, on inner dorsal margin.

Uropod 3 (Fig, 23); one-articulate (rami absent),

almost “ length telson, rounded [n cross-section

wilh one (right) oc two {left) euler and one short

inner sela al tip,

felson (Fig. 24): entire, subrectangular, length

about 44 width, distal margin slightly concave with

(wo small setae at cach corner,

Figs 14-24. Phreatoctilinmia anapathatme pen, csp, nov, lolowpe o, (3-19, snuthepuds Lb & 2, perenpous 3-7,

2, pleapods I-3 (feathered setae on 2 .& 3 nue shown); 24-23, uropods 1-3, 24, telsan. Scale bar = 6.2 mm.

NEW SPECIFS NF PHREATIC AMPHIPOD

Deseription of .allaiype: Female 3.8 mm.

ovigernus With 28 eggs in brood-pouch, Jike male

except for the following

Antenna 1) flagellum length 14% peduncle, of

eight articles with one ventral acsthetase at base ol!

each of last three articles.

Antenna 2: Magellum length slightly more than

iJ peduncle, of six articles

Gnathopod | (Fig. 25): article 5, posterodistal

lobe with clase-sel row of nine, stout, pectinule

spines; article 6 more reetangular, width ahour 0,6:

length, stout spines on: posteradistal corner cither

side of dacty! shghtly larger; dactyl a little longer

than width article 6,

Gnathopod 2 (Fig, 26): like Gl but leneth about

12» GL, and article 5 posterodiscal lobe with close-

scl yow Of seven, stoul, pectinaly spines.

Pereopod 3! (Fig. 27); length about 12% G2.

Persopod 4 (Pig, 28): coxa slightly wider than in

male, leneth only 0.8 width.

Percopod 5 (Fig. 29); coxa slightly wider than In

male, width about 1.8 width article 2; article 2

with small posterodistal lohe; article 4 slightly

longer than article 5,

Pereopod 6 (Fig. 30) and pereopod 7 (Fig. 31);

as for holotype, but. article 2 slightly narrower and

with very small posteradistal lobe in P6.

Oostegites (Figs 32-45): on coxae 2-5, all with

¢urled margins and numerous small hooks, together

forming tight marsupium. First is heart-shaped,

slightly shorter ihan G2, length L.&x* maximum

width; second is irapezoid, length 0.8% P3, 2.7%

width proximally and 2x wadth distally: thind is

similar to second bus swith concave posterior

margin: fourth is subovare with very convex anterior

margin and oblique distal margin, length about

06% PS, L4% maximum width,

Pleopods, uropods and telsan as in Hololype.

Evinology: The specilic name anophrfalma

refers ta the absence af eyes,

Variation

Apart from variations due lo size (e.g, number

of Flagellar articles of antennae), paratypes and

other material examined was remarkably similar ro

either the holotype or allotype Minor differences

nored were as follows. The number of aesthetascs

on Al varies with some males having an additional

one (five}, and some females an additional one

(four) or rarely two {five}, In the male paratype

from the AM collections [AM P40445) G2 Irom the

Figs 25-

32-15, postegites from cowa 2-5. Seale bar — 0.2 mm.

left is noticeably smatler than trom the nght, in all

other specimens homologous pereopads are of”

similar size, The spination of U1&2 varied slightly

with a few specimens with more or less spines

(usually only one, al most (wo), In one specimen

(female), from spring ES, U3 from the right is two-

articulate (unitamous) as found in Austracéilfonia

avsivalis (Sayce, 1901), Oostegites of females vary

considerably mm size, but are expanded, as illustrated,

in ovigerous specimens.

‘The possibility that speciation may have occurred

between springs without any obvious morphological

changes was considered, and specimens for

allozyme electrophoretic analysis were collected

from springs Db4 (type locality), E8 and Gad. A

preliminary analysis of this material wsing methods

outlined by Richardson ef aj, (1986) indicated fixed

genetic differences of less than 10% (for 16 loci),

thus supporting the morphological evidence of one

species with little variation, Given these results, a

more detailed analysis was considered unnecessary.

Diseussion

The new species described here closely resembles

an undescribed species that | have collected from

springs Scattered throughout the Flinders Ranges

in northern South Australia, sympatric with one or

more undesvribed Crangonyctoid species. However,

this other species a8 nol desenbed here as

considerably more work is required (o UeteTmine

whether or nor one of more species are present,

Phreatochitenia anephihalma has been found

outly ia Tew, relatively cold, seeps amongst the 10)

or 50 springs and mounds knows as Dalhousie

Springs, anos of which were sampled in 1985

(Zeidler & Ponder 1989). Many of the active springs

at Dalhousie are warm (> 30°C) with large

outflows (Smith 1989), and since freshwater

amphipods prefer cooler waters (Barnard & Barnard

1983), il is nol surprising that no phreatic

amphipods were found in these springs. However,

a species of Austrochilionia was found in two of

these warm springs, but only amongst sedges along

the edges of outflows where the water was

considerably cooler (Zeidler 1989).

The habitat of B anophthalma is very restricted

and animals were seen at the surlace, burrowing in

and out of the substrate, only near points of water

discharge, or were found under plant debris nearby.

The species was moderately abundant at most sites

execpt springs Cb] and G2, which had tittle ur nu

surface water present, The habitay at the other sites

35, Phreatochiltonia anophihatia gen. et sp. now. allorype 9G. 25-31), pnathopods L & 2, pereopods 3.7:

ts) W. ZBINDER

was very limited, consisting of small seeps about

3m long by 20 cm wide, and only a few millimetres

deep. At Cb! one specimen Was collected incidently

with hydrobiid molluses in May 1983, frum # small

shallow pool on the top of the mound, [n 1985, this

spring was completely dry, but two specimens were

collected from 4 mop trap placed in a hole dug to

the water level, adjacent to the mound, This

evidence suggests that these amphipods are

essentially sublerranean in habit and are only seen

at the surface when underground water pressure is

sufficient to breach the surface of the mound. The

nowion that these animals, and perhaps others,

inhabit the cool interstitial water beneath and

between mounds is an interesting prospect that

neals further investigation.

Factors that determine the distribution af this

species are unknown, but since the habirat appears

very limited, it may be more vulnerable to external

influences than are the habitats of larger springs.

Presence of surface water is probably unnecessary

for the survival of the species (eg, al Col), and on

a brief visit in April (986, all habitats had

contacted, animals were difficult to collect, and

hall of the habitat of E& had dried up. However

when surface water is present, they may concentrate

at the point of discharge

Nothing is known about the lite history of this

species, but of the 366 females collected, 97 were

ovigerous, and recently hatched juveniles were

common.

One can only speculate as to the evolutionary

origin of thi§ species, but mts occurrence af

Dalhousie Springs on the edge of the Simpson

Desert may indicare that it is a descendent of a

species which was more widespread! during a time

when central Australia was much wetter than it as

today (Krieg 1989). The phreacie habit having most

likely evulved in response to selective pressures in

an arid environment,

Acknowledgments

Lam most grateful to Dr W. F. Ponder (AM), who

dared to suggest thal phreatic amphipods might

exis| at Dalhousie Springs, and who collected the

first specimen in May 1983, provoking a oiure

intensified search in 1985. He is also thanked for

his assistance inthe field and for collecting more

specimens, as is Ms L. Winn (AM). Mrs K, F.,

Gowltit-Holmes (SAM) assisted erently with field

work and the collection of specimens, and alsy

typed the martuscript. Mr M. Adams, Evolutionary

Biology Unit, SAM, conducted the electrophoretic

analysis, and his expertise is gratefully

acknowledged. | also acknowledge constructive

comments of an anonymous referee witich markedly

improved this paper.

This study was supported by funds from the

South Australian Musturm.

References

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Zealand: Algae-living littoral Gammaridea (Crustacea;

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______ (1972b) Gammaridean Amphipoda of Australia,

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& BarnaArb, €, M. (1983) “Freshwater

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BousFiFLn, K. Lb. (1942) Malacostraca pp. 254-294-

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—___ (1986) Mound springs of the Great Artesian Basin

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, HERSHIER, R. & Jenkins, B. (1989) An endemic

radiation at tydrobiid snails from artesian springs In

monhern Souch Australia: their taxonomy, physiology,

distribution and anatomy. Mulacologia 31(1), 1-140.

Rronakuson, B., Baverstock, P, Ro & Apams, M.

(1986) “Allazyme Tlectrophoresis.” (Academic Press,

Sydney),

NEW SPECIES OF PHREATIC AMPHIPOD 187

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fresh-water Amphipoda. Proc. R. Soc. Vict. 13(2),

225-242.

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Zeidler & W. F. Ponder (Eds) “Natural History of

Dalhousie Springs.” (S.A. Museum, Adelaide),

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the United States. Rep. Comm. Fish & Fisheries Part

II, Report for 1872-73, 637-661.

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Copenhagen Museum and other sources. Part 2. Trans.

Linn. Soe. Lond. 7(8), 395-432, pls 30-35.

WILLIAMS, W. D. (1986) Amphipoda on land-masses

derived from Gondwana pp. 553-559. Jn L.

Botosaneanu (Ed.) “Stygofauna Mundi. A faunistic,

distributional and ecological synthesis of the World

fauna inhabiting subterranean waters (including the

marine interstitial).” (E. J. Brill/Dr W. Backhuys,

Leiden).

ZEIDLER, W. (1988) A redescription of Afrochiltonia

capensis (K. H. Barnard, 1916) with a review of the

genera of the family Ceinidae (Crustacea, Amphipoda).

Ann. S. Afr. Mus. 98(5), 105-119.

______ (1989) Crustacea pp. 79-87. In W, Zeidler & W. F.

Ponder (Eds) “Natural History of Dalhousie Springs.”

(S.A. Museum, Adclaide).

& PONDER, W. F. (1989) (Eds) “Natural History

of Dalhousie Springs,” (S.A. Museum, Adelaide),

PARAPHAULOPPIA (ACARIDA: CRYPTOSTIGMATA: ORIBATULIDAE)

AND ITS OCCURRENCE IN SOUTH AUSTRALIAN SOILS

BY D. C. LEE* & C. M. BIRCHBY}

Summary

Paraphauloppia Hammer is redefined and compared with other oribatulid genera. The description

of Paraphauloppia novazealandica Hammer is extended and four species are described:

Paraphauloppia acutinotata sp. nov., P. globata sp. noy., P. obtusinotata sp. nov., P. triforata sp.

noy. These mites are from soil and plant litter or moss at four of nine sites sampled in

South Australia. A key to adults is given for these five species: this is the first record of

Paraphauloppia from Australia.

KEY WORDS: Paraphauloppia, Paraphauloppia acutinotata, Paraphauloppia globata,

Paraphauloppia obtusinotata, Paraphauloppia novazealandica Hammer, Paraphauloppia triforata,

new species, Australia, plant litter, soil.

Hranseeiiin af thet Raval Saucier af & Aust (1991), 19SE4), 18% 194.

PARAPHAULOPPIA (ACARIDA; CRYPTOSTIGMATA: ORIBATULIDAE)

AND ITS OCCURRENCE IN SOUTH AUSTRALIAN SOILS

by D.C, Lee* & C. M, BIRCHRYT

Summary

Lit, DC, & BIRCHBY, C. M. (1991) Peraphauoppia (Acatida: Cryptostigmalta: Oribatulidae) from South

Australian soils. Tran Ro See. S.-Aust. (Sid), 184 19% 29 November, 1991.

Paraphanioppia Hammer is redefined and compared wirh other oribarutid genera, he description of

Paraphaulopple novaceatendica Hammeér ty extended and four spevies are described! Padraphialeppia

ucutinolaia sp. nov., PR. slobuta sp. nov., P obtusinatata sp. nov., 2 [riforata sp. noy. These mites are from

soil and plant litter or moss at four of ning sites sampled in South Australia, A key to adults is given for

these Jive species: this is the first record of Paraphanloppia trom Australia,

KEY Worns; Paraphauloppia, Paraphauloppia acutineiera; Paraphauloppia slobata, Paraphautappia

oftusinatata, Puraphaxloppia novasealandica Hamner, Paraphauloppia trifarata, new species, Australia,

plant litter, soit,

Introduction

Paraphauloppia Hammer, 1967 is examined as

part of a current study of sarcoptiform mites in

South Australian soils, sampled from nine florally

diverse sites, and for which Lee (1987) provided an

intraduetion to the relevant work on the advanced

oribate mites (Planofissurac).

Paraphuuloppia is allied 1o Oribatula Berlese,

1895, Phaulappia Berlese, 1908, Zveeribatula

Berlese, 1916 and Jornadia Wallwork, 1984 within

the Oribarulidae Thor, 1929, which is applied here

to a taxon approximating to the Oribatulinac of

Balogh & Balogh (1984). The Oribatulidae in this

restricted sense have been subdivided into the

Oribatulinae, Pscudoppiinae Mahunka, 1975 (see

Lee 1987) and Fovoribatulingae Lee & Birchby, 1991.

The Fovoribatulinae have since been transferred to

the Cragsoribatulidae, when the latter was newly

given family rank (Lee 1991), The original definition

of Poraphavloppia is modified to accommodate the

new species and distinguish it from Jornada, but

also to delineate if trom similar family-group taxa,

because the definition of the Orlbatulinae as

currently used is unpublished (Lee in prep.) 2

aovazealandica Hammer, 1967 is newly recorded

Trom Australia, and four new species are described

from South Australia,

Materials and Methods

New marcrial examined here, collected by DxC.L,,

is deposited mostly in the South Australian Museum

(SAMA), but also in the British Museum (Natural

History) London (BMNH), the Field Museum,

* South Australian Museum, Notih Terrace, Adeluide, S

Aust,, 5000 ;

+ State Herbartim, North Terrace, Adelaide, S Aust., O01

Chicago (FMNH) and the New Zealand Arthrapod

Collection, DS.1.R., Auckland (NZAC), whilst

previously described material is. deposited in the

Zoological Museum, Copenhagen (7MC). The

morphological notational system follows Lee (1987),

Ihe somal chaetotaxy of which is summarized in

Figs 3 and 6, with the total sciae present in each

file (eg. 6Z) indicated by number coming [irst,

whilst a particular seta (eg. 76) would have the

number last. The abbreviations for zoogeographical

regions follow Lee (1970, fig. 427). The descriptions

of eggs refer to those within the female soma, .All

Material was examined using a Nomarski

interference contrast device. All measurements are

in micrometres (um) and were made using an

eyepiece micrometer at «250 magnification.

Systematics

Paraphauloppia Hammer

Paraphauloppia Hammer, 1967, p. 45 (type species by

monotypy; FPurapkauloppia nevasealandica Hammer,

197); Coetzer, 1968, p. 58; Balogh & Balogh, 1984, p.

