CONTENTS

BAEHR, M.

A new species of the genus Lestignathus Erichson from Tasmania, with a

note on the Tasmanian species of Mecyclothorax Sharp (Insecta: Coleoptera:

Carabidae: Licininae, Psydrinae).

CORBETT, D. W.

A staunch but testing friendship: Douglas Mawson and T.W. Edgeworth David.

GENTILLI, E.

The Paracymus of Australia (Coleoptera, Hydrophilidae).

JONES, P. G.

Words to objects: origins of ethnography in colonial South Australia.

KRISTENSEN, R. M. & MACKNESS, B. S.

First record of the marine tardigrade genus Batillipes (Arthrotardigrada:

Batillipedidae) from South Australia with a description of a new species.

NELSON, R. C.

Why Flutes on Boomerangs and Throwing Sticks?

PRIDEAUX, G. J.

Simosthenurus newtonae sp. nov., a widespread sthenurine kangaroo

(Diprotodontia: Macropodidae) from the Pleistocene of southern

and eastern Australia.

WATTS, C. H. S.

Revision of Australian Chaetarthria Stephens (Coleoptera: Hydrophilidae)..

WATTS, C. H. S.

Three new species of Tiporus Watts (Coleoptera; Dytiscidae) with redescriptions

of the other species in the genus.

WATTS, C. H. 8. & HUMPHREYS, W. F.

Six new species of Nirridessus Watts and Humphreys and Tjirtudessus Watts

and Humphreys (Coleoptera: Dytiscidae) from underground waters in Australia.

WATTS, C. H. S. & PINDER, A.

Two new species of Antiporus Sharp from Western Australia (Coleoptera: Dytiscidae).

ZEIDLER, W.

Note on the origin of freshwater crayfish occurring on Kangaroo Island, South Australia

Volume 33(1) was published on 21 August 2000.

Volume 33(2) was published on 20 October 2000.

ISSN 0376-2750

PAGES

123-126

49-70

101-122

33-47

73-87

21-27

1-15

29-31

89-99

127-144

17-19

71-72

SIMOSTHENURUS NEWTONAE SP. NOV., A WIDESPREAD STHENURINE

KANGAROO (DIPROTODONTIA: MACROPODIDAE) FROM THE

PLEISTOCENE OF SOUTHERN AND EASTERN AUSTRALIA.

BY GAVIN J. PRIDEAUX

Summary

PRIDEAUX, G.J. (2000).Simosthenurus newtonaie sp. nov. is described from the Pleistocene of

southern and eastern Australia. Its cranium is similar in size to Simosthenurus occidentalis, but is

less brachycephalic and has narrower, more elongate rostrum with a less inflated frontal region.

The moderately high-crowned molars are distinctive among the species of Simosthenurus, because

they bear very few to no fine enamel crenulations, and primary crests that bear strong contacts with

cusp apices. In these respects, the molars resemble Sthenurus andersoni and Hadronomas

puckridgi.

SIMOSTHENURUS NEWTONAE SP. NOV., A WIDESPREAD STHENURINE KANGAROO

(DIPROTODONTIA: MACROPODIDAE) FROM THE PLEISTOCENE

OF SOUTHERN AND EASTERN AUSTRALIA

GAVIN J. PRIDEAUX

PRIDEAUX, G. J. 2000. Simosthenurus newtonae sp. nov., a widespread sthenurine kangaroo

(Diprotodontia: Macropodidae) from the Pleistocene of southern and eastern Australia. Records

of the South Australian Museum 33(1): 1-15.

Simosthenurus newtonae sp. nov. is described from the Pleistocene of southern and eastern

Australia. Its cranium is similar in size to Simosthenurus occidentalis, but is less brachycephalic

and has a narrower, more elongate rostrum with a less inflated frontal region. The moderately

high-crowned molars are distinctive among the species of Simosthenurus, because they bear very

few to no fine enamel crenulations, and primary crests that bear strong contacts with cusp apices.

In these respects, the molars resemble Sthenurus andersoni and Hadronomas puckridgi.

Gavin J. Prideaux, School of Biological Sciences, Flinders University of South Australia, GPO

Box 2100, Adelaide South Australia 5001, Australia. Current address: Department of Earth

Sciences, University of California,

Riverside,

CA 92521, USA. Email:

gavin.prideaux @ucr.edu. Manuscript received 11 November 1999.

Sthenurines were a diverse group of robust

browsing kangaroos common throughout

southern and eastern Australia in the Pleistocene.

Simosthenurus newtonae sp. nov. represents the

seventh new sthenurine described since 1992.

Although uncommon in most assemblages, it is

one of the more widely distributed sthenurines,

occurring in 21 localities (Fig. 1). The

description of this new species forms the subject

of this paper. A review of the phylogenetic

relationships, chronology, zoogeography and

evolution of the sthenurine kangaroos is

currently in preparation.

MATERIALS AND METHODS

Specimens referable to Simosthenurus

newtonae sp. nov. are housed in the vertebrate

palaeontological collections of the following

institutions: Australian Museum, Sydney (AM);

South Australian Museum, Adelaide (SAM); Field

Museum of Natural History, Chicago (FMNH);

Flinders University of South Australia, Adelaide

(FU); Museum of Victoria, Melbourne (NMV);

Queensland Museum, Brisbane (QM); Queen

Victoria Museum, Launceston (QVM); Western

Australian Museum, Perth (WAM). Serial

designation of the cheek dentition follows Flower

(1867), Wilson and Hill (1897) and Luckett

(1993), except that the third adult premolars are

now recognised as the only second generation

cheek teeth in marsupials (Cifelli et al. 1996;

Luckett and Woolley 1996). Dental nomenclature

follows Tedford and Woodburne (1987), Ride

(1993) or is standard. Mensuration follows

Tedford (1966) and Wells and Murray (1979). All

measurements are in millimetres.

SYSTEMATICS

Order DIPROTODONTIA Owen 1866

Superfamily MACROPODOIDEA Gray 1821

Family MACROPODIDAE Gray 1821

Subfamily STHENURINAE (Glauert 1926)

Genus Simosthenurus Tedford 1966

Simosthenurus newtonae sp. nov.

Sthenurus oreas DeVis (in part), 1895: 97.

Sthenurus atlas Glauert, 1912: 64.

Sthenurus andersoni Bartholomai (in part), 1963:

58, fig. 3.

Sthenurus sp. Lundelius, 1963: 77, fig. 2.

Sthenurus sp. cf. S. gilli Merrilees, 1965: 29-30.

Sthenurus andersoni Tedford (in part), 1966: 25.

Sthenurus sp. Il Marcus, 1976: 71, 74, fig. 27c-d.

Simosthenurus sp. II Pledge, 1980: 137, table 3.

Sthenurus sp. Williams, 1980: 107, site 30.

Sthenurus sp. cf. S. atlas Williams, 1980: 107,

site 37.

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G. J. PRIDEAUX

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FIGURE 1. Localities yielding remains of Simosthenurus newtonae sp. nov. (black squares): 1, Tight Entrance

Cave; 2, Balladonia; 3, Madura Cave; 4, Lindsay Hall Cave; 5, Curramulka Quarry; 6, Black Rock Gravel Pit; 7,

Baldina Creek; 8, Comaum Forest Cave; 9, Haystall Cave; 10, Henschke’s Fossil Cave; 11, SOS Cave; 12, Victoria

Fossil Cave; 13, Goulden’s Hole; 14, Green Waterhole Cave (Type locality); 15, Kilsby’s Hole; 16, McEachern’s

Cave; 17, Scotchtown Cave; 18, Teapot Creek; 19, Kandos; 20, Wellington Caves; 21, Darling Downs. Triangles

denote localities yielding Simosthenurus occidentalis.

Sthenurus sp. Lundelius & Turnbull, 1989: 2, 4,

fig. la.

Simosthenurus sp. nov. Prideaux & Wells, 1994:

227.

Sthenurus ‘P17250’ McNamara, 1994: 111, 115.

Sthenurus ‘P17250’ Prideaux & Wells, 1997:

191, 194.

Holotype

SAM P17249-P17250, partial adult cranium,

and fused left and right dentaries (Figs. 2, 3).

Type Locality

Green Waterhole Cave, a submerged cave near

Tantanoola, southeastern South Australia. Faunal

composition suggests a late Pleistocene age

(Pledge 1980; Newton 1988).

Paratypes

FU 0227, near complete adult cranium; FU

0252, juvenile cranium devoid of occipital region;

SAM P20255-P16632-P16633, partial adult

cranium, incomplete left and right dentaries; SAM

NEW STHENURINE KANGAROO 3

P28969, near complete, but crushed adult

cranium; FU 0179, right dentary. All paratypes

are from the Fossil Chamber in Victoria Fossil

Cave, Naracoorte, southeastern South Australia.

This locality is considered to be late Pleistocene,

but is probably older than 212 000 years (Wells et

al. 1984; Ayliffe et al. 1998).

Referred Specimens

Victoria Fossil Cave, Naracoorte, SA: FU 0205,

partial right juvenile dentary; FU 0226, right M,;

FU 0259, partial left juvenile dentary; FU 0293,

partial left adult dentary; FU 0887, partial left

juvenile maxilla; FU 1084, partial left juvenile

maxilla; SAM P16550, partial juvenile cranium;

SAM P20243, partial left adult maxilla; SAM

P27631, left I, M,,; SAM P28149, partial left

adult dentary; SAM P28478, right P,; SAM

P28479, right M,, M,; SAM P28518, left I; SAM

P28671, left juvenile dentary; SAM P28996, left

juvenile dentary, SAM P32533, left M,; SAM

P32541, left I; SAM P32542, left I, M,,; SAM

P32545, partial right juvenile dentary.

Haystall Cave, Naracoorte, SA: SAM P36624,

right dP.,.

Henschke’s Fossil Cave, Naracoorte, SA:

SAM P17837, partial left juvenile maxilla, right

dP’, right M'; SAM P18554, right M' metaloph;

SAM P34807, right P?; SAM P34808, left P?;

SAM P34809, right P?; SAM P34810, right P,;

SAM P34811, left P?; SAM P38788, dP,_,, P,,

M,.,, right P,, M,; SAM P38789, left and right

M*; SAM P38790, left P,; SAM P38791, right

M,; SAM P38792, left P’; and SAM P

unregistered, several loose teeth.

SOS Cave, Naracoorte, SA: SAM P33476,

partial right adult maxilla.

Comaum Forest Cave, near Penola, SA: SAM

P31967, fused left and right premaxillae and

maxillae.

Goulden’s Hole, near Mount Schank, SA: SAM

P36620, right P*; SAM P36621, left P,; SAM

TAA AL

ie) a 2

FIGURE 2. Right lateral view of partial cranium and dentaries of Simosthenurus newtonae sp. nov. holotype, SAM

P17249 / P17250. Small gradations on scale bar are millimetres.

4 G. J. PRIDEAUX

P36622, left M'; SAM P38780, left M*; SAM

P38781, left M,.

Kilsby’s Hole, Mount Gambier, SA: SAM

P38782, left P?, SAM P38783, right lower molar,

SAM P38784, left upper molar, SAM P38785,

right dP.,.

Black Rock Gravel Pit, near Orroroo, SA: SAM

P23166, left and right adult dentaries.

Baldina Creek, Burra, SA: SAM P21035, partial

juvenile dentary.

Curramulka Quarry, Yorke Peninsula, SA:

SAM P38786, right P®, SAM P38787, right P,.

McEachern’s Cave, near Nelson, VIC: SAM

P17319, partial right adult dentary; NMV

P198434—-P198438—P198440, left and right adult

maxillae; NMV P198435—P198436, left and right

adult maxillaj NMV P198439, partial left adult

dentary; NMV P198449, partial left adult maxilla;

NMV P198450, right P? in maxilla fragment.

Teapot Creek, Monaro Region, NSW: AM F

unregistered, partial right adult dentary.

Wellington Caves, NSW: AM F18872, left P?.

Kandos, NSW: AM F73721, partial left adult

dentary.

Darling Downs, southeastern QLD: QM F2978,

partial right adult maxilla.

Scotchtown Cave, near Smithton, TAS:

QVM:1992:GFV:232, right M,; QVM:1992:

GFV:238, right M?; QVM:1992:GFV:242, right

M!.

Madura Cave, Madura, WA: FMNH PM4356,

right P.,.

Lindsay Hall Cave, near Madura, WA: WAM

92.12.3, right M'; WAM 92.12.10, right My’;

WAM 00.1.1, partial right juvenile maxilla.

Balladonia Soak, near Balladonia, WA: WAM

63.11.2-63.11.3, right adult dentary.

Tight Entrance Cave, southwestern WA: WAM

FIGURE 3. Palate and cheek dentitions of Simosthenurus newtonae sp. nov. holotype, SAM P17249 / P17250: A,

palatal view (stereo); B, occlusal view of right lower cheek tooth row (stereo). Abbreviations: c, precingulum; i,

inflation of posterior face of hypolophid. Scale bars = 10 millimetres.

NEW STHENURINE KANGAROO 5

97.5.312-97.5.314-97.5.314—99.11.142, left adult

dentary.

Species Diagnosis

Cranium similar in size to Simosthenurus

brownei Merrilees 1967 and Si. occidentalis

Glauert 1910, but less brachycephalic, with

narrower, more elongate rostrum, longer diastema

and less inflated frontal region. Masseteric process

short, narrowing distally and twisted posteriorly.

P? short for width, crown inflated anterobuccally

and thus only slightly wider posteriorly than

anteriorly. Upper and lower molars relatively

high-crowned with very few to no fine enamel

crenulations. Crests on molars very well-

developed, generally maintaining strong contact

with cusp apices. Precingulum abruptly terminated

at position of preprotocrista. Pterygoid fossa

narrow and dorsoventrally deep. Lower molars

bear pronounced anterior turn of lophid ends and

strongly inflated posterior face of hypolophid.

Etymology

The new species is named after Cate A. Newton

who studied the fossil fauna of Green Waterhole

Cave and first recognised the morphological

uniqueness of the holotype.

Description

Cranium (Figs 2-5; Table 1).

Premaxilla rather slender in lateral view, but

flaring dorsally for extended contact with nasal.

Incisor-bearing portion of premaxilla rather

shallow and elongate (Fig. 5). Diastema

moderately elongate, maxilla-premaxilla suture

elongate and zigzagged. Cranial diastema

deflected anteroventrally relative to cheek tooth

row. Incisive foramina long with anterior border

Opposite or just posterior to posterior extremity of

I alveolus. Rostrum narrow, deep and elongate.

Anteorbital/buccinator fossae on maxilla very

shallow, resulting in reduced mesial curvature of

diastema ridge, and rather flat-sided maxilla

anteroventral to centre of orbit. Nasals long, broad

posteriorly, narrowing anteriorly. One to three

infraorbital foramina anterior to ventral border of

orbit. Masseteric process short, moderately wide,

flaring slightly towards distal end, twisted

posteriorly under anterior portion of jugal (Figs

2-5). Process consists primarily of maxilla with

only modest jugal contribution. Frontal region

rather elongate in dorsal view, only moderately

inflated laterally (Fig. 4A). Palatine vacuities

extend anteriorly to opposite M! precingulum.

Postpalatine bars form a thin bridge across palate

opposite or posterior to M* metaloph (Fig. 3A).

Basicranial plane markedly elevated above

palatal plane. Cranial pterygoid fossa wide and

deep. Basioccipital slightly flexed posterodorsally

relative to basisphenoid, and bears well-developed

medial keel. Zygomatic arch deep, with a very

wide ectoglenoid process at posterior extremity of

jugal. Postglenoid process and glenoid fossa very

large. Temporal crests not fully convergent on

sagittal suture and only moderately developed.

Occipital region broad, but not especially deep,

and oriented at 90° relative to dorsal surface of

neurocranium. Vertical medial occipital crest

TABLE 1. Mean dimensions of the adult cranium and dentary of Simosthenurus newtonae sp. nov. compared with

the dimensions of the holotype (SAM P17249-P17250) and mean dimensions of Simosthenurus occidentalis and

Sthenurus andersoni. Standard deviation is given in parentheses; sample size in brackets.

Dimension Simosthenurus

newtonae

Condylobasal Length 216 (3.0) [n=3]

Diastema Length

% Diastema Length: Palatal Length

Palatal Length

Palatal Width between M! Protolophs

Palatal Width between M‘ Protolophs

Max. Width across Zygomatic Arches

Maximum Width across Frontals

Width across Paroccipital Processes

Dentary Depth

Dentary Width

Dentary Depth / Width

40.3 (1.04) [n=4]

30.5 (0.85) [n=3]

132 (1.2) [n=3]

37.2 (2.27) [n=5]

39.2 (4.05) [n=3]

132 (5.6) [n=4]

77.5 (3.05) [n=4]

76.3 (5.12) [n=4]

34.8 (3.35) [n=5]

20.5 (1.20) [n=5]

1.70 (0.13) [n=5]

Holo- Simosthenurus Sthenurus

type occidentalis andersoni

- 198 (4.9) [n=4] 221 (8.5) [n=2]

= 31.8 (1.42) [n=4] 47.2 (4.38) [n=2]

- 25.2 (0.01) [n=4] 36.0 (2.83) [n=2]

- 126 (4.1) [n=4] 132 (2.1) [n=2]

39.0 35.0 (0.46) [n=3] 38.8 (0.92) [n=2]

43.1 41.2 (0.53) [n=3] 42.1 (0.14) [n=2]

136 143 (1.53) [n=3] 114 (2.1) [n=2]

82.0 91.8 (3.22) [n=4] 63.4 (1.98) [n=2]

82.0 67.7 (2.67) [n=3] 62.7 (3.25) [n=2]

39.3 37.6 (1.70) [n=13] 28.4 (1.82) [n=10]

20.6 23.4 (2.03) [n=13] 17.2 (2.04) [n=10]

1.91 1.62 (0.14) [n=13] 1.65 (0.13) [n=10]

6 G. J. PRIDEAUX

—

FIGURE 4. Cranium of Simosthenurus newtonae sp. nov. paratype, SAM P28969: A, dorsal view; B, palatal view.

Abbreviations: f, frontal; m, masseteric process. Scale bar = 10 millimetres.

slight to well-developed, and leads ventrally to

wide foramen magnum bordered by moderately

large occipital condyles. Paroccipital processes

deflected posteroventrally. Nuchal crests strongly-

developed and extended posteriorly (Fig. 4A).

Upper Dentition (Figs 2—5; Table 2).

I' quite low in crown height and rather rounded

in cross-section (Figs 4-5). Vertical occlusal facet

faces posteriorly. Strongly curved anterior surface

of crown extends forward well beyond anterior

extremity of premaxilla. ? round in cross-section,

one-third size of I'. I? unknown, but alveolus

suggests elongate crown.

Second upper deciduous premolar (dP?)

reminiscent of P? in general morphology, but

NEW STHENURINE KANGAROO t

FIGURE 5. Right lateral view of partial cranium of Simosthenurus newtonae sp. nov. paratype, SAM P20255.

Scale bar = 10 millimetres.

much shorter relative to width. Outline of tooth

very rounded, especially on lingual side. Main

crest straight and aligned with antero-posterior

plane of tooth, lingual crest very curved and much

lower in height than main crest. Anterior basin

absent. Posterior basin well-developed, half size

of longitudinal basin and separated by marked

transverse ridge. Third upper deciduous premolar

(dP?) completely molariform, narrower across

protoloph than metaloph and smaller than molars.

Small precingulum restricted to buccal two-thirds

of protoloph face, terminating at position of

preprotocrista. Unworn crest of protoloph very

convex anteriorly with very thick, united

postparacrista and premetacrista. Remaining

aspects very similar to molars, except for fine

postlink and crest centred on posterior face of

protoloph. P? usually short for width, only slightly

wider posteriorly than anteriorly (Figs 3-5; Table

2). Outline rounded, with anterobuccal aspect

strongly inflated and lingual side convex.

Posterobuccal accessory cusp absent, poorly

differentiated or slight, very occasionally with

small, poorly differentiated cuspule anterior to it.

Lingual crest lower and usually more curved than

main crest. Anterior basin small and not well

differentiated. Longitudinal basin short, of modest

width and depth, and occupied by fine to coarse

transverse ridgelets. Posterior basin well-

developed, and separated from longitudinal basin

by strong transverse ridge formed by two strong

ridgelets united at tooth midline, or by buccally

curved end of lingual crest.

Upper cheek tooth row moderately curved

buccally, with P? turned in slightly relative to

molars (Figs 3A, 4B). Upper molars sized

M'<M?<M?>M‘ (Figs 2-5; Table 2). Metaloph

narrower relative to protoloph on M‘ than on M!?.

Lophs moderately high-crowned, with unworn

crests slightly convex anteriorly. Loph faces bear

few very fine or no enamel crenulations.

Preparacrista strongly developed, and maintains

strong, direct contact with paracone apex.

Precingulum smoothly confluent’ with

preparacrista and terminates at tiny remnant of

preprotocrista, after extending across two-thirds of

anterior face of protoloph (Fig. 3A). Prominent

postprotocrista ascends posterobuccally into

interloph valley, but only extends onto base of

metaloph face as very short, fine crest. Well-

developed postparacrista and moderately

developed premetacrista curve in lingually, their

union forming a distinct notch. Posterior face of

metaloph dominated by well-developed

postmetaconulecrista, which curves buccally

across approximately three-quarters of posterior

face of metaloph to meet similarly-developed

postmetacrista. Tiny cuspule positioned at union

of postmetaconulecrista and postmetacrista most

prominent on My’, and probably represents stylar

cusp E. Posterior face of metaloph markedly

inflated above postmetaconulecrista, especially

more buccally.

Dentary (Figs 2, 3, 6; Table 1).

Ramus deep for width, especially posteriorly

due to large digastric eminence (Figs 2,6), which

curves in mesially along ventral border. Digastric

8 G. J. PRIDEAUX

a de

FIGURE 6. Right dentary of Simosthenurus newtonae sp. nov. paratype, FU 0179: A, lateral view; B, mesial view;

C, occlusal view of excavated right P, (stereo). Abbreviations: b, buccal crest; de, digastric eminence; ds, digastric

sulcus; mc, main crest; mhg, mylohyoid groove; pf, pterygoid fossa. Scale bars = 10 millimetres.

sulcus large and extends anteriorly to beneath M,

hypolophid or M, protolophid (Fig. 6B). Dentary

decreases in depth anteriorly, with diastema

somewhat procumbent relative to cheek tooth row

(Figs 2, 6). Symphysis rather slender and

procumbent, with dorsal border parallel with

alveolar margin of cheek tooth row. Posterior

portion of symphysis underlies small genial pit,

and only extends to beneath posterior root of P,.

Median dorsal groove deep and narrow. Anterior

mental foramen located just anteroventral of

buccinator sulcus. Buccinator sulcus shallow

anteriorly, but gradually deepens posteriorly,

terminating beneath M, hypolophid. Posterior

NEW STHENURINE KANGAROO 9

mental foramen mid-depth on ramus beneath M,

hypolophid.

Ascending ramus rather short in lateral profile

with anterior root located adjacent to M,

hypolophid (Figs 2, 6). Anterior root forms buccal

aspect of wide, deep postalveolar fossa.

Masseteric foramen large and elliptical, leading

into deep vertical masseteric canal which does not

extend into body of ramus. Inferior mandibular

foramen usually fairly large, with dorsal border at

level of alveolar margin of cheek tooth row.

Pterygoid fossa high and narrow, bearing

pronounced upward projection of angular process

(Figs 2, 6). Adjacent to ventral border of inferior

mandibular foramen and at anterior extremity of

pterygoid fossa, two sharp processes overhang

mylohyoid groove; a posteroventrally-oriented

process (cf. lingula in humans) and an

anterodorsally-oriented process extending from

anterior end of mesial border of pterygoid fossa

(Fig. 6B). Mandibular condyle large.

Lower Dentition (Figs 2, 6; Table 2).

I, large, robust, elongate (Figs 2, 6). Occlusal

surface slightly higher than, but oriented close to

parallel with diastema. Level of I, occlusal surface

well below level of cheek tooth occlusal surfaces.

Second lower deciduous premolar (dP,) similar in

overall morphology to P,, but shorter relative to

width. Main crest runs obliquely across tooth from

posterolingual corner to central position

anteriorly. Short buccal crest trends buccally from

posterior extremity of main crest, then curves

anteriorly to run parallel to posterior part of main

crest for short distance. Third lower deciduous

premolar (dP,) completely molariform, with

protolophid slightly narrower than hypolophid and

tapered markedly toward relatively narrow unworn

crest. Small paraconid positioned at anterior

extreme of trigonid, lingual to end of paracristid.

Premetacristid runs posteriorly from paraconid,

terminating midway up metaconid anterior face.

Thick ?parametacristid descends smoothly from

metaconid apex terminating in middle of trigonid

basin. ?Preprotostylocristid runs anteriorly from

topographic protoconid apex to point where

paracristid turns lingually. Cristid obliqua

strongly-developed, continuous with buccal

extremity of hypolophid, meets protolophid

slightly posterolingual and ventral to protoconid.

Preentoconid low, curves from entoconid apex

into interlophid valley to near midline of tooth.

Very slight posterobuccally-accentuated inflation

present on posterior face of hypolophid. Lophid

faces lack enamel crenulations.

P, small and moderately short relative to

molars, usually similar to or slightly shorter than

M, in length (Figs 2, 3, 6; Table 2). Posterobuccal

aspect of tooth inflated and bearing short, thick

buccal crest separated from posterior end of main

crest by very shallow, narrow median valley in

holotype (Fig. 3B). In paratypes and referred

specimens, inflation of posterobuccal corner less

marked; buccal crest usually slightly longer and

bears a thin, crescentic buccal crest separated from

main crest by small and shallow median valley

(Fig. 6C). Cuspules of main crest poorly

differentiated posteriorly, but three usually

distinguishable anteriorly. Very few to no

ridgelets traverse median valley.

Lower cheek tooth row shows slight to

moderate buccal curvature, with main crest of P,

oriented slightly anterobuccally relative to molars

(Fig. 3B). Moderately high-crowned lower molars

sized M,<M,<M,>M, (Figs 2, 3, 6, Table 2).

