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SOUTH

AUSTRALIAN

MUSEUM

±±j/ j/

DESCRIPTION OF TWO NEW LARVAL SMARIDIDAE (ACARINA) FROM

AUSTRALIA

R. V. SOUTHCOTT

Summary

Two new Australian larvae of Smarididae (Acarnia) are described, Smaris arenicola sp. nov. and
Sphaerotarsus monticolus sp. nov., both from South Australia, from soil and litter. Each is the
second species known as larva in its genus; both earlier described species were also Australian. The
term AME is introduced in the coding of the metric data of the larval dorsal scutum.

DESCRIPTION OF TWO NEW LARVAL SMARIDIDAE (ACARINA) FROM AUSTRALIA

R. V. SOUTHCOTT

SOUTHCOTT, R. V. 1997. Description of two new larval Smarididae (Acarina) from Australia.
Records of the South Australian Museum 30(1): 1-12.

Two new Australian larvae of Smarididae (Acarina) are described, Smarts arenicola sp. nov.
and Sphaerotarsus monticolus sp. nov., both from South Australia, from soil and litter. Each is
the second species known as larva in its genus; both earlier described species were also
Australian. The term AME is introduced in the coding of the metric data of the larval dorsal
scutum.

R. V. Southcott, Honorary Research Associate, South Australian Museum, North Terrace,
Adelaide, South Australia 5000. Manuscript received 1 May 1996.

The mite family Smarididae Vitzthum, 1929 is
cosmopolitan, most members being known as
adults. The adults (and deutonymphs) are small
mites, of a generally flattened stature, living as
predators in litter or under the bark of trees. These
mites are mostly known from their adult instars.
The larvae are probably all ectoparasitic on small
arthropods, as is the case with most other
members of the superfamily Erythraeoidea. Thus
one Australian larva, Smarts prominens (Banks,
1916) , has been found as an ectoparasite of small
Psocoptera (Womersley & Southcott 1941). A
taxonomic and historical review of the family was
given by Southcott (1961a). Seven genera are
known as larvae (Southcott 1995). Of these two
are Australian, each with a single described
species; that of Smarts prominens (Banks, 1916)
(by Womersley & Southcott 1941), and of
Sphaerotarsus leptopilus Womersley & Southcott,
1941 (by Southcott 1960). The present paper
describes a further new species in each of these
genera, both from South Australia, and known
only as larvae.

Materials And Methods

Specimens described in this paper were
obtained by Berlese funnel extractions of samples
of soil and litter in South Australia. They were
collected as active larvae, and each was confined
individually in a small tube and various small
arthropods were introduced as potential hosts, in
an effort to elucidate life histories.

The larvae were eventually mounted in Hover's
medium after first being cleared in 50% lactic acid
solution. Microscopy was done with the use of

Leitz Ortholux/Laborlux microscope equipped
with phase contrast and polarizing facilities.
Drawings were made with the use of a drawing
apparatus.

Chaetotaxic and other nomenclature and coding
are as in Southcott (1961b, c, 1963, 1992, 1995).
The code AME is introduced as a measure of the
anterior central emargination of the dorsal scutum.
(In my previous papers I had misinterpreted the
use of this term as ASBM, which was introduced
by Fain & Elsen 1987). AME may be formally
defined as the distance between the anterior end
of the scutum, in the midline, to the central
anterior point of the scutum; hence AME=ASBa -
ASBM (Fain & Elsen).

The terms protorostral, deutorostral and
tritorostral will be used for the anterior setae of
the gnathosoma, following Newell (1957).

All measurements are in micrometres (]im)
unless otherwise stated.

All specimens will be deposited in the South
Australian Museum (SAM).

Systematics

Family SMARIDIDAE Vitzthum

Smarididae Vitzthum, 1929: 69, 70; Womersley
& Southcott 1941: 61; Southcott 1946: 173; 1948:
252; 1960: 149; 1961a: 174; 1961b: 413; 1961c:
133; 1963: 159; 1995: 57; Shiba 1976: 189.

Erythraeidae Womersley, 1934: 218 (part).

Smarisidae Grandjean, 1947: 1.

For other synonymy see Southcott 1961b: 413—
414.

R. V. SOUTHCOTT

Genus Smarts Latreille

Smarts Latreille, 1796: 180; Womcrsley &
Southcott 1941: 63; Southcott 1946:175; 1948:
252; 1960: 155; 1961a: 176-177; 1961b: 438:
1961c: 133; 1963: 163; 1995:63.

Sclerosmaris Grandjean, 1947: 3, 53.

For other synonymy see Southcottl961b: 438-
439.

Key To Larval Smaris

— PDS 27-29 jam long. AL scutalae blunt-
ended. Sternala I thickened, well setulose

S. prominens (Banks, 1916)

— PDS 28 45 |im long. AL scutalae pointed.

Sternala I narrow, lightly setulose

5". arenicola sp. nov.

Smaris arenicola sp. nov.
(Figs 1A-D, 2A,B)

Material examined

Holotype. South Australia: Loveday, 7
8.ix.l985, R. V. Southcott, larva ACA2158 from
soil and litter sample TX244, site Map:
Department of Lands, South Australia , Barmera-
Loveday, 2nd Edition 1:10,000 6929-26, MR
466055 (140°25'03 M E, 34°17 , 25"S), extracted by
Berlese funnel live 6.x. 1985; to Hover's medium
28.1.1991. SAM.

Paratypes. Same data as holotype, 21 larvae
from same site and sample, ACA2150, 21 52—
2156, 2159-2173, extracted live similarly
24.ix.l985-10.x.l985; Hover's medium. SAM.

Diagnosis of larva

Anterior scutalae pointed. Gelli 65-72 um
long. PDS 28^15 um long.

Description of larva (from slide-mounted
holotype, supplemented by paratypes)

Colour in life red. Idiosoma 245 long, 180
wide; total length of animal 330.

Dorsal scutum transverse, oval, with slight
central anterior emargination, minutely porose.
Scutalae slender, pigmented, AL pointed, PL
blunt-ended, with adnatc to slightly outstanding
setules. AL scutalae placed anterolaterally on
scutum; PL placed posterolateral^; all near edge
of scutum. Anterior sensillary setae placed about
midway between levels of AL and PL scutalae
bases; sockets of posterior sensillary setae lightly

protruding, placed at about middle of posterior
border of scutum. Anterior sensillary setae thin,
but thicker than the filamentous posterior
sensillary setae; all with slight setules in distal
two-thirds, longer distally.
Metric data as in Table I.

Dorsal idiosomal setae about 41 in number,
pigmented, blunt-ended, with short, barbed
setules; anterior setae in indistinct rows across
dorsum; posterior setae form a denser group.

Ventral surface of idiosoma: two sternalae
between coxae I, pigmented, pointed, lightly
setulose; sternalae II vestigial, presenting as small
domes c, 5 across, between coxae II; between
coxae III two intercoxalae 28 long, similar to
sternalae I. Behind coxae III c. 1 1 setae similar to
dorsals, 27-36 long, arranged approximately 2, 4,
4, 1, with a denser grouping towards posterior
pole.

Legs fairly short and thick, similar to those of
Smaris prominens (see Womersley & Southcott
1941); lengths (including coxae and claws): I 380,
II 370, III 410. Coxalae and other leg scobalae
pigmented, setulose, pointed. Supracoxala to leg I
a slender, blunted peg, 5 long.

Leg specialized setae: SoGeI.77pd(23),
VsGeI.90pd(4), SoTiI.50d(38), SoTiI.70pd(33),
CpTi.75d(8)+SoTiI.76d(28), SoTil.78d(12),
VsTiI.83pd(6). SoGeII.51pd(15), SoTiIL03d(16),
SoTiII.78d(10). SoGeIII.52d(14), SoTiIII.03d(12).
Tarsus I with SoTaI.37d(45), curved, strong,
pointed; FaTaI..40ad(6); tarsus I also with an
accompanying trichobothriala to the solenoidala,
with several long setules, at .44d, seta 40 long.
Tarsus II with SoTaIL35d(18), slender, nearly
recumbent. FaTall appears as a minute vestigial
structure immediately alongside, i.e. posterior to,
SoTalL All tarsi with a long, slender, blade-like
empodium, 23 -27 long, over-reaching lateral
claws, which are paired on each tarsus, about 18
long, each a pad with a number of long ventral
onychotrichs.

Gnathosoma robust, pyriform, 77 long by 75
wide (combined). Cheliceral digits S-shaped, 16
long, sharp-pointed. Protorostral (galeal) seta
simple, slender, pointed, 14 long. Deutorostral
seta simple, conical, 7 long. Tritorostral setae
simple, slender, pointed, 21 long. Palpal setal
formula 0, 0, 1, 1, 3, 7 (including solenoiodala
and terminala (eupathidala)); all setae (except
palptarsal solenoidala) pointed; femorala, genuala
and tibialae 1 and 2 setulose. Odontus pointed,
hook-like, with two dorsal supplementary spurs
behind tip. Palpal supracoxala dorsal in position,
a clavate rod 5 long.

TWO NEW LARVAL SMARIDIDAE

FIGURE 1. Smarts arenicola sp. nov., larva. A, B holotype; A, Dorsal view, on left legs omitted beyond trochanters;
B, Dorsal idiosomal seta ('b' in A). C, D Paratypes: C, Teratological dorsal scutum of ACA2156; D, teratologics!
dorsal scutum of AC A2 161. (Each to nearest scale.)

R. V. SOUTHCOTT

FIGURE 2. Smaris arenicola sp. nov., larva, holotype. A, Ventral view, on left legs omitted beyond trochanters. B,
Tip of palp, ventral view, further enlarged.

TWO NEW LARVAL SMARIDIDAE

TABLE 1 . Metric data for Smarts arenicola sp. nov. larvae (*for maximum values)

Holotype

range

mean

s.d.

c.v.

Character

43^19

45.5

1.37

3.0

42-53

48.3

2.35

4.9

SBa

19-23

22.1

1.01

4.6

SBp

13-16

15.0

0.921

6.1

6-12

8.81

1.21

13.7

ASBa

15-22

17.8

1.83

10.3

AME

1-3

2.10

0.436

20.8

1SD

14-17

15.0

0.805

5.4

34^39

35.8

1.34

3.7

58-72

63.7

3.44

5.4

AAS

14-16

15.1

0.831

5.5

A-P

15-20

16.9

1.64

9.7

45-64

55.1

3.54

6.4

36^7

42.1

2.83

6.7

ASE

36-44

39.7

1.93

4.9

PSE

60-73

66.2

3.92

5.9

31-38

38^5*

41.0*

1.79*

4.4*

'Oc'

36^2

38.1

1.98

5.2

MDS

32-38

34.4

1.99

5.8

PDS

31-38

38-45*

40.8*

2.05*

5.0*

Gel

60-69

64.7

1.93

3.0

Til

60-69

64.7

2.12

3.3

Tal(L)

64-73

68.5

2.39

3.5

Tal(II)

27-35

32.2

2.56

7.9

Til/Gel

0.97

0.92-1.07

1.00

0.0340

3.4

Gell

55-64

59.3

2.44

4.1

Till

62-68

65.6

1.89

2.9

TaII(L)

47-60

52.8

3.15

6.0

TaII(H)

20-31

26.4

3.06

11.6

Till/Gell

1.06

1.02-1.19

1.11

0.0490

4.4

Gelll

65-72

67.9

1.77

2.6

Tilll

86-93

89.9

2.04

2.3

Talll(L)

49-61

56.3

2.64

4.7

Talll(H)

20-29

25.0

2.29

9.2

Tilll/Gelll

1.37

1.28-1.38

1.32

0.0360

2.7

AW/ISD

3.13

2.75-3.36

3.04

0.187

6.1

ISD/A-P

0.75

0.75-1.07

0.887

0.0913

10.3

AW/A P

2.35

2.15-3.00

2.71

0.260

9.6

StI

18-29

22.4

2.97

13.3

Cxi

40-53

44.0

2.59

5.9

CxII

20-31

23.3

2.51

10.8

CxIII

26-36

32.1

2.78

8.6

Til/AW

1.40

1.32-1.56

1.42

0.0668

4.7

Tilll/AW

1.98

1.83-2.12

1.98

0.0666

3.4

AW/AL

0.84

0.70-1.02

0.829

0.05^9

7.2

AL/AAS

3.50

2.81-4.27

3.65

0.341

9.3

Tilll/Til

1.41

1.28-1.44

1.39

0.0402

2.9

Till/PW

1.32

1.22-1.57

1.36

0.0768

5.6

LAV

0.60

0.47-0.64

0.56

0.0416

7.4

PW/AW

1.06

0.93-1.18

1.06

0.0545

5.1

AL/PL

1.30

1.13-1.52

1.31

0.112

8.5

R. V. SOUTHCOTT

Etymology

The specific epithet arenicola means 'sand-
dwelling' (Latin), a reference to the site of capture.

Biology

All larvae were captured free. In an attempt to
find possible hosts larvae were introduced
individually into small tubes, and various small
arthropods introduced as potential hosts. These
included various insects: small moths, small
hemipterans (aphids, psyllids, a ?mirid), small
Diptera and wingless Psocoptera; also a small
spiderling. However, no attempt at paras itization
by any of the mites was observed.

Genus Sphaerotarsus Womersley

Sphaerotarsus Womersley, 1936: 119;
Womersley & Southcott 1941: 63, 73; Southcott
1960: 149; 1961a: 177; 1961b: 144; 1963: 211;
1995:57,63.

Definition of larva

One eye on each side. Anterior sensillary bases
anterior to AL scutalae bases; posterior sensillary
bases posterior to PL scutalae bases. Palpal tibial
claw (odontus) apically simple or with narrow
terminal split, and an outstanding basal accessory
tooth. Genu I with 3-6 solenoidalae. Tibia I with
four solenoidalae.

Key To Larval Sphaerotarsus

PDSto 127 umlong

.. S. leptopilus Womersley & Southcott, 1941

PDS to 86 um long S. monticolus sp. nov.

Sphaerotarsus monticolus sp. nov.
(Figs. 3A,B, 4A,B)

Material examined

Holotype. South Australia. Mt. Lofty, 5
20.x. 1991, R. V. Southcott, larva ACA2556, in
damp soil and litter in Eucalyptus obliqua forest,
sample TX320, site at Map: Adelaide 6628 111,
1:50,000, MR 897282 (138 41'47" E., 34°58'03 M
S.), sample collected 5.x. 1991, extracted live by
Berlese funnel 20.x. 1991. Cleared with 50% lactic
acid, mounted in Hover's medium.

Paratypes. South Australia. Hope Forest,
ll.ix.1980, RVS, in moss under Eucalyptus
baxteri forest, at Map Reference Milang 1:63,360
617393 (138°36'19 M E., 35°18'06"S.), site TX198,

extracted by Berlese funnel 1 4-23. ix. 1980.
Stirling, at Map Reference Adelaide 1:50,000, 2nd
Edtn., 934258 (138 o 44*0"E., 34°59 , 30"S.),
ll.xi.1984, RVS, sample TX232 of soil, litter and
grass under Eucalyptus hueberiana y extracted by
Berlese funnel 17.xU984.

Diagnosis of larva
PDS to 66 u.m long.

Description of larva (from slide-mounted
holotype)

Colour in life: idiosoma orange, legs pale
orange. Idiosoma 325 long, 220 wide. Total length
to tip of cheliceral digits 410; in life these
dimensions were 300, 200, 380 respectively.

Dorsal scutum trapezoidal, with rounded
anterior end and acute posterior end; scutum
minutely porosc. Scutum has a sclerotized ridge
enclosing the anterior and posterior sensillary
areas, of a tear-drop shape, the point posteriorad,
and within the bases of the four scutalae. Scutalae
pointed, with many fine, pointed setules. Anterior
sensillary setae thicker than the filamentous
posterior sensillary setae; all sensillary setae with
fine setules in distal half of setal shaft.

Metric data as in Table 2.

Eyes 1+1, lateral to scutum, 18 across.

Dorsum of idiosoma with 24 setae, pigmented,
expanding slightly distally, blunt-ended, with
many fine setules; setae arranged approximately
6,4,2,4,2,4,2.

Ventral surface of idiosoma: sternalae I pointed,
setulose; sternalae II absent; anterior to coxae III a
pair of intercoxalae similar to sternalae I, 36 long,
60 apart. Behind coxae III are 10 pointed, setulose
setae, arranged 4, 2, 2, 2.

Legs somewhat longer than idiosoma; lengths
(including coxae and claws): I 400, II 395,, III
495. Leg scobalae pointed, lightly setulose.
Supracoxala to leg I blunted, 4 long.

Leg specialized setae: TbGeI.44ad(25),
TbGeI.44pd(22), SoGeI.60pd(23), SoGeI.67d(16),
SoGeI.76pd(17), VsGeI.90d(3), TbTiI.48pd(28),
TbTiI.53ad(29), TbTiI.54pv(27), TbTiI.54av(29),
SoTiI.61pd(30), SoTiL67d(30), SoTiI.77pd(30),
CpTiI.80ad(9)+ScTiI.82ad(30), SoTiI.86d(25),
VsTiI.95pd(2). VsGeII.88d(l), SoTill. 1 ld(14),
SoTilI.86d(10). SoTiIII.09d(18).

Tarsus I with SoTaI.17ad(20), TbTaI.20pd(22),
CpTaI.40ad(9)+SoTaI.42ad(22), FaTaL58ad(l),
CpTaI.70d(6)+TbTaI.71d(29). Tarsus II with
SoTaII.39d(20). Lateral tarsal claws with a few
slender onychotrichs; empodium slender, over-
reaching lateral claws.

TWO NEW LARVAL SMAR1DIDAE

¥<v

FIGURE 3. Sphaerotarsus monticolus sp. nov., larva, holotype. A, Dorsal view, legs omitted beyond trochanters. B,
Dorsal idiosomal seta (V in A), further enlarged.

R. V. SOUTHCOTT

100

FIGURE 4. Sphaerotarsus monticolus sp. nov., Larva, holotype. A, Ventral view, legs omitted beyond trochanters. B,
Tip of palp, ventral view, farther enlarged.

TWO NEW LARVAL SMARIDIDAE

FIGURE 5. Sphaerotarsus monticolus sp. nov., larva, holotype. Legs I, II, III. (All to scale shown.)

R. V. SOUTHCOTT

TABLE 2. Metric data for Sphaerotarsus monticolus sp. nov. larvae (*for maximum values)

Holotype

range

mean

s.d.

c.v.

Character

36-47

41.7

2.98

7.2

47-66

56.8

6.44

11.3

SBa

6-13

10.6

2.17

20.5

SBp

11-14

12.2

1.40

11.5

15-23

20.1

3.00

14.9

ASBa

9-16

13.4

2.76

20.6

AME

0-1

0.10

0.316

3.2

ISD

30-38

34.5

2.99

8.7

63-78

72.1

5.24

7.3

59-75

67.3

6.38

9.5

AAS

15-22

17.5

2.01

11.5

A-P

14-30

21.7

5.29

24.4

57-78

66.6

7.88

11.8

57-80

67.8

9.26

13.7

ASE

21-29

25.9

2.85

11.0

PSE

49-68

57.5

6.43

11.2

42-66

55-86*

73.5*

10.5*

14.2*

'Oc.'

45-73

58.5

9.92

17.0

MDS

40-73

54.4

11.8

21.8

PDS

55-86*

73.5*

10.5*

14.2*

Gel

72-82

76.5

3.37

4.4

Til

82-97

89.6

4.93

5.5

Tal(L)

52-67

60.4

5.19

8.6

Tal(H)

22-33

27.2

9.15

33.6

Til/Gel

1.25

1.11 1.25

1.16

0.0372

3.2

Gell

56-85

62.0

4.00

6.5

Till

76-86

82.3

4.27

5.2

TaII(L)

51-60

55.7

3.71

6.7

Tall(II)

18-26

22.4

2.41

10.8

Till/Gcll

1.02

1.02 1.40

1.28

0.107

8.3

Gelll

72-90

82.0

6.86

8.4

Till!

124

114-140

127.0

8.56

6.7

Talll(L)

62-75

66.8

4.54

6.8

Talll(H)

17-24

21.5

2.22

10.3

Tilll/Gelll

1.59

1.48-1.74

1.56

0.0707

4.5

AW/ISD

1.23

1.05-1.67

1.24

0.184

14.9

ISD/A-P

1.84

1.20-2.29

1.66

0.340

20.5

AW/A-P

2.26

1.54-3.36

2.04

0.596

29.2

StI

29-35

32.9

1.79

5.4

Cxi

49-62

55.6

4.03

7.3

CxII

34-48

38.0

4.22

11.1

CxIII

33—41

37.6

3.57

9.5

Til/AW

2.12

1.81-2.28

2.16

0.184

8.5

Tilll/AW

2.88

2.66-3.50

3.08

0.242

7.9

AW/AL

0.74

0.51-0.82

0.635

0.0901

14.2

AL/AAS

3.05

2.59-4.59

3.85

0.608

15.8

Tilll/Til

1.36

1.36-1.50

1.44

0.0505

3.5

Till/PW

1.55

1.27-1.61

1.43

0.144

10.0

LAV

1.03

1.03-1.17

1.07

0.0572

5.3

PW/AW

1.26

1.01-1.61

1.40

0.198

14.2

AL/PL

1.02

0.88 1.09

0.988

0.0601

6.1

TWO NEW LARVAL SMARIDIDAE

Gnathosoma comparatively slender; cheliceral
bases 64 long by 50 wide (combined); cheliceral
blades 16 long, slender, curved, pointed.
Protorostral setae simple, pointed, 9 long.
Deutorostral setae small, pointed, c. 5 long.
Tritorostral setae slender, simple, pointed, c. 18
long. Palpi fairly robust, with setal pattern 0, 0, 1,
1, 3, 7 including solenoidala and eupathidala
(terminala). Palpal femorala and genuala blunt-
ended, setulose; palpal tibiala 1 pointed, setulose;
palpal tibialae 2 and 3 simple, pointed. Odontus
simple, with strong ventral, conical, blunted tooth.
Supracoxala not identified.

Etymology

The specific epithet is an adjective, meaning
'mountain dwelling' (Latin).

Biology

Some observations were made on the living

larva ACA2121. On capture it appeared active
and healthy; it was identified in life by being
weighted down on a slide by a small cover glass,
and examined at 400 diameters magnification.
After release it became quite active again. It was
placed in a small dry tube, and droplets of water
given .Various small arthropods recovered from
the berlesate were offered as potential hosts over
several days: a small coccid (Hemiptera), 7-8
Collembola (?Isotomidae) and an immature
psocopteran. However, no attempt at parasitization
was observed during the nine days it survived in
the tube.

Acknowledgments

The work was done with the support of a grant from
the Australian Biological Resources Study.

References

BANKS, N. 1916. Acarians from Australian and
Tasmanian ants and ant-nests. Transactions of the
Royai Society of South Australia 40: 224-240.

FAIN, A., & ELSEN, P. 1987. Observations sur les
larves du genre Leptus Latreille, 1795 (Acari,
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LATREILLE, P. A. 1796 'Precis des Caracteres
generiques des insectes, disposes dans un ordre
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de la R.

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in Southeast Asia 7: 83-229.

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(Acarina). Proceedings of the Linnean Society of New
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264.

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SOUTHCOTT, R. V. 1961b Studies on the systematics
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revision of the genera and subfamilies. Australian
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SOUTHCOTT, R. V. 1961c . Description of two new
Australian Smarididae (Acarina), with remarks on
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12 R. V. SOUTHCOTT

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A NEW SHRIMP OF THE GENUS RHYNCHOCINETES FROM THE
GREAT AUSTRALIAN BIGHT (CRUSTACEA : DECAPODA :

RHYNCHOCINETIDAE)

Junji Okuno

Summary

A new shrimp of the family Rhynchocinetidae, Rhynchocinetes enigma, is described and illustrated
on the basis of two males and an ovigerous female from the Great Australian Bight. This species is
readily distinguished from the other eleven congeners by the absence of an arthrobranch from the
third maxilliped or lack of an arthrobranch on the pereiopods. It represents the fourth
rhynchocinetid species from depths exceeding 100 m.

A NEW SHRIMP OF THE GENUS RHYNCHOCINETES FROM THE GREAT
AUSTRALIAN BIGHT (CRUSTACEA: DECAPODA: RHYNCHOCINETIDAE)

JUNJI OKUNO

OKUNO, J. 1997. A new shrimp of the genus Rhynchocinetes from the Great Australian Bight
(Crustacea: Decapoda: Rhynchocinetidae). Records of the South Australian Museum 30(1):
13-18.

A new shrimp of the family Rhynchocinetidae, Rhynchocinetes enigma, is described and
illustrated on the basis of two males and an ovigerous female from the Great Australian Bight.
This species is readily distinguished from the other eleven congeners by the absence of an
arthrobranch from the third maxilliped, or lack of an arthrobranch on the pereiopods. It
represents the fourth rhynchocinetid species from depths exceeding 1 00 m.

J. Okuno, Natural History Museum and Institute, Chiba, 955-2 Aoba-cho, Chuo-ku, Chiba
260, Japan. Manuscript received 9 September, 1996.

Caridean shrimps belonging to the family
Rhynchocinetidae are distinguished from other
carideans in the combination of having a typically
movable rostrum, fine transverse striae on the
surface of the carapace and abdominal somites,
the first two pairs of pereiopods robust, with
fingers bearing long lateral and terminal spines,
and the second pereiopod with the carpus entire,
not subdivided. To date, this family includes two
genera, Rhynchocinetes H. Milne-Edwards and
Cinetorhynchus Holthuis (Okuno 1996b, in
press).

Through the courtesy of Ms K. Gowlett-Holmes
of the South Australian Museum, Adelaide
(SAM), I was able to examine two male and one
ovigerous female specimens of an unfamiliar
Rhynchocinetes species collected by trawling from
the Great Australian Bight at depths between 1 1 3
and 170 m. These specimens do not possess
arthrobranchs, although the previously described
congeners at least have arthrobranchs on the third
maxilliped and the first pereiopod. Because of
these differences, they are described herein as new
species.

The postorbital carapace is abbreviated as CL,
and the type specimens are deposited in SAM.

Rhynchocinetes enigma sp. nov.
(Figs. 1-3)

Material Examined

Holotype. Male (SAM C5599, 6.8 mm CL),
34°17'S, 132°42'E, Great Australian Bight,

approximately 1 5 km west-south-west of Pearson
Islands, 140-160 m depth, 'F. V. Comet', April
16,1989.

Paratypes. Ovigerous female (SAM C5598, 9.0
mm CL), 33°12'S, 130°53'E, Great Australian
Bight, 250 km south-south-west of Ceduna, 113
m depth, <F. R. V. Soela', August 5, 1981. Male
(SAM C5600, 7.6 mm CL), 33°18'S, 127°38'E,
Great Australian Bight, approximately 115 km
south-west of Eucla, 170 m depth, 'F. V. Comet',
January 17, 1989.

Diagnosis

A species of Rhynchocinetes without
arthrobranchs on maxillipeds and pereiopods.
Dorsal rostral margin armed with three
subterminal teeth. Fourth abdominal somite with
distinct posteroventral tooth on pleuron.
Ambulatory pereiopods with meri bearing three
spines, and dactyli with four accessory claws
posterior to terminal claw. Endopod of male first
pleopod with distinct lobe on outer margin.

Description

Carapace (Fig. 2A) covered with fine transverse
striae, armed with two acute teeth on dorsal
median carina, anterior tooth just behind rostral
articulation, posterior tooth feebly articulated with
carapace; supraorbital spine acute, considerably
longer than spines on dorsal median carina,
directed anteriorly; antennal spine acute,
supported by a feeble carina, directed anteriorly;
pterygostomial spine small, distinct. Rostrum
(Fig. 2A) articulated with carapace, 1.10 - 1.28x
as long as carapace; dorsal margin armed with

J. OKUNO

FIGURE 1 . Rhynchocinetes enigma sp. nov., holotype male (SAM C5599, 6.8 mm CL). Scale bar = 5 mm.

two teeth in the basal half, and with three small
teeth subterminally; ventral margin armed with
eleven teeth, large proximally, elongate,
decreasing distally.

Abdominal somites (Fig. 1) covered with fine
striae; first three somites with the pleuron
marginally rounded; pleura of fourth and fifth
somites with distinct posteroventral tooth; sixth
somite 0.48 0.50x as long as carapace, with
acute posteroventral spine, with strongly hooked
anal spine between bases of uropods, directed
posteriorly. Telson (Fig. 2B) 0.57 - 0.63x as long
as carapace, 1.16 - 1.30x as long as sixth
abdominal somite, armed dorsally with three pairs
of spines; midpoint of posterior margin
triangularly produced, with three pairs of spines,
median pair longest.

Eyes well developed, with large, globular
cornea; stalk much more slender than cornea.

Antennular peduncle (Fig. 2C) reaching
midlength of scaphocerite; stylocerite long,
reaching or slightly shorter than level of distal
margin of antennular distal segment; proximal
segment with distolateral spine reaching distal
margin of antennular median segment and small
proximal lateral tooth, ventrally with acute spine

at mesial margin; thickened part of upper
antennular flagellum slightly overreaching rostral
apex.

Antenna (Fig. 2D) with scaphocerite 0.84 -
0.93x as long as carapace, 4.20x as long as
maximum width, distolateral spine acute,
distinctly overreaching end of lamella; carpocerite
reaching proximal quarter length of scaphocerite;
basicerite with acute ventrolateral spine and with
rounded protrusion just above the spine.

Mandible (Fig. 3A) with robust three-
segmented palp, distal segment rounded, with
dense setae marginally; incisor process well
developed, with six marginal teeth; molar process
obliquely truncate distally, with finely ridged
distal end.

First maxilla (Fig. 3B) with slender palp armed
distally with single long stout spiniform setae;
proximal endite distinct, rounded, marginally with
numerous setae; distal endite armed with ten stout
spines on mesial margin.

Second maxilla (Fig. 3C) with well developed
tapering palp; proximal endite slightly truncate;
distal endite bilobed, upper lobe rather broader
than lower lobe; scaphognathite large,
overreaching level of tip of palp, anterior lobe with

A NEW RIIYNCHOCINETID SHRIMP

FIGURE 2. Rhynchocinetes enigma sp. nov., holotype male (SAM C5599, 6.8 mm CL). A, carapace with rostrum; B,
telson, dorsal aspect; C, right antennular peduncle, dorsal aspect; D, right antenna, ventral aspect; E, right firstpereiopod;
F, right second peretopod; G, right third pereiopod; H, same dactylus; I, endopod of right first pleopod; J, right second
pleopod; K, right uropod, dorsal aspect. Scale bars; A = 2.5 mm; B, C = 1 mm; D, K - 2 mm; E, G = 1 .5 mm; F = 3 mm*
H-J = 0.5mm.

J. OKUNO

TABLE 1 .Rhynchocinetesenigma,T\ew species. Branchial
formula.

Maxillipeds

Pereiopods

III

II III TV V

Pleurobranchs

Arthrobranchs

Podobranchs

Epipods

Exopods

1
1

1
1
1

1
1

1111
1 1 1

feebly quadrate distal end, posterior lobe tapering,
with elongate plumose setae posteriorly, mesial
margin convex.

First maxilliped (Fig. 3D) with endites
distinctly separated, distal endite concave
marginally, distinctly larger than proximal endite
with slightly rounded margin; palp long,
apparently two-segmented; exopod with long
flagellum, caridean lobe distinct.

Second maxilliped (Fig. 3E) with well
developed podobranch; epipod slightly pointed
distally; exopod well developed, tapering; dactylar
segment with truncate distal margin; propodal
segment with external margin rounded, mesial
margin expanded; ischiomeral segment distinct.

Third maxilliped (Fig. 3F) slightly shorter than
tip of scaphocerite; exopod slightly shorter than
distal margin of antepenultimate segment,
tapering, with dense long setae; ultimate segment
with six spines terminally, 0.59 - 0.62x as long as
carapace, 2.00 - 2.24x as long as penultimate
segment; penultimate segment 0.28 - 0.29x as
long as carapace.

Branchial formula as shown in Table L

First pereiopod (Fig. 2E) chelate, moderately
robust, falling slightly short of midlength of
scaphocerite; chela 0.44 - 0.45x as long as
carapace, 2.00 - 2.24x as long as carpus, tips of
both fingers with dark terminal claws; carpus 0.21
- 0.25x as long as carapace, with acute spine at
distal end of dorsal margin; mcrus dorsodistally
acute.

Second pereiopod (Fig. 2F) chelate, much more
slender than first pereiopod, reaching distal sixth
of length of scaphocerite; chela 0.40x as long as
carapace; carpus entire, 0.59 - 0.65x as long as
carapace, 1.50 - 2.03x as long as chela.

Third pereiopod (Fig. 2G) reaching distal end of
scaphocerite; ischium with single spine; merus
0.74 - 0.8 lx as long as carapace, 1.50 - 2.03x as
long as carpus, with three almost equidistant
spines; carpus 0.37 - 0.4 lx as long as carapace,

with two spines on outer surface; propodus 0.69x
as long as carapace, 1.68x as long as carpus, with
about six short spinules on flexor margin; dactylus
(Fig. 2H) with four accessory claws posterior to
terminal largest claw, decreasing in size
proximal ly.

Fourth pereiopod reaching distal fifth of length
of scaphocerite, spinulation resembling that of
third pereiopod; merus 0.67 - 0.72x as long as
carapace, 1.86 -1.94x as long as carpus; carpus
0.34 - 0.38x as long as carapace; propodus 0.69x
as long as carapace, 1.81x as long as carpus.

Fifth pereiopod slightly shorter than midlength
of scaphocerite; spinulation resembling those of
two anterior ambulatory pereiopods; merus 0.56 -
0.62x as long as carapace, 1.57 -1.72x as long as
carpus; carpus 0.32 - 0.37x as long as carapace;
propodus 0.58 - 0.66x as long as carapace, 1.79 -
1.80x as long as carpus.

Endopod of male first pleopod (Fig. 21) with
distal end rounded; well developed appendix
interna at midlength of mesial margin, distal end
of appendix with dense cincinnuli; distinct lobe at
distal third of outer margin of endopod.

Endopod of male second pleopod (Fig. 2J) with
appendices masculina and interna at distal two
fifths of outer margin; appendix masculina broad,
with distal margin rounded, fringed with dense
setae; appendix interna considerably more slender
and shorter than appendix masculina, with dense
cincinnuli at distal end.

Uropodal exopod and endopod (Fig. 2K)
slightly overreaching distal end of telson, exopod
with a fixed and a movable spine at distal fifth of
outer margin, the former considerably shorter than
the latter.

Coloration
Unknown.

Distribution

Known only from the Great Australian Bight.

Discussion

Most rhynchocinetid shrimps are found around
coral or rocky reefs in shallow waters, although
three species, R. australis Hale, 1941, R. balssi
Gordon, 1936 and R. ikatere Yaldwyn, 1971 are
known from deep waters at depths in excess of
100 m (Hale 1941, Yaldwyn 1971, Holthuis
1972). The present new species represents the
fourth rhynchocinetid shrimp obtained from such
deep waters. Okuno (1996a) suggested that the

A NEW RHYNCHOCINETID SHRIMP

FIGURE l.Rhynchocinetes enigma sp. nov., paratype male (SAM C5600, 7.6 mm CL). A, right mandible;B, right first
maxilla; C, right second maxilla; D, right first maxilliped; E, right second maxilliped; F, right third maxilliped; G, right
branchial region. Setae omitted on C, D, E. Scale bars; A-E = 1 mm; F = 2 mm; G = 3 mm.

J. OKUNO

TABLE 2. Specific list of Rhynchocinetes accompanied with number of arthrobranch (max. = maxilliped; per. =

pereiopod).

3rd max.

1 st per.

2nd per.

3rd per.

4th per.

5th per.

R. australis Hale

„

R. balssi Gordon

R. brucei Okuno

—

R. conspiciocellus Okuno et Takeda

R. durbanensis Gordon

R. enigma sp. nov.

R. ikatere Yaldwyn

R. kuiteri Tiefenbacher

R. rathbunae Okuno

R. serratus (H. Milne-Edwards)

R. typus H. Milne-Edwards

R. uritai Kubo

temperate and subtropical Rhynchocinetes species
all have a limited distributional range. Therefore,
R. enigma probably occurs in southern Australian
waters only.

Eleven species belonging to the genus
Rhynchocinetes were previously known (Okuno
1996a). Rhynchocinetes enigma differs distinctly
from the other species by the absence of an
arthrobranch on all the maxillipeds and
pereiopods (Fig. 3G). The other Rhynchocinetes
species always have two small arthrobranchs on
the third maxilliped and a developed arthrobranch
on at least the first pereiopod (Table 2). In both
Rhynchocinetes and Cinetorhynchus, the number
of arthrobranchs varies at the species level (see
Table 2; Okuno 1996b), and except for the number
of arthrobranchs, there is no unique morphological

character that distinguishes the new species from
other Rhynchocinetes species. Therefore, I am
inclined to place the new species in
Rhynchocinetes, rather than establishing a new
genus to accommodate it.

Etymology

The species name enigma refers to the
enigmatical status within the genus, because of
the absence of all arthrobranchs.

Acknowledgments

I am grateful to Ms K. Gowlett- Holmes for sending
me the specimens on loan. I also thank Dr L. B. Holthuis
for critically reading the manuscript and giving me
valuable suggestions.

References

GORDON, I. 1936. On the macruran genus
Rhynchocinetes, with description of a new species.
Proceedings of the Zoological Society of London
1936: 75-88.

HALE, H. M. 1941. Decapod Crustacea. B. A. N. Z.
Antarctic Research Expedition 1929-1931, Reports-
Series B (Zoology and Botany) 4(9): 257-286.

HOLTHUIS, L. B. 1972. The Crustacea Decapoda
Macrura (the Alpheidae excepted) of Easter Island.
Zoologische Mededelingen, Leiden 46: 29 54, 2 pis.

OKUNO, J. 1996a. Rhynchocinetes rathbunae, a new
shrimp from the Hawaiian Islands (Crustacea:
Decapoda: Rhynchocinetidae). Pacific Science 50(3):
309-316.

OKUNO, J. 1996b. Cinetorhynchus manningi, a new
shrimp (Crustacea: Decapoda: Caridea:
Rhynchocinetidae) from the western Atlantic.
Proceedings of the Biological Society of Washington
109(4): 725-730

OKUNO, J. in press. Review on the genus
Cinetorhynchus Holthuis, 1995 from the Indo-West
Pacific (Caridea: Rhynchocinetidae). in 'Le Benthos
des Fonds Meubles des Lagons de Nouvelle-
Caledonie'. Etudes et Theses, Ed. B. Richer de
Forges, ORSTIOM, Paris.

YALDWYN, J. C. 1971. Preliminary descriptions of a
new genus and twelve new species of natant decapod
Crustacea from New Zealand. Records of the
Dominion Museum 7: 85-94.

POMPHORHYNCUS HERONENSIS SP. NOV. (ACANTHOCEPHALA:

POMPHORHYNCHIDAE) FROM LUT JANUS CARPONOTATUS

(LUTJANIDAE) FROM HERON ISLAND, AUSTRALIA

SYLVIE PlCHELIN

Summary

Pomphorhynchus heronensis sp. nov. is described from Lutjanus carponotatus (lutjanidae) from
Heron Island, Australia. The new species can be distinguished easily from all other species of the
genus by the combination of the following characters: a relatively short neck with a symmetrical
bulb (bulb/total neck length ratioo = 1: 0.5), 15-17 longitudinal rows with 13-16 hooks per row of
which the apical hooks (7-8 per row) have roots and are larger than the basal hooks and are
arranged in slightly spiral rows whereas the basal hooks (6-8 per row) have no roots and are
arranged in straight rows, a proboscis receptacle that does not extend beyond the bulb and atypical
cement glands. This is the first species of Pomphorhynchus described from Australia and represents
one of the few species of that genus known from a marine host.

POMPHORHYNCHVS HERONENSIS SP. NOV. (ACANTHOCEPHALA:

POMPHORHYNCHIDAE) FROM LUTJANUS CARPONOTATUS (LUTJANIDAE)

FROM HERON ISLAND, AUSTRALIA

SYLVIE PICHELIN

PICHELIN, S. 1997. Pomphorhynchus heronensis sp. no v. (Acanthocephala:
Pomphorhynchidae) from Lutjanus carponotatus (Lutjanidae) from Heron Island, Australia.
Records of the South Australian Museum 30(1): 19-27.

Pomphorhynchus heronensis sp. nov. is described from Lutjanus carponotatus (Lutjanidae)
from Heron Island, Australia. The new species can be distinguished easily from all other species
of the genus by the combination of the following characters: a relatively short neck with a
symmetrical bulb (bulb/total neck length ratio - 1: 0.5), 15-17 longitudinal rows with 13-16
hooks per row of which the apical hooks (7-8 per row) have roots and are larger than the basal
hooks and are arranged in slightly spiral rows whereas the basal hooks (6-8 per row) have no
roots and are arranged in straight rows, a proboscis receptacle that does not extend beyond the
bulb and atypical cement glands. This is the first species of Pomphorhynchus described from
Australia and represents one of the few species of that genus known from a marine host.

S. Pichelin, South Australian Museum, North Terrace, Adelaide, South Australia 5000.
Manuscript received 29 April 1996.

There are four genera of Pomphorhynchidae:
Pomphorhynchus Monticelli, 1 905, Longicollum
Yamaguti, 1935, Tenuiproboscis Yamaguti, 1935
and Paralongicollum Amin, Bauer and Sidorov,
1991. Of these, only species of Longicollum have
been recorded in Australia (Edmonds 1989,
Roubal 1993). There are about 25 species of
Pomphorhynchus (see Amin 1985, Ortubay et ah
1991, Wang & Guo 1983). This study describes a
new species of Pomphorhynchus from Lutjanus
carponotatus (Lutjanidae) from Heron Island,
Australia.

Materials And Methods

dried, gold sputter-coated and examined under a
scanning electron microscope operating at 5-
lOkV. Some worms were serially sectioned and
stained with Mayer's haematoxylin and eosin.
Drawings were made with the aid of a camera
lucida and added to by hand. Measurements,
presented as the range with the mean followed by
the standard deviation in parenthesis, are given in
micrometres; where measurements are presented
in paired sets separated by a multiplication sign,
the first figure is length, the second width. Sample
size is given after each measurement. Type
material has been deposited in the Australian
Helminthological Collection (AHC), South
Australian Museum.

Twelve Lutjanus carponotatus were collected
using handlines in the Wistari Channel, Heron
Island and dissected for parasites.
Acanthocephalans were removed from the
intestine; host tissue surrounding the neck
including the bulb of the worms was removed
using micropins and forceps. Specimens were
placed in a cavity block in tap water and
refrigerated to encourage the eversion of the
proboscis and then fixed in Berland's fluid and
stored in 70% alcohol. Specimens were stained
with Mayer's haematoxylin and mounted in
Canada balsam. Lactophenol was used to clear
specimens temporarily for description and
measurement. Some worms were critical point

Systematics

Class PALAEACANTHOCEPHALA

Family POMPHORHYNCHIDAE Yamaguti,
1939

Genus Pomphorhynchus Monticelli, 1905

Pomphorhynchus heronensis sp. nov.
(Figs 1-7)

Type host: Lutjanus carponotatus- (Lutjanidae).
Other hosts: two specimens from intestine of 1 of

S. PICHELIN

FIGURE 1 . a, Proboscis showing two distinct hook types,
b, apical hooks, and c, basal hooks on proboscis. Scale
bars: a = 200, b and c = 20.

5 Cheilinus trilobatus (Labridae) from Heron I.,

Qld, Aust.

Type site: Rectum - neck and proboscis of

parasites bulging through rectal wall into body

cavity of host; infections easily diagnosed from

outside gut.

Type locality: Heron I. (23°27' S, 151°55' E),

Qld, Aust.

Material: holotype (male) AHC 30345; paratypes

AHC 27665-27683, 30346.

Infection rate: 9 of 12 infected.

Description

General

Proboscis cone to pear shaped with slight
anterior swelling; 2 distinct types of hooks (Figs
la-c, 2) present. Apical proboscis hooks large
29.5-51.0 (43.2 ± s.d. 6.6) (n=19) (Fig. lb); root
25.3-48.5 (39.6 ± s.d. 4.7) (n=15) with slight
indentation of base; hooks arranged in 15-17

slightly spiralling rows of 7-8 hooks per row.
Basal proboscis hooks small 14.8-23.5 (18.2 ±
s.d. 2.3) (n=10) (Fig. lc), slender with vestigial or
no roots arranged in 15-17 straight rows of 6-8
hooks per row. Neck short (neck length: trunk
length = 1: 0.32-0.61 (1: 0.44 ± s.d. 0.09)
(n=20)), uniformly cylindrical from trunk level
until abruptly swells to form symmetrical bulb
encircling neck; bulb without protrusions or other
irregularities, may be either spherical or slightly
wider than long. Proboscis receptacle 923-1307
(1100 ± s.d. 123) x 161-231 (188 ± s.d. 23)
(n=10) (Fig. 3) double walled, extends from base
of proboscis to base of bulb; does not extend into
trunk. Trunk widest anteriorly, tapering
posteriorly. Lemnisci 2, slender, 980-2674 (1696
± s.d. 702) (n=6) long.

Males

Total length 7780-13500 (10724 ± s.d. 2213)
(n=15) (Fig. 4). Trunk 552O-10000 (7409 ± s.d.
1426) x 1000-2120 (1572 ± s.d. 357) (n=ll) at
widest point. Neck length including bulb 2250-
3440 (2735 ± s.d. 522) (n=6); neck width
excluding bulb 280-620 (392 ± s.d. 100) (n=10);
bulb 950-1900 (1362 ± s.d. 369) (n=6) x 1360-
2500 (1856 ± s.d. 441) (n=5); bulb/total neck
length ratio 1: 0.42-0.59 (1: 0.5) (n=6). Lemnisci
do not extend beyond anteriormost testis. Testes 2,
ovoid, 360-840 (609 ± s.d. 124) x 246 520 (377
± s.d. 90) (n=14), in tandem or diagonal, in
anterior third of trunk. Cement glands number
undetermined, probably 2 (Figs 5a -c); glands
pyriform at level of seminal vesicle but become
tubular and convoluted as they extend to
posteriormost margin of testis. Proximal part of
cement glands, seminal vesicle and Safftigen's
pouch contained within genital sheath (Fig. 5a);
ligament sac envelops convoluted tubular part of
cement glands, sperm duct and testes. Safftigcn's
pouch pyriform, widest part measures 538 (n=l),
thick-walled, partly enveloped by cement glands.
Sperm duct emerges from seminal vesicle, runs
anteriorly to testes; union of sperm duct and testes
not seen. Seminal vesicle small, obscured by
proximal part of cement glands. Bursa with
pointed digitiform rays and 2 concentric rings of
circumbursal receptors (Fig. 6).

Females

Total length 6630-14000 (9593 ± s.d. 2317)
(n=21) (Fig. 7a). Trunk 4300-8540 (6157 ± s.d.
1194) x 1200-2960 (1761 ± s.d. 310) (n=16) at
widest point. Neck length including bulb, 2000
3700 (2703 ± s.d. 445) (n=14); neck width

POMPHORHYNCHUS HEROENENSIS SP. NOV.

Wt/M

^JmT^ ^.

nnnMHnn

FIGURE 2. SEM of proboscis showing apical hook (arrow) and basal hook (arrowhead). Scale bar = 50.

excluding bulb 300-500 (373 ± s.d. 42) (n=14);
bulb 900-1900 (1333 ± s.d. 350) x 1250-2400
(1699 ± s.d. 444) (n=16); bulb/total neck length
1: 0.41-0.59 (1: 0.50) (n=14). Female
reproductive system consists of ovarian balls,
ligament sac, uterine bell, uterus, well formed
vaginal funnel, 2 consecutive vaginal sphincters
and bulb (Fig. 7b); ligament sac broken in all
specimens examined. Eggs ovoid to fusiform with
small polar prolongations of middle membrane
(Fig. 7c), 98 129 (112 ± s.d. 7) x 27-35 (33 ±
s.d. 3) (n=15); eggs may be present in neck and
bulb.

Discussion

The present specimens belong to the genus
Pomphorhynchus according to acanthocephalan
keys by Amin (1987a) and Amin el al. (1991)
because they have a neck with a true bulb
anteriorly and lack the spirally twisted neck which
is characteristic of Longicollum.

The number of rows of hooks and the number
of hooks per row are similar in most species of
Pomphorhynchus including Pomphorhynchus
heronensis sp. nov. However, all species of
Pomphorhynchus except P. heronensis appear to

S. PICHELIN

FIGURE 3 . 1 Iorizontal section through the proboscis recep-
tacle. Scale bar =500.

have a proboscis receptacle that extends well
beyond the bulb and, in many instances, into the
trunk. The atypical shape and number of the
cement glands in P. heronensis (see below) also
distinguishes it from all other pomphorhynchid
species. Further features that can differentiate this
new species from other similar species of
Pomphorhynchus are given below. The new
species can further be differentiated from P.
patagonicus Ortubay, Ubeda, Semenas and
Kennedy, 1991, P. sebastichthydis Yamaguti,
1939 and P. yamagutii Schmidt and Hugghins,
1973 because they possess asymmetrical bulbs
(Ortubay et at 1991, Schmidt & Hugghins 1973,
Yamaguti 1939). Pomphorhynchus yunanensis
Wang, 1981 differs by having 12 rows of hooks
with only 5 6 proboscis hooks per row (Wang

1981). P. lucyi Williams and Rogers, 1984 differs
by having a considerably longer neck (Williams
& Rogers 1984) relative to that of P. heronensis.
Pomphorhynchus cylindericus Wang and Guo,
1983 and P. bosniacus Kiskaroly and Cankovic,
1969 have fewer than 15 rows of proboscis hooks
with 9 or fewer hooks per row (Wang and Guo
1983, Kiskaroly & Cankovic 1969); also the testes
of P. cylindericus are more posterior than those of
P. heronensis. Pomphorhynchus laevis (Zoega in
Muller, 1776) and P. intermedins Engelbrecht,
1957 have the last basal hooks on the proboscis

-peg

FIGURE 4. Male specimen (holotype), bulb slightly col-
lapsed. Scale bar ~ 2000. Legend: 1, lemnisci; t, testis; teg,
tubular part of cement glands; s, Safftigen's pouch; peg,
posterior part of cement glands; b, bursa with bursal rays.

POMPHORHYNCHUS HEROENENSIS SP. NOV.

FIGURE 5. Transverse sections through a male specimen showing: a, the proximal ends of Safftigen's pouch, cement
glands and seminal vesicle; b, the cement glands partially enveloping Safftigen's pouch (160u further anterior from
previous section);c, the widening of Safftigen's pouch (70u from previous section). Fig. 5a-c drawn to same scale. Scale
bar = 500. Legend: s, Safftigen's pouch; peg, posterior part of cement glands; sv, seminal vesicle.

larger than those immediately preceding (Golvan
1969). P. heronensis does not have this feature;
the last basal hook only appears longer when the
other hooks of P. heronensis are not fully exposed.
P. intermedins, like many other species of the
genus, does not appear to possess the two very
distinct sizes of hooks on the proboscis seen in the
new species. Two distinct sizes of hooks can be
seen on the proboscis of P. laevis as shown in

Brown et ah (1986, Fig. 5) but in addition to the
differences mentioned above P. laevis has fewer
hooks per row (13-20 rows with 8-13 hooks)
(Brown et ah 1986) than P. heronensis. Thus, the
new species can be distinguished easily from all
other species of the genus by the combination of
the following characters: a relatively short neck
with a symmetrical bulb (bulb/total neck length
ratio 1:0.5; neck including bulb/trunk length ratio

S. PICHELIN

..■ '

\ v

FIGURE6.Lightphotomicrographofcloacalbursashowingbursal rays (arrow) and two circles of circumbursal receptors
(arrowheads). Scale bar = 250.

1:0.44), 15-17 longitudinal rows with 13-16
hooks per row of which the apical hooks (7-8 per
row) have roots and are larger than the basal
hooks and are arranged in slightly spiral rows
whereas the basal hooks (6-8 per row) have no
roots and are arranged in straight rows, a
proboscis receptacle that does not extend beyond
the bulb and atypical cement glands.

Some females, but not males, of the new species
show deformities similar to those described by
Am in et al. (1991) for specimens of
Paralongicollum nemacheili Amin et al, 1991.
Other than the enlarged dorsal trunk region,
deformed females did not appear to differ from
normal females. Both normal and deformed
females had eggs that were dispersed throughout
the trunk and occasionally into the bulb; no eggs
were present in the deformed part of the gravid
female. The presence of eggs in the bulb seems
uncommon in other pomphorhynchids but has
been reported in P. kashmirensis Kaw, 1941 by

Kaw (1941) and in P. sebastichthydis by
Yamaguti (1939). The presence of eggs may well
be an artefact of the method of fixing or preserving
and is certainly not a useful taxonomic character.

The cement glands of P. heronensis appear
atypical for the family. Normally,
pomphorhynchids are said to have six cement
glands. In this species however, there appear to be
only two very long glands. A study of serial
sections showed that the proximal part of the
glands are pyriform and closely surround
Safftigen's pouch and the seminal vesicle. The
glands then narrow out and run alongside the
sperm duct up to the testes. Monorchic specimens
of P. heronensis (n=2) appear to have the same
arrangement of cement glands as normal males.
Cement glands in species of Pomphorhynchus are
not usually long but there is at least one other
species, Filisoma longcementglandatus Amin and
Nahhas, 1994, that has cement glands that may
reach up to 16.00 mm in length (Amin & Nahhas,

POMPHORHYNCHUS HEROENENSISSV. NOV.

FIGURE 7.a,Femalespecimen, bulbslightlycollapsed,b, uterine bell,c, egg. Scale bars: a=2000,b==500,c = 50. Legend:
I, lemnisci; o, ovarian ball; ub, uterine bell; ut, uterus; v, vaginal runnel; sph, sphincters, bu, bulb.

1994). Why P. heronensis should have cement
glands so different from other pomphorhynchids
is unknown. It is proposed here that the generic
diagnosis be amended to include the variable
nature of the cement glands seen in P. heronensis.
Two circles of circumbursal receptors similar to
those described by Brown (1987) were seen in one
male specimen. Bursal rays were also noted in the
same specimen. Neither circumpenial receptors
nor muscular suckers in the bursa as those

described by Brown (1987) and Doyle and
Gleason (1991) could be seen in the new species.
Although Brown (1987) considered the receptors
may be useful taxonomically, these structures
were not used as such in this paper because they
were difficult to see. Brown (1987, Fig. 3) also
showed the presence of vesicles in the bursa.
However, Doyle and Gleason (1991) reported
muscular suckers on the inner bursal surface and
suggested that the vesicles noted by Brown (1987)

S. PICHELIN

could have been a ballooning out of the muscular
suckers caused by the method of preparation. A
male with an hyperextended bursa would probably
not be able to hold the female in position and thus
fail to copulate.

Many acanthocephalans show size dimorphism;
noteworthy is that both males and females of P,
heronensis appear to be the same length. Also,
there appeared to be an absence of very young
stages. Although some juvenile pomphorhynchids
appear able to detach from the gut, move and
reattach to another region (Amin 1987b) no
pomphorhynchids in the present study were found
other than in the rectum. The lack of juvenile
stages may indicate that hosts are not continuously
infected throughout the year or that there is some
mechanism restricting establishment of new
parasites. Brown (1986) provided evidence to
show that populations of P. laevis could reach a
ceiling level and that parasite establishment could
be density-dependent. However, in some cases it
appears that this ceiling level is not reached
because the critical resource, e.g. space, is only
limited briefly (Bates & Kennedy 1991).

Pomphorhynchids have been recorded from a
wide range of teleosts (see Golvan & de Buron
1988). Amin (1987c) recognised three categories
of hosts for P. bulbocolli Linkins in Van Cleave,
1919: principal, accessory and occasional hosts.
In this study, L. carponotatus is considered as a
principal host for P. heronensis because female
pomphorhynchids with ovarian balls, immature
and mature shelled acanthors were regularly
recovered, all parasites were firmly attached and
none were found extraintestinally. This is
consistent with the findings by Brown et al.
(1986) and Amin (1987c) who considered the
regular presence of mature gravid females as an
indication that the parasite is in its preferred
definitive host. It is also consistent with the
observations made by Kennedy (1984) that P.
laevis in its non-preferred hosts were smaller,
immature, weakly attached and many were
recovered in extraintestinal sites.