272; Luxton, 1985, p. 68,

Definition: Hysteronotum with 10 pairs (2.4, 67, 28)

of setae and three or four pairs of multiporose

foramina, pteromorphs absent. Dorsosejugal

lurrow entire (sometimes /gint), arched, not

extending forward ta level of setae 2. Proter-

onotum withour translamella, prelamelta,

sublamella or tulorium, narrow costate lamella

(between setae zl-z2) present. Ventrosejugal

apodeme forming single, continuous bar across

midsternal line. Coxite seta /¥1 about level with

/Zg\, Genital shield with three or four pairs of

setae. Discidium furms low costate ridge. Femora

Land 1) with five setae (0,2/2,1). Tarsi long (total

WQ)

leneth of tarsi, HW, LS ane 1V more than 7.5% of

idiosomal length) and slini (height less than 33%

of length). Pretarsus with three claws, usually fateral

claw conspicuously slimmer than central claw

lexception P triforaia sp. nov., all three claws slim)

Geeeral morphelogy: Somal leneth range tor

adults: 259 a5 (South Amerean species larger,

330-650; South Australian species smaller,

259 165). Sumal chaetotaxy: 2), 22, Is; 24, 62, 2S;

YoU, BIA, ATV usually 4/72 fexceplion: #

novaredandiva WMammer 3/22), Se, 2/2a, 38a,

Leo chaetotaxy (solenidia in purenthesest: Lb - L 3,

ath 4(2), 22) LE - by 5 AUN) A(t). 1OE7); UL - 2,

31), 3), HAQ2) PY 1, 2, 2,310), 12. Totes tiiment

mainly smooth, gerateguinent inconspicuous,

sometimes (Australian species excep! P

nevazvalundica Ua mer) fine longiudinal lines in

ponite region, Hysteronotal setae Setose, Weakly

pilose, sety <2 clavate or plabose. Subpedal and

circuimpedal ridves merged intu single continuous

line Pemorca Jack ventral flanees, femur | without

caput collar

Distribution South America (INic; Argentine,

Bolivia, Chile, Peru), Ausiralia (Au; South

Australtal and New Zealand (An: South Island).

Within South Australia, Pommphauloppie iy recorded

from four sites each representing a habitat type

{mallee-broombush, piallee-heath, savannah

woodland, «clerophyll forest) with native vegetation

and in a Mediterraneanaype climatic cegion. In

South Aimerica and New Zealand Puraphiuloppia

occurs in master, cooler regions. Therefore, despite

its absence from the South Australian most

southerly, moist, evoasial site, Panphuulopoia

probably also occurs in the moister, temperate

environments of Australia.

Remarks: Peraphauloppia was considered similar

tn Subphavloppia Hammer, 1967, and so lo

Phawlapnia Berlese, }908, when itavas established

with one speeies from New Zealand (An). Now, it

could, be considered imost similar bo ferctwdio

fallwork, 1984, known from North and South

America, which differs in having five genital setae.

Coetzer (1968) included in Parechauleppie a lurther

vight of Hammer's South American species thal had

been grouped in either Oribatyla Beclese, 1895 or

fporbaruia Selinick, 1928. One of these species,

P austealis (Hammer, 1962), has since heen

mistakenly tsed (Balogh & Balogh, 1984, fig 70)

wi quder co illustrate Aporibaiuda, despite its being

excluded from thal genus in the same publication

en te basis of having ten pains of hysteronotal

setae. Until the present publicmion, oo other

changes have heen made to the genus. A paper (lee

DBC. LEF aC

M, BIRCTIBY

i99}) of the higher classification of jhe

Oripodoides, however, gives more Weight to the

presence of a central gap in fhe yentrosejgal

apodeme in delineating subfamilies, Consequentty,

Diphavlappia Balogh & Balogh, 1984, Gerlawbia

Coetzer, 1968 and Subphanlonpie were wranslerred

from the Oribatulinae to the Pseudoppiinae, so that

although previously regarded as similar to

Paraphauloppia, (hey are now included in another

subtumily, Phas #araphanloppia and Jornudia ave

the only oribatuline genera with ten pain of

hysteronotal setae, all the others having M-14 pairs.

Paraphauloppia cordylinasa Higgins & Woolley,

1978 is listed in ihe North American fauna by

Marshall ef a/ 41987), noting it as a problematic

combination; since it has at least 12 hysteronoral

sctal pairs it is excluded here from Paraphavloppia.

South Australian and South American species of

Paraphuuionpia differ. The South American species

are larger, except for P graces (Hammer, 19558),

which is similar to A novezealandica in somal

shape, size and form of notal serae, and posutionmy

of nolal seta S§ behind 274. The South Americun

species also have smaller sensory sctac (22) and

hysteronotal foramina, and back fine striations on

their comailes.

Parephauloppta includes 13° species: £2

altimontana (Hammer, 1958 B ecudinarata <p.

nov, FP altimontanoides (Hammer, 1958) 1!

australis (Hammer, 1962); Bo glvbeter sp, nov: PB

gracilis (Hammer, 1958): 2 mnegniporosa (Hammer,

1958); PB moreno: (Hammer, 1962); FP

novazealandica Hammer, 1967 (type-species); PF

obtusinofata sp, nov. P pisacensis (Hammer, (961),

P. quadrisetosa (Hammer, 1961); PB trifarate sp. Ty.

Key to Australlan Paraphanloppra species tadalts)

1 Hysteronotal setae shorter (23 and 24 nol eachiny

22 or SS). (Fig. 7). Veutrosejuval apodeme well

separated from margin of genital orifice, distance

between setae WH-/7 21 at least as grear as length of

conte sora VAT |Fig. 8).

Syterante al setae longer (73 and Za reaching 72 or

SS) (tig. 1). Ventmosejugal apodeme abviting onto

margit of genital anifiee op close to i, distane

between see L-J Zeb loss than Jength ef cosile seta

POA Pig Sooo. Te ee a oe ee

2. Larger soma (length > 300 ain), Three pains of zeriital

setae Pour pairs of hysleronotal foramina, Notal sete

JL and Zt shorter than 22 ar 73... .....0.-----

tithe A novazealandica Hammer IS?

Smaller soma (length < 3) pm), Four pairs of genisal

setac, Three paics of hysteronotal foramina. Notal

sere J2 and 7! Jonger than 22 or 23 (Pig, R)--

PB rriferaty sp, nav

a 1 areer soma (lenuth > 350 yin). Flysteroporna

Subelubular, Pysterangtal setae long (23 lengih a

PARAPHAULOPPIA,

ee We Wino denn dit A478 P gicbate sp. nov.

Smaller soma (length < 350 ym), Hysterosoma ovend.

Hysteronotal setae medium length (Z3 length

er ere

Paraphauloppia acutinotata sp. nov,

FIG. 1

Type material: Holotype female (SAMA N1990733),

plant litter, sparse moss and calcareous sandy-soil,

under ridge-fruited mallee (Eucalyptus incrassata)

amongst broombush shrubs (Mela/euca uncinata),

open scrubland, Ferri¢és-McDonald Reserve

(35°1S'S, 139°09'E), 20.vi.1974, Paratypes, four

2 2 (SAMA, N1990734 ~ N1990737) and four oc

(SAMA, N1990738 - N1990741), same data as

holotype.

Female; Soma oval, light brown, [diosomal length,

314 (5, 298-329), Leg lengths (femur-tarsus for 316):

1-169, H-(42, Ifl-142, 1V-180. Tibial maximum

heights (for 316): I-17, 11-12, Mfl=11, [V-12.

Proteronotum with terminal tooth to rostrum

forming narrow point, not as broad as long.

Anterior foramen (F1) absent. Seta j2 reaching

midway between /l-zl. Sensory seta (zz) clavate,

usually with caput longer (over 66% of total setal

length) than exposed stalk. Hysteronotum with

mainly medium length setae, but some peripheral

setae substantially longer, Z] usually longer than

22, S6 usually 1.25% length of 74. Seta SS well

forward su that level with seta Z4 and foramen /74

behind line between 24-55.

Idiosternal setae with inconspicuous cilia, long,

seta /2 reaching anterior margin of sternal tectum.

Coxite region striated from unterior margin back

to seta Sel, posteriorly striae superimposed on weak

reticulations. Discidial ridge with straight edge. Slit-

like pore Saf nearly longitudinal, less than 45° from

longitudinal axis. Egg subellipsoidal, exochorion

smooth, size 170 » 77 (1), length 52% of somal

length, eggs per female — 1 (1).

ORIBATULID MITES I

100.um

Fig. 1. Paraphaulappia acutinotaia sp. nov., G notum-of

soma, For sttal notation see Fig. 5_

Legs medium length (mean femur-tarsus length:

50% of somal length) with medium girth (mean

maximum tibial height 34% of mean length).

Central pretarsal claw with 23 depth of lateral

preiarsal claws,

Male> Similar to fernale but idiosama shorter, mean

length, 285 (4, 275-296).

Remarks: Vhe specific name acutinotata is derived

from the Latin for ‘pointed’ and ‘back’ and refers

D, C. LEE & C. M. BIRCHBY

192

*g pur ¢ SALZ 3a8 UONPIOU [eds JOJ “RUIOS JO WINUIAIS *¢ “BIOS JO WINjOU ‘7

wrvooL

“sou “ds oynqojs niddojnnydping *¢-7 Say

PARAPHAULOPPIA, ORIBATULID MITES {03

_di

Fig. 4. Paraphauloppia slobata sp. nov., 9, posterior aspect of ftemur-pretarsus of right legs 1, I, IIL and LV. all

setae in femora [ and II illustrated, d — dorsal, p = posterior, v = ventral.

to the sharp rostral tooth and anterior hysteronotal

margin. These distinguish it from the similar P

obtusinotata, sp. nov. as do the longitudinal slit-

like pore Saf and greater length of some peripheral

notal setae. P. acutinotata and P. obtusinetatia both

have medium sized hysteronotal setae and are

distinguishable from the other Australian species

which have either clearly longer or shorter relevant

setae.

Paraphauloppia globata sp. nov.

FIGS 2-4

Type material: Holotype female (SAMA N1990742),

plant litter, sparse moss and calcareous sandy soil,

under ridge-fruited mallee (Eucalyptus inerassata)

amongst broombush shrubs (Melaleuca uncinata),

open scrubland, Ferries-McDonald Reserye

(35°15'S, 139°09’B), 20.xi,1974. Paratypes, 16 9 9

He bo C, LEE & c.

(SAMA, NI9S07TSS -— NIg9IS2: I-BMINH;

|-FMNH)} L-NZAC; 3 lost) ind five a (SAMA,

NI990753 - NI9907S6; | lost), samme data as

holotype.

ornate: Soma hrowlly oval ur subspherieal in whapte,

light brown colour, Idiosomal length, 3459 117,

339-365), Lee feneths {femur — tarsus for 365):

1-213, 11-193, TH-188, 1V-239, Tibial maximum

heyshts (for 365): 1-21, 1-16, Pl-14, 1V-16.

Proteronotum with. terminal tooth ta rostrum

forming narrow point, not 4s broad ws long (in Fig.

Z not evident because directed downwards),

Anterior foramen (71) absent. Sela j2 reaching

anterior to zl to about O.8« distance between setae

/l ~ zl. Sensory seta (22) clavate, slim and long,

usually with caput subequal in length to exposed

stalk. Hysteronotum wath long setae, lengtli of setae

73 and #4 mare than 2« distances Irom 42 and

S35 respectively. Seta.85 nearly far enough forward

io be level with seta 24, but foramen F4 anterior

10 line between setac 24-85.

Tdhosternal setue with minute But neticesable cua,

lung sela 72 reaching anterior margin of slernal

levtuim, Coxe rewen with central area striated from.

anterior margin to seta ///1, laterally around seta

f2 lines faint and broken up into rows of short striae

aud dots, Discicdial ridge with tubercle bearing seta

FV, Slit-like pore Saf nearly transverse, more than

45 from Jongitudinal axis. Egg subellipsoidal,

exachonon rugose, mean size 166 80(10), length

47% of somal length, egas per female | (1), 2 (41,

33) or 4 (2).

Legs long (mean lemur-tarsus length: 57% of

somal length) and slim (mean maximum tibial

heieht 31% of mean length). Central pretarsal claw

with 2s depth of lateral pretarsal claws,

Male: Similar Wy female but idigsuma shorter, mean

length 342 (3, 337-347),

Remarks! The specific name globuta is derived frum

the Latin for globe’ and refers to Ne Avbspherical

shape of the hysteronorum. FP glebata ts

distinguishable from the other Australian species

by this shape, its relatively large size and long nolal

setae. ft is smaller chan all bur one of rhe South

American species, and differs from them in haying

such a long clavate sensory seta (22).

Purdphindoppia nevazealandica Yammer

Parapheuloppia nevazcalendica Wammer,

1967, p. 45, 46, fig. 59.

Tipe material examined: OF LL specimens recorded

with original description, dectotype co (labetled

“wpe” in vial af aleohol) and four paralectolypes

Qabelled “paratypes”, ane > and three ocr onvone

M, BIRCHEY

slide) examined (7MC), thick mass and bone-diy

lichens and Lycapodiwh, ope Murvku and

Nothufazus lorest, Tew 100 leet above lake, Lake

Ratoili, South Island, New Zealand, M. Hammer,

1962.

females Sama oval, straw colour (New Zealand

specimens) or light brown (South Australien

specimens), Idiosemal length (original descripuon:

“abour 0.34mim™}, 320 U1, New Zealand), 337 (25,

Sclerophyll forest, 326-346) or 363 (5, Savannah

woodland, 450-370), Leg lengths (femiur-tarsus,

Sclerophyll forest, (or 3413: L-E70, 1 151, Fi-142,

IV=l78, Tibial maximum heights (for 341): 1-19,

1-17, TH-92, LY-12.

Proreronotum with terminal tooth to rostrum

forming broad point, as broad as long {hot

ilusated by Hanmer, 1967; fig 59). Anterior

foramen (Al = “ap”) small but conspicuous. Shorr

ridge between setae /2-12 present (New Zealand

specimens) or absent (South Australian specimens),

Seta /2 as long as.0.75% distance between selae

J2-al. Sensory sela (z2) globular, with capur

subequal to or shorter than exposed stalk, two parts

clearly delineated. Anterior hysteronotal margin

completc, although faint, revoguisable between

dorsasejugal apophyses. Hysteronvotum with short

setac, Z], Z2 and 44 length subequal to distance

22-23, Seia $5 well posterior to seta Z4, and nearly

directly behind it, so foramen F4 lateral to line

berween setae 44-SS,

Idiosternal setae with inconspicuous cilia,

mecivm length, seta /2 not reaching anterior margin

of sternal feoclum, Coxile region not striared. weak

reticulations near midsternuot. Apodeme J!

present, sinall, thickened Jusion to coxite limited

10 shart linear strut or sub-circuwlar tubercle,

Discidial ridge with tubercle bearing sela PV3. Slit-

like pore Saf nearly longitudinal, Jess than 45° from

longitudinal axis. Ege subelfipsoidal, exvchurion

smooth, mean size 145 « 86 (11), lengih 46% of

somal length, eggs per female - | (9), 2 (1) 0631.