Lophid faces often lack enamel crenulations, but a

few very fine crenulations may be present on

anterior faces. Anterior turn of lophid ends very

pronounced (Fig. 3B). Paracristid and cristid

obliqua well-developed, with former maintaining

contact with topographic protoconid apex, and

latter contacting or very close to hypoconid apex

(Fig. 3B). Precingulid/trigonid shelf narrow and

moderately short. Transverse (anterior) portion of

paracristid broadly U-shaped in unworn teeth,

curving out to anterior extremity of tooth, above

precingulid, then posteriorly, usually uniting with

fine, low premetacristid. ?Parametacristid either

slight or absent. Cristid obliqua on minimally

worn teeth divided into anterior and posterior

components, although moderate wear produces

one continuous crest. Lower posterior component

curves smoothly anteriorly from hypoconid apex

and terminates on buccal side of anterior

component, located closer to midline of tooth and

oriented less obliquely. Anterior component of

cristid obliqua extremely slight anterior to

interlophid valley, fining and terminating midway

up posterior protolophid face. Preentoconid low

and very slight. Shelf-like postcingulid absent, but

buccal two-thirds of posterior aspect strongly

inflated more ventrally, and overlapped by

trigonid of succeeding molar (Figs 3B, 6A).

Intraspecific Variation

Cranium

There is very little variation in the general size

and morphology of known adult crania that cannot

be ascribed to differential preservation. The only

noteworthy variation is a slight difference in

G. J. PRIDEAUX

TABLE 2. Cheek tooth dimensions of Simosthenurus newtonae sp. nov., showing mean and standard deviation (in

parentheses), dimensions of the holotype (SAM P17249-P17250), and the Wellington Caves and Kandos specimens

(AM F18872, AM F73721).

Tooth Length

UPPER DENTITION

dP? 10.3 (0.91)

dP? 11.5 (0.20)

Mean 16.6 (0.72)

Holotype 16.9

AM F18872 18.3

Mean 13.2:(0:32)

Holotype 13.2

Mean 14.5 (0.60)

Holotype 14.7

Mean 15.0 (0.59)

Holotype 15.0

Mean 14.1 (0.67)

Holotype 14.8

LOWER DENTITION

dP, 9.5 (0.68)

dP, 10.4 (0.21)

Mean 15.3 (0.74)

Holotype 15.3

AM F73721 17.5

Mean 13.1 (0.67)

Holotype 13.1

AM F73721 14.3

Mean 15.2 (0.61)

Holotype 15.7

AM F73721 16.0

Mean 16.3 (0.45)

Holotype 16.6

AM F73721 17.3

Mean 14.6 (0.79)

Holotype 15.8

AM F73721 17.0

Width

7.7 (0.51)

9.8 (0.66)

10.5 (0.82)

10.6

10.0

12.9 (0.50)

13.1

13.9 (0.82)

13.7

13.7 (0.53)

13.8

12.9 (0:36)

13.0

6.1 (0.49)

8.3 (0.57)

7.6 (0.43)

Hell

8.4

10.2 (0.37)

10.1

11.4

11.3 (0.38)

11.5

12.5

12.1 (0.29)

12.1

13.5

11.5 (0.28)

113

13.0

Posterior

Width

9.6 (0.97)

10.9 (0.42)

12.3 (0.52)

12.9

13.4

12.6 (0.38)

12.8

13.0 (0.65)

13.0

12.9 (0.63)

13-3

11.1 (0.42)

11.5

8.1 (0.31)

8.8 (0.53)

9.3 (0.33)

O07

9.9

10.5 (0.35)

10.7

11.7

11.6 (0.34)

Pl?

13.0

12.3 (0.42)

12.4

13.8

10.4 (0.44)

10.7

12.2

Height

7.1 (0.47)

6.2 (0.85)

10.4 (0.61)

11.0

10.3

7.7 (0.79)

8.3

8.2 (0.71)

9.0

7.9 (0.48)

8.3

6.6 (0.55)

7.9 (0.95)

7.1 (0.89)

10.1 (0.79)

10.0

9.2 (0.98)

hil

10.0 (1.38)

9.2

9.5 (1.43)

9:5

10.1

8.4 (0.74)

8.6

10.8

Posterior

Height

7.8 (0.81)

7.0 (1.15)

10.1 (0.86)

10.8

9.8

9) (0.71)

8.0

8.4 (0.71)

8.7

7.9 (0.37)

8.3

6.2 (0.52)

6.6

7.1 (0.83)

7.2 (0.90)

9.4 (1.16)

10.1

9.7 (1.47)

10.4 (1.35)

9.6

9.4 (1.25)

9:5

11.7

7.3 (0.55)

7.3

10.3

Sample

Size

NEW STHENURINE KANGAROO 1]

zygomatic arch depth, with SAM P28969

shallower than SAM P17249, SAM P20255 and

FU 0227.

The juvenile cranium, FU 0252 (dP?3, M'

erupted), clearly differs from the adult crania in

proportion. The rostrum is relatively longer, the

ventral orbital border of the jugal is less laterally

expanded, the frontal region is less inflated, the

temporal crests are less convergent upon one

another, and the width across the zygomatic arches

is comparatively less.

Upper Dentition

No appreciable variation is visible in I' size or

morphology between the four specimens

preserving that tooth. These same specimens also

preserve I? but, in contrast to I', ? of SAM

P20255 is slightly smaller in cross-sectional area

than in the other specimens, even taking into

consideration its greater degree of wear. Minimal

size or morphological variation has been observed

for dP’, aside from the presence of an incipient

posterobuccal accessory cusp in SAM P16550, a

low, slight posterobuccal cingulum in FU 1084,

and the complete absence of any such feature in

FU 0252 and FU 1084. The longitudinal basin is

slightly larger in FU 0252 than FU 1084, but

SAM P16550 is too worn to determine. No

notable size or morphological variation is visible

in dP*. Upper molar variation is also very limited,

with the slightest fine enamel crenulations visible

on the unworn molars of some specimens.

Marked variation exists in both size and

morphology between the P? of the holotype,

paratypes and referred specimens. Size differences

are present in absolute length and width, crown

height, and width relative to length, although no

geographically-correlated size variation is evident.

Morphological variation is displayed in: a) degree

of posterobuccal accessory cusp development

(ranging from clearly defined with small anterior

cuspule, to poorly differentiated, to completely

absent); b) degree of curvature of lingual crest

and degree to which both crests are divided into

cuspules; c) nature of coarse ridgelet separating

posterior and longitudinal basins (composed of

continuation of lingual crest or two transverse

ridgelets); d) general inflation or roundness of

tooth outline.

It is worth singling out a rather large, unworn

P? from Wellington Caves (AM F18872) for

special consideration, because it is much wider

posteriorly than anteriorly. Although this conflicts

with one of the defining characteristics of Si.

newtonae, all other features of the tooth fit within

the recognised Si. newtonae morphospace. These

features include the inflation of the anterobuccal

corner, shape of the lingual crest relative to the

main crest, presence of one small cuspule

immediately anterior to a poorly differentiated

posterobuccal accessory cusp, and buccal

curvature of the posterior end of the lingual crest,

such that it partially separates the longitudinal and

posterior basins. For these reasons, and because

Si. newtonae is known from a dentary with rather

large molars from nearby Kandos (AM F73721), I

am confident that the Wellington Caves specimen

is Si. newtonae.

Dentary

Few morphological differences are visible

between the adult dentary specimens apart from

slight variation in the posterior extent of the

symphysis (beneath the anterior and posterior

roots of P,), and degree of development of the

digastric eminence and sulcus. As a consequence

of the latter variable feature, dentary depth varies

slightly between specimens (e.g., compare the

holotype dimensions with mean dimensions in

Table 2). Variation is also present in the size of

the inferior mandibular foramen and development

of the processes overhanging the mylohyoid

groove. Relatively, the holotype has the deepest

dentary, largest inferior mandibular foramen and

largest mylohyoid associated processes.

Disregarding overall size differences, comparison

of adult and juvenile specimens reveals a trend

with age for increased dentary depth relative to

width, and development of the digastric eminence

and sulcus.

Lower Dentition.

No significant variation is visible in I, and dP..

Similarly, there is little variation in dP,, with only

a slight difference between specimens in overall

size, relative length of the buccal crest, and the

variable presence of a ridgelet linking the anterior

end of the buccal crest to the second cuspule of

the main crest. While some size variation is

evident in P,, the considerable degree of

morphological variation mirrors that observed in

the P®. Variation is displayed in: a) general tooth

outline (rounded and gently narrowing anteriorly

with minimal differentiation into anterior and

posterior portions, posterobuccal corner inflated

with much of buccal side and lingual side near

parallel); b) shape of main crest (longitudinally

straight, slightly sinusoidal); c) degree to which

main crest cuspules are differentiated; d) relative

length and shape of buccal crest (very short and

12 G. J. PRIDEAUX

thick, short and crescentic); e) low, fine ridgelet

connecting buccal crest to second cuspule of main

crest (absent, present); f) width of median valley

and degree of development of contained ridgelets.

The P, also varies slightly in the degree to which

its longitudinal axis is deflected anterobuccally

relative to the curvature of the molar row. The

minimal variation noted for the upper molars also

holds for lower molars.

The Kandos dentary (AM F73721) varies

slightly from typical Si. newtonae, because its P,

and molars are around 10% larger. This is

noteworthy in view of the marked similarity in

dental size between specimens of Si. newtonae

from across its wide range, which includes

western and Tasmanian representatives. Although

the dentition is considerably worn, no

morphological differences are evident between the

cheek teeth of AM F73721 and similarly worn

specimens of Si. newtonae. While it is important

to note that the digastric region, ascending ramus

and anterior portion of the ramus are missing in

AM F73721, the dentary of this specimen only

differs from other individuals of Si. newtonae by

possessing a buccinator sulcus that is slightly

deeper anteriorly. In view of the marked variation

in size observed within other Simosthenurus

species (Prideaux 1999), I have very little

hesitation in referring this specimen to Si.

newtonae.

Comparison with other taxa

Cranium.

The cranium of Simosthenurus newtonae is

more dolichocephalic than that of any other

Simosthenurus species. In relative skull length it

is intermediate between the other Simosthenurus

species and Sthenurus Owen 1874, resembling

Hadronomas puckridgi Woodburne 1967 in this

regard. Among the species of Simosthenurus, the

Si. newtonae cranium is most similar to Si.

occidentalis, but it differs by having a more

elongate rostrum and diastema, and a less

posterodorsally-flexed basioccipital region.

Despite the otherwise similar occipital and

basicranial proportions of the two species, the

direct effect of these differences is that the portion

of the cranium posterior to the end of the maxilla

is longer in Si. newtonae than it is in Si.

occidentalis. Inflation of the frontal region and

development of the supraorbital crests are less

pronounced in Si. newtonae than in Si.

occidentalis and Si. brownei, but greater than in

Si. gilli Merrilees 1965 and Si. baileyi Prideaux

and Wells 1998. The masseteric process is shorter

and much narrower than Si. occidentalis, and is

closest in morphology to Si. maddocki Wells and

Murray 1979, but more twisted posteriorly.

Moderate development of the temporal crests

is similar to Si. baileyi. The shallow

anteorbital/buccinator fossae and reduced mesial

curvature of the diastema border resemble Si.

baileyi as well as Si. gilli, but both of these

species have much shorter rostra.

Upper Dentition.

The general shape of the Si. newtonae I' is

typical of most Simosthenurus species, but the

tooth is quite low-crowned, akin to that of Si.

brownei. Although the elongate I? alveolus

probably indicates a relatively elongate crown, I

have observed no I? which may be confidently

ascribed to Si. newtonae. The dP? of Si. newtonae

is smaller than in southeastern Si. brownei and Si.

occidentalis, but larger than in Si. maddocki and

Sthenurus andersoni Marcus 1962. Morpho-

logically, the tooth recalls Si. brownei and Si.

baileyi, but it is less inflated posteriorly. Although

quite variable in form, the P? of most Si. newtonae

individuals is quite dissimilar to the other

Simosthenurus species. This is especially so

because, relative to its length, the tooth is usually

quite wide anteriorly as well as posteriorly. The

manner in which the posterior end of the lingual

crest curves buccally to partially or wholly

separate the longitudinal and posterior basins is

only seen elsewhere in S. andersoni, as well as a

P? fragment from the early Pliocene Bow Local

Fauna of central eastern New South Wales (see

Fig. 1A in Flannery and Archer 1984). Marked

inflation of the anterobuccal corner of P? is only

seen in rare individuals of other Simosthenurus

species.

The upper molariform teeth of Si. newtonae are

unique among Simosthenurus, and cannot readily

be confused with any other species. They are

similar to Si. baileyi in size, but are easily

distinguished by being higher-crowned, lacking

any noteworthy enamel crenulations, and having

the primary crests strongly connected to cusp

apices. The smaller Si. gilli and Si. maddocki

upper molars bear some resemblance to Si.

newtonae in this latter feature, as well as the

curved nature of the postparacrista and

premetacrista. However, all crests on the molars

of Si. maddocki are more weakly developed than

in Si. newtonae, while the majority of crests are

more weakly developed in Si. gilli. The

postprotocrista, which is divided into two

components, and the better developed upper molar

NEW STHENURINE KANGAROO 13

midline crest of Si. gilli are the two exceptions.

Si. maddocki may also be distinguished from Si.

newtonae by the many very fine enamel

crenulations that coat its loph surfaces. Overall,

the molars of Si. newtonae are most similar to S.

andersoni, but they differ by having the

preparacrista much more strongly connected to the

paracone apex, a stronger premetacrista, a stronger

postmetaconulecrista and no continuation of the

precingulum beyond the preprotocrista. Although

a precingulum that does not extend lingually

beyond the preprotocrista on the upper molars is

unique to Si. newtonae, this condition is observed

on the dP? of S. andersoni.

Dentary.

Si. newtonae is most similar in dentary size and

morphology to Si. brownei, but the digastric

eminence of the latter species is usually larger, as

is the gradient of decreasing dentary depth

anteriorly. In addition, the masseteric fossa is

longer in Si. brownei, and the anterior root of the

ascending ramus lies opposite the M,—-M,

boundary or M, protolophid. In contrast, the

anterior root in Si. newtonae leaves the ramus

adjacent or just posterior to the M, hypolophid.

The slightly procumbent diastema of Si. newtonae

is not observed in any other Simosthenurus

species, but is observed in Sthenurus, in S.

tindalei Tedford 1966 and S. stirlingi Wells and

Tedford 1995. The slender symphysis of Si.

newtonae is most similar in size and form to that

of Si. brownei, but it does not extend under the

genial pit to the same degree, and its dorsal

surface is nearly horizontal rather than

anterodorsally-oriented. The narrow and deep

median dorsal groove present in Si. newtonae is

also characteristic of Si. maddocki.

Viewed posteriorly, the pterygoid fossa of Si.

newtonae is narrower than in any other

Simosthenurus species. Marked development of

the processes overhanging the mylohyoid groove

is similar to Si. oreas De Vis 1895, but the groove

in the latter species is deeper and narrower.

Lower Dentition.

In size and morphology, the I, of Si. newtonae

is intermediate in morphology between Si.

occidentalis and S. andersoni. In this sense, the

tooth resembles the I, of Si. pales De Vis 1895,

but is much smaller. Size and general outline of

the dP, is similar to Si. occidentalis, but the tooth

is relatively narrower anteriorly. The shortness of

the buccal crest is similar to that observed in Si.

maddocki, but the dP, of this species is narrower

and the main crest cuspules are more distinct. As

with the P?, the anterior width of P, is not

markedly exceeded by the posterior width of the

tooth. Overall, Si. newtonae is particular similar

to Si. brachyselenis Prideaux and Wells 1997 in

size and general morphology, but differs by being

longer relative to the molars and having a slightly

longer buccal crest. While the P, of the Si.

newtonae holotype is similar in outline to that of

Si. brachyselenis, other specimens of Si. newtonae

often narrow more gradually anteriorly. Compared

with ‘Simosthenurus’ cegsai Pledge 1992, the Si.

newtonae P, is more inflated posterobuccally, the

median valley is usually wider, and the buccal

crest is longer.

Although the protolophid base of the Si.

newtonae dP, is narrower than the hypolophid

base, the unworn crest of the protolophid is much

narrower, similar to that of Si. pales. Si. newtonae

appears to retain a paraconid lingual to the

paracristid in the anterolingual corner of the dP,

trigonid. A similar cusp is often observed on the

dP, of other sthenurines, such as Si. gilli, S. atlas

(Owen 1838) and Hadronomas puckridgi. Aside

primarily from the stronger connection between

the cristid obliqua and hypoconid apex,

morphology of the Si. newtonae dP, is very

similar to the succeeding molars. Among the

species of Simosthenurus, the morphology of the

Si. newtonae lower molars is quite unique. In

some respects, their form more closely resembles

species of Sthenurus, most especially S.

andersoni. Similarities include the paucity of

enamel crenulations on the lophid faces, the

proximity of the paracristid and cristid obliqua to

the buccal cusp apices, and the anterior turn of the

lophid ends. Within Simosthenurus, the Si.

newtonae lower molars most resemble Si. gilli in

crown height and paucity of enamel crenulations,

but are easily distinguished by their larger size,

markedly inflated posterior face of the

hypolophid, more curved transverse portion of the

paracristid, and thicker, more buccally situated

paracristid and cristid obliqua. A curved

transverse portion of the paracristid is also

observed in Si. euryskaphus Prideaux and Wells

1997 and many specimens of Si. occidentalis, but

the paracristid and cristid obliqua of these two

species are shifted more lingually, the lophid faces

bear distinct fine enamel crenulations, and the

lophid ends are less markedly turned anteriorly. In

this latter feature and in the marked posterior

inflation of the hypolophid, Si. newtonae is easily

distinguished from all other Simosthenurus

species.

14 G. J. PRIDEAUX

Geographic Distribution

Simosthenurus newtonae is one of the most

widely distributed Pleistocene sthenurine

species. Overall, its distribution pattern most

closely resembles that of Si. occidentalis (Fig.

1). Both species occur in late Pleistocene cave

deposits in southwestern Australia (Merrilees

1979; Gully 1997), and were probably

distributed across the southern periphery of the

continent during periods when woodland or

forest was more extensive. However, of these

two species, only Si. newtonae has so far been

recorded from Balladonia (= Sthenurus atlas in

Glauert 1912) and the Nullarbor Plain

(Lundelius, 1963; Lundelius and Turnbull 1989;

Prideaux 1994; Aplin et al. 1995). Conversely,

only Si. occidentalis is known from the Eyre

Peninsula.

Remains of Si. newtonae and Si. occidentalis are

commonly encountered in the cave faunas of

southeastern South Australia (eg., Pledge, 1980;

Wells et al., 1984), although the latter species is

much better known in Victoria. Together, they

represent the only sthenurines known from late

Pleistocene cave deposits in Tasmania, where Si.

occidentalis is again by far the more abundant

(Murray and Goede 1977; Goede and Murray

1979). While Si. newtonae is also known from

southeastern Queensland (= S. andersoni in

Bartholomai 1963), its only other occurrence north

of the Monaro region in southeastern New South

Wales is in the form of a large-toothed variant in

the Kandos and Wellington Caves deposits.

ACKNOWLEDGMENTS

The work presented here formed part of a Ph.D.

degree undertaken at the Flinders University of South

Australia on the systematics and evolution of sthenurine

kangaroos. For discussions and advice I offer my

sincerest thanks to Rod Wells, Dick Tedford, Peter

Murray, Jim McNamara and David Ride. A great many

of the specimens listed or described herein exist only

because of the efforts of countless Flinders University

students and volunteers who have assisted with the

excavations in Victoria Fossil Cave over many years.

Peter Daenke and Ed Bailey are commended for their

excellent preparation of much of this material. I am

indebted to Neville Pledge (South Australian Museum),

Alisanne Ramsden (Queen Victoria Museum), Bob

Jones (Australian Museum), John Long (Western

Australian Museum), Ralph Molnar and Joanne

Wilkinson (Queensland Museum), and Tom Rich and

Betty Thompson (Museum of Victoria) for extended

loans of fossil material. Bernie Cooke and Dick Tedford

kindly reviewed the original manuscript and provided a

number of constructive comments.

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PRIDEAUX, G. J. & WELLS, R. T. 1998. Sthenurus

baileyi sp. nov., a new fossil kangaroo from the

Pleistocene of southern Australia. Transactions of

the Royal Society of South Australia 122: 1-15.

RIDE, W. D. L. 1993. Jackmahoneya gen. nov. and the

genesis of the macropodiform molar. Memoirs of the

Association of Australasian Palaeontologists 15:

441-459.

TEDFORD, R. H. 1966. A review of the macropodid

genus Sthenurus. University of California

Publications in Geological Sciences 57: 1-72.

TEDFORD, R. H. & WOODBURNE, M. O. 1987. The

Ilariidae, a new family of vombatiform marsupials

from Miocene strata of South Australia and an

evaluation of the homology of molar cusps in the

Diprotodontia. Pp. 401-418 Jn ‘Possums and

Opossums: Studies in Evolution’. Ed. M. Archer.

Surrey Beatty & Sons and the Royal Zoological

Society of New South Wales: Sydney.

WELLS, R. T. & MURRAY, P. F. 1979. A new

sthenurine kangaroo (Marsupialia, Macropodidae)

from southeastern South Australia. Transactions of

the Royal Society of South Australia 103: 213-

219.

WELLS, R. T., MORIARTY, K. & WILLIAMS, D. L.

G. 1984. The fossil vertebrate deposits of Victoria

Fossil Cave Naracoorte: an introduction to the

geology and fauna: The Australian Zoologist 21:

305-333.

WILLIAMS, D. L. G. 1980. Catalogue of Pleistocene

vertebrate fossils and sites in South Australia.

Transactions of the Royal Society of South Australia

104: 101-115.

WILSON, J. T. & HILL, J. P. 1897. Observations on

the development and succession of the teeth in

Perameles. Quarterly Journal of Microscopic

Science 39: 427-588.

TWO NEW SPECIES OF ANTIPORUS SHARP FROM WESTERN

AUSTRALIA (COLEOPTERA: DYTISCIDAE).

BY C.H.S. WATTS AND A. PINDER

Summary

WATTS, C.H.S. AND PINDER, A. (2000). Two new species of Antiporus Sharp, A. pennifoldae and A.

mcraeae, are described from the south —west of Western Australia. Both species appear to be

restricted to an area threatened by rising salinity.

TWO NEW SPECIES OF ANTIPORUS SHARP FROM WESTERN AUSTRALIA

(COLEOPTERA: DYTISCIDAE)

C. H.S. WATTS AND A. PINDER

WATTS, C. H. S. and PINDER, A. 2000. Two new species of Antiporus from Western

Australia (Coleoptera: Dytiscidae). Records of the South Australian Museum 33(1): 17-19.

Two new species of Antiporus Sharp, A. pennifoldae and A. mcraeae, are described from the

south-west of Western Australia. Both species appear to be restricted to an area threatened by

rising salinity.

C. H. S. Watts, South Australian Museum, North Terrace, Adelaide, South Australia 5000. A.

Pinder, Department of Conservation and Land Management, PO Box 51, Wanneroo, Western

Australia 6065. Manuscript received 11 October 1999.

Two new species of Antiporus Sharp have

recently been identified from the south-west of

Western Australia. These were among material

collected in a biological survey of wetlands in the

wheatbelt and adjacent areas by the Department

of Conservation and Land Management (CALM).

The survey is part of the department’s response to

problems (particularly salinity and waterlogging)

associated with rising groundwater in the region

as a consequence of extensive clearing of native

vegetation.

Members of the genus Antiporus are common

in still, or relatively still, water in southern

Australia. One species, A. femoralis Boheman, is

particularly common in the South-West. In a

recent revision of the genus, Watts (1997)

described two additional species from the region,

both seemingly rare with one, A. pembertoni

Watts, known only from one specimen from near

Pemberton and the other, A. hollingsworthi Watts,

from 15 specimens. Since then further collecting

has provided two additional localities for A.

hollingsworthi (30 km N Perth, coll. Watts, 15 km

S Northcliffe, coll. Pederzani; both in the South

Australian Museum), but none for A. pembertoni.

We herein report on and describe two additional

species which, like the above, are known from

very few specimens. The species were not

collected during a recent more intensive survey of

wetland invertebrates in the Lake Muir/Poorginup

region by A. Storey (personal communication) and

will probably prove to have a restricted

distribution. The area in which they were found is

generally threatened by rising salinity.

We describe the two species here to enable

them to be included in the ongoing investigation

of this important wetland region.

SYSTEMATICS

Antiporus mcraeae sp. nov.

Type

Holotype: male; ‘Melaleuca Swamp, Kodjinup

Nature Reserve, 50 km ESE Manjimup,

34°23'45"S 116°39'01"E, W.A. Coll. A. Pinder

and J. McRae (CALM), 2/10/98’ Registration

number WAM 26607, Western Australian

Museum.

Description (number examined, 1) (Figs 1-4)

Habitus. Length 4.4 mm. Elongate-oval, dark

red-brown, sides of pronotum broadly paler,

appendages lighter particularly towards

extremities.

Dorsal surface. Punctures rather small, regular,

those on head smaller than eye facet; row of larger

punctures adjacent to suture on elytron; a few

small depressions along base of elytron;

microreticulation fine, moderately impressed.

Pronotal margin narrowly beaded, elytron weakly

so.

Ventral surface. Evenly rugose-punctate,

punctures somewhat larger than on elytron.

Prothoracic process relatively narrow, parallel

sided, ridged, tip blunt. Metacoxal lines parallel in

posterior quarter, broadening to about 2 times

their narrowest width anteriorly, area between

them and forward onto mesosternum depressed.

Male. Protarsi moderately expanded, basal

segment round, third very deeply bilobed; single

claw relatively stout, bent at right angles, with

small basal tooth (Fig. 4). Mesotibia quite strongly

indented on inner margin near apex; mesotarsi

only slightly less expanded than protarsi.

18 C.H.S. WATTS & A. PINDER

6 7

FIGURES 1-9. 1, Dorsal view of median lobe of aedeagus of A. mcraeae; 2, ditto lateral view; 3, ventral view of

male metafemur of A. mcraeae; 4, lateral view of proclaw and apical tarsal segment of male A. mcraeae; 5, ditto A.

pennifoldae; 6, dorsal view of median lobe of aedeagus of A. pennifoldae; 7, ditto lateral view; 8, ventral view of

metafemur of male A. pennifoldae; 9, dorsal view of elytron of female A. pennifoldae.