Most species of Pomphorhynchus occur in

freshwater fishes. Some exceptions such as
Pomphorhynchus laevis have been recorded from
both freshwater and marine fishes (see, for
example, Kennedy 1984). Although, it has been
suggested that the freshwater P. laevis is a
different strain from the marine P. laevis and that
each strain has a preferred host (see Kennedy
1984); currently there is thought to be three strains
(see Kennedy et al 1989, Kennedy 1996). The
preferred position in the preferred marine host of
P. laevis is the rectum whereas the preferred
position in freshwater host is more anterior (see
Kennedy 1984). All the specimens of P.
heronensis recovered for this study were found in
the rectum of its host, L. carponotatus. Lutjanus
carponotatus is not known from freshwater
habitats (Allen & Talbot 1985) and thus the
posterior position of the parasite cannot be
correlated with a response to changing salinity as
shown by Kennedy (1984) for P. laevis under
certain circumstances.

Specimens of Pomphorhynchus heronensis
were also recovered from one of five Cheilinus
trilobatus in the present study and from one of
five from another study of helminths in fishes of
Heron Island (Cribb pers. comtn.). In these
specimens the parts of the worms that had
penetrated the gut wall (i.e. the neck including the
bulb) were decayed. Had this deterioration
resulted from the worms being reproductively
exhausted followed by death in situ, then worms
in similar condition should have been found in L.
carponotatus; none were. It may be that C.
trilobatus is an unsuitable or at best an occasional
host for P. heronensis but further material is
needed to negate the possibility of other factors
influencing the presence or absence of these
parasites, such as the distribution of intermediate
hosts and seasonality.

Acknowledgments

I am grateful to Dr T. H. Cribb for donating specimens
of this species and reading this manuscript

References

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AMIN, O. M. 1985. Classification. Pp. 27-72 in

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AMIN, O. M. 1987a. Key to the families and subfamilies
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POMPHORHYNCHUS HEROENENSIS SP. NOV.

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AMIN, O. M., BAUER, O. N. & SIDOROV, E. G. 1991.
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BROWN, A. F. 1986. Evidence for density-dependent
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infected Salmo gairdneri Richardson and its bearing
on wild populations in Leuciscus cephalus (L.).
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FIELD IDENTIFICATION OF FEMALE LITTLE BROWN BATS
VESPADELUS SPP. (CHIROPTERA : VESPERTILIONIDAE) IN SOUTH

AUSTRALIA

L. F. QUEALE

Summary

Five morphologically similar species of little brown bats occur in South Australia; Vespadelus
finlaysoni, V. darlingtoni, V. regulus, V. baverstocki and V. vulturnus. As males can be identified
by the shape of their penes, the emphasis of this paper is on methods to differentiate the females of
these similar species. Characters are listed here for identifying female Vespadelus bats, particularly
for use in the field, using external measurements and geographic distribution data. Vespadelus
finlaysoni and V. darlingtoni were found to be generally distinguished on size alone and V.
vulturnus to be distinguished on a wing measurement ratio. Vespadelus regulus and V. baverstocki
presented the greatest difficulty for identification. However this study shows that these two species
can be distinguished 90% of the time. Areas of allopatry, parapatry and sympatry are also discussed.

FIELD IDENTIFICATION OF FEMALE LITTLE BROWN BATS VESPADELUS SPP.
(CHIROPTERA : VESPERTILIONIDAE) IN SOUTH AUSTRALIA

L. F. QUEALE

QUEALE, L. F. 1997. Field Identification of female little brown bats Vespadelus spp.
(Chiroptera: Vespertilionidae) in South Australia. Records of the South Australian Museum
30(1): 29-33.

Five morphologically similar species of little brown bats occur in South Australia; Vespadelus
finlaysoni, V. darlingtoni, V. regulus, V. baverstocki and V. vulturnus. As mates can be
identified by the shape of their penes, the emphasis of this paper is on methods to differentiate
the females of these similar species. Characters are listed here for identifying female Vespadelus
bats, particularly for use in the field, using external measurements and geographic distribution
data. Vespadelus finlaysoni and V. darlingtoni were found to be generally distinguished on size
alone and V. vulturnus to be distinguished on a wing measurement ratio. Vespadelus regulus
and V. baverstocki presented the greatest difficulty for identification. However this study shows
that these two species can be distinguished 90% of the time. Areas of allopatry, parapatry and
sympatry of species are also discussed.

L. F. Queale, South Australian Museum, North Terrace, Adelaide, South Australia 5000.
Manuscript received 20 June 1 996.

The bats once referred to as a single species of
Eptesicus (E. pumilus) are now regarded as nine
species of Vespadelus (Kitchener et al. 1987,
Volleth & Tidemann 1991). This diversity has
only recently been recognised and many of the
species' morphologies are very similar, so that
practical problems in the field of specimen
identification have not yet been completely solved.

The current taxonomy follows Kitchener et al.
(1987) who revised the genus using a combination
of morphological features. Their conclusions were
confirmed by allozyme electrophoresis (Adams et
al. 1987) which showed a minimum of nine
species. Males of the five South Australian
species, Vespadelus finlaysoni, V darlingtoni, V
regulus, V. baverstocki and V. vulturnus, can be
unambiguously identified using penis glans
morphology (Reardon & Flavel 1987) but a
reliable method is not available for female
morphology. Many of the characters used by
Kitchener et al (1987) were internal features and
cannot be used for field identification. Geographic
variation within species further complicates
identification.

The aim of the present study was to provide
simple field methods for identifying female
Vespadelus spp. using external characters.
Outside South Australia other workers have found
that there is more overlap in forearm lengths
between species (Tidemann 1981, Lumsden &
Bennett 1995). I found that limiting this study to

South Australia, the task of identification can be
made easier. These methods will prove useful in
other states, although because of geographical
variation there may be some differences.

Materials And Methods

Initially, 127 adult female Vespadelus bats (9
V. darlingtoni, 11 V. finlaysoni, 50 V. regulus, 38
V. baverstocki and 15 V vulturnus) from the spirit
preserved mammal collection of the South
Australian Museum were examined for this study.
Adults were distinguished by ossification of wing
bone epiphyses. Specimens in the collection were
collected from widely separated localities to take
account of regional variation within South
Australia. Based on the results of previous
investigations (Carpenter et al. 1978, McKean et
al 1978, Tidemann et al. 1981, Kitchener et al.
1987) and the ease of measurement and
assessment, the following characters were selected
for examination:

1 ) forearm (= radius) length

2) ratio of phalanx I to metacarpal length on
digit 111

3) geographic distribution

4) colour of skin and fur

Wing bone length measurements (to the nearest
0.1 mm) were taken, using dial calipers, from
joint to joint (outside) when the joints were bent
(see Reardon and Flavel 1987).

L, F. QUEALE

Distribution maps were compiled for each
species using females that had been reliably
identified by electrophoresis (n=45) and males
identified by glans penes (n=133), following the
method of Reardon and Flavel (1987).

General colour of skin and fur were also noted
to aid identification.

To test the reliability of the characters described
above, an additional 40 bats were analysed using
electrophoresis of liver enzymes. The
discriminating enzymes used were: aspartate
aminotransferase, carbonate dehydratase, p-
hosphogluconatc dehydrogenase, dipeptidase,
tripeptide aminopeptidase, glucose-6-phosphate
isomerase, mannose-6-phosphate isomerase (see
Adams et al. 1987).

30,0 -1

it H buventvcki

__•—, Hi v - regulus

« 20.0 —
6*

O 15.0 -

5.0 -

0.0 -

26.5 270 ti.% 2X.0 2X.5 29.0 29.5 30.0 30 5 310 315 32. B 31.5 33 33 5

Forearm length (mm)

FIGURE 1 . Size ranges in forearm length of female V.
haverstocki and V, regulus in South Australia, showing
the degree of overlap.

Results And Discussion

Forearm Length

Forearm ranges of Vespadelus spp. from South
Australia were smaller than those of Kitchener et
al. (1987). Two size groups were evident (see also
Kitchener et al 1987): Vespadelus finlaysoni and
V. darlingtoni were separated from other species
by their greater forearm length (Table 1). Females
of these two species exceded 33.0 mm whereas
the other three were usually less than 33.0 mm.
The three smaller species (V. regulus, V.
vulturnus and V. haverstocki) form a species
group which is referred to here as the "regulus
complex'.

Within the 'regulus complex' in South
Australia there were two subgroups based on
forearm length, with a larger species, V. regulus,
and two smaller species, V, haverstocki and V.
vulturnus. There was considerable overlap in
forearm length between V. haverstocki and V.
vulturnus. Among adult females, 90% of V.
regulus had forearm lengths greater than 30.5
mm, while all V. vulturnus and 95% of V,
haverstocki measured less than 30.5 mm. Fig. 1
shows the overlap in forearm length of females.

Ratio of Phalanx I: Metacarpal of Digit III

The ratio of phalanx I to the metacarpal on the
third wing digit was used by Kitchener et al.
(1987) to distinguish V, vulturnus from other
species. Table 2 shows ratio ranges calculated for
the three "regulus complex' species using only
specimens identified by electrophoresis. Since sex
does not affect this character (Kitchener et al.
1987), sexes were combined in the table. The data
demonstrate a clear distinction between V.
vulturnus (ratio > 0.396) and the other two species
(ratio < 0.395) and confirm previous findings
(Kitchener et al. 1987). Vespadelus haverstocki
and V. regulus cannot be distinguished using this
ratio.

Distribution and Habitat

Figs 2 to 6 show the geographical distribution
of each species. Initially these maps were drawn
using only bats that were identified by
electrophoresis or penis morphology. Bats
identified on wing measurements were examined
later and all fell within the distribution ranges
already recorded for the species. Habitat
information has been taken from Carpenter et al.
(1978) Reardon and Flavel (1987) and Lumsden
and Bennett (1995).

TABLE 1. Means and ranges of forearm length (mm) of
female Vespadelus species in South Australia.

Species

Range

V. darlingtoni

35.80±0.99

(34.7-38.1)

V. finlaysoni

34.45±0.86

(33.0-35.5)

V. regulus

31.37±1.14

(29.3-33.1)

V. haverstocki

29.12±0.95

(27.0-30.9)

V, vulturnus

28.47±1.24

(26.2-30.6)

TABLE 2. Range of ratios of phalanx I : metacarpal of digit
III of Vespadelus spp. of the ' regulus complex' in South
Australia. Males and females included.

Species

Ratio

V. regulus
V, haverstocki
V. vulturnus

0.34O-0.395
0.340-0.395
0.396-0.420

50
38
15

LITTLE BROWN BATS IN SOUTH AUSTRALIA

134°

26°

30°

34°

26°

30°

FIGURE 2. Distribution of V. darlingtoni in South
Australia.

Vespadelus darlingtoni (Fig. 2) is confined to
forested areas in the wetter southern and eastern
regions of the State;

Vespadelus finlaysoni (Fig. 3) occurs in arid
northern areas of South Australia in mountain
ranges and rocky country in association with
mines and caves;

FIGURE 4. Distribution of V. regulus in South Australia.

Vespadelus regulus (Fig. 4) occurs in the south
and east of South Australia in mallee woodlands
and forests. It roosts in tree hollows and buildings.
It is the only species found on Yorke and Eyre
Peninsula;

Vespadelus baverstocki (Fig. 5) is found over a
wide area in the arid and semi-arid woodlands
and shrublands of the State and roosts in tree
hollows and buildings;

26°

30°

-38°

FIGURE 5. Distribution of V. baverstocki in South
FIGURE 3 . Distribution of V. finlaysoni in South Australia. Australia.

L. F. QUEALE

26°

30°

34°

FIGURE 6. Distribution of V. vulturnus in South
Australia.

Vespadelus vulturnus (Fig. 6) is found in
woodlands and forests of the south-eastern region
of the State, and along the River Murray. It roosts
in tall trees along the River Murray and in the
South East (Reardon & Flavel 1987).

Vespadelus regulus and V. darlingtoni are the
only species of the genus to date recorded on
Kangaroo Island.

There are several areas of sympatry for
Vespadelus species in the State. The range of V.
finlaysoni is shared with that of V. baverstocki
(Figs 3 and 5) but the former species is not found
in the regions occupied by the other two 'regulus
complex' species. The northern limits for V.
regulus and the southern ranges of V. baverstocki
overlap in central Eyre Peninsula and the Murray
Mallee (Fig. 4). In this region it has been
suggested that V. baverstocki prefers Acacia spp.
open woodland and V. regulus, the mallee (C.
Kemper, T. Reardon pers. comm., Lumsden and
Bennett 1995). Vespadelus vulturnus inhabits
wetter, tall forested areas. Considerable overlap
occurs between all three 'regulus complex'
species in the River Murray region (Figs 4-6).
Here a corridor of sympatry follows the 250 mm
isohyet which coincides with the separation of
eucaiypt (mallee) and Acacia spp. woodland
(Griffin & MacCaskill 1986) and the well-watered
riverine eucaiypt forest.

Colour

Differences in fur colour have been noted
between Vespadelus species (McKean et al. 1978;
Kitchener et al. 1987; Reardon & Flavel 1987).
The fur of V. darlingtoni is very dark brown, as is
the skin on the forearms. By contrast, the fur of V.
finlaysoni is grey. The 'regulus complex' species
share a third colouration type in which the fur is
bicoloured, the hairs having brown bases and pale
tips. The ventral surface of the body is lighter than
the dorsal surface. Hence colour is not useful for
distinguishing between the species of the 'regulus
complex'.

Electrophoresis

To test the key (below) an additional sample of
female bats, with accompanying frozen tissue, was
identified using the key as 26 V. baverstocki, 10
V. regulus and four V. vulturnus. Then allozyme
electrophoresis was performed on this sample. The
allozyme comparisons confirmed that the
morphological characters had correctly predicted
the identifications of all 40 specimens. This
confirmation indicates that despite overlap in
some characters, the key can be used with
confidence to distinguish the species.

Species summaries

Vespadelus darlingtoni may be easily
distinguished from the 'regulus complex' species
by its greater size (forearm length) and from V.
finlaysoni by its dark colour and distribution.

Vespadelus finlaysoni is similarly distinguished
by its large size from the ^regulus complex'
species. There is a size overlap with V regulus
but there is no overlap in distribution.

Vespadelus vulturnus is separated from the two
larger species, V finlaysoni and V. darlingtoni on
size r with no overlap in forearm measurements. A
size overlap occurs rarely with V regulus and
more commonly with V. baverstocki and may be
distinguished from these on the basis of the
phalanx I: metacarpal ratio.

Vespadelus regulus may be distinguished from
V. darlingtoni on size and colour, from V.
finlaysoni on distribution and partly on size, and
from V. vulturnus partly on size and wing ratios.

Vespadelus baverstocki can be readily
distinguished on size from the two largest species,
V. finlaysoni and V. darlingtoni. It can be
separated from V. vulturnus on the basis of
phalanx I: metacarpal ratio. In many cases it can
be separated from V. regulus on size. When this is
not successful electrophoresis must be employed.

LITTLE BROWN BATS IN SOUTH AUSTRALIA

Key To Female Vespadelus In South Australia

1. — Forearm greater than 33 mm 2

Forearm less than 33 mm 3

2, — Latitude south of 33°S V. darlingtoni

Latitude north of 33°S V. finlaysoni

3 — Ratio of phalanx I: metacarpal of digit
111 greater than 0.39; latitude south of
33°S V. vultumus

Ratio of phalanx I; metacarpal of digit
ITT less than 0.39; locality variable 4

4. — Forearm usually greater than 30.5 mm;
mallee habitat/coastal or latitude south
of 34°S V. regulus

Forearm usually less than 30.5 mm; arid
habitat/inland or usually latitude north of
34°S V. baverstocki

Note on males

Although males can be identified by penis
morphology, this can prove difficult in the field.
The characters discussed above can be used to
identify males. However my data indicate that
male forearms tend to be shorter than females.

TABLE 3. Ranges of forearm length (mm) of male
Vespadelus species in South Australia.

Species

x SD

Range

V. darlingtoni

34.9±0.51

(33.5-38.1)

V. finlaysoni

33.2±0.64

(32.4-34.4)

V. regulus

30.3±0.96

(28.5-33.1)

V. baverstocki

29.2±0.96

(26.7-30.7)

V. vultumus

27.7±1.33

(25.2-29.5)

Thus the key can be used with some discretion as
an aid in identification of males. The one
exception is that the identification key and species
summaries do not distinguish easily between male
V. regulus and V. baverstocki on forearm lengths
(see Table 3).

Acknowledgments

I wish to thank M. Hutchinson for his advice and
encouragement. I am also grateful to K. Bowshall-Hill
for drawing the maps. I wish to thank the two referees
whose comments greatly improved the final paper, and
my husband B. McHenry for encouragement and help in
completing the manuscript.

References

ADAMS, M., BAVERSTOCK, P. R., WATTS, C. H. S.
& REARDON, T. B. 1987. Electrophoretic resolution
of species boundaries in Australian Microchiroptera.
L Eptesicus (Chiroptera: Vespertilionidae).
Australian Journal of Biological Sciences 40: 143-
162.

CARPENTER, S. M., McKEAN, J. L. & RICHARDS,
G. C. 1978. Multivariate morphometric analysis of
Eptesicus (Mammalia: Chiroptera) in Australia.
Australian Journal of Zoology 26: 629-638.

GRIFFIN, T. & MacCASKILL, M. (Eds) 1986. k Atlas
of South Australia'. South Australian Government
Printer: Adelaide.

KITCHENER, D. J., JONES, B. & CAPUTI, N. 1987,
Revision of Australian Eptesicus (Microchiroptera:
Vespertilionidae). Records of the Western Australian
Museum 13: 427-500.

LUMSDEN, L. F. & BENNETT, A. F. 1995. Bats of a
semi-arid environment in south-eastern Australia:

biogeography, ecology and conservation. Wildlife
Research 22: 217-240.

McKEAN, J. L., RICHARDS, G. C. & PRICE, W. J.

1978. A taxonomic appraisal of Eptesicus
(Chiroptera: Vespertilionidae) in Australia. Australian
Journal of Zoology 26: 529-537.

REARDON, T. B. & FLAVEL, S. 1987. 'A Guide to the
Bats of South Australia'. South Australian Museum:
Adelaide. 85pp.

TIDEMANN, C. R., WOODSIDE, D. P., ADAMS, M.
& BAVERSTOCK, P. R. 1981. Taxonomic separation
of Eptesicus (Chiroptera: Vespertilionidae) in
southeastern Australia by discriminant analysis and
electrophoresis. Australian Journal of Zoology 29:
119-128.

VOLLETH, M. & TIDEMANN, C. R. 1991. The origin
of the Australian Vespertilionidae bats, as indicated
by chromosomal studies. Zeitschrift fiir
Saugetierkunde 56: 321 330.

FOUR NEW SPECIES OF ANTIPORUS SHARP (COLEOPTERA,

DYTISCIDAE) FROM AUSTRALIA, WITH NOTES ON A. FEMORALIS

(BOH.) AND A. INTERROGATIONS (CLARK)

C. H. S. Waits

Summary

Four new species of Antiporus are described from Australia: A. jenniferae sp. nov., A.
hollingsworthi sp. nov., A. willyamsi sp. nov. and A. pembertoni sp. nov. The distribution of A.
femoralis (Boh.) and A. interrogationis (Clark) in Australia is reviewed. A key to the Australian
species is provided.

FOUR NEW SPECIES OF ANTIPORUS SHARP (COLEOPTERA, DYTISCIDAE)

FROM AUSTRALIA, WITH NOTES ON A. FEMORAHS (BOH.)

AND A. INTERROGATIONS (CLARK)

C. H. S. WATTS

WATTS, C. H. S., 1997. Four new species of Antiporus Sharp (Coleoptera, Dytiscidae) from
Australia, with notes on A. femoralis (Boh.) and A. interrogaiionis (Clark). Records of the
South Australian Museum 30(1): 35-42.

Four new species of Antiporus are described from Australia: A. jenniferae sp. nov., A.
hollingsworthi sp. nov., A. willyamsi sp. nov. and A. pembertoni sp. nov. The distribution of A.
femoralis (Boh.) and A. interrogaiionis (Clark) in Australia is reviewed. A key to the Australian
species is provided.

C. H. S. Watts, South Australian Museum, North Terrace, Adelaide, South Australia 5000.
Manuscript received, 28 February 1996.

The species of Antiporus were last revised by
Watts (1978) who recognised six species. In 1984
Brancucci (1984) resurrected A. interrogaiionis
(Clark) from synonymy with A. femoralis (Boh.).
In this paper I take the opportunity to describe
four additional species of Antiporus Sharp from
Australia and to review the distribution of A.
femoralis (Boh.) and A. interrogaiionis. One of
the new species, A. jenniferae, is close to A.
simplex Watts, one, A. willyamsi, has unusual
sexual hairs on the front and middle trochanters
and the other two belong to a group of dark
coloured Antiporus with enlarged metafemora in
the males (A. femoralis, A. interrogaiionis, A.
wilsoni Watts, A. hollingsworthi n. sp. and A.
pembertoni n.sp.). Males of this A. femoralis
group are easily separated on the shapes of the
metafemur, proclaw and aedeagus. Females are
difficult to distinguish, other than A.
interrogaiionis which can be recognised by the
colour on the pronotum.

The new species arc rare in collections,
although this may in part reflect a lack of
collecting particularly in south-west Australia.

The collections from which specimens were
examined are listed under the following
abbreviations:

AM Australian Museum, Sydney.
ANIC Australian National Insect Collection,

Canberra.
BMNH Natural History Museum, London.
MV Museum of Victoria, Melbourne.
SAMA South Australian Museum, Adelaide.
WAM Western Australian Museum, Perth.
UQIC Entomology Department Queensland
University, Brisbane.

Systematics
Key To Australian Species Of Antiporus Sharp

— Protarsus with 2 claws Females 2

— Protarsus with 1 claw Males 10

: — Relatively large (> 6mm), predominantly
reddish-yellow, elytra with 5-6 often
incomplete long dark lines

A. gilberti (Clark)

— Not with above combinations of characters
3

— Protarsi black. Relatively small (<4mm) ..
A. jenniferae sp. nov.

— Protarsi reddish-yellow, same colour as
rest of leg 4

At least the basal segment of mesotarsi
enlarged in comparison with protarsi 5

— Segments of mesotarsi similar to those on
protarsi or smaller 6

Elytron with small and large punctures,
all mesotarsal segments larger than those
of protarsi, elytra with lighter streaking in
younger specimens ...A. willyamsi sp. nov

— Elytra with uniform punctuation, only
basal segment of mesotarsi enlarged,
elytra uniformly dark red-brown to black .
A. wilsoni Watts

— Smaller (< 3.5mm) 7

— Larger (>3. 5 mm) 8

36 C.H.S. WATTS

7 — Elytron dark, usually with three reddish- — Protarsi reddish-brown, same colour as

yellow spots, pronotum reddish-yellow rest of leg 15

with dark front and rear borders

A. bakewelli (Clark) 15 — Elytron unicolorous, except for slightly

. r i . paler sutural region and sides. Metacoxal

— Elytron and pronotum uniformly red- ,. , A - , ■ , 01

, J r ,.,,,, lines evenly diverging anteriorly. Claw

brown A. simplex Watts , ,J , , ■ /, -.,

£ on protarsus relatively straight, with

8 _ More or less uniformly dark reddish- small basal tooth (Fig. 21) ..

brown, elytron tip often appears truncated A ' sim P lex Watts

A. blakei (Clark) — Elytron with one or more pale patches.

— Elytra normally black with or without Metacoxal lines much less divergent in

lighter markings. Tip of elytron pointed . 9 antenor ^f than '" mid ^- Cla f on

protarsus sickle-shaped, with large basal

9 — Black portion in centre of pronotum not tooth (Fig. 18) A. bakewelli (Clark)

reaching front border

A, interrogationis (Clark) 16 — A - femoralis complex. The species are

best separated by characters of the

— Black portion in centre of pronotum aedeagus and melafemur given in Figs

reaching front border !_ 15 In addition the b i ack portion in

.... A. hollingsworthi sp. nov. (W.A. only), thc centrc of fe pronotum does not reacn

1A. pembertoni sp. nov. (W.A. only, the front border in A interrogationis

female unknown), A.femoralis (Boh.) whereas it does in the other species.

10 — Prominent cup-like tuft of long golden

setae on mesotrochanters (Fig. 28).

Segments of mesotarsus larger than Antip or us femoralis (Boheman, 1858)

segments of protarsi ( Fl § s 7 ' 8 & 9 )

A. willyamsi sp. nov. ^

Number examined, 83.

— Mesotrochanters without such a tuft of Brancucci (1984) clarified the relationships
setae. Segments of mesotarsus smaller between this widespread and common species and
than those of protarsus 11 A interrogationis (Clark). It can be separated

r , , . ., from this species by characters of the aedeagus

11 — Metafemur produced on inside near apex , . e a u • *u a i * u

r r T - and metafemur and in having the dark patch on

" ** the middle part of the pronotum reaching the

— Metafemur simple 13 anterior border. In this last character it is similar

to the other members of the A. femoralis group. It

12 — Relatively large (>6 mm), predominantly differs from these in cha racters mentioned under

reddish-yellow, elytron with several long ±c diffcrcnt spccics and in ust ratcd m thc figures.

black lines. Metafemur with prominent

narrow triangular process on inside near Distribution

a P ex A - gUberti (Clark) Australian Capital Territory: Black

— Predominantly dark-brown to black. Mountain, ANIC; Hackett, ANIC; Mt Coree,
Metafemur broadly produced on inside ANIC; O'Connor, ANIC. New South Wales:
near apex 16 Allyn River, ANIC; Galston, ANIC; Queanbeyan,

ANIC; Sydney, MV, ANIC; Waratah, SAMA;

13 — Larger (length > 4.5 mm). Mesotibia Williams River, ANIC. South Australia:

strongly triangularly produced in middle Bcachport, AM; Eyre Peninsula, SAMA;

on outside A. blakei (Clark) Kangaroo Island, SAMA; Lucindale, SAMA; Port

— Smaller (length < 3.7 mm). Mesotibia Lincoln, AM, SAMA. Tasmania: 11 km NNW
simple or thickened and angular but Blackland, ANIC; Eaglehawk Neck, AM;
never as above 14 Georgetown, MV; Hobart, MV; Launceston, MV;

" ' Liffey Valley, ANIC; Waldheim, UQIC. Victoria:

14 Protarsi black in contrast to red-brown 4 km W. Ballan, MV; L. Barracoota, MV; Cape
colour of rest of leg Otway, MV;10 km NW Dartmouth Dam, MV; La

A. Jennifer ae sp. nov. Trobe, SAMA; L. Lcarmonth, ANIC;10 km NW

FOUR NEW SPECIES OF ANT1PORUS

FIGURES 1-15. la, dorsal view of aedeagus of A. hollingsworthi; lb, lateral view of aedeagus of A.
hollingsworlhi showing apical appendage; 2, proclaw of male A. hollingsworlhi; 3, postfemur of A. hollings-
worthi. 4, dorsal view of aedeagus of A. interrogationis; 5, proclaw of male A. interrogationis; 6, postfemur of
A. interrogationis. 7, dorsal view of aedeagus of A. femoralis; 8, proclaw of ma\t A. femoralis; 9, postfemur of
A.femoralis. 10, dorsal view of aedeagus of^. pembertoni; 11, proclaw of male A, pembertoni; 12, postfemur
of A. pembertoni. 13, dorsal view of aedeagus of A. wilsoni; 14, proclaw of male^. wilsoni; 15, postfemur of.4.
wilsoni.

C. H. S. WATTS

Lexton, MV; Montmorency, SAMA; Mt Buffalo,
MV; Mudgee, ANIC; Myrtleford, MV; Oakleigh,
MV; Reservoir, MV; Werribee, MV. Western
Australia: 9 mi NE by E Esperance, ANIC;
Geraldton, MV; King George Sound, ANIC;
Mullewa, SAMA; Ravensthorpe, ANIC; Stirling
National Park, ANIC; Swan River, SAMA;
Wilga, ANIC.

Antiporus interrogationis (Clark, 1862)
(Figs 4, 5 & 6)

Number examined, 62.

As pointed out by Brancucci (1984), A.
interrogationis is clearly distinct from A.
femoralis with which I had previously
synonymised it (Watts 1978). Apart from the
aedeagus (Fig. 4) and metafemur (Fig. 6), A.
interrogationis differs from the other members of
the A. femoralis group by the dark patch on the
middle part of the pronotum usually not reaching
the anterior border.

Distribution

Australian Capital Territory: Black
Mountain, ANIC; Paddy's River, ANIC. New
South Wales: 37 km E Hay, SAMA. South
Australia: Naracoorte, UQIC. Victoria: 4 km W
Ballan, MV; Bright, SAMA, MV, AM;
Buninyong, MV; Delatite, MV; Delley's Dell
Grampians, SAMA; Dividing Range, SAMA;
Eltham, MV; Glen Valley, MV; Hamilton, MV;
L. Purrumbete, MV; L. Wendouree, MV; Linga,
MV; Melbourne, ANIC, MV; Merrimans Creek,
MV; Mitta Mitta River, MV; Montmorency,
SAMA; Mt Buffalo, SAMA; Nelson, SAMA;
Noorinbee North, MV; Six Mile Creek
Dartmouth, MV; 8 km NE Toolangi, MV;
Warburton, MV; Werribee, MV; Yarra River,
MV; Yarrawonga, MV.

Antiporus jenniferae sp. nov.

(Figs. 25, 26 & 27)

Description (number examined 63).

Length 3. 4-3. 6mm. Oblong-oval, convex,
widest behind middle, elytron somewhat elongate
posteriorly, tip sharp. Reddish yellow, portions of
elytra a bit darker, sutural lines narrowly darker
and bordered with pale strip, apical portions
lighter; protarsi darker almost black in some
specimens. Strongly, densely, and evenly
punctured throughout. Pronotum and elytron

narrowly margined, elytron moderately serrate
towards apex, except for short distance near tip.
Apical segments of both maxillary and labial palpi
weakly bifid at tips. Prothoracic process narrowly
lanceolate, rounded, almost keeled in cross
section, slightly narrowed between procoxae.
Metacoxal lines raised, moderately separated,
diverging slightly posteriorly, widening to about
twice their narrowest width anteriorly. Some long
setae arising from pit in centre of sternite numbers
three and four. A line of long setae towards sides
of elytron.

Male: Pro- and mesotarsi moderately expanded,
first and second segments as long as wide, claw
on protarsus thickened, strongly bent near base,
abruptly narrowing to sharp point at apex, with
large basal tooth. Mesotibia robust, broadly but
weakly indented on inner side in middle. Seta tufts
on mesotrochantcrs somewhat thicker than on
female.

Female: Mesotarsi a little expanded, protarsi
less so, Protarsus with two simple claws.

Remarks

A. jenniferae can be distinguished from all
other Antiporus by its relatively small size, and its
black protarsi. It appears closest to A. simplex and
A. bakewelli. From A. simplex it differs by its
black protarsi, elytral colour pattern which
although less marked than in A. bakewelli is
clearly present, the aedeagus having a broader tip
and widening slightly in the middle compared
with the more parallel-sided aedeagus in A.
simplex and the spine at the base of the claw on
the male protarsi being larger than in A. simplex.
Based on the small numbers of specimens of both
species available it appears to be a little larger.
From A. bakewelli it differs in its black protarsi,
less marked and more diffuse elytral colour
pattern, the aedeagus with a broader tip, and the
claw on the male protarsi being straightcr in
comparison with the smoothly sickle-shaped claw
of A. bakewelli.

Distribution

Most specimens are from Cape York Peninsula,
but it is also known from Adelaide River, N.T. (in
BMNH) and Synnot Creek in northern W.A. (in
ANIC).

Types

Holotype male: "25K. N. Coen Qld. 29/9/84 C.
Watts" in SAMA.

Paratypes: 1, "12.40S 142°40E Qld Batavia
Downs Hmsd. 17-23 Jun. 1992 T.A. Weir, at

FOUR NEW SPECIES OF ANTIPORUS

A 16

* *

I I

■; y

FIGURES 16-27. 16 & 17, dorsal and lateral view of aedeagus of A. bakewelli; 18, lateral view of male^
hakewelli proclaw. 19 & 20, dorsal and lateral view of aedeagus of ,4. simplex; 21, lateral view of male A.
simplex proclaw. 22 & 23, dorsal and lateral view of aedeagus of ,4. willyamsi; 24, lateral view of male A.
willyamsi proclaw. 25 & 26, dorsal and lateral view of aedeagus of A. jenniferae; 27, lateral view of A.
jenniferae proclaw.

C. H. S. WATTS

light", in ANIC; 2, "12.40S 142.40E Batavia
Downs Qld 03-10 Mar 1993 at light, I.
Cunningham", in ANIC; 1, "16.318 126. IE
CALM site 25/1 Synnot Ck W.A. 17-20 June
1988 T.A. Weir", in ANIC; 1 1 "25k N. Coen Qld
29/9/85 C. Watts", in SAMA; 1, "24M.S.
Musgrave N.Q. 20.5.72 J.C. Brooks at light", in
ANIC; 1, "13.34S 142.32 E Qld, 17 km N.W. by
W. of Rokeby (Vardons Lagoon) 27 Oct. 1992, T.
Weir, P. Zborowski, Small pool with debris", in
ANIC; 56, "Qld Mt. Molloy 2 km S. 30/3/96, C.
Watts", in SAMA.

Antiporus pembertoni sp. nov.

(Figs 10, 11 & 12)

Description (number examined 1).

Length 4.6mm. Oblong oval, dark red brown -
black, front of head, sides of pronotum broadly
and sides of elytron narrowly but diffusely light-
testaceous. Ventral surface testaceous, sides of
meso and metasterna darker, apical tips of
metatibiae darker. Dorsal surface rather finely
punctured, stronger laterally, punctures on head a
little smaller than eye facets, punctures at rear of
pronotum towards sides and on adjacent areas of
elytra tending to join in roughly longitudinal lines.
Ventral surface much more strongly punctured,
strongly rugose-punctate laterally. Pronotum
narrowly but clearly margined, elytron weakly so.
Prothoracic process narrow, strongly raised, sides
sub-parallel, bluntly tipped. Metacoxal lines well
separated, sub-parallel in posterior third and
anterior quarter, diverging in between so that
anteriorly about twice the distance apart as
posteriorly, area between coxal lines and forward
onto mesosterum depressed.

Male: Protarsi moderately expanded, protarsal
claw large, slender, with well developed slender
basal tooth. Mesotibia weakly indented on inner
margin towards apex. Metafemur broadly
expanded on inner edge at apex, some strong
punctures and weak longitudinal ridging on
expanded area.

Female: Unknown

Remarks

More weakly punctured than the other species
in the A. femoralis complex, with the large
punctures along the front of the pronotum distinct
and more than twice the size of others on the
pronotum. Characters of the male aedeagus and
metafemur are diagnostic. Known only from a
single specimen I collected in a small stream in
cleared land near Pemberton.

Distribution

Known only from unique type from near
Pemberton, S.W. Australia.

Type

Holotype male: "W. Aust. 15 km N.W.
Pemberton, 17 May 1987, C. Watts" in SAMA,

Antiporus hollingsworthi sp. nov.
(Figs 1,1a, 2, & 3)

Description (number examined 15).

Length 4.1^. 6mm. Oblong oval, convex. Dark
red-brown, appendages lighter. Strongly evenly
and densely punctate throughout, punctures
smaller on head than elytra, punctures at rear of
pronotum towards sides and on adjacent areas of
elytra tending to join in linear longitudinal lines,
punctures towards sides of metacoxae and
sternites and on elytral epipleura very close
together and tending to coalesce. Pronotum and
elytra narrowly margined except for short distance
at tips of elytra, lacking serration, sides of elytra
before apex appear weakly and broadly flanged
from some angles. Prothoracic process narrow,
blunt, strongly carinate. Metacoxal lines sub-
parallel in anterior and posterior third, rapidly
diverging in middle.

Male: Pro- and mesotarsi moderately expanded.
First and second segments wider than long. Claw
on protarsus strongly bent near base, straight with
parallel sides in middle, rapidly narrowing at apex
basal tooth subobsolete. Mesotibia a little
thickened, weakly and broadly indented on inner
edge in middle. Apical ventral edge of mesofemur
expanded a little. Inner ventral edge of metafemur
with large, broad, triangular expansion in apical
third. Parameres robust, broad, aedeagus arrow-
headed with thin protuberance above tip arising
from neck of arrow.

Female: Pro- and mesotarsi a little expanded;
claws, meso- and metafemur and tibia simple.

Remarks

Antiporus hollingsworthi is a member of the A.
femoralis group. It appears closest to A. wilsoni
from the south-east of Australia which it
resembles in the pronotal and elytral grooving,
thickened male mesotibia and expanded
metafemur.

Antiporus hollingsworthi can be separated from
both A. femoralis and A. interrogationis by the
lack of pronounced paler areas on the head,
pronotum and elytra, although some specimens of

FOUR NEW SPECIES OF ANTIPORUS

A. hollingsworthi do have head, pronotum and
elytral patches paler than the rest of the upper
surface. The tendency of the punctures on the rear
edge of the pronotum and the adjacent areas on
the elytra to link up and form grooves is more
pronounced than in either A. femoralis or A.
interrogationis and the triangular expansion on
the male metafemur is more rounded than in either
of these species. A. femoralis and A.
interrogationis also lack the thickened and
indented mesotibia present in A hollingsworthi^
A. wilsoni and to a lesser extent in A. pembertoni.
Antiporus hollingsworthi is more strongly
punctured than A. pembertoni and the males have
differently shaped metafemora (Fig. 3). It differs
from A. wilsoni by characters of the aedeagus, by
the indentation on the male mesotibia being more
central, and in having a triangular rather than a
rounded expansion to the male metafemur.
Antiporus hollingsworthi differs from all other
Antiporus species (and all Australian Dytiscidae)
by the apical appendage on the aedeagus.

Distribution

So far A. hollingsworthi is known only from
Perth and Carrington, W.A. and from two
unlocalised localities in the South-west. However,
of all the Australian regions this is the most poorly
sampled for water beetles so it is likely to be more
abundant than the few known specimens indicate.

Types

Holotype male: "W. Aust. Maidavale, 27th
April 1990, C. Watts", in SAM A.

Paratypes: 1 male, 7 females, same data as
Holotype, in SAMA; 1 female "Perth 10/65 DE",
in SAMA; 1 female "S.W.A. ?Rotnest, Edward",
in SAMA; 1 male "JT201", in SAMA; 1 male, 1
female "R.P. McMillan, Carrington, 19.7.53", in
WAM.

Etymology

Named after Rod Hollingsworth, a strong
supporter of Natural History at the South
Australian Museum.

Antiporus willy amsi sp. no v.

(Figs 22, 23, 24 & 28)

Description (number examined 4).

Length 3.4-3.6 mm. Oblong-oval. Widest
behind middle. Testaceous, head, lateral quarters
of pronotum, patches on elytra and appendages

lighter, metafemur and tibia tend to be a little
darker towards their apices, lighter patches on
elytron tending to form broad longitudinal streaks.
Strongly, densely and evenly punctate, punctures
on discs of pronotum and head weaker,
particularly on head, a few scattered much smaller
punctures on disc of pronotum and elytra.
Pronotum narrowly but distinctly margined, elytra
less so. Prothoracic process with central keel,
narrow between procoxae. Metacoxal lines raised,
sub-parallel in posterior half and anterior quarter,
diverging in between, about twice the distance
apart anteriorly compared with posteriorly,
reaching mesofemur. Some long setae arising
from pits in centre of sternites three and four, and
a line of long setae on dorsal surface of elytron
close to side.

Male: Protarsi expanded, second segment
broadest, mesotarsi strongly expanded, first and
second segments largest, sub-equal. Mesofemur
and mesotibia stouter than profemur and protibia.
Protarsus with single claw, strongly bent, long and
narrow, with well developed basal tooth and weak
bulbous area in middle. Protrochanters with well
marked cup-like tuft of golden setae,
mesotrochanters with a much larger scallop-
shaped tuft of similar setae. Metafemur stout,
ventral inner apical region with scattered large
punctures and about four longitudinal grooves/
ridges.

Female: Protarsus with two weak claws.
Mesotarsi moderately expanded, first and second
segments largest, sub-equal. Mesotibia stout.

Remarks

The species is easily recognised by the
spectacular development of the setae on the
trochanters of the male which is unlike anything I

FIGURE 28.
willyamsi.

Meso- and metatrochanters of A,

C. H. S. WATTS

have seen on other beetles. The very broad
mesotarsi in the male and the striped elytral colour
pattern in teneral individuals are also distinctive.

Since collecting the original specimens in 1983,
I have revisited the locality (a deep, narrow,
spring-fed drainage ditch in acidic soil) a number
of times without success. Biologically the area has
now changed considerably from a healthy habitat
with numerous species to one that is almost
devoid of insect life although physically little
changed.

Distribution

Known only from the type locality in the south-
east of South Australia and from Healesville,
Victoria.

Types

Holotype male: "10 km S. Robe, S.A., 1/83, C.
Watts", in SAMA.

Paratypes: 2 females, same data as holotype, in
SAMA; 1 female, "Healesville, V. 12/68, C.
Watts", in SAMA.

Checklist Of Australian Antiporus Sharp

A. bakewelli (Clark, 1862)
A. hlakei (Clark, 1862)
A.femoralis (Boheman, 1858)
A. gilberti (Clark, 1862)
A. hollingsworthi sp. nov.
A. Jennifer ae sp. nov.

A. pembertoni sp. nov.
A. simplex Watts, 1978
A. willyamsi sp. nov.
A. wiisoni Watts, 1978
A. interrogationis (Clark,

1862)

Acknowledgments

References

The curators of the collections listed earlier are
thanked for allowing me to examine specimens in their
care. Mrs Vicki Wade typed the manuscript and Mr R.
Gutteridge drew the illustrations. All these are thanked
for their support and help.

BRANCUCCI, M. 1984. Notes on some species of the
genus Antiporus (Coleoptera: Dytiscidae). Aquatic-
Insects 6: 149-152.

WATTS, C. H. S. 1978. A revision of the Australian
Dytiscidae (Coleoptera). Australian Journal of
Zoology, Supplementary series No 57: 1-166.

THYLACiSUS MEGiRL4.\f< A NEW SPECIES OF THYLACINE (MARSUPEAL1A:
THYLACINIDAE) FROM THE ONGKVA LOCAL FAUNA OF CENTRAL AUSTRALIA

PETER F. MURRAY

MURRAY* P. F. 1997. Thyhwitws megirianr, a new species of thylacitie (Marsupialia:
Thylaeinidae} from the Oneeva Locai Fauna oi' Central Australia. Records of the South
Australian Atu.xeum 3G< I): 43-61 r

Approximately 10% larger than a maximum-seized Tasmanian wolf, the laic M ioccne' early
pliocene Thylaciniu megirinni n. sp, exhibits some similarities to the Sate Miocene thylacinid,
T. potens Woodbwnc l9o7 arid early to mid-Miocene ^imiyaanus ificksom Muirhead and
Archer 1590, while simultaneously expressing several derived features in common with the
Recent ihylacLtiu. 7" cy^OLtphuius. Other ihan the presence of a tiny stylar cusp-Hlce strucuue
distal to stylar cusp D on M\ which could be an artefact of wear, T. megiriani shows no
defi title uniquely derived characters Lhat would exclude i\ from ancc-slry of the recently extinct
T. cyn&cepttalus.

Peter P. Murray, Museums and Art Galleries of the Northern Territory, PO Box 2109 Alice
Springs, Northern Territory 0871, Austraha. Manuscript received 14 October, 1996, revised 3
Marth. 1 997.

Fossil vertebrates were first described from the
Alcoota Local Fauna al Alcoota (Fig. 1), northeast
of Alice Springs, by Woodburne (1967). The
strati graphically superposed Qngcva Local Fauna
contains certain taxa lhat are more derived than
those from the nearby Alcoota Local Fauna
(Murray, Megirian & Wells 1993, Megirlon,
Murray & Wells 19%). Among its characteristic
species is a zygomaturine diprotodontid, Kolopsis
yperus Murray* Mcgirian & Wells, which is
probably synonymous with the Chcltenhamian-
aged Zygoma f urns gtilt Stirlon 1967. A", yp&us is
iiiore similar lo Zygomaturus species than is the
Akoota Local Fauna species Koi&psis torus
Woodburne 1967. Thylacinus megiriani appears
to be analogously derived in comparison to the
Alcoota Local Fauna thylacinc, T. patens.

A direct ancestor of T. cynocephatus has not
been identified, I! rosfroiis, T. spelaean and T.
major are considered to be synonymous with T.
cynocephatMS (Dawson J9S2). Other thylacinids
are considered lo be too primitive or too
specialised (Archer 1982, Muirhead & Archer
1990, Muirhead 1992, Wroc pcrs. com.).
Ambiguity in the homology of the posterior stylar
euspules on (he molars of T. cynocephcttus has
resulted in a less than definitive resolution of late
Tertiary thylacinid phylogcny. T. megiriani
provides some additional: evidence pertaining to
the homology of the cusps and some hypotheses
of ihc phytogeny o\^ Ihe recent species of
Thvlacinw are discussed.

Matejuals And Methods

As is often the case with Ongeva LF fossils,

NTM P961S is not an ideal specimen, having
been found in a number of pieces scattered
throughout a hard nodule of calcareous sandstone.
As fragments were extracted more or less
individually, including fragments of the molar
teeth, the specimen was glued together at ihe
excavation silc along what appeared to be the
likely contacts.. The fragile condition of the
specimen dictated lhat several coals of Synocryl
(Bcdacryl) be applied to harden and hold it
together securely. However, this compound
sometimes has ihe disadvantage of obscuring
important morphological details, particularly
when fine particles of the matrix become
incorporated with the solution, sometimes
suppressing surface detail and occasionally even
mimicking small structures.

So it was with this specimen, that certain
features of the restoration seemed untenable, such
as a slight overlap of thy P~- alveolus with that of
the P 1 . As Thylacinus potcn.i has no appreciable
diastema between these teeth, this arrangement
appeared to be plausible, though further cleaning
of the matrix and excess Synocryl from the
specimen, in response to an observation by one of
the referees, proved that it had not been restored
corrcclly. This revelation provoked a more radical
program of disassembly and cleaning of the
specimen which I was initially afraid to attempt

135°E

FIGURE 1 . Locality map; the Dngcva LF ih located un Akoota Station, near Alice Springs Northern Territory.

due lo its extreme fragility. The new restoration
has greatly improved the resolution of the

specimen, to the extent thai many of my initial
observations can be slated more confidently;
though as will be discussed some features of the
specimen remain equivocal.

As the first draft of (his paper was based on the
original restoration, I have briefly described and
illustrated the modifications [ made lo the
specimen to avoid confusion with, any errant
copies of the earlier manuscript. 1 ani extremely
grateful to my referees fur alerting me to (he
possibility thai the specimen might be distorted,
and for drawing to my attention several other
morphological features, resulting, I believe, in a
much improved account of the species.

Systemaiics

Family THYLACINIDAE Bonaparte 1838
Thyluiinus me&iritmin. sn,

Hoiatype
NTM P9618 (field ref. no, SQ107), left maxilla

with P 1 , P\ M 1 " 1 .

Type focatity

Hill l h South Quarry, approximately 2 km
southwest of the junction of Ongeva and Waite
Creeks, Alcoota Station, Northern Territory,

latitude 22 a 52'5, Longiludc, L34*2TE The Type

THYLA ONUS MEWRIAM N .SP.

FIGURE 2. Photograph of the fcft maxilla of Thylaeinu* megiriani n. sp. (NTM P9G18); A, lateral aspect; B,
palatal aspect. This photograph depidLs iht: field reiteration of the specimen in which the fragment containing P 1
and P 2 had been glued approximately S.O mm posterior 10 its Lorrea position, lcadiri|f to the initial impression rhat
the posterior root of P ] was adjacent to, or sightly overlapped thai of P*. Note- also thai the M'istwistcd medially
(H) and is nearly a centimetre out of alignment with MP.

was recovered from a sandy siltstone channel

deposit in the Watte Formation, approximately
10m above the fossitiferous lacustrine scdimenls
containing the Alcoola Local Fauna (Murray,
NU'iiiruin &. WdLs H'93. Megirian, Murray &
Wells 1996), (Fig, I).

Fauna
Ongeva Local Fauna (Murray et at. 19931.

Rock unit and age

Waire Formation, Cheltenhamian equivalent,
laic Miocene-?cailv Pliocene.

Specific diagnosis

A Eargc, long-snouted thylacinid, equal to, or
larger than T- cynocephahs bul less massive than
r. p&tens; much larger than N. dicks on?
(Muirhead and Archer 1990) and T. macknesxi
(Muirhead 3992). P 1-2 shorter and narrower than
in T. potem, P in line with P 2 \ contrasting with
T. poterts in which P 1 is obliquely oriented and
situated medial to the midline of the canine
alveolus, P 5 large as in T. patens. Premolar
diaslcmata longer than in T. potens and in
Ntmhacinus dicksoni? more comparable to T.
cynocephalus. "M 1 J longer and narrower than in

FIGURE 3. Outlines depicting changes made to the initial restoration of the specimen; solid outline represents the
new restoration; compare with Fig. 2. A, lateral aspect; B, palatal aspect. The specimen was taken apart at its
primary joins, cleaned of excess Synocryl and glue, and reassembled. The anterior fragment of the rostrum was
carefully cleaned revealing a labial expansion representing the anterior alveolus of P 3 . It was glued in place at what
is probably the minimum distance from P 3 , as no definite contact could be identified. Removal of additional matrix
from a fragment of palate that previously did not seem to fit anywhere on the specimen, revealed the inner lining
of the anterior P 3 root and was glued in position. The M 3 was twisted out of alignment with the molar row by about
15°. This region of the specimen, which consisted primarily of impacted matrix and bone fragments was
embedded in modelling clay, then softened in acetone and pushed gradually back into its correct position. The
area above and internal to the M 3 was then reinforced with dental plaster. These changes resulted in a significant
transformation of the occlusal outline of the specimen, which subsequently lost some of its resemblances to T.
potens.

THYLACINUS MEGIRIANI N.SP.

1 j

ALV/C

ALV/M4

FIGURE 4, Drawing of corrected restoration of T. megiriani n. sp. (NTM P9618) left maxilla; A, lateral aspect;
B, palatal aspect. Abbreviations: AC, precingulum; ALV/, alveolus; B, stylar cusp B; C, canine; D, stylar cusp
D; E, stylar cusp E (brackets [E] indicate doubtful, or if worn, the approximate position of a structure); IEM,
interdental embrasure; IOF, infraorbital foramen; MI. ..3, molars; ME, metacone; MJS, maxillojugal suture;
PI ...3, premolars; PA, paracone; PC, postcingulum.

T. potens though relatively shorter in proportion to
their width than in T. cynocephalus. Agrees
meristically with T. cynocephalus in that M 3 is
slightly longer than M 2 , differing from T. potens
and Nimbacinus dicksoni in which M 2 is slightly
longer than M 3 . Length of molars exceeds anterior
width from protocone to parastyle; stylar shelf

considerably narrower than in T. potens;
metastylar spur much larger than parastylar corner
on M\ contrasting with T. potens in which they
are more nearly equal-sized; stylar cusp D present
on M 1- " 3 as in Nimbacinus dicksoni and probably
as in T, potens; rudimentary stylar cusp E present
on M 2 , absent in T. potens and in T. cynocephalus

P. F. MURRAY

but present on M 1 2 in Nimbacinus dicksoni.
Rudimentary anterior cingulum present on M 2 in
contrast to Nimbacinus dicksoni in which it is
well-developed on M'~\ T. potens in which it is
present on M 2 ~ 3 and T. macknessi in which it is
present on M 1 , the only upper molar known for
that species. A small precingulum is present on
M 2 of T. megiriani. Ectoflex of M 2 ~ 3 less
pronounced than in T. potens but greater than in
T. cynocephalus. Paraconal reduction on M'~ 3
comparable to that of T. cynocephalus; protocones
on M 23 reduced, similar to T. cynocephalus.
Infraorbital foramen opens above posterior half of
M 2 as in T. cynocephalus, unlike T. potens in
which it opens above M 1 ; maxillojugal suture
nearly reaches the margin of infraorbital foramen,
approaching the condition in T. cynocephalus and
in contrast with Nimbacinus dicksoni in which it
terminates considerably short of the margin of the
foramen.

Etymology

It is my privilege to name this species after Dirk
Megirian, Geologist, Museums and Art Galleries
of the Northern Territory, Darwin, NT; who
discovered and meticulously extracted the
specimen.

Description
Maxilla

Thylacinus megiriani n. sp. consists of a
maxilla that was recovered from nodular
calcareous sandstone matrix in about a dozen
pieces (Figs 2-4, 7). All but a few small
fragments have been assembled into the left side
of the animal's snout which includes the base of
the zygomatic arch to just below the orbit and the
infraorbital foramen and canal, to the posterior
margin of the canine alveolus. The full extent of
the missing jugal is clearly inscribed by its suture.
In palatal aspect the maxilla is shallowly concave
on the labial side and the snout was constricted
behind P ! much as in T. cynocephalus. The base
of the zygomatic arch is convex and marked by a
pair of low, irregular crests. The higher one,
corresponding to the maxillojugal suture, extends
obliquely upwards to immediately behind the
opening of the infraorbital foramen.

The infraorbital foramen is a large,
approximately 9.0 mm by 10.5 mm opening
situated 21.0 mm above the approximate middle
of the posterior half of M 2 (Figs 4, 7). Its position
in T. megiriani is similar to that of T.
cynocephalus and differs from the more anteriorly
situated opening in T. potens, above M 1 . Anterior

to the infraorbital foramen, a shallow, oval fossa
extends to above P 3 , anterior to which, the lateral
margin of the rostrum becomes convex.

A fragment of the maxillary palate extending
from P 1 to P 2 is broken at the median suture, A
prominent median palatal ridge is apparent,
parallel to which runs a deep, 5,0 mm wide
palatal groove. The palatal width at between the
posterior roots of P 1 would have been about 24.0
mm, approximately 5.0 mm wider than in T.
potens. The palate is narrower at this point in T,
potens because the posterior roots of P 1 are angled
inwards and are also larger, whereas they are
aligned with the median base of the canine
anteriorly and the P 2 posteriorly in T. megiriani
and T. cynocephalus (Figs 2-5).

Three additional fragments of the palate can be
approximated to adjacent structures medial to P 3 -
M 1 (Figs 3, 4) as additional removal of matrix and
Synocryl revealed the inner margin of the P 3
alveolus on the largest of the fragments. The
premolar diastemata are similar to those of T.
potens in that the P 1 and the P 2 are widely
separated, whereas the diastema between P 2 and
P 3 is shorter (Figs 5, 6). In the original restoration
of T. megiriani (Figs 2, 3) it appeared that the
posterior root of P 2 might have overlapped the
labial side of the anterior root of P 3 by about 3.0
mm. The specimen is broken at this point and the
contacts were not quite congruent, causing the
anterior part of the snout to twist out of alignment.
This contact was taken apart and cleaned,
revealing more of the P 2 alveolus and a straight
section of diastema, indicating that a P 2 /P 3
diastema of at least 5.0 mm is actually present
(Fig. 6). The premolars of T. megiriani are large
and posteriorly directed, though in proportion to
the molars, the First and second premolar crowns
and their alveoli are absolutely and relatively
smaller than those of T. potens (Figs 2-6, Table
1) but the P 3 crown, though not the alveolus, is
similar in size to that of T. potens. The lengths of
the premolar alveoli of T. megiriani are as
follows: P 1 , 10.5 mm; P 2 , 15.0 mm; P 3 , 18.5 mm.
Corresponding measurements of T. potens are: P ! ,
-15 mm;P 2 , 17.5 mm; P 3 , 20.5 mm (Fig. 6).