Legs medium length (mean femttu-rarsus length:

47% of somal length) with mmedium einlh trmean

maximun) tibial height 34% of mean length)

Central preiarsil claw with 2« depth of lateral

pretarsal claws.

Male: Similar to female hin idiwsoma usually

shorter, mean length, 328 (4, rypes.ex New Zealund,

320-334), 323 (25, Sclerophyll forest, 312-336) or

341 (6, Savannah Woodland, 326-355).

Referred enatertal: AL 9 -F (SAMA 1990999 - N1990944,

N1990955 - N1990964, NI99O09K} =INT990992, 4-BM NH,

4-PMNH, 4-NZAC) and 50 ero (SAMA NI9U0034

NIPY9093K, N1990046 - N 1990048 N}990965 - NI9900H,

1900803 - 819901006, 4. BMNI, 4.1'MNH, 4-NZAC),

plant litter, sparse moss Ofd calcareous sandysoil, vnde

195

PARAPHAULOPPIA, ORIBATULID MITES

“BWOS JO LUNUIOYS ‘9 “BUIOS Jo WIN]OU

wirooL

& “sou ‘ds piojoulsnigo viddojnnydping “9-5 satq

IW DC.IPE& 4,

selerophytlous shrubs amongst messmute stringyhark

(Eucatypius vbliqua), dry sclerophyll forest near sumimir

of Mt Lofry (3459'S, 198°45°E), Cleland Cunservanon

NIY9OLOO7, NIG9OLON$) and six of ot (SAMA NI990952

= S1990954, NIO9OTO09 ~ NISSOIOIY, wrass, mass, Jeal

litter and loamy soil under manna Lum Lives LEycalipius

viminaliy), savannah woodland, Chambers Gully

(34558'S, (3841 °b), Cleland Cansemvanon Park.

121974,

Renmarks: Paraphauleppia mavazealandica is

distinguishable within the genus by the presence of

foramen. Fl and only three pairs of JZg setae on

the genital shields. The Sauth Australian material

is more like New Zealand type specimens thin rhe

original description indicates, in that the anteriwr

margin of the hysteronotal shield is complete, the

notal setae arc longer and the rostrum has a tooth.

Two small differences, the larger sama and the

absence of a short provrenotal ridge on the

Australian material, however, sugeest chat there any

lie two subspecies present. £2 novazealandica is

superficially simllal to P obtusinoiute sp. nov. bul

is distinguished From it and other South Australian

speci’s, by its generically unique character stares

and ion having a globular sensory seta and no striae

on its Coxites, The most similar species to itis

gracihs CHaminjer. 1958) from Bulivia, which also

shares (he posilioning of hysterunotal seta $5, well

separated from 74, but directly behind it.

Paraphadivppis obfusinotata sp. nav.

PIGS 5,6

type materi: Holowpe 9 (SAMA NI990757),

plint litter, sparse toss and silaceuus sandy soil,

wider selerophyllous sheubs amongst messmate

stringy hark (iucalyplus obliquay, dry sclerophy ll

forest, near snmmit of Mt Lofty (34°59’S,

138°45'E), Cleland Conservation Park, 94,1974,

Paratypes, 105 9 O (SAMA, NI9M)758 NIY90773

and 1990785 — NI990858; 5-BMNH; 3-FMNH;

S-NZAC) ald Ol oo (SAMA NIYSh774 -

NIYOOTRE and NIQYORS9 N1I99Q933, 5 BMH,

5-FMNIIL, S-NZAC), same data as lolompe

Yentule: Soma with owal shape and light brown

colour, Idiosomal length, 308 425, 278-329), Lee

fength (ferourtarsus for 288): 1-155, 11-149, HT-142,

[V-168. Tibial maximum heights (for 288): 1-19,

L)-14, Hi-i4, FV-12.

Proteronatum with terminal tooth to: rosteum

forming broad. point, as broad as long. Antenor

foramen (F1) absent. Seta 2 length subequal to

distance /2-al. Sensory seta (72) clavale, mediua

sive with caput subequal in length to exposed stalk,

Hysteronotum with medium length setae, some

peripheral setae sttehtly Joager, 21 shorter than 22,

AM. BIRCHRY

S6 subequal in length 19 74. Seta SS posterior to

seta 74 so foramen JY justanterior ta line between

serge 24-55.

lfusternal setae with wconspicuous cilia, long,

seta /2 reavhing anterior margin of sternum. Coxite

renon with siritlions broken up into short lines or

dots, from anterior margin to ventrosejugal

apodeme, not superimposed on posteriur

reliculations. Discidial ridge with tubercle at base

of seta TV3, Slit-like pore Su/ nearly transverse, mare

than 45° from longitudinal axis, Begs ellipsoidal,

exochanon smooth, mean size 139 « 70, length

48%) of sonral length, egus per female - 1 ¢9), 2 (4)

or 3 (2).

Legs Jong (mean femurtarsus leneth: 53% of

somal lewgth) with mextium girch (meat maximum

libial height 34% of mean length). Central pretarsal

claw with 2% deprh of lateral pretarsal claws.

Male: Similar ta female but idjosoma shorter, mean

length, 293 (23, 278-293),

Remarke The specifie name obfusinotata is derived

from the Latin wards for blunt’ and ‘back’ and

reférs 10 the broad rostral tooth and rounded

anterior hysteronotal margin, which distinguishes

il trom the similar Ro gevrinorata sp. mov. Other

distinguishing characters are the longitudinal slit-

like pore Sef and the shorter peripheral hysterunaral

selac: # ohtusinarata ind Pacutiriotata are medium

sized amongst other Australian species which have

either clearly longer or shorter hysteronaral setae

Pamphauloppin triforata spr. Wo),

FIGS 7, &

Type material) Holutype & (SAMA NIS9OL012),

plant liter and calcareous sandy soil, under banksia

shrubs (Benda ornare) amongst sclerophylous

shrubs and sparse brown stringy bark mallee

(Auculyples baxlert) on ridge, tall open-shrubland,

Tamboore (35°S7'S, 140°29'H), aii 1974

Pararypes, 12 9 9 (SAMA SL990/013 — NT9901024)

and eight ac (SAMA N19901025 — N19901()32),

same data as holotype.

Females Suma narrowly aval, straw eolous,

Idiosumal length, 279 (13, 270-288). Leg lengih

(fomur-tarsus for 278}: 1-137, If 127, U1 -tag,

1V-144. ‘Vibial maxinvuni heiehts (for 278); 1-17,

(1-12, [1L0, 1-10,

Proteronotum wath terminal Looth to rostrum

forming broad poitit, as broad as long. Anteriot

foramen (Fl) absent. Seta 2 reaching just beyond

seta 1. Sensory seta (2) clavate, mediyim size, caput

subequal in Jenech ta exposed stalk, Hlysteronotum

with short sctac, 23 and 44 teeth less than distance

PARA PHAULOPPIA, ORIBATULID MITES 107

100um

Figs 7-8. Paraphauloppia triforata sp. nov., 9 7, notum of soma, 8, sternum of soma. Vor sctal notation sce Figs

5 and 6.

from 22 and S5 respectively. Seta S5 almost level

with seta 24, and foramen F4 absent,

Idiosterna! setae with shori, distinct cilia, short

seta /2 not reaching anterior margin of sternum.

Coxite region with striations reaching hack

posteriorly to seta /¥2, anterior striae longitudinal,

whilst posterior to ventrosejugal apodeme siriae

angle (owards genital shield. Discidial ridge straight.

Slit-like pore Saf nearly longitudinal, less than 45°

from longitudinal axis. Egg subellipsoidal,

exochorion smooth, mean size 131 x 6, length

47% of somal length, eggs per female - 1 (4), 2 (3)

or 3 (2).

Legs medium length (mean femur-tarsus length:

46% of somal length) with medium girth (mean

maximum tibia height 37% of mean length).

Central pretarsal claw only slightly greater in depth

than lateral pretarsal claws.

Male: Similar to female but idiosoma shorter, mean

length, 264 (8, 259-270).

Remarks: The specific name tri/ora/a is prefixed by

18 D.C. LER & C. M. BIRCHBY

a derivation of the Latin for ‘bree’ and refers to

presence of only three pairs of hvsterdnotal

foramina, a state unique in Parephauloppia. P

triforata is the smallest member of the genus, and

is pale in colour with relatively short legs.

Acknowledgments

We ate indebted to the Australian Biological

Resources study for funding the salary of C.M.B.

ina grant to D.C.L,, and to Dr Henrick Enghoff

(Zoological Museum, Copenhagen) for making

available types of the type-species, Thanks are also

due to Ms Kirstie Jamieson for the notation and

presentation of the figures and Mrs Debbie Lowery

and Ms Debbie Van Weenen for typing the

Manuscript.

References

BatocH, J. & Bao, P. (1984) Review of the

Oribatuloidea ‘Thor, 1929 (Acari: Oribatei). Acta zool,

Aung. 30, 257-313.

BeRLESE, A. (1895) Acari, Myriopoda et Scorpiones

hucusque in Italia reperta. Padua, 77, 5.

—____ (1908) Elenco di géneri @ specie nuove di Acari.

Redia 5, 1-15.

(1916) Centuri prima di Acari nuovt. /hid, 12,

19-67.

Coetzer, A. (L968) New Ornibatulidae Thor, 1929

(Oribatei, Acari) fram South Africa, new combinations

and a key to the genera of the family. Memis Inst. Invest.

cient. Mocainil,, Sér A, 9, 15-126.

HAMMER, M. (1958) Investigations on the oribatid fauna

of the Andes Mountains, Part I. The Argentine and

Bolivia. Biol, Skr. 10(1), 1-129, 34 pls,

_____ (1961) Investigations on the oribatid fauna of the

Andes Mountains, Part IT. Peru. /Bid. 13(1), 1-157, 43

pis.

_____ (962) Investigations on the oribatid fauna of the

Andes Mountains. Part Il, Chile. /bid. 13(3), 1-37, 11

Is,

on (1967) Investigations on the oribalid fauna of New

Zealand, Part W. fbid. 20(3), 1-70, 29 pls:

Hicoins, H, G. & Woottey, T, A. (1975) New mites

from the Yampa Valley (Acarcina: Cryptostigmata:

Oribatulidae, Passalozetidac). Gt. Basin Nar. 35,

1003-108.

Lee, D. C. (1970) The Rhodacacidse (Acar:

Mesostigmata); Classilication, external morphology and

Wistribution of genera, Rec, S. Aust, Mus. 16(3), 1-219.

(1987) Introductory study of advanced oribate

mites (Avarida: Cryptostigmata: Planofissurae) and a

redescription. of ihe only valid species of

Constrictobuies (Oripodoidea). Rec, §. Aust Mus.

24(1), 45-42.

(1991) 17.19. A modified classification of the

Oripodoidea (Acarida: Cryptostigmuta) based on a

phylogenetic model, pp 80-85 Jn F. Disbabck & V.

Bukva (Eds); “Modern Acarology”, Academic

Publishing and Academia, Prague.

—__— & Eirncusy, C. M, (1991) Cervribatula gen. nov.,

Fovoribatula gen. nov, and Fovoribatulinae sf. nav.

(Acarida; Cryptostigmata: Oribatulidae) from South

Australian sous. Rer, 5, Aust, Mus, 24(2), 71-89.

LUNTON, M. (1985) Cryptostigmata (Arachnida: Acari) -

a concise review, Fauna N.Z, 7, 1-106.

MAHUNKA, S. (1975) Neue und interessante Milben aus

dem Genfer Museum, XIII, Neve Oribatiden-Arten

(Acari) aus Senegal. Bull. lst. Fandurm. Afri. noire

S7(:A2), 288-296,

MARSHALL, V.-G., REEVis, R, M. & Nortos, R. A.

(1987) Catalogue of the Oribatida (Acarida) of

coriunertal United States and Canada, Mem. ent. Suc.

Can. 139, 1-418.

Sen enick, M, (1928) Formenkreis Hornmilben, Oribatei.

Tierwelt Mittcleur, 3(a), \-42.

Tsor, S, (1929) Uber die Phylogenie und Systematik der

Acarina mit Beilragen 7ur ersten

Entwicklungsgeschichte einzelner Gruppen. Nvt, Mag.

Naturvid, 67, 145-210.

Waruwork, J. A. & Weoms, D.C. (1984) Jornadia

larreaé nh. gen. on. sp., a new genus of oribacid mite

(Acari: Cryplostigmata) from the Chikhuahuan desert.

Acarotozia 25, 77-80,

LAKE FROME (SOUTH AUSTRALIA) ABORIGINAL TRAILS

BY J. C. MCENTEE*

Summary

This paper traces a previously unrecorded ceremonial route associated with the Flinders Ranges

Aboriginal groups, and their interaction with the people from the Barrier Ranges area in western

New South Wales. While most of the groups within the study area culturally practised circumcision,

the Wilyakali did not. This cultural difference appeared not to interfere with other activities such as

trade in ochre and grindstones or competitive social gatherings. The routes used for the interaction

have been interpreted in terms of language and landscape recognition. The nature of the group

gatherings has been pieced together with the assistance of the few remaining informants.

KEY WORDS: Lake Frome Plains (South Australia), interaction trails, circumcision, language

Srensivions ef the Roval Seecwery ats.

lash (99D). DES(A). 199-205

LAKE FROME (SOUTH AUSTRALIA) ABORIGINAL TRAILS

by J.C. MCENTEL*

Troprimatur

JOHNA-] LU INAWATANA VIPANA WANDU YUND-ANG-AI\),

Whatever John has pul down in tis paper is very goad. If is all right for anybody to read. | agree with

John's use and menning of the place names whith C have learned from my family,

signed: P v7 sted: nie aad liad

(Pear) McKenyie) 14.x1,90)

Summary

McEnree, J.C. (1991) Lake Frome (South Australia) Aboriginal trails. Trans. R. Soc. S, Aust W514),

199-205, 29 November, 1991,

This paper traces a previously unrecorded ceremonial route atsaciated with the Flinders Ranges Abiirivinal

groups, and their interaction will the people trom the Barrier Ranges dred in western New South Wiles.

While most of the groups within the sludy area culturally practised circutmeision, the Wilyakali did nen.

This cullural differance appeared nor to interfere with other activities such as trade in ochre and erindstanes

ar competitive social karherings, The routes used tor the interaction have been inlerpreted in terms of language

and landscape recognition. The nature of the group walherings has been picced together with the assistance

ol the few remaining informants,

kre Worbs: Lake Trome Plains (South Australia), interaction trails, circumcision, language

Introduction

The existing word lists for rhe complex phonetic

sysiem of the ABNAMATANA language contained

¢ighty 10 one hundred words prior to the work of

Svhebeck (1974), By the early 1970s it was realized

that language use had declined markedly. In

response Lo a special request from John and Pearl

McKenzie of Hawker, South Ausiralia, « word lis!

approaching three thousand words was compiled

eradually.and a pronunciation guide with selective

vocabulary was published by McEntee (1976).