Mesofemur considerably broader towards apex,

anterior margin near apex straight (Fig. 3).

Median lobe of aedeagus relatively simple but

with a thin lateral piece arising from near base on

both sides (Fig. 1).

Female. Unknown.

Remarks

The size, colour pattern, relatively broad

metacoxal lines with the area between them

depressed, male mesotibia indented and male

metatibia expanded, place this species in the A.

femoralis complex. Among these, the relatively

weak punctation, the shape of the male metatibia

and aedeagus ally it with A. pembertoni Watts. It

differs from this species in the shape of the male

proclaw which is a little more robust and the

central lobe of the aedeagus having an additional

piece on each side. In this unusual character it

resembles A. pembertoni which has a single

additional piece arising dorsally.

In the key given to Australian Antiporus in

Watts (1997) it (male) will run to the A. femoralis

complex. Further identification within this

complex depends on characters of the male

proclaw, metafemur and aedeagus (see Figs 1-4,

and those in Watts (1997).

Habitat

The single specimen was collected from a

NEW SPECIES OF ANTIPORUS 19

small, slightly acidic (pH 6.0), fresh (1.0 ppt)

Melaleuca swamp.

Etymology

Named after Jane McRae, the co-collector of

the specimen.

Distribution

Known only from the type locality in the wheat-

belt region of south-western Western Australia.

Antiporus pennifoldae sp. nov.

Types

Holotype: male; ‘Lake Poorginup, 62 km SE

Manjimup, 34°32'56"S 116°44'29"E, W.A. Coll.

A. Pinder and J. McRae (CALM), 2/10/98’.

Registration number WAM 26606. Western

Australian Museum.

Paratype: female; same data as holotype, South

Australian Museum.

Description (number examined, 2) (Figs 5—9)

Habitus. Length 3.3-3.4mm. Elongate-oval,

reddish-brown, appendages lighter.

Dorsal surface. Punctures dense, moderately

sized; those on head weaker and sparser, a little

smaller than eye facet; a row of large punctures

on elytron adjacent to suture, forming weak

groove. Pronotum and elytron with narrow but

well marked lateral beading. Microreticulation

fine, moderately impressed.

Ventral surface. Punctures similar to those on

elytron. Pronotal process blunt, sides weakly

bowed, moderately ridged. Postcoxal lines parallel

in apical quarter, weakly diverging to about 1.4

times narrowest width, area between them flat, not

depressed.

Male. Protarsi moderately expanded, proclaw

relatively stout, evenly curved with moderate

lateral rather than ventral basal spine (Fig. 5).

Mesotibia normal, mesotarsi similar to protarsi

except that third segment a bit shorter and two

claws present. Metafemur a little stouter than in

female with a small triangular spine on hind

margin in middle (Fig. 8). Apical third of elytron

weakly flanged. Median lobe of aedeagus with

asymmetric tip in dorsal view, distal portion

setiferous (Fig. 6).

Female. Protarsi weakly expanded, two claws.

Mesotarsi moderately expanded, more so than

protarsi. Metatibia simple. Elytron very widely

flanged, beginning in middle and expanding until

same width as rest of elytron near apex, then

ending abruptly, a short apical portion of elytron

not flanged (Fig. 9).

Remarks

A relatively small reddish species, the female of

which is instantly recognisable by the strongly

flanged elytra. In the male only a rather indistinct

narrow flange is present. The distinctly asymmetric

and setiferous median lobe of the aedeagus are

characters not shared by any other Antiporus.

In the key to Australian Antiporus in Watts

(1997) the females of A. pennifoldae can be taken

out at the start by the presence of strongly flanged

elytra. The males will run to couplet 15, where

they can be distinguished from both alternatives

by the presence of a small triangular spine on the

hind edge in the middle of the otherwise simple

metafemur.

The average size, relatively uniform reddish-

brown colour and essentially simple metafemora

suggest that the species does not belong in the A.

femoralis complex. It appears to be a rather

isolated species.

Habitat

Recorded only from a fresh (0.2 ppt), sedge-

filled peat swamp surrounded by Melaleuca

shrubs.

Distribution

Known only from the type locality.

Etymology

Named after Melitta Pennifold who was the

first to recognise the specific distinction of the

specimens.

ACKNOWLEDGMENTS

We thank Mr R. Gutteridge for the preparation of the

figures, Ms J. McRae for help in the field and Ms M.

Pennifold for help in the laboratory.

REFERENCE

WATTS, C. H. S. 1997. Four new species of Antiporus

Sharp (Coleoptera: Dytiscidae) from Australia, with

notes on A. femoralis (Boh.) and A. interrogationis

(Clark). Records of the South Australian Museum 30

(1): 35-42.

WHY FLUTES ON BOOMERANGS AND THROWING STICKS?

BY R.C. NELSON

Summary

NELSON, R.C. (2000). At first glance it may seem that the longitudinal flutes on the distinctive

throwing sticks and hunting boomerangs of Central and Western Australia must diminish their

aerodynamic performance. Nothing could be further from the truth. These surface features enhance

the performance. Wind tunnel tests show large drag force reductions in the case of throwing sticks

and large lift force increases in the case of boomerangs. The fluting alone is responsible for the

improved performance. In fact it can be shown that, without the fluting, some boomerangs may not

have flown at all.

WHY FLUTES ON BOOMERANGS AND THROWING STICKS?

R. C. NELSON

Nelson, R. C. 2000 Why Flutes on Boomerangs and Throwing Sticks? Records of the South

Australian Museum 33(1): 21-27.

At first glance it may seem that the longitudinal flutes on the distinctive throwing sticks and

hunting boomerangs of Central and Western Australia must diminish their aerodynamic

performance. Nothing could be further from the truth. These surface features enhance the

performance. Wind tunnel tests show large drag force reductions in the case of throwing sticks

and large lift force increases in the case of boomerangs. The fluting alone is responsible for the

improved performance. In fact it can be shown that, without the fluting, some boomerangs may

not have flown at all.

Ray C. Nelson, School of Civil Engineering, University College (UNSW), Australian Defence

Force Academy. Manuscript received 30 August 1999.

NOTATION

Projected area onto a plane normal to V.

Projected area onto a plane parallel to V.

Drag coefficient (dimensionless).

Lift coefficient (dimensionless).

Diameter.

Drag force.

Lift force.

Roughness height.

Reynold’s number (dimensionless)

Fluid free stream velocity.

Fluid density.

Fluid dynamic viscosity.

HD <P FT ONAS>

INTRODUCTION

The aerodynamics of boomerangs has received

considerable attention, particularly those of

returning boomerangs. One of the earliest attempts

to explain the reason why boomerangs return was

that of Wilkinson (1841: see reprint 1973:250—

254). This was a totally inadequate explanation. It

disregarded the fact that the vanes of the

boomerang were airfoils that created lift, and it

overlooked the gyroscopic nature of a rotating

boomerang and the consequent effects of the

precessional moment created by the lift force

differential between the two ends of the

boomerang. The first rigorous and complete

aerodynamic study of boomerangs was that of

Walker (1898). His analyses were not limited to

returning boomerangs. He was able to explain

why boomerangs which had strongly developed

convex curvature on both faces (the more curved

surface uppermost when thrown) and which also

had a small negative twist, produced the required

characteristics of a non-returning boomerang if

thrown with rotation in a horizontal plane.

Returning boomerangs often have positive twist

and are thrown with the rotation in a near vertical

plane.

The best known study of boomerang

aerodynamics is that of Hess (1968). It deals

exclusively with returning boomerangs. The main

contribution was a theoretical study of the

boomerang flight path with comparisons to

photographic records of boomerang flights. In a

later work, Hess (1975:545-549) did provide a

brief theoretical analysis of some characteristics

of the non-returning boomerang. Cotteral and

Kamminga (1990:175-180) drew heavily on the

work of Hess (1968) and did extend their work to

a brief consideration of the non-returning

boomerang.

No published work is available that relates

specifically to the aerodynamics of throwing

sticks used in Aboriginal Australia. However, the

discussion of maximum and minimum principal

axes by Cotteral and Kamminga (1990:179-180)

and the flight stability provided by the bend in a

boomerang, applies equally well to the flight

stability provided by the bend in a throwing stick.

The bend prevents spin about the longitudinal

axis, eliminating any undesired flight

characteristics that might result from phenomena

such as the Magnus effect (Roberson and Crowe

1997:446-447).

In recent times, Western fascination for the

returning boomerang has been responsible for hi-

tech design innovations used in modern sporting

22. R. C. NELSON

boomerangs but these are not relevant to this

study. The above summary does therefore

represent the current aerodynamic knowledge

applicable to the traditional throwing sticks and

boomerangs once used in Aboriginal Australia.

What is clear is that while the impact of surface

roughness on the performance of these

implements has been the subject of some

conjecture (e.g. Hess 1975:31, 37), it has never

been tested and quantified. Roughness elements

can have a profound influence on the

characteristics of the boundary layer as air flows

over the surface. This in turn determines the

pressure distribution around the implement and

the size of the wake formed behind the

implement. That is, the behaviour of the boundary

layer will determine the lift and drag forces on the

implement, which in turn directly affect its

performance. It is not only a difference between a

rough and a smooth surface that is important but

the size and type of the roughness is also

significant.

On relatively large (dimension parallel to the

flow greater than about 1 m if air is the medium),

highly streamlined objects, with very gentle

surface curvatures, it is necessary to have the

surface as smooth as possible to reduce the skin

friction drag. However, on relatively small objects

(dimension parallel to the flow of less than 100

mm if air is the medium) with relatively tight

surface curvatures, significant improvements in

the aerodynamic performance can be obtained by

the introduction of surface roughness. This results

in drag force reduction and lift force increase and

is technology used in modern every day life. For

example, the dimples on golf balls greatly reduce

the drag forces resisting flight, thereby increasing

the distance a well hit ball can travel by about 50

percent (Fox and McDonald 1998: 449-451,

Young et al. 1997: 410-412). Further, the

roughness produced by the stitching on a smooth

shiny cricket ball, enables a well bowled ball to

generate a lift force normal to the flight path and

produces what is known as swing (Brown and

Mehta 1993). This is done by slanting the seam at

an angle to the direction of travel so that on one

side only does the air pass over the roughness of

the stitching to create a turbulent boundary layer.

The reason for these aerodynamic

improvements is that the roughness elements

initiate transition from a laminar boundary layer

to a turbulent boundary layer closer to the leading

edge than would occur naturally on a smooth

surface. This natural transition would not

normally occur within 100 mm of the leading edge

of a smooth, streamlined object moving in air at

subsonic velocities. If a smooth, streamlined

object has a dimension parallel to the flow of less

than 100 mm, the boundary layer can only be

turbulent if it is initiated by roughness elements.

A turbulent boundary layer adheres longer to the

surface, into what is an adverse pressure gradient

on the rear portion of the object, before the

boundary layer separates from the surface. This

creates a narrower wake resulting in a smaller

drag force. It also reduces the overall pressure

force on that surface to below that which would

FIGURE 1. Typical specimens of hunting boomerangs

and fluted throwing sticks.

FLUTES ON BOOMERANGS 23

24mm

FIGURE 2. Cross-section of smooth foil.

exist if the boundary layer remained laminar.

Refer to Street et al. (1996:231-256, 489-418)

for further information on boundary layers, drag

forces and lift forces.

It is now a small step to suspect that the fluting

on the distinctive throwing sticks and hunting

boomerangs of Central and Western Australia may

have a significant utilitarian purpose. That is, a

purpose other than just decorative, ceremonial or

sacred in nature, namely aerodynamic. Typical

specimens of these artefacts are shown in Fig. 1

while Davidson (1936) has delineated the region

where these implements were in use. The

specimens shown were collected during the 1920s

and belong to the B.L. Hornshaw Collection. Both

implements are 640 mm along their axes. The

stick has a mass of 521 g (weight 5.11 N), is

circular in cross section and has an average

diameter of 30 mm. There is longitudinal fluting

around the whole circumference. The boomerang

has a mass of 314 g (weight 3.08 N) and has a

lenticular cross-section that is 48 mm in average

width and 12 mm in average maximum thickness.

Both faces of the boomerang have a convex

curvature, the longitudinal fluting being only on

the more highly curved face. In both cases the

flutes are about 5 mm wide and 0.2 mm deep. The

author perceived that the fluting on the throwing

stick was analogous to the dimples on golf balls

(drag reduction) while the flutes on the

boomerang were analogous to the stitching on a

cricket ball (lift enhancement), particularly as the

fluting is on one face only.

4mm 8mm

A series of experiments were designed and

undertaken to test these hypotheses. It is these

experiments and their results that are the subject

of this paper.

EXPERIMENTAL SETUP

The experiments were undertaken in a low

turbulence wind tunnel with a square test section

of side length 457 mm. Wind speeds were

measured using a Pitot tube connected to a micro-

manometer. The maximum wind speed possible in

the tunnel was 37 m/s. Models of the implements

were supported in the test section by a balance

system capable of measuring both the lift and drag

forces exerted on the models. The models were

455 mm long with cross sections modelled at full

scale. The models were located at mid height in

the tunnel with the longitudinal axis normal to the

airflow.

The models were manufactured from

aluminium on a computer controlled milling

machine to dimensions representative of the actual

implements shown in Fig. 1. Both smooth and

fluted models were manufactured. The selected

diameter of the throwing stick was 30.5 mm. The

selected cross section of the boomerang (smooth,

without flutes) is shown in Fig. 2. The fluted

models were made from smooth models using a

15 mm radius router to manufacture grooves 5

mm wide and 0.214mm deep. Photographs of the

fluted models are shown in Fig. 3 and Fig. 4.

24 R. C. NELSON

FIGURE 3. Fluted cylinder model.

THROWING STICK EXPERIMENTS

A throwing stick approximates a circular

cylinder for which considerable experimentation

has already been undertaken, especially for

smooth cylinders. Provided the throwing stick is

bent, and therefore prevented from spinning about

its longitudinal axis, the only aerodynamic force

exerted is that of drag. The magnitude of the drag

force is given by Eq. 1 where C, is the

dimensionless drag coefficient and A, is the

projected area onto a plane normal to the airflow.

F, =C,4, (1)

Clearly, the larger the drag coefficient, the

larger the drag force at any given velocity. If the

value of C, is known, the drag force on any

diameter cylinder, travelling at any velocity in any

fluid can be determined from Eq. 1. However, if

C, is not already known it must be determined

experimentally. This can be done using one fluid,

one cylinder of given D and series of test runs

using a range of V. A, and p would be known, V

and F', would be measured and hence C, derived

from Eq. 1.

The value of C, is also related to R,, a

dimensionless number defined in Eq. 2 that can

also be evaluated from the experimental

procedures described above.

_ pVD

The function of C, versus R, for a smooth

circular cylinder is given in Fig. 5 after

Schlichting (1968). The near horizontal portion

of the curve at C, values of about 1.2 is the

region where the boundary layer is laminar. At

R, values between 2x10° and 4x10° the boundary

layer goes through transition from a laminar to a

turbulent boundary layer until at R, values greater

than about 4.5x10°, the boundary layer is

completely turbulent and a very significant

reduction in C, is observed. The advantage of

having a turbulent boundary layer is obvious.

Unfortunately, this could never be achieved on a

30.5 mm diameter, smooth throwing stick, as it

would require airflow velocities in excess of

about 200 m/s (720 km/hr). The highest value of

R, possible in the wind tunnel was 7x10* for a

stick diameter of 30.5 mm.

Achenbach (1971) showed how roughness

modified the function of C, versus R,. The results,

shown as the dotted lines in Fig. 5, indicate that

the introduction of surface roughness induces

transition to a turbulent boundary layer at much

lower values of R, than would occur for a smooth

cylinder of the same diameter. The k/D values are

the relative roughnesses of the cylinders tested,

larger values representing rougher surfaces in

relative terms, the roughness type used being sand

grains. For example, k/D = 0.0090 is equivalent to

R (2)

FIGURE 4. Fluted foil model.

FLUTES ON BOOMERANGS 25

Smooth

_. k/D=0.0090

= k/D=0.0045

\ k/D=0.0011

FIGURE 5. Cd versus Re functions for cylinders.

a sand grain roughness diameter of 0.27 mm on a

30.5 mm diameter cylinder.

The results for the 30.5 mm diameter fluted

cylinder are shown in Fig. 5 as the heavy solid

line and indicate that the fluted roughness is

equivalent to something greater than a sand grain

roughness of 0.27 mm. The minimum drag

coefficient occurs at R, equal to 4x10*, one order

smaller than that for a smooth cylinder.

The dimensionless functions shown in Fig. 5

are universally applicable to any cylinders in any

fluid. The impact of flutes on a specific aboriginal

throwing stick is best illustrated in Fig. 6 where,

for any given velocity less than 37 m/s, the ratio

° Cylinders in air,

= dia. 30.5 mm.

pl Oe

oe)

et ade

wo |

cos

[Ess

Sy ! !

(=)

10.0 20-0 50:0 40.0

Velocity (m/s)

FIGURE 6. Drag force ratios — fluted cylinder/smooth

cylinder.

of the drag force on a fluted stick to that on a

smooth stick of the same length and diameter, is

given. Drag force reductions of up to 40 percent

are achieved at velocities of between 20 m/s and

30 m/s with reductions of 20 percent at velocities

of 40 m/s. These velocities are achievable by

throwing. The velocity of translation of a thrown

stick on leaving the hand was probably between

20 m/s and 30 m/s with a superimposed rotation

of about 5 rps (revolutions per second) adding a

further 10 m/s at the extremity of the forward

moving arm. The flutes appear to be the exact

type of roughness required to obtain maximum

drag force reduction on aboriginal throwing sticks.

BOOMERANG EXPERIMENTS

The lenticular cross section of a boomerang,

with one surface more highly curved than the

other, enables the arms to behave as rudimentary

airfoils that will generate a lift force in a direction

normal to the direction of the air stream. Drag

forces are also generated but, due to the more

streamlined nature of the cross section, they are

very much smaller than those generated by the

throwing stick. They are of secondary importance

to the lift forces in the context being considered

here. The lift force counteracts the gravity force

and prolongs the distance over which the

boomerang can be propelled. It is this ability to

generate lift that distinguishes a boomerang from

a throwing stick.

The magnitude of the lift force is given by Eq.

3, which in form is very similar to the equation

F, =C,4, 7 (3)

for drag force (Eq. 1) but with different

definitions of some terms. C,is the dimensionless

lift coefficient and A, is the projected area of the

airfoil onto a plane parallel to the airstream

direction.

Clearly, the larger the lift coefficient the larger

the lift force at any given velocity. However, as

with the cylinder, if the lift coefficient is unknown

it must be determined experimentally. With A_ and

p known and V and F, measured, the value of C,

can be derived.

The value of C, varies with the angle of attack,

the angle between the chord of the airfoil and the

direction of the airstream. The results for the

airfoils tested are given in Fig. 7 and show the lift

coefficient to be much larger for the fluted foil

26 R. C. NELSON

1.0

Foils

Lift Coefficient

FIGURE 7. Lift coefficient versus angle of attack for

foil models.

than for the smooth foil at all angles of attack

tested. The smooth foil did not generate positive

lift (upward lift) until the angle of attack exceeded

about +5 degrees. The angle of attack for most of

the length of a hunting boomerang is zero degrees.

A small negative twist (negative angle of attack)

is sometimes placed on the outer tip to lessen the

lift force in that region to reduce the precessional

moment that might otherwise cause the

boomerang to climb excessively (Walker 1898

and Cotteral and Kamminga 1990:180).

In the absence of flutes, the boomerang

modelled would not have flown at all as

evidenced by the very significant negative lift

coefficient (-0.5) at zero angle of attack. Flow

visualisation techniques indicated that on the more

gently curved underside, flow separation was

delayed until very near the trailing edge, while

flow separation occurred early on the rear half of

the more highly curved topside. The result was an

efficient negative (downward) force on the

underside but a lesser, inefficient, positive

(upward) force on the topside, the sum total being

a negative (downward) lift force.

The flow separation on the topside of the fluted

foil was delayed until very near the trailing edge,

thereby generating an efficient, positive (upward)

force that exceeded the magnitude of the negative

(downward) force on the smooth underside, the

sum total being a net positive (upward) lift force.

The flutes induce a turbulent boundary layer that

can adhere longer to the more tightly curved

Foils In air,

o _ chord length 48 mm,

— | zero angle of attack.

cured

(oe 1 | }

~ 1 ee ite 30. 40.

= ~~ Velocity (m/s)

oe Is

ae E Mo

| “<6

FIGURE 8. Lift forces on foil models.

surface, as described in the Introduction. It should

be noted that roughness on the underside would

have little impact on the lift characteristics. The

airflow was able to adhere satisfactorily to the

more gently curved, smooth surface without the

assistance of roughness. Therefore, the fact that a

boomerang may have some roughness on the

underside (adze marks for example) is not

important.

Fig. 8 shows the overall lift forces generated on

the two models, while Fig. 9 shows the increase

in lift force generated by the presence of flutes.

The values are given in Newtons per metre length

Foils in air,

chord length 48 mm,

zero angle of

es A attack.

o = IP

2 fo | |

OE ZOK 30, 40.

Velocity (m/s)

FIGURE 9. Increase in lift force attributable to flutes.

FLUTES ON BOOMERANGS 27

of foil (N/m). A hunting boomerang typically

weighs between 4.5 N/m and 5.0 N/m. Therefore,

the fluted foil tested has the lift characteristics

required of a boomerang launched at a velocity of

translation of between 20 m/s and 30 m/s and a

rotation of about 5 rps, values thought to be

characteristic of those actually used in Aboriginal

Australia (refer to section Throwing Stick

Experiments).

COMMENTS AND CONCLUSIONS

The testing of fluted and non-fluted samples of

throwing sticks and boomerangs has shown that

their aerodynamic performance is notably

improved by the fluting. The wind tunnel tests

show large drag force reductions for throwing

sticks and considerably increased lift forces for

boomerangs. The fluting alone is responsible for

this improved performance. The fluted boomerang

tested appears to have the optimum lift

characteristics. Without the fluting it has negative

net lift and therefore, becomes less efficient than

a throwing stick which has zero net lift. The flutes

on the throwing stick seem to be the optimum

roughness to maximise drag reduction benefits.

The use of roughness elements in pre-European

Australia (flutes in this case) appears to be

consistent with their use in contemporary times as

a technology for enhancing aerodynamic

performance. Fact must, however, be separated

from conjecture. It is fact that the fluting

improved the performance of these artefacts as

hunting tools and fighting weapons. As to whether

there was an empirical process that resulted in a

conscious decision to keep the flutes for this

purpose is in the realm of conjecture, because the

same flutes are on items that had no aerodynamic

requirements, such as wooden bowls. However,

there is the possibility that, along with many early

inventions, the fluting was a serendipitous

discovery implemented for advantage.

If the fluting on a boomerang created too much

lift near the outer tip, a larger than desired

precessional moment would be generated and the

boomerang would climb excessively. This can be

remedied by omitting the fluting near the tips.

Perhaps this accounts for those boomerangs

similar in all respects to that shown in Fig. 1, but

with fluting only over about the middle one third

of the longitudinal length. The precessional

moment can also be reduced by the negative twist

often present on hunting boomerangs.

REFERENCES

ACHENBACH, E. 1971. Influence of surface roughness

on the cross-flow around a cylinder. Journal of Fluid

Mechanics 46(2): 321-355.

BROWN, B. & MEHTA, R. 1993. ‘The Seamy Side of

Swing Bowling.’ New Scientist, 139(1887): 21-24.

COTTERAL, C. & KAMMINGA, J. 1990. ‘Mechanics

of Pre-Industrial Technology.’ Cambridge:

Cambridge University Press.

DAVIDSON, D. S. 1936. Australian throwing sticks,

throwing clubs, and boomerangs. American

Anthropologist, New Series 38(1): 76-100.

FOX, R. W. & McDONALD, A. T. 1998. ‘Introduction

to Fluid Mechanics.’ Sth ed. New York: John Wiley

& Sons, Inc.

HESS, F. 1968. The aerodynamics of boomerangs.

Scientific American 219(5): 124-136.

HESS, F. 1975. ‘Boomerangs, Aerodynamics and

Motion.’ Doctoral Thesis, Rijksuniversiteit Te

Groningen.

ROBERSON, J. A. & CROWE, C. T. 1997.

‘Engineering Fluid Mechanics.’ 6th ed. New York:

John Wiley & Sons, Inc.

SCHLICHTING, H. 1968. ‘Boundary Layer Theory.’

6th ed. New York: McGraw Hill.

STREET, R. L., WATTERS, G. Z. & VENNARD, J. K.

1996. ‘Elementary Fluid Mechanics.’ 7th ed. New

York: John Wiley & Sons, Inc.

WALKER, G. T. 1898. On boomerangs. Philosophical

Transactions Royal Society London, Series A 190:

23-41.

WILKINSON, H. 1841. ‘Engines of War.’ London:

Longman, Orme, Brown, Green, and Longmans.

Reprinted in Richmond by Richmond Publishing Co.

Ltd., 1973, with new introduction by W. S. Curtis.

YOUNG, D. F., MUNSON, B. R. & OKIISHI, T. H.

1997. ‘A Brief Introduction to Fluid Mechanics.’

New York: John Wiley & Sons, Inc.

REVISION OF AUSTRALIAN CHAETARTHRIA STEPHENS

(COLEOPTERA: HYDROPHILIDAE).

BY C.H.S. WATTS

Summary

WATTS, C.H.S. (2000). The genus Chaetarthria in Australia is revised. The synonymising of

Chaetarthria australis Knisch and C. sjostedti Knisch with C. nigerrima (Blackburn) by Balfour-

Browne is confirmed. This species is the only one present in Australia. It is patchily distributed in

coastal northern and eastern Australia.

REVISION OF AUSTRALIAN CHAETARTHRIA STEPHENS

(COLEOPTERA: HYDROPHILIDAE)

C. H.S. WATTS

WATTS, C. H. S. 2000. Revision of Australian Chaetarthria Stephens (Coleoptera:

Hydrophilidae). Records of the South Australian Museum 33(1): 29-31.

The genus Chaetarthria in Australia is revised. The synonymising of Chaetarthria australis

Knisch and C. sjostedti Knisch with C. nigerrima (Blackburn) by Balfour-Browne is

confirmed. This species is the only one present in Australia. It is patchily distributed in coastal

northern and eastern Australia.