Reconstruction

The reconstructed snout of T. megiriani is
compared with that of T. potens and T.
cynocephalus in Fig. 5. This reconstruction, based
on modifications of the original restoration of the
specimen (Fig. 3) indicates that although it is
comparable in size and robustness to T. potens, its
shape is more similar to that of T. cynocephalus

THYLACINUS MEGIRIANIN. SP.

10 20 30 40 50
l I I

FIGURE 5. Reconstruction of the rostrum of T. megiriani n. sp. for comparison with those of T. potens and T,
cynocephalus, drawn to scale. Comparative lengths of the premolar alveoli are indicated by brackets. T. megiriani
n. sp. is approximately the same size as T. potens, both species being considerably larger and more robust than
the Recent Tasmanian Wo\f,Thylacinus cynocephalus. T. megiriani differs from T. potens in having a wider, less
constricted palate at the level of P 1 , in its alignment of the premolars, and in having smaller P'~ 2 alveoli and longer
P^diastemata, all features shared with T, cynocephalus. Distinctive features of the rostrum and palate of T. potens
include the large size and oblique emplacement of P 1 , distinct narrowing of palate at the level of P 2 , separate rather
than confluent P' and canine alveoli, emplacement of the P' internal to the canine base, the incisive foramina are
formed within a deep, oval pit, (apparently absent in T. megiriani) and the palatal fenestrae are small.

P. F. MURRAY

15 6.2 17.5 2.0 20

j — i i

FIGURE 6. Diagram comparing proportional features of the cheek tooth rows of A, Thylacinus potens, B, T.
megiriani n. sp. andC, Thylacinus cynocephalus, all drawn to the same length from anterior P l to the metastylar
tip of M 3 , measurements in millimetres. Refer to scale and actual measurements given above, as brackets show
relative rather than scaled dimensions of the alveoli and the length of the metastylar spur from ectoflexus to tip.
The P 3 alveolus in?", potens, damaged on both sides, is a composite restoration; note the relatively and absolutely
longer diastema between P 1 and P 2 in T. megiriani, suggesting that the P 2 /P 3 diastema was probably slightly
longer than restored. The two pieces were glued at this point because no additional surface was available to
provide a secure union. Observe that the diastemata in T. cynocephalus are approximately equal lengths.

in being more drawn out in the premolar region
and in the premolar rows lying essentially parallel
to each other, rather than converging at the level
of P 2 . As the restored P 2 /P 3 diastema is a minimum
dimension, the snout of T. megihani may actually
have been slightly longer than depicted. The
distinctive median pit for the incisive foramina in
T. potens appears to have been absent in T.
megiriani. Note also that the P 1 alveolus is
confluent with the canine alveolus in T.
cynocephalus and T. megiriani, whereas in T.

potens they are separated by a bony ridge.
Longitudinal palatal crests define the lateral
margins of the palatine fenestrae in T. potens.
Similar crests in T. megiriani are situated further
apart, indicating that the palatal fenestrae were
probably wider.

Premolars

P 1 is a small two-rooted premolar closely
resembling that of T. cynocephalus. The crown is
8.5 mm long and 4.5 mm wide. The anterior

THYLACINUS MEGIRIANI N.SP.

FIGURE 7. Diagram comparing the position of the infraorbital foramen relative to interproximal MVM 2 and its
relationship to the maxillojugal suture in A, Thylacinus cynocephalus; B, Thylacinus megiriani n. sp.; C,
Nimbacinus dicksoni (NTM P907-3) and D, Thylacinus potens. In this series, the infraorbital foramen in
Thylacinus potens is positioned more anteriorly than in the other species under consideration. A more anteriorly
situated infraorbital foramen is a primitive feature in didelphoids, dasyurids and probably thylacinids. However,
the infraorbital foramen in Nimbacinus is situated more posteriorly than in T, potens, an otherwise much more
derived species. Note that the maxillojugal suture lies well posterior to the foramen in Nimbacinus, whereas it
reaches or nearly reaches the posterior margin of the foramen in T. potens, as it does in T megiriani and T.
cynocephalus. This relationship is a derived feature, suggesting that the forward position of the foramen in T.
potens is a secondary, rather than a primary condition, indicative of a derived state. Abbreviations: IOF,
infraorbital foramen; LJS, lacrimojugal suture; MJS, maxillojugal suture.

P. F. MURJRAY

margin of the crown commences with an enamel
bulge about 3.0 mm above the alveolus. The
posterior profile is steeper, longer and slightly
concave, terminating in an enamel thickening 1.0
mm above the alveolus (Figs 2-4, Table 1)

P 2 is represented by its anterior root and
posterior alveolus. Its alveolus is about 15.0 mm

long, 2.5 mm shorter than that of T. potens. The
posterior alveolus is slightly larger in diameter
than the anterior one.

P 3 is considerably the largest premolar. The
length of the crown at the base of the enamel is
about 16.5 mm. The circular, heavily worn
posterobasal cuspule is 8.5 mm transversely,

FIGURE 8. Photographs of the left M 1 " 3 ofThylacinus megiriani n. sp.; A, M 1 " 3 , taken before correction of the
position of M 3 on the specimen; B, enlargement of M 3 showing stylar cusp D and possible aberrant stylar cusp
E, which may actually be a peculiar wear artefact; C, paste-up showing the approximately correct alignment of
the M 2 metastyle with the parastylar corner of M 3 . Because of its dislocation, the occlusal surface of M 3 is tilted
more lingually than those of M 1-2 , revealing the entire buccal side of the crown and resulting in the misleading
impression that the metacone is situated more posterolingually than it actually is. Note the faint, postcingulum-
like structure on M\ the small precingulum on M 2 and absence of same on M 3 ; The remnant stylar cusp E on M 2
is indistinct in A, but is faintly visible in C.

THYLACINVS MEGIRIANI N.SP.

while the width across the anterior enamelled part
of the crown immediately above the posterobasal
cuspule is 7.0 mm. The crown is heavily worn on
the anterior and posterior surfaces where distinct
wear facets result in a steep, slightly biconcave
profile.

Molars.

M 1 is heavily worn (Figs 2, 4, 8A, 9C). Both
the paracone and metacone have been ground
flush with the stylar shelf. The remnant base of
the paracone indicates that it was probably
reduced relative to the metacone. A remnant V-
shaped 'wall' of vertical facets above the
premetacrista and centrocrista are faintly visible
on the specimen. Stylar cusp D, situated
immediately posterior to the labial cleft, is the
highest relief on the worn occlusal surface. Stylar
cusp E is not apparent. The protocone is round
and slightly constricted near its base. A faint
cingulum-Iike crest extends from a low bulge
located about mid-way between the base of the
protocone and the metastylar tip, though its exact
form and extent is obscured by a layer of calcite.
The length of the tooth along the stylar crest is
14.7 mm. Its transverse width from the protocone
to the parastyle is 12.3 mm and its posterior width
from protocone to metastyle is 16.0 mm. The M'
of T. megiriani is similar, but relatively longer
than the equivalent tooth in T. potens (Fig. 9B)
differing from that of N, dicksoni (Muirhead &
Archer 1990) in its reduced stylar shelf, reduced
paracone and absence of an anterior cingulum
(Fig. 9A) and from that of T, macknessi
(Muirhead 1992) in its retention of a distinct stylar
shelf remnant with a large stylar cusp D, in the
reduction of the paracone, in the absence of an
anterior cingulum, and in having a steep facet or
'wall' above the paracone and metacone bases
(Fig. 9D).

M 2 is considerably larger than M 1 and broader
transversely, though slightly shorter across the
stylar wings than M 3 (Figs 2, 4, 8A, C, 9C; Table
1). Its length from parastyle to metastyle is 16.8
mm. Its anterior transverse dimension from
protocone to parastyle is 15.0 mm and its posterior
transverse dimension from protocone to metastyle
is 19.5 mm. The parastylar corner is small
compared to the metastylar corner. The smaller,
lower and narrower paracone is cleanly divided
from the metacone by a mesial continuation of the
labial cleft. Approximately 3.0 mm distal to the
labial cleft and 2.0 mm labiad of the metacone is
a prominent stylar cusp D. It is connected to the
metacone by a short crest. Separated by a shallow

labial groove from stylar cusp D is a second, much
smaller enamel expansion situated about 3.0 mm
anterior the metastyle, which appears to represent
a remnant stylar cusp E. Although not prominent,
the tiny cusp has a small wear facet and a labial
bulge on the stylar shelf is evident. The
anterotingual surface between the protocone and
parastyle is distinctly convex and emarginated by
a short, though strong precingulum. The protocone
is smaller than on M 1 and is more U-shaped than
rounded. Remnants of the pre-and post-
protocristae extend from either side of the apex of
the protocone to the bases of the paracone and
metacone respectively. A small, but distinct lobe
is situated immediately above the dentine-enamel
junction, approximately midway between the
metastyle and the protocone. In overall shape, the
M 2 of T. megiriani is structurally intermediate
between Nimbacinus dicksoni and T. potens, but
differs from both in having a much reduced
parastylar wing, reduced paracone and relatively
reduced height of the stylar shelf. There is no
indication of stylar cusp E on T. potens, but the
cusp is well -developed, though small, in
Nimbacinus dicksoni,

M 3 is slightly longer and more slender across
the metastyle and protocone than M 2 . The
anterolingual and posterolingual emarginations
are reduced and the precingulum is rudimentary
(Figs 2, 4, 8B, C, 9C; Table 1). The parastylar
spur is smaller in proportion to its metastylar
component than in the previous molar, though the
ectoflex is more pronounced on M 3 than on M 2 .
The length of the crown from parastyle to
metastyle is 17.4 mm. Its anterior transverse
dimension is 16.1 mm and its posterior transverse
dimension is 20.3 mm. Though heavily worn, it is
apparent that the paracone was originally low,
narrow and much smaller than the metacone,
which in turn, is high, transversely elongated and
anteroposteriorly narrow. As in Thylacinus
cynocephalus, the preparacrista is short and the
internal angle formed by the preparacrista and
postmetacrista is more obtuse than in T. potens.
Prominent pre- and postprotocristae meet at the
apex of the narrow, V-shaped protocone. Both
crests extend to near the cusps of the paracone
and metacone accentuating a furrow-like occlusal
fossa between them. Stylar cusp D is situated -3.0
mm posterior to the labial cleft in the same
position relative to the posterolabial base of the
metacone as in M 2 . It is connected to the metacone
by a low crest. A possible rudiment of stylar cusp
E is situated about 4.0 mm anterior to the
metastyle. Though small, it is more obvious than

P. F. MURRAY

C if D [

12 3 4 5

_L_I l_l

C CCR

B PPC PRMC

12 3 4 5

ME I I I I LJ

LLLL
MM

POPC PRMC

POPC / PRMC
CCR

FIGURE 9. Comparison of M 1 " 3 of T. megirianin. sp. with thylacinid species: A, Nimbacinus dicksoni (NTM
P907-3); B, Thylacinus potens; C, Thylacinus megiriani; D, Thylacinus macknessi (M 1 reversed for
comparison); E, Thylacinus cynocephalus. Abbreviations: AC, precinguium; B, stylar cusp B; C, stylar cusp
C; CCR, centrocrista;D, stylar cusp D; E, stylar cusp E;ef,ectoflexus;gv, natural orwear groove in stylar shelf;
MCL, metaconule; ME, metacone; MS, tip of metastyle; PA, paracone; PC, postcingulum; PCL, paraconule;
PMC, postmetacrista; POPC, postparacrista; PR, protocone; PRMC, premetacrista; PSC, crest connecting
metacone and stylar cusp D.

THYLACINVS MEGIRIANI N.SP.

TABLE 1. Measurements of cheek teeth of Thylacinus megiriani compared with those of T. potens and T.
cynocephalus.

T. megiriani

T. potens

T . potens

T. cynocephalus

NTMP9618

CPC6746

UCMP 66971

sample means
(Woodbume 1967)

8.5

— _

—

6.2

P 1

4.5

—

3.3

<12.0

12.4

—

8.3

P 2

<5.0

5.5

—

3.8

16.5

16.0

—

10.6

P 3

8.5

8.8

—

5.0

14.7

12.0

—

10.9

M 1

12.3

12.8

—

11.5

w 2

16.0

13.5

—

11.5

16.8

15.7

—

13.2

M 2

w i

15.0

13.9

—

10.0

w 2

19.5

17.5

—

15.0

17.4

15.2

14.6

15.1

M 3

w t

16.1

15.9

14.7

12.0

w 2

20.3

19.0

17.5

17.8

the structure in the equivalent position on M 2 . It
consists of a nearly circular raised enamel ridge
with an oval wear facet or pit in the centre (Fig. 8
B, C). After long deliberation of the structure
under the microscope, I have concluded that it
could represent a freakish wear pattern, but that it
also indicates that thicker, more obdurate enamel
is located where stylar cusp E would be
anticipated. The enamel on the molars of this
specimen is uniformly quite thick and it may be
the case that some of these small structures,
including the postcingulum-like crests on M 12
might be the result of an exceptionally active
enamel organ. A -3.0 mm wide attrition furrow
between stylar cusp D and the equivocal cusp E
extends vertically some 4.0 mm up the labial side
of the stylar crest.

Meristic gradients of the molars are as follows:
metastyle to parastyle length, M'<M 2 <M 3 ;
ectoflex, M'<M 2 <M 3 ; Protocone base diameter,
M'>M 2 >M 3 ; Paraconc, M l >M 2 >M 3 ; Metacone
diameter, M ! ?M 2 >M 3 ; Metacone height,
M l ?M 2 <M 3 ; Parastyle length, M'>M 2 >M 3 ;
Metastyle length, M'<M 2 <M 3 ; Precingulum, M 1
nil M 2 present M 3 nil; Stylar cusp D, M'>M 2 >M 3 ;
Stylar cusp E, ? M 1 , weakly expressed on M 2 , ?
M 3 ; ?Postcingulum length, M'>M 2 nil on M 3 .

Discussion

The similarity in size and robusticity of T.
megiriani to T. potens contributed to some of the
errors in the initial restoration and interpretation
of the fragmentary specimen. The new restoration
of the fossil indicates that T. megiriani was a
long-snouted species, proportionally more similar
to T. cynocephalus than to T. potens. The length
of the diastema between P 2 and P 3 is a minimum
estimation, as no definite contact between the two
fragments of the snout could be identified. The
posterior margin of the fragment containing P 2
shows a definite lateral expansion for the anterior
root of P 3 , but the actual alveolar margin and
internal lining of compact bone is absent (Fig. 3).
Consequently, Thylacinus potens appears to have
undergone proportional changes in the rostrum,
possibly a forward adjustment of the zygomatic
root to shift the line of action of the masseter
muscle more anteriorly. Apparently related to
differences in the proportions of the snout is the
position of the infraorbital foramen in T. potens.
In that species the foramen is situated more
anteriorly than in Nimbacinus dicksoni, yet the
maxillojugal suture terminates close to the margin
of the foramen as in T. megiriani and T.

P. F. MURRAY

cynocephalus (Fig. 7). In Nimbacinus, the
maxillojugal suture terminates a considerable
distance posterior to the foramen. This particular
combination of characters suggests that the
condition in T. potens is secondary, or specialised
rather than primitive, having occurred in
conjunction with telescoping of the base of the
rostrum.

Another specialisation in T. potens, now
apparent, is the obliquity and rather large size of
the P l alveolus, which is part of an overall
proportional shift of the cheek tooth row in
relation the rostrum. In Nimbacinus dicksoni, the
P 1 is relatively small and lies in direct line with
P 2 -3 as in T, megiriani and T. cynocephalus
indicating that the condition is derived. However,
the position of the P 3 relative to the canine in T,
potens is like that of Nimbacinus dicksoni in lying
medial to the canine and in being separated from
the canine alveolus by a small bony crest. These
latter features are probably symplesiomorphic,
whereas the relationship of the P 3 to the canine
alveoli in T. megiriani and T. cynocephalus is
probably synapomorphic.

Comparison of the molar morphology among T.
potens, T. megiriani and Nimbacinus dicksoni
shows a similar mixture of plesiomorphic and
derived character states (Fig. 9A-E). In some
aspects, such as molar shape and development of
the stylar shelf, T. megiriani resembles
Nimbacinus dicksoni more than it does T. potens.
In T. potens the stylar shelf is high and wide, yet
appears to show more reduction of the stylar cusps
than T. megiriani and certainly more reduction
than in Nimbacinus in which the stylar cusps D
and E are well-developed on M 1 2 , though cusp E
is absent on M 3 . T. potens shows a much greater
development of the ectoflex on M 2 ~ 3 than either T.
megiriani or Nimbacinus dicksoni. These appear
to be derived characters of the dentition of T.
potens, which however, also shows some
plesiomorphic states relative to T. megiriani and
T. cynocephalus in retaining a well-developed
precingulum on M 2 ~\ in retaining large protocone
bases and in the length of M 2 exceeding the length
of M 3 . T. megiriani is derived relative to
Nimbacinus dicksoni and T. potens in M 3 being
slightly longer than M 2 , in having more slender
and proportionally longer metastylar spurs, in
having reduced protocones and more reduced
paracones, in the loss of the precingulum on M 3
and in having a more obtuse angle between the
preparacrista and the postmetacrista. These
features of T. megiriani trend towards the more
derived states found in the molars in T.

cynocephalus. However, T. megiriani is
plesiomorphic relative to T. potens in retaining a
vestige of stylar cusp E on M 2 and if the structure
on M 3 represents a stylar cusp, it is autapomorphic
(?reversal, eg. Marshall et al. 1994) among all
known thylacinids in that respect.

The heavily worn condition of M' of Thylacinus
megihani renders comparison with the unworn
equivalent of T. macknessi (Muirhead 1992)
somewhat less informative. Thylacinus macknessi
possess a well-developed anterior cingulum,
which is clearly absent in T. megiriani. In T.
macknessi the pre-and postprotocristae do not
ascend the paracone and metacone bases to form
acute crests as they do in all other thylacines
including the M 1 of T. megiriani. The stylar shelf
is well-developed on the M 1 of T. megiriani and
much reduced in T. macknessi. The molar is
broken in a critical position in relation to the
position of stylar cusp D. In T. megiriani, stylar
cusp D is very large and lies immediately adjacent
to the metacone. This suggests that the cusp may
have been reduced in T. macknessi. This
comparison agrees in ail aspects with Muirhead's
(1992) observations that T. macknessi is more
derived in the development of these features than
T. cynocephalus, yet plesiomorphic in the
retention of the anterior cingulum and
autapomorphic in its extension from stylar cusp B
to the base of the protocone.

Stylar cusp homologies among thylacines were
more ambiguous when T. potens was the only
fossil form of any antiquity known (Archer 1982).
The expression of the cusps in Nimbacinus
dicksoni is now well-documented, and accords
with other members of the family Thylacinidae
(Muirhead & Archer 1990). in Nimbacinus
dicksoni, stylar cusps B, D and E are present on
M'~ 2 and stylar cusp D is present on M 3 whereas
cusp E is absent from that tooth. In T. potens,
stylar cusp D is definitely present on M 3 and
almost certainly was present on M'~ 2 , though the
crowns are damaged in the region that the cusps
are usually situated. There is no sign of stylar cusp
E on M 2 ~ 3 in that species. In T. megiriani, stylar
cusp D is well-developed on M 1-3 whereas stylar
cusp E is rudimentary on M 2 . 1 am disinclined to
attribute any phylogenetic significance to the
cusp-like structure in the position of stylar cusp E
on the M 3 of T. megiriani, although its putative
existence is duly noted should future finds serve
to either verify or negate its presence.

In T. cynocephalus, stylar cusps are absent on
M 2-3 in conjunction with the reduction of the stylar
shelf A small stylar cusp occupying the usual

THYLACINUS MEGIR1ANI N.SP.

Ni. dicksoni T. macknessi L potens T, megiriani
1 \ — \ \

phalus

FIGURE 10. Character distribution in species ofThylacinidae. 1-16 are apomorphic, 17-23 are plesiomorphic:
1, loss of 'wall' below postparacrista and premetacrista; 2, transverse narrowing of molars and loss of sty lar cusp
D on M 2 ~ 3 ; 3, widened, elevated stylar crest, suppression of stylar cusp E at least on M 2 ; 4, increased ectoflex; 5,
lengthened premolar diastemata; 6, confluent P'-C diastema; 7, elongation of metastyle, reduced parastyle,
obtuse angle between preparacrista and postmetacrista, M i longer than M 2 ; 8, reduced stylar shelf; 9, reduced
protocone; 10, loss of precingulum and increased concavity of anterior margin of M 3 ; 1 1, reduced ectoflexus; 12,
posterior position of infraorbital foramen above middle of M 2 ; 13, reduced paracone; 14, loss of precingulum on
M 1 ; 15, forward extension of maxillojugal suture to posterior margin of infraorbital foramen; 16, large body size;
17, (plesiomorphic) absence of diastema between P 2 /P 3 ; (plesiomorphic) notched precingulum on M 2 -M 3 ; 19,
(plesiomorphic) P 1 situated lingual to canine; 20, (plesiomorphic) retention of precingulum on M 1 ; (plesiomorphic)
21, large paracone; 22, (plesiomorphic) metaconid on M 3 ; 23 (plesiomorphic) small body size.

topological position of stylar cusp E is present on
M 1 . This cusp was initially identified by Archer
(1982) as stylar cusp E. Subsequently, Muirhead
and Archer (1990) identified the cusp as stylar
cusp D that had become reduced and migrated
posteriorly in conjunction with the elongation of
the metastyle. The predisposition of the stylar
cusps in T. megiriani supports the latter
conclusion, in that if stylar cusp E was present, it
was already extremely reduced relative to stylar
cusp D. Moreover, there is no satisfactory

explanation of the fate of stylar cusp D had cusp E
been retained, as its amalgamation with the
metacone, analogous to Sarcophilus (Crabb 1982)
appears to have been highly unlikely (Stephen
Wroe, pers. com.).

PhyJogenetic position of T. megiriani

The character evaluations employed here are
based on those of Muirhead and Archer (1990).
As the new species exhibits additional structures,

P. F. MURRAY

the character polarities pertaining to those
structures are based on the observations made in
the discussion. An hypothesis of the phylogenesis
of T. megiriani is drawn from the following
postulations (Fig. 10):

Thylacinus cynocephalus-T. megiriani
synapomorphies: lengthened premolar
diastemata, confluent canine-P' alveoli,
posteriorly situated infraorbital foramen (in
line with middle of posterior half of M 2 ),
elongation and narrowing of the metastyle of
M 3 (including M 3 longer than M 2 , reduced
width of parastyle, obtuseness of angle of
postmetacrista and premetacrista and
reduction of height of stylar shelf), greatly
reduced paracone particularly on M 2-3 ,
reduction of diameter of protocone base, loss
of precingulum and concave profile of
anterior surface of M 3 , reduction of ectoflexus
on M 2 " 3 .

Thylacinus potens-T. megiriani-T.
cynocephalus synapomorphies; loss of
precingulum on M 1 , forward extension of
maxillojugal suture to margin of Infraorbital
foramen, possibly large size.
Thylacinus potens-Nimbacinus dicksoni
symplesiomorphies; absence of diastema
between P 2 and P\ presence of precingulum
on M 2 ' 3 , P 1 situated lingual side of canine.
Thylacinus macknessi-Nimbacinus dicksoni
symplesiomorphies: retention of anterior
cingulum on M 1 , large (unreduced) paracone,
possibly small size, remnant metaconid on
M 3 .

Thylacinus megiriani-Nimb acinus dicksoni
symplesiomorphy: expression of stylar cusp
EonM 2 .

Thylacinus megiriani apomorphies: possible
expression of stylar cusp E on M 3 (reversal?),
possible expression of anterobasal cuspule or
small postcingulum on M' 2 (?neomorphic).
Thylacinus potens apomorphies: extreme
development of labial emargination of
molars, stronger ectoflexus on M 2 ~\ forward
position of infraorbital foramen, obliquity and
large size of P 1 , enlarged trigonid and/or
reduced talonid in lowers.
Thylacinus macknessi apomorphies: extreme
reduction of stylar shelf, absence of crests
ascending lingual bases of paracone and
metacone, postparacrista and premetacrista
connect low in the crown basin consequently
the lingual wall below the paracone and
metacone is absent.
Thylacinus cynocephalus apomorphies:

extreme elongation of metastyle on all
molars, extreme reduction of stylar shelf, loss
of stylar cusp D on M 2 " 3 .
Probably because there are so few species and
relatively few characters due to the fragmentary
nature of the fossils, the resulting phytogeny
seems to be fairly straight-forward (Fig. 11).
Among the species considered, Nimbacinus
dicksoni is by far the least derived form. In
contrast, T. macknessi, while retaining more
plesiomorphic features than any of the remaining
species, is simultaneously more specialised than
T. cynocephalus. It is therefore the least likely
candidate for ancestry of any of the three later
species (T. potens, T. megiriani and T.
cynocephalus). T. potens is also highly derived in
its extreme thickening of the labial emargination
of the molars, deep ectoflex, relatively narrow
talonid and possibly in some degree of rostro-
facial proportional sliding. This species shows
subtle modification of the relative proportions of
the snout that are probably secondary because the
forward extent of the maxillojugal suture relative
to the infraorbital foramen appears to be
synapomorphous with T. megiriani and T.
cynocephalus. In several respects, T. megiriani is
the least derived of the large species in retaining a
rudimentary stylar cusp E on M 2 , in retaining
large stylar cusps D on M 1-3 and in retaining a
general proportional similarity in its molars to
those of Nimbacinus dicksoni. However, other
characters of T. megiriani show closer
resemblances to those of T. cynocephalus than
observed in any of the other species.

In general appearance, the differences between
T. megiriani and T potens are quite subtle. Its
large size and robustness, relatively small, broad
molars, large P 3 , presence of stylar cusp D and
deep maxilla all bear a close resemblance to T.
potens, to the extent that when I first saw the
specimen in the field, I thought it might represent
an extreme individual variation of the latter
species. This resemblance biased my initial
restoration of the specimen, although in spite of
this, its detailed differences considerably
outweighed the introduced and genuine
similarities. It is an important feature of the
specimen that while the state of many of its
characters show an unambiguous trend towards T.
cynocephalus, it is probably further removed
structurally from the modern species than it is
from T. potens. A reasonable stage of evolution
estimation for the time of separation of T. potens
and the lineal ancestor of T megiriani would be
in the later part of the mid-Miocene.

THYLACINUS MEGIRJANI N.SP.

T. cynocephalus T. meairiani T. potens
8 7

T. mackness

Ni. dicksoni

FIGURE 1 1 . Subjective dendrogram depicting hypothesis of phylogenetic relationships among thylacinid species;
1, loss of stylarcusp E on M 3 , loss of high postcingulum M'~\ plesiomorphic states of large paracone and small
size; 2, more posterior position of infraorbital foramen; 3, reduced sty lar shelf, loss of 'wall' above postparacrista,
centrocrista and premetacrista, extended precingulum M 1 ; 4, loss of precingulum on M 1 , forward extension of
maxillojugal suture to margin of infraorbital foramen, large size, reduction of stylar cusp E on ?all molars; 5,
widened, elevated labial emargination of molars, increased ectoflex on M 2-3 , loss stylar cusp E on at least M 2 ; 6,
increased length premolar diastemata, reduced stylar shelf, reduced parastyle and paracone, lengthened metastyle,
reduced ectoflex, more posteriorly situated infraorbital foramen; 7, possible neomorphic faint, low postcingulum
on M 1 " 2 , possible stylar cusp E on M 3 ; 8, increased reduction of stylar shelf, increased elongation of metastyle,
stylar cusp D confined to M ! , reduced ectoflex.

Paleobiology and biochronology

The Ongeva Local Fauna shows a distinct
faunal change from the underlying Alcoota LF. A
zygomaturine, close to, if not synonymous with
Zygomaturus gilli is the dominant large herbivore,
apparently replacing Plaisiodon centralis which it
resembles somewhat in size and general
morphology. A species of the crocodile Quinkana
is relatively common, whereas the dominant
Alcoota crocodilian, Bam, is absent. Though the
sample is too small to be definitive, it would

appear that T. megiriani had replaced the Alcoota
LF thylacinid species, T. potens by Ongeva times.
A new small ratite bird, possibly a dromornithid,
is also present in the Ongeva LF (Megirian,
Murray & Wells 1996).

Continuity with the Alcoota LF is indicated by
the presence of Kolopsis torus Woodburne 1 967,
Dorcopsoides sp., Dromornis sp. cf. D. stirtoni
and Ilbandornis sp. The Ongeva K. torus appears
to be little changed from the Alcoota population,
suggesting a fairly short interval between the two
local faunas. Because the depositional

P. F. MURRAY

circumstances of the two faunas differ, it is not
entirely clear as to whether the samples represent
distinct communities within a basically stable
environment or are a reflection of more general,
possibly fairly rapid changes in the biotic
attributes of the region.

Thy/acinus patens is a highly specialised
thylacinid showing as least as much, if not more,
derivation in its molar morphology from
Nimbacinus dicksoni than T. megiriani. The
increased width of the stylar area and relative
widening of the trigonids and narrowing of the
talonids of the lower molars, combined with its
robust skeletal attributes, indicate that T. potens
was capable of shearing and crushing highly
resistant tissues, and may have been somewhat
hyena- or Osteoborus-Yike in its habits. T.
megiriani shows an incipient trend towards the
extreme camassialisation that distinguishes T.
cynocephalus; consequently it may have been
more actively predacious in its habits than T.
potens. It is likely that the two species would have
considerable overlap in their habitat preference
and behaviour. Given their close similarities and
large size, it seems improbable that they would
have occurred sympatrically. Such large predatory
species usually have a considerable range, and it
is therefore a more plausible speculation that T.
megiriani represents a successional replacement
of T. potens rather than an addition to the
previously existing fauna.

and the small -si zed, extreme shearing
modifications of T. macknessi. The addition of
this specimen to the small existing collection of
fossil thylacines seems to help clarify and simplify
our current understanding of the phylogeny of the
group.

Thylacinus macknessi indicates that a highly
derived shearing complex had evolved in the early
to mid Miocene among thylacinids, but did not
lead to the development of the Recent thylacine
species, T. cynocephalus, which appears to have
originated in the late Pliocene. It is likely therefore
that extreme camassialisation in the larger
thylacinid species took place in the late Cainozoic,
perhaps in response to the radiation of
macropodine species that commenced in the early
to mid Pliocene (Flannery 1989). A tempting
speculation, based on the circumstantial evidence
for periodic drought in Alcoota times (Murray &
Megirian 1992) is that there may have been an
abundance of carrion over a sufficient duration to
have selected for a large mammalian scavenging
species. Following subsequent climatic
amelioration in the terminal Miocene, the hyena-
like T. potens was out-competed by the more
predacious T. megiriani. The subsequent loss of
Thylacinus potens as a possible scavenging
predator may have left the somewhat less
productive scavenging niche open, recruiting a
smaller species; namely, the ancestor of the living
Tasmanian Devil, Sarcophilus harrisi.

Conclusions

As in T. potens, the new species has a large P 3 ,
retains stylar cusp D and has transversely wide
molars. T. megiriani differs from T. potens in
many of the features that also serve to distinguish
the latter from T. cynocephalus. These include
reduced ectoflex of M 2 * 3 and stylar shelf on M 3 ,
increase in the length relative to width of M 3 ,
narrow protocones, reduction of P 1-2 relative to the
molars, alignment of the premolar row and
infraorbital foramen opening above posterior M 2 .
T. megiriani shows no definite structural features
that would preclude it from ancestry of T.
cynocephalus and shows a number of
synapomorphous features including reduced
paracone and lengthened postmetacrista. T.
megiriani is less specialised within the
morphological extremes of the large size and
powerful shearing/crushing dentition of T. potens

Acknowledgments

I am indebted to Jeannette Muirhead and Alex
Baynes for their useful comments on the first draft
of this paper. Their observations led to the correction
of a number of important details relating to the
original restoration of the specimen and several key
morphological features of which I was unaware were
generously and courteously pointed out to me. I
thank Dirk Megirian for his outstanding contribution
to all aspects of the excavation, interpretation and
documentation of the Alcoota/Ongeva Locality and,
of course, for giving me the opportunity to describe
and name this interesting species after him. I am very
grateful to Ian Archibald, the NT Museum's
Preparator/Taxidermist who photographed and
printed the bromides under difficult circumstances. I
also thank Rod Wells and the Flinders University
students who participate in the Alcoota excavations
annually. The field work at Alcoota is partially
supported by an NT Heritage Grant.

THYLACINUS MEGIRIAN! N.SP.

References

ARCHER, M. 1982. Review of Miocene thylacinids
(Thylacinidae, Marsupialia), the phylogenctic position
of the Thylacinidae and the problem of apriorisms in
character analysis. Pp 445-76 In 'Carnivorous
Marsupials'. Ed. M. Archer. Royal Zoological Society
of New South Wales: Sydney.

CRABB, P. L., 1982. Pleistocene dasyurids
(Marsupialia) from southwestern NewSouth Wales.
Pp. 512-16 h 'Carnivorous Marsupials'. Ed. M.
Archer. Royal Zoological Society of New South
Wales: Sydney.

DAWSON, L. 1982. Taxonomic status of fossil
thylacines (Thy/acinus, Thylacinidae, Marsupialia)
from Late Quaternary deposits in eastern Australia.
Pp. 527-36 In 'Carnivorous Marsupials'. Ed. M.
Archer. Royal Zoological Society of New South
Wales: Sydney.

FLANNERY, T. 1989. Phylogeny of Macropodoidea; a
study in convergence. Pp. 447-506 In Kangaroos,
Wallabies and Rat Kangaroos. Ed. G. Grigg, P.
Jarman and I. Hume. Surrey Beatty and Sons and the
Royal Zoological Society of New South Wales,
Sydney.

MARSHALL, C, RAFF, E. & RAFF, R. 1994. Dollo's
law and the death and resurrection of genes.
Proceedings of the National Academy of Science,
USA. 91:12283-12287.

MEGIRIAN, D., MURRAY, P. & WELLS, R. 1996. The
Late Miocene Ongeva Local Fauna of Central
Australia. The Beagle, Records of the Museums and
Art Galleries of the Northern Territory. 13: 9-38.

MUIRHEAD, J. & ARCHER, M. 1990. Nimbacinus
dicksoni, a plesiomorphic thylacine

(Marsupialia:Thylacinidae) from Tertiary deposits of
Queensland and the Northern Territory. Memoirs of
the Queensland Museum 28(1): 203-221.

MUIRHEAD, J. 1992. A specialised thylacinid,
Thylacinus macknessi, (Marsupialia: Thylacinidae)
from Miocene deposits of Riversleigh, Northwestern
Queensland. Australian Mammalogy 15: 67-76.

MURRAY, P. & MEGIRIAN, D. 1992. Continuity and
contrast in middle and late Miocene vertebrate
communities from the Northern Territory. The Beagle,
Records of the Northern Territory Museum of Arts
and Sciences 9(1): 89-110.

MURRAY, P., MEGIRIAN, D. & WELLS, R. 1993.
Kolopsis yperus sp. nov. (Zygomaturinae,
Marsupialia) from the Ongeva Local Fauna: new
evidence for the age of the Alcoota fossil beds of
central Australia. The Beagle, Records of the
Northern Territory Museum of Arts and Sciences
10(1): 155-172.

STIRTON, R. A. 1967. New species of Zygomaturus
and additional observations on Meniscolophus,
Pliocene Palankarina Fauna, South Australia. Bureau
of Mineral Resources, Geology and Geophysics,
Australia, Bulletin 85: 129-150.

WOODBURNE, M. O. 1967. The Alcoota Fauna,
Central Australia: an integrated palaeontologicai and
geological study. Bureau of Mineral Resources,
Geology and Geophysics, Australia, Bulletin 87: 1-
187.

SOUTH

AUSTRALIAN

MUSEUM

VOLUME 30 PART 1
JULY 1997
ISSN 0376-2750

CONTENTS

ARTICLES

1 R. V. SOUTIICOTT

Description of two new

13 J. OKUNO

A new shrimp of the genus Rhynchocinetes from the Great Australian Bight
(Crustacea: Dccapoda: Rhynchocinetidac).

19 S. PICHELIN

Pomphorhynchus heronensis sp. nov. (Acanthoccphala: Pomph
Lutjanus carponotatus (Lutjanidac) from Heron Island, Australi;

udae) trom

29 L. F. QUEALE

Field Identification of female iittlc brown bats Vespadelus spp. (Chiroptera:
Vcspertilionidae) in Souti

35 C. H. S. WATTS

Four new species of Antiporus Sharp (Colcoptera, Dytiscidae) from Australia, with
notes on A. femoralis (Boh.) and A. intcrrogationis (Clark).

43 P. F. MURRAY

Thylacinus megiriani, a new species of thylacine (Marsupialia
the Onseva Local Fauna of Central Australia.

ylacinidae) trom

Published by the South Australian Museum,
North Terrace, Adelaide, South Australia 5000

SOUTH

AUSTRALIAN

MUSEUM

\JtnE .IT MR ^fi

JANUARY

A NEW SPECIES OF MULTIPHASED CORBULIPORA MACGILLIVRAY, 1895
(BRYOZOA: CRIBRIOMORPHA) FROM SOUTHWESTERN AUSTRALIA.

P. E. BOCK & P. L. COOK

Summary

A new Recent species of the cribriomorph genus Corbulipora, C. inopinata, is described from
several localities in the western part of the Great Australian Bight. All species of Corbulipora, both
Recent and fossil, are known to occur in several phases. The form of the subcolonies, and the
characters of the component zooids may be very different in each phase, and subcolonies are
capable of separate existence. C. inopinata occurs in an encrusting phase, in a flustriform, ovicellate
phase, and a bilaminar rooted phase. All zooids have some form of cribriomorph frontal shield,
unlike some of those of the closely related C. tubulifera. The shield has, however, different
characters in each phase.

A NEW SPECIES OF MULTIPHASED CORBULIPORA MACGILLIVRAY, 1895
(BRYOZOA: CRIBRIOMORPHA) FROM SOUTHWESTERN AUSTRALIA.

P. E. BOCK & P. L.COOK

BOCK, P. E. & COOK, P. L. 1998. A new species of multiphased Corhulipora MacGillivray,
1895 (Bryozoa: Cribriomorpha) from southwestern Australia. Records of the South Australian
Museum 30(2): 63-68.

A new Recent species of the cribriomorph genus Corbulipora, C. inopinata, is described
from several localities in the western part of the Great Australian Bight. All species of
Corhulipora, both Recent and fossil, are known to occur in several phases. The form of the
subcolonies, and the characters of the component zooids may be very different in each phase,
and subcolonies are capable of separate existence. C. inopinata occurs in an encrusting phase,
in a flustnform, oviccllate phase, and a bilammar rooted phase. All zooids have some form of
cribriomorph frontal shield, unlike some of those of the closely related C. tubulifera. The shield
has, however, different characters in each phase.

P. E. Bock, Royal Melbourne Institute of Technology, Latrobe Street, Melbourne, Victoria
3001, and P. L. Cook, Associate, Museum of Victoria, Swanston Street, Melbourne, Victoria,
3000. Manuscript received 2 April 1997.

The occurrence of distinct phases of correlated
colony growth form and zooid morphology in
species of Corbulipora has been described in
some detail by Bock and Cook (1994; in press).
Essentially, the function of the phases appears to
be the same in all species. The first, ancestrulate
phase is minute, encrusts shells and shell
fragments, and may even have an interstitial
existence. It establishes the colony and results
from the settlement and metamorphosis of a
motile larva. The second, erect phase is cellularine
or flustrine, and arises from stalk-like kenozooids
and autozooids growing from the peripheral pore-
chambers of the zooids of the primary phase. This
phase develops ovicells and large interzooidal
avicularia. The second phase gives rise in various
ways to a small, bilaminar or frontally budded
third phase, which is known, in Recent species, to
be anchored by numerous rhizoids. As the
ovicelled second phase tends to be thinly calcified,
the delicate subcolonies may easily be destroyed,
or at least detached from their origins. The
bilaminar phase maintains the position of the
colony and develops further ovicellate
subcolonies, which may be able to alternate with
it more than once (Bock & Cook in press).

Collections of bryozoans, made by the R.V.
Franklin, using an epibenthic sled, from localities
in the western Great Australian Bight in July
1995, have produced a wealth of specimens and

species. These include subcolonies of C.
tubulifera and of an undescribed species of
Corbulipora, which has several distinctive
features.

Materials And Methods

Specimens are stored in the Collections of the
Museum of Victoria (MOV) and South Australian
Museum (SAM). Specimens for scanning electron
microscopy were coated with gold.

The Stations from which specimens of
Corbulipora were obtained were as follows:

C. inopinata sp. nov.

GAB 118 34°59'S, 1 19°00'E, 85m. Young non-
ancestrulate colonies encrusting worn adeonid and
lunulite colonies of bryozoans.

GAB 098 34°39'S, 122°26'E, 156m. 1 phase-2
subcolony with numerous avicularia.

GAB 093 34°32'S, 122°58'E, 95m. 1 phase-1
subcolony and 1 phase-2 subcolony with ovicells.

GAB 083 34°21'S, 124°08'E, 180m. 2 phase-1
subcolonies encrusting Turhtella shells.

GAB 049 33°53*S, 125°22'E, 156m. 4 phase-2
subcolonies, developing into phase-3 at tips, one
repeating phase-2.

GAB 056 33°19'S, 125°43'E, 73m. 2 large phase-

P. E. BOCK & P. L. COOK

2 subcoionies, one with ovicells, developing from
phase-3 subcoionies.

GAB 045 33°25'S, H$*5WE, 143.5m. 3 small
phase-3 subcoionies with stalks.

GAB 020 33°20'S, 129°18'E, 157m. 9 small
phase-2 subcoionies. 2 developing from phase-3
subcoionies.

GAB 013 33°06*S, 130°00'E, 101m. 2 phase-2
subcoionies, one with ovicells, and one phase-3
with numerous rhizoids and one phase-2
subcolony.

GAB 014 33°16'S, 130°00'E, 155m. 2 phase- 3
subcoionies developing into phase-2 subcoionies,
plus 5 isolated stalks.

C. tuhulifera (Hincks)

GAB 019 33°22'S, 129°19'E, 301m. 7 phase-2
subcoionies, and 2 phase-3 subcoionies.

Systematics

Order Cheilostomatida Busk, 1852

Superfamily Cribrilinoidea Hincks, 1879

Family Cribnlinidae Hincks, 1879

Genus Corbulipora MacGillivray, 1895

Corbuiipora MacGillivray, 1895:60; Wass
1975:168; Bock & Cook, 1994:33; in press.

Corbulipora inopinata sp.nov.

(Figs 1-6)

Material Examined: HOLOTYPE MOV,
F80665, GAB Stn 049, subcolony including
phase-2 developing into phase-3 and then
repeating phase-2.

FIGURES 1-2. Corbulipora inopinata sp.nov. 1, subcolony showing phase-2 autozooids and two ovicelled zooids
at the proximal end (arrowed). Intermediate zooids leading to two astogenetic generations of phase-3 zooids at the
growing tip (GAB Stn 049) X22; 2, phase-3 subcolony with marginal pore chambers and autozooids with occluded
orifices (GAB Stn 020) X40.

NEW SPECIES OF MULTIPHASED CORBULIPORA

FIGURES 3-4. Corbulipora inopinata sp.nov. 3, phase-2 zooids with ovicells and simple frontal spines (GAB Stn
049, see also Fig.l). X80; 4, phase-3 zooids with complex frontal shields (GAB Stn 049, see also Fig.l) X70.

Paratypes: Rest of material
L755, GAB Stn 049 part.

including SAM

Description

Corbulipora with colony including three
subcolony phases. Primary phase encrusting,
ancestrula not seen; primary triad zooids with
frontal shield of 16 simple spines. All zooids
with a distinct, smooth gymnocyst, and
communicating through small pore-chambers.
Remaining primary-phase zooids with frontal
shields with 16-18 spines, separated by 3-4
series of small lacunae and with two concentric
series of raised pelmatidia. Calcified orifice with
3 spines, lateral pair often bifid. Second, flustrine
phase inferred to arise from primary phase.
Fronds developing from stalk kenozooids and
elongated autozooids, often in quadriserial
groups; series bifurcating to form fronds 10-12
zooids wide and 150 astogenetic generations
long, bifurcating up to 4 times. Autozooids with
distinct gymnocyst and 14-16 flattened spines

overarching the frontal membrane. Brooding
zooids with enlarged, curved, sometimes
medially fused oral spines and large
hyperstomial ovicell. Ovicell frontal with median
suture and paired entooecial areas. Aviculana
scattered, rare in presence of ovicells. Rostrum
elongated, narrows not raised distally; slightly
expanded and rounded terminally, mandible
hinged on a delicate bar, orientated distally. Tips
of fronds with 3^4 astogenetic generations of
zooids with intermediate morphologies leading
to the third, bilaminar phase subcolonies. These
form small, expanded groups of zooids with
shields similar to phase-1 subcolonies, with two
concentric series of raised pelmatidia. Zooids
have a small orifice, sometimes partially
occluded by the fusion of enlarged oral spines.
Numerous rhizoids develop from marginal pore-
chambers, together with stalk kenozooids and
elongated stalk autozooids with a small opesia
and marginal spines, which are the origin of
repeated subcolonies of phase-2.

P. E, BOCK & P. L. COOK

JW^

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-f ■

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■

▼ 1^^^

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FIGURES 5-6. Corbulipora inopinata sp.nov. 5, phase-2 autozooids, note enlarged pair of distal spines (GAB Stn
098) XI 00; 6, phase-2 autozooids and avicularium (GAB Stn 098) X80.

Etymology'

fnopinus L. - - unexpected: referring to the
occurrence of a second Recent species of
Corbulipora with multiphase growth.

Remarks

The three phases of C inopinata resemble those
of C. tuhulifera closely, but differ in several
respects. The most noticeable difference is in the
occurrence of a spinous shield in the autozooids of
the second, flustrine phase. The spines are
flattened and generally do not fuse centrally,
although the enlarged suboral spines of marginal
zooids may be fused. In addition, the avicularia of
C inopinata are much narrower, and are not
raised distally, as are those in C. tuhulifera, and
the mandibles are hinged on a delicate, but
complete bar. The palate is much longer in
proportion to the proximal gymnocyst than in C.
tuhulifera. Other differences are small, but
distinct and consistent. Although no ancestrula
has been preserved, one of the small, phase- 1
colonies encrusting Turhtella from Stn GAB 083

possesses a post-ancestrular triad of very small
zooids. These all have a shield of simple spines,
very similar to the one post-ancestrular zooid of
the triad in C, ornata (Bock & Cook, in press,
fig. 1). Subsequent zooids have two series of
pelmatidia, which do not develop until later in
astogeny in C. tuhulifera. The orifices of this
stage are also wider than those of C. tuhulifera.
No stalk zooids have been preserved arising from
the encrusting phase m C inopinata, but by
analogy with C. tuhulifera, these are presumed to
be identical with those which arise from phase-3
bilaminar subcolonies. These consist of one or two
pairs of kenozooids 2.5mm long, often
accompanied or succeeded by one or more pairs of
elongated zooids 2mm long, each with a small
opesia surrounded by 8-12 short, simple spines.
Thus the stalks differ from those of C. tuhulifera
in being more robust, with more zooidal elements,
and in the occurrence of autozooids with an
opesia, rather than kenozooids, early in astogeny.
The zooids of intermediate morphology at the tips
of the fiustnne phase-2 fronds attain a complete

NEW SPECIES OF MULTIPHASED CORBULIPORA

shield of phase-3 autozooids within 4 astogenetic
generations. First, the flattened spines fuse
medially, leaving a row of small lacunae. A single
series of pelmatidia develops on the spines of the
next generation of zooids, and a paired series, with
3-4 lacunae, develops on the shields of
subsequent generations. The small calcified orifice
is complete within the next generation or two.
Although the general appearance of the autozooids
of the bilaminar phase is very like those of C.
tubulifera, they have a smaller orifice. Although
these are often partially occluded by the oral
spines, they do not usually exhibit the series of
changes resulting in complete closure of the
orifice which occurs in C. tuhulifera (Wass 1975).
In one specimen from GAB Stn 020, the spines
are enlarged and nearly occlude the orifice (Fig.

2).

One of the specimens from GAB Stn 013
illustrates the relationship and probable functions
of phase-2 and phase-3 subcolonies particularly
well. It consists of a small bilaminar phase-3
subcolony comprising approximately 60
autozooids. More than 50 rhizoids arise from the
marginal pore-chambers, and on one face of the
subcolony these are involved with and adhere to
minute shell and bryozoan fragments. On the edge
of the other face, one quadriserial stalk, 15mm

TABLE 1. Comparative measurements in mm of
Corbulipora inopinata sp.nov. and C. tubulifera
(Hincks). Lz, lz length and width of autozooid; lo, width
of orifice; Lov, lov, length and width of ovicell; Lav, lav,
length and width of avicularium; Lp, length of
avicularian palate.

Character

C. inopinata

C. tub\

ulijera

Phase 1

(including primary

zooids in mms)

0.41 -0.70

0.45-

-0.74

0.35-0.50

0.38-

-0.50

0.12-0.17

0.10-

-0.13

Phase 2

0.98-1.30

0.78-

-1.32

0.26-0.29

0.25-

-0.33

Lov

0.21-0.23

0.23-

-0.27

lov

0.24-0.26

0.28-

0.33

Lav

0.60 - 0.75

0.65-

-0.96

lav

0.15-0.17

0.22

0.26

0.43-0.51

0.31

0.35

Phase 3

0.50-0.75

0.54

0.87

0.37-0.50

0.32

-0.54

0.09-0.10

0.10

0.14

long, consisting of two generations of quadriserial
kenozooids and 10-12 generations of elongated
autozooids, gives rise to a phase-2 flustrine
subcolony about 30mm long, with more than 40
astogenetic generations. It seems almost certain
that the one face of the bilaminar phase was
buried in the surface sediments, whereas the
flustrine frond extended above the surface. This
subcolony is quite small; two flustrine subcolonies
from Stn 056 extend up to 70mm and include
10,000 to 12,000 zooids, many of which are
ovicelled. These would protrude well above the
surface of the bottom sediment, allowing the
larvae to be released easily.

The similarities in phase structure found in C.
inopinata and C. tubulifera reinforce the
inferences previously made by Bock and Cook
(1994; in press) as to the nature and relationships
of the known phases in C. ornata and C.
suggerens, from the Victorian Tertiary. They also
strongly suggest that the ovicellate phase of C,
suggerens was a thinly calcified cellularine
subcolony type, which has not been preserved as a
fossil.

C. inopinata occurs in the western Bight
together with C. tubulifera, but from shallower
waters. In Bass Strait, and from Tasmania, C.
tubulifera is known from a wide range in depth
from 40m to 800m. It appears to be at the limits
of its range south east of Eucla, in deep water.
Wass (1975) first described Corbulipora
oriparma, which is the bilaminar phase of C.
tubulifera, from three localities. The holotype was
from northeast Tasmania, the other specimens
were from the eastern part of the South Australian
coast. Subsequently, Wass and Yoo (1983)
recorded C. oriparma from 12 localities,
extending from near Perth to southwest of Eucla.
These are similar to many of the localities from
which specimens have been recently examined,
and which have produced C. inopinata. None of
Wass and Yoo's (1983) localities was from a
greater depth than 158 m, far shallower than the
most westerly known locality for C. tubulifera.
Only examination of the specimens recorded by
Wass and Yoo (1983) can decide their identity.

Acknowledgments

We should like to thank Dr Yvonne Bone (University
of Adelaide) and the Master and crew of CSIRO R.V.
'Franklin' for the opportunity for one of us (P.E.B.) to
take part in the sampling programme in July 1995,
which led to the discovery of the species described here.
We are also grateful to Mr D. McDonald for his help in
preparation of this paper.

P. E. BOCK & P. L. COOK

References

BOCK, P. E. & COOK, P. L. 1994. Occurrence of three
phases of growth with taxonomically distinct zooid
morphologies. Pp 33-36 in Hayward, P. J., Ryland, J.
S. & Taylor, P. D. (eds): Biology and Paleobiology
of Bryozoans. Olsen & Olscn: Fredensborg.

BOCK, P. E. & COOK, P. L. in press. A revision of the
genus Corbulipora MacGillivray (Bryozoa:
Cribriomorpha). Memoirs of the Museum of Victoria.

BUSK, G. 1852. Catalogue of Marine Polyzoa in the
Collection of the British Museum. Part 1
Cheilostomata (Part).: viii, 1-54, vi. Trustees of the
British Museum, London.

HINCKS, T. 1879. On the classification of British
Polyzoa. Annals and Magazine of Natural History
(5)3:153-164.

MACGILLIVRAY, P. H. 1895. A Monograph of the
Tertiary Polyzoa of Victoria. Transactions of the
Royal Society of Victoria, 4:1 166.

WASS, R. E. 1975. A revision of the Bryozoan Genus
Corbulipora MacGillivray. Proceedings of the Royal
Society of Victoria 87:167-173.

WASS, R. E. & YOO, J. J. 1983. Cheilostome Bryozoa
from the Southern Australian Continental Shelf.
Australian Journal of Marine and Freshwater
Research 34: 303-354.

THE ARCHAEOLOGY OF LAKE SYSTEMS IN THE MIDDLE COOPER
BASIN, NORTH-EASTERN SOUTH AUSTRALIA.

ELIZABETH WILLIAMS

Summary

This paper presents the final report of a study of the archaeology of a number of lake systems
located in the middle section of the Cooper Basin and neighbouring areas near Innamincka, South
Australia. The paper outlines the results of archaeological fieldwork carried out during 1987 in one
area within the region and relates this to other research undertaken in 1986 and 1989 in other parts
of the middle of the Basin.

THE ARCHAEOLOGY OF LAKE SYSTEMS IN THE MIDDLE COOPER BASIN
NORTH-EASTERN SOUTH AUSTRALIA.

ELIZABETH WILLIAMS

WILLIAMS, E. 1998. The archaeology of lake systems in the middle Cooper Basin, north-
eastern South Australia. Records of the South Australian Museum 30(2): 69-91.

This paper presents the final report of a study of the archaeology of a number of lake systems
located in the middle section of the Cooper Basin and neighbouring areas near Innamincka,
South Australia. The paper outlines the results of archaeological fieldwork carried out during
1987 in one area within the region and relates this to other research undertaken in 1986 and
1989 in other parts of the middle of the Basin.

E. Williams, World Heritage Unit, Department of Environment, Sport and Territories, Canberra,
Australia Capital Territory, 2601. Manuscript received 3 1 June, 1995.

The lakes forming the main focus of this study
are located within an extensive linear dunefield,
the Strzelecki desert, which forms part of
Australia's continental dunefield. Although
archaeological research has been undertaken along
the main channel of the Cooper in the middle
Cooper basin, particularly the Innamincka area
(Hughes & Lampcrt 1980, McBryde 1987), the
archaeology of the region's series of lake systems
had not been studied until this research
programme. Given that at least a number of the
lakes hold fresh water for some considerable time,
and that this is a strikingly unusual feature of arid
areas generally, it is very likely that the lakes
would have been the major focus of occupation for
at least the recent period and possibly for extended
periods in the past. As well, it is possible that
landforms associated with the lakes such as
source-bordering dunes preserve information
about Pleistocene climates and changing
environmental conditions, and possibly also about
Pleistocene human occupation. A study of the
lakes therefore had the potential to contribute to
our knowledge of not only the recent occupation
of the region but also that of past settlement, and
possibly, the wider question of how Australia's
arid zone itself was occupied.

In structuring the project, a preliminary analysis
of the region revealed considerable variation of
the nature and types of lakes. The study was
designed to examine this variability in relation to
human settlement. The study was designed as
follows. I examined the archaeology of examples
of the three major types of lake systems in the
region. These are the semi-permanent lakes of the
Coongie system, fed by the Cooper's North- West

Branch; small isolated salt lakes marking the edge
of the Cooper Basin, and a series of ephemeral
lakes to the north of the Basin fed by local
drainage and precipitation. The results of this
work are presented below. In the latter sections of
my discussion I compare this information with
that obtained from sites located in a relatively
waterless core of the dunefield located in the
southern part of the Strzelecki desert. The
concluding part of the paper outlines a number of
speculative hypotheses about the occupation of the
region and of Australia's arid zone in general.