The link belween general anthropology and

Aboriginal language provides the answers to many

perplexing questions (McEntee unpublished). ln

particular, there are several AVNAMATANA

lanpuage ternis connected with mythical storycelling

and navigation, For example, the ward Y4STA

WANDA- meaning ‘to tell a story or myth’ or fiterally

‘to story tell the land’ is derived trom the words

YATA imcaning ‘ground, country, land’, and

WANDA- meaning ‘to tell a story’, NARRU NARRU

means ‘straight to a place’, ‘as tle crow thes’

YALPA-RI- means ‘10 be all in line’ and YUWAL-ATA-

means to find one’s way across country’,

This paper clarifies these meanings with reference

to ceremonial and social interaction among the

various groups that lived on and around the Lake

Frome Plains (Fig. 1), and to the local importance

of landmarks and stories.

* Enidina Station, via Yurnta S.Aust. S40

Gale (1986) reinforces this view concerning

navigation:

“berigital peaple developed a nunber-af ways of

depicring the countryside and defining routes to be

laken,.. [they] constructed stylised and very practical

maps of their environment? (Gale 1986 p. 41)

Berndt 41987) described mythical ancestral

travelling routes and Neadjuri place names in the

north of South Australia-and McEntee (unpublished

tris,) noted that at various stages, .a number of these

mythical story lines were placed in remarkably

straight lines.

Requests for possible meanings of the Ngadjuri

place pames ip Olary Province, mentioned in

Berndi’s paper (1987), because of the close

relationship with ADNAMATANA, led to the idea

of putting together some of the knowledge given

by word of mouth by John McKenzie who died in

1986, He was a WILARU or fully initiated

ADNAMATANA elder who had told the author

about some of the interaction between the Flinders

Ranges people and those in western New South

Wales,

The generally accepted region where groups who

did, and did not, practise circumcision shows trails

of dnterartipe across the Lake Frome Plains (Figs

, 2; see also Beckett 1978). lh was noted by Dix

(1883) that the Boolcoomata people (ie, the

Wilyakali) called the groups that practised

circumcision Buerndoppa. This term should be

compared with Yadliyawarra — Bardnapa, and

ADNAMATANA - VADN-APA, Since the observation

fon

“Aieaiinaok: Hii!

St Marp2 Psab

Broken Hill

Elavation

Fig. J. Study até to the Flinders Ranges, Olary Uplands

wind Biterter Ranges. LG. — Luke Gregory; £.B. = Lake

Bignebe, LO = Lake Callabouna, LT = Lake

Yorrens, PEA - Port Augusta.

by Dix, there had heen little investigatory work

carried out in the area until thar of Beckett (1967)

whi summarised language and dialect change in

groups located in north west New South Wales.

Beckett alse showed that these groups displayed

variations in the fulfilment of initiation

requirements (Beckett 1967).

Informants and the Trails

In the 1880s John McKenzie’s father Frecl was a

lad working on Wirrealpa Station in the Flinglers

Rahes. 1h tater years he passed on fis knowledge

ta John who over the years refold many of |hese

stories, In an interview in 1984, John desetibed a

meeting and a ‘challenge match’ between

neighbouring groups as far off ay western New

* Lampert, R. (1984) Unpublished tape recardings, held

by collector

' Mountford, C. P. (1937-1541) The Legend of the Native

Cat VERUITTATITA and Gaanna VARADINA. Research

on the Flinders Ranges. Mountlord - Sheard Calteenon,

41 myths, State Library of South Australia. Special

Collection unpublished notebaoks.

JO, MCTNTER

South Wales, which he likened ta Ihe Olympic

Games’, The following is a deseriplion by John af

this challenge matel with waddies, held semen

last ceutury,

“There's been a challenge match, Some af tie

Ahorigines.tram New South Wales, from Milourinka

and Wileannia, had a challense mutch, They made

a sort af challenes that they were past going to walk

to Mr derleand that the New South Hales peaple

wotld beat em.

These New South Wales peaple came cerays co

challenge the Mt Serle trite —, Ldoat Known hew

muny of ten... quite a mob, They meade up their

waddies [WIRRT] and thev had to pe daw to Weeds

Sprite [WIDNAT] in Mt Serle countey where they held

uchallenge match tosee which side would fwinf but

the Mt Serle moh wiin with the waddies they made."

(Lampert Tapes 1984!)

It is possible, even with sketchy information, to

outline the route taken to travel from New South

Wales to Mt Serle. Qe source of material is the

legend of the Goanna VADNA and the Natwe Cit

IDNA (Mountford 1937-1941"), The legend

concerns the illicit relationship between Goanna

aud Native Cat that caused them to flee from

punishment. The story began at Poolamacca in the

Barrier Ranges, north of Broken HIll. Goatina and

Natwe Cat were very skilful at hiding and they

hurried across che Mundi Mundi plain as they were

being pursued by the anery eklers of the group.

Much of the storyline is lost but the trail would

surely inchide known water sources in (his arid

plain, The couple reached the vicinity of Billerow

Watcthole on the Billeroo Creck KUMBILANA

PARI. Prom there the (rivellers. continued to

Coombes Spring YUDE LAWADLU-NA meaning ‘the

bush Scwevola spinescens hollow’ ngar the south»

east shore of Lake Frome MUNDA (McEntee 1978,

1986). Munda means ‘a trap net" in lwo ways - firstly

from the physical shape of the lake and secondly

from the fact that if one walks onto the lake there

is a possibility of breaking through the salt crust

and becoming trapped in the oozy mud. It is unclear

whether the storyline goes across the lake hur in

view of the passibility of beconiing, rapped, it is

likely that the storyline Follows the southern

shoreline (9 VURA-KARRA-NA waterhole in the Big

John Creek (Fig, 2), The next place mentioned in

the legend as Prism Hill VADNA WAIATI-NA

meaning ‘guaina deep" or ‘voanna has due irself’

deeply into the ground", This explains the origin of

the large sand dune on the eastern side af Prism

Hill. The hill is south of Wertaloona WATALU-RA

meaning ‘shadow coming over From there the

storyline enters the Flinders Ranges to The Jolin

waterhole IRRARA-NA. !1 passes near Mt McKinlay

ABORIGINAL TRAILS

ae rene

201

—

SOUTH aUSTRAUA

WALES

Few sain

Weapamitga Hu

§ Ewatere fimir ef sircuntcdaton

EL Niiee Timely eT a 4

wees Dp tiie oe oF arab

Travelling tacts

Mrtnieal teeeatting route

. - sa

2 Sirrerone

f IN

3 = Hiioe Miche !

il Roura 1

az =f af «

ba -7 aaltaarsiae je he someon ¢erd

z Wear Boley bees erica HNN Tr

E _ = mu

§ on

' 1 ite

128430 mail Wrasre hill Gemma! Wares

all wed

& grt?

o 50km ot

i; 3

32”, '30° <

mine

Pal

Fig. 2. Map of the Lake Frome Plains and surrounding ranges showing the vountry occupied by the Aboriginal Groups

mennioned in the text and the ‘ne of sight’ tail. The fine on the map marking che castern limit of circumeision

is taken from Tindale (1974).

WAYANA meaning ‘dodge this place’ or ‘don't go

there’, The story ends near Arcoona Bluff ARKU-

NA meaning ‘ved ochre’ where Goanna and Native

Cat were speared many times at a sandy patch

known as YANGAVUTLYUTIL meaning ‘dusty liver’,

because their livers were pounded to dust by the

elders as punishment. The spearings are an

explanation of the spots on Goannas and Native

Cats, Fram Arcoona Bluff it is not very far to Mr

Serle ATU-WARAPA-NA. [tis likely (hat (he people

from Milparinka, the Malyangapa vroup, travelled

10 Mt Serle via Yandama Creek which enters Lake

Callabonna just north of Lake Frome, Once the

northern extremity of Lake Frome is reached M1

McKinlay would be visible. The people from the

Darhinyg, River probably followed the storyline [rom

Poolamaccva.

John McKenzie told of another ‘challenge match’

with the VAWUNDI people - likely the Wilyakali

group — who travelléd across to Blinman frorn the

country around Bimbowrie: These people certainly

hud ties with Poolamacca, since some years after

European setticment the Bonbowrie people returned

to their former country al Poolamacca (Mawson

& Hossfeld 1926). John Mekenzie did net

remember much detail abour the VAWUNDI people,

only that they did not practise circumneision, The

ADNAMATANA term for such groups was

YADLARRA, It is of interest to compare this ward

with ‘talara’? mentioned by Dix (1883).

The following is a description. of a competition

using a specially curved throwing club known as the

WAVA (fig. 3b). The WAVA possessed # straight

handle and as John described:

“The point of it had a bit af a curve in at, rot

a sharp curve, half straight. It had to be levelled off

Jot MG@ENTEE 402

Fig. 3. Aborainal Throwing Clubs. A - MUKURA, B

WAV. Scale = 30 ems, Photogcaph conrlesy F, Teague,

al the point. hdd ta pill o Cérigin Point omit!

{Lampert Tapes 1984 )

There was a second straight piece of green wood,

the name of which had been forgotten. This was

laid Mat on the ground; as opposed to green brush

ot @ bush which was used in another conipelition

where the (hrowing suck was named KURURA (Fig,

3a), The nen were divided into two teams and took

icin turns to throw the WAVA so thar the point hit

und then ricocheted off the wood lying on the

ground until it came to rest with the point of the

club facing away from the thrower, I! the WAVA

landed and came to rest at right wngles to the

thrower, there was no score, Likewise a foul throw

was nol counted and that particular competitor

could not pick up the WAVA for a second try, The

competitors would stand in a line, one belind the

other, but alternatively, onc team member and then

the other team member. Two mentbers of rhe same

team could nov stand logether. There would be two

men (rom each team standing at the place where

the WAVA was going to land, to keep the scare, Ef

for cXample two mates from one leam made a

mistake like throwing consecutively, Lhe scorekeeper

would take away wwo points, That would conteavene

the rules.

Jolin McKenzie gaye the following account ofa

WAVA competition as told to him by his father;

From Plamaka [Poolamacca} there way a tot af

dhorigines . .- thatis like the New Sowth Wales mob.

Somehow or other they sent a message like tl might

he specially two young fellaws that ga. They would

Just walk... gete bivafimeker to live on in benveen,

lo bring We message across, The New South Wales

mah sent-a wessaye to Bliniman, that they [the

(MW) have du coare across, Well they came from

Plamuka to Wirreatpa, to Glinman. They played

there... Oh played there for about a week and won

the vane. A the Bliamoe people hed 1a do was to

pay them with rations. .@ bi Of Hour, fed, suman

witchern\ arubs aad cooked meat ,

(Lambert Tapes (984°)

eureror willaba

Before returning lo New South Wales they:

“Set dhewr arranging the next challenye. Lhe lor fran

Wirrealpu and Blininan had to gu across 10 Plamaka

hecause the others had come across to Rhinman. Ther

sent a message to say there wold be another game

af ea ky Ther way anew horse and care asa prive

at Plamaka, That was the challenge «if the Bliamian

moh could win the game they eauld take RAMe the

horse and cart” [which in fact they did]

(Lampert Tapes 1984!)

The possible route from Blinman to the

Plumbago Hills and thevce to meet the people al

Poolamacca is gontained ina fragmentary legend

remembered by Pearl and Myra MeKensie, and

recorded again by Tunbridge (988). A family

travelled from Bhoman to Wirrealpa WIRA-WALPA

meaning River Red Gum (McEntee 1984),

buenivptus camaldilensis —‘dried vegetable maticr'

(Fig. 2). From Wirrealpa the line of travel led to

Limestone Spring VANDATA-NA meaning ‘hard,

precipital J limestone’, then through Nob Gap

VILI-MALKA meaning ‘ventral scales of a snake,

marks’, and then to Tooths Nob Ruins close to

Reaphook Ed! VULIWARU NA mening ‘ventral

scales of a snake, lacing” This isin reference torhe

peak of Reaphook Hill which resembles the head

and part of the underbelly of a Yellow-laced

Whipsnake Demwnsia psanmmephis or WIPARL.

Wiperu also happens to be the Neadjun name fn

Reaphook Hill (Berndt 1987).

From Reaphook Hill the track Jed to the vicinity

of Erudina Station WIRAVUDNU) meaning ‘{hickel

ol E. cumatdulensis! From Erudina the trail ted to

the waterhole at Curnaniona homestead WiIkA-

MALDA niecating vlump of &. camaldilensis;

referring to the group of trees representing the

family waiting at rhe waterhole, which according

to che legend was dry at that time. Ifthe waterhale

at Curnamona happened to be dry, there was a

chance of finding water by deviating a few

kilometres from the straight track 10 a large swanip

known as Katchiwilleroo KAT DAWADLU meaning

the ‘the Emu Bush, Fremophila duttonit hollow’.

This plant grows around the swamp to this day. This

swamp is capable of holding (resh water for twelve

months alter good rainfall. However, the legend

describes the husbitnd Icaving his family at

Curngmona and heading for Batata Spring

VARARINA meaning ‘playing 4 joke’, where he had

to catch a kangaroo to skin lo make a waterbag for

his perishing, family who lad ‘travelled in’ the

meantime to Toweroo Spring (Aboriginal worl

unknown).at Spring Hill. The lewend ends with the

ABORIGINAL. TRAILS a4

husband eventually catching up with his family and

Irving in vain to save rhem. In times of reasonable

raintall, a more direct route could be taken from

Curnamona homestead to Toweroo Spring ar Spring

Hill. From this place the track could lead ro ia

number of locations On Plumbago, Bimbowne and

Outalpa seations. Billeroo Hill is yinble from Spring

Hill and shows (he way into the granite country

where there would be assured water supplies,

However the next feature in travelling the straight

track from Spring Hall is Windamerta WINDA-

MALTA meaning ‘Teo elha, the Barn Owl, thick or

stout’ (McEntee 1986). The hill vertainly looks like

the forepart of an owl especially when viewed from

the norrh west. It is Uncertain from Berndt’s notes

whether Windamerta and Windagudna are separate

places but Windasudna could possibly refer to the

Camel Hump. WINDAKUDNA means “Bari Owl

droppings’, On top of Camel Hump is a white

ourerop which looks very much as if a legendary

owl left droppings acthat place. The white outecop

is plainly visible from the north west of Windamerta

and an Aborigiial painting sire t§ located at the

south western base of Camel Hump, John

Mckenzie ofien referred to the Barn Qwl as a

special bird.