C. H. S. Watts, South Australian Museum, North Terrace, Adelaide, South Australia 5000.

Manuscript received, 22 October 1999.

The hydrophilid genus Chaetarthria Stephens,

1835 contains numerous species, predominantly

in the Neotropical and Neartic regions (Hansen

1991). The extensive New World fauna has been

reviewed by Miller (1974) and Spangler (1977).

In comparison the Australian fauna is

depauperate with only one recognised species, C.

nigerrima (Blackburn, 1891), although two

others, C. sjostedti Knisch, 1922a and C.

australis Knisch, 1922b, have been described.

None are described from nearby Indonesia or

New Guinea but I would expect the genus if not

the species to also occur there. Apart from the

original descriptions nothing has been written on

the Australian species.

The genus belongs in the tribe Chaetarthriini,

readily recognised by their globular shape and

either divided eyes or the first and second

ventrites having a large cavity normally filled with

a hyaline mass and covered by long setae arising

from the front edge of the first ventrite (Hansen

1991). The tribe is represented in Australia by two

genera: Amphiops and Chaetarthria. From

Amphiops, Chaetarthria can readily be separated

by their undivided eyes, small size (< 2.0 mm.)

and, in the Australian species at least, virtual lack

of punctures.

The collections from which specimens were

examined are listed under the following

abbreviations:

ANIC Australian National Insect Collection,

CSIRO, Canberra

BMNH Natural History Museum, London

MV Museum of Victoria, Melbourne

NTM _ Northern Territory Museum, Darwin

NRS Naturhistoriska Riksmuseet, Stockholm

SAMA South Australian Museum, Adelaide

DPIM Queensland Department of Primary

Industries, Mareeba.

UQIC University of Queensland Insect

Collection, Brisbane

SYSTEMATICS

Genus Chaetarthria Stephens, 1835

Chaetarthria nigerrima (Blackburn,1891)

Paracymus nigerrimus Blackburn, 1891.

Chaetarthria nigerrima (Blackburn, 1891); J.

Balfour-Browne, 1938; Gentili, 1993.

Chaetarthria australis Knisch, 1922b; J. Balfour-

Browne, 1938.

Chaetarthria sjostedti Knisch, 1922b; J. Balfour-

Browne, 1938.

Types

Paracymus nigerrimus Blackburn. Lectotype:

‘Australia Blackburn Coll B.M. 1910-236’

‘Paracymus nigerrimus, Blackb’ ‘T 3566’ BMNH.

Blackburn gave the locality as ‘Mountains of

Victoria’ in his original paper. Herein designated.

Chaetarthria sjostedti Knisch. Lectotype: ‘Ma-

landa’ ‘Queensl. Mjoberg’ ‘Type’ ‘Chaetarthria

Sjostedti m. Nsp. A.Knisch 1921’ ‘5348 E91 +’

with red TYPUS label, NRS. Herein designated.

Paralectotypes: 1, ‘Ma-landa’ ‘Queensl.

Mjoberg’ ‘5347 E91+, (missing head and thorax),

NRS. 1, ‘3566’ ‘Victorian Alps Blackburn’

‘Paracymus nigerrimus, Blackb. Co-type’, SAMA.

Herein designated.

30 C.H.S. WATTS

Chaetarthria australis Knisch. Syntypes: Not

located. Type locality given as Gayndah,

Queensland. Knisch 1922b gives the locations of

the types as Museum Godeffroy No 10696 and

10701 and a further example in the Hamburg

Museum.

Description (number examined, 296) Fig. 1

Length 1.5-2.5mm. Round, deep bodied, height

of elytra a bit less than half length; dorsal surface

shiny black, sides of pronotum, edges of elytra

and apex of elytra light testaceous-yellow, ventral

surface dark-testaceous, appendages a little

lighter.

Dorsal surface: Head relatively narrow, sides

converging somewhat in front of eyes, finely

reticulate, very weakly punctate; labrum large,

front edge straight or very weakly concave, with

moderate punctures stronger than on rest of head.

Pronotum smooth apart from weak fine

microreticulation and sparse weak punctures at

sides and along front margin, disc virtually

impunctate. Elytron shiny, microreticulation weak,

about twenty rows of serial punctures (these are

very weak and hard to trace other than from the

ventral surface with transmitted light), sutural

groove strong in apical half, then progressively

weaker forwards but still traceable almost to

scutellum.

FIGURE 1. Dorsal view of aedeagus of Chaetarthria

nigerrima (Blackburn) from Millstream, W.A.

Ventral surface: Shiny, microreticulation weak;

metasternum with quite strong setae in middle;

first ventrite with confluent row of very long,

strong, golden setae along front edge reaching to

third ventrite, a similar row of much shorter setae

on rear edge of second ventrite pointing

backwards, short inward-pointing setae at sides of

second ventrite; other ventrites with quite dense

covering of short setae/spines except for front half

of third ventrite; epipleuron broad in front, rapidly

narrowing to middle, absent behind, almost in

same plane as sides of elytron.

Appendages: Maxillary palpi relatively short,

apical segment about 2.5 times length of

penultimate, with 3-5 small, elongate sensillae on

the outside at base. Protibia relatively narrow,

parallel-sided for most of length, with numerous

blunt spines towards apex but without modified

setae; procoxa with around eight strong spines on

ventral surface near base. Ventral surface of pro-

and mesofemora covered with short golden setae

except close to base, metafemur without such

setae except along front edge.

Male: Basal piece of aedeagus short, paramere

tips flared, truncated, apical portions and inner

edges semi-membranous and less chitinised than

rest of paramere (Fig.1).

Remarks

Chaetarthria nigerrima has a wide distribution

around northern Australia from the Pilbara region

of Western Australia to eastern Victoria. It is not

common but when present is often abundant in a

small area. I suspect that the species might be

semi-colonial. One such aggregation that I found

in the Northern Territory was living in small

tunnels in wet sand just above, and possible also

below, the waterline at the edge of a sandy pool in

a drying river bed. Other specimens are recorded

as having been collected among gravel at the

water’s edge. It also comes to light which is how

most of the specimens were collected. Miller

(1974) likewise recorded the genus in North

America as predominantly living in sand and

gravel at the edge of still or relatively still water,

and also flying to light. The larva of C. nigerrima

is unknown but that of the European C.

seminulum Herbst, 1797 has been described by

Bo6ving and Henriksen (1938).

Knisch, who was unaware of Blackburn’s

species, described two additional species, one

from Gayndah, Queensland and the other from

Malanda, Queensland. The type of P. nigerrimus

Blackburn and the type of C. sjostedti belong to

the same species. The types of C. australis would

AUSTRALIAN CHAETARTHRIA 31

appear to have been lost but there is nothing in

the description that would clearly distinguish it

from C. nigerrima and this and the fact that all

modern specimens appear to belong to the one

species lead me to agree with Balfour-Browne

(1938) that C. australis Knisch, 1922b is a junior

synonym of both C. nigerrima (Blackburn) and C.

sjostedti Knisch, 1922a.

Note on priority: The description of C. sjostedti

was published on the 24th of January 1922

(Knisch 1922a), not in 1921 as given in Knisch

1924. In June the same year Knisch published the

description of C. australis and at the same time

also reprinted his earlier description of C. sjostedti

(Knisch 1922b) as a new species.

Distribution

Australian Capital Territory: Bendora Dam,

ANIC; Black Mountain, ANIC. New South

Wales: 17 km SE Braidwood, ANIC; Cabbage

Tree Creek, Canberra-coast road, ANIC;

Chichester State Forrest, ANIC; Valery, ANIC.

Northern Territory: Bessie Springs, ANIC;

Nourlangie Creek, 20 km SSW Jabiru, SAMA; 19

km E by S Mt Borradaile, ANIC; Muirella Park,

Kakadu National Park, DPIM; U. D. P. Falls,

Kakadu National Park, NTM; Upper South

Alligator River, ANIC. Queensland: Bushland

Beach, 20 km N Townsville, SAMA; Bushy

Creek, Mossman-Mt Lewis road, ANIC; Cairns

District, SAMA; 25 km N Cooktown, ANIC; 30

m N Cooktown, UQIC; 70 km SW Greenvale,

SAMA; Henrietta Creek, Palmerston National

Park, UQIC; Iron Range, DPIM; Kennedy Creek

S of Laura, DPIM; 25 km N Laura, DPIM;

Kuranda, ANIC; 30 km W Laura, DPIM; 22 km S

Mareeba, DPIM; Millaa Millaa Falls, UQIC;

Mossman-Mt Lewis road near Julatten, ANIC; Mt

Surprise, DPIM; 22 km N Mt Molloy, ANIC;

North Pine River, UQIC ; Palmerston National

Park, UQIC; 15 km NNW South Johnstone,

DPIM; 20 km S Townsville, SAMA; Stewart

Range, SAMA; Walkamin, DPIM; Windsor

Tableland, DPIM. Victoria: Genoa, ANIC;

Meredith, MV; Victorian Mountains, BMNH.

Western Australia: Fitzroy River, ANIC;

Millstream, ANIC; 1 km N Millstream, ANIC;

Wittenoom Gorge, ANIC.

ACKNOWLEDGMENTS

I would like to thank the curators of the collections

listed earlier for allowing me ready access to specimens

in their care. Ms D. Churches put the final touches to

the manuscript, Mr R. Gutteridge prepared the figure

and the librarians, Mrs M. Anthony and Mrs J. Evans,

helped with the library references particularly the dating

of Knisch’s two publications.

REFERENCES

BALFOUR-BROWNE, J. 1938. A contribution to the

study of the Palpicornia.l. The Entomologist’s

Monthly Magazine 74: 102-107.

BLACKBURN, T. 1891. Further notes on Australian

Coleoptera, with descriptions of new genera and

species. Transactions of the Royal Society of South

Australia 14: 65-153.

BOVING, A. G. & HENRIKSEN, K. L. 1938. The

developmental stages of the Danish Hydrophilidae.

Videnskabelige Meddelelser fra den Dansk

Naturhistoriske Forening i Kjgbenhavn 102: 27-

162.

GENTILI, E. 1993. Paranacaena Blackburn, 1889: a

valid genus (Coleoptera, Hydrophilidae). Giornale

Italiano di Entomologia 6: 285-296.

HANSEN, M. 1991. The Hydrophiloid Beetles.

Phylogeny, Classification and a Revision of the

Genera (Coleoptera, Hydrophiloidea). Biologiske

Skrifter 40: 1-368.

KNISCH, A. 1922a. Results of Dr E. Mjéberg’s

Swedish Scientific Expeditions to Australia 1910—

1913. 29. Hydrophilidae. Arkiv fér Zoologi 14,

number 17: 4pp.

KNISCH, A. 1922b. Hydrophiliden-Studien (Op. 10.)

Archiv fiir Naturgeschichte 88: 87-126.

KNISCH, A. 1924. Hydrophilidae, in ‘Coleopterorum

Catalogus pars 79’. Eds. Junk and Schenkling.

Berlin, 306pp.

MILLER, D. R. 1974. Revision of the New World

Chaetarthria (Coleoptera: Hydrophilidae).

Entomologica Americana 49:1-123.

SPANGLER, P. J. 1977. Three new Ecuadorian species

of the aquatic beetle genus Chaetarthria (Coleoptera:

Hydrophilidae). Proceedings of the Biological

Society of Washington 90: 566-578.

WORDS TO OBJECTS: ORIGINS OF ETHNOGRAPHY IN COLONIAL

SOUTH AUSTRALIA.

BY PHILIP G. JONES

Summary

JONES, P.G. (2000). During the two decades preceding the establishment of the South Australian

Museum in 1856, colonial ethnography was an ill-defined project, influenced by the diverse aims of

social philanthropists, Christian missionaries and those intent upon creating optimum conditions for

the colony’s economic growth. This paper notes the significant role of individual ethnographers

such as Eyre, Grey, Angas and Cawthorne in producing descriptive ethnographies and in collecting

associated series of artefacts for a British, rather than a local Australian public. This factor slowed

any impetus towards a local ethnographically-oriented museum. The paper documents another

factor; the efflorescence in philological studies of South Australian languages during the 1830s and

1840s, which overshadowed local interest in assembling museum collections of ethnographic

material.

WORDS TO OBJECTS: ORIGINS OF ETHNOGRAPHY IN

COLONIAL SOUTH AUSTRALIA.

PHILIP G. JONES

JONES, P. G. 2000. Words to objects: origins of ethnography in colonial South Australia.

Records of the South Australian Museum 33(1): 33-47.

During the two decades preceding the establishment of the South Australian Museum in

1856, colonial ethnography was an ill-defined project, influenced by the diverse aims of social

philanthropists, Christian missionaries and those intent upon creating optimum conditions for

the colony’s economic growth. This paper notes the significant role of individual ethnographers

such as Eyre, Grey, Angas and Cawthorne in producing descriptive ethnographies and in

collecting associated series of artefacts for a British, rather than a local Australian public. This

factor slowed any impetus towards a local ethnographically-oriented museum. The paper

documents another factor; the efflorescence in philological studies of South Australian

languages during the 1830s and 1840s, which overshadowed local interest in assembling

museum collections of ethnographic material.

P. G. Jones, Department of Anthropology, South Australian Museum, North Terrace, Adelaide,

South Australia 5000. Manuscript received 1 November 1999.

PHILANTHROPISTS AND FRINGE DWELLERS

Meeting in London during 1834, the South

Australian Literary and Scientific Association had

shown an interest in Australian Aborigines even

before the new colony was proclaimed. Expressed

in lectures on the phrenological and physiological

character of the Aborigines, or their treatment at

the hands of British colonists, this interest had

less to do with ethnographic theory than with

colonialist practice. Phrenological technique was

perceived as a means, not only of discovering

more about a strange people for the sake of

science, but also of managing these encounters

more efficiently and, if possible, more humanely

than on the colonial frontiers of New South Wales

or Tasmania (Reece 1974: 85-93). South

Australia’s Adelphi planners were aware that their

treatment of the Aborigines would test their stated

humanitarian objectives for the new colony.

South Australia’s first Chief Justice stated the

case in these terms: ‘The system hitherto adopted

in the immediate neighbourhood of this Province,

towards the native population, is one at which

humanity shudders ... But such is not the system

which will be adopted towards them here, where I

trust, under Providence, that a new era is about to

dawn for them’. Interestingly, this opinion was

expressed in an 1837 court case in which two

white men had been charged with the theft of a

jacket and some ‘warlike implements (spears and

waddies) from some of the aboriginal inhabitants

of this Province’—one of the first documented

cases of direct appropriation of Aboriginal

artefacts in South Australia.2 This optimism

prevailed at least until 1839 when the Southern

Australian expressed the view that ‘[flor the first

time in the history of colonisation, the civilised

and uncivilised man have met without collision,

and emigration has brought with it a blessing

rather than a curse’.

By 1841, following the massacre of the ‘Maria’

shipwreck survivors and the Rufus River affray,

attitudes of South Australian colonists towards

Aborigines had noticeably hardened (Pope 1989).

Controversy surrounding the legality of the ex

judice executions of two Coorong Aboriginal men

for their part in the ‘Maria’ massacre helped to

crystallise the new status of South Australian

Aborigines as British subjects (ibid 144-45). With

the privileges attached to this legal standing, such

as the capacity to serve as witnesses in criminal

' Sir John Jeffcott, address to Grand Jury, quoted in the Register, 3 June, 1837, p.5.

2 Ibid: 4-5. The offence was committed in an Aboriginal hut at Glenelg.

> Editorial comment, Southern Australian, 16 June, 1838, p.4.

34 P. G. JONES

trials, came an entanglement of legal regulation

directed at defining and circumscribing Aboriginal

autonomy and movement in the Adelaide region.

As Colonel Light’s grid of streets and squares

took shape, so Aboriginal hunting and gathering

practices near settled areas became increasingly

problematic. Aborigines were drawn into the local

European economy and their previously unfettered

range was restricted to defined living areas such

as the ‘Native Location’ or semi-permanent camps

such as those at Glenelg or at Kensington.*

During 1847 a plan was drawn up to create

restricted areas within the city’s parklands for the

Aborigines and their ‘wurlies’.° Aboriginal

children were encouraged or coerced into the

missionaries’ day school, while their parents were

engaged as hewers of wood and fetchers of water.

As early as 1839 Robert Gouger had observed that

‘a little sugar, biscuit, or bread is sufficient

inducement for them to bring wood, water or

stone for building, and several instances have

occurred of ten or twelve of these poor fellows

working during six hours consecutively for an

individual for biscuit [sic]’.° During 1840 the

Adelaide Chronicle published an article

discussing a ‘scheme for the indoctrination of

industrious habits amongst the Aborigines’.’

Three years later, ‘the best means of civilising

Aborigines’ was discussed at one of the early

meetings of the Adelaide Literary and Scientific

Association and Mechanics Institute, the precursor

of the colony’s Library and Museum.®

Aborigines in colonial Adelaide had been

effectively marginalised by the later 1840s. The

frontier, and the danger which it represented to

the young and vulnerable capital, had receded

beyond everyday sight and discussion. This period

saw the image of the ignoble mendicant and the

“comic savage’ displace that of the noble savage

encountered by South Australia’s First Fleet.

Edward Snell’s first impression of Adelaide

Aborigines in 1850 reveals this shift: ‘Plenty of

natives stalking about the streets half naked—

most of the women with nothing on but a blanket

and nearly all of them the ugliest wretches it is

possible to conceive’ (Snell 1988: 49).

The progression in European depictions of the

Aborigine observed by Bernard Smith for

Sydney’s first two decades is also evident in

Adelaide (Smith 1984: 174-75, 220-21). With the

increasing tendency to reduce Aborigines to

Stereotypes in commentary, journalism and art,

came an inclination to apply more general policies

which took even less account of specific

Aboriginal requirements. Of course this trend was

not universal; the exotic character of Aboriginal

life remained an important ingredient of colonial

Adelaide for at least a decade after settlement.

Images of the noble savage continued to surface—

at the palti or kuri dances held at full moon, or on

the battlefield during periodic confontations

between the Adelaide, Encounter Bay, and River

Murray groups. Even these occasions were subject

to European surveillance or control; the

corroborees were usually attended by European

voyeurs (invited or not), while the battles were

often either terminated or forestalled altogether by

police intervention.? One of William Cawthorne’s

most telling watercolour depictions of the

Aboriginal subject at this time was his study of a

pile of decorated shields and spears, smashed by

police horses to prevent tribal fighting near

Adelaide.'° Cawthorne was sufficiently disturbed

by this event to write an article for the Register,

on behalf of one of the principal Adelaide elders,

King John (Mullawirraburka):

On Monday last, a fight was to have taken place

between Moorundee, Encounter Bay, and Adelaide

natives. Great preparations are accordingly made.

The young men were all in high glee—tattooed

[printed in error for ‘karkooed’, ochred], oiled, and

all ready for the coming amusement, but

unfortunately they were disappointed; for, as they

were marching to meet each other on the old Bay

See Foster 1991. For example, in June 1837 the Protector of Aborigines noted that he had removed ‘certain Aborigines who had settled on Dr Wright's

town acre’ and a few weeks later John Morphett complained that Aborigines were cutting down trees on his town acre (GRG 24/1/1837/210; 246). In

December 1838, the Superintendent of Police reported the ‘habit of the Aborigines of disfiguring trees in the Park Lands, yet during the same year colonists

were employing Aborigines to remove timber from the same areas (GRG 24/4/1838/20; 24/1/1838/8, SRO). For a broader discussion of this issue, see

Reynolds (1990: 129-163).

GRG 24/61847/440, State Records, South Australia

Gouger 1838: 56. By 1847 Adelaide Aborigines were being employed to break stones for paving the city’s streets (GRG 19 24/4/1847/273, State Records,

South Australia).

Adelaide Chronicle, 18 & 25 February, 1840, p.3; p.2.

The meeting canvassed a range of strategies, ranging from the establishment of a native police force to the dual operation of a ‘House of Correction’ and

a ‘Native Location’. See the Southern Australian 28 February, 1843, p.2; Adelaide Examiner 25 February, 1843, pp.3-4.

See Angas (1847a, vol.1: 102-108) and Leigh (1839: 143) for descriptions of corroborees attended by Europeans at this time.

Cawthorne Diary, Monday, 22 April, 1844, Foster (ed.) 1991: 46. The original drawing is contained in the Cawthorne mss., Mitchell Library, Sydney

WORDS TO OBJECTS: 35

road, three horse-police very unceremoniously

stopped them, and had every spear and shield laid

on the ground, and broken up. The astonishment that

this act produced, was truly remarkable—some

looked quite aghast, others were confounded, and

many for the moment, I dare say, doubted their

senses, whether such a collection of beautiful

uwindas and shields, kylahs and midlays, were

absolutely to be destroyed."

By the later 1840s these events, and Aborigines

themselves, had become no more than a

picturesque backdrop to the bustling activity of

colonial Adelaide. Already in 1843 Aborigines

were tried for ‘appearing naked’ in Gawler Place.

‘Sunday corroborees’ were finally forbidden

during 1847." The careful, individual portraiture

of George French Angas and William Cawthorne

was gradually supplanted by the caricatures and

drawings of S. T. Gill, Alexander Schramm, and

Edward Snell, or by landscape painting in which

‘the aborigine [was] relegated increasingly to

fulfil the function of a pictorial embellishment to

topographic landscape, providing a local touch

and pointing the contrast between primeval life

and the busy progress of the town’.

During the years following European settlement

the curiosity value of Aboriginal objects, like their

owners, steadily diminished. Familiarity bred

contempt. Some collections of Adelaide Plains

artefacts continued to make their way to Europe,

evoking, like trophy displays, the raw experience

of the colonial frontier, but little attention was

paid to the preservation or classification of such

artefacts in Adelaide itself during the 1840s or

1850s. A factor of some significance, particularly

in the light of the marked effect on local

collecting practice triggered by the demand for

Aboriginal artefacts for the International

Exhibitions of the 1870s, may lie in the British

Museum’s focus (and that of British collectors)

on the Assyrian antiquities, first acquired in 1849

(Bohrer 1994). The British Museum’s

preoccupaton with antiquities during the late

1840s and 1850s may well have been mirrored in

colonial Australia, to some extent.

Despite the pioneering roles of Governors

Gawler and Grey, the exigencies of colonial life

in Adelaide meant that the practice of ethnography

was important only so far as it contributed to the

colonists’ image of a new and strange land, or to

an overlapping phase in which it was necessary to

understand the Aborigines in order to control them

more effectively. The early descriptions of

Aboriginal life in South Australia and the

published lithographs of watercolour studies by

Angas catered for the first category of interest,

primarily directed to a British readership. The

ethnography of the missionaries and early

Protectors such as Wyatt and Moorhouse was

directed towards the more practical ends of the

second phase. Moorhouse’s Murray River

vocabulary (1846) was constructed well after his

Adelaide work, and was made at Grey’s behest. In

this sense it does not provide an exception to the

general rule, by which other ethnographers

followed their publications on grammar and

vocabulary with more discursive works on

manners and customs. Moorhouse and

Teichelmann produced a joint ‘Report on the

Aborigines of South Australia’ (published in

January 1842) and Moorhouse was solely

responsible for the ‘Annual Report of the

Aborigines’ Department’ (unpublished until 1991,

see Foster 1991). The concentration of

ethnographic research and publication in Adelaide

during this second phase was markedly higher

than in other colonies, a fact which throws the

decline in ethnographic enquiry during the 1850s

and 1860s into greater relief. Following the

publication of Moorhouse’s vocabulary in 1846,

eighteen years passed before the next South

Australian linguistic publication appeared—

George Taplin’s first work on Lower Murray

River languages (1864)."*

The reasons for this concentrated burst of

activity are two-fold. In the first place, because of

the concern expressed in England during the late

1830s by Lord Glenelg and the Colonial Office

about the mistreatment of indigenous peoples, and

partly because of the concentration on this issue

by the key individuals associated with the colony

of South Australia, Aboriginal matters were of

greater concern for South Australian policy-

"Register, 24 April, 1844, p.3. See also Adelaide Observer, 27 April, 1844, p.5.

"2 Southern Australian 5 September, 1843, p.2; GRG 24/4/1847/195; 205; 1430; 1462, State Records, South Australia.

3 Smith 1984; 220. Smith makes this point in relation to Sydney artists of the 1790s and 1800s; the same shift can be observed in Adelaide’ s colonial period,

three decades later. See also Dutton (1974).

Teichelmann produced an unpublished, extensive, revised version of his Adelaide vocabulary during this period; this was sent to George Grey in Cape

Town in 1857 (Teichelmann 1857 ms.). The vocabulary published by E. M. Curr in 1886 (Teichelmann, Schurmann & Wyatt 1886) is a much shorter

wordlist.

36 P. G. JONES

makers.'> This sensitivity accounts for the

extended but ultimately unsuccessful efforts to fill

the post of Protector of Aborigines with the

‘conciliator’ of Tasmanian tribes during the

1820s, George Augustus Robinson.'® It also

explains the trouble to which George Fife Angas

went to secure the services of Lutheran

missionaries prepared to learn the languages of

the South Australian Aborigines in order to impart

Christian principles to their children. In their

attempts to explore the mental world of South

Australian Aborigines, Teichelmann, Schurmann

and Meyer went further than assembling simple

word lists. Their expanded publications on the

‘manners and customs’ of Aborigines of Adelaide,

Port Lincoln and Encounter Bay followed their

scholarly work on the languages of these groups.

For example, Teichelmann and Schurmann’s

grammar and vocabulary of the Adelaide

Aborigines (1840) preceded Teichelmann’s work

on their manners and customs (1841) published in

the following year, and Schurmann’s vocabulary

of the Parnkala people of Spencer’s Gulf (1844)

preceded his general work on the Aborigines of

Port Lincoln (1846). Schurmann’s ethnographic

work with Port Lincoln Aborigines can be

regarded as part of his duties as Protector of

Aborigines there. Meyer’s vocabulary of South

Australian Aborigines (1843) preceded his work

on the manners and customs of Encounter Bay

Aborigines (1846).