Work carried out in the Coongie lake system
has previously been reported on in Williams
(1988) and that undertaken in the Strzelecki
dunefield summarised in Smith, Williams and
Wasson (1991). The current study draws this
research together to present a comprehensive view
of the region's archaeology.

General Environmental Features Of The Region,
Including Those Important For Human Occupation

The study area lies in one of the most arid parts
of Australia. The region is covered by an extensive
linear dunefield and receives the lowest rainfall
reading of any area of the continent (125 mm per
annum). Despite the aridity there are reliable
water sources here in the form of the Cooper
Creek channel, its overflow system - the North-
West Branch, and associated lakes fed by the
channel systems. The amount of water in this
drainage complex does not reflect local rainfall
patterns and predominantly derives from received
precipitation falling many hundreds of kilometres

E. WILLIAMS

to the north-east during the wet season in the
upper parts of the Cooper catchment in north-west
Queensland.

Given the arid setting of the region, water was a
valuable resource and one of the most important
features for human occupation. This is confirmed
by the distribution of archaeological material
within the Coongie lake system - site size and
stone artefact density increases in relation to the
location of permanent and semi-permanent water
sources (Williams 1988: 59).

Another important feature for occupation was
the availability of good quality stone suitable for
artefact manufacture. Stone sources are extremely
localised here and are either scarce or entirely
absent within most of the linear dunefield itself.
Some stone is available in the form of pebbles
carpeting the surface of the gibber plains or 'stony
deserts' located on the edges of the study area but
this source only exists in the form of small
nodules, not necessarily suitable for all types of
artefact manufacture. The most extensive areas of
stone are found in outcropping areas of hard rock
within areas of dissected tablelands located on the
margins of the study area.

Although the availability of resources such as
water and stone were crucial, the two resources
do not always coincide geographically. In the area
immediately surrounding Innamincka where are
found the extensive deep waterholes of the main
Cooper channel and the abundant good quality
stone resources of the locality's mesas and
dissected tablelands, the two resources are
abundant and lie close together. In the Coongie
lakes system located within the stone-free linear
dunefield, although water resources are extensive,
good quality stone lies at least 60 km away.
Therefore one of the major features of the
archaeology of the region relates to how people
balanced their need for stone with that for water
and how this varied from one area to another.
Because this feature is responsible for much of
the patterning of the archaeology of the Basin and
neighbouring areas, it is the major focus for this
study.

A second and related focus concerns the
antiquity of human occupation of the region. This
topic is reported on in detail in Smith, Williams
and Wasson (1991) and is also discussed here.

The structure of the paper is as follows. Given
that the environment places such strong
constraints on human occupation in the region, a
general overview of climatic and geographic data
is presented first, including material on
environmental chronologies. This is followed by a

brief summary of historical accounts of how
Aboriginal people were observed to be using the
area at the time of European contact, in order to
present a picture of the occupation of the area at
least for the recent past. The paper then reports on
archaeological sites investigated, including an
analysis of stone artefacts from a selected sample
of sites. I conclude with a brief discussion of some
of the theoretical implications of my work.

The Contemporary Environmental Srtting

A number of the environmental parameters for
the region have already been presented in
Williams (1988). This section does not repeat this
material and instead provides more detailed
information on localities previously discussed and
new data on areas not referred to in other
publications.

The main focus of the study area is a series of
lake systems within the Strzelecki desert. On the
east and west the Strzelecki is bounded by stony
country, comprising gibber plains and areas of
dissected tablelands. The latter consists of the low,
but locally steep escarpments and small mesas of
the geological features known as the Innamincka
and Cordillo domes.

The aridity of the area has been noted above.
Precipitation in the region is unpredictable and
erratic, with no systematic seasonal patterning.
Evaporation is extremely high, at least one order
of magnitude greater than precipitation (Reid and
Gillenl988: 16).

Three of the lake systems which form part of
the study area are discussed in some detail here.
They are the freshwater lakes of the Coongie
system, a ring of salt lakes which mark the
boundary of the Cooper drainage system, and
another chain of lakes which lie to the north of the
system and to the west of Cordillo Downs station.
For convenience I refer to this latter group of lakes
as the 'Cordillo lakes'.

The Coongie system

The Coongie system comprises a series of large,
shallow, irregularly-shaped lakes lying within the
linear dunefield. The lakes are fed by a tributary
of the Cooper, the North- West Branch. The system
has a complex hydrology and is described here in
its constituent sections from south to north.
Closest to the North-West Branch feeder channel
are three lakes: Coongie, Marroocoolcannie and
Maroocutchanie, which are fed directly from the
North-West Branch. Two lakes lying further

ARCHAEOLOGY OF THE MIDDLE COOPER BASIN

JUT

Stony desert
(Cordillo Dome)

* Cordillo Downs if]

Stony desert

■ \K l f\A !\v\\\\0*to Lake Koonoomoorinna ,} v \ ' L 1 ' ^h>\sVi' i*
t ffy \y%TiIlie Lakes y\ \ \\\\\\ \M*> I wllUw \

Stony desert
(Innamincka Dome)

FIGURE 1. Regional map of north-east South Australia showing the sites mentioned in the text

E. WILLIAMS

^5 Salt Lakes ^"^ Source bordering dunes

FIGURE 2. Map of the Coongie and Cordillo lake systems.

ARCHAEOLOGY OF THE MIDDLE COOPER BASIN

north, Toontoowaranie and Goyder, are fed by
interconnecting creeks from overflow from the
first three lakes. To the north again are a further
three lakes: Marradibbadibba, Lady Blanche and
Sir Richard, which fill from overflow from Lake
Goyder. In this paper I refer to all of the lakes
collectively as the 'Coongie system'.

A further series of lake and channel systems
also have some connection either to the Coongie
system or the North-West Branch channel. It is
noted that although they do hold water at certain
times (as observed by the explorer, John McKinlay
in 1862), the Coongie system, at least in the recent
past, held the most extensive and most reliable
sources of water. Little is currently known about
their hydrology and for this reason the lakes do
not form part of this study.

The North-West Branch which joins the Cooper
west of Innamincka diverts most of the flow of
the main Cooper channel northwards. For the area
west of Innamincka relatively little water thus
flows down the main Cooper channel on to Lake
Eyre (Mike Steele, Innamincka store, pers.
comm.). The Coongie system therefore catches
most of the water coming down the Cooper
channel. Flow down the Cooper derives from two
sources: local precipitation and received rainfall
which falls as far as 500 km upstream in the
channel country of western Queensland. This
latter precipitation falls mostly during the summer
monsoon and contributes the major portion of the
Cooper's flow. In contrast to local rainfall its
timing is relatively predictable. Water from the
summer monsoon rains takes about three months
to come down the channel, first reaching the
Coongie about late autumn (Barry Saunders,
Innamincka region, pers. comm.).

Despite this level of predictability, the amount
of water which actually comes down the channel
and the exact time of year when it arrives
markedly fluctuates in relation to variability in the
timing and effectiveness of rainfall in the upper
Cooper catchment. Nonetheless, there are
predictable water sources in the form of deep,
permanent waterholes which are often a couple of
kilometres in length, located along the channels
of the main drainage systems such as the Cooper
and the North- West Branch.

With the possible exception of Coongie, the
lakes do not hold water on a permanent basis.
Oral tradition amongst current residents of the
Innamincka region suggests that Coongie Lake
itself holds water for most of the year, although
the nature and amount of water it holds is difficult
to quantify because of a lack of historical records.

The other lakes in the system hold water for
months at a time but the extent and duration is
highly variable and depends on a number of
factors. These include the amount of water coining
down the Cooper from rainfall in the northern
catchment, local precipitation, the position of the
lake relative to the North-West Branch and the
season of the year (evaporation is highest during
summer).

Despite variability in the amount of water held
in the Coongie system, the presence of often
extensive supplies of fresh water in an otherwise
extremely arid environment is extremely rare
within Australia. The region consequently
supports a very productive and diverse ecology,
but one which varies in relation to water levels in
the lakes and channels. The variability does not
decrease overall productivity, and in fact is
responsible for the high biodiversity of the region.
The chain of lakes, with its complex sequence of
rising and falling water levels is biologically
highly productive. The lakes provide a diverse
range of constantly changing habitats and have
the potential to support the largest number and
widest range of resources in the region. The major
resources found here are: birds, especially
waterbirds; fish, crayfish and mussels; dry-land
and aquatic plants, and some mammal species.

Biological studies of the system reveal this very
high productivity (Reid and Gillcn 1988, Reid and
Puckeridge 1990). Reid and Gillen for example
recorded 161 separate bird species in the Coongie
system and estimated that during the year under
study (1986 - a wet year when all lakes in the
system held water for much of the time) 20 000
waterbirds permanently occupied the lake chain.
(1988: 184, 198).

Because of the interconnectedness of the
system, rising and falling levels in one lake affect
other lakes in the chain. Given that most of the
plant and animal species in the region are oriented
to specific environmental niches, variation in
water levels consequently provides a much wider
range of habitats than would otherwise be
available along a permanent waterhole where
water levels remain relatively constant. For
example, although water levels in one lake may
begin to fall with evaporation, certain species of
waterbirds whose habitat is that of more shallow
water will migrate in to the lake while those
whose habitat is that of deeper water will move to
another lake where water levels are rising. The
differential fluctuation of water levels means that
the lakes have the potential to carry the largest
biomass of plants and animals in the region

E. WILLIAMS

because they are providing a wide range of
ecological zones. The permanent waterholes on
channels and rivers carry a relatively lower
biomass because their environments are much less
variable. Water levels fluctuate significantly less
in these deep waterholes, supporting a smaller,
but more predictable range of resources.

Salt lakes or 'salinas '

These irregular-shaped lakes lie in a ring along
the northern and western boundary of the Cooper
drainage basin and mark the boundaries of that
system. They are located in the midst of the linear
dunefield and are deflation hollows cut to
groundwater level. This type of feature has been
termed a 'salina' (Macumber 1980). Their salinity
derives from the fact that in the region the
groundwater is saline. Although no hydrological
studies of the salinas in this area have been
undertaken, it appears that for the recent past at
least, they are permanently saline. In contrast to
the Coongie system they therefore support
relatively few plant and animal resources.

The Cordillo lakes

To the north of the Coongie system, beyond the
salt lakes, lie another series of irregularly-shaped
lakes. As with the salinas, no formal studies of
these have been undertaken and there are no
records of how often they fill.

Cartographic and field observations of the
locality suggest that there are at least ten lakes in
the system, connected by a series of channels. It
appears that they fill from local run-off from the
stony country of the Cordillo dome to the east.
Their hydrology, unlike the Coongie system,
therefore reflects local precipitation. The lakes
were dry when fieldwork in the area was
undertaken (1987), despite the years 1986 - 1987
having been much wetter than average in terms of
local rainfall. This suggests that considerable local
precipitation is required to fill them and they thus
remain dry for very long periods of time. They are
not salinas though, and small pools of fresh water
were present in 1987 in a few places along the
channels linking the lakes. The predominantly dry
character of the lake system suggests that it
supports relatively few plant and animal resources
in comparison to the Coongie, but the presence of
some areas of freshwater indicates that more
resources are supported than the saline lakes.

The region's current environment is not
necessarily the same as today as in the past. Over
time there have been significant changes in
climate, hydrology and landforms, with

consequent effect on the human occupation of the
region.

Past Environments - Dunefield And Floodplain
Research

Much of the information on past regional
climates has been summarised and presented
elsewhere (Williams 1988) and so here I briefly
refer to this work and also discuss new data which
has arisen as a result of the study. R. Wasson has
developed an environmental chronology for the
region, based on stratigraphic sequences derived
from longitudinal dunes and floodplain sediments
(for references see Williams 1988). A summary of
his findings and a discussion of their implications
for the hydrology of the lakes follows.

Wasson's work indicates that before 22 000
years ago the Cooper had a higher velocity than
today and was flooding out to a much greater
extent. Although geomorphological studies of the
Coongie lakes themselves have not been
undertaken, Wasson's data suggest that if the
Coongie system was in operation during this
period it would have held considerably more water
than today. As well, groundwater levels for the
region as a whole would have been higher,
indicating that the lake depressions which are now
salinas may have held fresh water, Current work
does not indicate whether local precipitation was
also high at this time, so that it is not possible to
model processes which may have been occurring
in the Cordillo lakes.

Around 22 000 years ago, the Cooper's velocity
decreased, Hooding of the outer floodplain
decreased and it is likely that the water table
lowered as well. Consequently it is likely that what
are now salinas would have changed from fresh
water at this time. Lake levels in the Coongie
system may have become lower. About 20 000
years ago conditions were windier and more arid,
activating a period of dune building throughout the
linear dunefield. The dune sediment was drawn
from the deposits formed from the continuing,
reduced flooding of the outer floodplain. These
climatic conditions and the consequent dune
building persisted until around 16 000 years ago.
At about 1 2 500 years ago the flooding-out of the
outer floodplain ceased, removing the sediment
available for dune building.

Dune building began again in the later
Holocene, although on a smaller scale than in the
late Pleistocene. Wasson (1986: 79) notes that this
event is enigmatic, since in contrast to the late

ARCHAEOLOGY OF THE MIDDLE COOPER BASrN

Pleistocene there seems to have been no climatic
change occurring on a scale large enough to be
able to mobilise sediments. He has speculated that
it is possible, although not yet conclusively
demonstrated, that the phenomenon could be
related to more intensive human occupation of the
region. If such intensive occupation took place,
the erosion resulting from Aboriginal firing of
vegetation to aid the procurement of food
resources, together with clearing of trees and other
vegetation to provide fuel for campfires and earth
ovens, may have caused greater mobility in
sediments.

Past Environments - Studies Of Lakeshore
Features

The environmental data presented above is
derived from a study of dunefield and floodplain
sediments and presents a broad regional
chronology. Is it possible to develop a chronology
for the lake systems themselves?

A formal study of the geomorphology of the
lake systems was beyond the scope of this study
so I developed a smaller project, more
complementary to the archaeological work. This
involved an examination of whether landform
features such as source-bordering dunes, which
have the potential to contain information on lake
chronology, had developed on lakeshore margins.
These transverse dunes, Twidale's 'leeside
mounds' (1972; 85-86), are similar to the lunettes
of semi-arid regions. They are formed when
longshore drift transports debris to beaches or lee
shores of lakes. This sediment is then locally
redistributed by the wind before being trapped by
vegetation close to the lake margin.

The work was complementary to the
archaeological survey in the sense that it was
anticipated that source-bordering dunes were also
likely to be a focus for human occupation.
Systematic survey of these features would not
only have the potential to reveal information
about environmental features but also about
human occupation, and the chronology of such
occupation.

The project comprised an examination of
examples of source-bordering dune features for
each of the lake systems discussed earlier. Aerial
photographs were used to identify the features and
distinguish them from the linear dunes of the
continental dunefield. The transverse dunes are
paler in colour and generally trend east-west in
contrast to the north-south longitudinal dunes.

Once they were identified on the aerial
photographs I travelled to them on the ground and
systematically surveyed all exposures for any
archaeological material.

Access to the features by vehicle was often
difficult and in a number of cases access was by
helicopter. Regarding the dating of the dunes it is
noted that at the time the work was undertaken
only radiocarbon dating was available to the
study. Thus if organic material suitable for such
dating was not present in-situ the dunes were
unable to be directly dated. The results of the
work are presented below.

The Coongie system

Information on a number of the source-
bordering dunes of the Coongie system is outlined
in detail in Williams (1988: 58-60). A summary
of the work is presented here along with new data
on other parts of the system.

The analysis of aerial photographs, maps and
the survey work itself revealed that while some
features such as colour of the source-bordering
dunes are similar across the system there is major
variability in the number, size and location of the
features. A common feature is that all dunes are
pale in colour and have little or no carbonate
formation within their profiles, and they contain
intermittent scatters of artefacts on their surface -
typologically the artefacts appear to be late
Holocene in age. All of these factors suggest the
dunes are recent features. When cores of the
dunes were exposed by erosion and these sections
were examined, no organic or archaeological
material was found in-situ, with the exception of
one site at Lake Marradibbadibba - Lake Goyder
1 . Details of this site are presented in Williams
(1988: 60) where it is noted that it dates to the
relatively recent past 810-130 BP (ANU 5424)
thus confirming the recent age of the transverse
features.

There is considerable variation in dune shape
and alignment across the system. Some lakes had
a low and amorphous single transverse dune (such
as Toontoowaranie); others had a higher and more
distinct single dune (Coongie, Marroocoolcannie).
Still others had multiple transverse dunes, one
behind the other (five in the case of Lake Lady
Blanche, yet none on the neighbouring Lake Sir
Richard, and five or more in the floodplain
between Coongie and Toontoowaranie). Location
of dunes relative to lakes ranged from north, to
north-east and east.

Given that the study of the dunes was only a
preliminary one, it has not been possible to

E. WILLIAMS

determine what factors are responsible for the
variation in dune features. It is likely that since
there is major variation in the shape and size of
the lakes themselves and how they fill relative to
channel features and other lakes, and marked
fluctuations in the amount of water coming into
the system and the timing of when these events
occur, it is probable that these factors are
responsible for much of the variation.

The morphology and colour of the dunes
indicates that they are of late Holocenc rather than
Pleistocene age. Two hypotheses could explain
their recent age. The first proposes that the lake
system was active in the Pleistocene, but since tt is
still in operation today, receiving what can be
major flows of water and often flooding out
extensively, any Pleistocene shore features have
either been swept away by previous high-water
events or are well buried below flood deposits. In
1974 for example, the entire system flooded out -
aerial and satellite photography reveals water
extending for hundreds of square kilometres with
only the tops of linear and transverse dunes visible.
Krieg and Callen note that the Coongic system is
one of net sediment accumulation (1990: 60) and it
is thus possible that many older lake-shore features
have been disturbed or buried by sedimentation.

The second hypothesis proposes that the lake
system may not have been in operation in the
Pleistocene. Recent work by Callen and Bradford
(1992) presents data showing that the current
situation, in which most of the ilow down the
Cooper channel west of Innamincka is diverted
north up the North-West Branch, may be very
sensitive to minor changes in topology resulting
from such factors as tectonic activity or shifts in
sedimentation or the location of dune features
(transverse and longitudinal). Slight movements
on faults or in sedimentary features of only 1-2
metres within a large, low angle fan formed by
the Cooper immediately west of Innamincka may
be sufficient to create major changes in channel
direction, and thus whether the Cooper flow is
diverted either along the Branch, along another
channel such as the main Cooper channel or even
southwards down Strzclecki creek. Given this
sensitivity to change it is possible that during the
Pleistocene the North-West Branch may not have
been carrying the main Cooper flow and the lakes
may not have functioned as they do today.

Because a detailed study of the geomorphology
of the Coongie system has not been undertaken it
is not possible to resolve these competing
hypotheses. It is nevertheless possible to conclude
that whatever the reason, the lake-shore features

currently exposed within the area do not provide a
window onto the Pleistocene.

The salt lakes

Two of the larger salinas in the region, Coorie
Coorie Tillie and Lake Deception, were chosen for
analysis. Although the lakes lie some 100 km
from each other, the form of their transverse dunes
is virtually identical, in direct contrast to the
variability of the Coongie system. As well, the
transverse dunes themselves are very different to
those of the Coongie.

Each lake has a low, flat, pale-coloured
transverse dune on its northern edge. These have
been extensively gullied to expose older-looking
sediments which are redder in colour and more
compact with a pronounced carbonate horizon. It
is likely that further analysis would confirm that
the sediments are of Pleistocene age.

A systematic survey of the length of each dune,
including an examination of the many sections of
dune core exposed by erosion, revealed that the
small amount of archaeological material present
all lay within recent sediments on the dune
surface, rather than in the Pleistocene core.

In contrast to the Coongie, there are no large
amounts of water coming into the system to
modify or distort landform features. The salinas
are groundwater features, and do not fill from any
channels. The dunes' consistency in nature and
form could result from similarity in formation and
hydrology operating on the lakes. Unlike the
Coongie, the transverse dunes appear to have the
potential to preserve Pleistocene sediments but no
stratified archaeological material is currently
visible within their profiles.

The Cordillo lakes

This system has a set of source-bordering dunes
different from both the Coongie system and the
salinas. Like the salinas and unlike the Coongie
system, there appeared to be a strong consistency
in the number, form and structure of the features
across the lake margins.

On the northern edge of the lakes is a sequence
of three transverse dunes trending east-west, and
located one behind the other. The dune closest to
the lake edge is quite low, less than a metre in
height and pale in colour. Behind this is another,
slightly higher, pale-coloured dune which is
beginning to blow out into small longitudinal
dunes. Behind this again is another higher dune,
distinctly orange in colour with a pronounced
carbonate horizon, which has almost completely
blown out into longitudinal dunes.

ARCHAEOLOGY OF THE MIDDLE COOPER BASIN

The outermost dune is heavily eroded and it is
likely that it is Pleistocene in age. Samples of
carbonate from within the profile of the dune from
Lake Koonoomorinna were dated using
radiocarbon techniques and returned the relatively
recent dates of 3930-80 BP (ANU 6182) and
4560-80 BP (ANU 6182). John Head of ANU
Radiocarbon Laboratory notes that the samples are
not dating the formation of the dune itself, only
the last major phase of mobilisation of carbonates
within the dune. Such mobilisation would have
taken place during the last period of stable high-
water levels in the region and the samples are
probably dating this event (pcrs. comm.).

A complete survey of all of the dunes from one
lake, Koonoomoorinna, revealed that stone
artefacts are scattered across the surface of each
dune, but that no artefacts or organic material
were present in-situ within the cores of the
features, including the Pleistocene dune. An
analysis of a number of samples of the artefacts is
presented later in the paper but it is noted here
that those seen had features characteristic of
artefacts dating to the mid-late Holocene.

It is suggested that the regularity in source-
bordering dune spacing and form for the Cordillo
lake system is, like the salinas, reflecting
consistency in processes involved in formation
and filling. The lakes fill from run-off within the
local catchment, and in contrast to the Coongie
system there is significantly more regularity in
their hydrological processes and many fewer flood
events to affect the morphology of lake-shore
features. Like the salinas and in contrast to the
Coongie system, the transverse dunes appear to
have the potential to preserve Pleistocene
sediments but, as with the salinas, no in-situ
archaeological material is currently visible within
their profiles.

It is not known at this stage why the lakes
preserve a group of three transverse dunes. Each
dune may reflect a period of relative stability in
lake levels so that the position of the dunes, one
behind the other, may indicate a shrinking lake
basin since higher levels in the later Pleistocene.
Alternatively, the position of the dunes may
reflect the fact that all lakes in the region,
including the Coongie system, are slowly
migrating southwards over time (Bob Wasson
pers. comm.). The presence of the dunes may
reflect both factors.

Following this presentation of climatic and
environmental information, the next section places
the region in a more recent context by
summarising historical accounts of how

Aboriginal people occupied the area at the time of
contact with European people. The section draws
on information presented in Williams (1988) and
Layton, Foley and Williams (1991: 258-259) and
also presents new material.

Historical Accounts Of Subsistence And
Settlement

Citing McKinlay (1862), who was the first
European to travel through the entire middle
Cooper lake system, the region was densely
populated at the time of contact. McKinlay's
expedition, which was one of those initiated to
search for Burke and Wills, involved the party
spending several months in the lake system in
1861-1 862. During this time they were
systematically examining each of the lakes and
channel systems of the North- West Branch in their
search for the missing explorers.

McKinlay saw more than 700 people in the
region during his travels, including more than 300
people camped along Hamilton Creek,
immediately west of Lake Toontoowaranie in the
Coongie system, 200-300 people along the North-
West Branch just south of Coongie Lake itself,
and at least 150 people around Lake Lady Blanche
(1862: 37, 38, 46). Because of the length of time
he spent in the region he was also able to observe
how people utilised fluctuations in water levels
across the lakes.

McKinlay 's information shows that population
densities were highest around the large,
permanent waterholes on the channels of the
Norlh-West Branch and the Cooper itself, and
those lakes within the North-West Branch system
which were currently holding water. Group
composition appears to have been relatively fluid.
Large aggregations of 1 50 or more were formed to
take advantage of the flush of food resources
which appeared as lakes began to fill. As they
dried out again, people moved to other lakes,
following the water, or split into small groups to
harvest other resources or move on to other areas.
Permanent or semi-permanent camps of between
twenty and forty people were found along the
permanent waterholes of the channel systems and
this observation is also consistent with that of
King, the surviving member of the Burke and
Wills party who was cared for by Aboriginal
people in the Innamincka region (Burke and Wills
1861). At these more settled camps habitations
comprised substantial huts.

Important food resources were fish, crayfish

E. WILLIAMS

and mussels; waterbirds, and the variety of plant
foods which germinated after flooding or local
rain, especially nardoo. People favoured the lakes
and rivers but after rain in the local catchment
dispersed into small groups and travelled out into
the drier dune country (McKinlay 1862: 49; Jones
1979). In these latter areas a flush of resources
including small mammals, reptiles and certain
types of plant foods, especially seed plants,
appeared after the rain.

Although there are no ethnographic
observations of how the Cordillo system was used,
it can be hypothesised that after local rain areas
such as this became a focus for occupation, the
water sources it now held resulting in a relative
abundance of food. It is also possible that if semi-
permanent pools of water remained for a
considerable time throughout the year in the
shallow channels between the lakes, some small
groups of people might have occupied the region
on a more permanent basis.

While historical accounts mention the existence
of the salt lakes there are no direct ethnographic
observations of people using these areas. This
implies that they were not favoured areas for
occupation on a day-to-day basis.

Dealing with periods of resource stress

Although the Cooper lakes and channel systems
contain a large range of food resources for much
of the year, it appears that periods of resource
stress were common, especially during droughts.
Although there are no direct observations of the
study area itself during these periods, historical
material in Reuther (1981) provides information
on strategies used by Aboriginal communities in
the north-east of South Australia generally, to deal
with the major fluctuations in environmental
conditions characteristic of the region.

Reuther noted that when resources were
abundant, for example in periods after rain,
communities utilised natural surpluses in a variety
of ways. Large gatherings were organised to take
advantage of the abundance of food and these
aggregations had the primary function of
satisfying social obligations through, for example,
trade and exchange (1981; vol. 1: 42, 47^8). As
well as these activities, people harvested more
than they needed for immediate consumption,
processing and storing the surplus for later use.
Seeds, fish and grubs were the resources most
favoured for this type of processing (1981 : vol. 1:
13, 47^18, 232, 273, 491; vol. 2: 547, 772, 835,
915, 927; vol. 4: 1852; vol. 7: 458, 549, 659).
As well as these measures to maximise the

productivity of a region, according to Reuther the
distribution of food resources was also artificially
managed. He recorded that rats and caterpillars
were moved from one area to another so as to
increase stocks in particular regions (1981 : vol. 2:
811-812; vol. 9: 232). Plant foods were
propagated through the broadcasting of seed and
the distribution of the branches of a particular vine
species (1981: vol. 1: 107; vol. 7: 450). Areas of
stream channels were dammed to increase stocks
of fish (1981: vol. 8: 23). As well, certain
localities were not utilised and were left in reserve
for use during lean times, access to especially
productive areas was strictly controlled and the
boundaries of group territories were rigorously
enforced (1981: vol. 1: 101, 123-128, 412; vol. 2:
559, 751, 1044; vol. 3: 1255, 1259).
Transgressions, such as cases of people trying to
obtain more food resources than their due, or
attempting to accumulate resources, were severely
punished according to Reuther, even by death
(1981: vol. 1: 12-13, 123-129; vol. 2: 793-794).
Further, instances were recorded of particular
groups attempting to appropriate others' territory,
sometimes successfully (1981 vol.1: 415; vol. 2:
896).

During times of drought people fell back to
those areas which had water, and fish and stored
food resources were the main foods consumed
(1981 vol. 1: 520; vol. 2: 547, 616, 758 ? 915, 950;
vol. 2: 1259; vol. 4: 1740). Although Reuther
doesn't specify these areas, it is likely that the
Coongie system and the Cooper channel near
Innamincka were used in this manner.

Despite measures to counteract the effects of
drought or a lack of surface water, local groups
still suffered considerable stress. Reuther noted
that many people perished at these times, from
thirst, heat exhaustion or starvation (1981: vol. 1:
424, 535; vol. 2: 994-995, 1036; vol. 4: 1741-
1742, 1926, 2108; vol. 8:75, 118).

Factors Other Than Food Resources Affecting
Settlement

It is a truism to state that it is not only the
acquisition of food resources which affects the
choice and nature of settlements. Access to raw
materials necessary for the manufacture of goods
and for trade, such as stone, wood, ochre, resin,
bone, shell and the narcotic pituri, was another
important determinant.

Historical accounts show that the Innamincka
region was an important centre both for the

ARCHAEOLOGY OF THE MIDDLE COOPER BASIN

production of technological items, especially
stone artefacts and for a related reason, as a
crucial node in the vast exchange network which
spanned the continent (Kerwin and Breen 1981:
286 and McBryde 1987). Innamincka was
especially renowned for its extensive grindstone
quarries (see Reuther 1981 vol.2: 899-900,
McBryde 1987). The quarries, other important
sites involved in the extraction of resources and
the manufacture of goods, and the exchange
networks of the Lake Eyre Basin generally, are
currently being studied by McBryde and are thus
not reported on here.

Although factors such as the utilisation of items
for artefact manufacture and the location of trade
and exchange networks are, as outlined above,
crucial factors involved in settlement, not all of
these features are equally archaeologically visible.
For this reason I have chosen to look at one aspect
that is more archaeologically visible than others,
namely the characteristics of stone artefacts on
lake-side sites and what these data, and other
characteristics of the sites themselves, can tell us
about Aboriginal use of the region. As outlined,
stone suitable for the manufacture of artefacts is
not uniformly distributed across the region and
with the exception of Innamincka, stone sources
and water sources do not generally coincide.

Two main sources of stone arc present in the
study area. Silcrete, chert and chalcedony are
found as smallish pebbles on the surface of gibber
plains, while better quality and more extensive
areas of hard rock, comprising outcrops of chert,
chalcedony, silcrete, quartzite and sandstone are
found within the geological features known as the
Innamincka and Cordillo domes (South Australia
Department of Mines and Energy staff: pers.
comm. and personal observation). Discrete areas
of outcropping hard rock also occur within the
linear dunefield south of the Cooper channel and
east of Strzelecki creek (see Smith, Williams and
Wasson 1991: 185).

Although there are no historical accounts of
how Aboriginal people in the region quarried and
used stone, apart from Reuther's observations on
the Innamincka grindstone quarries, field
observations and geological data can be used to
determine what may have been favoured sources.
It appears that while gibber pebbles were utilised,
this took place on a relatively limited basis
compared to other sources because of the
relatively small size of the nodules of stone that
could be worked. Outcropping areas of hard rock
were thus favoured over gibber pebbles because
they provided larger fragments of stone and were

more uniform in quality. As well, certain types of
stone such as sandstone were not available as
pebbles and were only located within hard rock
localities.

Using data from geological mapping of the
region and advice from South Australian
Department of Mines and Energy staff, it is
estimated that the Coongie system lies at least 60
km from good quality stone sources while the
Cordillo lakes lie at least 20 km distant.
Particular stone sources which would have been
favoured include: the silcrete (including
chalcedony, cherts, cherty silcretes and
quartzites) of the Cordillo surface which caps
the mesas and cuestas of the Cordillo and
Innamincka regions and also includes the
silicified Eyre formation and the silicified
Winton formation; the Eyre formation which
includes sandstones suitable for grindstone
manufacture, and also small pebbles of agate and
fossil wood, and the Cadelga limestone which
includes a cherty limestone.

Surrounding these outcrops of hard rock and
extending outwards for five kilometres or so are
areas of gibber containing pebbles and larger
blocks of stone eroded from these formations.
Some of this material would have been utilised as
well.

A Model For Archaeological Site Distribution
WiTHrN The Lake Systems

Historical observations can be combined with
information on the availability of stone to present
a model of how the lake systems were utilised.
The model makes a number of assumptions,
namely that there is a relationship between the
availability of water and food resources; and
intensity of occupation as reflected by the size and
nature of archaeological sites. Cutting across these
factors are others relating to the availability of
stone, the most common material found on sites.
Using models developed by other researchers such
as Hiscock (1987), it is hypothesised that there
will be a general correspondence between the
morphology of artefacts on a site and distance
from the stone source. These models predict that
as one moves significant distances from stone
sources, characteristics of extreme reduction of
stone material will be found in assemblages such
as recycling, rejuvenation and rationing.
Combining these different models, we can develop
a unified set of hypotheses regarding the
patterning of archaeological material.

E, WILLIAMS

The Coongie system

The Coongie lakes, with their abundance of
food sources, and availability of water for at
least part of the year, were areas with a high
population density relative to other lakes in the
region. Large and diverse occupation sites will
be found in those areas which contain the most
predictable and extensive areas of water. Such
places include the permanent waterholes on the
channels, the inlets to lakes, and those parts of
the lakes that are deeper relative to the rest of
the lake basin and thus hold water for a longer
period of time. These latter areas are found on
the southern margin, since as outlined above, the
lakes in the region are slowly migrating
southwards, cutting into longitudinal dunes on
their southern shore. Their southern end is thus
their deepest section.

It is anticipated that given the known high
population densities of at least the recent past,
there will be a correspondingly high density of
archaeological material (especially stone
artefacts) on the large, favoured occupation
sites. It is likely that on these sites people
would not have moved as much compared to
areas which contained less predictable
resources. We can therefore expect evidence
for the manufacturing and maintenance of
tools such as stone artefacts. Given that the
sites were used on a longer-term basis we can
also expect a higher proportion of formal tool
types than on sites with more unreliable water
sources, and a greater proportion of finer-
grained, good quality stone. As well, it is
likely that a relatively larger number of
artefacts will have edge damage from activities
such as trampling, because of high population
densities at the sites and residential stability
(Scott Mitchell pers comm).

Because it is possible that people from other
areas may have been using these localities as
refuges in times of resource stress, it is also
expected that these places will contain a wider
range of raw material than at other sites,
including stone brought in from other regions.
Because stone sources lay some considerable
distance away (60+ km), primary reduction was
unlikely to have been undertaken here and the
stone may have been 'rationed'. Cortex, an
indication of primary reduction, will be rare on
pieces. Flakes and cores will be small in size
and noticeably reduced. High population
densities and length of site occupation may have
exacerbated the incidence of rationing, so that
these features will be particularly marked at the

large sites near permanent or semi-permanent
water.

The salt lakes

Given the absence of fresh water in these
localities it is anticipated that they were not
favoured as occupation sites. In times of high local
rainfall it is possible that the lakes may have filled
to some extent with rainwater run-off from the
surrounding dunes. At these times salt levels may
have decreased and they might have been used on
a temporary basis as people pushed out into the
dunefields after rain.

The salt lakes may have had another role, as
navigation aids for people travelling through the
dunefield. They are often the only notable features
present within extensive areas of dunes and are
visible for some distance because of the sun
sparkling on the salt surface.

I conclude that the sites were used, albeit on a
much more limited basis than the Coongie and
Cordillo system. It is likely that groups using them
were small in size and highly mobile. There is
likely to be a correspondingly low density of
artefacts, relative absence of formal tools, a more
limited range of raw material types present and a
lower proportion of good quality fine-grained raw
material on occupation sites. Little manufacture or
maintenance of artefacts will have been carried
out on site. Given that the lakes are often well
within the dunefield and thus some distance from
stone sources, the stone discarded here will have
been noticeably reduced.

The Cordillo lakes

With their better resources than the salt lakes
the Cordillo lakes are likely to have been the
subject of more intensive occupation but on a less
pronounced basis than the better-watered Coongie
system. It is anticipated that sites will be smaller
in size than in the Coongie and fewer artefacts
will be present. Because people occupied these
sites for shorter periods of time it is expected that
there will be less evidence for the manufacturing
and maintenance of stone artefacts. A lower
number and a smaller range of formal tool types
will be present, and there will be less variation in
raw material type. Larger sites are more likely to
be located in areas of more predictable water and
food resources such as the inlets to lakes or the
small waterholes on channels.

Given that the Cordillo lakes are closer to raw
material sources than the Coongie it is expected
that stone artefacts will not be reduced as much.
Cores will be more common and cores and flakes

ARCHAEOLOGY OF THE MIDDLE COOPER BASIN

will be heavier and larger in size, on average. A
higher proportion of flaked pieces will contain
cortex.

The Archaeological Surveys: General
Archaeological Features Of The Lake Systems

Before presenting material on the
archaeological sites examined as part of the study
I will outline some of the factors affecting the
location and recording of archaeological material
in the field.

The visibility of archaeological material,
especially stone artefacts, is reasonable compared
to many other areas. The aridity of the region
means that dense vegetation is often absent and
much of the ground surface is exposed. This also
means that stone sources are relatively
straightforward to detect. Open sites are the major
archaeological site type in the region because rock
formations which form rock shelter sites are
extremely rare here. Only one shelter (a very small
and shallow one located near Innamincka) is
known for the whole region. Although open sites
arc the major site type available for analysis, given
the good ground surface visibility this is not
necessarily the problem that it is in other regions.
Stratified sites are extremely rare nevertheless and
it is often very difficult to determine whether
archaeological material is unequivocally in-situ,
has been deflated down from other units, or
otherwise re-deposited.

I will present general features of the
archaeology of the lake systems first and compare
them with my model. In the following sections I
will discuss more detailed analyses of
assemblages from a sample of collected sites so as
to make a comparison between localities within
the region.

The Coongie system

The results of this work are presented in
Williams (1988) and are briefly summarised here,
along with other material from 1987 surveys.
Before outlining this information it is noted that
not all of the lakes in the Coongie system were
surveyed for archaeological sites. Coongie itself
was not examined because it was already being
systematically surveyed by a team of
archaeologists from the South Australian
Aboriginal Heritage Branch, (now within the
Department of State Aboriginal Affairs), in
relation to the development of a management plan
for the region. The results of that study are not yet

available for analysis. Lake Goyder was not
examined because in the years which formed the
subject of my fieldwork (1986-1987) the lake was
flooding out and thus conditions were not
conducive to archaeological survey.

The most common site type around the lakes
surveyed for the study comprised scatters of stone
artefacts. In all sites surveyed these scatters were
either located within upper recent sediments or
appeared to have deflated down from such units
to sit as a lag on more compacted surfaces. On
many of the sites, scatters of freshwater mussel
shell were also present. Although not all the
margins of every lake were examined, it appeared
that stone and shell was present as an intermettent
scatter intermittently around the margins of
lakes that were surveyed (Toontoowaranie,
Marroocutchanie). These were often present as
small scatters of shell with numbers of artefacts,
similar to Meehan's (1982) 'dinnertime camps'.
Artefact density and site size increased as one
approached permanent water sources and the
largest sites were found in the areas outlined
earlier - permanent waterholes on the channels,
the inlets to lakes, and the deeper, southern
margins of the lakes. These latter sites often
incorporated extensive scatters of shell as well.
Compared to the large sites, site size and density
of material was comparatively low on the pale-
coloured transverse dunes. On these sites stone
artefacts and scattered termite mound hearth
fragments were the most common form of
material.

Typologically, most, if not all artefacts dated to
the mid to late Holocene. The main types noted
were tula adzes and adze slugs, points, backed
blades, small scrapers, cores, flakes and fragments
of large, flat grindstones of the type described by
Smith (1986). As well, single examples of large,
cube-shaped silcrete cobbles which were
commonly ground on one or more surfaces and in
some cases also had flakes removed from edges
and had thus been used as large cores, were often
also present. Raw materials favoured for artefact
manufacture were chalcedony, chert, cherty
silcrete, a coarser-grained silcrete, and quartzite.

Other artefacts such as edge-ground axes and
flaked pieces of quartz crystal were also found but
appeared to be restricted in distribution to the
lakes most difficult to access by vehicle. It appears
that such artefacts were formerly present on other
lake-side sites but, together with other types of
artefacts such as complete grindstones, have been
removed by collectors (a conclusion supported by
oral tradition in the region and also personal

E. WILLIAMS

observation of collections held by individuals
resident in the region).

Apart from the occasional large silcrete cobbles
and remaining grindstones left on sites, stone
artefacts were mostly small in size and appeared
to be noticeably reduced. As well, although
density of stone material varied with location, even
on sites with comparatively higher numbers of
artefacts, density was still relatively low compared
to artefact scatters found in many other parts of the
continent. On the larger sites average density for
collected sites was of the order of 6-8 artefacts/
square metre while on the smaller sites maximum
densities were only 2-A artefacts/square metre.

Occasionally larger flakes and horsehoof cores
were found on sites or located as isolated finds
within the dunefield or near lakes. There appeared
to be no consistency in their location though and
these were often in association with artefacts that
typologically dated to the mid to late Holocene.
As well, they were not noticeably patinated or
weathered, and did not appear to be derived from
Pleistocene sedimentary contexts, suggesting that
most of these are mid to late Holocene in age.

Hearth material, usually fragments of burnt
termite mound, was often scattered across sites
and isolated hearths were also present. The
amount of hearth material rose in density on the
larger sites and on these latter places fragments of
bone (mostly that of fish and small mammals)
were often found as well. The large sites are
indeed extensive - examples up to 10 000 square
metres and 7 000 square metres in extent
respectively are described in Williams (1 988: 59).

Dating of material on sites confirms that
implied by the typology of artefacts, that is, of the
late Holocene. Details of radiocarbon dates are
provided in Williams (1988).

The archaeological data is consistent with the
ethnographic observations that the Coongie lake
system was extensively used and densely
populated. The presence of good quality raw
material and a range of artefact types, including
formal tools and manufacturing debris, suggests
that residential stability could have been fairly
high in comparison to other areas. This is also
supported by the wide range of archaeological
material present, including bone and shell. The
small size and marked reduction of artefacts
indicates rationing of raw material was taking
place and this is consistent with the distance of
the area from stone sources.

The salt lakes
As expected, within the areas sampled for sites

for Coorie Coorie Tillie and Lake Deception,
archaeological material existed in very low
densities. At Coorie Coorie Tillie for example,
along the 5 km stretch of the transverse dune on
the northern margin of the lake only five isolated
artefacts were seen: three amorphous flakes, one
tula slug and one small core. All artefacts were
manufactured from silcrete. Archaeological
material was similarly rare at Lake Deception.
Sites comprising occasional small, discrete groups
of artefacts were found in the vicinity of both
lakes but not within the transverse dune systems.
Instead they were set back one or two longitudinal
dunes from the lake margin, on the surfaces of
small claypans.

The sites found in these areas comprised small
(mostly less than 30 artefacts), discrete scatters of
stone artefacts. Density was very low, much less
than 1 artefact/square metre. Neither mussel shell
nor animal bone was present on these sites or on
other parts of lake margins but occasionally
examples of isolated termite mound hearths were
found in association with the stone artefact
scatters.

One of the termite mound hearths at Coorie
Coorie Tillie was dated. To examine particular
features of the region's environment which may
be affecting dates, the material was separated into
a humic and non-humic fraction and each sample
separately dated. The humic fraction returned a
date of 1150 180 BP (ANU 6181) and the non-
humic: 710 170 BP (also ANU 6181). The
disparity in the dates can be explained as follows.
John Head, ANU Radiocarbon Laboratory,
confirms that the non-humic fraction, comprising
clay particles, binds to younger carbon and clay
material, biasing the date and making it too
young. To account for this the non-humic
fraction, which although comprising polymers
which can incorporate younger carbon, was
treated with solvents so as to extract the older
carbon which is more likely to be reflecting the
true age of the sample (pers. comm.). The older
carbon is dated and the result, 1 150 180 BP, is the
best estimate of the sample's true age.

Apart from the differences in density and site
composition from the Coongie sites, other
distinguishing features were also apparent. Most
of the artefacts were manufactured from a
relatively restricted range of raw material,
assemblages being dominated by a grey,
comparatively coarse-grained silcrete. Some cores
were present and also occasional formal tool types
such as tulas, scrapers or points. Artefacts
appeared to be noticeably larger than those at

ARCHAEOLOGY OF THE MIDDLE COOPER BASIN

Coongie. Because of the low density of material,
and also given the difficulty of access to the sites,
collections of artefacts were not made when the
sites were visited and so a more formal
comparison between these features and other lake
systems is not possible. The low density of
material present on these lakes suggests that while
they were utilised, they were not favoured for
occupation. The restricted range of raw material
found, its contrast in quality to the fine-grained
material found on the Coongie system and the
relative absence of manufacturing debris, indicates
people used these areas on an opportunistic basis.
The low numbers of artefacts present also
suggests that group sizes were relatively low.

The Cordillo lakes

Vehicular access to the lakes was difficult
because of their remote location and thus only one
lake. Koonoomoorinna, was surveyed in detail.
Reconnaissance trips to other lakes in the region
revealed that the general patterning of sites on this
lake is similar to that on others.

As for other parts of the study region, the most
common site types seen were scatters of stone
artefacts. Formal tool types were similar to those
observed on the Coongie sites, comprising tulas
and other adzes, scrapers, points, fragments of
grindstones and the large silcrete cobbles with
ground and/or flaked surfaces. Formal types did
not appear to be as common within assemblages
as for the Coongie and no shell or bone was
present on sites. The range of raw material did not
seem as extensive and Finer grained raw material
such as chert and chalcedony was found in lower
densities. Artefacts appeared to be larger in size
and not as heavily reduced.

Hearths were also not as common and no hearth
sites were observed on any of the sites ringing the
immediate margin of the lake. A small number of
hearths were found in the inter-dune corridors one
or more dunes back from the lake margin. One,
comprising burnt calcrete nodules, was dated and
returned a 'modern 1 radiocarbon age (100.2, 1.8
%M, ANU 6183), although this sample may have
been contaminated by younger carbon from the
surrounding locality (John Head pers. comm.).

The densest, most extensive scatters of artefacts
were found in those areas with the most reliable
sources of water - inlets, the southern margin of
lakes, and on the edges of possibly semi-
permanent waterholes along the channels which
connect the lakes. A large scatter along one of the
channel waterholes contained a number of hearths.
Intermittent, low density scatters of artefacts were

found along the lake margins and on the
transverse dunes to the north.

The archaeological patterning is consistent with
the hypothesis that the Cordillo lakes were utilised
more than the salt lakes but less than the better-
watered Coongie area. This is supported by the
fact that the Cordillo sites contained a smaller
range of archaeological material than the Coongie
sites (no shell or bone, hearths not as common, a
lower density of formal tool types and of better
quality stone). The relative lack of reduction of
artefacts could indicate either less residential
stability or a closer distance than Coongie to raw
material sources, or both.

Now that I have presented general observations
of the archaeology of the lake systems I will
outline the results of the more detailed
comparisons between sites. This work provides a
more fine-grained analysis of differences that may
exist across lake systems and between systems.

More Detailed Comparison Of Assemblages
From Individual Sites

Because of the relatively low density of
artefactual material within the region, for reasons
of statistical analysis I only made formal
collections from large sites. The sites analysed in
this way are outlined below, along with the results
of the analysis.

Before discussing the sites it is noted that
although some of the analysis of the material was
published in a preliminary form by Smith,
Williams and Wasson (1991, see for example
Tables 1 1 and 12), a number of the figures in that
paper vary slightly from those presented in this
publication. This is due to the fact that the results
of the analysis of the Coongie sites outlined in the
1991 paper were only at a preliminary stage. The
material presented in the current paper reflects the
completed analysis of the assemblages.

Sites chosen for analysis

Five sites were chosen for analysis:
Marroocoolcannie 2 (MCO 2), Toontoowaranie 1
(TTW 1), Lake Lady Blanche midden (LLB),
Koonoomoorinna inlet (KOI) and
Koonoomoorinna second white transverse dune
(KSW). The first three sites are located on
Coongie lakes - their name indicates on which
lake they are situated, and the latter two sites are
located on the southern-most of the Cordillo lakes,
Koonoomoorinna. All sites comprise large,
discrete scatters of stone artefacts, together with

E. WILLIAMS

other material such as shell and bone in some
cases.

The lakes were chosen for sampling and
analysis to determine whether there were any
major changes in stone artefact manufacture and
discard across the Coongie system, Each site was
chosen because it was the most extensive seen for
the sampled areas of that particular lake. MCO 2
lies along the western margin of the channel
feeding from Lake Marroolcoocannie to Lake
Marroocutchanie. It covers an area of
approximately 2 600 square metres and comprises
a scatter of artefacts and shell deflating down from
the uppermost recent unit of a pale, longitudinal
dune. The material exists as a lag on a blow-out
within the dune. At the western end of the site is a
localised area of burnt shell and bone, the latter
comprising the remains offish and small mammal
species. A hearth, which lies on the eastern part of
the site, dated to 1 130 1 10 BP (ANU 429).

A baseline was laid through the middle of the
scatter so as to bisect both the area of burnt bone
and the hearth. All artefacts along a strip 60 m
along the baseline and 1 m either site of it were
collected for analysis.

TTW 1 lies further to the north than MCO 2,
located on the southern margin of Lake
Toontoowaranie. It covers an area of 7000 square
metres and comprises stone artefacts, shell and
the remains of the collapsed, wooden framework
of a domed hut, all lying on recent sediments on
the top of a longitudinal dune. A sample of shell
from the site returned a date of 330 80 BP (ANU
5425). A 20 x 2 metre grid was laid out within the
centre of the scatter and all artefacts within this
were collected.

LLB lies to the north and west of MCO 2 and
TTW 1, on the inlet channel to Lake Lady
Blanche, one of the terminal lakes in the Coongie
system along with Lake Sir Richard. These two
lakes hold water for a shorter period of time than
those closer to the North-West Branch. When the
lake was visited in August 1987 it was dry and
had a salt crust on the lake floor. This was in
contrast to the other Coongie lakes which were all
full of water.

The site comprises a large scatter of stone
artefacts, shell and bone extending for 12 000
square metres, eroding out of the uppermost units
of a recent, white transverse dune associated with
the inlet channel and flood-out zone of the lake.
The bone comprises fish bone and golden perch
otoliths. It is the largest and richest site in the
inlet area. A 15 x 15 metre grid was laid out on
the eastern edge of the exposure, where the

densest scatter of shells, otoliths and artefacts was
eroding out of the sand. All artefacts within the
grid were collected.

The relative lack of water in the lake compared
to others in the Coongie system reflects the
distribution of sites in the area. Large sites such
as this one, which are not only extensive but
contain a variety of material, are restricted to the
margins of the inlet channel. Isolated scatters of
stone artefacts are found on the surface of the
complex of at least five separate transverse dunes
lying to the north-east of the lake, but these sites
are small, discrete scatters of stone artefacts and
do not contain shell, bone or hearth material.

Although the lake contains water on fewer
occasions than others in the system, it was much
favoured by Aboriginal occupants. McKinlay
visited the lake in January 1862 and found it full
of water, * between five and six feet deep, and
seven and three quarter miles in circumference ...
and tens of thousands of pelicans on it ... with
innumerable other birds ... and plenty of fish*
(1862: 38). There was a large camp of at least 150
people on the lake when McKinlay was there and
he observed that fish and 'nardoo 1 was their chief
sustenance (1862: 42).

For the Cordillo lake, Koonoomoorinna, two
areas were sampled for artefacts. Both sites, KOI
and KSW were the most extensive discrete
scatters of artefacts seen within the vicinity of the
lake and this is why they were chosen for analysis.

KOI is a very extensive scatter of artefacts
which follows the inlet to Lake Koonoomoorinna
for several hundred metres. The site comprised
stone artefacts only - no shell, hearths or bone
were present. A small part of the site (10 m by 2
m) where the densest collection of artefacts was
found, was gridded and a collection made.

KSW comprises a discrete scatter of artefacts
lying as a lag within a blow-out in the second
transverse dune north of the lake. Like KOI, only
stone artefacts were present within the site.

MCO 2 and TTW 1 lie 65 km from sources of
outcropping hard rock in the Innamincka dome
and 60 km from the Cordillo dome sources. LLB
lies 40 km from the Cordillo dome sources and 65
km from the Innamincka dome. KOI and KSW lie
30 km from the Cordillo dome and 90 km from
the Innamincka dome.

For comparative purposes, the five lake-shore
sites are compared with each other and also with
one from the waterless core of the dunefield, site
JSN. This latter site, which is reported on in
Smith, Williams and Wasson (1991), comprises a
scatter of stone artefacts and hearth material, and

ARCHAEOLOGY OF THE MIDDLE COOPER BASIN

Table 1. Characteristics of artefacts within assemblages,
excluding ground pieces

Site mean weight % flakes % cores % formal

in gm, (sd) tool types

Table 3. Mean weights of cores (gm)

Site

mean

KSW

7.55 (17.57)

70.4

3.4

3.8

n=291

KOI

5.14(20.53)

48.4

4.2

2.6

n=308

LLB

2.65 (7.11)

45.2

0.6

11.9

n=336

TTW1

1.74(3.33)

59.5

—

4.2

n=264

MC0 2

2.17(8.32)

50.1

0.6

5.6

n=661

JSN

6.6 (sd not

37.3

2.7

not

n=407

available)

available

is the only known Pleistocene site in the region.
Hearths from the locality date to around 14 000,
10 000 and 2 500 BP. Associated with the hearths
and lying as a lag on two inter-dune pans within
longitudinal dunes is an extensive scatter of
artefacts. Despite the Pleistocene dates for the
hearths, analysis of the stone artefacts strongly
suggests that they are late Holocene in age (Smith,
Williams and Wasson 1991: 184).

JSN was chosen as a comparative site for the
lakeside sites because it comprises the only known
extensive scatter of stone artefacts for the core
dunefield region. Most core dunefield sites only
consist of an isolated artefact or very small
scatters of stone artefacts (personal observation).
The large size of the artefact sample (N = 407)
from JSN and the fact that the artefacts appear to
be late Holocene in age, similar in age to the
lakeside sites, makes it the only comparative
material currently available for the Coongie and
Cordillo sites.

KSW

KOI
LLB
TTW1
MC0 2

JSN

58.29

50.49

42.62

81.75

13.7

1.90

7.65

6.56

89.2

57.9

% artefacts

with cortex

12.4

11.0

4.29

9.2

3.2

20.0

Table 4. Percentage of artefacts larger than 1 cm in
length which contain cortex.

Site

KSW
KOI
LLB
TTW 1
MC0 2
JSN

Analysis of the sites

Broad characteristics of the stone artefact
assemblages are presented in Table 1. As
predicted by the model, the Coongie assemblages
have features reflecting an extreme degree of
reduction. Artefacts (all pieces of stone on the site
worked in some way, excluding ground pieces)
are extremely small in size (measured by mean
weight in grams) and cores are very rare, or even
non-existent in the case of TTW 1 . The Coongie
sites are very similar to each other but are different
to both the Cordillo sites and JSN, which contain
larger artefacts and a greater number of cores
respectively.

The differences in mean weight between the

Table 2. Values of t
weights of artefacts

for comparison between mean

Table 5. Raw material composition of the assemblages
(excluding small shatter < 1cm in length and including
ground pieces)

Site KSW

KOI

LLB

TTW 1 MCO 2

Site

% Finer

% Coarser

% Very coarse

KSW

KOI not

grained

grained (eg
sandstone)

significant

— —

KSW (n-278)

47.1

41.7

11.2

LLB t = 4.71

t = 2.11

- -

KOI (n-239)

37.2

59.0

3.8

TTW1 t = 5.33

t = 2.72

not

LLB (n=223)

72.6

19.3

8.1

significant

- -

TTW 1 (n-223)

66.4

27.4

6.3

MC0 2 t = 6.48

t = 3.22

not

MC0 2(n=659)

59.8

23.7

16.5

significant

JSN(n-401)

59.5

33.2

7.4

E.WILLIAMS

Table 6. Mean weights and mean lengths of complete flakes for different raw material types. Standard deviations are
shown in brackets. Note: Only those artefacts > 1 cm in length have been included in this analysis because of the
difficulty in determining whether smaller pieces were derived from finer or coarser-grained raw material. Ground
pieces and sandstone have also been excluded from the sample.