Figs 2 & 4 show the landmarks standing in line

of sight. By moving just 1 kilometre to the South

west of Curnamona honiestead, Spring Hill and

Windamerta merge. By moving 1 kilometre to the

north east of Curnamona on area of plains country

shows hetween Windamerta and Spring Hill, Ifthe

travellers kept Spring Hill in sight from the

Reaphook and Curnamona, they would have been

assured of arnvying at water supplies on the

otherwise featureless plain, Conversely, by keeping

the Reaphook in sight and the correct shape (

NALKA means shape) the reverse trip could be

neeouitted just as safely, Pearl MeKenzie’s mece,

Myra ulso told of people who would gather at

Willippa and head east. following the low hills to

Spring Fhllto meet up with the VAWUNDI people:

Hardy (1976) mentioned Gearge Dutton who was

horn at Yancannia, but was brought up by his

Malyangapa step-father as betie:

“niore strangely qware of what remained of the ale

landmurks that hued guided his peanle”

(Flardy 1976 p. 202)

Untortunately the interconnecting knowledge has

died with the descendants of the Barrier Ranges

people who ventured to the Flinders Ranges,

Two very similar legends concerning “Moon

Man" are contained in Isaacs (1980), One version

is told by George Dutton (Barkindji Story) and the

other is told by May Wilton (ADNAMATANA

] phook

& a =

¢ “8 =

iS = ;

a 6 6@ 5

ag &§

B 3 uy)

17)

£ =

2 >

5 8

2 S

o = Shy

ee a es

Fig. 4. Silhouettes of the Reaphook, Windamerta, Spring,

Hill and Toweroo Spring. A. I'he Reaphook. B. The

Straight line view from Cumamona Waterhole, C. The

view | kilometre south west of Curnamona. D, Fhe view

| kilomeire north east of Curnamona, Note thal the

precise location at Toweroo Spring in Views C and D

is diffieult 19 determine.

Story). Hereus (1982) recorded an identical southern

Baagandji story. Basically the stories describe how

4 greedy uncle Monn Man with two wives, forbids

his nephew to eat emu meat that had been caught

during a hunt. The nephew decides to get even with

his uncle by tricking the latter into climbiny a tree

having many witchetty grubs. ‘he nephew keeps

blowing on the tree lo make il grow taller. He then

ashs Moon Man to try to touch the sky, As Moon

Man grabs at the sky the nephew quickly makes the

tree shrink, leaving his uncle stranded in the sky.

The nephew then takes off with Moon Man’s wives

for himself.

Conelusions

These stories, travelling routes and the map show

the relationship between country, cravel iand

mitts

communication among Aboriginal people in the

Flinders Ranges, the Olary Upland and the Barrier

Ranges (Fig. 1).

Diseussions over rhe vears with John and Pearl

McKenzie have shown rhat links between the people

of western New South Wales and the people of the

Flinders Ranees were even stronger than the lirks

belween (he latter and the closely related Kuyani

People 1o the west.

Acknowledgements

Since 1972 | have varned out extensive linguistic

survey work among the elders of the

ADNAMATAN.A people of the North Jlinders

Ranges and am much indebted to them for rheir

co-operation. Chief teachers have been May Wilton

and her son-in-law John MeKenaty, both of whom

wre now deceased, und John's wite Pearl of Hawker,

South Australia,

lam indebted to the work of ihe late Ronald

Bernadi and his Ngadjuri informant Barney Waria.

Dr Loutse Hereus has assisted in many ways with

usclul information [rom her greal experience with

Aboriginal languages. The tapes made by Dr

Ronald Lampert and willingly made available, were

of great assistance. Thanks are due to Prof. Isabel

MeBryde for her advice and support. Ralph

Grandison, Ron and Aileen Moorhouse and Fred

Teague and his family have been a source of helpful

discussion over the years, Thanks to Margaret

Nobbs who helped with encouragement and typing,

to Kelly Maurice-lones who drafted the figures and

to Margaret Davies for her very helpful advice.

(ilossary

Stany lelvers in (he transeription used by the author for

ADNAMATANA Words have similar corresponding sounds

in Eaglish. However consonants witha superinseribed dot

are lamino-paltial. Consonants with a subinscribed dot

are retrofiex and consonants with & superimsuribed

eireumfles are lamino-dental, A bar over N denotes the

velar nasal, us ‘ng’ in sing, The four infervocalic rhotics

are Lranscribed. thus: D and R for the flaps, RR for the

trilland R for the ghde. The three vowels A, Land U are

similar ta Utalian in their pronunviatwem, Verbs appear im

‘dictionary forme and therefore end in hyphens.

Nhe words listed are ADNAMATARA Words unless

otherwise stated.

ALISA stone, rovk

ARI ved ochre

INS Nate Cat, most dikely Dasvurrns

seoffrov’ (Gould, 1840)

AL DA Yadlivawarra word Corresponding to

JC. MCENTEE

RLIRLAA

KUDNA

KUMBILANA

NALKA

NARRU NARRU

PARI

VADNA

VAIIN-APA

VANDATA

VAKARI-

VAWUNDE

VIPA

viii

VUT] VUTI

VUDSU

VURA-KARKANA

MATA

MATA, MATABRA

MAL KA

MAIDA

MUNIIA

VANGIA

YATA

YAOLARRA

YALPA-RI-

YUDEI

YUNOU

VUWAL-ATAS

ADNAMATARA ward ALDA

‘Eremophila duttonit F Mueltl.", also

‘E, freelingii F. Muelt’,

name af straight competition

throwing stick (Yadliyawarra 7)

Neadjuci and Yadliyawarra, word

corresponding lo the ADNAMATAR A

word UDNA ‘excrement’

Yadliyawarra place name, wirh

unknown meaning, Part ol Billerou

Creek

shape or form

straight to (4 place), direct, express

Yadlivawarra Malyangapa word

corresponding to AQNAMATANA

word VARI ‘creck’

the sand godnna bMeranus couledil

Cray

first stage initialed sale person

tard, preciphaled hmestone

to play a joke ar prank

appellation for group living around

Bimbowric and castwards

1, ADNAMATANA form of English

wold “paper

2. May also mean ‘thin sheet'as it i4

contained in the word ARRU VIPA

‘cirrus Cloud’, lilerally ‘grey thin shoct!

ventral scales of a snake

dusty, powdered, pulverised

thieket of treés

Bottom John Waterhole in. rhe Biz

John Creek. Possihly derived fram

VURA ‘knee" and Yaulivawarra

KARRA ‘high’ cf, AQNAMATANA

word ARRA ‘hich’

thick, stout, large

plural-persan-wordl attached to

Bounds, ‘rou’, ‘mob’, hence

ADNAMASANA “rovk group! or

‘people of the tavks”

1. the piilga tree 4eacia aneura F,

Mucell, ex Benth.

2. jnurks

1, cheek

2, plural-plant-word aliached io

nouns. “chinip"

1. Lake Brome

2, trap net nse in hunting

liver

earth, ground. COunLry

term far groups nol

circumcision

intransitive verh meaning to be

stacked", lo be all in fine?

the bush Sevevola spinescens RBr,

‘Fan blower’ This word is alse

pronounced ¥LPLI and YUINL

lo pul down, to place

composilé verb meaning 4o Find

Ohe's Woy across country” derived

front PUWALY 'sizam" and probably

VATA: a Stretoh®

TWaCUSH

ABORIGINAT. TRAILS 205

WADLU Yadliyawarra word for ‘hallow’ or = WALPA dried vegetable matter

‘hole’ a5 in waterhole WIDNA reeds

WATATI low down WINDA the Barn Owl, Tv/e alba Scopali

WATALU shadow coming over WIPA ant, /ridomyratex spp.

WANDLU 200d WIPARLT the Whip Snake, Demansia

WANDA- to tell a story psammophis Schlegel

WAVA. name of curved competition WIRA the River Red Gum, Eucalypius

throwing stick cumaldulensis Dehnh. (1823)

WAYA: to dodge, to steer clear, avoid WIRRI waddy, club with knobbed end

WARU facing, facing towards WILARU fully initiated male person

References

Beckett, J. (1967) Marriage, Circumcision and Isaacs, J. (1980) “Australian Dreaming: 40,000 years of

Avoidance among the Maljangaba of North-west. New

South Wales. Mankind 6 (10) 456-464,

(1978) George Dution’s Country: portraits of an

Aboriginal drover Pp 2-31 in “Aboriginal History" 2.

(ANU Press, Canberra).

BeRnot, R. M, (1987) Panaramitiee Mawic. Rec. S. Aust,

Mus, 20, 15-28.

Dix, W. J. L. (1883) Boolcoomata Tribe. Pp, 176 in E,

M. Curr (Ed) “The Australian Race”, 2. Melbourne J.

Ferress, London Trubner & Co.

GALE, F. (1968) Art as a Cartographic Farm. The Globe

(24).

Harpy, B. (1976) “Lament for the Barkindji”, (Rigby,

Adelaide).

Hexcus, 1. A. (1982) The Baayandji Language. Pp

257-262. Lingiustics Series B, (67) (Research School of

Pacific Studies, ANU Canberra).

Aboriginal History”. (Lansdowne Press, Sydney).

McENTFF, J. C. (1976) “Pronunciation Guide ta the

ADNAMATANA Language”. Privately published.

. (1986) “WITIITA-NANALPILA; Plants and Birds

of the North Flinders Ranges and adjacent Plains with

Aboriginal names”, Privately published.

Mawson, D. & Hossrecp, P, (1926) Relics of Aboriginal

Occupation in the Olary-District of South Australia.

Trans. R. Soc. S. Aust, 1: 17-25,

SCHEBECK, B. (1974) ‘Texts on the Social System of

Atynyamatana; with grammatical Notes. Pacilic

linguistics Series 21, (Australian National University

Press, Canberra).

TUNBRIDGE, D. (1988) “Vlinders Ranges Dreaming”.

(Aboriginal Studies Press, Canberra.)

TINDALE, N. B. (1974) “Aboriginal Tribes of Australia"

{Australian National University Press, Canberra,)

A NEW TROMBELLID MITE (ACARINA: TROMBELLIDAE) FROM

SOUTH AUSTRALIA

BY R. V. SOUTHCOTT*

Summary

Austrotrombella leprosa gen. et sp. noy. is described, for the adult and deutonymph, from

south-eastern South Australia; the new genus is monotypic. Revised definitions are given for

Parathrombella Andre, 1958 and Durenia Vercammen-Grandjean, 1955. Two species placed by

Andre in Parathrombella are here reclassified as Durenia vilhenae (André), comb. noy., and

D. gracilipalpe (Andre), comb. nov.

KEY WORDS: Taxonomy. Acarina. Trombellidae. Austrotrombella, South Australia, adult,

deutonymph. Durenia, Parathrombella.

Transactions af the Roval Society af S. Aust. V991), TUS(4), 207-212.

A NEW TROMBELLID MITE (ACARINA: TROMBELLIDAE) FROM

SOUTH AUSTRALIA

by R, V, SouTHCOTT*

Summary

Saurneari, R. V, (1991) A new trombellid mite (Acarina: Trombellidae) From South Ausiralia. Trans: R,

Soc. §. Aust. 145(4), 207-212, 29 November, 1991.

Ausiro(rombella leprosa gen. et sp. nov, js desertbed, for (he adult and deujonymph, tram south-eastern

Sourh Australia; the new genus is monotypic. Revised definitions are given for Purathrombella Andre, 1958

and Durenia Vercammen-Grandjeau, 1955, Two species placed by André in Parathrombella are hove

reclassified as Durenia vilhenae (André), corfnb, nov.,.and D, gracilipalpe (André), comb, nov.

Key Woros: Taxonomy, Acarina, Trombellidac, 4ustrotrombella, South Australia, adult, deutonympl.

Durenia, Parathrombella.

Introduction

The mite family Trombellidae Thor, 1935, as

restricted by Southcott (1986a, 1987), contains three

genera known only as adults: Parathrombella

André, 1958 (Africa), Neanothrothrambidium

Robaux, (968 (South America), and

Maiputrombella Southcott, 1986a (South America).

One genus, Womerslevia Radford, 1946 (Maldive

Islands, southern Asia) is known only from the

larva. For Thombella Berlese, 1887 (Europe, Africa,

Australia) the adult, larval and deutonymphal

instars have been described (sce Southcott (986a,

b). For Dufenia Vercanimen-Grandjean, 1955

(Africa, South America, south-eastern Asia) the

same instars are known (Vercammen-Grandjean,

1955) Robaux, 1968; Vercammen-Grandjean &

Audy, 1959). For Nothrotrombidium Womersley,

1954 {adult) (Europe, south-eastern Asta, North

America) larvae have been described by Feider

(1958) and Southcott (1987),

This paper records the discovery of a new genus

and species of trombellid, for the adulr and

deutonymphal instars, from. south-eastern South

Australia, described here In consequence generic

concepts in the Trombellidae are re-examined, re-

definitions offered for Durenia‘and Parathromibella,

and the reclassification of two species of

Parathrombella as Durenia.

Materisls and Methods

Samples of wet soil With growing grasses and

dicotyledonous herbs were collected from a site

beneath a stand of Leplospernmum lanigerum

(Aiton) Smith near Robe, South Australia, in

March 1990, 'The site, near a Swamvp edge, had been

studied for four years for life history studies of

trombiculid mites, Samples weighing 5-6 kg were

* 2 Taylors Road, Mitelyam, S, Aust. 5062.

placed in six large plastic pols and returned to

Mitcham, S,A,, Where they were placed on a cement

patio away from other vegetation and kept moist,

At interyals oF one month, half the contents of a

pot were spread on a coarse wire mesh in a large

Berlese funnel, and allowed to dry at ambieut

temperatures, The arthropod and other fauna were

extracted live and examined daily. Among these were

three adult and ohe deutonymphal trombellid mite,

of a new genus and species, One adult (the holotype

its life history was made, by confining it in a tube

with some of the site soil; however it died eight days

later, without ovipositing. The other three

specimens were fourid dead in the extraction jars,

despite daily examinations.

The mites were cleared in 50% lactic acid and

mounted in Hoyer's medium (Krantz, 1978). Fig.

| was taken with a Leica M3 camera with bellows

and ring extensions, with a 25 rm lens and flash

Ulumination, All drawings were made with the aid

of a camera lucida, All measurements are in

micrometres (um) unless otherwise specified. Setal

and other terminology follows Southevott (1986a, b,

1987).

Material is deposited in the South Australian

Museum, Adelaide (SAM).

Genus Austratrombella gen, nov,

Definirian: Adult: Dorsum of idiosoma with four

longitudinal columns of almost conliguous

depressions lined with basal plates, heavily

sclerotized and sculptured with irregular pits, from

rounded to oblong and polygonal, cach plate with

a smaller more or less central area of much smaller

pits. Anteromedian plate single, transverse, with two

sensillary pits, well-separated, in its posteniar part,

each bearing one fine sensillary seta, All dorsal

idiosomal plates lack normal selae (scobalae).