PHILOLOGY AND “THE ABSTRACT SCIENCE OF MAN’

For a decade following the creation of the

colony in 1836, the level of South Australian

ethnographic activity exceeded that of any other

colonial outpost throughout the world. At least

thirteen vocabularies, four grammars and six

descriptive works on South Australian Aboriginal

manners and customs were produced. The

vocabularies were by Bromley, Stevenson and

Finlayson (unpublished and untraced); Gell

(Assistant Secretary to Governor Gawler) 1841;

Meyer (1843), Moorhouse (1846), Piesse (1840),

Schurmann (1844), Teichelmann (Teichelmann,

Schurmann and Wyatt 1886), Teichelmann and

Schurmann (1840), Weatherston (n.d.; see Hunt

(1985: 38)), Williams (1839), Wyatt

(Teichelmann, Schurmann and Wyatt 1886). The

descriptive ethnographies were by: Angas (1847a;

1847b), Cawthorne (1926), Eyre (1845), Grey

(1841a), Meyer (1846), and Schurmann (1846).

Schayer’s (1844) Berlin publication on the

language, manners and customs of South

Australian Aborigines was probably based on the

work of the Dresden missionaries. In addition, it

appears that other concerted ethnographic and

linguistic work was undertaken, such as that by

the surgeon Richard Penney in 1841 and by the

police-trooper and Sub-Protector George Mason

during the early 1840s (Lendon 1929: 24: & 31:

20-33). Proportionately, this level of output in

Australian linguistics has not been equalled

since.”

South Australia’s evolution as a colony

coincided with the early development of

ethnography as an internationally accepted

scientific discipline. The first documented use of

the term ‘ethnography’, to mean ‘the scientific

description of nations or races of men, their

customs, habits and differences’ was in 1834, the

foundation year of the South Australian Literary

and Scientific Association.'* From the beginning

though, ethnography was far more than a

descriptive exercise: like other branches of

Enlightenment natural science it held great

promise as a means of discerning the origins of

humankind. Australian Aborigines provided a new

and exciting field for ethnographers, but seldom

as a people to be studied in their own right. Their

primary ethnographic value lay in the data which

an analysis of their characteristics could contribute

to wider debates.

The focus of ethnographic study shifted back

and forth several times during the nineteenth

century, from physical description to philology, to

The correspondence files of the South Australian Company, particularly those of its Director, George Fife Angas, are full of references to this crucial

issue (PRG 174, Mortlock Library, Adelaide). See also further discussion of this point in Pope (1989). Following the recommendations of the 1837 ‘Report

from the Select Committee on Aborigines’ (1968 (1837)), Standish Motte (himself a member of the South Australian Literary Association), published

his ‘Outline of a System of Legislation for Securing Protection to the Aboriginal Inhabitants of All Countries Colonized by Great Britain’ (Motte 1840).

‘© John Brown, a founding committee member of the South Australian Literary and Scientific Association in London, was a cousin of the celebrated

missionary George Augustus Robinson who worked among Tasmanian Aborigines during the 1830s. During 1836 Brown agitated successfully for

Robinson to become South Australia’s Protector of Aborigines, but Robinson eventually decided against the position.

‘8 ‘The Shorter Oxford English Dictionary’: 685.

The 1960s-70s produced a very high volume of linguistic publications but from a much wider pool of specialists (J. Simpson pers. comm. 1991).

WORDS TO OBJECTS: 37

kinship studies and religion, to archaeology and

material culture. Internationally, the emphasis

during the first half of the century centred firmly

on philological research.'? This promised a level

of methodological certainty and a potential for

deductible results which other branches of

anthropology lacked. For newly arrived settlers in

the New World and Australia the question of how

the native inhabitants came to be there was of

particular interest. Posing this question in his

‘Notes on the State of Virginia’ in 1804, Thomas

Jefferson had predicted that language would offer

‘the best proof of the affinity of nations which

can ever be referred to’. Forty-two years later,

Henry Schoolcraft’s ‘Plan for the Investigation of

American Ethnology’, presented to the

Smithsonian Institution in 1846, stressed the

resilience of this notion:

Philology is one of the keys of knowledge which, I

think, admits of its being said that, although it is

rather rusty, the rust is, however, a proof of its

antiquity. I am inclined to think that more true light

is destined to be thrown on the history of the Indians

by a study of their languages than of their traditions,

or of any other feature (Jefferson and Schoolcraft

quoted in Hinsley (1981: 23).

Comparative philology had the added advantage

of offering a method for tracing the origins and

diffusion of the world’s peoples within the

relatively short chronology accepted within

biblical orthodoxy. South Australia’s third

Governor, George Grey, was a significant

contributor to debate on this subject, which

remained the ‘queen of the human sciences’ until

the late 1850s, when biblical chronology was

undermined by the emerging acceptance of

Darwinian theory, geological time and the new

‘prehistoric’ archaeology (Chapman 1985: 21—22;

Crawfurd 1863). In the meantime though, the

careful, scientific methodology of philologists

such as Johann Forster, Samuel Marsden, Lord

Monboddo, and Wilhelm von Humboldt

paralleled the techniques employed by the rising

generation of earth scientists, future evolutionists

included. Charles Darwin’s influential friend, the

geologist John Herschel, drew the analogy:

“Words are to the Anthropologist what rolled

pebbles are to the Geologist—battered relics of

past ages often containing within them indelible

records capable of intelligible interpretation’

(Desmond and Moore 1992: 215).

In Adelaide during the 1840s the science of

philology held similar promise for the colony’s

‘utopian socialists’. Addressing an audience of

1000 in the Queen’s Theatre, even the austere

Advocate-General, William Smillie, reserved a

prominent place for philological studies in

Adelaide’s cultural life:

The student of human nature will also find an

appropriate chapter on the aborigines. Many of their

peculiar customs and superstitions, the analogical

sense contained in their terms, and the structure of

their speech, illustrate the abstract science of man,

while etymologies may be collected to throw some

light on their origins and history, as connnected with

other races... (Smillie 1842: 437).

Another of South Australia’s ‘utopian

socialists’, Robert Gouger, devoted considerable

space to the Aborigines in his publication about

the colony, referring to the linguistic similarity

between South Australian Aborigines and the

‘Malays of Dampier Straits’ as perceived by Mr

Donovan, chief officer of the Katharine Stewart

Forbes (Gouger 1838: 52-53). The emerging

science of ethnography drew its data neither from

social traits nor from a comparison and analysis

of material culture, but from linguistic

‘specimens’. All the attributes of scientific

practice were satisfied by the new field of

linguistics. Given the correct direction and

technique, useful data could be gathered by

amateurs who would gain the reward of knowing

that each fragment collected would assist their

scientific mentors in constructing a new picture of

human origins. Unlike collectors of ethnographic

objects, whose arrays of curiosities still bore no

evident relation to European artefacts, amateur

and scientific linguists shared the conviction of

natural scientists that they were actively

collaborating in a grand scheme.

With the decline of philological research and

deductive enquiry after mid-century, more

tangible types of evidence seemed called for. The

historian William Chapman has noted that for

some, ‘these were the evidence of skeletal and

cranial forms, either as measured or uncovered

from the ground; for others, the evidence of

archaeology more generally; but for Pitt Rivers,

the privileged evidence was to be that based on

the comparison of artifacts’ (Chapman 1985: 22).

Even so, ethnographic collecting did not achieve

quick popularity. It was not until the final decades

See Gascoigne (1994: 160-176) for an account of philology within the Enlightenment context.

38 P. G. JONES

of the century that the role of Australian museums

as repositories for ethnographic objects came to

be validated in terms of the contribution which

those objects might make to the pursuit of

ethnography, or to science in general. During the

decade after South Australia’s proclamation, the

collection of linguistic information had been

validated in just those terms. As an ‘exact science

of mental objects’, philology provided the impetus

for the first systematic application of Western

scientific principles of classification to

Aborigines. As the French philosopher Renan put

it, philology was a scientific method to be used

for arriving at ‘the very system of things ... [it is]

... the exact science of mental objects. It is to the

sciences of humanity what physics and chemistry

are to the philosophic sciences of bodies (Renan

1890 (1848): 149).

The multitude of Aboriginal vocabularies and

basic grammars collected from the colonial

frontier during the middle years of the nineteenth

century can be contrasted with the relatively

casual collection of Aboriginal artefacts during

the same period. Susan Pearce has suggested that

such an inverse relationship between language and

objects may have been characteristic of this era,

implicit in the dual status of ethnographic objects

as both ‘real’ and ‘constructed’ artefacts:

The long-term trend of European thought,

increasingly cogently expressed from the late

seventeenth century onwards, is to give a low value

to the material world as such, and to regard it as the

fit place for the exercise of human reason and

enquiry through which real knowledge will be

constituted. On this reading, objects in general are

the passive result of social action, and museum

collections enshrine the results of objective enquiry

which has yielded real understanding; in other

words, the metaphorically constructed

understandings have been seen as superior to the

concrete, contextual reality of the things. An

important aspect of this is the tendency to regard

language, the prime medium for classification and

reason, as the faculty which creates social structure,

although as we have seen there is not an exactly

parallel relationship between language and the

material world (Pearce 1992: 257).

The rigour which characterised linguistic

studies in this early period has its direct analogue

in natural science, as Edward Said has observed:

Science gives speech to things; better yet, science

brings out, causes to be pronounced, a potential

speech within things. The special value of linguistics

(as the new philology was then often called) is not

that natural science resembles it, but rather that it

treats words as natural, otherwise silent objects,

which are made to give up their secrets. Remember

that the major breakthrough in the study of

inscriptions and hieroglyphs was the discovery by

Champollion that the symbols on the Rosetta Stone

had a phonetic as well as a semantic component. To

make objects speak was like making words speak,

giving them circumstantial value, and a precise

place in a rule-governed order of regularity (Said

1978: 140).

But this ‘circumstantial value’ was not attached

to ethnographic objects by museum scientists until

the third quarter of the nineteenth century. The

shift of status which eventually occurred during

that period is best symbolised by the replacement

of the descriptive term ‘curio’ by the phrase

‘ethnographic specimen’, carrying the implication

that the classifications and strategies applied to

natural objects by museum scientists could equally

be applied to artefacts. Until that point was

reached philology or linguistics remained the only

branch of ethnography accorded scientific status.

With few exceptions this field remained beyond

the confines of the newly established natural

history museums in Europe and Australia.

South Australia’s early ethnographic studies,

and similar researches in other Australian

colonies, have been enlisted by historians of

anthropology in describing the developing picture

of the country’s anthropological discipline (See

for example, McCall 1982; Mulvaney 1964; 1993;

Peterson 1990; Tindale 1986). In his subdivision

of Australian anthropological history, A.P. Elkin

placed the work undertaken during the colonial

period within an ‘incidental phase’, implying little

continuity with succeeding periods (Elkin 1970:

6; 1963). McCall (1982) and Peterson (1990)

accept this characterisation of early Australian

anthropology. While there is no doubt that later

ethnographers and anthropologists built upon the

results of this early work, the phase does bear a

distinct character, apart from the sense employed

by Elkin. The phase can be distinguished by the

pragmatic, utilitarian nature of the research which

it generated. Almost all of the ethnographic

pamphlets and booklets appearing in Adelaide

during the 1840s were published in the name of

science. Most of this output was directed towards

practical, short term ends; to facilitate the tasks of

administrators and missionaries in dealing with

Aborigines in the colonial situation. This early

ethnographic work can be regarded as a precursor

of the applied anthropology undertaken for

colonial administrations during the early twentieth

century.

Each of the early Protectors of Aborigines

WORDS TO OBJECTS: 39

produced vocabularies of the Adelaide Plains or

adjoining regions, not primarily as a contribution

to the growing international corpus of such

material, but as a means of undertaking their

assigned duties. The English Parliamentary Select

Commission on Aborigines (British Settlements)

of 1837 had officially recommended that

Protectors of Aborigines ‘should be expected to

acquire an adequate familiarity with the native

language’ and that ‘the Protectors should be

furnished with some means of making to the tribes

occasional presents of articles either of use or

ornament’ (‘Reports from the Select Committee

on Aborigines’. 1968 (1837), vol.2, p.83). By the

time that this directive was issued, South

Australia’s first two Protectors, George Stevenson

and Captain William Bromley, had already

prepared working vocabularies. By August 1837,

when Bromley was replaced by William Wyatt,

the colonial government had adopted the Select

Commission’s directive regarding Aborigines and

advised the new Protector that ‘no time should be

lost in acquiring a knowledge of their native

tongue’.”°

Wyatt and his successor Matthew Moorhouse

prepared vocabularies of the Adelaide and Murray

River Aborigines (Wyatt 1879, 1886; Moorhouse

1846).2! The Governor’s secretary wrote to Wyatt

in August 1839 following the Protector’s enforced

resignation, informing him that the governor ‘had

peculiar opportunities for observing the patient

and scientific research with which you have

investigated their language, and he has now in his

possession an extensive and very valuable

vocabulary of it compiled by you’ (Lendon (n.d.)

ms.: 212).

The Colonial Storekeeper, William Williams,

who also collected Aboriginal artefacts at this

time for the Colonization Commissioners,

recognised the utility of publications which might

assist government officials and employers to

benefit from the labour of ‘idle natives’, while

encouraging the civilising process. During early

1839 he compiled his own vocabulary of the

Adelaide Aborigines and offered this for sale to

subscribers, including Governor Gawler and other

government officials and notables.” Until his

removal from office in 1841, Gawler played a

vital role in sponsoring ethnographic studies of

this kind, encouraging and assisting the work of

the Protectors (notably Moorhouse), the Lutheran

missionaries, and Edward John Eyre.

The arrival of George Grey during 1841 to

replace Governor Gawler lifted South Australian

ethnography out of its short-term, utilitarian

mould. An influential anti-slavery and Church

Missionary Society advocate, the young cavalry

captain had already undertaken substantial

ethnographic and linguistic studies of Western

Australian Aborigines before his South Australian

appointment (Grey 1841a). On the basis of his

Western Australian experiences he had written

carefully on the issue of ‘promoting the

civilisation’ of Australia’s Aborigines (1841b). In

the year of his arrival in South Australia, and two

years before the foundation of the Ethnological

Society in London in 1843, Grey published a

paper on administering native peoples, his major

anthropological conclusions on Western

Australian Aborigines and a detailed dictionary of

South-Western Australian Aboriginal dialects

(Grey 1841a; 1841b; 1841c; see also Mulvaney

1964: 23-25). During his term as South

Australia’s Governor from 1841 to 1845, Grey’s

first interest remained Australian linguistics and

his 1845 paper on this subject summarised the

state of knowledge in this field, noting that South

Australian researchers had adopted a common

system of orthography (Grey 1845). The linguistic

evidence suggested to Grey that ‘this continent

was peopled from the north-west, and that the

lines of migration were along the coast and the

great water drainages of this country’ (ibid: 366).

Together with Threlkeld in New South Wales,

Grey was a leading Australian figure in

investigating the Indo-European hypothesis of

Aboriginal origins. His conclusions were

reinforced with the publication of J. C. Prichard’s

findings that the Australian language had affinities

with that of the Tamils of southern India

(Mulvaney 1964: 22). He was a strong critic of

inconsistent or inferior research, noting that ‘up to

the present time we have had only very meagre

vocabularies, collected by passing strangers, each

of whom adopted his own system of orthography,

and the comparisons formed from such

® Colonial Secretary's Instructions to Protector of Aborigines, South Australian Gazette and Colonial Register, 12 August, 1837, p.1.

Government Gazette, 7 June, 1839.

Although Wyatt's vocabulary was not published until 1879, it was available for use in government circles on its completion in 1839. See South Australian

Williams’ advertisement appears in the Southern Australian, 27 March, 1839, p.2. A copy of this vocabulary is listed among the contents of Grey's papers

held in the South African Library, Cape Town (Williams 1839). It was later published in Parkhouse (ed. 1923).

40 P. G. JONES

compilations must necessariy have been erroneous

in the highest degree’ (Grey 1841a, vol.2: 215-

16). He stressed the importance of establishing

consistent principles of orthography in recording

Aboriginal languages as a means of adducing

reliable evidence to support the Indo-European

hypothesis and in this respect his encouragement

of the linguistic work of South Australian

Lutheran missionaries was a major contribution.

This initiative began before his appointment as

Governor, during his first visit to Adelaide in

1840. Tindale (1974: 3) asserts that Grey guided

the Lutheran missionaries in the production of

their linguistic work. Some evidence for this lies

in Teichelmann’s acknowledgment of discussions

with Grey about Australian ‘dialects’

(Teichelmann & Schurmann 1840: vii, viii). But

Schurmann also recorded the fact that although

Grey gave valuable encouragement to

Teichelmann and himself in publishing their

grammar and vocabulary, it was Moorhouse who

assisted in reading and editing their English text

(Schurmann 1987: 104; Teichelmann &

Schurmann 1840: viii). In fact, it is possible that

Grey incorporated some of Teichelmann’s and

Schurmann’s vocabulary within his own 1841

publication without acknowledgement (Grey

1841a: 212-15). Grey was also quick to adopt the

modified Threlkeld ‘South Sea’ orthography used

by Teichelmann and Schurmann in their work, and

argued for its adoption by the Royal Geographical

Society (J. Simpson, pers. comm. 1991).

Until Grey’s arrival in South Australia the

colony had no ethnographer of equivalent

standing to Threlkeld, who combined local,

practical linguistic studies with a commitment to

ethnography as a developing international science

(Threlkeld 1834: 185). By promoting

ethnographic enquiry as a branch of science linked

to other fields of philosophical enquiry, Grey

helped to give the study of Aboriginal manners

and customs a new relevance in the colony. His

significance in Australian anthropological history

lies in the fact that he bridged the gap between the

applied ethnography of colonial administrators

and the more scholarly, enquiring approach

promoted by the Royal Society, subsequently

adopted by both the Royal Geographical Society

and the Anthropological Society of London.

Grey’s style of ethnography, like that of the

explorers Charles Sturt and Edward Eyre, had its

origins in the British naval and exploration

tradition. The Royal Society’s ‘Directions for

Seamen, Bound for Far Voyages’ were first

published in 1665. Cook’s instructions, based on

these directions, included the obligation to

‘observe the Genius, Temper, Disposition and

Number of the Natives’ (Smith 1984: 8,16). The

earliest opportunity for British ethnographic study

in South Australia arose on Matthew Flinders’

voyage of exploration in 1801-2. In view of his

thorough investigation of northern Australian

Aboriginal implements, rock paintings and burial

modes, it is unfortunate that Flinders had little

contact with Aborigines in South Australia. The

single vocabulary published in his expedition

report was obtained at Caledon Bay, on the west

coast of the Gulf of Carpentaria (Flinders 1886).

The French voyage of Baudin and Peron to

Australia coincided with Flinders’ voyage and the

two expeditions met in Encounter Bay on April 8,

1802. But the French and British observations of

South Australian Aborigines were minimal in

comparison to the wealth of data on Tasmanian

Aborigines recorded by Baudin’s expedition. As

Rhys Jones has explained, during this period the

French displayed a much greater commitment than

the British to anthropological research, through

the Societé des Observateurs de 1’ Homme. This is

despite the fact that the Baudin expedition rarely

approached the standards set in Degerando’s

detailed anthropological instructions and

Suggestions (Degerando 1969) or by Cuvier

(quoted in Jones, R. 1988: 37).?3

The earliest record of South Australian

Aboriginal languages was made in Western

Australia, by a colleague of Dumont d’Urville

during his voyage in the Astrolabe (1826-29).

During his 1829-1830 Murray River expedition

the explorer Charles Sturt made little use of his

numerous opportunities to record Aboriginal

language and customs, despite having been

directed to ‘note the description of the several

people whom you may meet, the extent of their

population, their means of subsistence, their

genius and disposition, the nature of their

amusements, their diseases and remedies, their

objects of worship, religious ceremonies, and a

For a useful discussion of the distinction between French and British anthropology at this time see Gascoigne (1994: 158-159).

4 Dumont d’Urville 1830-35. Dumontd’ Urville obtained his word list froma colleague, Gaimard, who had recorded several words froma South Australian

Aborigine in Western Australia, possibly brought there by sealers (Simpson, 1996). See Gaimard (1833) for this vocabulary. Dumontd'Urville also made

an ethnographic collection in Australia, identified by Sylviane Jacquemin and referred to in Kaufmann (1994: 125).

WORDS TO OBJECTS: 41

vocubulary of their language’ (Sturt 1833, vol.1:

187-88). While he adhered to his other

instructions regarding the collection and recording

of natural science objects, Sturt collected very

little ethnographic material and made few

observations of Aboriginal life. Writing after his

Central Australian Expedition of 184446, Sturt

admitted: ‘It might be thought that having been in

the interior for so many months I ought to have

become acquainted with many of the customs and

habits of the people inhabiting it, but it will have

been seen that they seldom came near us’ (Sturt

1847, vol.2: 139-40):

Sturt’s only records of exchange with

Aboriginal people during the earlier expedition

were references to bartering tomahawks, knives,

pieces of iron and coloured ribbons for fish (see

for example Sturt 1833, vol.1: 180; vol.2: 113).

During his inland expedition though, two

members of his party, Daniel George Brock and

John Harris-Browne, made small collections, and

part of the latter collection may have been

received by the South Australian Museum in

1949.*° Brock’s contact with Aborigines on the

expedition was relatively close and frequent, in

contrast to his leader. Sturt’s attitude towards

Aborigines, like that of the influential surveyor G.

W. Goyder twenty-five years later, appears to have

been friendly but firm:

if one or two of them were a little forward, I laid it

to the account of curiosity and a feeling of

confidence in their own numbers. But a little thing

checked them, nor did they venture to touch our

persons, much less to put their hands in our pockets,

as the natives appear to have done, in the case of

another explorer. It is a liberty I never allowed any

native to take, not only because I did not like it, but

because I am sure it must have the effect of lowering

the white man in the estimation of the savage, and

diminishing those feelings of awe and inferiority,

which are the European’s best security against ill

treatment (Sturt 1847, vol.1: 77-78).

George Grey produced several manuscripts on

Aboriginal material culture subjects, including

‘basket making’, ‘utensils for carrying water’,

‘ornaments’, and ‘shields’, presumably directed

towards a major publication on South Australian

Aborigines, never completed.*° These papers were

not simply descriptive. In his attempts to ascribe

origins to styles and design motifs Grey

prefigured the work of later ethnographers who

elevated material culture studies to scientific

status. Here again, Grey encouraged similar work,

particularly that of George French Angas (as a

protegé) and Edward John Eyre (as a rival). The

twenty-three year-old Angas accompanied Grey

on his vice-regal tour of the colony’s south-east

during January 1844, and his written observations

reveal something of the more experienced

ethnographer’s influence. Even taking account of

William Cawthorne’s unacknowledged assistance,

Angas’s fine-grained artistic depictions of South

Australian Aborigines and their artefacts set new

standards for ethnographic realism within

Australia. They met his stated objective of

‘preserving true and life records of man and

scenes, so quickly passing away ... by pictorial

representation, to describe the most interesting

and peculiar features of South Australia and its

aboriginal inhabitants’ (Angas 1847b: preface).

That verisimilitude was heightened in Angas’s

first major exhibition, at London’s Egyptian Hall

during April 1846, by the inclusion of ‘costumes,

implements, weapons and utensils, belonging to

the Australians and New Zealanders’.”” Angas had

collected these artefacts during his travels in order

to illustrate them at the journey’s end. A critic

from The Times commented on the exhibition’s

value as a documentary record, together with the

accompanying ‘antiquities’: ‘The views and

portraits are far beyond the common class of

pictures; as works of art they possess very great

merit, but as connected with the antiquities and

present character and manners of the country in

which they were taken, they are almost

invaluable’ .”8

A similar emphasis on the range and variation

of Aboriginal material culture in southern South

Australia was evident in Angas’s narrative,

“Savage Life and Scenes in Australia and New

Zealand’ (Angas 1847a). The accent was further

TC oO TF EE

See Jones file on the Gilbert Collection. The provenance of the collection is not clear and several objects appear to date from later in the century and

from regions not visited by Sturt. Harris-Browne and his brother had pastoral interests throughout South Australia and the Northern Territory.

Nevertheless, several of the objects conform to the type and style of manufacture observed for the north-east of South Australia, through which Sturt’s

second expedition passed during 1845.

Cawthorne during 1843. See Foster (ed.) 1991: 22.

Grey manuscript collection, South African Library, Cape Town. Copies held in AIATSIS library, Canberra. The publication is referred to by William

7” {llustrated London News, 18 April 1846, p.253, quoted in Tregenza (1982: 13).

*% The Times, 6 April, 1846, p.3c, quoted in Tregenza (1982: 17).

42 P. G. JONES

marked in the work of Edward John Eyre. The

explorer’s ethnographic investigations were

advanced considerably following his appointment

as Resident Magistrate at Moorundie on the River

Murray in 1841. Here Eyre had the task of

regulating relations between Aborigines and cattle

overlanders following the bloody events of the

Rufus River affray earlier in that year. Eyre used

the opportunity to supplement the notes gathered

on his Central and Western Australian

explorations with detailed observations on the life

and material culture of the Aborigines of the

lower River Murray. Returning to England a

matter of days before Angas’s arrival in Adelaide,

Eyre took with him a large collection of

Aboriginal artefacts, stuffed animals, an aviary of

parrots and an emu. He had also obtained the

permission of his main informant, Tenberry, to

take his son Warrulan and another Aboriginal boy,

Koar, with him.” Perhaps this example provided

Angas with an additional incentive to emulate the

American watercolourist George Catlin’s ‘Indian

Gallery’ which had presented paintings, artefacts

and Native American performers in London’s

Egyptian Hall during 1840 and 1841.

Many of Eyre’s artefacts were collected by

Edward Scott, his old friend, a fellow explorer

and an associate at Moorunde. Scott shared Eyre’s

ethnographic interests and this collection, which

included weapons, nets, fishing gear and

ornaments, represented more than a casual

assemblage of souvenirs. Uncertain of his next

appointment, Eyre was probably serious when he

informed Scott by letter that, ‘when I get all odds

and ends together, I shall almost be enabled to

open a museum in Regent Street’.*° Eyre’s was

among the first to recognise the value of

collecting Aboriginal objects as empirical data

which could supplement linguistic evidence for

cultural origins and diversity. This recognition

was at least partly inspired by George Grey. Eyre

wrote:

as Captain Grey judiciously remarks, if the

comparison in such [linguistic] cases be extended,

and the vocabulary of each enlarged, there will

always be found points of resemblance, either in the

dialects compared, or in some intermediate dialect,

which will bear out the conclusion assumed. This

view is still further strengthened, by including in the

comparison the weapons, habits, customs, and

traditions, of the various tribes ... No one individual

can hope personally to collect the whole material

required; but if each recorded with fidelity the facts

connected with those tribes, with whom he

personally came into contact, a mass of evidence

would soon be brought together that would more

than suffice for the purpose required (Eyre 1845,

vol.2: 398, 411).