Site

mean weight (gm)
of complete flakes

mean weight (gm)

of debitage>l cm

in length

Mean length (mm)
of complete flakes

finer

coarser

finer

coarser

finer

coarser

grained

KSW

3.9(4.1)

12.0(18.6)

3,2 (2.7)

7.4(14.1)

25.0 (9.4)

31.7(13.6)

KOI

2.3 (2.6)

6.71 (15.6)

2.3 (2.9)

5.0(7.1)

17.8(7.1)

25.2 (14.60)

LLB

1.3(2.0)

2.1 (2.8)

3.1 (4.6)

9.3 (21.8)

15.8(5.1)

19.1 (6.8)

TTW1

0.9(1.1)

4.8 (6.4)

1.0(1.7)

7.4(14.9)

15.8(6.1)

26.3 (14.7)

MC0 2

0.9(1.5)

8.0 (24.3)

0.9 (0.9)

3.5 (5.3)

15.2(5.3)

23.5(16.2)

JSN

3.1 (3.2)

4.7 (10.6)

5.6(13.0)

3.5 (6.4)

not available

Coongie and Cordillo assemblages are significant
at the 0.05 level (Table 2). The figures for JSN are
not available for comparison but it is anticipated
that if they had been available, the mean weight
of the assemblage would be statistically different
to the Coongie sites but of the same order as the
Cordillo assemblages.

As predicted by the model, the three groups of
sites are very different in relation to the reduction
of cores (Table 3). The cores from the Coongie
sites are extremely small and, as outlined in
Smith, Williams and Wasson (1991:187), very
close to the theoretical limit at which bipolar
techniques become necessary for further reduction.
Consistent with this is the observation that in a
further attempt to ration raw material, worked-out
or small, broken cores have been recycled into
small scrapers in the Coongie assemblages (N=9 S
mean weight = 8.2 gm, sd = 4.9). The Cordillo
sites contain cores that are noticeably larger and
heavier than those on the Coongie sites, while
JSN contains on average larger cores again. There
is no evidence that cores have been recycled as
implements at either the Cordillo sites or at JSN.

The Cordillo sites contain proportionately
greater number of flaked pieces with cortex than
Coongie (Table 4), which is as predicted given
that they lie closer to raw material sources,
although the figures for TTW seem high in
comparison to the other Coongie sites. The JSN
assemblage contains an unusually high number of
flakes with some degree of cortex. This feature is
unexpected and as well as the relatively larger size
of cores and flakes, indicates that the JSN
assemblage shows neither the effect of relatively
large distance from stone sources, nor mediating

responses offsetting a diminishing stock of stone
(see Smith, Williams and Wasson 1991: 187).
This issue is discussed in more detail later in this
paper.

Regarding raw material composition for
assemblages, Table 5 shows that the Coongie sites
contain proportionately more finer grained
material (classified here as chalcedony, chert and
cherty silcrete) than either the Cordillo sites or
JSN and this is as expected (coarser-grained
material has been classified here as granular
silcrete and quartzite). Table 6 shows that flakes
and debitage from fine-grained material arc on
average smaller in size and lighter in weight than
those from coarser-grained material, but this docs
not explain all the variation in size between
assemblages between sites in different areas. Fine-
grained flakes and debitage from the Coongie sites
are in virtually every case smaller in size and
lighter in weight than fine-grained pieces from
either the Cordillo sites or JSN.

As well as these trends, the Coongie sites also
show a wider range of raw materials as predicted.
MCO 2 has 13 major types of raw material, TTW
1 has eight types, LLB has seven, KOI has five
and KSW has seven.

In terms of the artefact analysis it was decided
not to differentiate artefacts with retouch from
those where this feature appeared to be absent.
This is because of the difficulty found in
unambiguously identifying examples of retouched
pieces within the Coongie and Cordillo
assemblages. Numbers of artefacts within the
assemblages contained what I recorded in my
analysis as 'edge damage', where I describe this
as an edge where very small flakes had been

ARCHAEOLOGY OF THE MIDDLE COOPER BASIN

removed and there is also a gloss or polish (visible
under low-power magnification). However given
the small size and thin cross-section of many of
the flaked pieces it was impossible to determine
whether this edge damage had occurred as a result
of use, or as deliberate retouch, or from some other
process such as flaking or trampling for example.
Accordingly it was not possible to examine one of
my hypotheses, that sites near more permanent
water sources would show more edge damage
from trampling.

Because of the difficulties in identifying what
had caused the edge damage, none of the statistics
for this feature have therefore been listed for this
analysis. It is noted that in the assemblages
examined there were very few artefacts with
pronounced retouch (where this involved larger
flakes being removed along an edge and/or very
obvious polish) which did not also comprise a
formal tool type such as a scraper, adze, point or
backed blade. For the very few artefacts which did
not fit in to a formal category they were
nonetheless listed under 'formal types' in my
tables.

Given that this classification methodology
differed from that used for the JSN artefacts, it
has not been possible to compare the two groups
in a formal sense. Nonetheless some comparison
are possible.

In terms of my category of 'formal tool types',
the Coongie sites contain a greater proportion of
these artefacts than the Cordillo sites (Table I), as
predicted. Adzes such as tulas and tula slugs
formed the largest proportion of formal types in
the Coongie sites (59% for MCO 2, 45% for TTW
1 and 55% for LLB). The remaining implement
types comprised scrapers, points and backed
flakes. For the Cordillo sites in contrast there is
greater variation in the formal types of stone tools
represented on sites. Adzes comprised 36% of the
formal types present at KOI but only 18% of those
at KSW. At KOI scrapers comprise the most
common formal type (50%) while at KSW, points,
backed flakes and backed blades are the most
common type at 64%.

Regarding the artefacts comprising formal
types, it was expected that examples will be more
heavily reduced at sites near permanent water.
Examining the largest sample of a formal type
available, tula adzes and tula adze slugs, it is
found that this is not the case. There is no clear
trend in mean weights between the Coongie sites
(MCO 2: mean - 8.5 gm, sd - 5.02, n = 15;
TTW 1: mean = 3.25 gm, sd - 1.35, n = 2; LLB:
mean - 14.18 gm, sd = 1 0. 1 , n = 1 9) or between

the Coongie and Cordillo sites (KOI: mean =
3.7, sd= 1.0, n = 2; KSW: mean - 6.8, sd = 2.7,
n = 2).

This suggests a number of factors. While
overall characteristics of the assemblages
presented in Table 1 show that stone was being
curated and rationed in a very intensive way for
the Coongie sites, the variability in size at discard
for tulas and tula slugs, especially the relatively
higher weights for the LLB site, suggest that
occasionally the site occupants were not behaving
as if raw material was as scarce as it appeared to
be. This suggests that while certain factors such
as the balancing of procurement of food and water
sources with that of raw material is responsible
for much of the variation seen on sites, people did
not always behave as if this was the only equation
affecting the nature of occupation and discard of
stone artefacts.

Although the JSN assemblage has been
classified according to a different typology, a
preliminary analysis of it reveals that formal types
are rarer than at either the Coongie or Cordillo
sites. This is consistent with the model. As well,
it is further observed that adzes are not present on
the site. A possible explanation for this relates to
the lack of hardwood trees within the core
dunefield region supplying suitable wood for
adzing activities. The absence of such trees is very
likely related to the availability of water -
hardwood specimens being found on the banks of
waterholes and lakes.

Comparison Of The Trends In TirE Assemblages
With Data From Other Parts Of The Arid Zone

The trends seen in the assemblages for the
region are also echoed in other arid zone work
available for comparison. The major study which
provides comparative data is that of Veth (1993)
for the Western Desert region, Western Australia.
His analysis shows similar patterning to that
outlined above, with features of assemblages from
sites at permanent waters indicating more
intensive stone reduction than for cither semi-
permanent waters or ephemeral water sites. His
data show that differences in intensity of reduction
are not as clear between semi-permanent and
ephemeral water sites as they are between these
sites and more permanent ones (see 1993: Tables
8.21-8.31). For a number of features, ephemeral
sites show trends characteristic of more intensive
reduction than semi-permanent sites, which is not
predicted by Veth's model. This appearance of

E. WILLIAMS

conflicting trends also appears to be echoed by
comparisons between the Cordillo sites and JSN.
The sites are more similar to each other than they
are to the Coongie sites but there is much
variation in the nature and direction of trends
between the two groups. This suggests that the
permanent sites, in Veth's sample and mine, form
clear groups but other sites, while showing some
similar trends, are more variable.

The association of higher proportions of formal
types in assemblages, especially adzes, with more
predictable water sources, is also a trend noted by
Veth (1993: 95-96). Veth also found that there
was some patterning in the amount of reduction of
tula adzes and relation to water sources but that
there was a considerable degree of variation
(1993: 98-101) and this is also found in my
samples.

In his discussion of his results of the artefact
analysis, Veth concludes that the timing and
intensity of site use are the major factors
responsible for variation in the assemblages,
rather than distance to raw material sources and
the operation of specific discard criteria (1993:
101). This also seems to be the case for the north-
east South Australian sites, as I will outline
below.

Major Factors Affecting The Patterning Of
Sites In The Region

One of the predictions of my model of site
patterning is that apart from water sources,
availability of stone sources is be one of the most
important factors responsible for trends seen and
that Aboriginal occupants favoured raw material
sources closest to the site.

An examination of the JSN assemblage reveals
that the people who visited the site did not behave
in this manner. Rather than using the closest stone
to the site (40-50 km to the east) for artefact
manufacture and discard, the sourcing of stone
artefacts from the site indicates that they instead
used stone from Della-Dullingari which lies 115
km away, to the north-east (Smith, Williams and
Wasson 1991: 185). Despite the use of such
distant material, the assemblage does not have
features characteristic of extreme reduction
(Smith, Williams and Wasson 1991: 185-189).
Cores have not been noticeably curated and have
not been recycled as other implements. Mean
weight of artefacts is significantly greater than for
the Coongie sites, despite the fact that utilised
stone sources are considerably further away than

for the Coongie. Flaked pieces contain much
higher proportions of cortex, also consistent with
less intense reduction.

The patterning of the JSN assemblage is
characteristic of a situation where raw material is
closer rather than further away. Yet the paradox
can be resolved if we instead see stone source
utilisation in terms of re-supply time rather than
just in terms of linear distance of stone source
from occupation site. For example, if people
stopped at JSN as part of a journey to the Della-
Dullingari region, they would have been aware
that they were only a couple of days away from
being able to replenish stone sources. As well, it
is likely that the duration of visits to the JSN were
relatively short, so that the demand for stone was
easily met by the stock-in-hand without recourse
to the recycling and the extreme reduction of stone
evident at the Coongie sites (Smith, Williams and
Wasson 1991: 188).

The JSN site shows that it is not necessarily
linear distance from the closest stone sources that
is a major factor in the patterning of stone artefact
manufacture and discard. The key factors are
instead the length of time spent at the site, which
in turn relates to the length of time before the
group could replenish stone sources.

This line of reasoning can be applied to the
Coongie sites as well. The extreme reduction of
stone reflected by the assemblages can be seen as
the archaeological correlate of residential stability
within the lake system relative to other
environments in the region. It appears that people
chose to stay closer to the lakes even if it meant
that they were restricting their access to stone,
thus triggering what appears to have been
significant pressure on the way they used the
resource. This is despite the fact that they could
have chosen to be more mobile, travelling more
often between stone sources and the lakes. In this
latter case, features of such extreme reduction
would not be visible in assemblages.

Therefore, like Veth for the Western Desert
region, I see the main determinant of site
patterning in the Coongie and surrounding areas
as relating to the timing and intensity of site
use rather than to distance from raw material as
such.

This indication of relative residential stability in
the Coongie, combined with the high population
densities observed in the area at the time of
contact implies that the system may have been
vulnerable to significant stress given the extreme
environmental fluctuations characteristic of the
region. It is likely that the system worked

ARCHAEOLOGY OF THE MIDDLE COOPER BASIN

smoothly when resources were abundant but in
cases when an anticipated flow of fresh water
failed to come down the Cooper or when there
was widespread drought, it is possible there were
major problems. The fact that the current situation
where the Coongie captures most, if not all of the
Cooper flow, is potentially very unstable and
channel flow could be completely switched on
and off as a result of only slight environmental
shifts, would exacerbate this.

Ethnographic observations support the notion
that stresses were operating on the system. The
rigid sanctions outlined earlier involving the
enforcement of group boundaries and access to
resources indicate this, along with the
information that many people in what is now the
north-east South Australian region perished
during droughts. Further, Wasson's observation
of a dune mobilisation event in the late Holocenc
that might be linked to the intensity of the
occupation of the region, is another feature that
suggests the system might have been under some
stress.

Evidence for 'intensification '?

It is tempting to suggest that the characteristics
of the mid-late Holocene occupation of the region
reflect an intensive settlement pattern and are
derived from a process of 'intensification' (see
also Williams 1988: 61). For example, all of the
dates for occupation sites for the Coongie system
fall within the late Holocene and on all sites
recorded within the region, including JSN, the
typology of artefacts suggests a mid-late Holocene
date. While it is tempting to conclude the region
does provide evidence consistent with the
operation of such a process, as in my 1988 paper I
am cautious about maintaining that these features
provide definite evidence. I take this position
because it is likely that the environmental
instability could be causing major biases in the
archaeological record. Such instability could be
selectively removing older sites from the
archaeological record, or rendering them
archaeologically invisible through sedimentation,
or even resulting in vast fluctuations in channel
flow thus affecting the filling of lakes. I therefore
conclude that while the evidence for
intensification is persuasive, further
geomorphological work is needed to more fully
expand on factors affecting the visibility of
archaeological material.

Given these issues, can any conclusions can be
drawn about the occupation of the arid zone,
especially that of the more distant past?

Early Occupation Of The Arid Zone

1 will conclude the paper with some brief and
speculative comments about models relating to
when the arid zone was first occupied on a
consistent basis.

As noted in Smith, Williams and Wasson
(1991: 190) the evidence from the JSN site
indicates regular use of the dunefield region
during the late Pleistocene, when there was
probably also occupation of the riverine corridors.
The presence of Velesunio shell at JSN in a
Pleistocene context is tangible evidence of some
link with these riverine habitats. The nature of the
archaeological material present on the site
suggests that late Pleistocene use did not involve
much on-site stone working and that this changed
some time during the mid-late Holocene when
visits to the area appear to have become more
prolonged and to have involved the grinding of
seeds and the manufacture and maintenance of
chipped stone artefacts (ibid).

Dating of hearths to the period around 12 000
BP for hearths on the lower reaches of Coopers
Creek (Veth, Hamm and Lampert 1990) is also
consistent with late Pleistocene occupation of
what is now north-east South Australia.

Despite the presence of the dunefield site of
JSN, Veth (1993) has developed a model
suggesting that the continent's core arid zones
were only colonised as recently as 5 000 years
ago. Veth maintains that the evidence for
Pleistocene occupation of JSN does not disprove
his model since the Strzelecki dunefield region is
located within an area of co-ordinated drainage,
thus implying that it was not part of the core arid
zone (1993: 111).

Veth's model is an interesting one, but it could
prove difficult to test. His 'core arid zones'
comprise the deserts of the Great Sandy Desert,
Great Victoria and Simpson Deserts. These
regions are by definition sandy deserts and are
thus areas where outcrops of hard rock are rare
and therefore rock shelters with potential
Pleistocene sediments are either non-existent or
uncommon. Rockshelters such as those examined
by Veth (1993) in the Gibson Desert, have late
Holocene occupation resting directly on bedrock
and thus do not provide contexts for the
examination of occupation (or the lack of it) for
the period before the late Holocene.

The fact that the presence of rockshelter sites
which preserve Pleistocene sediments is so crucial
for the examination of whether Pleistocene
occupation took place or not, is shown by Veth's

E. WILLIAMS

discussion of his model (1993: Chapter 9). All of
the eleven Pleistocene sites and early Holocene
sites which lie a considerable distance from co-
ordinated drainage, and thus form the basis of his
model, are rockshelters.

Knowing that rockshelter sites which preserve
Pleistocene sediments are likely to be extremely
rare in the regions important for Veto's model, it is
also unlikely that Pleistocene open sites will be
easy to locate. Extensive outcrops of hard rock are
not common in the sandy deserts and thus stone
suitable for the manufacture of artefacts is also a
scarce commodity. It is therefore likely that stone
artefact manufacture, one of the maj or
archaeological indicators of human occupation, has
to take place on a relatively intensive basis before it
becomes archaeologically visible in these regions.
The JSN site for example indicates that Pleistocene
occupation is not as archaeologically visible as
regards stone artefact discard as is mid-late
Holocene occupation. This evidence is
complemented by my findings for the study area
that artefacts do not appear to be present within
visible Pleistocene contexts. This is despite that fact
that we know from the evidence of JSN that people
were using the region during the late Pleistocene.

Further, the logistical difficulties of carrying out
systematic archaeological surveys for sites in
remote locations such as sandy deserts, compound
these problems. JSN was not found as the result
of systematic archaeological sampling but was
instead located virtually by accident during the
course of a geomorphological study of the
dunefield by Wasson.

The question of when the core arid zones were
first occupied on a systematic basis thus remains
a challenging one. Until this question is resolved,
JSN remains the continent's sole example of a
Pleistocene site located within the midst of the

continental dunefield, albeit a dunefield in the
vicinity of co-ordinated drainage. The spur to the
finding of Pleistocene sites within Veth's 'core
arid zones' will be to expand further on the nature
of the continent's Pleistocene occupation and to
also determine the antecedents of the patterning
seen in the mid-late Holocene.

Acknowledgments

The project reported on in this paper was funded
jointly by the National Research Fellowship Scheme and
the Department of Prehistory, Research School of Pacific
Studies, (now the Division of Archaeology and Natural
History), the Australian National University. I thank
these bodies for support. The Australian Heritage
Commission also provided some assistance and I thank
Betty Meehan particularly in this regard. Bob Wasson
was instrumental in encouraging me to initiate the
Coongie project and I especially thank him for his advice
and support during the course of the work. SANTOS
provided much logistical support for fieldwork and I
particularly thank Oleg Morozow and Steve Tunstall for
their assistance. The Aboriginal Heritage Branch gave
permission for collection of sites and the excavation of
JSN. Chris Dodd (then Aboriginal Liaison Officer, SA
NPWS) gave approval for the excavation of JSN and I
especially thank him for his assistance with work on the
site. I wish to thank all of the people who provided
assistance in the field: Doreen Bowdery, Nick Drayson,
Ros Fraser, Parry Kostiglou, Gary Dunnett, Pam and
Colin McDonald, Ben Smith, Barney Stevens, Steve
Sutton, and Keryn Walsh. I am also indebted to Barry
Saunders and Vince for their advice and support in the
field, and the Royal Flying Doctor Service whose radio
network was invaluable in such a remote area. I
particularly thank my co-worker at JSN, Mike Smith;
useful advice and discussion on ideas in this paper were
provided by him, Bob Wasson and Barry Cundy. I am
also especially grateful to Win Mumford for her work
on the maps and illustrations.

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ARCHAEOLOGY OF THE MIDDLE COOPER BASIN

KREIG, G. W. & CALLEN, R A. 1990. Coongie Lakes
Control Zone - a map and note for the surface
geology and terrain. Mines and Energy Review,
South Australia 157: 59-61.

LAYTON, R, FOLEY, R, & WILLIAMS, E. 1991. The
transition between hunting-and-gathering and
specialized husbandry of resources. Current
Anthropology 32: 255-274.

MCBRYDE, I. 1987. Goods from another country:
exchange networks and the people of the Lake Eyre
Basin. Pp. 252-273 in 'Australians to 1788'. Ed. D.
J. Mulvaney & J. P. White. Fairfax, Syme and
Weldon Associates: Sydney.

MCKINLAY, J. 1862. 'McKinlay's Journal of
Exploration in the Interior of Australia'. F. Balliere:
Melbourne.

MACUMBER, P J. 1980. The influence of groundwater
discharge on the Mallee landscape. Pp. 67-84 in
'Aeolian Landscapes in the Semi-arid Zone of South-
eastern Australia. Eds. R. R. Storrier & M E
Stannard. Australian Society of Soil Science: Wagga
Wagga.

MEEHAN, B. 1982. 'Shelibed to Shell Midden'.
Australian Institute of Aboriginal and Torres Strait
Islander Studies: Canberra.

REID, J. R. W. & GILLEN, J. 1988. The Coongie
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Planning: Adelaide.

REID, J. R. W. & J. T. PUCKERIDGE. 1990. Coongie
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REVISION OF AUSTRALIAN AMPHIOPS ERICHSON, ALLOCOTOCERUS
KRAATZ AND REGIMBARTIA ZAITZEV (COLEOPTERA: HYDROPHILIDAE)

C.H.S. WATTS
Summary

The Australian members of the Hydrophilid genera Amphiops (five species), Allocotocerus (three
species) and Regimbartia (one species) are revised and redescribed. Keys to species of Amphiops
and Allocotocerus are given. Amphiops austrinus, Amphiops micropunctatus and Allocotocerus
yalumbaboothbyi are described as new.

REVISION OF AUSTRALIAN AMPHIOPS ERICHSON, ALLOCOTOCERUS KRAATZ AND
REGIMBARTIA ZAITZEV (COLEOPTERA: HYDROPHILIDAE)

C. H. S. WATTS

WATTS, C. H. S. 1998. Revision of Australian Amphiops Erichson, Allocotocerus Kraatz and
Regimbartia Zaitzev (Coleoptera: Hydrophilidae). Records of the South Australian Museum
30(2): 93-106.

The Australian members of the Hydrophilid genera Amphiops (five species), Allocotocerus
(three species) and Regimbartia (one species) are revised and redescribed. Keys to species of
Amphiops and Allocotocerus are given. Amphiops austrinus, Amphiops micropunctatus and
Allocotocerus yalumbaboothbyi are described as new.

C. H. S. Watts, South Australian Museum, North Terrace, Adelaide, South Australia 5000.
Manuscript received 15 April 1997.

The three genera included in this revision are
grouped for convenience not phylogeny although
both Allocotocerus Kraatz, 1883 and Regimbartia
Zaitzev, 1908 belong in the same tribe (Berosini)
and they and Amphiops Erichson, 1843 are
frequently mixed together in collections. All are
wet-tropical beetles with, at the generic level, a
wide distribution outside Australia (Hansen
1991). They are all relatively small very deep-
bodied insects with Amphiops and Allocotocerus
having almost spherical bodies. They occur
commonly in pools and swamps around the
tropical coast from the Kimberley to about
Sydney.

The most recent taxonomic work on the
Australian species was by J. Balfour- Browne who
briefly commented on them and described
Allocotocerus tibialis and Amphiops
queenslandicus (Balfour-Browne 1939) and
Hansen, 1991 who discussed their generic
placement.

Material was examined from the following
collections.

AM Australian Museum, Sydney

ANIC Australian National Insect Collection
BM(NH) Natural History Museum, London
NMV Museum of Victoria
NTM Northern Territory Museum

QDPIM Queensland Department of Primary

Industries, Mareeba
QM Queensland Museum, Brisbane

SAMA South Australian Museum, Adelaide
UQIC University of Queensland Insect

Collection, Brisbane
WAM Western Australian Museum, Perth.

Systematics

The three genera can be separated from other
Australian aquatic Hydrophilids by the following
characters (after Hansen 1991).

Amphiops: Size 2.5^.5 mm. Dark brown to
black. Eyes divided into upper and lower portions
by extensions of side of head. Elytra
approximately as high as long, without striae.
Meso- and meta-tibiae without swimming hairs.
First ventrite very short, with a fringe of fine setae
rising from its basal margin.

Allocotocerus: Size 3.5^.5 mm. Black. Meso-
and meta-tibiae with swimming hairs. Eyes
normal. Elytra as high as long, virtually without
striae.

Regimbartia: Size 3.5-5.0 mm. Black. Meso-
and meta-tibiae with swimming hairs. Eyes
normal. Elytra high, about 2.8x longer than
height, with distinct striae.

For more detailed descriptions and discussion
of the affinities of these genera see Hansen, 1991.

Amphiops Erichson, 1843

Australian Amphiops are all very similar and
are best separated by the male genitalia. Indeed I
have found it impossible to reliably separate A.
queenslandicus J. Balfour-Browne and A.
duplopunctatus Blackburn by any other means.
The best general character is the form of the
clytral punctation, in particular the interstrial
punctation on the sides of the elytra. There are
three more or less distinct size classes of elytral
interstrial punctures: 1) large, about the size of
the strial punctures; 2) small, between about 20-

C. H. S. WATTS

60% of the larger ones; 3) micro, which are
normally no more than pin pricks even under
moderate magnification.

Key to Australian Amphiops

Interstrial punctures consisting of a few large
punctures and a few to numerous micro
punctures, the small size seemingly absent
(Figs 1&3). Systematic punctures on pronotum
large and distinct 2

Interstrial punctures consisting of large, small
and micro punctures with small predominating
(Figs 2 & 4) 3

Central lobe of aedeagus not hooked (Fig. 13).
First elytral stria (close to suture) distinct,
traceable (anteriorly) well beyond apex of
scutellum. Scutellum always moderately
punctate A. micropunctatus sp.nov.

Central lobe of aedeagus hooked at tip (Fig.
14). First elytral stria virtually absent.
Scutellum lacking punctures or weakly

punctate with very small punctures

A. australicus Blackburn.

Central lobe of aedeagus hooked at tip (Fig. 9).
Small punctures on sides of elytra relatively
small, < 1/4 diameter of adjacent large
punctures (Fig. 4). Serial punctures easily
traceable at apex A. austrinus sp.nov.

Central lobe of aedeagus rounded, parameres
straight. Small punctures on sides of elytra
relatively large, many 1/2 size of adjacent
punctures (Fig. 2). Serial punctures may be
hard to trace at apex 4

Basal portion of aedeagus shorter than
parameres (Fig. 10). Size <3.9 mm. Elytra
often with a vague series of darker patches in
alternate interstriae

A. duplopunctatus Blackburn

Basal portion of aedeagus nearly twice length
of parameres (Fig. 11). Size > 3.5 mm. Elytra
without series of dark patches

A. queenslandicus J. Balfour-Browne.

Amphiops austrinus sp. nov.

Description (number examined 5) Figs 4, 9

Length 3.4-4.0 mm. Broadly oval, elytra deep,
almost as high as long. Reddish-brown, disk of
pronotum a little darker as are serial punctures.
Head broad, strongly punctate with variously
sized punctures, confluent at front. Pronotum
broad, punctures of various sizes, weak on disk

grading to strong laterally, systematic series
distinct. Elytron very weakly punctured near
suture, grading to strong laterally; in middle at
side serial punctures strong, large, somewhat
difficult to trace; large punctures separated by
own diameter or a bit less, small punctures, which
alternate with larger ones, are very much smaller;
interstrial punctures of three distinct sizes, largest
as large or larger than those in serial lines,
separated by own width or less, small punctures
more numerous, very much smaller, 1/4-1/3
diameter of large ones, in places third size group
of minute (micro) punctures present.

Aedeagus squat, basal portion shorter than
parameres. Apical portion of central lobe narrow,
weakly tapering, considerably shorter than
parameres, ending in small broad hook. Parameres
broad in basal half, narrow in apical, strongly
curved downwards near tip, tips truncated and a
little enlarged.

Distribution

Only known for the type localities in south-east
Queensland.

Types

Holotype: 'Brisbane 1/64, CW\ SAMA.

Paratypes: 4, 'Qld Pctrie, 10 km W, 23/1 1/95,
C. Watts', SAMA.

Remarks

In most characters similar to A. duplopunctatus
and A. queenslandicus. Apart from the male
genitalia A. austrinus differs from both of these
by the greater contrast in size between the large
and small interstrial punctures on elytra, caused
primarily by the comparatively small size of the
small punctures. In A. duplopunctatus and A.
queenslandicus the small interstrial punctures
become quite prominent towards the side and
apex, often reaching half the size of the larger
ones. In A. austrinus, apart from very near the
elytron edge, the small punctures remain much
smaller than the larger ones. The downward curve
of the parameres and the apical hook on the tip of
the central lobe of the aedeagus, also clearly set it
apart from these two species.

Amphiops australicus Blackburn, 1898

Description (number examined 234 ) Figs 3, 14

Length 2.7^.0 mm. Broadly oval, elytra deep,
about as high as long. Brown, sometimes with
distinct darker blotches on elytron, to black with

AMPHIOPS, ALLOCOTOCERUS AND REGIMBARTIA

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FIGURES 1^4. Pattern of punctures near anterior-lateral angle of elytron. 1, Amphiops micropunctatus; 2, Amphiops
duplopunctatus; 3, Amphiops australicus; 4, Amphiops austrinus.

some vague lighter areas. Head broad, moderately
to strongly punctate, punctures mainly small with
scattered much larger ones, becoming stronger
and denser in front, confluent and rugose along
front margin. Pronotum broad, weakly punctured,
punctures almost obsolete on disk but becoming
stronger laterally, much smaller than eye facet,
systematic punctures small, about size of eye
facet. Elytra weakly punctured on disk, becoming
stronger laterally. At side in middle, serial
punctures about puncture-width apart, lacking
alternating small punctures. Interstrial area usually
only with large punctures which are
approximately same size, and approximately same
size as serial punctures, about a puncture-width

apart or somewhat greater. Areas of elytra near
the scutellum and laterally may have indistinct
micro punctures.

Aedeagus elongate, basal piece about same
length as parameres. Central lobe narrow,
narrowing to point, tip with upward hook, shorter
than parameres. Parameres elongate, broad in
basal half, narrow in apical, tips weakly expanded,
curved downwards in apical fifth.

Type

Holotype female: '5074T', 'Adelaide R, NW
Australia, J. J. Walker', 'Blackburn Coll 1910-
236', 'Amphiops australicus, Blackburn', with
round type label, BM(NH). Seen.

C. H. S. WATTS

Distribution

Northern Territory

11 km SW by S Borroloola, ANIC; Cahills
Crossing, ANIC; 7 km NW by N of Cahills
Crossing, ANIC; Gimbar Stn, NTM; Howard
Springs, SAMA; Holmes Jungle, NTM; Jabiru,
NTM; Junction of Arnhem Hwy and Oenpelli Rd,
NTM; Kakadu NP, NTM; Koongarra, ANIC;
Magela Ck, ANIC; McArthur R, ANIC; 19 km E
by S of Mt Borradaile, ANIC; 30 km WSW Mt
Cahill, ANIC; 9 km N of E Mudginbarry HS,
ANIC; Nourlangie Ck, ANIC; 18 km E by N
Oenpelli, ANIC; Tindale, ANIC, NMV; UDP
Falls, NTM; 12°40S 132°54E, ANIC; 12°06S
133°04E,ANIC.

Queensland

12°08S 142°16E, ANIC; 12°22S 142°37E;
12°06S 142°33E, ANIC; 14°51S 142°58E,
ANIC; 18°33S 138°11E, ANIC; Archer Bend,
SAMA; Batavia Downs HS, ANIC; Barron
Falls, ANIC; Bertiehaugh Ck, ANIC; Cairns,
ANIC; 110 km S Coen, NMV; East Claudie R,
UQIC; 110km S Coen, NMV; 70 km SW
Greenvale, SAMA; Hann R, ANIC; Lakefield
NP, QDPIM; 50M N. Laura, UQIC; 72 km NW
Laura, ANIC; 21 km E Mareeba, QDPIM;
Moreton, ANIC; 2 km NNE Mt Tozer, ANIC; 9
km ENE Mt Tozer, ANIC; 3 km NE Mt Webb,
ANIC; 10 km WNW Rokeby, ANIC; Wenlock
R, ANIC.

Western Australia

14°25S 126°40E, ANIC; 14°52S 125°50E,
ANIC; 14°52S 125°50E, ANIC.

Remarks

Among Australian Amphiops, A. australicus
can only be confused with A. micropunctatus due
to the virtual lack of small and micro punctures on
the elytra. Amphiops micropunctatus has more
extensive micro punctures over the elytra but these
can be missed if the specimen is at all dirty or the
light poor. The two species are best separated by
the much more strongly punctate scutellum and
the quite well developed first elytral stria on A.
micropunctatus, compared to the virtually
impunctate scutellum region in A. australicus. I
have not seen any black specimens of A.
micropunctatus but light coloured A. australicus
are occasionally encountered, usually with well
marked dark patches on elytra. The tip of the
aedeagus in A. australicus is hooked, in A.
micropunctatus it is not.

Amphiops duplopunctatus Blackburn, 1898

Description (number examined 65 ) Figs 2,10

Length 3.0-3.9 mm. Broadly oval, elytra deep,
as high as long. Dark reddish-brown, elytra with
vague lighter patches. Head broad, strongly
punctate, punctures of varying size, often
becoming confluent along front margins.
Pronotum broad, weakly punctured on disc,
becoming stronger laterally, punctures of various
sizes, systematic series moderately distinct,
relatively small, about same size or a little larger
than eye facets. Elytra weakly and shallowly
punctured on disc, becoming much stronger
towards sides. At side, in middle, serial punctures
large, easy to trace, each separated by own width
or less, alternating with a small puncture.
Interstrial punctures of three sizes, largest a bit
smaller than serial punctures and most separated
by more than their width, small punctures much
more numerous and 1/3-1/2 diameter of large
ones, numerous and relatively large micro
punctures.

Aedeagus with basal piece shorter than
parameres, apical portion of central lobe
narrowing to thin point, lacking any apical
thickening or hook, shorter than parameres.
Parameres wide in basal half, narrow in apical
half, tips rounded not curved.

Types

Holotype female: '6370 Qu. T.', 'Blackburn
Coll 1910 236', 'Amphiops duplopunctatus,
Blackburn', BM(NH). Seen.

Lectotype: 'Queensland Blackburn's Coll.',
'Amphiops duplopunctatus BL. co-type',
dissected and remounted, SAMA. Seen.

Distribution

Northern Territory

Daly R Crossing, NTM, ANIC; Darwin,
ANIC, NTM, SAMA; 11km NE of E of
Darwin, ANIC; Gimbat Stn, NTM; Holmes
Jungle, ANIC, NTM, SAMA; Howard River,
NTM; Howard Springs, NTM, SAMA;
Humpty Doo, NTM, QDPIM; 10 km N of
Jabiru, NTM, QDPIM; 29 km W by N Jabiru,
NTM, ANIC; Keep River NP, NTM, ANIC;
Magela Ck, ANIC; 12 km NNW Mt Cahill,
NTM, ANIC; 19km WSW Mt Cahill, ANIC;
9km N by E of Mudginbarry, ANIC; Nabarlek
Dam, ANIC; 6km SW by S Oenpelli, ANIC;
Sth Alligator R, ANIC; 15 km SE Wangi,
NTM; Yungaburra, NTM, QDPIM; 16°02S
130°23E, NTM.

AMPHIOPS, ALLOCOTOCERUS AND REGIMBARTIA

FIGURES 5-8. Ventral view of mesofemur (top) and metafemur (bottom). 5, Male Allocotocerus yalumbabooihbyi,
6, Female Allocotocerus yalumbaboothbyi; 7, Male Allocotocerus tibialis; 8, Female Allocotocerus tibialis.

Queensland ANTC; Caloundra, SAMA; Claudie R, UQIC;

Bamaga, SAMA, UQIC; Barringtonia Lagoon Dalhousic R, ANIC; East Claudie R, UQIC;

W Cape York, QM; Batavia Downs HS, ANIC; Flinders Is, SAMA; 12 km SW Heathlands,

Bertihaugh Ck, ANIC; 8 km N Bluewater, ANIC; Holroyd R, ANIC; Iron Range, UQIC;

SAMA; 9km N Bluewater, SAMA; Burster Ck, Jordine R, UQIC; Kuranda, SAMA; Lawn Hill,

C. H S. WATTS

ANIC; Lake Barrine, AN1C; Lake Tinaroo,
ANIC; 73km NW of W Laura, ANIC; Lockerbie,
UQIC; 9 km NNW Lockhart R, ANIC; 18 km N
Mareeba, QDP1M; Mazlin Ck, QDPIM, ANIC; 3
km ENE Mt Tozer, ANIC; 9 km ENE Mt Tozer,
ANIC; 11 km ENE Mt Tozer, ANIC; Murphy's
Ck, UQIC; Normanby R, ANIC; Pascoe R, ANIC;
N Pine River, QM; Redcliffe, UQIC; Ripley,
SAMA; 27km NW by W Rokeby, ANIC;
Schramm Ck, ANIC; Townsville, SAMA, 20km S
Townsville, SAMA; 3 km WSW Ussher Point,
ANIC; Wenlock R crossing, ANIC; 18°35S
138°03E, ANIC; 18°40S 138°20E, ANIC.

Western Australia

Carson Escarpment, ANIC; Drysdaie R, ANIC;

Remarks

If reference material is available A.
duplopunctatus can be relatively easily separated
from A. austrinus by its stronger interstrial
punctation, but lacking reference specimens it can
only be separated with certainty from this species
by its aedeagus.

Typical A. duplopunctatus are smaller, higher
and often darker than A. queenslandicus and have
stronger large punctures on elytra. In many A.
duplopunctatus, particularly from the Northern
Territory, many of the large serial punctures on
the sides of the elytra are less than a puncture-
width apart, a condition I have not seen in any A.
queenslandicus. Except for particularly dark
specimens, A. duplopunctatus usually has quite
regular darker blotches in alternate elytral
interstriae variably distinct. Amphiops
queenslandicus lacks these and has more
uniformly coloured elytra although the punctures
are often darker particularly laterally. Amphiops
duplopunctatus is also smaller than A.
queenslandicus (<3.9 mm as compared to >3.5
mm).

Balfour-Browne (1939), considered A.
duplopunctatus to be possibly no more than a
subspecies of A. australicus, an opinion based
mainly on a perceived close similarity of the male
genitalia. The male genitalia of what I take to be
A. australicus and A. duplopunctatus are clearly
distinct (the types are female). The markedly
different punctation also differentiates the two
species.

Amphiops micropunctatus n. sp.

Description (number examined 30) Figs 1,13
Length 2.7-4.3 mm. Oval, elytron deep, a little

longer than high. Reddish-brown, punctures on
elytron darker. Head broad, moderately strongly
punctate, punctures tending to be either big or
small, smaller predominating, becoming
confluent on front margin. Pronotum broad,
moderately punctate, punctures of varying sizes,
stronger laterally, systematic series distinct,
comparatively large, many over twice size of
adjacent punctures, nearly twice size of eye facet.
Elytra weakly and shallowly punctured on disc,
becoming much stronger towards sides. Laterally
in middle serial punctures large, easy to trace,
about a puncture-width apart; interstrial area
shiny, punctures of two sizes, the larger sparse,
most > 2x width apart, somewhat smaller than
strial, the smaller, more numerous, very small
(micro), in contrast to other punctures, becoming
obsolete laterally.

Aedeagus with basal piece about same length
as parameres, parameres wide in basal half,
narrow in apical, tips broadly rounded, central
lobe narrow, tip bluntly pointed, reaching nearly
to end of parameres.

Remarks

In colour and size resembling A.
duplopunctatus but differing from it in punctation
and in the shape of the aedeagus which has the
central lobe larger than in A. duplopunctatus. The
absence of the 'small' size of interstrial punctures
in the species resembles A. australicus and readily
separates these two species from the other
Australian Amphiops which have the interstrial
areas much more punctate. From A. australicus
(and most other Australian Amphiops) it differs
by having the first (sutural) elytral stria quite well
developed as well as a greater development of
very small (micro) punctures on the elytra,
although an occasional A. australicus may have
quite extensive micro punctation.

This species is only known from a limited area
of North Queensland. Judging from the localities
it is possible that it is a closed forest species. The
specimens from the West Claudie basin and
Lockhart River were taken in October from still
pools in closed forest. No other habitat
information is available.

Types

Holotype male: 'Australia, N Qld 15 km WNW
of South Johnstone "Light Trap' 11 , 'Fay and
Halfpapp 1986', QDPIM. **

Paratypes: 2 same data as holotype, QDPIM; I,
'Australia N Qld Tolga 3', '1986 J. D. Brown
light Trap', QDPIM; 1, 'Cow Bay, N of Daintree

AMPHIOPS, ALLOCOTOCERUS AND REGIMBARTIA

FIGURES 9-14. Ventral view of aedeagus. 9, Amphiops austrinus; 10, A. duplopunctatus; 11, A. queenslandicus\
12, Regimbartia attenuata\ 13, Amphiops micropunctatus; 14, /*. australicus.

100

C. H. S. WATTS

N Qld, 15-22nd March 1984 I. C. Cunningham',
QDPIM; 2, '12°43S 143°17E, 9 km ENE of Mt
Tozer 5- 10th July 1986 T Weir and A. Calder\
ANIC; 3, '12.44S 143. 14E 3 km ENE of Mt
Tozer 28th Jun - 4th July 1986 T Weir and A.
Calder', ANIC; 10, '15°03S 145°09E 3 km NE
Mt Webb 30th April - 3rd May 1981 A Calder', 8
ANIC, 2 SAMA.

Distribution

Queensland

Dead Horse Ck, ANIC; East Claudic R, UQIC;
Gordon Ck nr Claudie R, UQIC; Iron Range,
UQIC; 9km NNW Lockhart R, ANIC; 3.5km SW
by S Mt Baird, ANIC; 2km NNE of Mt Tozer,
ANIC; 6km ENE Mt Tozer, ANIC; 11km ENE
Mt Tozer, ANIC; Mt Webb Nat. Pk, ANIC; West
Claudie R, ANIC.

Amphiops queenslandicus Balfour-Browne,
1939

Description (number examined 34 ) Fig. 1 1

Length 3.5-5.3 mm. Broadly oval, elytra deep,
almost as high as long. Dark reddish-brown, elytra
with vague darker/lighter areas and many serial
punctures black. Head broad, strongly punctate,
punctures of varying sizes becoming confluent
along front margin. Pronotum broad, moderately
punctate on disc becoming stronger toward sides,
systematic punctures moderate, about 3x size of
adjacent punctures, a little larger than eye facet.
Elytra with small but sharp punctures on disc
becoming much stronger laterally. At sides in
middle, large serial punctures relatively small,
most separated by more than their widths, small
alternating punctures relatively large, some
approaching 1/2 width of larger ones; interstrial
punctures of three sizes: large, larger than serial
punctures, separated by more than their width,
small much more numerous, relatively large, 1/4-
1/2 diameter of large ones, micro punctures
present over most of elytra.

Aedeagus elongate, basal piece nearly twice
length of parameres. Central lobe shorter than
parameres, narrowing to a point, apex not swollen
or hooked. Parameres broad basally, becoming
narrower apically, tips rounded, weakly curved
downwards.

handwritten locality (which I cannot read),
BM(NH). Seen.

Paratypes: 6, same data as Holotype (except
handwritten label and Type designation); 1, male,
'Rockingham austral', 'Australia' Amphiops
queenslandicus M. J. Balfour-Browne det., all in
BM(NH) with round paratype labels. Seen.

Distribution

Northern Territory

S Alligator R, QM; Black Point Coburg Pen,
ANIC; 5km NNW Cahills Crossing, ANIC; 7km
NW by N Cahills Crossing, ANIC; Coastal Plains
Research Station, ANIC; Daly R Mission, ANIC;
Darwin, SAMA; Fogg Dam, NTM; Howard
Springs, SAMA; Humpty Doo, ANIC; Jabiru,
NTM; Jim Jim, ANIC, NTM; Kakadu NP, NTM;
Kapalga, QM; Koongarra, ANIC; 12km NNW Mt
Cahill, ANIC; 19km WSW Mt Cahill, ANIC; 9km
N by E of Mudginbarry HS, ANIC; Oenpelli, AM;
6km SW by S Oenpelli, ANIC; 12°40S 132°22E,
QM.

Queensland

Ashgrove, QM; Ayr UQIC; Bowen, SAMA;
Brisbane, UQIC; Bulburin St Forest via Many
Peaks, UQIC; Caloundra, SAMA; Cooloola, QM;
East Claudie R, UQIC; Gatton, UQIC; Homehill,
SAMA; Ingham, UQIC; Ipswich, AM; Iron
Range, UQIC; Lockerbie UQIC; Many Peaks,
UQIC; S Pine R, QM; Ripley, SAMA;
Rockhampton, ANIC, SAMA; 20 km S
Townsville, SAMA; 37 km S Townsville, SAMA.

Remarks

The distinctive male genitalia readily identify
this species. Amphiops duplopunctatus is smaller
with the elytra not quite as deep, elytral punctures
weaker and usually differently coloured (see under
A. duplopunctatus).

Amphiops queenslandicus also closely
resembles A. austrinus but in this case the weaker,
smaller punctures on the elytra of A, austrinus
seem different enough to enable reliable
separation. However, the male genitalia should be
used wherever possible.

Regimbartia Zaitzev, 1908

Types

Holotype male: 'Queensland Australia',
'Amphiops queenslandicus Type J. Balfour-
Browne det.', with round Type label and a

Regimbartia attenuates (Fabricius,1801)

= Volvulus scaphiformis Fairmaire, 1879
Synonymy after d'Orchymont, 1932.

AMPH10PS. ALLOCOTOCERUS AND REGIMBARTIA

101

Description (number examined 89) Fig. 12

Length 3.6-5.0 mm. Narrowly boat-shaped,
high, sides of elytra subparallel, perpendicular,
black, shiny, appendages testaceous. Head
deflexed, eyes large, punctures moderate
scattered, weaker on disc, those forward of eye
about size of eye facet, systematic punctures
inward from eye relatively small, about size of
eye facet. Pronotum broad, weakly and sparsely
punctured on disc, more strongly towards sides
where they are about same size as on front of
head; systematic punctures few and hard to trace.
Elytra more strongly punctured, somewhat
stronger at sides than on disc, each elytron with
10 serial lines, inner two incomplete anteriorly,
for the most part serial punctures joined forming
sharp grooves. Mesosternum with strong narrow
keel, apical sternite with short sharp spine in
midline at apex.

Male: Protarsi with two basal segments
moderately expanded dorso-vcntrally.

Types

Hydrophilus attenuate! Fabricius. Ceylon. Type
not located.

Volvulus scaphiformis ? Holotype. Rockhampton
Fairmaire, ?in Museum Godeffroy. Not seen.

Distribution

Northern Territory

Bessie Springs, ANIC; 11 km SW by S of
Borroloola, ANIC; 30 km NE by E of Borroloola,
ANIC; 48 km SW by S Borroloola, ANIC; 1 km
N of Cahills Crossing, ANIC; 7 km NW by N
Cahills Crossing, ANIC; Daly River, NTM
Darwin, ANIC, SAMA; Elsey Creek, ANIC
Fcrgusson R, ANIC; Fogg Dam, ANIC, NTM
Howard Springs, SAMA; Jabiru, NTM
Katherine, NTM; Keep River NP, ANIC; Lake
Bennett, NTM; Magela Ck, ANIC; Manton
Reservoir, NTM; 2 km N Mudginberri HS, ANIC;
6 km N by E Mudginberri, ANIC; Nourlangie Ck,
ANIC; Roper R, ANIC; South Alligator R, QM;
U.D.P. Falls, NTM;

Queensland

Archer Bend, SAMA; Ayr, UQIC; Brisbane.
MV, QM, UQIC; Cairns, ANIC, QM; Calliope R,
ANIC; Caloundra, SAMA; Cape Flattery, DPIM;
Chillagoe, DPIM; Claudie R, UQIC; East Claudie
R, QIC; Clermont, AM; Cooktown, MV; 40M N
Cooktown, UQIC; Duaringa, AM; Edungalba,
ANIC; Einasleigh, DPIM; Frenchman's Creek via
Rockhampton, UQIC; Gin Gin, UQIC; Gregory
River Hotel, DPIM; Homehill, SAMA; Kirrama

Rng, QM; Karumba, UQIC; Lakefield NP, ANIC;
Mackay, SAMA; Mareeba, ANIC, DPIM; 18km
N Mareeba, DPIM; Mary Ck, ANIC; Mclvor R,
UQIC; Mt Garnet, SAMA; 3.2 km SW of Mt
Inkerman, ANIC; Mt Molloy, ANIC, QM; 3 km
ENE Mt Tozer, ANIC; 6 km ENE Mt Tozer,
ANIC; Nordcllo Lagoon, DPIM; 11km WSW
Petford, DPIM; N Pine R, QM, UQIC;
Rockhampton, SAMA; 67 km E Roma, QM;
Samford, UQIC; Silver Plains CP, DPIM; 15km
WNW South Johnstone, DPIM; Starbright HS,
DPIM; Strathmore Stn, DPIM; Tolga, DPIM;
Townsville, QM; Walkamin, DPIM; 40km S
Weipa, DPIM; Yaamba, UQIC; Yeppoon, UQIC;
18°38 S 138°11 E, ANIC; 18°34 S 138°08 E,
ANIC.

Western Australia

Channley R, ANIC; Derby, SAMA; Fitzroy R,
ANIC; 12 km S Kalumburu Mission, ANIC;
Kunanurra, DPIM; 1 km NNE Millstream, ANIC;
Synnot Ck, ANIC; 14°53 S 125°45 E, SAMA.

New South Wales

Eccleston, UQIC.

Remarks

Once known, this distinctive water beetle is
unlikely to be confused with any other Australian
species. It is relatively common in shallow
swamps and dams along the coast from northern
New South Wales to the Kimberlcy. Beyond
Australia the species has a wide distribution in
South-east Asia as far west as Sri Lanka.

I have followed d'Orchymont (1932 p. 709) for
the name of this widespread species.

Allocotocerus Kraatz, 1883

The three Australian species of Allocotocerus
(- Globaria Latreille) are all moderately sized,
highly spherical insects with the ventral surface
shiny black. Within Australia they can only be
confused with species of Amphiops, particularly
A. queens landicas which is of a similar size and
shape. The even, regular punctation, uniformly
black ventral surface, and swimming hairs on the
legs in Allocotocerus readily separate them from
Amphiops.

There is little to separate the species other than
the sexual characters of the aedeagus, trochanter
setae and the labrum.

The first described and best known species, A,
punctatus, is common and widespread in slowly

102

C. H. S. WATTS

moving water and in swamps in eastern Australia bit larger than size of eye facet, systematic

from northern New South Wales northwards. It punctures virtually absent. Pronotum narrower

appears to be absent from the Northern Territory than elytra, punctures on disc much weaker than

and northern Western Australia where its place is on head, becoming considerably stronger towards

taken by A. tibialis and A. yalumbaboothbyi. front comers, systematic punctures few, sparse

Conversely these species likewise have not been and hard to find. Elytra covered with very even

recorded from the east coast. punctures in same area but much stronger laterally

An additional species (the type species, A. than on disc, serial punctures 1.5-2x diameter of

bedeli Kraatz 1883) occurs in New Guinea, adjacent punctures, reduced to 2-3 short lines

Unfortunately the holotype does not appear to be midway along side of elytron. Midline of

in the Deutsches Entomologisches Institut in mesosternum with strong, tall spike between

Eberswalde. In its place is a label written by mesocoxae. First ventrite constricted in middle

Korschefsky in 1937 indicating it was lost (Lothar with rather wide strong midline carina.

Zerche pers. com.). The recognition of this species Metasternum with two strong carinae in midline

and any possible influence on the nomenclature with deep gap between them, the anterior one

of Australian species must await the collection of cylindrical and angled backwards, the posterior

further specimens from New Guinea. one anvil shaped in lateral view.

Male: Labrum prominent, front edge deeply

bifid. Protibia triangular in cross-section.

Key to Australian Allocotocerus Aedeagus with basal portion strongly bent, 2.5x

length of parameres which are broad for whole
Labrum strongly extended forward, strongly | engthj central lobe broad e t for short

blfld male ^ P mctatUS (B^kburn) narrow port]0n at apcx MeSQ _ and ^

Labrum normal, often hidden beneath clypeus, trochanters with widely scattered short, stout setae

not blfid - 2 on ventral face.

Protibia light yellow with darker apical portion Female: Labrum little exposed, front edge not

A.punctatus (Blackburn) bifid. Protibia of normal shape, contrast between

Protibia lighter than other legs but without dark a P ical P ortion and yellow on rest of proleg

darker apical portion 3 often less pronounced than in male. Meso- and

meta-trochanters with scattered, sparse, short,

Setae on outer posterior angle of ventral stout setae on ventral face.
surface ot mesofemur more developed than

those on metafemur (Figs 7 & 8). Aedeagus Type

with parameres broad with rounded tips, \j , . n^n«oi *™ », ™ *^*^ .***,>,

central [lobe sharply pointed, much shorter than Holotype: T2328' 'Blackburn coll 1910-236',

parameres (Fig. 16) Volutus punctatus, Blackburn', BM(NH). Seen.

A. tibialis (Balfour-Browne) _ .

„ c ., . Distribution

Setae on metafemur equally poorly or slightly

more developed than on mesofemur (Figs 5 & Queensland

6). Aedeagus with parameres narrowing to 8 km N Bluewater, SAMA; Cardstone, ANIC

point, central lobe broad with narrow spine at 40 km N Coen, SAMA; 70 km N Coen SAMA"

tip, as long as parameres (Fig. 17) Caloundra, SAMA; 29 km NW by W Cooktowni

A -y^rnbaboothbyi, V .no,. ANJC; 4Q km N Cooktown? SAMA; , ngham

ANIC; Lakefield NP, QDPIM; 18 km N Mareeba,
QDPIM, ANIC; Mary Ck, ANIC; Mc Ivor R,
ANIC; Townsville, MV; Moorehead R, ANIC;
Moreton, ANIC; Mt Coolum, ANIC; 20 km S
Townsville, SAMA; 37 km S Townsville, SAMA;
Weniock R Crossing, ANIC; 15°17S 145°10E,
ANIC.

1 —

2 —

Allocotocerus punctatus (Blackburn, 1888)

Description (number examined 112) Fig. 15

Length 3.5-4.5 mm. Oval, elytra high, height
only a little less than length. Black, shiny,
appendages lighter, yellowish with darker terminal
portion to tibia. Head narrow, front margin
broadly and quite deeply concave, sides of
clypeus narrowly margined, rather evenly covered
with moderate punctures which are weaker in
middle than at sides, those forward from eyes a

Remarks

In overall shape, punctation and colour closely
similar to A. tibialis and A. yalumbaboothbyi but
the male characters readily separate it from these
species. Male A. yalumbaboothbyi and A. tibialis

AMPHIOPS, ALLOCOTOCERUS AND REG1MBARTIA

103

FIGURES 15-17. Ventral view of aedeagus.
Allocotocerus valumbaboothbvi.

15, Allocotocerus punctatus; 16, Allocotocerus tibialis; 17.

lack the very distinctive labrum and protibia of
male A. punctatus. In A. punctatus the dark apical
portion of the protibia separates both male and
female from the other species which have more
uniformly coloured protibiae. In both male and
female A. tibialis, the number and density of setae
on the trochanters are much larger than in A.
punctatus.

The species is relatively common in coastal

dams and swamps from northern Southern
Queensland to Cape York, often found together
with the similarly shaped Amphiops.

Allocotocerus tibialis (Balfour-Browne, 1939)

Description (number examined 61) Figs 7, 8, 16
Length 4.5-5.0 mm. Oval, elytra high, height

104

C. H. S. WATTS

only a little less than length. Black, shiny,
appendages dark-testaceous. Head narrow, front
edge widely and moderately concave, sides of
clypeus margined, strongly punctured,
punctures in front of eyes larger than eye facet,
systematic punctures inwards from eye few and
hard to trace, punctures on disc somewhat
weaker than elsewhere, about size of eye facet.
Pronotum narrow, somewhat more weakly
punctured than head particularly on disc,
systematic punctures few and hard to find.
Elytron punctured rather more strongly than on
pronotum and head, punctures much weaker on
disc than on sides where they are strong and
less than a puncture-width apart. Serial
punctures few or absent, if present little larger
than adjacent punctures. Mesosternum with
moderately tall, sharp spine in midline. Midline
of metasternum with two distinct carinae,
separated by deep gap, anterior carina
cylindrical and angled strongly backwards,
posterior one broader and curving backwards
and upwards when viewed from side. First
abdominal ventrite narrowed in middle half,
midline with rather narrow and strong carina,
ventrites two and three narrowed in central half,
weakly bulbous/carinate in midline. Area
between ventrites deeply grooved. Fourth
ventrite weakly narrowed anteriorly, smooth,

Male: Labrum simple. Protibia a little flattened
and widened, tarsi unmodified. Mesotrochanters
stout with a well-marked group of long golden
setae on anterior apical ventral angle, inner half
of ventral face lacking punctures/setae.
Metatrochanters stout with a less well-developed
group of setae in same place, metatibia stout.
Aedeagus long, sinuate, relatively broad, basal
piece about 1.5x length of parameres, parameres
broad, subparallel, rounded at tips, central lobe
narrowing progressively to tip, shorter than
parameres.