20% KR

Crista ubsent. Two sensory setae on each side,

underneath anterior edge of anteromedian plate, the

lateral ones resembling the prodorsal sensillary

setae. In between the dorsal plates the integument

carries simple chitinized structures (‘cupolae’), each

bearing one strong, simple, pointed seta, Ventral

surface with coxae | and tl, and JI. and 1V of each

side fused in a sclerotized plate; coxae bearing

simple, pointed setae, Genital plate triangular, with

several rounded pits al anterior and posterior ends

of each lateral valve; setation of genital plate

reduced. Legs with coarse selae, many robust,

lanceolate. Hysterosoma with plates similar to

dorsal, asetose, with cupolae on intervening

integument.

Eyes apparently 2+2, with outer surfaces

somewhat irregular,

Deutonymph: As for adult, but with genital plate

reduced, each lateral valve with one large rounded

pit at anterior and posterior ends.

Larva: nol known.

Recognition: Austroirambella may be separated

from other adult (and deutonymphal) Trombellidae

by replacing Caption 4 of Southcoit (986a, p. 164)

with the following:

4(3) Four depressions in lateral dorsal idiosomal

column... ,.Parathrombella André, 1958

Five or six depressions in lateral dorsal

idiosomal column, .....--... Lei 4A

4A (4) Five depressions in lateral dorsal idiosomal

column, depressions without setae

Tee bee then Pd Austrotrombella gen. nov.

Six depressions in lateral dorsal idiosomal

column, depressions with normal idiosomal

setae (scobulae). 2.0... ane htt oa

she ods't fetes Trombella Berlese, 1887.

Austrotrombella leprosa gen. et. sp. nav.

FIGS 1, 2, 3A-E, 4A-€

Hololype 2 SAM N1991112 (adult)

Paratypes: SAM N1991113, NI99I1114 (adults)

Description of aduli (Figs 1, 2, JA-E): Colaur of

idiasoma in life reddish brown dorsally, red

ventrally, legs and palpi orange-red. ldiosoma

oblong: in life 1220 long by 970 wide, toral length

to tip of chelicerae 1350; after mounting on slide

idiosoma 1495 long by 1125 wide, total length 1855,

Details not covered in generic definition:

Anteromedian plate transverse, 255 long by 364

wide, wider than two smaller plates immediately

posterior; with a large central pit 28 long by 36 wide,

lateral to Which on éach side a sensilligerous pit

bearing a filiform seta 130 long, centres of

sensilligerous pits 202 apart, Similar seta 145 long

V. SOUTHCOTT

1mm

Fig. |. Austroprombella leprosa gen. ch sp. nov, adult,

holotvpe, live, dorsul view, fram a Kodachrome

originates anterolaterally under anteromedian plate,

and medial to this seta another seta 84 long; each

seta arises from a small recessed pit. A series of

narrow sculptured plates lies laterally along the

idiosoma,

Ordinary setae (scobalae) absent frony all dorsal

and other idiosomal plates, but individual setae set

in inter-plate integument, cach arising from the

summit of astrongly chitimzed bell-shaped ‘cupola’

(see Figs 2, 3A)

Fyes 242, with irregular (ic. non-spherical)

external surfaces, set tn small ocular plate

underlying anteromedian and contiguous

anterolateral dorsal plate; strong, pointed seta 102

long on adjacent cupola medial to cyes.

Ventral surface: coxal plates coarsely pitted,

bearing simple pointed setae. Genital plate 336 long

by 315 wide, each lateral valve beanng three rounded

pits at anterior end and six at posterior end

(paratype AC B1i182 with 7-8 pits at posterior ends),

Row of simple pointed setae, 34-50 long along

medial edge of each valve, and few similar setae

elsewhere On valves (see Fig 3B). No acetabula seen

within genital aperture.

Anal plale roughly triangular, with rounded

NEW TROMBELLID MITE

als j

ah '

n a é .

SS Fig. 2. Austrotrombella leprosa gen, et sp. nov., adult, holorype,

\ dorsal view, entire.

Naty

2M) R. V. SOUTHCOTT

fiv. 3. Auscerotrombelle leprose ven. et yp. nov., adult,

holotype. A. Cupola, bearing seta (‘c’ in Fig. 1).

6 External genitaha. C Tip of chelicerae, dorsal. D Palp

tibia and tarsus, dorsal. E Same, yentral. (Each to

nearest scale).

angles; posterior angle obtuse; length 140, width

235; with pitting as for coxal and hysterosomal

plates.

Hysterosoma with symmetrical plates similar to

dorsal idiosomal plates, not seen clearly owing ta

thickness of preparations, but similar to those of

deutonymph, inf.

Legs robust, lengths (including trochanter to tip

of tarsal claws) [ 1185, If 1255, HI 1260, [V 1695,

Legs weil provided with coarse, pointed selae,

mostly lanceolate to pointed-conical, generally

simple, but some of the conical setae with fringing

setules. Tarsi, particularly 1, with reduced setation,

setae mostly simple, lanceolate, pointed; few setac

(also on tibiae) slender, Several minute solenoidalae

on genua, tibiae and tarsi, Tarsal claws simple,

falciform_ Leg segmental measurements (genua~

tarsi}.as in Table 1.

Gnathosoma; cheliceral blades 125 long, pointed,

straight, with 12-13 retrorse teeth, more outstanding

peripherally. Palpi stmplitied, particularly tibia and

tarsus; general setation similar to that of legs, much

reduced on tibia and almost absent on tarsus; tibial

claw strong, blunted,

Description of deutonymph (fram paratype

N1991115); (Fig. 44-C).

Colour and general morphology simular to adult,

but less heavily sclerotized. Idiosoma (mounted) 585

long by 405 wide; total length to tip of chelicerae

700. Anteromedian plate of dorsum 150 long by 270

wide, with its sensillary setae c.60 long; centres of

sensilla 132 apart.

Eyes apparently 2+2, abortive, corneal surfaces

aspherical,

Ventral surface: coxal plates as for adult, with

setation of simple pointed hairs, 27-70 long, as

figured. External genitalia trapezoidal, 104 long by

86 wide where widest, toward posterior end (Fig.

4B); each valve with one large rounded pit at each

(anterior and posterior) end; medial edges of valves

with few simple pointed setae 25-36 long. Anal

plate obscured in preparation. Hysterosoma with

large, pirted plates as figured. Integument between

plates with seta-bearing cupolae as figured.

Legs similar to those of adult; lengths (including

trochanter to tip of tarsal claws) 1 670, 11 480, EIT

665, [V 850. Leg segmental measurements as io

Table |. Lez setation similar to adult.

Gnathosoma: cheliceral blades similar to adult,

with ¢. 13 retrorse tecth. Palpi similar to adult but

Fig. 4. Austrotrombella feprosa gen. et sp. nov,

deutonymph, paratype, A Ventral view, legs. omitted

beyond trochaniers. B External genitalia, C Palp, femur

To tarsus, ventral. (Each ro nearer scale).

NEW TROMBELLID MITE 21

TABLE |. Lex segmental measurements for Austrotrombella leprosa gen. et sp. nov. Data shown as mean | standard

deviation, with the range in parentheses.

Adults

Holotype nh

ACBII32 9

Gel 235 3

Til 277 3

Tal(L) 2358 3

Tal(H) 67 3

Til/Gel 118 3

Gell 242 3

Til 308 3

Talf(L) 283 3

Tall¢H) 62

Till/Gell 1.27 3

Gell 244 3

Till 319 3

Tall (Lh) 288 3

Talll(H) 48 3

Till /Gelll 1.31 3

GelV 382 3

TilV 369 3

Tal V(L) 309 3

Tal V(H) 43 3

TilW/GelV 1.04 3

Deutonymph

Paratype

ACBI216

238.7422.72 125

(218-263)

277.04 25.00 153

(252-302)

259.34+19.04 154

(241-279)

64.347.37 36

(56-70)

1.1634 0.0153 1.22

(1,15-1,18)

249.3425.79 120

(228-278)

312.7 +33,25 156

(282-348

291.04.14,73 168

(282-308)

61.3 = 3.055 av

(58-64)

1.25340.0153 1,30

(1.24-1.27)

249.7 430,89 116

(222-283)

328.3439.83 160

(294-372)

294.3 427,06 165

(271-324)

48.74+7.02 31

(42-56)

1.313+0.00577 1.38

(1.31-1,32)

390.34 56,96 174

(338-451)

387.0+ 51,42 196

(347-445)

314.7422,05 174

(296-339)

48.049.54 uM

(42-59)

1.023 + 0.0208 1.13

(1.00-1,04)

less setose, e.g. palpal tibia with four pointed setae,

palpal tarsus with about five flattened, simple setae.

Material examined: South Australia: Robe district,

Map Reference (Penola 1; 250,000) 283411, in wet,

alkaline, shellgrit — containing soil near swamp edge,

under a stand of Leptospermum lanigerum (Aiton)

Smith, collected 22.ii/.1990, R. V. Southcott, and

successively extracted by drying through Berlese

funnels. Holotype SAM N1991112 (ACB 1132)

obtained on 2.iv.1990 (adult); died 10.iv.1990.

Paratype adults SAM NI1991113, N1991114 (ACB

1152, ACB 1182) obtained dead on 5.v.1990 and

12.vi,1990 respectively, Paratype deutonymph SAM

NI9S9ILIS (ACB 1216) obtained 7,xti.1990.

Etymology: Austrotrombella trom ‘australis

(southern) and ‘Australia’, and Trombella? leprosa

refers to the appearance in life, as though the

dorsum is covered with ulcers,

Remarks an taxonomy: Austrotrombella is very

distinct from its nearest genera (see the rubrics

above), The palp is more obsolete than that in any

other described trombeilid,

Remarks on biology; All specimens came trom wet

soil. Although the extraction jars were examined

daily, only the holotype adult was obtained ative.

Evidently this species is sensitive to desiccation.

Comment on classification of Trombellidue

André (1958) described Parathrambella with type

species P nasuta André, 1958, and (/c.) two other

a2 KV. SOUTEICOTT

species from Angola, 2 vilkenae and P. gracilipalpe.

From Andre's descriptions itis clear that the latter

two species lack the dorsal plates present in B

nasuta, od bear dorsally and anterolaterally on the

idiosomia the two small pits Which Vercammen-

Grandjean (1955) recorded in Durenia Vercammen-

Grandjean, 1955; they also have similar leg structure

to chat of 2. bukavuensis Vercammen-Grandjcan,

1955 (Africa) and D. glandulosa Rabaux, 1968

(South America). These two species of André are

heré reclassified as Durenia vilhenae (André), comb.

noy,, and D. sreci/ipalpe (André), comb. nov.

Ihe following revised generic definitions are

niffered:

Durenid Vercamnmien-Grandjean, 1955

Defiitiion: AdwWt Wromoelhidae. Eyes 242, Dorsum

of idiosoma with two small anterolateral pits, but

livking colunmins of depressions or depilate areas.

larva; Trombellidae. Pyes 2+2, Pedocoxal setal

formula 2, 1, 1. Pedorarsal claws 1, |, 2, Nasus of

dorsal scutum large, triangular, its lateral border

continuous with anterolateral border of seutum,

with ar most only small constriction. Lee tibia 1

without a large solenvidala,

Type species: Durenia bukavuensis Vercammen-

Grandjean, (955,

Remarks: Durenia is at present known for three

African and one South American species in post-

larval instars, and (as a larva only) for B

singdporensis Vercammen-Grandjean & Audy, 1959

(south-eastern Asia). Larva to deutonymph

correlation was established by Vercammen-

Grandjean (1955) with the type species.

Parathrambella Andre, 1958

Definition: Adult: Trombellidae. Eyes 1 +1, Dorsum

of idiosoma with three longitudinal eolumns of

depilate depressions, four in each lateral column.

Larva: Not known,

Remarks: At present restricted to the type species

P. nasuta André, fron: Angola,

Acknowledgments

I thank Messys P. R. and J. Enright, of

‘Hermitage’ property. for access (a the collecting

area, and Dr R. N. McCulloch for field assistance,

Lam indebted to Dr J. B, Jessop, State Herbarium

of South Australia, for plant identifications. The

work was supported by the Australian Biological

Resources Study-

References

Arnone, M. (1958) Aeariens thronbidians (adultes) de

VAngola. PubL cult Diameng No, 4S, 1-125,

Wiki rsh, AL (LRRT) Frombeta glandulasa Berl, n. sp. (et

genus) tyviih plate). Acari, Myriapoda et

Pscodoscorpiones hucusque in fralia reperta, Number

40, Pr. 2 (Vol. IV). (A. Bertese, Patayii)-

Fringe, 7, (958) Prima larva dio familia Trombellidae

(Scarina) objinurs prin culturh gi despre noua

ciracterizare a familiel, lead. R, PR. Fil tape Std,

Cer. Sti Biol, $i. \eric. HI), 265 282.

Krasiz, GW. (1978) “A manual of acarology, 2nd edn,”

(Oregon State University Book Stores, Corvallis,

Grepzon).

Raprorn, CoD. (1946) Larval and nymphal mites

i Acarina; Trombieuhdacy tran. Ceylon and the Maldive

Islands. Parasnoloz® 37(1,2), 46-34.

Rowan. PB. (LM68) Dhrombidiidae d’ Amérique di sud. |

— Vanaupodinae, Johnstomaninae, Thrombellint

(Acanina ~ TVhrombidiidae), Acarologia t0(3), 450-466.

Saytkoory, RV. (t98hay Gn Trombicula alpha n. sp.

{Agarina: Trombellidae) trom Australia: correlarion,

descripnon, developmental abnormialirics, syolematios

und possible auditory structures, Rec. S. Aust, Mus,

MF), 145-168,

—____ (1886) Australian licvae of the genus Trombella

(Acarina: Trombidividea), 4use f Zoul. 34(4), 611-646.

(1987) The classification of the mite families

frombellidae and Johnstonianicae and related groups,

with the descripiion of a new larva (Acarina:

Trombellidae; Nothratrambidium) trom Narth

America. Trans, Ro Sac, S$. Aust V1), 25-42.

Thow, S, (1938) Whersicht und Kintcilung der Familie

trombiudiidae W. 0, Leach 1814in Untertamilien, Zool,

Anz W9(S-6), 107-182.

VERCAMMEN-GRANDIEAN, P. H, (1955) Un weure

nouveau: Duvrenia, dans lta sous-famille des

Trombellinae. Rey, Zoo/, Bor Afr, $2(3 4), 252-260.

2 & Anny, J. R. (959) Une seconde espéece

apparlenant au genre Dyrenia Vercammen 1955 et

originaire de Malaisie: Durenia singaporenstsy 1. sp.

(Acarina; Trombidiidae), Biologisch Jearb. (Dodonaes),

Ghent 27, 98-101,

Womerscey,, H. (1954) On the sablamily. Trombellinue

Sig Thor 1938 (Acarina, Trombidiidae) with the

diagnosis of the nymph of Audyene tharpsunt

Womersley, 1954, Kee. 8. Aust Wis, U0), 121-128.