Even so, this passage makes plain the restricted,

corroborative role of native artefacts in the

developing science of ethnography at this time.

Artefacts were still characterised as ‘curiosities’,

not yet amenable to the levels of investigation and

analysis which ‘specimens of language’ were

receiving from philologists. Exceptions could be

found: during his survey of the Australian coast

undertaken between 1818-1822 Philip King

collected and described material culture objects in

some detail, notwithstanding the fact that his

primary ethnographic instruction was to compile

comparative vocabularies of all the tribes

encountered.*! Eyre’s ethnography provides

another significant example.

At the time of Eyre’s publication, the British

scientific expedition vessels H.M.S. Fly and

H.M.S. Rattlesnake were cruising Pacific waters.

The naturalist John MacGillivray accompanied

both voyages and his Australian linguistic data

gathered on the latter voyage contributed to one

of the most influential works in philology, R. G.

Latham’s ‘Elements of Comparative Philology’

(1862). Despite the wealth of material culture

objects encountered throughout the Pacific and

northern Australia the anthropological research

undertaken on the voyages was largely restricted

to philology. Mulvaney notes the fact that despite

the sixth volume of information published by the

expedition being titled ‘Ethnology and Philology’,

linguistics occupied all but a few pages: ‘a few

lines sufficed in every instance for a superficial

account of the rich material culture of

contemporary Pacific peoples’ (Mulvaney 1964:

26). Yet just five years later in his account of the

Rattlesnake’s voyage, J. McGillivray published

the first investigation into comparative material

culture (Mulvaney 1964: 30). His discussion of

the similarity between the Australian boomerang

and an Egyptian boomerang in the British

® See Eyre’s correspondence to E.B. Scott, August to December 1844, PRG177/2, Mortlock Library, Adelaide.

3% ibid, 30 November, 1844

31 Both this and the following example are drawn from Mulvaney’s 1964 survey of the history of Australian anthropology (Mulvaney 1964: 18-19, 22,

26).

WORDS TO OBJECTS: 43

Museum had two effects. It signalled the

admissibility of material culture data as evidence

to be investigated in tracing the history of peoples.

More significantly perhaps, it drew attention to

the future relevance of museum collections as a

site for this investigation.

Grey, Angas and Eyre helped shift the character

of Adelaide ethnography from its restricted basis

in administrative practice to a more empirical style

linked to wider trends. Grey’s and Eyre’s

approaches in particular foreshadowed the ‘survey

method’ of anthropology which became so

popular later in the century. In contrast to the

vocabularies and short works on ‘manners and

customs’ produced in Adelaide, the work of these

individuals was published in London for a wide

readership. Unfortunately for the development of

Adelaide ethnography, the nature of their talents

and connections made it inevitable that their stay

in Adelaide would be brief. While Grey can be

said to have built on the work of Governor

Hindmarsh in fostering the beginnings of

Adelaide anthropology, he left no successor. A

gap of almost two decades separated the

publications of Eyre and Angas from the next

detailed research on South Australian

Aborigines—George Taplin’s work on the

Ngarrindjeri groups.

The trio left Adelaide within months of each

other: Grey to become governor in New Zealand

and then South Africa, Angas to travel further and

to exhibit and publish his Australian and New

Zealand watercolours in London (adding a further

series of South African studies during 1847), and

Eyre to England before becoming a lieutenant-

governor in New Zealand two years later under

Grey. Each of these men had formed important

collections of Aboriginal material, but almost

nothing is known of the subsequent history of

these objects or, indeed, of the original

circumstances of their collection. This is despite

their extensive writings on Aboriginal material

culture.

There is little doubt that Eyre, like Angas and

Grey, collected Aboriginal objects partly to

illustrate subjects discussed in his publications.

He may have retained several of the items figured

in his 1845 two-volume account within his private

collection. But he may have had an additional

motive for collecting ethnographic material,

linked to his official role as Protector. By 1840

his counterpart in the Port Phillip Protectorate,

George Augustus Robinson, was advising his sub-

protectors in each Victorian district to encourage

the manufacture of native artefacts for sale

through agents appointed for the purpose. A Mr

Lilly, based in Melbourne, acted as the main agent

for this purpose until his retirement during

September 1840. By July 1840 the volume of

artefacts had provoked Robinson to advise that a

scale of purchase prices be devised, so that

Aborigines could ‘receive in provisions or useful

articles the full value of the money realised ...

money on no account should be given to the

Natives’.*? Following Lilly’s retirement the

scheme appeared to lose momentum, and

Robinson advised that the ‘baskets, skins and

other articles ... of Aboriginal industry are in

future to be retained in store at the station, until a

sufficient quantity be collected for transmission to

Melbourne ... to be disposed of ... either by private

bargain, public auction, public bazaar, or any

other mode’ .*?

Eyre’s advocacy of a similar scheme suggests

that he was aware of Robinson’s initiatives in this

regard. He wrote: ‘The elder natives should be led

as far as could be, to make articles of native

industry for sale, as baskets, mats, weapons,

implements, nets etc., these might be sent to

Adelaide and sold periodically for their benefit’

(Eyre 1845, vol.2: 489). The idea went no further

apparently; with Eyre’s departure the link between

ethnography and administration of Aboriginal

affairs was broken. It was to be another two

decades before South Australian Aborigines were

actively encouraged to produce artefacts for sale,

under George Taplin’s administration at the Point

MacLeay Mission.*

Angas’s published account of his January 1844

expedition with Governor Grey refers to only two

occasions on which he collected ethnographic

items. Of the first Angas wrote: ‘Mooloo, the

native whom I met near the junction of the lake,

parted with his mother’s skull for a small piece of

tobacco!’. This object was noted by William

Cawthorne on Angas’s return to Adelaide, shortly

after Angas had drawn it for publication.

Cawthorne described it incredulously as ‘a human

2G. A. Robinson to C. W. Sievwright, 20 July, 1840, quoted in Lakic and Wrench (1994: 37).

3 G. A. Robinson to C. W. Sievwright, 18 July, 1840; 21 September, 1840, quoted in ibid: 36, 37.

™ One of the first indications that Point MacLeay Aborigines were producing artefacts for sale appears in the Observer, 4 February, 1860, p.6.

at P. G. JONES

skull plastered up here and there with gum for a

PITCHER. The natives put a string through one

part of the skull ... and so carry their water. The

skull is cleanish outside but very black on the

inside. It looks a curious thing for a domestic

utensil. They are as bad as Lord Byron’.*®

On the second occasion Angas recorded without

comment the requirement of reciprocity implicit

in dealing with objects on the frontier of contact:

about twenty young men and boys came up to us,

and lighted their fires close to our encampment.

Their hair was mostly in curls, and had it not been

for the grease and ochre with which they had

bedaubed their heads, many of them would have

displayed beautiful hair. We obtained specimens of

it, and they insisted upon having locks of ours in

return, which they carefully stowed away in their

rugs (Angas 1847a, vol.1: 94, 134).

It is unlikely that the Aborigines regarded this

occurrence as extraordinary; their own use of

human hair for hair-string would have provided a

context.

Immediately prior to his appointment as

Secretary to the Australian Museum in Sydney

in 1853 Angas donated twenty-one artefacts to

that institution. Several of these may have been

Aboriginal, although it is likely that the bulk of

this and other ethnographic donations made by

him consisted of African and New Zealand

artefacts (Specht 1980: 8-9). Angas returned to

South Australia by the time of the Museum’s

foundation but by then he had abandoned his

interest in ethnology for the scientific

description of shells. He acted as temporary

Curator of the Adelaide Museum during

Waterhouse’s absence on MacDouall Stuart’s

expedition of 1861-62 and assisted in

describing the shells collected on that

expedition (Gill 1886: 44).

Four small grinding stones ‘from Southern

Australia, carried by the females in a bag slung at

their side’ were presented by Grey to the British

Museum during his term as Governor of South

** Cawthome Diary, Saturday, 3 February, 1844. Foster (ed.) 1991: 32-33.

Australia, possibly as part of a larger collection.*°

Grey also shared the nineteenth-century

collector’s interest in burial customs and artefacts.

During his January 1844 expedition to the

colony’s south-east he removed two mummified

bodies from burial platforms near Lake

Alexandrina, later sending them to the Royal

College of Surgeons in London (Adelaide

Observer, 6 April, 1844, p.5). The skull of a

South Australian Aborigine, used as a drinking

vessel, was collected by Grey and sent to Owen at

the British Museum during the early 1840s; this

may have been the very object mentioned by

Cawthorne (Gill 1907-8: 232-33). Grey

continued to correspond with South Australian

linguistic workers after his departure from

Adelaide, receiving a revised version of

Teichelmann’s Adelaide vocabulary as late as

1858.*” As Governor of South Africa, Grey made

presentations of copper ores and ‘a collection of

[mineral] specimens’ to the Adelaide Museum in

1857, and donated a New Zealand greenstone axe

thirty-five years later from his South African

home (Hale 1956: 6-7).*8

The departure of Eyre, Grey and Angas marked

the close of the first phase of South Australian

ethnographic activity. The period had been

characterised by a remarkable level of linguistic

research and publication. While the products of

this research helped to make evident the

differences between local Aboriginal groups, and

highlighted something of the complexity of their

social structures, the results barely touched the

sensibilities of those European immigrants who

rapidly outnumbered and displaced the

diminishing Aboriginal population. Against the

governing perceptions of Aboriginal people as

undifferentiated, marginal and primitive, the work

of the colony’s early ethnographers seems more

exceptional. In that light, the subsequent lack of

attention towards the collection and description of

Aboriginal material culture was_ hardly

remarkable.

*6 Museum of Mankind specimen documentation no. 1840-12-1 (1-4), Jones 1985 ms. Grey’s note also reads: ‘The magnet is sensibly affected by them’.

It is not certain that these were collected by Grey in South Australia. The 1840 date, if correct, may indicate that they were obtained on his first, brief

trip to the colony, or that they were collected in Western Australia, which would be consistent with the description, ‘Southern Australia’. Artefacts

which Grey collected during his governorship in New Zealand were presented to the British Museum in 1854 (Braunholtz 1938: 7). Grey was also

an enthusiastic natural history collector, with several species named after him. See Appendices C,D,E,F in Grey (184 1a).

Pers. comm. J. Simpson, 1991.

*8 On taking up his post as Governor of South Africa in 1855, Grey became a firm advocate of the South African Museum in Cape Town (founded in

1825). He sought copper ores from South Australia for it, in exchange for those sent to South Australia.

WORDS TO OBJECTS: 45

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1840. ‘Outlines of a Grammar, Vocabulary, and

Phraseology of the Aboriginal Language of South

Australia, Spoken by the Natives in, and for Some

Distance around, Adelaide’. Thomas & Co, Adelaide.

TEICHELMANN, C. G., SCHURMANN, C. W. and

WYATT, W. 1886. Adelaide and its neighbourhood

[vocabulary]. Pp. 148-151, in E. M. Curr (ed.) ‘The

Australian Race’. 4 vols. John Ferres, Melbourne,

vol.3.

THRELKELD, L. E. 1834. ‘An Australian Grammar

Comprehending the Principles and Natural Rules of

the Language as Spoken by the Aborigines in the

Vicinity of Hunter’s River, Lake MacQuarie, New

South Wales’. Sydney.

TINDALE, N. B. 1974. ‘Aboriginal Tribes of Australia.

Their Terrain, Environmental Controls, Distribution,

Limits and Proper Names’. University of California

Press, Berkeley.

TINDALE, N. B. 1986). Anthropology. Pp.235-249 in

C. R. Twidale, M. J. Tyler and M. Davies (eds.)

‘Ideas and Endeavours—The Natural Sciences in

South Australia’. Royal Society of South Australia,

Adelaide.

TREGENZA, J. 1982. ‘George French Angas. Artist,

Traveller and Naturalist 1822-1886’. Art Gallery

Board of South Australia, Adelaide.

WILLIAMS, W. 1839. ‘A vocabulary of the language

of the Aborigines of the Adelaide district, and other

friendly tribes, of the province of South Australia’.

MacDougall, Adelaide.

WYATT, W. 1879. Some Account of the Manners and

Superstitions of the Adelaide and Encounter Bay

Tribes. Pp. 157-181 in J. D. Woods (ed.) ‘The

Native Tribes of South Australia’, E. S. Wigg & Son,

Adelaide.

WYATT, W. 1886. Adelaide District: Vocabulary. Pp.

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vol. 2. John Ferres, Melbourne, vol. 3.

A STAUNCH BUT TESTING FRIENDSHIP: DOUGLAS MAWSON AND

T.W. EDGEWORTH DAVID.

BY D.W. CORBETT

Summary

CORBETT, D.W. (2000). Sir Douglas Mawson and Sir T.W. Edgeworth David enjoyed a long and

fruitful friendship which began when Mawson was a student at the University of Sydney and

continued until David’s death in 1934. The relationship developed from that of teacher-student to

expedition colleagues in the Antarctic and on to academic equals bonded by a passion for geology

and the pursuit of scientific truth. Differences in character and temperament made the two men

unlikely friends, but a strong bond of respect and affection enabled them to overcome the not

infrequent tests of equanimity, particularly on Mawson’s part. The connections they made and the

influences they exerted across Australia in the days before air travel and e-mail mark an important

phase in the development of a continental perspective on Australian geology and its emergence on

the world stage.

A STAUNCH BUT TESTING FRIENDSHIP: DOUGLAS MAWSON

AND T. W. EDGEWORTH DAVID.

D. W. CORBETT

CORBETT, D. W. 2000. A staunch but testing friendship: Douglas Mawson and T.W.

Edgeworth David. Records of the South Australian Museum 33(1): 49-70.

Sir Douglas Mawson and Sir T. W. Edgeworth David enjoyed a long and fruitful friendship

which began when Mawson was a student at the University of Sydney and continued until

David’s death in 1934. The relationship developed from that of teacher-student to expedition

colleagues in the Antarctic and on to academic equals bonded by a passion for geology and the

pursuit of scientific truth. Differences in character and temperament made the two men unlikely

friends, but a strong bond of respect and affection enabled them to overcome the not infrequent

tests of equanimity, particularly on Mawson’s part. The connections they made and the

influences they exerted across Australia in the days before air travel and e-mail mark an

important phase in the development of a continental perspective on Australian geology and its

emergence on the world stage.

D. W. Corbett, Honorary Research Associate, South Australian Museum, North Terrace,

Adelaide, South Australia S000. Manuscript received 26 August, 1999.

T. W. Edgeworth David (1858-1934) was

Australia’s foremost geologist in the early decades

of the twentieth century (Fig. 1). He was

appointed Professor of Geology and Physical

Geography at the University of Sydney in 1891

and his department was the first in Australia to

train students for a professional career in geology.

David was a gifted and dynamic teacher, an

adventurous and energetic field worker and a

prolific writer on geological themes, popular as

well as scientific. His most ambitious project, a

synthesis of the geology of Australia remained

unfinished at his death in 1934. Aspects of

David’s life and achievements are covered in

important reviews by Branagan (1973, 1981,

1985) and Carey (1990).

Douglas Mawson’s long association with David

began in 1899 when he entered the University of

Sydney as a sixteen year old undergraduate. For

the next thirty five years, until his death in 1934,

David was in turn Mawson’s teacher, mentor,

colleague, confidant and friend. Their relationship

was not always smooth, in part a measure of their

age difference (twenty-four years), and their

marked contrast in character. David was the

quintessential Victorian gentleman, courteous,

charming and erudite, wearing his heart on his

sleeve and with a flair for publicity. Mawson

although less outgoing was generous hearted with

a friendly disposition, but inclined to reticence

and introversion at times. Both were sensitive by

nature, David particularly so, and at times this led

to misunderstanding and over-reaction to real or

imagined slights. Yet despite their differences, the

friendship stood firm, surviving the hardships of

Antarctic exploration, the upheavals of the First

World War, and academic lives lived half a

continent apart. It was a friendship sustained by

regular contact at scientific meetings around

Australia and David, an inveterate traveller, was a

regular visitor to Mawson’s home in Adelaide.

Their correspondence, drawn upon freely in this

paper, throws much light on the relationship

between the two men. Unfortunately the surviving

correspondence is markedly unbalanced, with few

of Mawson’s letters surviving in the David

archive.

Mawson and the University of Sydney 1899-

1905

Mawson began his studies at the University of

Sydney as an engineering student. At that time the

Engineering Department was part of the Faculty

of Science and there were strong ties between

Engineering and the newly established, and

government supported, School of Mines. By

Mawson’s time the Geology Department was well

housed and equipped and attracting a growing

number of students, many of them, like Mawson,

from Engineering; first year Geology being a

compulsory subject for all engineering students.

David himself was responsible for most of the

50 D. W. CORBETT

FIGURE 1. T. W. Edgeworth David, (1858-1934).

(Photo: University of Adelaide.)

geology teaching with assistance in mineralogy

and petrology from W. G. Woolnough (from his

graduation in 1898 until 1901 when he moved to

Adelaide) and H. S. Jevons (1901-05) (Branagan

1985: 129). Mineralogy courses, formerly part of

Chemistry had been transferred to Geology with

the re-organisation of the Geology Department so

that Mawson in his early University years was

exposed to a curriculum strong in geology,

mineralogy and mining — all subjects which were

to influence his later career.

David’s wide-ranging interests and early

experience, both in Britain and later with the New

South Wales Geological Survey before joining the

University, were reflected in his courses which

covered physical and general geology and

included lectures on Australian geology.

Laboratory studies, both microscopic and

chemical, were also introduced. Courses were

strongly field-oriented, students being introduced

to the pleasures of field work at an early stage of

their careers.

One of David’s prime interests was in the

geological evidence for past glaciation. When he

was a student at Oxford in the late 1870s the

nature of glacial processes was still a new and

hotly debated topic on which there was little

general agreement. His first published papers (in

the Journal of the Cardiff Naturalists’ Society)

were on the evidence for glaciation in South

Wales and based on his own fieldwork.

Subsequently, in 1887, when working for the New

South Wales Geological Survey, he identified

glacial sediments in the Permo-Carboniferous

rocks of the Hunter Valley (David 1897), while in

1901 he confirmed the evidence for glacial action

on Mount Kosciusko (David et al. 1901). In the

same year, during Mawson’s third year at the

University, Walter Howchin, in South Australia,

announced the discovery of glacial rocks of

presumed Cambrian age in the Sturt Gorge south

of Adelaide (Howchin 1901).

David had a strong interest in South Australian

geology: he and Howchin had discovered the

first Cambrian fossils in the Mount Lofty Ranges

in 1896 (Howchin 1897). A Permo-

Carboniferous age for the Hallett Cove and

Inman Valley glacials having been established

by this time (David & Howchin 1897), the

recognition of two widely separated glacial

events focussed attention on the significance of

glaciation in the South Australian geological

record, which was to have important implications

for Mawson. Given David’s strong interest in

ancient glaciation it would surely have been a

hot topic of discussion in the Sydney department

and it seems likely that Mawson’s own interest

was kindled at this time. However his early

enthusiasms were for mineralogy and petrology.

He was awarded the Petrology Prize in 1901 and

completed the coursework for his B.E. in Mining

and Metallurgy in 1901. With David’s support

he was appointed Junior Demonstrator in

Chemistry in 1902 and collaborated with T. H.

Laby on a geochemical investigation of the

radioactive properties of Australian minerals

(Mawson & Laby 1904)

In 1903, on David’s recommendation,

Mawson took six months leave of absence to

undertake a ship-based reconnaissance survey

of the New Hebrides archipelago, a highly

adventurous trip into a little known area. The

expedition gave him valuable field experience

under trying conditions and he proved himself a

resourceful and perceptive field worker,

qualities developed later in his career as an

explorer and academic geologist (see Corbett

STAUNCH BUT TESTING RELATIONSHIP 51

FIGURE 2. Douglas Mawson, (1882-1958).

1997: 110-112). His published report (Mawson

1905) was the first comprehensive account of

the geology of the islands. Mawson’s

correspondence with David began on this trip

and his enthusiasm and persistence are well

shown in a letter from Tangoa on Santo (3 July

1903): ‘...have seen a good deal of S. Santo but

have found no trace of any really old rocks’. He

confides to David that he and his companion

Harold Quaife deceived Captain Rason, the

expedition commander, pretending to stay at a

missionary’s house knowing they would have

been refused permission to climb the highest

mountain on Santo. In the event conditions

proved difficult and they did not reach the

summit, but the geology was interesting and the

mountain was found to be ‘the ruined cone of

an old volcano’. Mawson returned to Sydney in

September 1903 to write his report and resume

his research projects and coursework towards

his B.Sc., which he completed, majoring in

geology in 1904, graduating in 1905.

New Fields

In 1904, W. G. Woolnough resigned from the

position of Lecturer in Mineralogy and Petrology

at the University of Adelaide and returned to

Sydney. With David’s backing, Mawson applied

for the position and was appointed, moving to

Adelaide and taking up his duties early in 1905.

David doubtless viewed the appointment of his

former student to the Adelaide Department as a

further strengthening of his ties with South

Australian geology. His old friend Walter

Howchin now became Mawson’s lecturing

colleague in the small Department. Differences in

age, temperament and areas of interest allied to a

strong perception that Howchin held a proprietary

interest over much of South Australia led Mawson

to choose a distant and relatively unknown area,

the border country between South Australia and

New South Wales in the Olary—Broken Hill region

for fieldwork and research. It proved an astute

choice; for it is a highly mineralised region of

ancient crystalline rocks which provided abundant

scope for his mineralogical and petrological

interests (Mawson 1912). Among the numerous

mineral occurrences was a recently discovered

radioactive deposit which Mawson was the first to

investigate fully and which he named Radium

Hill. A new and complex mineral which he found

there—a titanate of iron, uranium and rare

earths—he named davidite after his former

Professor (Mawson 1906).

Of particular significance in the Olary region,

the igneous and metamorphic basement rocks are

overlain by a sedimentary sequence of great

antiquity which Mawson was able to relate to

similar rocks closer to Adelaide, thus

complementing some of Howchin’s stratigraphic

work in the Mount Lofty Ranges. It was his

identification of thick glacial deposits within the

sedimentary sequence which not only suggested

correlation with the Sturt Tillite but re-kindled

Mawson’s interest in glacial rocks and processes

leading to his first Antarctic adventure and the

beginning of a lifelong concern with glaciation.

That Mawson dated his involvement from that

time is confirmed in a letter written to David many

years later (25 June 1934), only two months

before David’s death, in which Mawson recalls a

meeting in Adelaide with Dr. Caldenius, a

European glacial geologist. Howchin was present

at the time and evidently irritated Mawson,

prompting the comment ‘He [Howchin] does not

appear fully to appreciate details of our older

glacial series—as he talks of measuring up all the

D. W. CORBETT

22.

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WO w]] ‘MO1 JOY) PIARG oe pepnjouy *LO6] Arenuer ‘aprejepy ‘ydesSojoyd dnoid (SyyY) 20U2IDg Jo JUSUZ.URApY OY) JOJ UONeIDOSsy UvISE[eNsSNY ‘¢ FUNO

STAUNCH BUT TESTING RELATIONSHIP 53)

varve beds in the State—that job is more than a

lifetime’s work. I explained that I had been

specially interested in these beds and regularly

collecting data thereon since December 1906—

and that it was my desire to better understand

fluvioglacial sedimentation that caused me to seek

a trip to the Antarctic with Shackleton’.

Mawson, perhaps prompted by David, seized

the opportunity to meet Shackleton when he

passed through Adelaide in January 1907 (see also

Fig. 3). In the event his involvement in the

expedition did not quite go the way he had

planned, as he was to note later in his diary: ‘In

the first place, believing the Prof. [i.e. David] to

be remaining in Antarctica for whole time, I asked

only to go on voyage down—and that in 1909...

Without any correspondence on subject the Prof

wired me: appointed Physicist to Expedition and

report at once. Now this is hardly what I wanted

as, though I should have embraced the opportunity

of stopping whole time as Geologist, I did not like

the idea of physicist. However, after asking the

advice of friends and now hearing from the Prof.

that he and Cotton were only going down with the

ship, I agreed to go... Now, when halfway to

Antarctica, the Prof. decides to stay— he can do

geology, I cannot’ (Jacka & Jacka 1991: xxvii).

South with Shackleton

Despite Mawson’s early misgivings about his

role down south, David involved Mawson in the

geological work from the start and the two shared

quarters in the expedition’s base hut at Cape

Royds at the foot of Mount Erebus. When David

convinced Shackleton that an attempt to make the

first ascent of the mountain before winter set in

would have scientific value as well as being a

useful exploratory exercise, Mawson was included

in the party which made the ascent. He was

responsible for collecting geological samples as

well as for the photographic work. It took the six

man party three days and two nights of blizzard

conditions to reach the rim of the old summit

wall. David and Mawson with McKay then

descended to the crater floor, and climbed the

inner, active cone, looking down into the vent

from which steam was rising accompanied by a

loud hissing noise and periodic dull booms (Fig.

4). They calculated the height of the mountain to

be 13 370 feet (4075m) above sea level.

The short but difficult journey yielded much

interesting geological information (see David &

Priestley 1914) and its success probably

influenced Shackleton in selecting the same team

of David, Mawson and McKay, designated the

Northern Party, to undertake the journey to the

South Magnetic Pole. There is a certain

ambivalence in Shackleton’s instructions to the

party, for while the achievement of the Magnetic

Pole is given first priority, the geological

objectives are also stressed, including a survey of

the coast of Victoria Land and the Western

Mountains (Fig. 5), while on the return journey

Mawson is given explicit instructions to

investigate Dry Valley as a matter ‘of supreme

importance’ and to prospect for minerals of

economic value. Shackleton, most probably under

David’s influence, was now utilising Mawson for

his geological skills, realising that the David-

Mawson combination made a formidable team.