Types

Holotype male; 'Adelaide River 92-2\ '4969',
'Globaria tibiale mihi J. Balfour-Browne det.'
with red type label, BM(NH). Seen.

Lectotypes: 1, 'Adelaide River, NW Australia
J. J. Walker', L G. C. Champion Coll. B,M. 1927-
409' with red (allotype) type label, BM(NH), 10
same data, in BM(NH); 3, same data as holotype,
BM(NH). Seen.

Distribution

Northern Territory

Gimbat Sin, NTM; 6 km E Humpty Doo,

QDPIM; Jabiru, NTM; 10 km N Jabiru, QDPIM;
Junction of Arnhem Hwy and Oenpelli Rd, NTM;
Magela Ck, ANIC; 19 km E by S Mt Borradaile,
ANIC; 8 km E Mt Cahill, ANIC; 46 km WSW Mt
Cahill, SAMA; Nabarlek Dam, ANIC;
Nourlangie, ANIC; UDP Falls, NTM; Wildman
R,NTM.

Remarks

Both male and female A. tibialis can be
separated from A. punctatus and A
yalumbaboothbyi by the more extensive
development of setae on the mesotrochanters. The
elytral striae are also usually much weaker or
absent in this species but at least traceable in the
others. The species has not been recorded from
the east coast and appears to have a rather
restricted distribution in coastal Northern
Territory. Here it occurs commonly, often with A.
yalumbaboothbyi.

Amphiops tibialis can also be separated from
the very similar A. yalumbaboothbyi by the
differently shaped posterior mesosternal
protuberance: in A. tibialis this is thicker and
curved upwards behind in lateral view whereas it
is straight in A. yalumbaboothbyi.

Allocotocerus yalumbaboothbyi sp. nov.

Description (number examined 51) Figs 5, 6, 17

Length 3.5^.5 mm. Oval, elytra high, only a
little longer than high. Black, shiny, ventral
surface and appendages dark-testaceous, proleg
yellowish. Head narrow, front margin broadly and
quite deeply concave, rather evenly covered with
moderate punctures which are a little weaker on
disc than at sides, those forward from eyes a bit
larger than size of eye facet, systematic punctures
virtually absent. Pronotum narrower than elytra,
punctures on disc much weaker than on head,
becoming considerably stronger towards front
corners, systematic punctures few and hard to
find. Elytra covered with very even punctures
which become much stronger laterally, serial
punctures reduced to 2-3 short lines on each
elytron towards sides in middle; midline of
mesosternum with tall sharp spike forward of
mesotrochanters, midline of metasternum with
two strongly raised carinae separated by short
deep gap, front carina in shape of backwardly
inclined broad spine, rear carina oval shaped on a
broad pedestal when viewed laterally. First
ventrite moderately constricted in middle with a
narrow but well marked central carina extending

AMPHIOPS, ALLOCOTOCERUS AND REGIMBARTIA

105

forward between metatrochanters. Second sternite
also constricted but without carina, fourth not
restricted. Sutures between ventrites wide, deep,
well-marked.

Male: Labrum small, usually contracted, not
bifid, protibia not expanded, basal two segments
of protarsi a little expanded with ventral hairs.
Mesotrochanters with a few scattered, short spines
on apical ventral surface. Mesotrochanters with
longer more numerous setae forming a sparse
brush along posterior ventral portion of
metatrochanter. Aedeagus long, sinuate with
relatively broad basal portion which is about 2.5x
length of parameres. Parameres narrow, evenly
narrowing to quite sharp tip. Central lobe as long
as parameres. In a number of specimens the
parameres are twisted and splayed out,
presumably an artefact of preservation, but one
that I have not seen in other species.

Female: Basal segment of protarsi not weakly
expanded, with ventral hairs. Mesotrochanters
with a few sparse, stout setae on ventral face at
apex, metatrochanter with similar setae restricted
to apical half.

Types
Holotype male: '12°22S 133°01E 6 km SW by

5 of Oenpelli NT, 30. v. 73, at light E. G.
Matthews', SAM A.

Paratypes: 4, 'W Australia Mitchell Plateau
14°40S 125°44E 23 Sept 1982 B. V. Timms\
SAMA; 4, '13°34S 132°15E\ 'Moline Rockhole'

6 km E by E of Mt Daniela 23. v. 1974, NT T.
Weir and T. Angeles', NTM; 6, '12°32S 132°50E
Koorgarra 15 km E of Mt Cahill NT, 15.xi.1972,
T. Weir and A. Allwood', NTM; 11, 'NT UDP
Falls, 18-19th July 1980, M. B. Malipatil
Freshwater Pool', NTM; 20, '12°49S 132°51E,
15 km E by N of Mt Cahill NT, 29.X.72, light
trap, E. Britton 1 , ANIC.

Distribution

Northern Territory

8 km E by N of Mt Cahill, NTM; 19 km E by N
Mt Cahill, NTM; Junction of Amhem Hwy and
Oenpelli Rd, NTM; Kambolgie Ck, SAMA;
Koongarra, NTM; Wildman R, NTM.

Remarks

Virtually indistinguishable from A. tibialis and
A. punctatus other than by sexual characters.
Males can be easily separated from A. punctatus
by the lack of a large bifid labrum, from A. tibialis
by the weaker development of setae on the
mesotrochanters. In A. tibialis the mesotrochanter
setae are longer and denser often to the extent of
matting together. They are also more restricted to
the apical portion of both meso- and meta-
trochanters than in A. yalumbaboothbyi.

Separation of female specimens is more
difficult. Female A, punctatus have the prolegs
pale yellow with a more or less obviously darker
terminal portion to the tibia. In both A.
yalumbaboothbyi and A. tibialis the tibiae lack
the darker apical portion. The setae on both meso-
and meta-trochanters of female A.

yalumbaboothbyi and A. punctatus are small and
sparse. In A. tibialis they are more developed
particularly on the mesotrochanter.

Acknowledgments

The curators of the collections listed earlier provided
ready access to specimens in their care. Ms D. Churches
and Ms C. Home typed the manuscript and Mr R.
Gutteridge prepared the drawings. Mrs M. Anthony and
Mrs J. Evans helped with library aspects. Dr E.
Matthews greatly improved the manuscript. I thank all
these people without whom my task would have been
much greater.

References

BALFOUR-BROWNE, J. 1939. Contribution to the
study of the Palpicornia Part III. Annals and
Magazine of Natural History 11(4): 289-310.

BLACKBURN, T. 1888. Notes on Australian
Coleoptera with descriptions of new species.
Proceedings of the Linnean Society of New South
Wales 111(2): 805-875.

BLACKBURN, T. 1898. Further notes on Australian
Coleoptera, with descriptions of new genera and
species. Transactions of the Roval Society of South
Australia 22: 221-233.

ERICHSON, W. F. 1843. Beitrag zur Insecten-Fauna
von Angola, in: besondere Beziehung zur
geographischen Verbreitung der Insecten in Afrika.
Archiv fur Naturgeschichte 9: 199-267.

FABRICIUS, J. C. 1801. Systema Eleutheratorum...I,
24+506pp. Kiliae.

FAIRMAIRE, L. 1879. Descriptions dc Coleopteres
nouveaux ou peu connus du Musee Godeffroy
(Hydrophilidae). Journal Museum Godeffroy 24:
80-83.

C. H. S. WATTS

HANSEN, M. 1991. The Hydrophiloid Beetles.
Phylogeny, Classification and a Revision of the
Genera (Coleoptera: Hydrophiloidea). Biologiske
Shifter 40: 367pp.

KRAATZ, G. 1883. Allocotocerus nov. gen.
Hydrophilidarum. Deutsche Entomologische
Zeitschnft 27: 14-15.

ORCHYMONT, A. 1932. Zur Kenntnis der

kolbenwasser Kafer (Palpicomia) von Sumatra, Java
und Bali. Archives Hydrobiologica. Supplement Band
IX (Tropische Binnengewassen 1 1): 623-714.

ZAITZEV, F. A. 1908. Catalogue des Coleopteres
aquatiques des families Dryopidae, Georyssidae,
Cyathoceridae, Heteroceridae et Hydrophilidae.
Trudy russkogo Entomologicheskogo Obshchestva
38: 283-420.

DOUGLAS MAWSON: THE GEOLOGIST AS EXPLORER

D. W. CORBETT

Summary

Douglas Mawson was a born explorer. His professional career as a geologist and academic in
Australia demonstrates this aspect of his character as clearly as his exploits in the Antarctic. Science
and exploration dominated his life and motivated his endeavours, and the inseparable link between
them was his pursuit of knowledge of glaciation-both ancient and modem. This paper reviews the
diversity and wide geographical extent of his geological investigations, principally in South
Australia, during the first half of the twentieth century at a time when the geology of large areas of
the State was still largely unknown. The manner of his achievement and his distinguished
contribution to the advance of geological knowledge in South Australia exemplify Mawson as the
explorer-geologist.

DOUGLAS MAWSON: THE GEOLOGIST AS EXPLORER

D. W. CORBETT

CORBETT, D. W. 1998. Douglas Mawson: the geologist as explorer. Records of the South
Australian Museum 30(2): 107-136.

Douglas Mawson was a born explorer. His professional career as a geologist and academic
in Australia demonstrates this aspect of his character as clearly as his exploits in the Antarctic.
Science and exploration dominated his life and motivated his endeavours, and the inseparable
link between them was his pursuit of knowledge of glaciation — both ancient and modern. This
paper reviews the diversity and wide geographical extent of his geological investigations,
principally in South Australia, during the first half of the twentieth century at a time when the
geology of large areas of the State was still largely unknown. The manner of his achievement
and his distinguished contribution to the advance of geological knowledge in South Australia
exemplify Mawson as the explorer-geologist.

D. W. Corbett, Honorary Research Associate, South Australian Museum. North Terrace,
Adelaide, South Australia 5000.

Douglas Mawson was a geologist with a broad
scientific background and diverse interests within
his discipline, as a review of his publications will
verify (see Innes and Duff 1990). Known and
honoured as one of the great Antarctic explorers,
the prime motivation behind his achievement was
the pursuit of scientific knowledge, which he
coupled with the innovative use of the latest
available technology. With the exception of Otto
Nordenskjold, geologist and leader of the Swedish
Antarctic Expedition (1901-03), Mawson was the
only science-trained expedition leader in the heroic
age of South Polar exploration and his approach
marked the beginning of the modern age of
Antarctic exploration.

Mawson's career as a geologist and his
scientific reputation were built in South Australia
during his many years in the geology department
at the University of Adelaide. While this paper
focusses on the academic aspects of Mawson's
life more than his Antarctic exploits, it is
impossible to disentangle the two strands of
endeavour. Down south he was the scientific-
explorer; in the arid outback of South Australia he
was the explorer-scientist. His major researches
inextricably linked the two vastly different
environments, for it was the opportunity to study
an ice-age at first-hand and to equip himself better
as an interpreter of past glaciations that inspired
Mawson to venture south with Shackleton in
1907,

The circumstances of time and place (early
twentieth century South Australia, much of it little

known geologically) and his life-long interests in
glaciation, ancient and modern, determined that
Mawson the geologist and Mawson the explorer
were one and the same and through his extensive
investigations in South Australia he personifies
both the adventurous and the enquiring spirit.

Mawson's Antarctic exploits have been
chronicled by historians, fellow-expedition
members and in his own classic account 'The
Home of the Blizzard'. His career as a
scientist, and notably an assessment of his
geological work, have been less well treated,
although aspects of them are covered in the
biography by his wife (Mawson 1962) and by
Alderman (1967); by Parer and Parer-Cook
(1983); in the biographical summary by Jacka
(1986) and by Sprigg (1986). This paper presents
a review, with emphasis on Mawson's career as a
geologist in South Australia and his contribution
to geological knowledge of the State.

Early Years

As one of the first generation of geologists
trained in Australia, Mawson brought an
Australian rather than European perspective to his
work. Although born in Yorkshire, England on
the 5th May, 1882, he was only two years old
when his family migrated to Sydney and he grew
up staunchly independent and thoroughly
Australian in outlook— as the promotion of his
Antarctic expeditions was to demonstrate. He

108

D. W. CORBETT

14°

16°

18 (

Torres
Islands

Q»° Banks

Islands

Espirit'
Santo

tu l ^A ™. IWaewo

SYDNEY

Pentecost I.

Malekula

CT^S Ambrim

o£>

Efate

100km

\^t\ \. j Eromanga

Tana

O Aneityum

I70 c

172 c

FIGURE 1. Locality map. The New Hebrides archipelago in the south-west Pacific.

DOUGLAS MAWSON GEOLOGIST

109

-5
u

110

D. W. CORBETT

entered the University of Sydney in 1 899 at the
age of 16, doing well in Physics, Chemistry and
Mining on the way to his Bachelor of Engineering
degree in 1901. But it was in geology that he
excelled, gaining first class honours in 1900 and
being awarded the petrology prize, and
subsequently, in 1905, completing a B.Sc in
geology. His inspiration came from the
charismatic personality and the dynamic teaching
of the Professor of Geology, T. W. Edgeworth
David, who in turn was quick to recognise the
abilities of the young science student. David was
to become Mawson's mentor, fellow expeditioner
and lifelong friend and confidant.

One of David's many preoccupations was
ancient glaciation. As a young geologist in Britain
he had followed with interest the debate on the
nature of the latest (Pleistocene) Ice Age and the
evidence of a much older glaciation (Permo-
Carboniferous) discovered in India. Working for
the New South Wales Geological Survey in the
late 1880s he confidently identified glacial
deposits in the Permo-Carboniferous rocks of the
Hunter Valley. In 1901 he visited Mount
Kosciusko where he immediately recognised the
work of glaciers in the tarns, moraines and ice-
sculpted valleys in the high country (David et al.
1901) — his views effectively settling a dispute
that had continued ever since W. B. Clarke had
reported evidence of glaciation there half a century
earlier. Meanwhile in South Australia Walter
Howchin had discovered glacial sediments, older
than any known to that time, in the Sturt Gorge
near Adelaide (Howchin 1901). These deposits
(the Sturt Tillite) were assigned to the Cambrian
but were later shown to be even older (late
Precambrian), extending the evidence of past
glaciation in both time and space and acting as a
spur to the search for further clues to a former
frigid earth.

David was Australia's first 'national'
geologist, with interests spread well beyond his
base in New South Wales. He had particularly
strong links with South Australia through Walter
Howchin, with whom he had collaborated in the
field on such diverse matters as Cambrian fossils
and glacial geology. It was perhaps inevitable
that Mawson should become interested in these
new developments which were extending the
frontiers of Australian geology but his studies
initially led him in other directions. In 1902 he
was appointed Junior Demonstrator in chemistry
and the following year he took six months leave
of absence to make a trip to the South West
Pacific.

The New Hebrides 1903

Mawson's first opportunity to explore and
engage in adventurous fieldwork was due to the
initiative and recommendation of David, who had
a long-sustained interest in the geology of the
Pacific basin. It was to prove a testing assignment
for Mawson, requiring him to work under arduous
and at times dangerous conditions, making the
most of general short excursions from the ship as
it sailed through the island archepelago, strung
out over a vast area of ocean. Little known
geographically away from the coastal fringe, the
interiors were rugged and thickly covered with
dense rainforest making access difficult while the
native inhabitants had a reputation for being
unpredictable and at times hostile.

Apart from a scanty record of previous
investigations, the New Hebrides (Fig. 1) were a
geological 'terra incognita' and for Mawson a
challenge he was eager to meet: 'A great field
was now open before us ... nothing has yet been
gleaned as to the true nature of the rocks forming
this extensive chain of islands'.

Mawson (and W. F. Quaifc, a biologist) arrived
at Vila, on the island of Elate and the chief town
in the group on 13 April 1903. His first
observations were made in the low hills around
the town where a raised coral reef overlay volcanic
sediments. This general geological situation was
confirmed at locations further north along the
coast where much thicker reef deposits overlay the
'foundation rocks' — a variable series of tuffaceous
volcanics including agglomerates and pumice-rich
beds (Fig. 2).

From Efate the ship steamed north to Malckula
where Mawson wrote: 'The natives of Malekula
are the most uncertain of any of the inhabitants of
the group, having not yet abated from cannibal
habits; indeed war was raging between the
Natives ... at the time of our visit. For this and
other reasons wc were unable to examine this
region very thoroughly'. Nevertheless he was able
to make some investigations and excursions into
the interior. Reaching Espiritu Santo, the largest
island, a base was established at the mission
settlement of Tangoa on the south coast and an
assault made on Losumbuna (5 520 feet), the
highest mountain in the New Hebrides, The
interior mountains had never been explored by
Europeans and the challenge was irresistible.
Accompanied by three native guides, Mawson and
Quaife negotiated two subsidiary peaks before bad
weather, difficult terrain and, crucially, their
guides' refusal to continue, forced a retreat.

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Although disappointed at not reaching their
objective, Mawson collected specimens and made
sketch sections of the rugged hinterland (Fig. 3).
Fieldwork proved more productive along the coast
where three separate pyroclastic units intruded by
a variety of igneous rocks were found. An earlier
report of older basement rocks (gneiss) in this part
of the island could not be confirmed despite
diligent searching.

The northern-most islands in the chain were
covered in a cursory reconnaissance, but enough
to confirm the volcanic nature and essentially
basic composition of the Banks Group. Arriving
back in Vila in mid-August after four months
away, the expedition was greeted by a minor
earthquake with another tremor following a few
days later — enough to indicate the seismic
instabilty of the region. Finally, investigations in
the Havannah Harbour area provided key sections
in the local sequence of volcanic sediments and
overlying reef deposits.

The New Hebrides expedition was inevitably
very much a reconnaisance and Mawson was
keenly aware of the limitations of his survey. One
of the strengths of his published report (Mawson
1905) was his inclusion of all previous references,
both published and unpublished, to the geology
and physical features of the islands. The general
seismicity of the region, already well documented,
and which Mawson himself experienced, he
attributed to local volcanic sources, recognising
also the importance of regional tectonic
movements along major faultlines. He wrote:
'before long numerous seismological stations will
be distributed through the South Pacific Islands,
as only by analysing such data as would result,
can definite decisions regarding the present earth-
movements in this much-troubled area be arrived
at*. It was to be fifty years before that hope began
to be realised.

An anticipation that because the old rocks of
nearby New Caledonia were mineral-rich, the New
Hebrides might be similarly endowed was quickly
dispelled because of the geological youthfulness of
the islands. Sulphur deposits and iron-bearing
sands were the only resources considered to be of
possible future commercial value. Within the
regional context of the Pacific basin, Mawson
viewed the New Hebrides island arc as of late
Tertiary age, one of a series of parallel arcs which
were remnants of a much fragmented continental
mass. Folding and over-thrusting, associated with
volcanicity, against a rigid foreland lying to the
west (New Caledonia), had been followed by major
faulting and sea- floor subsidence.

The evidence for elevation which most
impressed Mawson were the raised coral reefs.
These capped the Tertiary deposits and ranged in
terraces rising to over 2 000 feet (609 metres).
They were estimated to be post-Pliocene in age,
and gave a clear indication of continued tectonic
activity since the middle Tertiary.

His published account of the New Hebrides
expedition put the islands on the map geologically
and established Mawson as a resourceful and
perceptive field worker, competent in many areas
of geology and with a flair for synthesis. He
foreshadowed many of the more recent
interpretations of south-west Pacific region
geology made in the light of Plate Tectonic theory,
with the New Hebrides viewed as an island arc of
volcanic lavas and pyroclastic rocks and intrusions
adjacent to a deep ocean trench and above a
steeply dipping seismic zone.

Early Researches

On returning to the University, Mawson was
soon involved in research (with fellow student
and later Antarctic explorer Griffith Taylor) on
the igneous rocks in the Bowral-Mittagong area,
south-west of Sydney. The Mittagong study
(Taylor and Mawson 1903) was the first of many
on the petrography and petrology of igneous
rocks in their regional setting which was to
remain a prime interest in the years ahead. About
this time Mawson also began investigations into
the radioactive properties of minerals. This was a
pioneer study in Australia, closely following the
discovery of radioactivity by Becquerel in 1896
and the follow-up work by the Curies which led
to the isolation of radium in 1898 and the
recognition that uranium and thorium-bearing
minerals were radioactive. Mawson reviewed
these recent researches and carried out tests on
natural waters, mud, soil and air. He was
required to construct his own analytical
equipment, notably a gold-leaf electroscope, and
a large number of Australian minerals was tested
including the only two known occurrences of
uranium-bearing minerals in Australia and which
gave the predicted high readings. The other
minerals found to test positive were mainly
monazites from various tin- fields around
Australia. His short paper (Mawson & Laby
1904) marked the beginning of a life-long
interest in the geological occurrence of
radioactive minerals and their economic
potential.

DOUGLAS MAWSON GEOLOGIST

113

South Australia 1905

Late in 1904, Mawson, with the support and
encouragement of David, was appointed to the
position of Lecturer in Mineralogy and Petrology at
the University of Adelaide. He arrived to take up
his academic duties full of enthusiasm and with
high expectations but was immediately confronted
with a lack of suitable accommodation and facilities
for geology due to the University's chronic shortage
of funds. Frustrations developed which were to dog
him for many years, including the necessity to forge
a working relationship with the only other member
of the Geology Department, Walter Howchin.
Already with an established international reputation
and with a great deal of experience of South
Australian geology, Howchin was 60 years of age
when Mawson, 23, arrived on the scene. It seems
there were tensions from the start. The question of
'spheres of activity' (Mawson's phrase) provided
an immediate and ongoing irritant, to be
temporarily removed when Mawson went south in
1907.

The geology of South Australia was known in
broad outline at the beginning of the century
(Corbett et al 1986) This was due predominantly
to the work of three men: Ralph Tate, the first
geologist appointed to the University (1875-
1901); Walter Howchin, Methodist Minister and
enthusiastic amateur until his appointment to the
University after Tate's death in 1901; and H. Y.
L. Brown, the first Government Geologist who
was to act as a one-man Geological Survey from
1882 until 1912. While Tate and Howchin worked
mainly, though not exclusively, close to home in
the Mount Lofty Ranges, Brown had been
carrying out some epic geological explorations in
the northern regions of the State which at that
time included the Northern Territory. He
published the first geological map of South
Australia in 1883, revising it periodically. His
1 899 map was a remarkable achievement and in
its essentials remains little changed today.

Tate had discovered glacial features at Hallett
Cove in 1877; Howchin subsequently
demonstrating the widespread evidence for this
glaciation outside the Adelaide region and
establishing a Permo-Carboniferous age for the
event by about 1900. In 1901 Howchin, as
mentioned previously, had discovered the much
older glacials in the Sturt Gorge, a few miles
south of Adelaide. But perhaps the most
significant of all the early breakthroughs was the
finding of Cambrian fossils in the old rocks on the
western flank of the Mount Lofty Ranges

(Howchin 1897). Howchin thereafter concentrated
much of his effort on determining the stratigraphy
of the western ranges.

It was Howchin' s clearly expressed
'prerogative' to work in most areas of the State
that prompted Mawson to choose the remote
Olary-Broken Hill area for his first extensive field
investigations. It was a decision both judicious
and timely. Not only did this work extend
Howchin' s researches on the supposed Cambrian
rocks of the Mount Lofty Ranges into the Olary
Spur and over the border to Broken Hill and the
Barrier Ranges, but the problems presented by the
region also satisfied his interests in mineralogy,
petrology, and stratigraphy and his discoveries
were to influence his future involvement with
glacial deposits both ancient and modem.

The explorer-geologist was severely tested in
the harsh environment near the north-east state
border. It was (and is) an arid, inhospitable and
sparsely inhabited country. Previous investigations
had centred on and around the Broken Hill ore-
body itself — the largest silver-lead ore body in the
world. While little was known of the origin of the
ore-body and its stratigraphic controls, Mawson
set himself the task of ascertaining the broader
regional context of the deposit. His methods
involved covering the ground in a series of
traverses on foot, horseback, bicycle and on
occasions, by motor vehicle. He was to determine
and define two major rock divisions in the Olary-
Broken Hill region:

1. An older basement complex of metamorphic
rocks including gneiss, schist, amphibolite and
granite of Precambrian age which he named
the Willy ama Complex, and

2. A younger series of sedimentary rocks, in part
metamorphosed, of assumed Cambrian age
including a wide variety of rock types, the most
notable and persistent being boulder-beds and
related sediments of glacial origin. This he
named the Torrowangee Series.

In January 1907, the Australasian Association
for the Advancement of Science (AAAS), held its
meeting in Adelaide and Mawson gave two
(unpublished) papers based on his recent field
work and particularly concerned with the glacial
beds. His interest in these beds and a desire to
reach a full understanding of their nature and
origin was developing rapidly. Many years later
he wrote to Edgeworth David that his commitment
to glacial geology dated from December 1906
when his Olary field investigations were well
underway. Matters moved very rapidly at the end
of 1907. Mawson heard, through David (who was

114

D. W. CORBETT

to join the British Antarctic Expedition), that the
leader, Ernest Shackleton, was to visit Adelaide
briefly on his way to New Zealand to join the
'Nimrod'. Mawson successfully requested a
meeting and volunteered his services for the
summer season. Within three days Shackleton had
offered Mawson the position as physicist for the
duration of the expedition. The opportunity to
realise his ambition to experience 'an ice-age in
being' was too good to miss. He accepted, quickly
arranging for a ' locum' to cover his absence from
the University, and left Adelaide before the end of
December bound for the Antarctic.

South With Shackleton - the British Antarctic
Expedition

David was senior geologist on the expedition
with Raymond Priestley, a young man with no
formal qualifications in science, as his deputy.
Mawson as physicist had no official role in this
area. Yet from the start David involved his old
student in the geological program and in two of
the three main journeys undertaken by the
expedition — the ascent of Mount Erebus and the
trek to the South Magnetic Pole. Mawson played
a crucial role, both as a young and vigorous team
member and a valuable contributor to the
geological work, which David was quick to
acknowledge in his final report (David & Priestley
1914). Here Mawson is referred to as
Mineralogist, Chemist and Physicist to the
expedition (the study of Antarctic ice — its
crystallisation, granulation and other properties- —
was considered by David as much a matter of
mineralogy as of physics). The report incorporates
much of Mawson's thinking and his contribution
to it was wide-ranging, covering mineralogy and
petrology and stratigraphy, as well as economic
geology implications and broad- scale
considerations of Antarctic geology within a
global context.

Mawson and David were both in the small party
which reached the summit of Mount Erebus
(3 794 m) on 8 March. Mawson made a traverse
map of the eight day journey which provided its
share of adventures (Shackleton 1911). He also
took photographs and collected rock specimens.
Later that winter (June 14) the volcano was seen
in eruption from their base, steam issuing from
the crater, although no lava was observed on the
flanks of the cone.

With the coming of spring, preparations were
made for the two major assaults of the expedition;

Shackleton's push for the geographic pole and an
attempt by a northern party (David, Mawson and
McKay) to reach the South Magnetic Pole. The
route was along the coast of Victoria Land as far
as the Drygalski Glacier Tongue, where they were
to turn inland climbing onto the Polar Plateau
towards the pole. A general survey of the geology
of the coast and where possible of the Western
Mountains was an important part of the program.
The early stage of the journey was along the edge
of the sea-ice with outcrops confined to occasional
rocky bluffs projecting from under the ice-cap.
Observations were made at Cape Bernacchi
(marble), Cape Irizar, Granite Harbour and Depot
Island. The rocks encountered were highly altered
(metamorphic) and included gneiss and schist
together with granite (itself metamorphosed) and
basic lavas. Looking inland through field-glasses,
these lavas were seen at a higher level to overlie
the granites and in turn to be succeeded by
stratified rocks which were taken to be the Beacon
Sandstone, discovered and described by Ferrar
(1907) on Scott's first expedition (1901 03).
David described Depot Island as \.. most
wonderful geologically and a perfect elysium for
the mineralogist'. Mawson relished the variety of
rocks and minerals they found, among them
epidote, pyrite, copper pyrites (chalcopyritc),
garnet and manganese oxides, some of which he
thought might prove of economic value. Certain of
the rocks, notably the gneiss and granite reminded
him of those in the Broken Hill area. Meanwhile
the ubiquitous glacial debris (moraine) dumped at
the melting ice-front was a rich source of
supplementary evidence of what lay further inland,
below the ice-cap. Erratics at one locality were
very similar to the Victor Harbor Granite.

The ascent of the plateau became an ordeal and
any thoughts of geology were replaced by
concentrating on the dash for the magnetic pole.
The elusive and ever-wandering point was reached
on 16 January 1909. The return journey proved a
battle for survival and the party was very fortunate
to make the rendezvous with the 'Nimrod' and
safety.

The information collected by the Magnetic Pole
parly made a valuable contribution to the
geological results of the expedition. Notably,
David and Mawson were able to build on Ferrar's
earlier work and show that Victoria Land
comprised a basement complex of metamorphic,
igneous and mineral-rich rocks of assumed
Precambrian age, overlain by a sedimentary
sequence, which, on the limited evidence
available, included rocks intermediate in age

DOUGLAS MAWSON GEOLOGIST

115

FIGURE 4. Mawson, seated, on his return from Antarctica, 1909. Howchin stands at his right shoulder. Photo: P.
Driver-Smith.

between the basement and the Beacon Sandstone.
For Mawson the expedition proved a
stimulating and scientifically rewarding
experience. His desire to experience modern
glacial processes in action had been achieved and
the insights gained were to be developed on his
next journey south and later applied to the ancient
glacial deposits of South Australia. Above all, his
participation in the Shackleton expedition had
determined his lifelong interest in the southern
continent.

Interlude

Mawson returned to Adelaide in April 1909 to
a hero's welcome (Fig. 4), but within a few weeks
he was back in the Olary-Broken Hill country,
hiring a horse and dray and continuing his
investigations among the mineralised rocks and
the ancient glacial beds of the arid north. After
completing his fieldwork, he spent some time in
Broken Hill writing up his results, and on
laboratory studies of the mineralogy and
petrography of the key rock types. He was
particularly impressed by the pegmatites which
were such a prominent feature of the region and
wrote: 'In no other part of the world can pegmatite
formations occur on a more extensive scale'. The

study of pegmatites was at that time in its infancy
and Mawson recognised that in Olary he had
evidence of their importance as 4 a connecting link
between the igneous rocks and the differentiated
products economically worked as ore deposits'.
His fieldwork, and the subsequent published
review (Mawson 1912a; Fig. 5) of all aspects of
the geology and economic geology of this
important and yet little known mineral province,
was a major contribution to South Australian
geology and earned him the degree of D.Sc. from
the University of Adelaide in 1909.

While in Broken Hill he examined and valued
several mineral collections from the mines on
behalf of the South Australian Museum. Some of
them included specimens of outstanding quality.
Mawson had first acted for the Museum in 1906,
when he had successfully acquired the Dunstan
Collection from the Wallaroo Mines. In 1908 he
was appointed Honorary Curator of Minerals, thus
beginning a long and productive association with
the Museum which was to last his lifetime, and
which became crucial in later years, as no full-
time curator responsible for the collection was
appointed until 1956. The Mawson-Museum
connection cannot be treated in any more detail
here.

Early in December 1909, Mawson left Adelaide
for London on board the SS 'Mongolia'. He was

116

D. W. CORBETT

FIGURE 5. MawsoTTs interpretation of the geology near
Olary, South Australia (1912b). Courtesy ANZAAS.

keen to go to the Antarctic again and full of ideas
for a scientific program that he hoped would
appeal to either Shackleton or Scott, both of whom
he knew had plans for expeditions south. Scott
was very keen to enlist Mawson (even offering
him a place in the final polar party), but was not
prepared to consider any modification of his own
plans which were already well advanced.
Discussions were long and frank but when
Mawson learned that the position of chief scientist
(which he might have accepted) had been filled
and his proposal for a western party to explore
King George V Land unacceptable, he declined
Scott's offer. He was simply not prepared to
compromise his purely scientific approach to
Antarctic exploration. As for Shackleton, initially
Mawson maintained friendly relations with his
former leader, even visiting a gold prospect in
Hungary which Shackleton was convinced would
make his fortune (and Mawson's too if he came in
with him as an investor). But becoming
increasingly disillusioned with Shackleton's
erratic behaviour and lack of firm plans, Mawson
followed him to America in an attempt to pin him
down; and the two signed an agreeement that an
expedition would leave late in 1911 and should
Shackleton be unavailable as leader Mawson was
to take over and receive all Shackleton's support
and assistance in raising funds. Ever the realist, it
was clear to Mawson that the responsibility for
mounting the expedition was most likely to be his
and so it proved. He was then faced with the
daunting task of raising funds quickly and in
competition with Scott who had a head start and
the advantage of being based in London. Mawson
returned to Adelaide. For the next eight years the
Antarctic was to dominate his life. But for a brief
interlude he returned to academic life and his
research activities and to these we now return.

Radioactive Minerals: Radium Hill and Mount
Painter

Mawson's involvement with radioactive
minerals had been renewed soon after his arrival
in South Australia when in May 1906 carnotite, a
highly radioactive mineral, was found in ore taken
from a mine 24 miles east-south-east of Olary. H.
Y. L. Brown visited the site and issued a report in
which he speculated that the mineral was a
decomposition product of primary uranium ores
occurring at depth. In August, Mawson obtained
samples which he passed over to E. H. Rennic,
Professor of Chemistry, for analyses which

DOUGLAS MAWSON GEOLOGIST

117

s g

FIGURE 6. Geological cross-section in the Mt Painter
Arkaroola area (1912b). Courtesy ANZAAS.

suggested that black minerals initially identified
as magnetite, were far more complex. This
prompted Mawson to visit the mine and make a
detailed survey of the exposed reefs which had
been pegged originally in the belief that the
prominent black mineral was a tin-bearing ore.
Further examination showed that while the
magnetic iron/titanium mineral was dominant in
the main reef, a rarer black mineral with a
distinctive and very brilliant lustre also occurred
as grains, streaks and cubic crystals. Analysis
shows that in addition to a high percentage of iron
and titanium the mineral also contained rare earth
elements together with uranium, vanadium and
chromium. Mawson proposed the name Davidite
for what he believed to be a new mineral species
in honour of his former teacher (Mawson 1906).
He also identified Roscoelite, a bright green
vanadium-bearing mineral. At a time when
knowledge of radioactive minerals was in its
infancy, Mawson recognised the deposit as a
significant discovery, possibly of economic
importance, and named the locality Radium Hill.
He was optimistic but cautious in his appraisal:
'This body of radio-active ore is, in the matter of
quantity, much the most important yet discovered
in Australia. Its low grade, however, introduces
serious difficulties to commercial enterprise in this
direction'. Radium Hill was to have a long but
intermittent history based on the working of its
radioactive lodes (Fig. 7), and Mawson was to
renew his interest in the deposit in the 1920s.

Mawson visited the Flinders Ranges for the first
time towards the end of 1910, during his short
spell in Adelaide between Antarctic expeditions.
He had received a visit from W. B. Greenwood,
the owner of Umberatana Station, who had
brought some specimens of a bright green mineral
which Mawson identified as torbernite, an
hydrated phosphate of copper and uranium.
Greenwood had been prospecting in the northern
ranges on behalf of the Mines Department for
several years and had become annoyed when
specimens, including the green mineral, which he
had left at the Department some time before had
not received attention; whereupon he had retrieved
them and taken them to the University. Excited by
the prospect of another radioactive mineral deposit
in South Australia, Mawson headed north and,
with Greenwood and renowned prospector Harry
Fabian, made an examination of the geology and
mineral occurrences in this most rugged and
inaccessible part of the Flinders, attempting to
place the local geology in the broader regional
context (Fig. 6). The short visit marked the

118

D. W. CORBETT

beginning of his long and fruitful association with
the geology and mineral resources of the region
and his field notebook for the trip is full of detail
of his observations and the excitement of
discovery; for example: 'The results of my recent
journey have been in the highest degree
satisfactory... My main objective was the
investigation of the metamorphic rocks eastward
of Yudnamutana in connection with the
interpretation of the Barrier Ranges, a subject in
which I have long been absorbed. . . '

With regard to the potential of the uranium-
bearing lodes he wrote : '... the outcrop is on the
whole low grade, though richer patches are met
with at intervals. Improvement may be expected
below. The ore can be treated very inexpensively
and this will offset the low-grade character. So far
as I am aware this is the most extensive
uraniferous lode formation in the world'. The back
of his field notebook contains estimates of costing
for ore extraction, transport, freight and handling,
including comparisons for treatment on the spot,
at Port Pirie or in England. There would be a need
for roads and wells and for camels to bring out the
ore. If the grade proved high enough, he foresaw
the setting up of a company to work the ore — the
Radium Extraction Company. He even worked out
the capital required - £20 000. These estimates
may have been doodlings around the campfire, at
the latest they were put on paper soon after his
visit, and they have an immediacy which reveal
Mawson's keen interest in the region, its
economic potential and his desire to be part of the
action. He was not to return north for more than a
decade. Over the next few years some intermittent
mining activity took place at Mount Painter, but
with the outbreak of World War I activity ceased.

South Again - The Australasian Antarctic
Expedition (AAE) 1911-14

In January, 191 1, Mawson gave a lecture to the
combined geology and engineering groups at the
annual meeting of the AAAS in Sydney. It was a
fund raising exercise designed to appeal to
national pride as well as pointing out the scientific
and future economic potential of the southern
continent. He stressed that: 'the collection of
scientific data (was) obligatory upon us ... its
mountains were blessed with mineral resources ...
(and) it holds among its fossil strata secrets
especially interesting to Australians'. Science was
to be linked to technology and the latest
developments — the wireless and motor sledges —

still to be tested down south, were seen as a vital
aid to scientific exploration. The burdens of
leadership were enormous leaving little time for
any geological researches of his own, but Mawson
recruited a strong team of four geologists who
were to carry out valuable surveys over a great
expanse of territory. Cecil Madigan, later to join
his leader at the University of Adelaide, was
appointed meteorologist.

Reaching the area selected for the expedition's
base, the expedition ship 'Aurora' had great
difficulty in breaking through the ice barrier and
was forced to cruise westward; Mawson had
hoped to find the Antarctic continent in these
latitudes bounded by a rocky and attractive coast
like that in the vicinity of Cape Adare, the nearest
well-explored region. The poor prospects of
carrying out extensive geological work under such
conditions must have been a blow, but Mawson
was impressed and awed by the sheer physical
presence and power of the ice and its dominance
of the environment: 'The land was so
overwhelmed with ice that, even at sea-level, the
rock was all but entirely hidden. Here was an ice-
age in all earnestness; a picture of northern Europe
during the Great Ice Age some fifty thousand
years ago' (Mawson 1915, 1996 p. 40).

The main base was established at Cape Denison
(Commonwealth Bay) and a western base party,
including two geologists, some 1 500 miles (2 400
km) further along the coast to the west on the
edge of the Shackleton Ice-Shelf. The vicinity of
Cape Denison was comparatively ice-free 7 and the
geologists were able to study glacial erosion and
deposition at first hand. The rocks outcropping
near the main base were predominantly
metamorphic — folded and contorted gneiss and
schist of presumed Precambrian age. Evidence of
ore-bearing minerals was also found, among them
iron, copper and molybdenum which Mawson
believed gave promise of larger bodies in the
vicinity and indicated the probability of mineral
wealth beneath the continental ice-cap. Around
the edge of the ice margin, the terminal moraines
were 'a veritable museum'. Rocks, showing every
variety in colour and form, were assembled,
transported from far and wide over the great
expanse of the continent ... The story of the buried
land to the south is in large measure revealed in
the samples brought by the ice and so
conveniently dumped' (Mawson 1915, 1996 pp.
74-76). After a particularly severe winter with
hurricane force winds, the summer journeys
commenced, and much valuable scientific work
carried out. A western party made a particularly

DOUGLAS MAWSON GEOLOGIST

119

interesting find, the first meteorite from
Antarctica, duly named the 'Adelie Land
Meteorite 7 , and the first of many to be found in
later years.

Mawson himself, with Ninniss and Mertz and
the dogs, set off on the longest and most
ambitious of all the journeys, striking east on a
southerly course in an attempt to cover as much
territory as possible in the time available. The
story of that harrowing journey which cost the
lives of both of Mawson's companions, and of his
miraculous return against overwhelming odds just
in time to see the 'Aurora' disappear over the
horizon, is an epic story of endurance, fortitude
and sheer will power which has been well told by
Mawson himself in L The Home of the Blizzard'
(Mawson 1915) and by Bickel (1977). Mawson
and the small remnant of the expedition left
behind to search for him had to face another
Antarctic winter. The 'Aurora' returned in
December 1913 and on the 26 February 1914,
Mawson arrived back in Adelaide.

The War Years and After

The expedition was home, but for Mawson
there was to be no respite and it was to be a
further six eventful years before he was able to
resume an ordered academic life again. The First
World War began as he was returning from an
overseas business trip to Europe concerned with
the wind-down of the expedition, and by 1916,
after travelling to the United States, he was back
in Britain where he spent the remainder of the war
on highly responsible government work with the
Ministry for Munitions. He did not return to
Adelaide until May 1919. Back at the University
he found that little had changed in his absence
except that Walter Howchin had been awarded the
title of Honorary Professor in Geology and
Palaeontology in 1918. Mawson (1944a) wrote
later: '1 was not a little confounded. 1 would have
packed up and left immediately but for the
supreme consideration of faithfully recording and
adequately publishing the immense mass of data
received during my Antarctic Expedition'. Early
in 1920 he talked with Howchin about the future:
'stating that I was not prepared to continue at
Adelaide for another year under the then existing
inadequate arrangements for geology'. There was
no prospect of the much-needed new building for
geology and Mawson tied himself to a
compromise twenty year agreement on additional
accommodation which proved increasingly

inadequate as years went by, and which he was
later to regret. Meanwhile, Howchin's continuing
presence at the University was perceived by
Mawson as a hindrance to the satisfactory
development of the Geology Department and it
must have been a great relief to him when
Howchin, then 75, decided to retire at the end of
the year.

In 1 92 1 Douglas Mawson was appointed to the
newly created Chair of Geology and Mineralogy,
and Dr Cecil Madigan, his old comrade from the
Antarctic, joined him as Lecturer the following
year. Mawson and Madigan were to be the only
two staff members in the Geology department for
much of the inter-war period. The establishment
of the new department and his teaching and
administrative duties (not to mention continuing
Antarctic matters) took precedence in the early
1920s but field investigations were resumed in
both the Olary region and the north Flinders
Ranges. While he built on his work in the
Boolcomatta area, locating several key sections
spanning the junction between the basement and
overlying sedimentary sequence, he was never to
resume a sustained fie Id work program in the
region, despite the urgings of David.

David's retirement at the end of 1923 and the
politics involved in choosing his successor proved
unsettling for Mawson who at that time was still
not fully committed to staying in Adelaide and
applied for the job, and although the best qualified
of the candidates, he was ultimately unsuccessful
because of David's influence on the course of
events and his (perhaps over-zealous) loyalty and
support to Leo Cotton, his deputy and locum for
many years, who was appointed to the Chair. The
outcome certainly posed strains on the David-
Mawson relationship, and further tensions and
distractions were to follow when David's
increasing obsession with the search for supposed
Precambrian fossils focussed on the old rocks in
South Australia (see also pl24, End of an Era).
On an earlier visit to Adelaide in 1921, David had
found what he believed to be arthropod remains in
the old rocks at Reynella (long considered by
Howchin as Early Cambrian) tentatively placing
them in the late Precambrian (Proterozoic) and
proposing a new division the Adelaide Series.

Meanwhile, Mawson turned his attention to the
north Flinders once again. He had published a
brief comparison of the Mount Painter and Olary
provinces (Mawson 1912b), but the results of his
investigations around Mount Painter in 1910 were
treated more fully in a long-delayed paper
(Mawson 1923) in which he recognised a core of

120

D. W. CORBETT

:•'

FIGURE 7. The Radium Hill Mine, soon after its discovery. Mawson, 1944b, pi. 19. Courtesy Royal Society of South
Australia..

* #:i

FIGURE 8. Mount Gee, viewed from the west. Mawson, 1944b, pi. 19. Courtesy Royal Society of South Australia.

DOUGLAS MAWSON GEOLOGIST

121

Precambrian igneous and metamorphic rocks
overlain by a sedimentary series similar in many
respects to those overlying the basement in the
Adelaide region. He followed David's lead in
applying the term Adelaidean (Adelaide Series) to
these sediments and suggesting a Proterozoic (late
Precambrian) rather than Cambrian age. Igneous
activity included two acid phases, the second with
pegmatites and aplites followed by a volcanic
phase characterised by vesicular basalts. The
mineralisation of the Mount Painter-Mount Gee
area, notably the quartz reef at Mount Gee and the
uranium minerals (torbernite and autunite) at
Mount Painter, was described in detail (Fig. 8).

Following Mawson's enthusiastic report on the
uranium potential of the area after his initial visit
in 1910, several companies had been floated and
prospecting carried out on Radium Ridge and
Mount Gee. Meanwhile the Mines Department
had been more cautious but in favour of further
prospecting. Sporadic attempts at developing the
prospects (notably the No. 6 workings) had

languished during the war, but in 1924 there was
renewed interest in radioactive ores world-wide
and a return to the Mount Painter field seemed
justified. Consequently, Mawson returned north in
November 1924 and spent much of his time in the
Mount Painter area on detailed study of the
mineralised rocks but also in broader scale
assessment of the regional context of the
mineralisation.

The regional geology of the Flinders Ranges
was little known at this time. Howchin had made
a pioneer traverse from west to east at the level of
the Parachilna Gorge in 1907 but the results were
not published until 1922. While on his return trip
from the north, Mawson had the good fortune to
make a particularly significant discovery just west
of the Italowie Gorge. It was a case of the
prepared mind recognising the significant
evidence and provides a good example of
Mawson's versatility as a geologist, for fresh from
the mineral riches of the Mount Painter country he
was now to show that he had a good eye for a

FIGURE 9. Entenng Yudanamutana. Mawson, 1923, pi. 33. Courtesy Royal Society of South Australia.

122

D. W CORBETT

fossil also. What he found were 'curious
markings' on roadside outcrops of limestone which
'on inspection ... was found to be due to massed
fossil heads of a Cryptozoon-Wkt alga ... developed
in a massive formation, the algal remains
constituting the bulk of at least some of the beds'
(Mawson 1925). Similar structures had been
reported from the MacDonnell Ranges by
Chewings and described by Howchin (1914) but
there was as yet no general agreement on their
stratigraphic position. Mawson inclined towards an
Early Cambrian age, but David, on hearing of the
find thought they might be older (Proterozoic). Very
little was then known of Precambrian life forms;
most of the obscure remains described being
referred to as Crpytozoon (hidden life)— but the
recently announced discovery of algal fossils in
Precambrian rocks in Montana by Walcott
suggested that the Flinders rocks might be of
similar age. David certainly believed so and urged
Mawson to investigate further. Mawson's

'Cryptozoons' are now recognised as algal
stromatolites, These columnar and dome-shaped
structures produced by the trapping of sedimentary
particles by mats of blue-green algae are the earliest
abundant fossils and are well developed in certain
of the Precambrian rock formations in the Flinders
Ranges. Mawson, however, was not to be
pressured into narrow specialisation and continued
his wide-ranging investigations of Flinders Ranges
geology (Figs 9, 10), publishing further papers in
the mid 1920s. One was on the volcanic rocks of
the Wool tana area (Mawson 1926a), first
encountered in Arkaroola Creek in 1910 and
subsequently examined near Wooltana on a brief
visit in 1924. There he found a thick sequence of
volcanics — vesicular lavas, tuffs and agglomerates
overlain by boulder beds and finer sediments which
he described as: 'most probably of fluvio-glacial
and glacial origin'. Large granite boulders higher
in the section were even more convincing, leaving:
\.. no doubt in my mind ... as to the glacial nature

FIGURE 10. The quartzitc of the Yudanamutana Gorge. Mawson, 1923, pi. 33. Courtesy Royal Society of South
Australia.

DOUGLAS MAWSON GEOLOGIST

123

of these beds,..' Sediments above the glacials,
unsuccessfully searched for fossils, were believed
of likely Late Precambrian age.

Another spin-off from the 1924 North Flinders
trip was a visit to the Paralana Hot Springs which
interested Mawson because of the radioactive
nature of the waters (Mawson 1927a). For a short
period about this time an enterprising doctor had
sought to capitalise on the supposed therapeutic
qualities of the spring by setting up a spa on the
site, although with no success. The springs are
situated close to the faulted junction between the
old rocks of the ranges and the younger sedimentary
basin to the east. While most geologists supported
an origin from deep-seated basin waters, Mawson
speculated on a possible contribution from the
highly uranium-rich rocks lying to the west. He
collected water and gas for examination, hoping the
latter would prove rich in helium. Unfortunately the
samples did not survive the journey back to
Adelaide. The highly active gas which at first
puzzled the analysts, has since proved to be rich in
radon; the water is much less radioactive.

Mawson had made a brief visit to the Willouran
Ranges on the north western flank of the Flinders
in 1920. From here he described and named the
Willouran Series—a thick sequence of
sandstones, slates and calcareous beds underlying
a dominant quartzite formation and forming a
synclinal structure (Mawson 1927b). Sediments of
probable fluvio-glacial origin were also found in
the eastern part of the area.

Mawson's activities in the Flinders Ranges
during the 1920s were essentially of a
reconnaissance nature and based on relatively
little time, overall, spent in the field. The
information included in the papers cited above
demonstrate how much he could gain from a brief
investigation, displaying his strength as a field
worker and skill in rapidly appraising the
geological situation in unknown country, and
revealing his pre-eminence as an explorer-
geologist. Work carried out during this period,
while not yet part of a coordinated research
programme, provided the basis for his later
systematic approach to regional studies in the
1930s and 1940s.

Away from his empirical work in the field, he
demonstrated a comparable ability to synthesise
existing knowledge in a lengthy treatment of the
igneous rocks of South Australia, delivered as a
Presidential Address at the AAAS meeting at
Perth in August 1926 (Mawson 1926b). This was
an important review, given the widespread
distribution and importance of igneous rocks,

notably in the older terrains of the State, but also
in those of younger age. By discussing them
region-by-region in their structural and
stratigraphic context, he produced what was
essentially a geology of the State. One issue of
particular interest, not to be resolved for many
years, was the age of the highly metamorphosed
sediments on the eastern flank of the Mount Lofty
Ranges and their associated intrusives. With
reference to Howchin's view that these rocks were
the equivalents of the Adelaide Series, Mawson
urged caution, pointing out: 'the geological
mapping of the Ranges is yet only in the early
stages (and) final conclusions are unsafe until
more detailed petrology and complete geological
mapping have been accomplished'. As for the
Victor Harbor granites Mawson comments
perceptively: 'there is yet no convincing evidence
against the assumption of an early Palaeozoic age
for this granite'.

Central Australia 1927

In November 1927, Mawson and Madigan
made a short visit to Alice Springs and the
Western McDonnell Ranges to check a mineral
occurrence which had excited local interest. A
white crystalline substance, long known to the
aborigines and which burned with a fierce flame,
was thought possibly to have some connection
with petroleum. Specimens had been brought to
Adelaide (initially to Madigan), and identified by
Mawson as potassium nitrate (saltpetre or nitre) a
rare mineral with some economic use in the
chemical and explosives industries. Realising the
opportunity for geological exploration in an area
still little known geologically, Mawson quickly
organised an expedition to investigate. They
travelled north by train to Alice Springs, and their
car journey out through the western ranges from
Alice Springs blazed a trail through country never
traversed by vehicle before. The deposit proved to
be of organic origin, the long-time accumulation
of animal droppings, of scientific but not
economic interest.

Much of the route west ran along the strike of
the well-exposed sedimentary beds which gave a
general indication of the southward dipping
succession between the Precambrian crystalline
basement and known Ordovician rocks of the
Horn Valley. Traverses were run and rough
sections made at several localities along the way,
including Ellery Creek, and over 10 000 feet of
strata were measured. In the lower beds

124

D W. CORBETT

'cryptozoal limestones' and organic structures
similar to those from the Flinders Ranges were
prominent in several areas, while worm tracks and
traces of mollusc shells together with algal
limestones distinct from those lower in the
sequence were found in the younger beds. On the
basis of his limited field evidence, the sedimentary
sequence was divided into two divisions — the
Pataknurra and Pataoorta Series, once again
showing his flair for the rapid assessment of the
general geological situation in a new area. He
made tentative and essentially correct correlations
between the central Australian rocks and the older
sequences in South Australia. This first and, as it
proved, only venture into central Australian
geology was a major contribution to knowledge of
the region, building on and revising the earlier
work of Tate and Watt (on the Horn Expedition),
H. Y. L. Brown, C. Chewings and L. K. Ward.

Mawson visited London the following year
where he exhibited photographs and specimens at
the Geological Society. The results of the
expedition were published by the Society
(Mawson & Madigan 1930). While circumstances
determined that Mawson never returned to the
Centre, it seems likely that he always envisaged
following up the promising results of his first trip.
Madigan, meanwhile, had been captivated by the
Centre and immediately began his own
independent and wide-ranging researches into the
geology of the region which were to produce some
of his best work. This perceived 'take-over' by
Madigan was resented by Mawson and led to bad
feeling between the two men, but Mawson was
soon to be occupied with other matters.

BANZARE 1929-31

Mawson was very much involved with
Antarctic matters from 1929 through into the early
thirties. BANZARE (The British, Australian and
New Zealand Antarctic Research Expedition)
differed from Mawson's previous expeditions in
being ship-based. The program was carried out on
two summer cruises (1929-30, 1930-31) and
while no land bases were established, limited
sorties were made ashore, notably on some of the
sub-Antarctic islands. With the emphasis on
marine science, much of the data was collected by
systematically sounding the depth of the offshore
waters, sampling the water column and dredging
the bottom for biological specimens. Because of
the nature of the expedition, geological work was
restricted, although sediments collected from the

sea-floor adjacent to the land and the variety of
ice-transported boulders dropped there provided
abundant evidence of sedimentation and
sedimentary processes in a modern glacial
environment. The BANZARE expedition further
inspired Mawson to continue his study of the
ancient glacial deposits in the Flinders Ranges,
and he was to do this with renewed vigour in the
years ahead.

The extended geographical and geological
knowledge of Antarctica resulting from the
BANZARE voyages enabled Mawson to be the
first to postulate the existence of an extensive
landmass lying beneath the southern ice-cap and
bounded by a wide continental shelf. Much of the
evidence for the geological make-up of this buried
continent came from the ocean-floor studies where
dredged boulders proved to be rocks of continental
type. Mawson envisaged that the mineral
resources of Antarctica, the existence of which his
expeditions had suggested but were yet to be
proven, would be developed at some time in the
future, and that underground mining operations
could be carried out successfully despite the
problems of climate and terrain.

End of an Era

Professor Edgeworth David died in 1934. In his
later years he had become increasingly obsessed
with his recognition of 'fossils' in the Precambrian
rocks of South Australia (David 1928). Most of
his finds had been made on visits to Adelaide at
times when Mawson was away overseas or
otherwise engaged on Antarctic business and had
no chance to control these incursions. For
Mawson it was particularly galling for David to
hint that his old student was sitting on a rich
treasure trove of Precambrian fossils while he,
David, made the pickings, even enlisting
Mawson's students and staff to help him collect.
Wisely, Mawson held his counsel and
endeavoured not to be drawn into the matter. It
soon became clear, to almost all but David, that
the supposed 'fossils' were in fact inorganic in
origin (a verdict upheld by later workers), but he
was to persist in his belief through to the end of
his life, considering his discoveries in South
Australia to be the highpoint of his career.