CRYPTIC SPECIES IN POPULATIONS OF GLOBOCEPHALOIDES

TRIFIDOSPICULARIS KUNG (NEMATODA: TRICHOSTRONGYLOIDEA),

PARASITIC INMACROPODID MARSUPIALS

BY D. L. OBENDORE*, I, BEVERIDGE*, & R, H, ANDREWS}

Summary

The technique of allozyme electrophoresis was applied to populations of the nematode

Globocephaloides trifidospicularis Kung from Macropus giganteus and M. rufogriseus from

Tasmania. Fixed genetic differences were found at four of 24 (17%) loci examined. Because the

nematodes and their hosts are in sympatry, these populations constitute two distinct biological

species. By comparison, both populations of Globocephaloides differed at 58% of loci from the

related genus and species Amphicephaloides thylogale, parasitic in Thylogale billardierii in

Tasmania.

KEY WORDS: Globocephaloides; Amphicephaloides; Macropus; Thylogale; allozyme

electrophoresis; cryptic species

framsacnons uf rhe Hovel Soetery of S Anse (1991), 1U5(S}, 21) 216

CRYPTIC SPECIES IN POPULATIONS OF GLOBOCEPHALOIDES

TRIFIDOSPICULARIS KUNG (NEMATODA; TRICHOSTRONGYLOIDEA),

PARASITIC IN MACROPODID MARSUPIALS

by D. L. OBENDORF*, £. BEVERIDGE**, & RH. ANibREWwSt

Summary

Osenbuer, Do La. Beverincn, 1

& Asnaews, R.A, (1991) Crymic species in populations at

lobocephaluides trifidaspicdlaris Kune (Nematoda: Trichostrongyloidea) parasitic in) mucropadid

marsupials, Trans, R, Soc. §. Aust. 154), 213-216, 29 November, 1991,

The technique of allozyme electrophoresis was applicd to populations of the nematode Glabacephelaides

tfidespicalaris Kung trom Macropus aizenteus and M. ry/ogriseus trom Tasmania. Fixe . genetic differences

were found al four of 24 (178+) loci examined. Because the nematodes and their hosts are 1m syhipatry,

these populations consriture two distinct biological species, By comparison, buth populations af

Glohacephaloides ditlered at 58% of logi fram the related genus and species Amphicenhatoides phytogale,

parasitic in Taydogele billerdierti jn Tasmania.

Key Worps. Glofioeephatoides, Amplicephaloides; Macrapus; Thylogale; allozyme elecinphoresis: cryptic

specics

Introduction

Globovephaloides rrifidaspicidards Kung, 1948 is

a common duodenal parasite of a number of

macropodid marsupial specics from south-eastern

Australia (Reveridge 1979). Because it feeds on (he

blood of the host, this nematode species is capable

of causing mortality in eastern grey kangaroo

(Macropus giganteus) populations, particularly in

Juvenile animals (Arundel e/ a/, 1990), In a

Taxonomic revision of the sub-family

Globocephaloidinae, Beveridge (1979) concluded

that 4 single, morphologically variable species

ovcurred in Macropus eiganteus, M. fuliginasus, M.

riufogriseus, M. eugenii, M, parryi and Wallabia

diewor in the south-eastern region of the continent.

During an investigation into mortalities nM.

gigantexs on Maria Island, Tasmania (42°38'S,

148°05’E), in which G. trifidospiculeris was

involved, it was observed that M. eufogriseus, also

parasitised by Glohocephaiojdes, temsained

unaffected (D. L. Obendorf unpubl. dara). These

observations suggested that the nematodes infecting

the twa macropodid hosts might belong to differenc

species. The present study aimed fo Jes! this

hypothesis using the technique of allozyme

electrophoresis, a powerful means of testing the

existence of sibling species (Richardson ef. al.

1986).

* Jasmantan Department: of Primary Industry, Mount

Measunt Laborstories, Kings Meadows, Tax 7249.

“* Deparlment of Veterinary Science; Universiy of

Melbourne, Parkville, Vic. 3082.

! Fvoluionary Biology Linu, South Australian Museum,

Advluide; current addvess: Department af

Micrabialagy and Immunolopy, University of

Adelaide. Adelaide, S Aust. 5000.

Materials and Methods

Specimens of Globocephatoides were collected

from the duodena of M. gizanteus from Mt

William, Tas. (40°53'S, 148°1S’E), and from M.

rufogriseus from Trevallyn, Tas. (417°27'S,

147°05'E), Nematodes were washed in saline and

frozen in the wells of micro-titre plates ac -80°C

until processing. As an outgroup, specimens of a

Telated penus and species Asnphicephaloaides

thylogale, a duodenal patasite of Thylogale spp.,

were utilised; they were derived from Thylogale

billardierii from the Launceston area, Tas.

Because of heir small size, pools of nematodes

rather than individuals were examined.

Homogenates were prepared by adding an equal

volume af homogenising solution (Richardson ef

al, 1986) to thawed samples, sonicating and

centrifuging at S(W0g for 10 min af 4°C,

Electrophoresis was conducted on cellulose acetate

(Cellogel”,, Chemetron, Milan) according to the

methods of Richardson ef al. (1986).

Forty-five enzymes Were screenéd for suitability

as enzyme markers, Thirty-two enzymes showed

activity following histochemical staining in at least

one sample of elther Globocephulvides or

Amphicephaloides, The enzyme names,

abbreviations and Enzyme Commission codes

(EC.) for these enzymes ure as follows: acid

phosphatase (ACP, E.c, 3.1.3.2.), adenosine

deaminase {ADA, E.C., 3.5.4.4.), alcohol

dehydrogenase (ADH, B.C, [11,1 adenylate

kinase (AR. B.C. 2.7.4.3), aldolase (ALD, EC,

4.12.13). diaphorase (DIA, E.C. 1.8.1.4), enolase

(ENOL, EC. 4.2.1.1]}, esterase (FST, B.C, 3.1.1.1),

furnarate hydratase (KUM, EC, 4,3,1.2), guanine

deaminase (GDA, E.C. 1,4.1,3), glulamate

dehydrogenase (GDH, E,C, 1,4,1,3), aspartate

7 DLL, OHLNDORE, |, BEVERTDGE & R. H. ANDREWS

aminotransferase (GOT, Ec. 2.6.1.1),

glucose-6-phosphate dehydrogenase (G6PD, EC.

1.11.49), glucose phosphate isomerase (GFI, EC.

$3.19), alanine aminotransferase (GPT, EB,

26.1.2), glutathione reductase (GSR, E.C. 1.6.4.7),

hexosaminidase (HEX, E.C. 3.$.1.52), isovitrate

dehydrogenase (IDH, E.C, 11.1.42), leucine

aminopeptidase (LAP, E.C. 3.4.11,1), lactate

dehydrogenase (LDH, E.C 1,1,1,27), malate

dehydrogenase (MDH, E.C. 1.1.1.37), mannose-

phosphate isomerase (MPI, EC. §.3.1.8), purine

nucleoside phosphanlase (NP, E.C. 2.4.2.1),

peptidase valine-leucine (PEP-A, EC, 34.13.19),

peptidase leucine-glycine-glycine (PEP-U, &.C,

3.4.11.4), phosphoglycerate mutase (PGAM, EC,

5.2.4.1), phosphogtyccrate kinase (PGK, EC.

2.7.2.3), phosphoglucomutase (PGM, ES*. 5.2.4.2),

pyruvate kinase (PK, E.C. 2,7.140), triose-

phosphate isomerase (TPE, EC, 3.3.11), URD-

glucose pyrophospharylase (UGPP, E.C, 2,7,7.9)

and xanthine oxidase (NO. B.C. 1.1.3.22),

Representative specimens of the rwo nematode

populations from: Maria 1, were fixed ui 10%

formalin and cleared in lactopheno] for

examination, Measurements wert made with an

ocular micrometer on five male and Tive female

Globovephatoides from the two macropodid hosts.

Representative specimens from both hosts from

various areas of Tasmania have been depusited in

the South Australian Museum: from M7. gizarfters

HO 9YI87, 14572, 16581-3, 16612; from M.

niufperiseus HC 9164, 10742, 14843-4, 10799, 16440.

Results

Sulficent stainig intensity and resolution For

reliable genetic interpretation was observed for 23

enzymes, encoded by a presumptive 24 loci, A

further nine enzymes stained but showed sub-

optimal activity. Of the 24 enzyme loc! scored (Table

1), fixed allelic dilferences occurred between the twa

populations of Globocephaloides at Tour loci

(Ada-2, ldh-2, Got, Pep-BY (17%). Fixed differences

between both samples of Globocephaloides and

Amphicephaloides ocoutred at 14 of the 24 (58%)

loci examined (Table 1),

The following enzyme loci were invariant between

samples: Ald, Dia, Gpt, Mdh, Peam and Tpi.

Detected morphological differences between the

two populations of Globocephaloides were slight

(Table 2), Specimens from M. gigantens were

marginally longer than those trom M. rufogriseus,

and the females were substantially stouter, The

spicules were slightly longer in specimens framy M.

giganteus, bul when expressed as a percentage of

total body length, were similar to specimens from

M. rufogriseus. Female nematodes from MM.

giganteus contained substantially more eggs in the

anterior and posterior uteri when compared with

females from M. rufogriseus, In nematodes from

M, giganteus, there were significantly more eggs in

the anterior interus.

Discussion

Although allozyme analyses were limited to

results obtained from pools of nematodes rather

than individuals (because of their small size), fixed

allelic differences were found at four af 24 (174)

of the lovi examined! for the two populations of

Globocephaloides. Typically, allopatric gene pools

having fixed differences at more than (5%. of

TasLe |. Summnury of allelic differences between pools df Globocephaloides from Macropus gigantens (4) and

Macropus rufogriseus (B/ and pools ef Amphicephaloides thylogale from Thylogale billarcierii,

Enzyme Locus*

Ypooes

Acp Ada-l Ada-2 Ak Enol Est Fum Gda Got Opi Gsr Id Lap Ldh Mpi

Pep-A Pep-B Ugpn

Glodocepheloides A — cd} b af d — 6b

Glokacephaluides Bob be c a bd b b&b

Amphicephaloides a ah a b ac a ab

a ob b a Bb a a/b) a a §

Mutltipte Jaci are designated numerieally according to Increasing tlectrophoreti¢ mability.

t Alleles are designated alphabetically, where a is the mast cathadally migratinu allele,

{ [lelerozygostly is represented by the two homologous alleles, ic ab, be ete,

L Where electrophoretic palterns did not conform: to expectations lor heterozygosity (eg MPL, a Monomeric enzyme

where heterozyzoies should be double-banded) samples were depicted as possessing a mixture (1) of allozymes

(eu forthe monomeric enzyme MPI, the Amphicephaloides sample had a mixture of three allozymes a/b/ch,

_ Staining intensity and resoludion was not sufficient ie allow unequivocal genetic interpretation

CRYPTIC SPECIES IN POPULATIONS OF GLOBOCEFPHALOIDES TRIFIDOSPICULARIS 215

Taatet 2, Measurements fin millimetres! af Globocephaloides trifidospicularis frem Macropus giganieus and M.

tufagriscus /rom Maria ¢, Tas. (news of five measurements tn parentheses).

Total length ¢f

Maximum with >

Spicule length >

Spicule length as Vo of body lengih

Mo. of eggs in anrerior uterus

Naw af evgs in posterior uterus

enzyme loci belong to different biological species

(sce Richardson e7 a! 1986). Thus the data presented

here sugyes! thar che two populations of

Globocephaloides examined represent distinct

species. However, there is reason to believe that the

two jematode (axa are in fact sympatrc, even

though pur samples were not collected at the same

site. The two hosts afe sympatric over muclLof their

geographical range tn Tasmania (Calaby 1983;

Kirsch & Poole 1972}, and in addition the same two

host species on the mainland have a substantial

overlap (749) in diet (Jarman & Phillips 1989),

Since G. rrifidospicularis infects its hast orally

(Beveridge 1979) the two populations of nematodes

in M, gizentevs and M. rufogriseus are essentially

sympatric. The morphological differences noted

between nemaloue specimens occurs irrespective of

the lovality of collection within Tasmania,

suggesting that the segregration is by host rather

than by geography, and that collecting from any

location in Tasmania would yield similar results. In

the case of a sympatry, a single fixed allelic

dilferetice is sulficient to indicate a lack of gene

flow and hence the presence of distinct biological

species (Richatdson er af 1986). Thus, the

demonstration of four fixed allelic differences

between these nematode populations would indicate

that they belong to distinct biological species. The

much greater proportion of fixed differences For

both species of Glohocephaloides and

Amphicephaloides (58%) supports the generic

distinclion made between them at the

morphological level (Beveridge 1979).

The genetic differences berween the two species

of Glaobovephaloides are matched by quite minor

morphological differences (Table 2). The latter

would probably not be considered significant in the

absence of genetic data. The difference in absolute

size in spicule length is ot marked when considered

aS & percentage of total body length, and in the case

of both male and female nematodes, differences in

averall body size may have been discounted as being

M. gigatteus Mi. mifogriseus

92111001

Ih Pel3.4 (11.8)

Q,38-0.51 (U4)

1).54.0.60 (0.56)

4,82-6,49 (5,61)

92-139 (119)

$0-132 (103)

7.6-8.6 (8.2)

99-11, 3 (10.7)

0.23-0,36 (0.30)

0.43-0.47 (0.45)

$,26-5.72 (3.51)

13-61 (43)

14-68 (47)

host-induced. Numbers of eggs in the uteri of

female worms are nor generally considered a reliable

laxonomic character in trichostrongyloid genera,

and a variely of Vactors, including exposure Ww

chemicals (Hotson ef ai, 1970), can influence the

number of cggs present. This feature as well as

dilferences in egg number between anterior and

posterior uteri have, however, been utilised

advantageously for taxqnomic purposes:in the case

of the trichostrongyloid nematodes of monotremes

(Durerte-Desset & Cassone 1983). It appears

therefore that in the case of G. tri/idospicularis,

relatively minor morphological differences may

indicate the existence of sibling species.

The results presented here raise questions as to

the status of records of G. trifidaspiculerts in other

macropodid host species, Unsuccessful altempts (0

infect worm-free M, eugenii with G, trifidaspicularis

derived trom M. givantews (1. Beveridge inpubl.

data), may indicate that the nematode in M, even

is an independent sibling species. The status of

specimens from M, parryi and W ficalor also

remains to be investigated. The type hos! of Ci,

trifidospicularis is M. rufogriseus, the original

description being based on specimens obtained

from wallabies housed at the Regent's Park Zoo

{Kung 1948), Beveridge’s (1979) redescription is

based on material from Af, giganreus from Victoria

and therefore probably represents an undescribed

species, No new names are proposed here, pending

the clarification of the status of G. trifidospicularts-

like nematodes in other matropuodid host species.