It was always going to be a tough assignment

and there were tensions in the party from the

beginning. Mawson was frequently frustrated and

irritated by David’s somewhat eccentric habits and

behaviour and his physical inability to keep up with

his younger colleagues, who consequently had to

literally shoulder more of the burden of sledge

hauling. There is no doubt that David did his best

(he was then fifty years of age) and that he in fact

overtaxed himself while carrying out his duties to

the best of his ability, but the fact remained that his

condition did place an enormous burden on his

colleagues. While Mawson confined his

frustrations to his diary and a rare verbal exchange,

it appears that McKay was more directly abusive to

his companion, reaching a point where he was

prepared to certify David insane, and urging

Mawson to take over the leadership of the party. It

was a step Mawson was reluctant to take. While

David at the end of their ordeal confided to

Mawson that he had considered Mawson as leader

from early in the expedition, in fact David (with

McKay) had decided to push for the Magnetic Pole

against the opinion of Mawson who was all for

abandoning this to concentrate on the scientific

objects of the journey (Jacka & Jacka 1991)

The Magnetic Pole journey was a gruelling

adventure which could well have had a less

satisfactory outcome. The three were perhaps

fortunate to survive and there is no doubt that it

was Mawson who held the small party together.

Nevertheless, despite the difficulties, the two

geologists were able to collect a considerable

amount of geological information during the

earlier part of the trip and Mawson is given due

recognition in the final report (David & Priestley

1914). Perhaps most significantly, at no time

during the entire expedition did Mawson lose his

54 D. W. CORBETT

FIGURE 4. The active crater of Mount Erebus, taken from the lower part of the crater edge. (Photo: D. Mawson,

from David & Priestley 1914).

respect for the man who was dubbed ‘The Prof.’

by the expedition, and he was quick to recognise

that, despite his idiosyncrasies, David always gave

of his best and had the success of the expedition

at heart. To Priestley, the other geologist on the

expedition, David was ‘a tower of strength’ who

was ‘in spirit the youngest man in a very youthful

company’.

The Stressful Years

Soon after the expedition returned to Australia,

David wrote to Mawson (7 September 1909),

discussing plans for the writing up of the results.

David had all the geological specimens in Sydney

and was relieved that Shackleton had not

requested any to be sent to London. Mawson was

to describe the southern journey rocks, and other

likely contributors had to be found. Meanwhile

Mawson still had to finish off his Olary-Broken

Hill work but was already arranging leave of

absence for a trip to Europe. For Mawson, the

Shackleton Expedition marked the beginning of a

lifelong involvement with Antarctica. For the next

ten years the southern continent and the First

World War were to occupy the greater part of his

time while University matters of necessity

STAUNCH BUT TESTING RELATIONSHIP 55

”’. GE ACIE

ee ee

aur

vi i eget

Apts be tf

PLATE IX

Numerous

4 high Ki

2 igh peaks

DRYGALSKI

ICE BARRIER

TONGUE

vor

+ $00 fathoms

Septe OF Statube Miles. ix 6.

CHEETHAM ICE BARRIER

TONGUE

lear

Map of Nansen-Drygalski area of South Victoria, Land

Showing Coast.Horst, outlet Glaciers, & Inland Ice.

Chiefly after Survey by DrMawson.

FIGURE 5. Map of part of the route taken by the South Magnetic Pole Party. (From David & Priestley 1914.)

assumed a subsidiary role. Before Mawson left on

his Australasian Antarctic Expedition in late 1911,

David wrote to him:

My dear Old Mawson,

Just a few words to bid you and all yours God

Speed, and wish you all safe return. You have a

great and perilous venture before you; may (you) be

spared to come through it all unscathed and to return

full of honour and gain to Science. My heart goes

with you all. Ever my dear man, your affectionate

old comrade.

T. W. Edgeworth David.

The AAE expedition did indeed prove a

perilous venture for Mawson and he was not to

come through it unscathed, but it marked an

important episode in Antarctic exploration with

significant geographic and scientific results which

established Mawson as an expedition leader of the

first rank and a pioneer in the use of the new

technology of radio. Mawson returned to Adelaide

on 26 February 1914.

In a letter (15 November 1913) David,

anticipating Mawson’s earlier return, had written:

‘There is so much to talk about that one hardly

56 D. W. CORBETT

knows where to begin’, and he ‘eagerly awaits’

details of Mawson’s expedition, noting that ‘the

whole word has been intensely moved by the two

great Antarctic tragedies’ (i.e. that of the Scott

party as well as Mawson’s loss of Ninnis and

Mertz). He was full of enthusiasm for the

geographic and scientific discoveries made by the

AAE, ‘which have added King George V Land

and Queen Mary Land to the map’; while ‘there

are all the biological, geological and glaciological

results, your physical observations and the

magnetic results, which should prove of the

highest interest’.

Both Mawson and David volunteered for war

service. David, though now 57 years old, was

attached as an officer to a Mining Battalion and

later as geologist attached to the General

Headquarters of the British 1* Army. On the

Western Front he was involved in drainage

tunnelling and water-supply problems and trenches

construction, and his expertise was recognised to

the extent that he was appointed Chief Geologist of

the British Armies (Carey 1990).

For Mawson, the war began before he had fully

recovered from the traumatic and debilitating

experiences of his expedition and while he was still

extremely busy with post-expedition matters. After

initial difficulties in securing a suitable position

(MacLeod 1988), he spent two years in a highly

responsible administrative post with the War Office

in London during which time David and Mawson

managed to keep up their correspondence. David

was particularly concerned with the publication of

the results of the BAE, writing to Mawson in

London from the headquarters of the Australian

Mining Corps in France (11 July 1916): ‘It is very

good of you to take on the work of seeing this Vol.

II of Antarctic Geology when you are otherwise so

busy. There is a lot of real good stuff in it, and it

has been unfair to the authors to keep back so much

good material from publication. It will be great

relief to me when the vol., is published.” On 23

August he mentioned the ‘nerve wearying work’

and the strain of being ‘more or less under fire

about every other day, and I have had four months

of it now, but am keeping very well, I am thankful

to say’.

Mawson replied (17 September 1916)

discussing BAE matters and criticising

Shackleton, relations with whom had deteriorated

after their return from the South and notably since

Shackleton’s ambivalence on plans for further

explorations, which had resulted in Mawson

taking charge and organising his own expedition.

Mawson no longer trusted Shackleton and he

confided to David that he had written to Lady

Shackleton: ‘to try and explain. I have stated that

Sir Ernest upset some of the government officials

in Australia with unwise policy. I stated finally

that I myself was sad to see that Sir E. had

dedicated the Geol Vol. of the 1907 expedition to

Caird [the donor to his present Expedition] when

Australia and Australians were the real

contributors. A case of off with the old love and

on with the new’.

On 13 October, David wrote to Mawson with

unwelcome news. He had been inspecting a well

on the Vimy Ridge when the windlass broke and

he fell 80 feet to the bottom, sustaining severe

internal and other injuries. Although he recovered

well, the after-effects were to cause him trouble in

later life. Nothing daunted, by 13 February 1917

he was once again writing of Antarctic matters,

thanking Mawson for information on the progress

of the AAE results—’ your own scientific results

seem to promise excellently’—as well as Vol. II

of the Shackleton expedition geology—’good to

hear that the latter is at last appearing thanks to

your untiring unselfish efforts’. The volume, a

compilation of individual contributions on aspects

of Antarctic geology investigated by the BAE,

includes one by Mawson on the petrology of rocks

from the mainland of South Victoria Land

(Mawson 1916).

Return to Academia

After the war, David and Mawson returned in

1919 on the same ship to Australia and to their

respective Universities, but to very different

futures. David was now approaching retirement

while Mawson was about to resume his academic

career after a very lengthy and stressful period

away from the University. Little had changed

during his absence: Walter Howchin was still

active in the Department at the age of seventy

four (although soon to retire), and facilities

remained at a pre-war level. So when Mawson

was appointed to the newly created position of

Professor of Geology and Mineralogy in 1921, he

had much work ahead of him to build what was

virtually a new department. The question of

adequate accommodation for teaching and

research was not to be satisfactorily resolved for

many years, and Mawson later regretted adopting

a compromise solution which proved increasingly

untenable as the years progressed. For most of the

inter-war years his only full-time teaching

colleague was Cecil Madigan, his Antarctic

STAUNCH BUT TESTING RELATIONSHIP 57

colleague for whom he had a high regard.

Unfortunately, the two were too similar in

temperament and ambition for them to be able to

sustain a successful personal relationship.

Inevitably most of Mawson’s energies in the

early 1920s were taken up with the development

of his new department, and it was to be some time

before he was able to pick up the threads of

research. David meanwhile was beginning to

relinquish some of his academic responsibilities

in Sydney, and to concentrate on his long-planned

work—the compilation of a ‘Geology of

Australia’. The success of such an ambitious

project depended heavily on the cooperation of

his network of fellow geologists around the

continent, a network he had worked long and

assiduously to develop with great success. In

Adelaide, Mawson, Howchin and L.K.Ward were

important members—Howchin, as we have seen,

since the 1890s. Mawson had hardly settled back

into academic life before he received a letter (31

May 1919) in which David expressed a hope to

see him soon and talk over a letter he had received

from Charles Schuchert at Yale dealing with: ‘...

the probable Proterozoic age of everything in SA

that is below the Archaeocyathae, or at any rate

below the Brighton Limestone—everything, that

is, that is not Archaeozoic. There is much to

support this view which was our old view until

the discovery of Archaeocyathinae by Howchin

and myself at Normanville’. While Mawson

would scarcely have had the opportunity to put

his mind to South Australian geology at this time,

he replied to his old teacher (although his letter

has not survived) for David wrote again on 27

July expressing his keenness in joining Mawson

on a trip north and interest ‘that you think certain

bands in the purple shales are acid tuffs, or

resemble acid tuffs. In my experience red beds are

often the result of the weathering of tuffs’. There

is no doubt that over the next few years there

were times when Mawson came to resent the

regular ‘pestering’ on geological matters to which

David subjected him. Yet David was always

solicitous of Mawson’s health, the concerned

friend and mentor interested in his work and

offering support and encouragement—as the

following extracts show.

On 21 January 1920 David replied to a (lost)

letter of Mawson’s in which he refers to Mawson’s

recent illness ‘[the] aftermath I fear of your really

appalling experiences on your Adelie Land

Expedition’. He continued: ‘In regard to the

comparative opportunities (to which you refer in

your letter) on elaborating your work on Broken

Hill as compared with the scientific work of your

AAE there can’t be the shadow of a doubt, I think,

that you chose aright, and the scientific world will

be greatly the richer through your expedition. I am

glad you are going to apply yourself to study of

the belt between Olary and Mt. Painter. It

must be a fascinating region geologically and

mineralogically’. He also refers to the agreement

Mawson had recently signed with the New South

Wales Government, which was to take over the

printing of the AAE results: ‘it will be a great load

off your mind when the last volume is finished and

in print.’ And referring to departmental matters he

wrote: ‘It was high time that the Geological

accommodation at Adelaide was improved. I am

glad your authorities recognise the need at last.’

In June 1921 David visited South Australia and

with Walter Howchin made an extensive trip north

through Marree and Oodnadatta to Crown Point

on the Finke River (Branagan 1981). The

objective was to gather information for his

‘Geology of Australia’ as well as for the AAAS

Glacial Committee. It was a good opportunity to

catch up with Mawson again, and responding to

an invitation he wrote (June 5 1921): ‘I have

promised also to stay with Howchin and Ward,

but will come to you first.’ He kept in touch while

on his travels, writing from Marree of the train

journey from Quorn ‘in the lap of luxury’ having

been given the use of a private rail car ‘fitted with

bunks, parlour compartment, cooking stove and

every luxury... From Quorn we have passed over

extraordinarily interesting country, both

geologically and botanically.’

David travelled extensively during 1921,

visiting Western Australia, Queensland and

Tasmania as well as parts of the northern and

western areas of South Australia to gather

information for his book. Back in Sydney towards

the end of the year, on 22 December, he wrote a

long letter to Mawson asking for information on

South Australian geology and especially on the

relative ages of the granites of which there were

two phases; but were both Precambrian or was

one younger? And was there any evidence of

intrusion into the Archaeo limestones? The

questions continued.

It was during 1921 that David revived interest

in the possibility that fossil remains might be

present in the Precambrian rocks of the Mount

Lofty Ranges, an interest which dated back to

investigations with Howchin twenty five years

earlier. The new ‘discoveries’, in siliceous

limestones below the Brighton Limestone at

Reynella south of Adelaide, suggested crustacean

58 D. W. CORBETT

remains and as David believed the horizon to be

Upper Proterozoic (Precambrian), he was

cautiously excited by the find: ‘If the crustacean

remains...are really, as I believe, Proterozoic in

age it would be of quite extraordinary interest to

secure a complete fossil specimen... There is...

convenient to Adelaide, a considerable thickness

of strata containing traces of obscure organisms,

and it is to hoped that patient search by local

geological workers will soon be rewarded...’

(David 1922). David pursued the search with

relentless vigour in the years ahead and was to tax

the patience of Mawson at a particularly eventful

period in his career.

Further Distractions

A more immediate development was

precipitated at the end of 1924 by David’s

retirement as Professor of Geology at the

University of Sydney (Branagan 1981, Corbett

1998). At the time Mawson was not fully

committed to a future in South Australia despite

his new responsibilities and he was keenly

interested in the position, not least because of the

advantages of being located in Sydney where the

Government Printer had recently taken over the

publication of the AAE results. He made enquiries

regarding the terms of office from the Registrar in

Sydney, to learn that David was strongly

supporting his locum and deputy Leo Cotton. In

deference to his former Professor, Mawson was

slow to throw his cap in the ring. But as it

transpired he almost won the chair, being the most

favoured of a small group of candidates who were

considered (including W. N. Benson from Otago

and C. E. Tilley from Cambridge) and losing on

the final vote 10-8 to Cotton. The complicated

background to the appointment has been discussed

by Branagan who concluded: ‘There is little doubt

that David swayed the Senate to ensure this

appointment...[and] the Senate vote clearly was

far from unanimous and some Professors were not

happy with the procedures, and felt David was not

as open as he might have been on the matter’

(Branagan 1981: 38). Certainly to many it seemed

that justice had not been done and Mawson was

foremost among these, writing to David on the 23

January 1925. David replied promptly and at

length justifying his role in events, and using all

his charm and powers of persuasion to help pacify

his aggrieved friend. He wrote (31 January, 1925):

‘In spite of what is clear from your letter viz. that

the judgment of our Senate in the matter of choice

of my successor, guided as it was a good deal by

myself, was in your opinion, one which did not do

justice to yourself. You still sign yourself “ever

yours sincerely”...Only a man of noble and generous

nature could under the circumstances have done that

and I know the words come from your heart. My

own feeling is that nothing in the world should ever

come between the friendship of you and me, and I

am humbly thankful that your friendship for me has

stood this tremendous test.’

He counters Mawson’s queries and criticisms

with his own views before conceding: ‘Possibly

my judgment and advice to the Senate has been at

fault, but I am sure you know me well enough to

be assured that if I have erred, it has not been

through any motive other than a bona fide desire

to do a fair thing.’

He concluded by taking the high ground and

turning teacher: ‘Have you tried to put yourself in

Cotton’s position, and imagined what you would

have thought if you had run your Department at a

University with conspicuous success for 6 years

and then were asked to keep on for a 7th year, in

order to stand down for someone else at the end

of that time. Both you and I would, I think, have

protested pretty vigorously—at least I know I

would...we must please, agree to drop the subject.

I want badly to write to you about your Antarctic

matters and about S. Australian geology.’

David became concerned when Mawson did not

reply to his letters and suspected that a Sydney

colleague, H. S. Carslaw, Professor of

Mathematics, who was also a correspondent of

Mawson, had been causing trouble between them.

On 17 March he wrote again: ‘[you have]

probably been away somewhere in the Flinders

Ranges and have not been getting your mail

regularly’ and continued ‘you may have allowed

the recent appointment to the Sydney Geology

Chair to estrange us.’ He refers to Carslaw as ‘a

notorious “‘sticky beak”’ and concluded: ‘If by any

chance he has been making trouble between us, I

would earnestly entreat you as one of my very

dear friends...to tell me straight out what the

trouble is.’

Two days later (19 March) David wrote again,

still concerned but supporting Cotton, referring to

his own disappointments (David had once, as a

young man and before leaving England, sought a

job in Canada), and pointing out:

‘in your position in Adelaide, as far as research is

concerned, you have one of the very choicest plums

in the British Empire. What more glorious field for

geological research could any one in the world have,

with your special geological tastes, than the Mt.

Lofty and Flinders Ranges. I can assure you, as a

STAUNCH BUT TESTING RELATIONSHIP 59

result of many years experience, that you would

undoubtedly have had distinctly less opportunity for

research at Sydney than you have at Adelaide.’

All this was very true but hardly likely, one

would imagine, to pacify a resigned but still

resentful Mawson. The letter ends on an

emotional note: ‘I realise that I am getting an old

man now, and in the eventide of life one clings

more than ever to early friends. It would be a

bitter disappointment to me to lose your friendship

now. I feel sure you will have the greatness and

goodness to brush aside all feelings of resentment

through disappointment, and will come back again

into good fellowship, and will help me with my

research in my old age, as you have helped me so

often in the years that are gone.’

This letter drew a prompt double response from

Mawson, for David wrote again on 1 April: “Your

kind letters have been an immense relief to my

mind. I am most thankful that our old friendship

stands firm. May it ever strengthen to one’s life’s

end! I never perhaps fully realised what your

friendship means to me until my experiences of

the last four weeks. The first thing in the morning

and the last thing at night you were in my

thoughts and every mail I was eagerly looking for

a letter from you. It was a great joy to me when it

came at last.’

The Search for Ancient Fossils

David was particularly active later in 1925,

travelling to Melbourne and Adelaide and on to

Perth from where he intended to continue on to

England. This extension of his travels was,

however, postponed and he retraced his steps to

Sydney. Although keen as always to see Mawson,

David missed him on his way west (Mawson was

in the field at Boolcoomata), while an invitation

for a stopover on the return journey was declined.

Nevertheless correspondence was kept up during

these travels and David’s letters were full of

geological questions with enthusiastic references

to the Cryptozoon fossils Mawson had found at

Italowie and the question of their age (Mawson

1925). On 16 July, before leaving Sydney, David

wrote : ‘[you] mention you consider the age

Ordovician, is it really clear that the horizon is not

Cambrian or even lower, such as the Adelaide

Series?’ And in response to Mawson’s reply

(missing) he wrote again on 9 August: ‘...glad to

hear that you consider your Crytozoa from the

Flinders Ranges at newest Cambrian, and

probably Proterozoic’. He also ‘hope[s] you will

soon be publishing your work on the

Boolcoomatta and Olary areas. It will be a very

important contribution to our knowledge of

Australian geology.’ Later, on the train near Port

Augusta on the way to Perth, he wrote again (25

August), musing on the similarity between the

Italowie Cryptozoons and the algae from the Belt

Series in Montana described by Walcott and

adding: ‘looks like strong confirmatory evidence

of a Proterozoic age for the Adelaide Series.’ He

mentions his own finds in the believed

Proterozoic rocks at Reynella and the *...hope now

that you will be able to get some complete forms

of those v. interesting small crustacea whose

remains are so numerous, but so fragmentary in

the Brighton Limestones.’

Before David left for England in late November

1925, he wrote twice to Mawson on the matter of

the age of the cryptozoal limestones from

Italowie, and their similarity to forms from the

Macdonnell Ranges described by Ward (1925)

and thought to be Ordovician in age. Very keen to

clarify the matter, David wrote on 27 October: ‘is

for example their relation to the Adelaide Series

established as by their relation to the tillites, the

Brighton Limestone or the purple shale series?’

Ten days later he wrote again trying to pin

Mawson down on the stratigraphic position of the

fossil-bearing beds.

By February 1926, David was in Cambridge

writing to thank Mawson for papers received,

touching on a range of matters and reviving the

Cryptozoon question with a comment on the

opinion of A. C. Seward (a_ leading

palaeobotanist) which cast some doubt (later

resolved) on the organic origin of the MacDonnell

Ranges specimens. While David comments that

his book ‘progresses at a painfully slow rate’, in a

later letter (1 July) from London, he is again

seeking Mawson’s opinions, this time on

Madigan’s recently published work on Fleurieu

Peninsula:

‘Do you think it really proved that S. of Yankalilla —

on towards Cape Jervis these limestones are really

the equivalents of the Archaeocyathinae Lsts, or

may they be equivalents of the Torrens etc

limestones? Obviously if Madigan’s views are

correct, they considerably modify one’s views about

the dates of igneous intrusions in the Encounter Bay

area. I cannot but think it probable that those

pegmatised quartzites and conglomerates close to

the rutile mine you showed us at Yankalilla are the

basal beds of the Adelaide Series and the equivalents

of the Aldgate basal beds’

David illustrated his letter with sketch sections

60 D. W. CORBETT

and was grappling from half a world away with

the interpretation of one of the key areas for the

elucidation of early Australian stratigraphy. He

admits that his ‘path is beset with doubts and

difficulties, and three months ago I had a rather

nasty collapse from overwork, but am better now’

Mawson meanwhile, apart from_ his

departmental responsibilities, remained very much

involved with Antarctic matters, both past and

future, to the extent that his own researches and

David’s probing questions on matters of local

geology became a secondary consideration. He

was himself overseas, in South Africa and

England later in 1926, and carried out a short

lecture tour in the United States, the proceeds of

which were to help defray the cost of publishing

some of the results from the Australasian

Antarctic Expedition (AAE) which were still

pending. He was also beginning to formulate

plans for what were to be his own last Antarctic

explorations, the BANZARE voyages of 1929-

31, and in 1928 was back in England seeking

support and a suitable expedition ship. (He was

able to lease Scott’s old ship ‘Discovery’ from the

British Government for two seasons of

exploration.)

While Mawson was away and pre-occupied with

Antarctic matters, David, now back in Sydney, was

still strongly focused on his self-imposed task of

bringing together all that was known about

Australian geology, well aware of the paucity of

information in many areas and equally sensitive to

the fact that time was against him—a truth which

only served to heighten his zeal.

The Search Intensifies

Mawson was away again overseas in 1928 and

in his absence David returned to South Australia

and, possibly alerted by observations made by A.

R. Alderman (D. F. Branagan, pers. comm.) made

further finds of (he believed) organic remains in

the Precambrian rocks of the Mount Lofty

Ranges. The enthusiasm with which he renewed

his forays into South Australian territory was

reminiscent of his discoveries with Walter

Howchin around the turn of the century. Then it

had been a case of a professional working with a

highly competent but unqualified amateur in what

was Virtually a ‘terra incognita’. Now thirty years

on, David’s actions seemed very much like an

intrusion into the preserve of a small but well-

established geological community, headed by a

long-time colleague and friend. Not surprisingly

Mawson was perturbed, and his concern was

compounded by the realisation (shared by most of

those who became aware of the investigations)

that David had been so carried away in his

enthusiasm that he had begun to lose touch with

reality.

The saga began on 26 September 1928, when

David wrote a very long letter to Mawson, in

London at the time, telling of his discovery of

fossils,which he believed to be eurypterids, in the

old Beaumont quarries and also at Tea Tree Gully.

The finds were ‘of immense scientific importance’

and he was preparing a paper for the Royal

Society (David 1928). David wrote

enthusiastically that ‘more specimens will be of

quite extraordinary palaeontological and

evolutionary interest.’ He also mentioned that A.

R. Alderman, one of Mawson’s staff, had assisted

with the collecting.

Impatient that he had received no response from

Mawson on the matter of the fossils, David wrote

again on 9 October in a state of high excitement:

‘To my mind the whole thing is stupendous. Some

5000 feet [1524 metres] (more or less) of strata from

just above the Blue Metal Limestone down and

halfway between the upper and lower Torrens

Limestone contain remains of Eurypterids of great

variety and in great abundance certainly on some

horizons, as for example in that of the 2 ft. [60 cm]

bed of quartzite at Tea Tree Gully. If only I could be

with you even for a few hours I’m sure I could fill

you with enthusiasm for the whole matter... It will

be for you and your Geology School to enter this

‘promised land’. One cannot be thankful enough that

one’s eyes have at last been blessed with the sight of

it after some 35 years of intermittent seeking... Of

course it [will] take a hundred years or more to get a

fairly complete acquaintance with the Adelaide

Series Fauna, but it should be quite possible in a

years time to collect sufficient eurypterid material to

warrant publishing a detailed description’

A Friendship Strained

Mawson while in England had heard

independently of David’s activities in South

Australia and returning to Australia n the SS

Mooltan he wrote on 26 September:

“Your letter re fossils in the Adelaide Series, near

Adelaide is interesting. I have, of course, poorly

indicated fossil remains from elsewhere in South

Australia. Further examination of these has been

held up until I can complete my section in the North

Flinders Range. I got some money for expenses for

this voted by the University Council late last year,

and if this present matter [proposed Antarctic

STAUNCH BUT TESTING RELATIONSHIP 61

expedition] had not cropped up would have been

there now. The ‘pellets’ in the shales associated with

the archaeocyathinae at Sellick’s Hill, were left to

you by agreement as you had first drawn attention to

them. When I left Australia in March, I asked

Madigan to write and ascertain whether you

intended to go on with them—if not he was to

examine them and the enclosing rocks for fossils... I

gather from your letter and a reference made in a

letter from Madigan that you examined other rock

formations of the Adelaide Series and found

abundant traces of life.’

‘I am wondering what you are including in your

operations in South Australia. I have clearly

explained to you by references on several occasions

both verbally and by letter, that for years past I have

had a plan of campaign laid for the eventual

elucidation of the stratigraphy of the older rocks of

South Australia and it has been going forward

according to plan. To that end I have encouraged

and trained various students—and allocated to them

and to Madigan problems which have been selected

as part of the scheme for the ultimate complete

unraveling of the stratigraphy. I have spent money

in travel to see formations in South Africa and

America and familiarise myself with glacial

petrography and the character of the fossil forms of

the plant and animal life of the oldest terraines.’

‘This problem I particularly decided upon as a major

life work after I found I was not wanted in Sydney

for the Chair there. I am not looking for the

limelight consequently have not published anything

but adventitious scraps of the knowledge of the said

rocks now accumulated. My plan has been to

publish little until’...(here the letter ends with the

remaining page(s) missing).