Deciphering the Flinders

In the mid- 1930s Mawson renewed his long-

DOUGLAS MAWSON GEOLOGIST

125

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127

standing but formerly sporadic association with
the Flinders Ranges. Now for the first time with
fewer distractions, he was able to develop and
pursue a sustained and systematic ficldwork
program the results of which constitute his most
notable contribution to South Australian geology.
Only a brief resume of this work is given here, to
be treated more fully in a later paper.

The Flinders Ranges were sparsely populated in
the 1930s, and moving around in the outback
more arduous than today. Roads and tracks were
in general poorly maintained and motor transport,
then in its infancy, was still an adventurous
undertaking. There were no photo-based maps or
accurate large-scale topographic maps available in
those days, the only maps available being pastoral
plans which were quite inadequate as a base for
geological mapping. Mawson's approach was to
select key sections which were measured and
described in detail. In a few of his papers, small-
scale regional maps were included. He first used
aerial photography as an adjunct to his ficldwork
in the late 1940s. Field-camps in the pre- and
early war years combined the instruction of senior
(third year) students in field techniques, including
section measuring, with the gathering of basic
geological information in country previously
geologically unknown.

By 1939, having spent time in the Wooltana,
Mount Carnarvon, Parachilna Gorge and Italowie
areas, Mawson's work in the Flinders had reached
a stage where review was warranted, and two
publications in that year, on the Late Precambrian,
and Cambrian successions respectively (Mawson
1939a, 1939b), detail rock sequences in the
Brachina Creek-Oraparinna and Wirrealpa Basin
areas of the central ranges (Fig. 11). The Pound
Quartzite, the dominant ridge-former of the
Flinders which occurs some distance
stratigraphically below the oldest fossiliferous
Cambrian rocks, was provisionally placed at the
base of the Cambrian on lithological grounds and
interpreted as marking the start of a new cycle of
sedimentation. Defining the Precambrian-
Cambrian boundary was to become of increasing
interest to geologists in the years ahead, a task
made no easier by the lack of fossils and by the
absence of major breaks in sedimentation in the
thick Upper Precambrian to Cambrian
sedimentary sequence in South Australia. In his
Wirrealpa Basin paper, two important
fossiliferous units are described: the
Archaeocyatha Limestone and a younger
limestone (the Wirrealpa Limestone) first
described by Howchin but here fully documented

for the first time with its fossil fauna of algal
remains {Girvanella), brachiopods (Oboiella),
pteropods and trilobite fragments (Redlichia).

Other papers were to follow dealing with
sequences in the Aroona, Copley, Brachina and
Wilpena areas and culminating in an important
review, 'The structural character of the Flinders
Ranges' (Mawson 1942), which brought together
the results of his fieldwork program to that time.
A small scale map (Fig 13) showed the location
of the key sections measured and the distribution
of the rocks spanning the four main periods of
deposition: the Middle Proterozoic and Late
Proterozoic, the Cambrian and the Tertiary-
Recent. Key marker beds, including the prominent
Pound and ABC Quartzites, tillites and
hieroglyphic limestones were also delineated. A
cross-section (Fig. 12) indicated the essentially
simple anticlinal structure of the central ranges. A
wider-ranging paper (Mawson 1947) extended
and amplified his earlier work, adding new
information from the western flank of the ranges
both in the north (near Copley) and in Mundallio
Creek to the south. In both areas Mawson
recognised thick quartzite formations at the base
of the sequence, pointing out the error made by
Segnit (1939), working for the Mines Department,
who had equated these ABC Quartzite sequences
with the Pound Quartzite. Mawson, however, was
himself misled in the area west of Quorn where he
mistook the prominent folded ABC Quartzite as
Pound Quartzite (here thinly developed). Such are
the dangers of making correlations on lithological
grounds in rocks without fossils to act as age
indicators.

Mawson also discussed the Flinders succession
in its broader tectonic framework which he
considered a major trough of deposition of Late
Precambrian to Cambrian age and referred to as
the 4 great synclinal basin' (later to be named the
Adelaide Geosyncline). His work had
demonstrated the great thickness of strata
deposited in the trough which extended south from
Central Australia to the Adelaide region and lay
between two ancient Precambrian massifs:
Yilgamia to the west and Willyamia to the cast.
The flat-lying sedimentary cover-rocks to the west
of Port Augusta, believed on lithological and
structural grounds to be correlates of the basal
quartzites in the western ranges are now known to
be ABC Range Quartzite equivalents. Wider
correlations of the Flinders Ranges succession
with the rocks of the western Mount Lofty Ranges
(comparisons with the still largely unresolved
sequence on the eastern flank of the Mount Lofty

128

D. W, CORBETT

THE CENTRAL AREA OF THE FLINDERS RANGES

\ \ \ \ \ \ N

FIGURE 13. Mawson's geological map of the central Flinders Ranges. Mawson, 3942. Courtesy Royal Society of
South Australia.

DOUGLAS MAWSON GEOLOGIST

129

Ranges still remain highly speculative), with
Central and Western Australia and also with
South Africa (Transvaal System) were also
suggested and discussed (Mawson 1947).

In the early 1940s Mawson returned to the
North Flinders investigating the metamorphic
effects of plug-like granite intrusions in the
Precambrian country rocks (Mawson & Dallwitz
1945a,b). This study of both the sedimentary and
igneous aspects of the geology of the region linked
his early work on the older mineralised terrains in
the Mount Painter area with his later concentration
on unravelling the stratigraphy of the overlying
sedimentary sequence,

Ancient Glaciation

Mawson first encountered evidence of ancient
glaciation on his trip to the north Flinders in 1910.
Subsequently he had demonstrated the widespread
distribution of glacial rocks in the ranges during

his extensive field investigations (Fig. 14). His
Antarctic experience of a glaciation-in-being, with
the diversity of its products and processes, from
the coarse boulder beds of terminal moraines to
the finely layered sediments laid down in tranquil
glacial lakes, had equipped him, as he had
intended it should, to recognise the varied and not
always easily interpreted evidences for former ice-
ages.

By the late 1940s, Mawson had established the
stratigraphic position of the Sturt Tillite in many
areas of the Flinders and also identified a younger
glacial episode in the Elatina area on the western
flank of the central ranges (Mawson 1949a). The
distinctive sediments— coarse sandy beds overlain
by a thick unstratified boulder bed — first
encountered in 1938, had initially been thought to
be of pyroclastic origin (Mawson 1938) because of
the preponderance of igneous pebbles in the rock.

Mawson then produced evidence for a third
glacial event, this time predating the Sturt Tillite
(Mawson 1949b), in the deeply eroded high

FIGURE 14. A large striated erratic weathered out of Proterozoic tillite, Yankarunna Station. Photo R. H. Jones.
Mawson, 1945, pi. 14. Courtesy Royal Society of South Australia.

130

D. W. CORBETT

country of the Bibliando Dome east of Hawker.
Here a thick sequence of glacials and associated
beds revealed two extended periods of glaciation
separated by a well-defined inter-glacial interval.
The younger glacial event was equated with the
Stmt Tillite while the older (lower) tillites were
referred to as the Bibliando stage.

The clear demonstration of a third glacial phase
in the history of the geosynclinal basin, with a
broad range of deposits from boulder tillites
through coarse fluvio-glacial sediments to fine
laminated shales, raised such questions as the
interpretation of past geographies, climates and
environments, and the cyclic nature and causes of
glaciation. Mawson, with his deep and extensive
knowledge of ice-action both ancient and modern,
was, more than any other geologist of his
generation, aware of the diversity, complexity and
the problems associated with interpreting glacial
evidence. He discussed some of these in his paper
on Bibliando, which marks the culmination of his
career-long investigation of glacial phenomena.
He took the opportunity to view this latest work in
a broad historical perspective. The early
discoveries of Selwyn, Tate and Howchin had
placed South Australia at the forefront of research
in the study and understanding of past glaciations.
His own work had extended and clarified that
knowledge, setting it within the wider context of
the development of a major Upper Precambrian
geosyncline, the importance of which had now
become apparent.

While the rocks in the Flinders are
magnificently displayed and accessible for study,
the lack of fossils in the ancient rocks have always
made the task of the stratigrapher difficult.
Mawson, the first geologist to systematically
approach the task of establishing an ordered
succession was acutely aware of this problem. As
his work proceeded and became recognised
beyond South Australia, he perceived the need for
a standardisation of geological nomenclature,
particularly of stratigraphic names, and he was a
member of the first committee set up by ANZ AAS
in 1947 which led to a Code of Stratigraphic
Nomenclature being formalised in 1950.

Mawson favoured dropping the long-
established but restricted Adelaide Series of
David and Howchin and replacing it with the
Adelaide System which he sub-divided into the
Torrensian, Sturtian and Marinoan Series
(Mawson & Sprigg 1950). The defining of the
Adelaide System marked the end of Mawson's
work on the old rocks of South Australia. The
discovery of jellyfish fossils by Reg Sprigg in the

Pound Quartzitc at Ediacara in 1946 (Sprigg,
1947, 1949), and the later finds of a diversified
animal fauna by Flounders and Mincham, led to
a reappraisal of the stratigraphy of the fossil-
bearing beds (Glaessner & Daily 1959).
Relegation of the Pound Quartzite from the base
of the Cambrian (as previously suggested by
Mawson) to the uppermost formation in the
Precambrian meant that the Ediacara fossils, now
confirmed as the oldest animals known, became
world famous.

The South-East

Contrasting strongly with the rugged grandeur
of the Flinders Ranges, the geologically youthful
and topographically unchallenging south-eastern
region of South Australia attracted Mawson from
the mid- 1930s. He became interested in the
problematic 'coorongite', a rubbery bituminous
substance first discovered in 1852 at Alf Flat east
of Salt Creek at the southern end of the Coorong
Lagoon (Fig. 15). A belief that the flammable
material might be a petroleum product led to
speculation that oil existed at depth; and several
(unsuccessful) drilling attempts were made in the
1880s and again in the 1920s. By then almost all
investigators had become convinced of the
vegetable origin of the substance (and it has since
been shown to be the product of algal blooms).
Mawson, however, was unconvinced; and he
inadvertently attracted overseas interest for a brief
period when he took a visiting petroleum expert
from Britain (Washington Gray) into the field to
view the material and Gray went home to
publicise the find, much to Mawson's chagrin.
Some biochemical work was carried out overseas
on the 'algal gel', but the matter was not pursued
any further.

More substantial and fruitful work in the region
led to a series of publications on the small and
isolated outcrops of granitic rock which protrude
through the thin Quaternary cover of the Upper
Southeast. Fieldwork involved exploration on a
minor scale, for many of the outcrops lay in
difficult country accessible only on foot or
horseback and where the outcrops were not
extensive. The studies were carried out in
association with students or former students and
covered a quadrilateral bounded by Meningie and
Coonalpyn on the north and Kingston and
Bordertown to the south. The field investigations
and subsequent analytical work led Mawson to
recognise a variety of intrusive and extrusive rock

DOUGLAS MAWSON GEOLOGIST

131

FIGURE 15. Mawson, with auger, drilling for coorongite, the Coorong, mid-1 930s. Washington Gray on right.
Source unknown.

types in the outcropping basement: granites;
adamellites and granodiorites; and quartz
keratophyres (volcanic). Similarities between the
intrusives of the South-East and those occurring
along the eastern flank of the Mount Lofty Ranges
and on Kangaroo Island supported the concept of
a large granitic batholith existing beneath the
sedimentary cover, and, on the available evidence.
Middle to Late Cambrian in age.

Finale

Mawson was sixty-three at the end of the
Second World War and approaching the normal
age for retirement. He continued on, however, to
the age of seventy and spent his last university-
years rounding off his own researches and
ensuring that his department was left in a healthy
and secure position to meet the challenges ahead.

132

D. W. CORBETT

THE MAWSON LABORATORIES

SCHOOL OF GEOLOGY

UNIVERSITY OF ADELAIDE

WOODS BAGOT I AT BCUR.KE -SWIT H £^ I R.WI M

A R. C H 1 T C. C T & .

FIGURE 16. The proposed Mawson Laboratories. Courtesy Woods Bagot Pty Ltd, Adelaide.

He had been acutely aware of the greatly increased
role that geology would be required to play in the
post-war world. In particular, well trained
geologists could only be supplied by the
universities which in turn would need to provide a
sufficient level of support in terms of
accommodation, equipment and staffing. In all
these areas Mawson had been struggling against
inadequacies for over twenty years. During this
time, geology had made the best of sub-standard
accommodation and increasingly inadequate
provision for research, whereas the rest of the
University had developed and prospered. It was
the urgent need for a better deal for geology that
prompted his memorandum to the University
authorities in February (1944a). In his
memorandum Mawson wrote: 'The department
suffers from insufficiency of room, from
inadequacy of the laboratories, and from the entire
absence of any planned layout suitable for a
science department. All other science departments
of the University now have buildings specially
planned for their needs though some are already
proving too limited in accommodation for the
flood of students now presenting themselves. Ours
is an anachronism'.

He went on to make a very strong case for the
role of geology in education, its application to

national development and significantly, South
Australia's importance as a source of future
mineral resources. He envisaged Adelaide
University as a future centre for teaching and
research in economic and mining geology and
itemised the basic needs of a modern School of
Geological Studies, estimating the cost of the new
development at around £25 000.

His argument was decisive. The University
responded appropriately, if belatedly, and the
Mawson Laboratories were constructed, largely to
his specifications, between 1949 and 1952 (Fig.
16). He just had time to move into the new
building before retirement. By then his staff had
increased to five and included E. A. Rudd who
was appointed to the new Chair of Economic
Geology in 1949, and Martin Glaessner, a
palaeontologist with an international reputation
who was to become the authority on the
Precambrian fossils from Ediacara. Arthur
Alderman, former student and junior lecturer in
the department, returned after a period away from
the University to take over from Mawson as
Professor of Geology and Mineralogy in 1953.

In the years following his retirement Mawson
remained active and influential in Antarctic
matters and continued his long association with
the South Australian Museum as Honorary

DOUGLAS MAWSON GEOLOGIST

133

Curator of Minerals and Member of the Museum
Board. He was appointed Chairman of the Board
in 1952.

Douglas Mawson's broad interests in all
aspects of geological science and his unfailing
encouragement and support of young people with
an enquiring mind and an interest in science is
well illustrated by the background to his last
published paper (Mawson 1958). It deals with
australites (small glassy objects of meteoritic
origin) collected by a young schoolboy, Mervyn
Pens, from the Murray Plains west of Morgan in
the mid-1 930s. Mawson had visited the site and
established a correspondence with young Pens,
which led to further finds and ultimately to the
public aquisition of a large and significant
collection of australites now housed in the South
Australian Museum. The paper was read before
the Royal Society of South Australia on 10
October 1957. Sir Douglas Mawson died on 14
October 1958.

Mawson's Legacy

Mawson's inborn qualities as an explorer, his
methodical approach to the job in hand, his
tenacity and resolution and an ability to inspire
and motivate others were revealed as much in his
career as an academic geologist as in his exploits
as an Antarctic explorer. He was essentially a man
of action, a trail-blazer, single-minded in the
pursuit of his goals, at times impatient for results,
and who could, at times, be protective to the point
of touchiness, of his own areas of interest. His
inherited characteristics, hard-headed Yorkshire
practicality, stubborness and reticence, with a
streak of romanticism (from his father) were
blended with a pioneer Australian belief in
opportunity and self-reliance to produce a complex
character of contrasting and at times conflicting
elements. He was a man not easily assessed or
categorised. Geology is arguably the most
adventurous of all the field sciences (the first
scientist to visit the moon was a geologist), and
Mawson in his life and career, both in the
Antarctic and in his geological investigations in
outback South Australia, exemplified both the
scientific-explorer and explorer-scientist.

Mawson's scientific career spanned, almost
exactly, the first half of the twentieth century.
When he entered the University of Sydney in 1899
the scientific world-view was of a cooling,
shrinking earth not more than one hundred million
years old, the limits having been set by the

physicists. Geologists, whose investigations had
by then spread to all parts of the globe, intuitively
sensed a more ancient earth but lacked the means
to prove it. Mountain belts were seen as a crustal
adjustment to a cooling interior and most
geologists accepted the relative permanence of the
continents and oceans.

In retrospect, the years 1900-1950 can be
viewed as a relatively quiet episode in the
development of geological science, enlivened by
two developments early in the century : the
application of the discovery of radioactivity to the
dating of rocks, and Wegener's hypothesis of
continental drift. Each in its own time, the first
rapidly, the second much more slowly, became a
vital element in the future Plate Tectonic
revolution. The recognition that the decay of
radioactive minerals produced heat freed
geologists from the shackles of a cooling earth
and a restricted time-scale by providing a source
of self-sustaining energy and a means of dating
the crustal rocks. By 1914, the estimated age of
the earth had been pushed back to over one
thousand million years. Mawson's work on
radioactive minerals in Australia as a young
student and his later discoveries in South
Australia gave him a pioneer role in this new
development and he was always very proud of his
work in this field (see Mawson 1944b). But in his
day Australia was effectively too remote from the
rest of the world for its small and largely
inaccessible deposits to be economically viable,
and its universities (certainly Adelaide) too poor
to engage in expensive age-dating equipment. In
his later years Mawson was keenly interested in
these developments overseas and their future
potential for dating the old rocks in Australia.

Communication between scientists was severely
curtailed during the two world wars which
dominated the first half of the century, although
technical developments during the second war
were to help revolutionise geological and
geophysical exploration both on land and in the
ocean basins. Mawson's own career was savagely
disrupted by the First World War and his more
active years were over before the benefits of post-
Second World War technical advances became
widely felt. The inter-war years and those
immediately after the second war were his most
productive period for research, more particularly
from the mid- 1930s when his active involvement
in Antarctic exploration had ended. Although the
general outline of South Australian geology was
well known by this time, no systematic work had
been attempted and for much of the State

134

D. W. CORBETT

information remained highly generalised and
lacking in detail. As previously indicated,
Mawson's early researches were essentially
exploratory reconnaissances, becoming more
systematic with his work in the Flinders from the
1930s. His detailed descriptions of rock units and
measured cross-sections of key sequences
provided the basis for the regional mapping
program to be instigated by the Geological Survey
in the 1960s.

Mawson's research was very much an integral
part of the growth and development of his
department. His field studies in the Flinders were
carried out with small groups of advanced
students as part of their course work, while in the
Southeast he published in collaboration with post-
graduate and former students as well as directing
them in projects of their own. His teaching is
remembered by past students as being of variable
quality; he was at his best, according to
Alderman, when extemporising on subjects that
particularly interested him. There is no doubt that
he could inspire his students and a considerable
number went on to make distinguished careers in
geology and mineralogy, among them E. A. Rudd,
L. W. Parkin, R. C. Sprigg, W. B. Dallwitz, D. E.
Thomas, E. R. Segnit, B. Skinner, R. L. Crocker,
M. Reynolds, B. Daily, R. Sweatman, A.
Kleeman, B. Forbes and G. Chinner.

Mawson, the man of action, always seemed to
have been fighting a lack of time to complete his
projects; always planning another field trip and,
one senses, chafing at the administrative burden
placed on him as a highly responsible and
consistently conscientious head of department.
Never an armchair geologist, Mawson did not
have the time or the inclination to become
involved with the theoretical issues of the day, the
most contentious of which was the idea of
continental drift. Sir Mark Oliphant, a science
student at the University in the 1920s, has recalled
attending a debate on the subject between
Mawson and Wood-Jones the anatomist, during
which Mawson scathingly dismissed the concept
of drift, citing the rigidity of the ocean floors and
the lack of any known mechanism that could
power the process.

Sir Douglas Mawson made an outstanding
contribution to South Australian geology. His list
of 102 publications, the majority published in the
Transactions of the Royal Society of South
Australia, provide the material evidence of his
work. He also left a strong department which he
had established and which was developing rapidly
in teaching and research. While it is idle to
speculate, one cannot help but wonder what he
might have accomplished as a geologist if so
much of his time and energy had not been devoted
to the Antarctic. He was not able to settle down to
an ordered academic life and an on-going research
program until he reached his fifties. From then on
his geological output was sustained until the end
of his career. It was a remarkable achievement.

Acknowledgments

The research on which this paper is based was begun
when the author was a staff member in the Mawson
Graduate Centre for Environmental Studies at the
University of Adelaide and completed as an Honorary
Research Associate at the South Australian Museum. It
relies heavily on a study of the Mawson Papers housed
in the Special Collections archive of the Barr Smith
Library, The University of Adelaide, and Mawson's
published papers. Research was also carried out on the
Edgeworth David papers held by the Fisher Library at
the University of Sydney and in the library of the Scott
Polar Research Institute, Cambridge.

For valuable assistance during the course of this work,
thanks are due to: Susan Woodbum and the staff of
Special Collections, The University of Adelaide; Mr Ken
Smith, Archivist, The Fisher Library and Professor
David Branagan, The University of Sydney; Mr Robert
Headland, Curator and Archivist, The Scott Polar
Research Institute, Cambridge.

For on-going encouragement and support for this
research and for much helpful discussion on matters
relating to Mawson, grateful thanks are extended to Mr
Richard Ferguson, Polar historian; Emeritus Professor E.
A. Rudd, The University of Adelaide; Dr Victor Gostm
of the Geology Department, The University of Adelaide
and Dr Allan Pring, Curator of Minerals, The South
Australian Museum. Particular thanks are due to Dr
Robin Oliver of the Geology Department, University of
Adelaide, for his critical review of the original draft of
this paper, and his many useful suggestions.

Selected Bibliography of Douglas Mawson

(See Alderman (1959) for a Memoir and Bibliography of
Mawson. Innes and Duff (1990) give a complete list
of his scientific and other papers together with
biographical works on Mawson).

903 (with T. G. TAYLOR). The geology of Mittagong.
Journal and Proceedings of the Royal Society of New
South Wales 37: 306-350."

DOUGLAS MAWSON GEOLOGIST

135

1904 (with T. H. LABY). Preliminary observations on
radioactivity and the occurrence of radium in
Australian minerals. Journal and Proceedings of the
Royal Society of New South Wales 38: 382-389.

1905 The geology of the New Hebrides. Proceedings of
the Linnean Society of New South Wales 3: 400-485.

1906 On certain new mineral species associated with
carnotite in the radioactive orebody near Olary.
Transactions of the Royal Society' of South Australia
30: 188-193.

1912a Geological investigations in the Broken Hill area.
Memoir of the Royal Society of South Australia 2(4):
211-319.

1 9 1 2b PreCambrian areas in the north-eastern portion of
South Australia and the Barrier, New South Wales.
Reports of the Australasian Association for the
Advancement of Science 13: 188-191.

1915 The Home of the Blizzard. Heinemann, London.
[Extracts quoted are taken from the abridged popular
edition (Hodder - Stoughton 1930), published as a
facsimile edition by the Wakefield Press, 1996.]

1923 Igneous rocks of the Mount Painter belt.
Transactions of the Royal Society of South Australia
47; 37^387.

1925 Evidence and indications of algal contributions in
the Cambrian and Pre-Cambrian limestones of South
Australia. Transactions of the Royal Society of South
Australia 49: 186-190.

1926a Wooltana basic igneous belt. Transactions of the
Royal Society of South Australia 50: 192-200.

1926b A brief resume of present knowledge relating to
the igneous rocks of South Australia. Reports of the
Australasian Association for the Advancement of
Science 18: 229-274

1927a The Paralana hot spring. Transactions of the
Royal Society of South Australia 51: 391-397.

1927b Geological notes on the area along the north-
eastern margin of the north-eastern portion of the
Willouran Range. Transactions of the Royal Society
of South Australia 51 : 386-390.

1930 (with C. T. MADIGAN). The pre-Ordovician rocks
of the McDonnell Ranges, Central Australia.
Quarterly Journal of the Geological Society of
LondonS6: 415^28.

1938 The Mount Carnarvon Series of Proterozoic Age.
Transactions of the Royal Societ\> of South Australia
62:347-351.

1939a The Cambrian sequence in the Wirrealpa Basin.
Transactions of the Royal Society of South Australia
63(2): 331-347.

1 939b The late Proterozoic sediments of South Australia.
Reports of the Australasian and New Zealand
Association for the Advancement of Science 24: 80-
88.

1942 The structural character of the Flinders Ranges.
Transactions of the Royal Society of South Australia
66(2): 262-272.

1944a Unpublished memorandum on housing the
department of geology, The University of Adelaide.
February 1944.

1944b The nature and occurrence of uraniferous mineral
deposits in South Australia. Transactions of the Royal
Society of South Australia 68(2): 334-357.

1945a (with W. B. DALLWITZ). The soda-rich
leucogranite cupolas of Umberatana. Transactions
of the Royal Society of South Australia 69(1): 22-
49.

1945b (with W. B. DALLWITZ). Scapolitised dolomites
of Yankaninna. Transactions of the Royal Society of
South Australia 69(2): 212-216.

1947 The Adelaide Series as developed along the
western margin of the Flinders Ranges. Transactions
of the Royal Society of South Australia 71(2): 259-
280.

1949a The Elatina glaciation: a third recurrence of
glaciation evidenced in the Adelaide System.
Transactions of the Royal Society of South Australia
73: 117-121.

1949b The late Precambrian ice-age and glacial record
of the Bibliando Dome. Journal and Proceedings
of the Royal Society of New South Wales 82: 150-
174.

1950 (with R. C. SPRIGG). Subdivision of the Adelaide
System. Australian Journal of Science 13(3): 69-
70.

1958 Australites in the vicinity of Florieton, South
Australia. Transactions of the Royal Society of South
Australia HI: 161-163.

References

ALDERMAN, A. R. 1959. Professor Sir Douglas
Mawson Memoir and Bibliography. Transactions of
the Royal Society of South Australia 82: 1-6.

ALDERMAN, A. R. 1967. The development of geology
in South Australia. Records of the Australian
Academy of Science 1(2): 30 52.

BICKEL, L. 1977. This Accursed Land. Macmillan
Australia.

CORBETT, D. W., COOPER, B. J. & MOONEY, P. M.
1986. 'Geology' in 'Ideas and Endeavours - the
Natural Sciences in South Australia'. Royal Society of
South Australia Occasional Publication Number 5.

DAVID, T. W. E. (with R. HELMS & E. F. PITTMAN)
1901. Geological notes on Kosciusko, with special
reference to evidences of glacial action. Proceedings of
the Linnean Society of New South Wales 26: 26 74.

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DAVID, T. W. E. & PRIESTLEY, R. 1914. Geology
Vol. 1. Reports, on the Scientific Investigations, the
British Antarctic Expedition, 1907-08.

DAVID, T. W. E. 1928. Notes on the newly-discovered
fossils in the Adelaide Series (Lipalian?), South
Australia. Transactions of the Royal Society of South
Australia SI: 191-209.

DAVID, T. W. E. & TILL YARD, R. J. 1936. Memoir
on fossils of the late Pre-Cambrian (Newer
Proterozoic) from the Adelaide Series, South
Australia. Angus and Robertson, Sydney.

FERRAR, H. T. 1907. Report on the field geology of the
region explored during the 'Discovery' Antarctic
Expedition, 1901-04. National Antarctic Expedition
1901-04, Natural History, 1, Geology (Field
Geology: Petrography). British Museum, London.

GLAESSNER, M. F. & DAILY, B. 1959. The geology
of the late Precambrian fauna of the Ediacara Fossil
Reserve. Records of the South Australian Museum
13(3): 369-401.

HOWCHIN, W. 1897. On the occurrence of Lower
Cambrian fossils in the Mount Lofty Ranges.
Transactions of the Royal Society of South Australia
20(2): 74-86.

HOWCHIN, W. 1901. Preliminary note on the existence
of glacial beds of Cambrian age in South Australia.
Transactions of the Royal Society of South Australia
25: 10-13.

HOWCHIN, W. 1914. The occurrence of the Genus
Cryptozoon in the ? Cambrian of Australia.

Transactions of the Royal Society of South Australia
38: 1-10.

HOWCHIN, W. 1922. A geological traverse of the
Flinders Range from the Parachilna Gorge to the Lake
Frome Plains. Transactions of the Royal Society of
South Australia 46: 46-82.

INNES, M. & DUFF, H. 1990. 'Mawson's Papers 1 . The
Mawson Institute for Antarctic Research, The
University of Adelaide.

JACKA, F. 1986. Mawson, Sir Douglas(1882-1958).

Australian Dictionary of Biography 10: 454-457.
MAWSON, P. 1962. 'Mawson of the Antarctic'.

Longmans, London.
PARER, P. & PARER-COOK, E. 1983. 'Douglas

Mawson the Explorer'. Alella Books ABC.

SEGNIT, R. W. 1939. 'The Pre-Cambrian Cambrian
Succession'. Bulletin of the Geological Survey of
South Australia 18.

SHACKLETON, E. 1911. 'The Heart of the Antarctic'.
Heinemann, London.

SPRIGG, R. C. 1947. Early Cambrian (?) jellyfishes
from the Flinders Ranges, South Australia.
Transactions of the Royal Society of South Australia
72(2): 212-224.

SPRIGG, R. C. 1949. Early Cambrian 'Jellyfishes' of
Ediacara, South Australia. Transactions of the Royal
Society of South Australia 73: 72-99.

SPRIGG, R. C. 1986. The Adelaide Geosyncline: a
century of controversy. Earth Sciences History 5(1):
66-83.

REVISION OF AUSTRALIAN ENOCHRUS THOMSON
(COLEOPTERA: HYDROPHILIDAE)

C.H.S. WATTS

Summary

The Australian members of the hydrophilid genus Enochrus are revised and redescribed. A key to
the fourteen species recognised is given. Six species are described as new: E. aliciae, E.
eubenangeei, E. isabellae, E. pseudoweiri, E. samae, and E. weiri. The following synonymies are
proposed: E. mjobergi Knisch, 1921 - E. deserticola (Blackburn, 1896); E. andersoni (Blackburn,
1896) = E. eyrensis (Blackburn, 1894); E. persimilis Regimbart, 1908 = E. eyrensis (Blackburn,
1894); E. pullus (Fauvel, 1883) = E. esuriens (Walker, 1858) and E. artensis (Fauvel, 1883) = E.
maculiceps (MacLeay, 1873). The New Caledonian species E. caledonicus (Fauvel, 1883) is
considered a synonym of E. elongatus (MacLeay, 1873).

REVISION OF AUSTRALIAN ENOCHRUS THOMSON
(COLEOPTERA: HYDROPHILIDAE)

C. H. S. WATTS

WATTS, C. H. S. 1998. Revision of Australian Enochrus Thomson (Coleoptera;
Hydrophilidae). Records of the South Australian Museum 30(2): 137-156.

The Australian members of the hydrophilid genus Enochrus are revised and redescribed. A
key to the fourteen species recognised is given. Six species are described as new: E. aliciae, E.
eubenangeei, E. isabellae, E. pseudoweiri, E. samae, and E. weiri. The following synonymies
are proposed: E. mjobergi Knisch, 1921 - E. deserticola (Blackburn, 1896); E. andersoni
(Blackburn, 1896) - E. eyrensis (Blackburn, 1894); E. persimilis Regimbart, 1908 * E.
eyrensis (Blackburn, 1894); E. pullus (Fauvel, 1883) = E. esuriens (Walker, 1858) and E.
artensis (Fauvel, 1883) = E. maculiceps (MacLeay, 1873). The New Caledonian species E.
caledonicus (Fauvel, 1883) is considered a synonym of E. elongatus (MacLeay, 1873).

C. H. S. Watts, South Australian Museum, North Terrace, Adelaide, South Australia 5000.
Manuscript received 19 March 1997.

The hydrophilid genus Enochrus Thomson,
1859 is world wide in distribution (Hansen 1991).
In Australia its members are the commonest and
most widespread hydrophilids and occur in
virtually all fresh water bodies from small
stagnant pools to the banks of major rivers and
lakes. Despite this prominence, their taxonomy is
so bad that no species can be identified with any
confidence. Thirteen species have been named and
several others have been recorded as being present
but until now no revision has been attempted.

Australian Enochrus are structurally very
similar and difficult to separate. Apart from the
aedeagi the only characters that I have found
useful are size, the punctation of the upper surface,
the slope of the mesosternal keel, and the colour
pattern of the head. The colour pattern of the head
is often a reasonable indicator of species, whereas
elytral and pronotal colour can vary from
predominantly black to totally light reddish in the
same species.

Within Enochrus six subgenera are recognised
(Hansen 1990). Three of these occur in Australia:
Hydatotrephis MacLeay, 1873 with one species;
Enochrus Thomson, 1859 with one species; and
Methydrus Rey, 1885 with 12 species.

At first sight E. (H.) mastersi is very similar to
the distantly related Limnoxenus zealandicus
Broun and is often confused with it in collections.
Both are moderate sized (10-15 mm), shiny black
and relatively common, particularly L.
zealandicus. A closer inspection readily separates
them: L. zealandicus has well defined punctate
elytral striae and swimming hairs on the meso-

and meta-tibiae, both of which are absent in E,
(H.) mastersi. Characters of the maxillary palpi
and mesosternum also separate the two genera
(Hansen 1990).

The one Australian Enochrus (Enochrus)
species, E. (E.) peregrinus Knisch, 1922, is small
and in general appearance resembles several
Australian Enochrus (Methydrus) species. It is
known from only one specimen labelled from
Sydney, although there were three in the original
collection. It may eventually prove to have been
mis-labelled and not Australian.

The remaining 12 Australian species are all in
the subgenus Methydrus. They are usually small
(2 8 mm), oval, rather flat species, shiny with
light-testaceous to black colouring with at least
the area immediately in front of the eyes
yellowish. Two species are larger and
predominantly black, superficially resembling E.
(H.) mastersi. Chasmogenus nitescens Fauvel
resembles some Enochrus (Methydrus) species
and is often confused with it in collections. Its
small size (< 5 mm) and lack of yellowish areas
in front of its eyes readily separate it from similar
sized Enochrus - as well as many structural
characters mentioned later under 'Systematics'.

One additional species, Philhydrus marmoratus
W. MacLeay, 1873, has been included in
Enochrus in the past but has been shown by
Gentili, 1981 to belong in Laccobius.

For a full discussion of synonyms and
subgenera of Enochrus see Hansen, 1991.

Despite their being ubiquitous in freshwater the
biology of Australian Enochrus is poorly known.

C. H. S. WATTS

Anderson (1976) described the larvae of E. (M.)
maculiceps and E. (M.) elongatus and gave brief
notes on their habitat and breeding. All species
come readily to light and are often collected in
large numbers in light traps, particularly in
northern areas during the wet season from
December to February. I know of no study of the
ecology of any Australian species other than
inclusion in general faunal lists of invertebrates
collected in various water bodies.

The collections from which specimens were
examined are listed under the following
abbreviations:

AM Australian Museum, Sydney

AN1C Australian National Insect Collection

BM(NH) Natural History Museum, London

IRSNB Institut Royal des Sciences Naturelles
de Belgique, Bruxelles

MNHN Museum National d'Histoire Naturelle,
Paris

NMV Museum of Victoria

NRS Naturhistoriska Riksmuseet, Stockholm

NTM Northern Territory Museum

QDPIM Queensland Department of Primary
Industries, Mareeba

QM Queensland Museum, Brisbane

SAMA South Australian Museum, Adelaide

UQIC University of Queensland Insect
Collection, Brisbane

WAM Western Australian Museum, Perth.

Systematics

Australian members of the genus Enochrus can
be separated from other Australian Hydrophilids
by the following combination of characters.
Length 2-9 mm, oval, more flattened than convex;
second segment of maxillary palpi more or less
curved outwards, apical segment slightly
asymmetrical with straighter inner face; without
contiguous ventral keel but with variable keel on
mesosternum; without swimming hairs on tibiae;
without strongly marked striae on elytron; with
systematic punctures (rows or fields of coarse
setiferous punctures usually distinctly larger than
others) on head and pronotum (masked in some E.
malabarensis); in all but E. mastersi and E.
aliciae at least the area immediately in front of
eyes lighter (see also Hansen 1991).

The subgenus Hydatotrephis MacLeay can be

separated from Australian members of the
subgenera Enochrus Thomson and Methydrus
Rey as follows: mesosternal protuberance
triangular, cone-shaped, abruptly truncated
posteriorly which does not reach the front edge of
mesocoxae (Fig. 19), sides of mesosternum evenly
converging anteriorly, apical and penultimate
segments of maxillary palpi subequal. Enochrus
(Methydrus) have a more keel-like mesosternal
process usually extending backwards between the
mesocoxae (Figs 15-18; 20), the apical segment
of the maxillary palpi shorter than penultimate and
sides of mesosternum subparallel to strongly
convex anteriorly. Apart from E. (M.) aliciae and
E. (M.) eubenangeei, the Australian members of
E. (Methydrus) are smaller (< 8 mm), often
testaceous in colour, and have at least a lighter
patch of colour in front of the eyes.

The aedeagus of E. (H.) mastersi differs from
that of other Australian Enochrus (Fig. 14;
Hansen 1990). In these there is a variably shaped
basal ventral plate beyond which projects a
thinner apical portion in all but E. (M.) aliciae
which lacks this apical portion (Fig. 11). In E.
(H.) mastersi the aedeagus is of more uniform
construction with a smooth dorsal surface and the
ventral surface consists of a broad open groove
formed by infolding of the two sides.

There is one species, E. (E.) peregrinus Knisch,
of the subgenus Enochrus in Australia. It is
similar in looks to the smaller Methydrus species
but is readily separated from them by the short
stout maxillary palpi with approximately equal
sized apical two segments and the diverging
elytral suture lines.

By utilising the male genitalia species can be
differentiated. However without these I have found
it impossible to reliably separate all species, in
particular E. weiri and E. pseudoweiri and the
lighter colour phase of E. deserticola and E.
maculipes. This is reflected in the keys and
descriptions that follow. 1 have restricted my
descriptions to the few characters that serve to
differentiate the species and I have provided two
keys, one utilising all available characters, the
other characters other than the male genitalia.

Key to Australian Enochrus Thomson 1859,
not using characters of the male genitalia

1 . — Maxillary palpi stout, first segment shorter than
distance from front edge of eye to front of
clypeus, apical segment subequal in length to

penultimate 2

Maxillary palpi more elongate, first segment

AUSTRALIAN ENOCHRUS

139

longer than distance from front edge of eye to
front margin of clypeus, apical segment about
2/3 length of penultimate (subgenus
Methydrus) 3

2. — > 6.0 mm. Black. Sutural lines on elytra

subparallel in anterior quarter ,

E. {Hydatotrephis) mastersi (MacLeay)

— < 5.0 mm. Dark testaceous. Sutural lines on
elytra diverging noticeably in anterior quarter ..

E. (Enochrus) peregrinus Knisch

3. — Large (>6 mm). Black (pronotum may be

reddish at edges) 4

Small (< 6 mm). With at least the areas in front
of eyes reddish-yellow 5

4. - Punctures towards sides of elytra well

impressed, not much smaller than others

E. (M.) euhenangeei sp. nov.

— Punctures towards sides of elytra subobsolete,
much smaller than those on disc

E. (M.) aliciae sp. nov.

5. — Dorsal surface reddish-yellow, rear of head at

most somewhat darker. Elytra moderately
punctate, length > 4.0 mm, pro-, meso- and

metaclaws modified in male

E. (M.) elongatus (W . MacLeay)

- Dorsal surface reddish-yellow to black, rear of
head dark and much of rest of head in many
species, males with only pro-claws modified 6

6. — Dorsal surface strongly punctured, punctures on

elytra as strong or stronger than those on head,
many punctures on elytra >half diameter of
serial punctures which are often hard to trace.
Front of head 1/3 or more black 7

— Dorsal surface weakly punctured, punctures on
elytra tend to be weaker than those on head,
punctures on elytra < half size of serial
punctures which are usually distinct 9

1, — Length >3.5 mm. Black morphs present.
Maxillary palpi with dark tips 8

Length < 4.0 mm. Black morphs not known.

Maxillary palpi usually pale throughout

E. (E.) malabarensis Regimbart

8. — Thinner outer portion of mesosternal keel well

developed (Fig. 16). Front of head

predominantly black

E. (M.) eyrensis (Blackburn)

— Thinner outer portion of mesosternal keel less
developed (Fig. 15). Amount of black on front
of head variable E. (M.) samae sp. nov.

9. — Length 2.2 mm - 2.6 mm. Mesosternal keel

weakly developed (Fig. 1 8). Front of head > 2/3
black E. (M.) esuriens (Walker)

— Length > 2.6 mm. Mesosternal keel well
developed (Fig. 17) 10

10. — Larger (4.5-6 mm). Black, except region in
front of eyes light. Elytral punctures small but

usually well marked

E. (M.) isabellae sp. nov.

— Smaller (3-4.5 mm) E. (M.) deserticola complex
(£. (M.) deserticola (Blackburn), E. (M.)
maculiceps (W. MacLeay), E. (M.) weiri sp. nov
and E. (M.) pseudoweiri sp. nov). (E. (M.)
deserticola only species with black morphs and
individuals > 4.2 mm long. Enochrus weiri and
E. pseudoweiri always have pale palpi.)

Key to Australian Enochrus incorporating
characters of the male glnitalia

— Maxillary palpi stout, pseudo first segment
shorter than distance from front edge of eye to
front of clypeus, apical and penultimate
segments subequal in length 2

— Maxillary palpi more elongate, pseudo first
segment longer than distance from front edge of
eye to front margin of clypeus, apical segment
noticeably shorter than penultimate (subgenus
Methydrus) 3

— > 6.0 mm. Black. Sutural lines on elytra
subparallel in anterior quarter

E. (Hydatotrephis) mastersi (MacLeay)

< 5.0 mm. Dark testaceous. Sutural lines on
elytra noticeably diverging in anterior quarter ..
E. (Enochrus) peregrinus Knisch

— Large (> 6.0 mm). Black (pronotum may be
reddish at edges) 4

— Small (< 6.0 mm). With at least the areas in
front of eyes reddish-yellow 5

— Aedeagus broad reaching little beyond half
length of parameres (Fig. 1 1). Parameres flat. ..

E. (M.) aliciae sp. nov.

Aedeagus with broad basal piece and long thin
apical piece bending upwards and reaching
nearly to end of parameres, which also bend

upwards (Fig. 10)

E. (M.) euhenangeei sp. nov.

— Tips of parameres hooked (e.g. Fig. 4) 6

— Tips of parameres not hooked (e.g. Fig. 1) 9

— Dorsal surface reddish-yellow, rear of head at
most somewhat darker. Aedeagus tip strongly
crotchet-shaped (Fig. 7). Pro-, meso- and
metaclaws modified in male

E. (M.) elongatus (W. MacLeay)

Variably coloured but always with rear of head
clearly darker than at least portions of front of
head. Aedeagus with little or no dorsal/ventral
expansion at tip, males with only proclaws
modified 7

140

C. H. S. WATTS

7. — Dorsal surface strongly punctured, many

punctures on elytra > half size of serial
punctures E. (M.) eyrensis (Blackburn)

— Dorsal surface weakly punctured, punctures on
elytra < half size of serial punctures 8

8. — Larger (4.5 - 6.0 mm). Black, except in front of

eyes. Aedeagus shorter and apical portion
broader (Fig. 6) E. (M.) isabellae sp, nov.

— Smaller (3.0-4.0 mm). Pale morphs common.
Aedeagus longer and apical portion thinner
(Fig. 4) E. (M.) deserticola (Blackburn)

9. — Length <2.8 mm. Front of head > 2/3 black.

Mesosternal keel weak (Fig. 18)

E. (M.) esuriens (Walker)

— Not as above 10

1 0. — Dorsal surface strongly punctured, punctures on

elytra as strong as, or stronger than, those on
head, many punctures on elytra > half diameter
of serial punctures 1 1

— Dorsal surface weakly punctured, punctures on
elytra tend to be weaker than those on head,
punctures on elytra < half size of serial
punctures 12

11. — Larger (4.4-5.6 mm). Black morphs present.

Aedeagus with moderate apical pad, inner edges

of parameres sinuate (Fig. 9)

E. (M.) samae sp. nov.

— Smaller (<3.8 mm). No black morphs.
Aedeagus narrow with, at most, a weak pad,
inner edges of parameres straight (Fig. 12)

E. (M.) malabarensis Regimbart

12. — Tips of parameres truncated or weakly bifid

(Fig. 8) E. (M.) weir i sp. nov.

— Tips of parameres rounded (Figs I, 2, 3) 13

13. — Aedeagus moderately broad with slight to well-

marked apical pad (Figs 1, 2)

E. (M.) maculiceps (W. MacLeay)

— Aedeagus narrow, sharply pointed, lacking
apical pad (Fig. 3)

E. (M.) pseudoweiri sp. nov.

Descriptions

Subgenus Enochrus

Enochrus (Enochrus) peregrinus Knisch, 1922

Types

Lectotype: 9 'Sidney Mus. Godeffroy No.
10705, coll. Knisch, coll. d'Orchymont'; 'ex. col.
Knisch, No. 5251438, coll. d'Orchymont';

'Knisch det. 1921, Enochrus peregrinus m';
'Coll. A. Knisch, TYPUS', on red card, in
IRSNB, herein designated.

Knisch (1922) mentioned three specimens (No.
10705) in the Hamburg Zoological Museum. I
have been able to trace only one. He mentioned
that one was labelled 'M. Regimbart det. 1905'
and the others with 'Wehncke determ'. The
surviving specimen, now in Brussels, lacks such a
label but would otherwise appear to be part of the
syntype series.

Description (number examined, 1)

Length 3.2 mm. Broadly oval. Elytra dark-
testaceous, pronotum very slightly lighter, head
black except for small indistinct testaceous patch
along margin in front of each eye, maxillary palpi
light-testaceous, tips darker. Punctures on head
moderate, approaching size of eye facet, most
separated from each other by at least puncture
width. Systematic punctures rather sparse, about
twice diameter of adjacent punctures. Pronotum
rather weakly and sparsely punctured, serial
punctures also relatively small, indistinct, about
2x diameter of adjacent punctures, punctures on
elytra stronger than on pronotum, sparse, many 2x
diameter or more apart, a little stronger laterally.
Serial punctures not traceable. Maxillary palpi
short, stout, pseudo basal segment not reaching
front edge of eye, apical and penultimate segment
subequal in size.

Male: Unknown.

Distribution

Known only from Sydney, the type locality,

Remarks

This unique specimen is separated from all other
Australian Enochrus, other than the much larger E.
mastersi, by the short stout maxillary palpi. The
masking of the elytral strial punctures is found only
in the much larger E, eyrensis and E. samae and in
E. malabarensis. Enochrus malabarensis differs
from E. peregrinus by the more normal (for
Australian species) maxillary palpi, the more
extensive testaceous areas in front of the eyes and
lighter coloured elytra. The elytral sutures continue
to diverge noticeably in anterior quarter whereas in
E. malabarensis they are subparallel. Most E.
malabarensis also have stronger punctures, but this
character is variable and tends to be relatively weak
in the few Sydney area specimens I have seen. The
pronotal punctures in E. peregrinus are weaker
than those on elytra whereas in E. malabarensis
they are similar sized.

AUSTRALIAN ENOCHRUS

141

The fact that only the type specimen is known
worries me. For all the other small Australian
Enochrus, specimens are relatively abundant.
Also it is the only known specimen in the
Australian region of the subgenus Enochrus
(Hansen 1990). This is the sort of situation where
mislabelling should be considered. This is beyond
the scope of this revision.

Subgenus Hydatotrephis

Enochrus (Hydatotrephis) mastersi (MacLeay)

Hydatotrephis mastersi MacLeay, 1871

= Parana simplex Knisch, 1922: Hansen, 1990

Enochrus (Hydatotrephis) mastersi (MacLeay):
Hansen, 1990

Types

Hydatotrephis mastersi: Lectotype: ( 9- ), (round
red label), 'K 1950T, "Hydatotrephis mastersi
Mc L.W. Gayndah'. 'Lectotype: Hydatotrephis
mastersi W. J. MacLeay designated by M. Hansen
1990', AM.

Parana simplex: Lectotype: (unsexed) 'Sidney
Mus. Godeffroy. No. 10704'; 'Philhydrus x M.
Regimbart determ. 1905'; 'Parana simplex
Knisch n.g. et sp. A. Knisch det. 1921', round
black label; 'Lectotype Parana simplex Knisch
designated by M. Hansen 1988', IRSNB.
Synonymy after Hansen 1990.

Description (number examined, 71) Figs 14, 19

Size 8.0 mm - 9.1 mm. Broadly oval. Relatively
flat. Black; lateral edges of elytra, pronotum and
head, palpi and apical portions of legs diffusely
dark-testaceous. Maxillary palpi stout, length
about width of head in front of eyes, apical and
penultimate segments subcqual in length, together
a tittle larger than pseudo first segment. Head
strongly punctured, each about size of eye facet,
most closer than a puncture width apart,
systematic punctures 2-3x size of normal
punctures. Pronotum with punctures a little
smaller, shallower and more separated than on
head, systematic punctures well marked, with
sharp groove sometimes weak or lacking on front
edge. Elytra punctured as on pronotum, each
elytron with four rows of larger serial punctures,
punctures in rows increasing in both number and
scatter towards sides, elytra weakly grooved and
flanged laterally, a few scattered large punctures
between puncture rows two and three counting
from suture. Mesosternal keel cone-like, strongly

raised, narrowly triangular from both lateral and
front views, sharply truncated behind just prior to
mesocoxal cavities so that no part of the keel
extends between mesocoxae.

Male: Aedcagus broad, lacking collar or ventral
plate, sides folded in ventrally tending to form
broad channel. Parameres stout, broad, narrowing
shortly before tips which tend to be turned inward
to varying degrees.

Distribution

Australian Capital Territory

Black Mt, ANIC; Canberra, SAMA, ANIC; 3
km E Piccadilly Circus, ANIC; 6 km NE
Piccadilly Circus, ANIC.

New South Wales

Allyn River, ANIC; Barrington River, ANIC;
Blue Mts, ANIC Chichester St Forrest, ANIC;
Coffs Harbour UQIC; Como, ANIC; Galston,
SAMA; Lansdowne via Taree, ANIC;
Muswellbrook, ANIC; Nerringa, SAMA; 12 km
NW Nellinger, ANIC; 2km NE by N Rousmill,
ANIC; Salisbury UQIC; Stanwell Park UQIC;
Sydney, SAMA, ANIC; Tooloom Plateau via
Woodenbong UQIC; Ulladulla, ANIC; Valery,
ANIC; Wee Jasper, ANIC.

Northern Territory

30 ml W Alice Springs, SAMA; Stanley
Chasm, SAMA.

Queensland

Brisbane UQIC; Bulburin State Forest via
Many Peaks UQIC; Bunya Mt, ANIC; Carnarvon
UQIC; Discot, ANIC; 13 km SW by S
Gordonvale, ANIC; Joalah Nt Pk Tamborine Mt,
ANIC; Mt Norman, via Wallangarra UQIC; Mt
Spec, ANIC; Upper Cedar Ck, via Sandford
UQIC; 8 km E Wallaman Falls, SAMA.

Victoria

Bagots Ck, SAMA; Ferntree Gully UQIC;
Healesville, SAMA; 10 mis N of Valencia Creek
via Maffra UQIC; Victorian Alps, SAMA.

Remarks

A widespread and relatively common species
which, within Australian Enochrus, can only be
confused with E. aliciae and E. eubenangeei. It is
readily separated from these species by the stout
maxillary palpi which have the last two segments
subequal instead of having the apical segment
about two-thirds the length of the penultimate as

142

C. H. S. WATTS

in all other Australian Enochrus. The cone-like
shape of the mesosternal keel is also diagnostic.

Enochrus mastersi is most frequently confused
with a very different, but similar looking and very
common species, Limnoxenus zealandicus, of the
subtribe Hydrobiina. Although very similar in
general appearance Limnoxenus can be separated
from E. mastersi by the presence of nine well-
marked series of punctures on each elytron,
swimming hairs on the meso- and metatarsi and
virtual absence of rugose portions on meso- and
metafemora.

Subgenus Methydrus

Enochrus (Methydrus) aliciae sp. nov.

Types

Holotype: o\ '15°16*S, 144°59'E, 14 km W by
N of Hope Vaie Mission, Qld, 8-10 Oct. 1980, T.
Weir', ANIC.

Paratypes: 13, same data as holotype, 10
ANIC, 3 SAMA; 1, '12*31% 132°54'E, 9 km N
by E of Mudginbarry Hs„ N.T., 10.vi.1973, T.
Weir and A. Allwood, M3985", NTM.

Description (number examined, 17) Figs 11, 20

Length 7.5 mm - 8.2 mm. Broadly oval,
relatively flat. Shiny, black, appendages, apical
and lateral edges of head, pronotum and elytra
dark-reddish. Maxillary palpi moderately long,
pseudo first segment longer than distance from
eye to front of clypeus, longer than maxillary stipe,
apical segment 2 I to 3 / 4 length of penultimate.
Head strongly punctate, punctures about puncture
width apart, about size of eye facet or a little
smaller. Systematic punctures large about 3x size
of adjacent punctures. Punctures on pronotum
similar to or a little weaker than those on head,
systematic punctures well marked, 3~4x diameter
of adjacent punctures, rear and lateral margins
weakly grooved and flanged, front margin grooved
to about level of inner edge of eyes. Elytral
punctures as on pronotum, more weakly
impressed towards rear. Serial punctures in four
loose rows, sparse and indistinct, more numerous
but more scattered laterally, elytra weakly grooved
and flanged laterally. Mesosternal pillar well
marked, triangular in both front and lateral views,
posterior edge often steeper near top of keel,
reaching a short distance between mesocoxae.

Male: Aedeagus very short, broad, rounded,
lacking narrower apical section. Parameres squat,
inner edge a little sinuate, tips obliquely truncated

on inside or curved weakly outwards. Proclaw
straightened with strong basal swelling, mesoclaw
thickened, curved with basal swelling, metaclaw
straightened and greatly swollen in basal half. In
female claws similar but with straightening and
basal swelling much weaker.

Distribution

Known only from the type localities in North
Queensland and the Northern Territory.

Remarks

Readily separated from all other Australian
Enochrus (Methydrus), other than E.
eubenangeei, by its large size and almost all-black
colouring. The very short, broad aedeagus, and
weak lateral punctures on the elytra separate it
from the otherwise very similar E. eubenangeei.

In general appearance it is similar to E. (H.)
mastersi but readily separated by its longer
maxillary palpi with the apical segment shorter
than penultimate. See also under remarks in E.
mastersi.

Enochrus (Methydrus) deserticola (Blackburn)

Philhydrus deserticola Blackburn, 1896

Enochrus (Lumetus) deserticola (Blackburn):
Knischl924

—Enochrus mjobergi Knisch, 1921: syn. nov.

—Enochrus (Lumetus) mjobergi Knisch: Knisch
1924

^Philhydrus temporalis Regimbart, 1908: syn.
nov.

^Enochrus (Lumetus) temporalis (Regimbart):
Knisch 1924

Types

Enochrus deserticola: Lectotype: '5487, Bl.
Paisley T' top specimen of two separately carded
on same pin, BM(NH), herein designated.

Paralectotypes: 1 unlabelled but mounted
below lectotype on same pin, BM(NH); 1 6 '5487
Palm Cr', ' Philhydrus deserticola Blackb. Co-
type', SAMA; 3, 'Reedy Hole', NMV; 2, 'Ellery
Ck\ NMV; 2, 'Paisley Bluff, NMV.

Enochrus mjobergi: Lectotype: 6 'Cape York
Penins'; 'Queensl. Mjoberg'; 'Type', ''Enochrus
(Lum.) mjobergi m. n. sp., A. Knisch 1921 ';
'Typus', on red label NRS, herein designated.

Paralectotypes: 1, 'Queensl. Mjoberg Cape
York Penins'; 'Sjostedt don 1921'; 'Ex Coll.
Knisch No. 5521544 Coll. d'Orchym'; 'Knis det

AUSTRALIAN ENOCHRUS

Enochrus (L.) mjobergi m'; Coll. A. Knisch
TYPUS', on red card. IRSNB; 3, 'Cape York
Penins'; 'Queensl. Mjoberg', NRS. Synonymy
after examination of types.

Enochrus temporalis: Holotype: 'Avon R';
'eaudouce'; ' Philhydrus temporalis, Reg.
TYPUS % MNHN. Synonymy based on
descriptions and examination of types (of E.
temporalis in 1964, not currently available).