{n northern Australia, G. trifiduspicularis is

replaced by two congeners, G. affintiy ogcurring

almost exclusively in the black-striped wallaby,

Macropus dorsalis, and Ci. rracropodis occurring

in a wide range of host species (Beveridge 1979;

Beveridge e7 af. 1984), The results obtained in this

study suggest that a similar analysis al G.

macropadis may be appropriate,

The slight morphological differences observed

here between the populations of Globocephaloides

216 D. L.. OBENDORF, I. BEVERIDGE & R, H, ANDREWS

from M. giganieus and M, rufogriseus, indicate the

care which needs to be taken when morphologically

similar parasites from different hosts are assigned

to the same taxon, The results have significant

practical implications, since in the mortalities

observed in M. giganfeus on Maria I., M.

rufogriseus was not acting as an alternative or

reservoir host for the parasite. Plans to reduce the

densities of kangaroos in order to control the

adverse effects of this nematode parasite probably

do not therefore need to take account of sympatric

M. rufagriseus populations.

Acknowledgments

This study was supported by grants from the

Clive and Vera Ramaciotti Foundation and the

Australian Research Council. We thank the

Evolutionary Biology Unit of the South Australian

Museum for laboratory facilitics, and Drs R. Speare

and D. M. Spratt for comments on the manuscript.

References

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BLAck, R. (1990) Epiderniglogical observations on the

helminth parasiles of Mucropus giganteus Shaw in

Victoria. Aust. Wildl Res. 17, 39-51,

BEVERIDGE, f. (1979) A review of the Globocephaloidinac

Inglis (Nematoda: Amidostomatidae) from macropodid

marsupials. Aust. J. Zool. 27, (51-75.

, SPFARF, R. & JOHNSON, P.M, (1984) New

records of Gtobocephaloidinac (Nematoda:

Trichostrongyloidea) from Macropodidae in north

Queensland. Trans. R, Soc. S. Aust, 108, 197-201.

Carasy, J. H. (1983) “Red-necked wallaby, Mucropus

rujogriseus”? pp. 239-241. In R. Strahan (Ed.)

“Complete Book of Australian Mammals”. (Angus &

Rabertson, Sydney).

Duretrre-Desset, M. C. & Cassone J. (1983) A

taxonomic revision of the trichostrangyloid nematode

parasites of the echidna, Tachyglossus gceuleatus

(Monairemata), Aust. J. Zoo/. 31, 257-284,

Horson, 1. K., Cammpent, N. J. & Smear, M. G-. (1970)

Anthelmintic resistance in Trichostrongylus

colubriformis. Aust, vet, J. 46, 356-360,

JARMAN, PB J. & PHILLIPS, C. M. (1989) Diets in a

community of macropod species. pp. 143-149 Jn G.

Grigg, P. Jarman & I. Hume (Eds). “Kangaroas,

Wallabies and Rat-kangaroas”, (Surrey Beatty & Sons,

Chipping Norton).

Kirescn, JOA. Wo & Pooir, W. E. (1972) Taxonomy ana

Jistribution of the grey kangaroos, Macropus giganteius

Shaw and Macropus fuliginosus (Desmarest) and their

subspecies (Marsupialia: Macropodidae). Austr. J Zool.

20, 315-339,

KUNG, C. C. (1948) Some new nematodes from the

Australian wallaby Macropus rufogriseus Jruticus with

a note on the syuonymy of the geneta Zoniolainus,

Labrostrong lus and Buccostrongylus, J. Helminthol,

22, 93-108.

RicHARbDSON, B. J, Baversrack, P. R, & Abams, M.

(1986) “Allozyme Electrophoresis: A Handbook for

Animal Systematics and Population Studies”.

(Academic Press, Sydney).

A NEW SPECIES OF ANTECHINIELLA QUENTIN & BEVERIDGE, 1986

(NEMATODA: ACUARIIDAE) FROM THE AUSTRALIAN WATER RAT,

HYDROMYS CHRYSOGASTER GEOFFROY, 1804.

BY LESLEY R. SMALES*

Summary

Antechiniella sertatum sp. nov. (Nematoda) is described from the Australian water rat, Hydromys

chrysogaster Geoffroy, 1804. /t differs from its single congener, A. suffodiax, in length of body and

oesophagus, length and disposition of cordons, length of spicule and proportions of the female tail.

The genus is one of only three in the family Acuariidae parasitising mammals. Dietary preferences

and feeding habits of the insectivore, dasyurid, procyonid and rodent hosts of these genera may be

important factors in host distribution and occurrence of infections.

KEY WORDS: Nematoda, Spirurida, Antechiniella, rodent, Hydromys

Vanneerdin i thet Rew Aoeiore af & Aust. (1194)), P1Si44, 217-2200

A NEW SPECIES OF ANTECHINIELLA QUENTIN & BEVERIDGE, 1986

(NEMATODA: ACUARLIIDAE) FROM THE AUSTRALIAN WATER RAT,

HYDROMYS CHRYSOGASTER GEOFFROY, 1804,

by LesLey R, SMALES*

Summary

SMALES, LR, (1991) A new species of Artféchnieffa Quentin & Bevendge, 1986 [Nenapada; Acuarndae)

from the Australian water rat. Hydravns cheysogaster Geoltroy. 1804. Trans, & See S. uss.) 1514),

27 220), 30 November, 141.

Antechiniely seriatum sp. nov, (Namatoda) is described from the Australian waler ral, Aydronys

chrysogaster Geoftroy, 1804, Tt differs from its single congener, ol. su/fudiex, in length af body and

oesophagus, length and disposizion of cordons, length of spieule and propartions of the female ail. The

genus is One at Only three in tbe (ansily Acuariidue parasitisinw mammals, Dielary preferences and fecding

habits of the insectivore, dasyurid, procyonid and rodent hosts of these genera may be importance tyctors

in hose distribution and occurrence of infections

Key Woros- Nematoda, Sparurida, Avifechinitet/a, rodent, Aydromys

Introduction

Although acuariid nematodes oecur principally

in) birds, a few oecur in mammals. These were

reviewed by Quentin & Beveridge (1986). They

concluded, on the basis of cephalic ontogeny that

the three species previously grouped in the genus

Stammerinema Osche, 1955 (Acuariinae) had

different phylogenetic origins. Only the species from

holarctic insecrivores should remain in the

Stummerinema, while the species ftom neotropical

procyonids should be referred to the Skrjahinoclava

Sobolev, 1943 (Acuariinac). The third, from

Australian marsupials and rodents, should be

referred to'a new genus Aritechiniella to be placed

in the Seuratilnae. .4. suffodiax (Beveridgc & Barker,

1975}, first described from the Australian marsupial

Antechinus stuartii Macleay, 1841, was later

reported from Rafius /urreolus and R. fuseipes in

Victoria by Obendor! (1979). This was woted by

Quentin & Beveridge (1986), but they did not

include rodents as hosts in their diagnosis of the

genus, nor did they indicate the reasons for this

omission. In recent collections of parasites from

another rodeni, the walter ral, Hydrovrps

chrysogester Geoffroy, 1804, a second species of

Antechiniella has been Vuund.

Materials and Methods

Nematodes were collecied irom one water mit

from Back Valley, South Austratia in August 1988

and from three water sais from the Inman Rives,

$.A,, In February 1989. Stamachs were digested tn

“0 pepsin (pH = 1) at 40°C for up to 2 hours, and

* Department of Biology, University College of Cenzral

Queensland, Rockhampton, Qld, 4702,

the nematodes were dissected free from surrounding

gastric tissues. Nematodes were fixed in 70%

ethanol, 5% formalin or glacial acetic acid.

Additional material, from S.A, and Victoria, held

in the Australian Helminthological Collection

(AHC), at the South Australian Museum (SAM),

lots 1682, 1698, 1764, 4610, 4611, 5326, 14829, was

also examined. Only two females and five males

remained intact. In addition anterior or posterior

ends of some males and females were preserved.

Accordingly measurements ate from 5-10 male and

2-6 female specimens. Specimens were cleared in

lactophenol. Measurements, in am unless otherwise

stated, with the range followed by the mean, were

made with the aid of an ocular micrometer, drawing.

tube and measuting wheel. L[lustrations were

prepared with the aid of a drawing tube, All

specimens have been deposited in the Sourh

Austrahan Museum,

Antechiniella seriatum sp. nov

PIGS 1-12

étymology: The specific name sertaturm is derived

from the Latin ‘a garland*and refers.to the cordons

around the anterior end.

Descriplion: Seuratiinve. Long worms with anterior

half of body considerably dilated, Cuticle thickened

irregularly on narrowest mid-section of body

adjacent to point of emergence from host stomach

wall, leading to slight widening towards posterior

end. Two large pseudolabia present; cordons

prominent, transversely striated, with deep wide

median groove. Cordons anastomosing, extend

dorsolaterally and ventrolaterally along pseudolabia

and sides of head, extend posteriorly almost to

nerve cord, turn towards one another and fuse,

recurt ta level of distal pharynx. Deirids bicuspid.

L. R. SMALES

21K

qaseanat ANS

a ai MINTS

LPG

My s>

re RUG 77,0

ANTECHINISLLA SERTATUM SP. NOV. FROM MYDROMYS CHRYSOGASTER IN

Male: Length 18-38 (31) min. Maatmum width

340-770 (627), minimum width 160-224 (189)

Corndons extend 100-240 (189) from anterior end.

Pharyox conies!, 64-90 (73) long. Anterior

muscular oesophagus 500: 7904656) und posterior

glandular oesophagus 2880—-4KM1 (3426) long. Nerve

ring around muscular ocsophaeus 220-30) 4273)

from anterioc end, exerclory pore 360-550) (478) and

deirids 248-380) (333) from anterior end. Left

spicule 620-1140 4927) long. flared at proximal end,

trisngular in section, alac not scen. Right spicule

blunt tipped 140-230 (193) long. Four pre-anal and

four post-anal pairs of pedunculated papiltae

present.

Female: Length 3-42 (36)aini, Maximum width

1425-1525 (1475), minimum width 250-275 (263).

Cordons extend 260-380 (320) from anterior end.

Pharynx conical, 60-100 (92) long. Anterinr

muscular pesophiwus 775-1000 (886) and posterior

glandular oesophagus 3250-4525 (3588) long. Nerve

mtg arqund muscular oesophagus 310-360 (337)

fram anterior end, excretory pore 490-720 (605) and

deirids 430-540 (48%) from anterior end. Vulvw

situated 7-13.5 (Simm from posterior end, Vagina

vera Very short about 250, Tall 268-380 (328) lang,

up rounded. Eggs smooth, thick shelled 42.5-47,5

(43.0) » 30.0-32,% (32.0).

Thpe specimens: Holotype SAM Va4lds. Alloeype,

SAM V4149_ Paratypes, AHC 18392,

Tipe fasts Hvdromys chirysegaster Gealfray, 186d,

the water tat.

Tepe facaliry: Inman River, SA.

Site of infeesiin: Stomach.

Discussion

This spevies differs from its congener, 4.

suffodiax, as follows (measurements of males are

given unless otherwise stated); greater length of

body 18.0-38.0 (31.0)mm compared with (7.2-24.6

(18.9)num: longer posterior exten, 100-240 (189} as

compared with 65-180 (100), and disposition of

cordons; and greater length of oesophagus, 500-790

(656) for the muscular portion and 2880-4000

(3426) for the glandular portion as compared with

34-860 (470) and 1200-2500 (200) respectively;

longer lef spicule, 620-1140 (927) as compared with

560-860 (700); longer tail of female 268-380 (328),

as compared with 170-200 (180) and longer distance

from vulva Lu poslerior end, 7.0-13.5 (9mm

compared with 5,0-7.2 (5.8)men-

Specimens of Arfechiniellu from water rats, held

inthe AHC, have been variously designated Sninieru

s. 1. sp. Johnson & Mawson 1952) 4. suffodiox and

Amechiniella sp, (Smales et a/. 1990), Same of the

specimens were postcnur ends af worms associated

with fibrous nodules in the stomach wall. Material

thal consisted only of pastentor portidas éf worms

could be idemified positively as 24, sergetun; by the

length of the spicule, proportions of the fernale taal,

and the presence of irregular cuticular thickening

on the mid-body in both sexes, Therefore, all rhe

material from water tats lodged in the AHC Is .4.

serlaswn,

The appearance of the nodules an the stomach

wall Was consistent with gross descriptions of

nodules given by Beveridge & Barker (1975) for 4,

shiffadiax. However the nature of the marcrial, old

and inadequately fixed for histologival sectioning,

precluded a more detailed examination.

Another acuarird Svarhinwntus e@ustratiensis

(Johnston & Mawson, 1942) also pecurs in WV,

vhrysogaster and A. stuertic These small worms can

readily be distinguished from A. seriadven by their

size, the disposition of the cordons and the spirally

coiled posterior extremity of the male SS

australicasis and 4, serfatum have been recorded

from jhe same localities in S.A, and in #.

oh vsegaster as mixcd infections inthe same hnsi.

Tite food preferences of the mammalian hosts of

acuanid nematode specics appear to be yery similar.

The insectivores Antechinus and Sorex will take a

variety of arthropods (Herter 1972; Statham 1982),

and in the case of Sorex, small vertebrares and

camion, A, “invsogester and Procvon cancrivarus,

the crab cating racoon, are opportunistic predators

taking aquatic insects and crustaceans as well as

larger animals and fresh carrion (Watts & Aslin

1981; Poglayen~Nenwall 1972), The latter two hoses

alsa demonstrate remarkable similarities in feeding

behaviour and food preferences. Both take food

from the water and cat it while sitting on the

riverbank. “Coon sign”, piles of crustacean

exoskeleton, lef at the feetling sires of PF

curcrivaris is paralleled oy similar midden heaps

left by H. chrprogaster at its regular feeditig sites.

These feeding patterns appea’ to ensure that

appropriate intermediate hosts form pari of their

dict, and may provide an insight into the selective

distribution and occurrence of host and parasite.

Since a. serfarum is found only in Australian

rodents and A. siffadiex is found in R. hytrenfies

Figs. 112. Anrechinielly sertatum sp.noy, froavstamach of Aydramys chrysagester. 1. Male amerinr end, lateral View,

2, Ceplialic end, enface view. 3. Cephalic cal ublique, mainly dorso-ventral view, +. Transverse section mid body,

irregularly thickened cuticle. 4 Mid bauy region, irregularly thickened cuticle 6, Deine. 7, Right spicule, vemral

view, 8. Left spoule. prosinzal cnc, lateral view, ¥. Female, vulval péeiol, 10, Letl spicule, distal end, lateral view:

1) Mate, posterior end, lateral view. 12. Female prosterine end, taleral view. Scale lings: Fig, |, 0Smms Fins 23,4501,

Shay; Pivs 6,8,L0. 25m; Figs 7,12, 40yin; Fin 4, 200m.

220

and R. fuscipes as well as the marsupial A. stuartii,

the generic diagnosis as given by Quentin &

Beveridge (1986) should be expanded to include

rodent hosts.

L, R. SMALES

Acknowledgments

I would like to thank Drs T. Cribb and I.

Beveridge for supplying material and Mrs P, M.

Thomas for the loan of material from the

Australian Helminthological Collection.

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