Description (number examined, >1000) Fig. 4

Oblong oval, length 3.3 mm - 4.9 mm. Two
colour morphs; one with dorsal surface black
except light portions in front of eyes, other with
elytra and pronotum light testaceous and head
dark testaceous to black except for light testaceous
area forward from eyes and a little wider than
eyes, leaving central darker panel of variable
width on front of head, underside dark testaceous,
appendages lighter towards extremities except for
maxillary palpi which usually have apical portion
of last segment darker. Head covered with small,
sharply impressed, well separated punctures a lot
smaller than eye facets, a group of large punctures
inwards from each eye, at least 4x diameter of
other punctures. Punctures on pronotum as on
head, but more shallowly impressed, systematic
punctures as in E. elongatus, four or more times
diameter of adjacent punctures. Elytral punctures
increase somewhat in strength apically and
laterally, serial punctures large, sparse and quite
distinct at apex, interstrial punctures varying from
weak to almost obsolete.

Mesosternal keel well developed, front edge
nearly perpendicular, ventral edge straight or
slightly convex.

Male: Aedeagus pointed, with weak dorsal pad
at tip, collar a bit closer to tip than base.
Parameres outwardly hooked at tips. Proclaw with
basal lobe and rest of claw somewhat
straightened. Meso- and mctaclaws, with slight
basal lobes, weakly curved.

Female: Claws only slightly weaker than males.

Distribution

New South Wales

Bonville, ANIC; Chichester SF, ANIC
Clarence River, NMV; Coffs Harbour, ANIC
Fowlers Gap, ANIC; 10 km N Jabrru, QDPIM
Lord Howe Island, SAMA; Moree, SAMA
Valcry, ANIC; Wootton, ANIC.

Northern Territory

Adelaide River, ANIC; E Alligator River, AM;
Bessie Spring, ANIC; Berry Spring, ANIC; 45

km SW by S Borroloola, ANIC; 7 km NW by N
Mt Cahill Crossing East Alligator River, ANIC; 8
km N Mt Cahill, SAMA; 12 km NNW Mt Cahill,
SAMA; 14 km S by W Cape Crawford, ANIC;
Cooper Creek, ANIC; Daly River Mission, ANIC;
170 km E Daly Waters, SAMA; Darwin, ANIC,
SAMA; Ellery Ck, NMV; 15 km S Elliot, SAMA;
Harst Bluff, NMV; 12 km NE Humpty Doo,
QDPIM; Jim Jim Creek, ANIC; Koogarra, ANIC;
4 km N McArthur R Stn, SAMA; 19 km SSE
Mataranka, SAMA; 5 km E Mataranka, SAMA;
Nourlangie Creek, ANIC; 18 km E by N Oenpelli,
ANIC; Paisley Bluff, NMV; Pine Creek, QDPIM;
Reedy Hole, NMV; 46 km S Renner Springs,
SAMA; Stanley Chasm, SAMA, ANIC; 100 km
W Tennant Creek, QDPIM; Tindal, ANIC;
Wessel Island, ANIC.

Queensland

Ashmore, QM; Atherton, ANIC, QDPIM; 31
km NE Aramac, SAMA; Bakerville, QDPIM;
Bamaga, SAMA; Brisbane, QM, UQIC;
Benaraby, ANIC; 5 mis N Bloomfield River,
ANIC; Bundaberg, ANIC; Bunya Mts, ANIC;
Burketown, QM; Cairns, ANIC, SAMA; Calliope
River, ANIC; Camooweal, QDPIM; Cape
Tribulation, ANIC; Cardstone, ANIC; Chillagoe
Creek, ANIC, QDPIM; Coen, QDPIM; 60 km S
Coen, SAMA; Mt Coolum, ANIC; Cooktown,
ANIC; 30 mis N Cooktown, UQIC; Crystal
Creek, ANIC; Cunningham's Gap, ANIC;
Dalhunty River, SAMA; Edungalba, ANIC; 5 km
NE Edungalba, SAMA; Eidsvold, ANIC; Emu
Park, UQIC; Fletcher, QM; Gayndah, UQIC; Mt
Garnet, ANIC; Greenhills, ANIC; Home Hill,
QDPIM; Ingham, ANIC, SAMA; Mt Inkerman,
ANIC; Iron Rng, QDPIM; Julatten, ANIC; Julia
Creek, SAMA; Kenilworth SF, UQIC; Kingaroy,
ANIC; Kuranda, ANIC, SAMA; Laura, QDPIM;
70 km N Laura, QDPIM; 25 km N Laura,
QDPIM; Mt Lewis, ANIC; Mackay, ANIC;
Magnetic Isl, SAMA; Malanda, SAMA; 23 km E
Marecba, QDPIM; Mary Creek, ANIC; Melvor,
ANIC; 21 mis S Miriam Vale, ANIC; 100 mis S
Miriam Vale, UQIC; Mission Beach, ANIC; 3
mis W Mourilyan, ANIC; Mornington Isl
mission, SAMA; 5 km W Mossman, QM;
Mossman George Nt Pk., UQIC; Mossman
George, ANIC; Normanton, QDPIM; Oxley Ck,
QM; Patuma Dam, ANIC; 9 km W Paluma,
QDPIM; Prince of Wales Isl, NMV; 8 mis NE
Proserpine, ANIC; Rocklea, QM; Stanthorpe,
QM; Mt Spec, ANIC; Stewart Rng, SAMA;
Tolga, SAMA; Townsville, ANIC, NMV, SAMA;
9 km ENE Mt Tozer, ANIC; Weipa, QDPIM;

144

C. H. S. WATTS

Windsor Tableland, QDP1M; Yamba, UQIC;
Yungaburra, QDPIM.

South Australia

Arkaba Creek, SAMA; Mambray Creek,
SAMA; Marne River., SAMA; Mt Remarkable,
SAMA; 6 km E Nilpinna, SAMA.

Victoria

Dimboola, ANIC; Genoa, ANIC; Sea Lake,

SAMA.

have, however, been unable to reliably separate
light coloured females. In general E. deserticola
are larger and more weakly punctured but
considerable character overlap occurs between the
two species.

In more southern areas E. deserticola overlaps
with E. eyrensis, E. samae and E. elongatus but
can be separated from all three by the much
weaker punctation and in the case of E. elongatus
by the well-marked dark patterning on the head
which E. elongatus lacks.

Western Australia

6 km NNW Broome, ANIC; Beverley Springs,
WAM; Cadjeput Rockhole, WAM; Charnley
River, 25 miles N Beverley Springs, WAM;
Derby, SAMA; Drysdale River, ANIC; 7 miles
NE Giles, WAM; Gill Pinnacle, WAM;
Hammesley Rng, SAMA; Koolan Island, WAM;
6.5 km N Mt Bell, WAM; Kununurra, ANIC,
QDPIM; Millstream, ANIC; 5 km SE Millstream
HS., ANIC: Mitchell Plateau, ANIC; West
Peewah River, ANIC; Synnott Range, WAM;
14°53'S, 125°45'E, SAMA.

Remarks

A relatively large species showing considerable
variation in size, colour and strength of dorsal
punctation making its identification difficult
without dissection. The two colour morphs are
well marked and occur together in the same
populations. Occasional specimens have an
intermediate dark-testaceous dorsal surface to a
variable extent. It is my impression that the dark
form is more common in the north where it is the
dominant form in many areas. The black morphs
are readily identified by their weak to moderate
punctation contrasting markedly with the strong
punctation in the two other species with black
morphs, E. eyrensis and E. samae. It is also the
only species with the combination of tips of the
parameres hooked, weak to moderate dorsal
punctures and black patterning on the head. In the
North-east the species overlaps with E. weiri and
E. pseudoweiri. These can be separated from most
E. deserticola by the lack of black morphs,
somewhat stronger punctation, pale tips to the
maxillary palpi and the small amount of black on
the front of the head. An occasional specimen of
E. deserticola may also have pale palpi tips and
much reduced black on the front of the head.

Over most of its range E. deserticola is
sympatric with E. maculiceps. The two species
are readily separated by the male genitalia and
presence of black morphs in E. deserticola. I

Enochrus (Methydrus) elongatus (W. MacLeay)

Philhydrus elongatus W. MacLeay, 1 873

Enochrus (Lumetus) elongatus (W. MacLeay)
Knisch, 1924

= Philhydrus caledonicus Fauvel, 1883: syn. nov.

= Enochrus caledonicus (Fauvel): Knisch, 1924

Types

Enochrus elongatus: Lectotype: 'Philhydrus
elongatus Gayndah McL W'; 'K19506'. Right-
hand specimen of two mounted on same card, AM
herein designated.

Paralectotypes: 1 , same data as lectotype, left-
hand specimen, AM; 3, on same card, same data
as lectotype, ANIC; 1, 'Philhydrus elongatus
McL', 'Griffith Collection Id by A. M. Lea', 'Co-
Type', SAMA; 1, 'Philhydrus elongatus M.
Queensland cotype 14629', SAMA.

Enochrus caledonicus: Lectotype: 'm',
'Nouvelle Caledonie. Noumea Marais Anse Vata,
Octobre Saves, ex. coll. Fauvel', 'Coll. et det. A.
Fauvel Philhydrus caledonicus FvP, '17— 479%
'lectotype' on red label. Remounted with genitalia
dissected out, IRSNB.

Paralectotypes: 1 , 'Nouvelle Caledonie Kanala,
Rec. Coste, ex coll. Fauvel'; 'Coll. et det A.
Fauvel Philhydrus caledonicus Fvl.' '17-479', 1,
'Nouvelle Caledonie. Anse Vata, Noumea Saves',
'Coll. et det A. Fauvel Philhydrus caledonicus
Fvl,' IRSNB. Synonymy after examination of
types.

Description (number examined, 330) Fig. 7

Elongate oval, body flattish, length 4.4 mm -
5.0 mm. Dorsal surface testaceous, diffusely
blotched, lighter towards sides, rear of head often
diffusely darker but never distinctly darker than
rest of head. Ventral surface dark red-brown,
apical portions of legs lighter, palpi light
testaceous, tips of maxillary palpi dark brown.
Head covered with small sharply impressed, well

AUSTRALIAN ENOCIIRUS

145

FIGURES l— 14. Dorsal views of apical portion of male genitalia. 1 & 2, E. maculiceps showing range of variation;
3, E. pseudoweiri; 4, E. deserticola; 5, E. eyrensis; 6, E. isabellae; 7, E. elongatus, lateral view of median lobe; 8,
E. weiri; 9, E. samae: 10, E. euhenangeei; 11, E. aliciae; 12, E. malabarensis; 13, E. esuriens; 14, E. mastersi.

146

C. H. S. WATTS

separated punctures, quarter to half size of eye
facet, a dozen or so much larger punctures
inwards from each eye. Pronotum similarly
punctate, a row of systematic punctures about 3x
diameter of adjacent punctures running inwards
from each front corner and smaller group of
similar punctures a little behind middle on each
side. Elytral punctures slightly stronger than
those on pronotum, stronger laterally than on
disc. Serial punctures, larger, sparse, hard to
trace towards apex. Underside evenly setose-
punctate, rugose portions of femur,
approximately three quarters length of femur.
Mesosternal keel well developed, anterior edge
curved somewhat, nearly perpendicular, ventral
edge flat or slightly convex. Notch on apical
stemite small but well marked.

Male: Aedeagus with crotchet-shaped hook at
tip. Tips of parameres outwardly hooked.
Metaclaws with enlarged basal lobe and rest of
claw straightened. Pro- and mesoclaws curved
with thickened basal portion

Female: Claws curved with thickened basal
portions.

Distribution

Australian Capital Territory

Black Mt, ANIC.

New South Wales

Albury, ANIC; Allyn River, NMV; Armadale,
ANIC; Balranald, ANIC; Bathurst, AM; Bogan
River, AM; 20 mis SSW Bourke, SAMA; Broken
Hill, SAMA; Coonabarabran, AM, ANIC; Cowra,
AM; Ecceleston, AM; Hay, ANIC; 37 km E Hay,
SAMA; Mt Kaputar Nat. Pk., ANIC; Kosciusko
Nt Pk., ANIC; Lake Menindie, SAMA; Menindie,
SAMA; Narrabri, ANIC; Norfolk Island, ANIC;
97 km S Tibooburra, NMV.

Northern Territory

53 km NE Alice Springs, ANIC; 12 km SW by
W Alice Springs, ANIC; Top of Ayers Rock,
WAM; 97 km N Barrow Ck, SAMA; 30 km NE
by E Borroloola, ANIC; Ellery Creek, NMV; Finke,
SAMA; Glen Helen, NMV; Stanley Chasm, ANIC;
15 km East, Vaughan Spring, HS, ANIC.

Queensland

Bollon Dist, AM; 42 km Boulia, ANIC: Bunya
Mts, UQIC; Camooweal, QDPIM; Edgbaston HS,
SAMA; 5 km NE Edungalba, SAMA; Glen
Alpin, UQIC; Helidon, UQIC; 6 km E, Kamma,
ANIC; Mackay, ANIC; MacPherson Rng, Nt Pk.,
AM; Marmor, ANIC; Mt Isa, QDPIM; Mt Spec,

ANIC; Normanton, QDPIM; Tolga, QDPIM;
Toowoomba, UQIC.

South Australia

Aroona Dam, SAMA; Mt Ban, ANIC; Old Billa
Kallma HS, SAMA; Blanche Cup Spring, AM;
Broken Hill, AM; Coopers Creek, SAMA; Coward
Sp., SAMA; 16 km E Curdimurka, SAMA; 36 km
ESE Curdimurka, ANIC; 1 1 km NE by Etadunna
HS, ANIC; Gawler Rngs, SAMA; Lake George
ANIC; Mt Lofty Rng, AM; Mannum, SAMA
Marne River, SAMA, McDovall Peak, UQIC
Murray Bridge, AM, SAMA; Olary, ANIC
Oodnadatta, SAMA; Paralana Hot Springs,
SAMA; Quorn, AM; Renmark, SAMA; Roonka
Stn, SAMA; Roseworthy, SAMA; Scorpion
Springs CP, SAMA; River Torrens, SAMA;
Warburton, River, ANIC; Wearing Gorge, SAMA;
Willochra Ck, SAMA.

Tasmania

15 mN Waratah, UQIC.

Victoria

Eppalock Res., NMV; Eustace Gap Ck, NMV;
King Lake Nt Pk., NMV; Kiata, NMV; La Trobe
River, NMV; Lake Hattah, ANIC; Lilydale,
SAMA; Little Desert, ANIC; Lake Hattah, NMV;
Marysville, SAMA; Melbourne, UQIC; Preston,
NMV; Shepparton, ANIC; Skeleton Ck, SAMA;
Stawell, SAMA; Swan Hill, NMV; Traralgon Ck,
NMV; Violet Town, ANIC; East Warburton,
NMV; Wyperfeld Nt Pk, ANIC.

Western Australia

15 km SSE Armadale, ANIC; 102 km SE by E,
Broome, ANIC; 12 km NE Broome, ANIC;
Bunbury, ANIC; 23 km NW by W Mt Arid,
ANIC; 7 miles NE Giles, WAM; Gill Pinnacle,
WAM; Great Victoria Desert, WAM; Mary sp.
HS, ANIC; 13 km ESE Mooka H.S., WAM;
Koolan Island, WAM; Serpentine River, NMV;
West Peawah River, ANIC; Wilga, ANIC;
Wmgello, ANIC.

Remarks

Enochrus elongatus differs from all other
Australian Enochrus by the pale coloured dorsal
surface including the head. It is the only
Australian species with the tip of the aedeagus
distinctly hooked upwards and with a basal
swelling on the meta- and mesoclaws as well as
the proclaws.

The rear of the head is variably coloured. In
many specimens it is darker than the front of the

AUSTRALIAN ENOCHRUS

head, but diffusely so and never black and sharply
delineated from the front as in most other species.

Enochrus elongatus superficially resembles the
pale morphs of E. deserticola and E. eyrensis in
size and colour. It also shares with them, and no
other Australian species, the hooked parameres.
Unlike them the black colour morph common in
these two species does not appear to occur in E.
elongatus. In both E. deserticola and E. eyrensis
the head is predominantly black with only small
patches of yellow-brown on the front angles. In E.
elongatus the head is uniformly yellow-brown or
the rear is diffusely darker. In E. deserticola the
dorsal punctation is much weaker than in E.
elongatus and in E. eyrensis it is stronger.

Enochrus elongatus is a widespread and
common species within Australia including
Norfolk Island. I can find no significant difference
between Australian specimens and the types of E.
caledonicus thus extending the range of E.
elongatus to New Caledonia.

Enochrus (Methydrus) esuriens (Walker)

Philhydrus esuriens Walker, 1858

Enochrus (Lumetus) esuriens (Walker): Knisch
1924

Enochrus (Methydrus) esuriens (Walker):
d'Orchymont 1927

= Philhydrus pullus Fauvel, 1883: syn. nov.

= Enochrus (Lumetus) pullus (Fauvel): Knisch
1924

Types

Enochrus esuriens: Type locality: Ceylon (Sri
Lanka). Type not seen.

Enochrus pullus: Lectotype: 6 'Coll. IRSNB,
Nouvelle Caledonie, Anse Vata, Marais Juill
Octobre Rec. Saves, ex. coll. FauveF; 'Coll. et det
A. Fauvel Philhydrus pullus Fvl. IRSNB 17-497',
IRSNB with red lectotype label, herein
designated. Seen.

Paralectotypes: 3 6 , 1 9 7 same data as
lectotype in IRSNB.

I have not seen type material of E. esuriens and
have relied on the work of Balfour-Browne (1939,
1945) and specimens identified by him in
BM(NH) for my concept of this widespread
species which is known from India, through
South-East Asia to Northern Australia and
Polynesia.

Description (number examined, 130) Figs 13, 18
Length 1.6 mm- 2.3 mm. Oblong-oval. Elytron

and pronotum light-testaceous, often blotched and
streaked darker, head black, area in front of eye,
to about width of eye light-testaceous, central
black panel two to three times width of lateral
light area. Ventral surface dark-testaceous,
appendages lighter towards extremities, whole of
apical segment of maxillary palpi usually darker.
Punctures on head moderately strong, systematic
punctures inwards from eyes about size of eye
facet and about twice diameter of adjacent
punctures. Punctures on pronotum somewhat
weaker, punctures on elytra somewhat stronger
than on head, those towards apex and laterally
about half diameter of adjacent serial punctures
which are often not easy to trace. Mesosternal keel
weakly developed.

Male: Aedeagus narrow, pointed, virtually
lacking terminal dorsal pad, shorter than
parameres, collar nearer base than tip, parameres
broad in basal half narrow for most of apical
half, inner edge only very weakly sinuate, tips
rounded. Proclaw straightened, strongly swollen
at base, meso- and metaclaws rounded, swollen
at base.

Distribution

New South Wales

Wootton, ANIC.

Northern Territory

Adelaide River, ANIC; E Alligator River,
AM; Darwin, ANIC; Humpty Doo, QDPIM; 12
km N Humpty Doo, QDPIM; 6 km N Humpty
Doo, QDPIM; Jim Jim Creek, ANIC; 9 km N
by E Mudginbarry HS, ANIC; 19 km E by S Mt
Borradaile, ANIC; 12 km NNW Mt Cahill,
ANIC; Nourlangie Creek, ANIC; Oenpelli,
AM.

Queensland

Annan Falls, ANIC; Ayr, ANIC; 29 km S
Bamaga, ANIC; Cairns, ANIC, SAMA; Calliope
River, ANIC; Cardstone, ANIC; 60 km S Coen,
SAMA; 40 km N Coen, SAMA; Ingham, ANIC;
Lansdowne St, ANIC; 73 km NW Laura, ANIC;
Laura, QDPIM; Macleod River, SAMA;
Mackay, ANIC; Mareeba, QDPIM; Mary Creek,
ANIC; Mt Baird, ANIC; Mt Coolum, ANIC; Mt
Ivor River, ANIC; Mt Molloy, ANIC; 2 km NNE
Mt Tozer, ANIC; 1 1 km ENE Mt Tozer, ANIC;
8 km E by N of Mt Tozer, ANIC; 12 km S
Normanton, SAMA; North Pine River, UQIC;
Port Douglas, ANIC; Redcliff, UQIC; 15 km
WNW South Johnstone, QDPIM; Townsville,
ANIC, SAMA; Weipa, QDPIM.

148

C. H. S. WATTS

Western Australia

Mitchell Plateau, AN1C; Nullagine, WAM.

Remarks

Enochrus esuriens' small size (<2.5 mm),
predominantly black head and weak mesosternal
keel make it one of the most readily recognised
species of Australian Enochrus. It is most easily
confused, not with other Enochrus, but with
species of Paracymus and Anacaena, These latter
genera lack the systematic punctures present on
the head, pronotum and elytra of Enochrus.

Enochrus (Methydrus) eubenangeei sp. nov.

Types

Holotype: <J , 'Qld. Babinda Eubenargee
Swamp, 1/4/96, C. Watts', SAMA.

Paratypes: 6, 'Bramston Beach via Innisfail,
N.Q., 15 Aug. 1987, A. Walford-Huggins, coastal
melaleuca swamp at light', ANIC; 1, 'N.T.
Corndorl billabong nr. Jabiru, M.V. light,
2.x. 1982, MB. MalipatiF, NTM.

Description (number examined, 8) Fig 10

Length 7.0- 7.5 mm. Broadly oval, relatively
flat. Shiny. Black; palpi and tarsi testaceous.
Maxillary palpi moderately long, pseudo first
segment longer than distance from eye to front of
clypeus, longer than maxillary stipe, apical
segment approximately three-quarters length of
penultimate. Head strongly punctate, most
punctures about a puncture width apart, almost
size of eye facet. Systematic punctures large, about
3x size of adjacent punctures. Punctures on
pronotum well marked, similar to those on head,
not greatly weaker laterally, systematic punctures
well marked about 3x size of adjacent punctures,
lateral margins grooved and flanged, front and
rear margins weakly grooved. Elytral punctures as
on pronotum, not greatly weaker laterally. Serial
punctures in four loose rows, relatively well
marked, more numerous but more scattered
laterally, elytra weakly grooved and flanged
laterally. Mesosternal pillar well marked, sharply
triangular in front view, front edge sloping, ventral
edge short slightly pointed at rear, rear edge
vertical in top half. Notch on apical sternite
shallow, relatively broad.

Male: Aedeagus with very broad basal portion
and narrow apical portion which is curved
upwards. Parameres relatively broad, hooked
outwards at tips, bent upwards in apical half.
Proclaws bent, broadly thicker in basal third,

thickened part ends abruptly, meso- and
metaclaws curved, basal third somewhat
thickened, thickened part ending smoothly.

Remarks

Enochrus eubenangeei closely resembles E.
aliciae. Apart from the genitalia it differs from E.
aliciae in the lateral punctures on pronotum and
elytra being almost as strong as more central ones
and a greater extent of black. Based on the
relatively few specimens of both species that are
available E. eubenangeei also differs from E.
aliciae by its black head, whereas in many E.
aliciae the head has some testaceous colour, and
by the broader but narrower exposure of the
membrane between frons and clypeus than in E.
aliciae where it is shorter but deeper.

The aedeagi are distinct. Enochrus eubenangeei
has a normal narrow apical portion to the aedeagus,
without apical pad but bending strongly upwards.
Enochrus aliciae has a very broad basal portion
but, uniquely among Australian Enochrus, lacks
the apical portion, resulting in the aedeagus
reaching little beyond half way along the
parameres. The parameres are rather similar but the
tips are more obviously hooked in E. eubenangeei
and they bend noticeably upward in this species.

In its relatively large size, rounded shape and
shiny, black appearance, E. eubenangeei also
resembles E. mastersi. It, and E. aliciae, can be
most readily separated from this more common
species by the longer maxillary palpi with the
apical segment shorter than the penultimate. The
only other species it could be confused with is E.
isabellae which is smaller, narrower, has light
areas in front of its eyes and has the mesosternal
process long and blade-like.

Distribution

Known only from type localities in Northern
Territory and Queensland.

Enochrus (Methydrus) eyrensis (Blackburn)
Philhydrus eyrensis Blackburn, 1894

Enochrus (Lumetus) eyrensis (Blackburn):
Knisch 1924

=s Philhydrus andersoni Blackburn, 1896: syn. nov.

= Enochrus (Lumetus) andersoni (Blackburn):
Knisch 1924

?= Philhydrus persimilis Rcgimbart, 1908: syn.
nov.

?= Enochrus (Lumetus) persimilis (Regimbart):
Knisch 1924

AUSTRALIAN ENOCHRUS

149

FIGURES 15-20. Mesosterna. 15-18, Lateral view of
mesostcrnal keel. 15, E. samae; 16, E. eyrensis; 17, E.
maculiceps; 18, E. esuriens. 19 & 20, mesostcrnum. 19,
E. mastersi (after Hansen 1990); 20, E. aliciae.

Types

Enochrus eyrensis: Holotype: 9, 'Blackburn
coll. 1910-236', 'Philhydrus eyrensis Black 1 ,
remounted with '178T' on original Blackburn
card, BM(NH),

Paratypes: 2 'Philhydrus eyrensis Blackb co-

type', 'Philhydrus eyrensis Blackb, C. Australia
co-type 8242', SAM A.

Enochrus andersoni: Holotype: ?sex.
'Blackburn coll. 1910-236% 'Philhydrus
andersoni, Blackb' remounted with '178T on
original Blackburn card, BM(NH).

Synonymy based on examination of types.

Enochrus persimilis: Holotype: ?Victoria Aust;
' Philhydrus persimilis Reg.' MNHN. Synonymy
based on descriptions and examination of types
(of E. persimilis in 1964, type not currently
available). It is possible that my E. samae will
prove to be this species.

Description (number examined, 146) Figs 5, 16

Oblong-oval, length, 3.2 mm - 5.0 mm' Two
colour morphs exist, one with elytra and pronotum
testaceous often with sutural lines and strial
punctures darker, occasionally with disc of
pronotum darker, head dark-testaceous to black
with lighter area forward from eyes to about the
same width as eyes giving a darker central panel
three to four times the width of one of the lighter
patches, clypeus dark. Second colour morph has
the elytra dark-testaceous or black, pronotum a

C. H. S. WATTS

similar colour but quite widely but diffusely
lighter laterally, head as for first colour morph.
Underside of both morphs dark-testaceous,
appendages lighter towards extremities, except for
maxillary palpi which have apical portion of
apical segment darker.

Head quite densely covered with relatively
strong punctures about half size of eye facet, a few
large punctures inwards from eyes, 2— 3x diameter
of surrounding punctures. Punctures on pronotum
slightly stronger, systematic punctures as in E.
elongatus, 2-3x diameter of adjacent punctures.
Punctures on elytra rather stronger than on
pronotum, towards apex approaching size of serial
punctures, which, as a consequence, are hard to
trace.

Mesosternal keel well developed, anterior edge
weakly curved, nearly perpendicular, ventral edge
straight or weakly convex, narrow vertical 'blade'
above an extensive broader base. Notch on apical
sternite moderately developed.

Male: Aedeagus narrow, pointed, virtually
lacking terminal dorsal pad, collar nearer base
than tip. Parameres outwardly hooked at tip.
Proclaw a little longer, less curved than in female,
otherwise little sexual difference in claws.

Distribution

Australian Capital Territory

Black Mt, ANIC; Jarvis Bay, ANIC.

New South Wales

Bonville, ANIC; 25 km N Bulahdelah, ANIC;
Maroota South, ANIC; 8 km SE by S Moruya,
ANIC; Norfolk Island, ANIC; Queanbeyan,
ANIC; 17 mis S Woodburn, ANIC; Wootton,
ANIC.

South Australia

Adelaide, SAMA; American River, SAMA;
Beachport, AM, SAMA; Bool Lagoon, SAMA;
Coonalpyn, SAMA; Fairview Park Cons Pk,
SAMA; Hindmash Park, SAMA; Lucindale,
SAMA; Naracoorte, UQIC; Taratap Stn, SAMA.

Tasmania

Scamanden River, NMV; Swansea, SAMA.

Victoria

East Pomborneit, ANIC; Lome, SAMA;
Maffra, UQIC; Otway Rngs, UQIC; Wyperfeld
Nt Pk, ANIC.

Western Australia

Albany, ANIC; Applecross, ANIC; Broomehill,

WAM; Dumbleyung, WAM; 5 km NE Esperance,
ANIC; 63 mis E Esperance, ANIC; 9 mis NE by
N Esperance, ANIC; 19 km E Esperance, ANIC;
5 km NW Esperance, ANIC; Fitzgerald River Nt
Pk, ANIC; Karridale, ANIC; Maidavale, SAMA;
Mt Margaret, ANIC; Mandurah, ANIC; 23 km
NW by N Mt Arid, ANIC; Perth, ANIC; 8 mis E
Pinjarra, ANIC; Stirling Rngs, SAMA; 12 km NE
Wanneroo, WAM; 155 km SW Warburton,
WAM; Wilga, ANIC.

Remarks

This relatively large species is common in both
the south-east and south-west.

The relatively strong dorsal punctation separates
it from all other Australian Enochnis except E.
samae and the generally much smaller E.
malabarensis. Enochrus malabarensis is also a
more northern species and lacks black morphs.
From E, samae, E. eyrensis differs in having the
parameres hooked at the tips rather than rounded
and in a higher, more blade-like mesosternal keel.
This latter character is, I believe, a good one but is
hard to see on many specimens and is difficult to
use if comparative material is not available.

Enochrus eyrensis appears close to E, tristus
(Broun) from New Zealand and some Pacific
Islands in the hooked parameres and strong
punctations. It differs from this species in having
the apical portion of the aedeagus shorter than the
basal and in rounded, not slightly straightened,
male metaclaws. Enochrus eyrensis differs from
E. abditus (Sharp) from New Zealand, in details
of the aedeagus and mesosternal keel.

Enochrus (Methydrus) isabellae sp. nov.

Types

Holotype: 6\ right-hand specimen on card,
'Qld. Townsville, 10 km NW, 23/3/96, C. Watts',
SAMA.

Paratypes: 7, same data as holotype, SAMA; 3,
'Ayr Qld., 10.ix.1990, W.B. Muir', in ANIC; 1,
'Ingham, N.Q., 27.L68, J.C. Brooks', ANIC; 1,
'Blighs Lookout, Cardwell Rng, N.Q. at light,
30.ix.67, J. Brooks', ANIC; 1, 'Bramston Beach
via Innisfail, N.Q., 1 5. viii. 1 987, A. Walford-
Huggins. Coastal mclaleuca swamp at light', in
ANIC; 6, 'Qld. Nardello's Lagoon nr. Mareeba
29/3/96, C. Watts', SAMA.

Description (number examined, 14) Fig 6

Length 4.5 mm - 5.5 mm. Oval, shiny. Black
apart from lateral areas of pronotum, region in

AUSTRALIAN ENOCHRUS

front of eyes and appendages which are reddish-
yellow. Tips of maxillary palpi darker. Head quite
densely punctate, with weak to moderately sized
punctures, systematic punctures rather sparse,
somewhat larger than an eye facet and 2-3x size
of adjacent punctures. Pronotum and elytra
similarly punctured except that systematic
punctures a bit larger in comparison with others.
Mesosternal keel well-developed, narrow, often
with small protuberance on anterior end.

Male: Claws on protarsi more bent than in
female with broad basal portion. Parameres
hooked apically, aedeagus with apical portion
shorter than basal, relatively broad, narrowing
rapidly at tip, slight but noticeable dorsal
thickening at tip.

Distribution

Known only from type localities in Queensland.

Remarks

A relatively large nearly totally black species. It
strongly resembles the black morphs of E.
eyrensis and E. samae but is much more weakly
punctured, has a shorter apical portion of the
aedeagus than in E. eyrensis and has hooked
parameres which distinguish it from E. samae.
Compared to the black morph of E. deserticola
which occurs sympatrically with it, it is larger,
more strongly punctured and has more extensive
black on the front of the head than is normal in E.
deserticola. The male genitalia are relatively
similar to those of E. deserticola but E. isabellae
has a broader and shorter apical portion to the
aedeagus with a more developed apical dorsal/
ventral thickening. From E. elongatus it can be
separated on colour, generally weaker punctation,
weaker apical hook on aedeagus which ends well
short of tips of parameres and lack of modified
meso- and meta-claws in the males.

The Townsville and Nardello's Lagoon
specimens were collected from shallow water
among thick drying reeds at the edge of extensive
shallow swamps.

Enochrus (Methydrus) maculiceps (MacLeay)

Philhydrus maculiceps MacLeay, 1873

Enochrus (Lumetus) maculiceps (MacLeay);
Knisch 1924

= Philhydrus laevigatus Blackburn, 1888: syn. nov.

- Enochrus (Lumetus) laevigatus (Blackburn):
Knisch 1924

= Philhydrus artensis Fauvel, 1883: syn. nov.

= Enochrus (Lumetus) artensis (Fauvel): Knisch
1924

- Enochrus (Lumetus) bryani d'Orchymont,
1927: syn. nov.

Types

Enochrus laevigatus: Holotype: 'Philhydrus
laevigatus Blackb', BM(NH), 'Blackburn coll.
1910-236'. Remounted with M552T' on original
Blackburn label. Synonymy based on examination
of types.

Enochrus maculiceps: Lectotype: Male.
'Philhydrus maculiceps McL W. Gayndah',
'K19505'. Right-hand specimen of two mounted
on same card, AM, herein designated.

Para lee totypes: 1, on same card as lectotype,
AM; 2, same data as lectotype and mounted on
one card, in ANIC; "Philhydrus maculiceps McL.
'Co-type' Griffith collection id by A.M. Lea" ; 1,
'Philhydrus maculiceps McL. Queensland co-type
14631', SAM A. Synonymy based on examination
of types.

Enochrus artensis: Holotype: 'Nouvelle
Caledonie He Art*, 'Rec. ex. coll. Fauvel', 'Coll.
et det. A. Fauvel Philhydrus artensis Fvl\ with
red Holotype label, IRSNB.

Paratype: 9 , 'Nouvelle Caledonie Kanata Re
Deplenche ex. coll. Fauvel'; 'Coll. et det. A.
Fauvel Philhydrus artensis Fvl, IRSNB 17-479',
with orange paratype label, IRSNB. Synonymy
based on examination of types.

Enochrus bryani: Holotype: 'Savaii Samoa',
'Salailua v-22-24', 'H. Bryan Jr, Collector', 6,
'Type', 'A. d'Orchymont det, Enochrus (Lumetus)
bryani\ Bishop Museum, Honolulu.

Paratype: 'Enochrus (Lumetus) bryani A.
d'Orchymont det'; 'Samoan Is. Apia Upolu Is';
'female' T Samoa Mars 1924 Coll. d'Orchymont',
IRSNB.

Both the types are mounted on card from the
same source and similarly labelled by
d'Orchymont. d'Orchymont clearly identified the
Bishop Museum specimen as the holotype.
Unfortunately all but a hind tibia and tarsus of the
holotype has been lost. I base my concept of E.
bryani on the Brussels paratype and
d'Orchymont' s description and illustration of the
aedeagus.

Description (number examined, 134) Figs 1,2, 17
Length 2.6 mm 4.0 mm. Elytra testaceous,
sutural lines, some serial punctures and a rough,
small patch on each humeral angle darker;
pronotum testaceous, disk rather darker than
sides; head with testaceous areas in front of each

152

C. H. S. WATTS

eye much wider than eye, central dark panel about
half width of lateral light patches, boundary
between areas often diffuse. Ventral surface dark-
testaceous, appendages only slightly lighter
towards extremities, maxillary palpi with tip of
apical segment darker. Punctures on head
moderately sized, a few larger punctures inwards
from eyes a little smaller than eye facets and about
4x diameter of adjacent punctures. Pronotum
similarly punctate, systematic punctures relatively
weak, smaller than eye facets. Interstrial punctures
on elytra weak, verging on subobsolcte, serial
punctures also relatively small and weak, both a
little stronger apically and laterally, on humeral
angle the serial punctures are about 3x the
diameter of adjacent punctures but both are small
and relatively weak. Mesosternal keel moderately
developed, ventral edge flat except for a well
developed anterior protuberance.

Male: Genitalia variable in elongation,
aedeagus usually relatively broad, narrowing to
rounded point, collar closer to base than tip.
Parameres relatively narrow, pointed. Male and
female claws similar.

Distribution

Australian Capital Territory

Blundello Creek, ANIC; Canberra, SAMA.

New South Wales

Albury, ANIC; Ashfield, AM; Balranald,
NMV; Berry, SAMA; Bogan River, AM; 20 mis
SSW Bourke, SAMA; Collector, SAMA; Congo,
ANIC; 9 km NNE Coonabarabran, ANIC; 14 km
W Coonabarabran, ANIC; Dc Burghs Rng, AM
Deniliquin, ANIC; 37 km E Hay, SAMA
Illadulla, ANIC; Jenolan Caves, SAMA
Kenilworth, ANIC; North Sydney, NMV
Tooloom Plateau, UQIC; Wallacia, SAMA
Wilcannia, SAMA; 3 km S Wingellow, ANIC
Wingham Scrub, ANIC; Wootton, ANIC; Valery,
ANIC.

Northern Territory

30 mis W Alice Springs, ANIC; Stanley
Chasm, SAMA; 100 mis W Mt Olga, WAM; 30
km N Wauchopc, ANIC; Yuendumu, SAMA.

Queensland

31 km NE Aramac, SAMA; Atherton, QDPIM;
Babinda, UQIC; Brisbane, UQIC, SAMA;
Bundaberg, ANIC; Byfield, ANIC; Cardstone,
ANIC; Clermont, AM; Edungalba, ANIC; Funnel
Creek, ANIC; 50 mis W Mackay, ANIC; Mt
Coolum, ANIC; 9 mis S Stanthorpe, ANIC;

South Australia

Arkaba Creek, SAMA; 80 mis S Cooper Pedy,
SAMA; 10 km SE Coward Springs, SAMA;
Murray River, SAMA; Wintana Station, SAMA.

Victoria

Healsville, SAMA; Maffra, UQIC; Nathatia,
SAMA; Portland, SAMA; 13 km SE Shepperton,
ANIC; 14 km NW Violet Town, ANIC.

Western Australia

Nr Mt Gibson, WAM; 3 km E by N Newman,
ANIC.

Remarks

A widespread and common species resembling
a small E. deserticola and difficult to separate
from light morphs of this species without
reference to the male genitalia. Somewhat more
strongly punctured than in most E. deserticola. In
the North-cast, E. maculiceps is sympatric with E.
weiri and E. pseudoweiri which it closely
resembles in general form. Both these species are
a little more strongly punctured and have pale tips
to their maxillary palpi and usually only a small
central triangular area of black on the front of the
head. However E, maculiceps with pale palpi and
the central black portion on the front of the head
reduced to a triangle, are not uncommon.

The male genitalia are variable in elongation,
from a squatter form with the aedeagus quite
curved in lateral view, with a strong apical pad
and with only a small amount of structure apical
to the pad, through to a more elongate form with
elongate parameres, nearly straight aedeagus with
a weak apical pad and a quite pronounced broadly
triangular portion apical to the pad.

Some more strongly punctured specimens of E.
maculiceps may approach weaker punctured
specimens of E. malabarensis. In E. malabarensis
the front of the head is nearly all black whereas in
E. maculiceps the black area is seldom more than
a third the width of the front of the head.

I have compared Australian specimens with the
holotype, a paratype and five other specimens of
E. artensis Fauvcl from New Caledonia in IRSNB
including two dissected males and consider that
E. artensis is a junior synonym of E. maculatus.

Enochrus (Methydrus) malabarensis

(Regimbart)

Philhydrus malabarensis Regimbart, 1903

Enochrus (Lumetus) malabarensis (Regimbart):
Kmsch, 1924

AUSTRALIAN ENOCHRUS

153

Types

Enochras malabarensis: Holotype. 'Make,
Calicut Ind'; 'Philhydrus ma lobar ens is Reg',
MNHN.

Description (number examined, 37) Fig. 12

Length 2.5 mm - 4.1 mm. Broadly oval.
Elytron and pronotum coloured as for E.
pseudoweiri. Head black, areas in front of eyes
light testaceous to about width of eye, central
black panel 3-4x width of lateral light area.
Ventral surface as for E. pseudoweiri, tips of
maxillary palpi sometimes a little darker than
rest of palpi. Punctures on head relatively strong,
often approaching the size of eye facet,
systematic punctures inwards from eye a little
larger, sometimes hard to trace. Pronotum
moderately to strongly punctured, systematic
punctures often hard to trace. Punctures on elytra
somewhat stronger towards apex and sides,
approaching size of serial punctures which, as a
consequence, may be difficult to distinguish.
Mesosternal keel moderately developed,
pronounced protuberance anteriorly, composed
almost entirely of broad basal portion.

Male: Aedcagus relatively short, narrow,
pointed with only slight apical pad, collar
equidistant from tip and base. Parameres broad in
basal half, narrow in apical quarter, inner edges
almost straight, tips rounded. Proclaw rounded or
weakly straightened strongly swollen at base,
meso- and metaclaws, rounded.

Distribution

New South Wales

Ashfield, AM; Bondi Heights, AM; Sydney,
NMV, SAMA.

Northern Territory

Black Point Coburg Peninsula, ANIC; Coastal
Plains Research Stn, ANIC; Crocodile Island,
SAMA; Darwin, ANIC, SAMA; Groote Eyland,
SAMA; Howard Springs, ANIC; Kakadu NP,
ANIC.

Queensland

Bowen, SAMA; Brisbane, UQIC; Bundaberg,
ANIC: Stewart River, SAMA.

Enochrus. The punctation is strongest in
specimens from the Darwin area where the
systematic punctures on pronotum and elytra are
virtually untraceable. The punctures get weaker to
the south were some specimens can approach the
situation in the most strongly punctate E.
maculiceps, E. pseudoweiri and E. weiri. The
greater extent of black on the front of the head
(the pale area reduced to the width of an eye) will
separate E. malabarensis from these species
where the black portion covers <l/3 of the head
width. Enochrus malabarensis has the unusual
combination of strongly black head with pale tips
to the maxillary palpi.

The male genitalia of E. malabarensis differ
from those of E. maculiceps in that the inner
margin of the parameres are straight rather than
sinuate to accommodate the apical pad on the
aedeagus. Although the tip of the aedeagus in E,
malabarensis is thickened dorsoventrally, it is not
laterally expanded (to varying degrees) as in E.
maculiceps.

The mesosternal keel is stronger than in E.
weiri and E. pseudoweiri and most specimens of
these species lack the pronounced anterior
downward protuberance found in both E.
malabarensis and E. maculiceps (Fig. 17). The
genitalia of E. weiri is distinctive, that of E.
pseudoweiri quite closely resembles E.
malabarensis but has a shorter and thinner
aedeagus.

Enochrus malabarensis differs from the two
other strongly punctate Australian Enochrus (E.
eyrensis and E. samae) by its smaller size, lack of
black colour morphs, usually pale maxillary palpi,
stronger punctation particularly on the head and
the strong downward protuberance at the front of
the mesosternal keel which is usually lacking in
E, eyrensis and E. samae. Enochrus malabarensis
has a generally more northern distribution than
the other two species but overlaps with E.
eyrensis on the east coast. The male genitalia
readily separate the species (Figs 5, 12).

I have not seen the type of E. malabarensis (it
is unavailable at this time) and have based my
concept of this species on specimens identified by
d'Orchymont from Sulawesi in IRSNB.

South Australia

Billa Kalina Station, SAMA.

Remarks

The moderate size and strong dorsal punctation
separate E. malabarensis from all other Australian

Enochrus (Methydrus) pseudoweiri sp. nov.

Types

Holotype: S, '16°03'S to 16°05'S, 145°28'E,
Cape Tribulation area, Qld., 21-28 Mar 1984, A.
Calder and T. Weir\ ANIC.

154

C. H. S. WATTS

Paratypes: All male. 4, "Cardwell Rng, N.Q.,
30.9.87, G.B. 'Q363'; 'J. G. Brooks Bequest
1976', ANIC; 2, '15°03'S, 145°09*E, 3 km NE of
Mt Webb, Qld., 1-3 Oct. 1980, T. Weir', ANIC;

1, 'Whitfield Rg., Rd., ca. 486 m, Qld., 3.ii.l970,
at light, J. G. and J. A. G. Brooks', ANIC; 1,
<12°44S, 143°17'E, 8 km E by N of Mt Tozer,
Qld., 7 July 1986, T. Weir and A. Calder', ANIC;

2, Mclvor Rv., 25 mis N Cooktown, N Qld, 6
May 1970, S. R. Curtis', one ANIC, one SAMA;
1, 'Australia, N Qld, Iron Range, 26-31. x.1991,
Wood, Dunn and Hasenpusch', QDPIM; 1, 'Cow
Bay, N of Daintree N Qld, 20-1 7-11 1984 1, C.
Cunningham', QDPIM; 1, 'Cow Bay, N of
Daintree, Qld., 27/7/82, C. Watts', SAMA; 4,
'Peach Ck, N Qld., 24/7/87, C. Watts', SAMA; 1,
'Cairns, C. J. Wild, Jan 91', QM.

ANIC; Cape Tribulation area, ANIC; 3km S by
W of Cooktown, ANIC; 25mls N Cooktown,
ANIC, SAMA; Cooktown, ANIC; 2mls N
Kuranda, ANIC; 8km E by N of Mount Tozer,
ANIC; 3km NE of Mount Webb, ANIC; Peack
Ck, SAMA; 15km WSW South Johnstone,
QDPIM; Whitfield Rg Rd, ANIC.

Northern Territory

Howard Springs, ANIC.

Remarks

Apart from the male genitalia and a tendency
for the darker portion at the front of the head to be
larger and more diffuse, I can find no difference
between this species and E. weiri (see under E,
weiri).

Description (number examined, 33) Fig. 3

Length 3.0 mm - 3.6 mm. Oblong oval. Elytra
testaceous to dark-testaceous, sutural region and
serial punctures darker; small darker patch on
humeral angles in many specimens. Pronotum
dark testaceous centrally, widely lighter laterally.
Head dark testaceous-black, frons area light
testaceous except for small triangularly shaped
dark area with apex often not reaching front of
head, boundary between darker area and light
areas indistinct, clypeus dark testaceous, ventral
surface dark testaceous, appendages somewhat
lighter towards extremities, maxillary palpi
lacking darker tip. Head sparsely covered by
small weak punctures, much smaller than facets
of eyes, a few larger punctures inward from eyes
about size of eye facet and 2 3x larger than
adjacent punctures. Elytral punctures a little
stronger, shallow. Serial punctures relatively
weak, hard to trace laterally except on humeral
angles where they are more distinct and 3-5x the
diameter of adjacent punctures. Mesosternal keel
strongly raised, ventral edge weakly convex,
usually with a distinct anterior protuberance,
broader basal portion making up most of keel
with only a narrow thinner section in lateral
view.

Male: Genitalia very stout to moderately stout
overall. Aedeagus very slender, lacking terminal
pad, collar nearer base than tip. Parameres broad,
tips rounded. Proclaws curved, strongly swollen
in basal half, more so than in female.

Distribution

Queensland

Cairns, ANIC; Cardstonc, ANIC; Cardwell,

Enochrus (Methydrus) samae sp. no v.

Types

Holotype: S, 'East Pombomeit, Vic. 24 km
ESE Camperdown. Temporary pond, Aug. 1978-
Feb. 1979, P.S. Lake coll.', ANIC.

Paratypes: 2 6, same data as holotype,
ANIC; 2 2, '29.01°S, 167.57°E, Norfolk
Island, Filmy Fern Walk, 14 Nov. - 2 Dec.
1984, I.D. Naumann ex ethanoP, ANIC; 4 9,4
6, 'Stonor, Tas: Lea', SAMA; 'Swansea, Tas.,
Jan. C. Watts', SAMA; 1 2, 1 S, M0 km S
Robe S.A., 1/83, C. Watts', SAMA; 2 & 9
'Grampians, Vic, 2.63, C. Watts', SAMA; 1
6, 'Chain of Ponds, S.A., December 1957, C.
Watts', SAMA; 1 6, 'Mylor, S.A., Aug. 1958,
C. Watts', SAMA.

Description (number examined, 47) Figs 9, 15

Length 4.0 mm - 5.0 mm. Oblong-oval. Elytra
and pronotum dark-tcstaccous, quite broadly
lighter at sides, dark area broken up with
longitudinal rows of spots towards apex, front of
pronotum often narrowly light-testaceous. Head
dark-testaceous to black except for broad light-
testaceous areas forward from eyes, leaving a
central dark panel which is about the same width
as one of the pale patches, boundary between the
dark central panel and pale patches diffuse.
Underside dark-testaceous, appendages only
slighter lighter towards extremities, apical portion
of apical segment of maxillary palpi darker.
Punctures on head relatively strong, not much
smaller than eye facet, a few larger punctures
inward from eyes, about 3x the diameter o£
adjacent punctures; pronotal punctures on elytra a

AUSTRALIAN ENOCHRUS

155

little stronger than on pronotum, towards apex and
laterally approaching size of strial punctures
which are consequently not easily traced.
Mesosternal keel moderately developed, triangular
shaped, broader basal portion makes up most of
keel.

Male: Aedeagus broad, well-marked terminal
dorsal pad, collar nearer to base than to tip.
Parameres with rounded tips. Male proclaws
greatly swollen in basal half, strongly curved in
apical half, female less swollen and more evenly
curved.

Distribution

Australian Capital Territory

Black Mt., ANIC.

New South Wales

Norfolk Isl., ANIC.

South Australia

Chain of Ponds, SAM A; Mylor, SAMA; 10 km
S Robe, SAMA.

Tasmania

George Town, SAMA; Stonor, SAMA;
Swansea, SAMA.

Victoria

Grampians, SAMA; Hamilton, NMV; Lome,
SAMA; Otway Rngs., UQIC; East Pombomcit,
ANIC.

Remarks

In general facies this species is very similar to
E. eyrensis and at least in the South-east occurs
sympatrically with it. The strong dorsal punctation
and patterned head readily separates these two
species from other Enochrus in the region.
Enochrus samae differs from E. eyrensis in
having rounded rather than hooked tips to the
parameres and in a squatter mesosternal keel. This
latter character is hard to use if comparative
material is not available

Enochrus (Methydrus) weiri sp. nov.

Types

' Holotype: o\ M2°44'S, I43°14'E, 3 km ENE of
Mt Tozer, Qld., 28 Jun - 4 Jul 1986, T. Weir and
A. Calder', ANIC.

Paratypes: All S. I, '12°44'S, 143°17*E, 8 km
E by N of Mt Tozer, Qld., 7 July 1986, T. Weir
and A. Calder', ANIC; I, '12°44'S, 143°14'E, 3

km ENE of Mt Tozer, Qld., 28 Jun - 4 Jul 1986,
T. Weir and A. Calder', ANIC; 2, M2°43'S,
143°17'E, 9 km ENE of Mt Tozer, Qld., 5-10 July
1986, T. Weir and A. Calder', one ANIC, one
SAMA; 1, 'Dejinghe Ck, N.Q., 3 mis SW of
Yarrabah, 26.x. 1966 E. Bntton, R Trumble',
ANIC; 1, M6°03'S to 16°05'S, 145°28'E Cape
Tribulation area, Qld., 21-28 Mar 1984, A.
Calder and T. Weir\ ANIC; 1, 'Australia, N Qld.,
Cow Bay N of Daintree R, 13. ix. 1990,
Cunningham and DeFaveri', QDPIM; 1,
'Australia, N Qld Cow Bay N of Daintree R,
14.xii. 1987-6.i. 1988, Storey and Cunningham',
QDPIM.

Description (number examined, 1 8 with male
genitalia extracted) Fig. 8

As for E. pseudoweiri except for male genitalia.

Male: Aedeagus broadest in middle, tapering to
a point, lacking any terminal dorsal pad. Tips of
parameres truncated, weakly bifid.

Distribution

Known only from type localities on Cape York
in North Queensland.

Remarks

A tropical species known only from the
Queensland east coast, north of Cairns. I have
found no reliable way to separate this species
and E. pseudoweiri, with which it is sympatric
over most of its range, other than by the
distinctively different male genitalia. These
species can also be readily confused with E.
maculiceps and with small brown colour morphs
of E, deserticola which also occur in the region.
The virtual absence of black on the front of the
head and pale tips to the maxillary palpi will
separate them from most examples of these
species. Most E. weiri lack or have only a small
downward protuberance at the front of the
mesosternal keel whereas most E. maculiceps
have it well developed. The differently coloured
head and weaker dorsal punctures will readily
separate E, weiri from the similarly sized E.
malabarensis.

Acknowledgments

I thank the curators of the collections listed earlier for
allowing me ready access to specimens in their care: Ms
D. Churches typed the manuscript, Mr R. Gutteridge
prepared the figures, Mrs M. Anthony and Mrs J. Evans
helped with the library references and Dr E. Matthews
helped improve the manuscript.

156

C. H. S. WATTS

References

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(Coleoptera). The biology of some Australian species
with descriptions of immature stages reared in the
laboratory. Journal of the Australian Entomological
Society 15: 219-228.

BLACKBURN, T. 1888. Notes on Australian Coleoptera
with descriptions of new species. Proceedings of the
Linnean Society of New South Wales 3: 805-875.

BLACKBURN, T. 1894. Further notes on Australian
Coleoptera, with descriptions of new genera and new
species. Transactions of the Royal Society of South
Australia 18: 139-168.

BLACKBURN, T. 1896. Coleoptera (exclusive of the
Carabidae). Pp. 254^-308 in B. Spencer (ed.) 'Report
on the work of the Horn Expedition to Central
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Slade: Melbourne.

FAUVEL, A. 1883. Les Coleopteres de la Nouvelle-
Caledonie et dependances avec descriptions, notes et
synonymies nouvelles. Revue d'Entomologie, Societe
Francaise d'Entomologie 2: 335-360.

GENTILI, E. 1981. The genera Laccobius and
Nothydrus (Coleoptera: Hydrophilidae) in Australia
and New Zealand. Records of the South Australian
Museum 18(7): 143-154.

HANSEN, M. 1990. Hydatotrephis MacLeay, a
subgenus of Enochrus Thomson (Coleoptera:
Hydrophilidae). Entomologica Scandinavica 21(1):
71-76.

HANSEN, M. 1991. The Hydrophiloid Beetles.

Phylogeny, Classification and a Revision of the
Genera (Coleoptera: Hydrophilidae). Pp. 348.
Biologiske Skriften 40. The Royal Danish Academy
of Science and letters, Kommissionaer: Munksgaard,
Copenhagen.

KNISCH, A. 1922. Results of Dr E. Mjoberg's Swedish
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KNISCH, A. 1922. Hydrophiliden-Studien (op. 10).
Archive fur Naturgeschichte 88: 87-126.

KNISCH, A. 1924. Hydrophilidae. Pp 306 in W. Junk &
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pars 79'. Berlin.

MacLEAY, W. 1873. Notes on a collection of Insects
from Gayndah. Transactions of the Entomological
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d'ORCHYMONT, A. 1927. Papers on Malayan Aquatic
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REGIMBART, M. 1903. Voyage de M. Maurice
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RfeGIMBART, M. 1908. Dytiscidae, Hydrophilidae et
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WALKER, F. 1858. Characters of some apparently
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Natural History 2(series 3): 202-209.

SOUTH

AUSTRALIAN

MUSEUM

VOLUME 30 PART 2
JANUARY 1998
ISSN 0376-2750

CONTENTS

ARTICLES

63 P. E. BOCK& P. L. COOK.

A new species of multiphased Corbulipora MacGillivray, 1895 (Bryozoa; Cribriomorpha)
from southwestern Australia.

69 E. WILLIAMS

The archaeology of lake systems in the middle Cooper Basin, north-eastern South
Australia.

93 C. H. S. WATTS

Revision of Australian Amphiops Erichson, Allocotocerus Kraatz and Regimbartia
Zaitzev (Coleoptera: Hydrophilidae).

107 D. W. CORBETT

Douglas Mawson: the geologist as explorer.

137 C. H. S. WATTS

Revision of Australian Enochrus Thomson (Coleoptera: Hydrophilidae).

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