It was a strong letter and the tenor of it was not

lost on David; but his reply on 18 October was

entirely in character, beginning with an

admonition of Mawson for not replying to his

earlier letters and going on to feign surprise, hurt

and indignation when it appears ‘I have, most

unintentionally, given you offense’, and ‘that

there is a suggestion implied though not expressed

in so many words, that I have been doing things

behind your back, and have in some measure been

jumping your claim. If any implication of this

kind is intended, I wish to state most emphatically

that it is most deplorable that you should entertain

any such idea...’

He then embarks on a lengthy review and

justification of his work in South Australia, dating

from his early associations with Tate and Howchin

and his publications in the Transactions of the

Royal Society. He refers to the results of his

Reynella discoveries (David 1922), when ‘You

certainly never gave one the slightest hint at the

time that you in any way resented my publishing

this paper’; and he reminds Mawson that the two

of them collected together at Brighton and Reynella

in 1925 ‘with the expressly affirmed object of

trying to discover further and more definitive traces

of life in the rocks of the Adelaide Series. Again

you did not give me the least hint that you did not

wish me to proceed with the work, and you most

kindly and generously helped me along with the

work of collecting.’

While Mawson was away earlier in the year

David had concluded from his microscopic studies

that the Reynella rocks ‘were not only swarming

with small fossil organisms, but that even their

branchidae, tentacles etc. were beautifully

preserved. This find greatly thrilled me and I wrote

over at once to Howchin, Ward and Madigan about

it and followed on myself. Had you been there of

course, I would have gone to you first and no

possible misunderstanding could have arisen.’

David was so carried away with enthusiasm for

his discoveries and his desire to prove their

authenticity that he labours the point of the

frequency of his visits to Adelaide, the practical

help he received from Mawson’s own staff

(Madigan, Alderman and Brock) as well as

Howchin; and the details of the search. Perhaps

part conscious of the effect this would have on

Mawson he continues:

‘I sincerely trust that if you have had the goodness

to read this lengthy letter so far, you will realise that

I was in no way prepared to find that you in the least

resented your old teacher and colleague having

ended his own intermittent search for fossils in the

Adelaide Series by coming upon forms which for

variety and structure have simply taken one’s breath

away... When you see one’s paper in print (I hope it

will be ready now in 2 or 3 weeks) I feel sure that

there is nothing in the paper of which you will

disapprove. One has, I hope, quoted fully and fairly

your own work on the Adelaide Series in that paper.

The occurrence of this fossil fauna so widely

distributed throughout the Adelaide Series is a really

stupendous thing teeming with possibilities, and the

surface of the problem has as yet been barely

scratched. It will be for you and your staff and

students surely to enter into the Promised Land, and

I’m sure that a grand future a really tremendous

future lies before you there.’ Finally he wrote:

‘I am convinced that you must have written that

letter to me of 29.9.28 [David has the date wrong

here. He is referring to the ‘Mooltan’ letter of

26.9.28] under some misapprehension. Of course it

is not nice for any scientist (such is human nature

alas! I confess guilty to the weakness myself) to

have some other scientist come along and discover a

gold reef under one’s front door step—personally I

62 D. W. CORBETT

Ta St

FIGURE 6. Edgeworth David at the Tea Tree Gully ‘fossil’

site excavation, with workmen, 1930. (Photo: E. Joyce.)

STAUNCH BUT TESTING RELATIONSHIP 63

don’t like that sort of thing at all, and it has several

times happened to me—but in your case while your

feelings would be I’m sure like mine—viz. regret

that one hadn’t found it all out oneself—you are

surely far too big a man, a world personage for all

time, whose Antarctic volumes are already making a

very creditable show alongside of the Challenger

expedition Reports, to resent your old Professor

finding after many years the bread he has cast upon

the waters!’

“Your “Mooltan” letter, in the light of your non-

reply to my friendly letters has caused me very much

pain. It is not in the interests of our friendship or of

Science. Please may I, only with the best possible

intentions and friendly feeling, be allowed to return

it to you? I trust...that you will kindly say yes.

Yours Very Sincerely

T. W. Edgeworth David’

This letter has been quoted at length because of

the way it illuminates David’s character and the

nature of his relationship with Mawson. He cannot

resist resorting to the ploy of one-upmanship—the

old Professor scoring a point off his old student

and having taken the prize, to magnanimously hand

it over to his worthy pupil and successor. Although

highly sensitive himself and conscious of the fact

that Mawson might have been disconcerted by the

news of the discoveries, he seems peculiarly

insensitive in the manner of his writing. He was so

overjoyed with what he believed to be the high

point of his career that he could not understand

others not being equally excited and responding

quickly and with enthusiasm. That Mawson, whose

opinion and counsel he valued so highly, did not

do so was devastating and David could not hide his

feelings or control his pen. Mawson could perhaps

have acknowledged the flurry of letters he received

more promptly, but he was very busy in London on

Antarctic matters and his response, when it came in

the ‘Mooltan’ letter, was entirely in character—

courteous and cautious while expressing a sense of

affront that was entirely justified.

But David was now a man obsessed, and with

no further word from Mawson, he wrote again on

28 October:

‘I am deeply distressed at not having any reply

from you to my last three letters. You are one of

my students of whom I am especially proud; and

for whom I have a warm and abiding affection,

which nothing can alter, and your present silence

is so distressing to me just by reason of the

affection I have for you. Were it not for this I

would just have said “Well if he won’t reply to

my courteous letters written with the simple desire

to remove any possible misunderstanding, why not

let him rip!”

He then, mentions his indebtedness to Howchin

during his recent researches, commenting: *... for

with all his faults—and I wish I had as few as

he—he really is a wonderful old man and has

done a monumental piece of work around

Adelaide and far beyond.’

On 1 November, 1928, Mawson finally put

David’s mind at rest writing with apologies for

not replying earlier and explaining that he had

been extremely busy but

‘Now I am beginning to breathe again lectures are

over. Alas! I am sorry to have worried you but I

have merely accumulated letters from friends whose

handwriting I know until the moment should arrive

when I could deal with them... Yours of course are

of the greatest interest. Since arriving back I have of

course heard of course what your discoveries and

work over here has been—and your letter of 26th

September is most interesting. From reports in the

English papers it was not at all clear what you had

discovered. Though it did indicate that we in

Adelaide had been sleeping on the most stupendous

museum of clearly preserved fossils...’

(The page ends here and the remainder of the

letter is missing). Courteous and cool in its

introduction, the letter was beginning to warm up

a little. But there was certainly no stopping David

in his quest. He returned to Adelaide several times

over the next few years, in 1929, 1930 and 1933,

to investigate the Tea Tree Gully quarry.

Standing Firm

Mawson remained aloof and did not become

seriously involved, but the fact that David was

regularly assisted by members of Mawson’s

department, staff and students, indicates a desire

to keep an eye on proceedings. David was assisted

by Madigan and Alderman in 1929, by Rudd in

1930 (Fig. 6), and on his last visit in 1933 when

he also visited the Beaumont Quarry, by Madigan,

Kleeman and Barnes. On this, his last visit to

Adelaide, he spent time with Madigan at the

University looking over material from Central

Australia including large masses of cryptozoa

which he found ‘very wonderful’. In September

1929, Mawson supported David and his

collaborator, R. J. Tillyard, in seeking a grant of

£100 from the Royal Society of London to open

up a quarry in the Precambrian rocks near

Adelaide to search for further fossils. The

successful outcome enabled David to carry on his

research. Mawson was asked to contribute a piece

on evidence for the Precambrian age of the

Adelaide Series and for an unconformity below

64 D. W. CORBETT

the fossiliferous Cambrian in the Flinders Ranges

for: ‘a joint paper by Tillyard, yourself and myself

for the Phil. Trans., which would be a nice little

“quid pro quo” for the 100 quids for which we

jointly applied’. He tries again to win Mawson’s

enthusiasm, noting that Tillyard is ‘...quite

definite as to the arthropod nature of the Teatree

Gully fossil’, a photograph and a drawing of

which he includes. Mawson was not to be

tempted, however, and in a cool letter (18

September 1930) he offered to run out for a

couple of hours to Beaumont but ‘My time,

however, is extraordinarily full’. He also declines

to collaborate on the proposed paper writing : ‘I

would not enter into a new subject without going

very fully into it’, while he has half a dozen

papers on the verge of publication.

By the middle of 1931, with the BANZARE

cruises behind him and the pressure off a little,

Mawson was able to catch up with his

correspondence. On 2 July he told David of Kerr-

Grant’s discovery of the Karoonda meteorite and

the public interest aroused; most reported finds

were ‘myths’ but one report had led to the visit by

Alderman to the previously unknown Henbury

meteorite craters and the collection of numerous

iron meteorites. On 14 July he wrote again telling

of Madigan’s recent seven week trek east of Alice

Springs and his discovery of archaeocyathae in

the same relations to the algal limestones that

Mawson himself had found in the Flinders in

1929. While David was worried by ill health and

his old war injuries during 1931, he was buoyed

by the publication of his geological map of

Australia and by the renewed and more regular

contact with his old friend. By 12 September he

was optimistic enough to approach Mawson again

regarding the Precambrian fossils. He wrote:

‘Anxious that you should join me in a stratigraphical

note on the horizon of those fossils in the quartzite

at Teatree Gully. You remember that Tillyard and

you and I joined in an application to the Royal

Society London for that grant of £100. The results

have exceeded even sanguine expectations, and

Tillyard has prepared a very able paper on the most

typical of those protoarachnoids. We have a

specimen nearly perfect about 6 inches long. I

should personally be extremely grateful if you would

help me with this stratigraphical note. If you will

say yes! I will forward at once some rough ideas of

my own for you to trim up.’

Mawson did not say yes. In his reply of 23

September he quite firmly distanced himself from

the Teatree Gully project:

“With regard to the fossil evidence you must rely on

your own judgment and that of Dr Tillyard. With

regard to the stratigraphical position of those beds,

Howchin is, I suppose, the best authority. In view of

the fact that Howchin has made a special feature of

the study of the old sedimentary rocks in the

neighborhood of Adelaide, I have never entered the

field in any critical fashion and have reserved my

efforts for more distant regions. Should Howchin

pass away, I would, of course, feel it my duty to re-

investigate the Adelaide region. Until then there is

such a wide field in this State for further work that

one feels one would do better for Geology as a

whole by extending observations elsewhere rather

than critically examining a field that has been given

very great attention by Howchin.’

A Shift in the Relationship

David’s last attempt to gain Mawson’s support

had failed; indeed it had merely provoked him,

arousing all the old resentment about Howchin’s

monopoly on the geology of the Adelaide region.

As far as Mawson was concerned the matter was

now closed, and in his firm and calculated

withdrawal from the field it seems that in the

twilight of David’s life, roles had been reversed:

Mawson is now the mentor and David the receiver

of counsel. While David continued to keep

Mawson informed on the development of the

Precambrian fossils saga, Mawson’s letters

focused on other topics including Madigan’s work

in the MacDonnell Ranges and Hossfeld’s in the

North Mount Lofty Ranges. Nevertheless David

missed no opportunity to raise the question of the

possibility of finding Precambrian fossils in the

Flinders and Central Australia where the rocks

were less altered, while he diplomatically kept

away from mentioning the Adelaide fossils.

Meanwhile David had written to geologists

world-wide, spreading the news of his discoveries.

Mawson’s skepticism regarding the Precambrian

fossils was shared by many in the geological

community, including J. W. Gregory (Glasgow

University), C. E. Resser (Smithsonian

Institution), and Calman and W. D. Lang,

biologist and geologist respectively at the British

Museum, and Woolnough at home. Lang was

‘sorry not to be able to go as far as you in

considering these supposed organisms referable to

annelids or arthropods. I cannot deny that they

may be organic, though even this I think rather

unlikely than probable’. The general feeling was

that the impressions were, as Woolnough put it,

‘mere concretions’; but with Tillyard now an

enthusiastic convert (as well as an expert on living

STAUNCH BUT TESTING RELATIONSHIP 65

FIGURE 7. The supposed Precambrian fossil Protadelaidea howchini. (From David & Tillyard 1936: plate I.

Angus & Robertson.)

arthropods) and cautious and non-committed

support from some of his correspondents overseas,

David pressed on. Tillyard remained less

confident than David of gaining acceptance for

their work, and sounded a warning in a letter (5

February 1932) referring to some of Calman’s

criticisms and continuing: ‘In science, as

elsewhere, nobody is to be allowed to run beyond

the pre-conceived limits set by the conservative

opinion of his own times. So, I much fear, you

and I are in for a severe castigation when our

paper comes out. Nevertheless, when we are dead,

complete fossils of the same Pre-Cambrian age

will be brought to light, and will to a large extent

justify the reconstruction which we have given’

The paper referred to was rejected by the Royal

Society of London in July 1932. Tillyard

meanwhile, in a letter of 7 September 1932, was

concerned about the fate of specimens sent over

to the British Museum in the light of the rejection

of the paper by the Royal Society. While the

Society would ‘return the typescript and the

drawings..., these are of little value without the

type fossils’. He was also concerned about

Mawson, and wrote: ‘I am very sanguine as to

Mawson’s reaction; do not forget the old complex

about your working in his field. He is a very nice

fellow and is always very decent to me, but I

wonder whether he has a big enough mind to see

your point of view in this particular case.’ Tillyard

was also keen that another application be made

for a research grant from the Royal Society.

Mawson had been an influential supporter who

had helped secure the initial grant, but this time

Tillyard is less than confident, writing: ‘As he

[Mawson] is going to London next month, I

should expect that he would be more inclined to

secure a grant for exploration in the Flinders or

MacDonnell Ranges.’ Ten days later Tillyard was

able to report that the specimens were to be

returned and the scientific fraternity were a little

less antagonistic. ‘Perhaps they now feel rather

ashamed of having turned us down, and are trying

to make amends’. Tillyard, still hopeful of

enlisting Mawson’s support, wrote to David (22

October 1932) ‘I do hope you will bring Mawson

round to our way of thinking’

David was still trying. On 2 September he had

written to Mawson with the Royal Society paper

rejection fresh in his mind and sending copies of

the referees’ reports (by Calman and Lang of the

British Museum) with his own critical asides. He

66 D. W. CORBETT

FIGURE 8. Reconstruction of Protadelaidea howchini,

dorsal view. (From David & Tillyard 1936: plate IX.

Angus & Robertson.)

clearly did not accept the judges verdict and had

found some of the comments offensive. Calman,

‘a mere zoologist’ had concluded his report : ‘with

great reluctance I have come to the conclusion

that from a zoological point of view this paper is

entirely without scientific value.’ Lang’s

criticisms, ‘though not rude like Calman’s... [are]

not brought forward at all... if only he had read

our paper carefully... His criticisms really in no

way weaken our position.’ David then appealed to

Mawson: ‘Unwilling as I am to trespass on your

time, which I know is so very fully occupied, I

wish you to kindly do me the favour of reading

through these notes on the Adelaide fossils very

carefully, if you will be so good. With the arthritis

becoming constantly more acute... one realises

alas! that that little bit of work at Teatree Gully

may be the last bit of field work of one’s lifetime

and which embodies the only discovery of any

consequence that I have ever made, I am naturally

anxious to see a reasonably good foundation laid.’

On 6 September he wrote again with further

descriptions of the fossils and concluded *...in our

opinion, the evidence for the organic origin of

these fossils is so conclusive as to be quite

overwhelming.’ A postscript hammered home the

point:

‘The Teatree Gully quartzite has already yielded so

many remarkable and entirely new forms of pre-

Cambrian animal life that in my opinion its further

exploitation in the way of quarrying and splitting

the blocks parallel to the cleavage is really of

enormous palaeontological importance. No other

strata of such an age anywhere in the world, have

yielded results in the least comparable with this.

Surely before trying out new areas in the northern

Flinders or the Macdonnells further work should be

done at Teatree Gully e.g. the discovery of a third

specimen of Protadelaidea howchini [Figs 7-8]

would surely satisfy even the British Museum

brahmans...’

On the 15 September he wrote advocating

further quarrying at Teatree Gully, adding ‘I

would immensely like to hobble over the ground

(while I can still hobble) in your company some

time before you leave for England.’ Included in

this letter is an annotated section from his own

paper (David 1928), indicating the position of the

supposed fossil bearing-beds (Fig. 9).

Mawson never made that visit with David, for he

left for England towards the end of 1932 and did

not return until July 1933. Before leaving however,

he forwarded to the Royal Society in London, at

David’s request, an application for a further grant

for the excavation of the fossil site. Despite

David’s continuing hope for unqualified support,

Mawson’s attitude was at once diplomatic,

uncompromising, yet supportive. He appended the

following statement: ‘[as the] wording on the

document does not properly express my views, I

am merely forwarding this supporting note... The

position is that Sir Edgeworth David is still

convinced that certain markings, impressions and

cavities in a thin bed...of ?middle Proterozoic age

are of organic origin. My view is that some of them

are difficult to account [for] on a purely inorganic

basis, but I see nothing indisputably organic in

them’. His conclusion was pure diplomacy. ‘If they

are organic their occurrence is of great importance.’

He supports the further grant to prove or disprove

the “fossils” and would like to see the matter

further investigated.

Disappointed to learn that Mawson had not

approved of the wording of the Royal Society

application (he claimed he thought that Mawson

had accepted Protadelaidea), but still undaunted,

he returned to South Australia in April 1933,

while Mawson was away, on what was to be his

last visit. On this occasion he was assisted by

Madigan, Kleeman and Barnes in the field and

stayed at Mawson’s home.

STAUNCH BUT TESTING RELATIONSHIP 67

The following vertical section is chiefly after Professor Howchin, with

additions from C.

850 limestone with Saltcrelia planceowuers; Tate.

horizon, proved as yet, for fossiliferous Cambrian rocks locally.

Black slates and shales, weathering to claret colour.

Glen Osmond clay slates. — seep

T. Madigan’s papers :—

Archacocyathinac limestones underlying Redlichia beds.

Black to dark-grey carbonaceous shales with phosphatic concre-

mge spicules, and 8 feet bed of

This is lowest

Purple slate series with thin quartzites and a little thin oolitic

Brighton limestone, rich in remains of annelids and arthropods.

Siliceous limestones of Reynella, Field River, Burra, ete.

Tapley's Hill laminated calcareous shales and slates, with thin

limestones at top and bottom.

“Blue Metal” limestone win hs, annelids and Bucpplesida

The thick quartzite of Black Hill, Stonyfell, Mount Lofty, Mount

yas

wank AE saftey hace be ;

Upper Saal nts = soe Crystal At act es Mylor, Tena-

feate Creek, Peer

Gully, very fossiliferous.

Lower Torrens limestone, much marmorise

Phyllitic rocks with 180 feet of basal grit with much detrital

tam ch, » <—— a)

we of Ses Ae Tea 5

Schists, etc. intruded by basic to intermediate rocks of the

we oy Fett Feet.

4 ei

Z25{ 830 50

Sea | sco tions with radiolaria and 3;

2. ==

Sh ————

Be 150

H| 1000-

<{ 1690 limestone.

“5 78s

150-

600

ases

1000-

3000

7588

z 1$0- Sturtian Tillite.

<] 1000

z 750 9383 Subglacial quartzite.

a} "000 Upper clay slate.

e :

co .,-Gken Osmond wma

f ou} srs Ke

= 8) 2500

hereaAronte |

SE] 280 "2

4 7A Fo uzes

Vipede a ooo Upper phyllites.

12265

000 Barker, etc.

13265

4 « 2 ™500 Lower retell

? Ax 14766 Tas 4

200.4965

V7 aX = — Quer!

eS Phyllites and quartzites.

5965 | sere ree

50 Site +=

+1008 . ilmenite.

em 17S Violent unconformity.

nS g

e| Houghton Magma.

< %y

< below the lowest Australian horizon of

*Thus the base of the Adelaide oerios ia about 16.275 feet (4,924 metres)

age.

proved Lower Cambrian

FIGURE 9. Annotated geological section (from David 1928), showing position of supposed fossil-bearing formations

in the Precambrian Adelaide Series. This was included in a letter from David to Mawson, 15 September, 1932.

Envoi

By the end of 1933, David’s health was giving

serious cause for concern but his letters to

Mawson on a range of geological matters

continued unabated. He wrote in December 1933:

‘My health has been v. poor lately and my

suffering from my old war injuries has been

increasing but I am thankful to say that there is

some “life in the old dog yet” and I may yet live

to be of some service to my very distinguished old

student for whom I cherish no small affection.’ In

March 1934 he was writing of Antarctic matters,

Mawson’s work on coorongite in the South East

and north Flinders Ranges geology. He also had

plans to visit his publisher Edward Armold in

England for discussions on the still uncompleted

‘Geology of Australia’. Meanwhile work on the

book was continuing, though very slowly, with

the help of W. R. Browne and other Sydney

University geologists. By July the trip to England

had been deferred to the end of the year. In a

letter of reply to one of Mawson dealing with

glacial matters on July 11, his old student was

asked about his two Precambrian glaciations with

the question: ‘Have you, please, published

anything about this?’, while his last letter to

Mawson (15 August 1934) was concerned with

the Henbury meteorite crater reserve.

David did not live to complete his book. He

died in Sydney on 28 August 1934 and W. R.

Browne eventually saw it through to publication.

This proved a monumental assignment as David

had been continually re-organising his material

68 D. W. CORBETT

and the book lacked a final plan. Also as the

project had been in progress for so long, not the

least of Browne’s tasks was to bring the

information up to date. ‘The Geology of the

Commonwealth of Australia’ in three volumes

was published by Edward Arnold in 1950, edited

and much supplemented by Browne. As Branagan

has written: ‘Browne had to write David’s

“Geology’—and he did it magnificently’

(Branagan 1981: 52).

Lady David began compiling a biography of Sir

Edgeworth after his death but her draft had been

rejected by Angus and Robertson. This was a

severe blow, and her daughter now set about

rewriting the manuscript before taking it off to

seek a publisher. She wrote to Mawson on 27

March 1936: ‘I have put in 18 months severe

work and amassed facts only about him—but he

had such a romantic life it is not easy for me to

“see the wood for the trees”.’ She concluded: ‘I

am so glad you loved him.’

Mawson felt the loss of his old professor keenly

and when asked by the Royal Society of London

to write an obituary he did so in prose that was

very high in its praise, not only of David’s

scientific achievements, but of his character and

personality. He wrote :

‘Attributes of his greatness were an endearing charm

of manner and a nobility of mind embodying high

Christian principles. He was an accomplished scholar

plentifully endowed with fine instinct and broad

vision... With a rare fund of anecdote and a keen sense

of humour, as well as a vivid dramatic appeal and

exceptional powers of narration, he was always good

company and the centre of attraction in any gathering.

By tact, influence, and a unique personality, he

promoted science, and especially geological research,

in Australia, for more than four decades.’

And with reference to David’s geological

achievements: ‘Next in interest to the investigation

of evidences of past glacial climates in the Australian

geological record, the subject which claimed David’s

special attention during the later years of his life,

was that of search for fossil evidences for life in the

Pre-Cambrian terrains of the neighborhood of

Adelaide. In what appears to be the upper limit of

the local Pre-Cambrian sediments, he, some years

ago, recorded impressions of a fragmentary nature

which he determined as relics of crustacean life. He

more recently came upon coarser and still more

obscure impressions at a much lower horizon; these,

also, he believed to be fragments of crustacean or

proto-crustacean life...he was still investigating this

fascinating problem at the time of his death.’

(Mawson 1935).

The abundant material that David had left on

the supposed fossils, was, at Lady David’s

request, organised by W. R. Browne and Leo

Cotton and, in cooperation with Tillyard,

published in book form by Angus and Robertson

(David & Tillyard 1936).

Precambrian Fossils — the Sequel

With David no longer there to practise his

persuasive advocacy for the Precambrian fossils,

the memoir did nothing to revive interest and there

were few geologists in the years ahead who were

prepared to keep an open mind on the subject. Fifty

years on, S. Warren Carey, who had seen some of

the original specimens, was a _ powerful

spokesperson for the skeptics. He wrote: ‘I don’t

know any palaeontologist today who accepts them

as organic remains, and little is said about them, as

though this skeleton in David’s cupboard is best

forgotten’ (Carey 1990: 46). But over the years

between they had not been entirely forgotten. Even

if David was wrong, palaeontologists since his day

have recognised that the relatively complex animals

found as fossils in the earliest Palaeozoic rocks

must have had Precambrian ancestors. In the mid-

1950s, a highly respected textbook of invertebrate

palaeontology made the point clearly: ‘The Phylum

Arthropoda comprises an unusually large and

varied group of highly developed invertebrates

whose clearly long and extensive history reaches

back almost certainly into the Pre-Cambrian’

(Shrock and Twenhofel 1953: 536), and although

they do not discuss David’s discoveries, his

Memorr is listed in their bibliography.

If David’s fossils were indeed ‘mere

concretions’, how could such an accomplished

scientist (and his specialist collaborator) have

made such a misinterpretation? Carey asks the

question and proffers a possible answer; perhaps:

‘David, working through so many thousands of

specimens, may have progressively filtered out

those with bilateral symmetry, and Tillyard,

working with such a biased set, could easily have

been convinced that they had to be organic’

(Carey 1990: 46).

On home ground in South Australia, David’s

torch had been kept burning since the late 1930s

when a young geologist appeared, who carried

many of David’s characteristics, notably

enthusiasm, flair and imagination. R. C. (Reg)

Sprigg stood in the same relationship to Mawson

as Mawson had to David four decades earlier. As

a student he found what he thought were

STAUNCH BUT TESTING RELATIONSHIP 69

arthropod remains in rocks close to the

Precambrian—Cambrian boundary at Sellicks Hill

south of Adelaide. Mawson was singularly

unimpressed and later, much to Sprigg’s chagrin,

had the key specimen thrown away (Sprigg 1989:

95). But within a few years serendipity took a

hand, and Sprigg, working for the South

Australian Department of Mines on a field

mapping project in the Ediacara Hills west of the

main Flinders Ranges, found a fossil jellyfish in

rocks at that time believed to be lowermost

Cambrian. Mawson, perhaps inevitably, was not

initially convinced, and there was a strong

skeptical element in the geological fraternity

regarding the validity of Sprigg’s discovery.

However, further finds by Sprigg and other

collectors, of more jellyfish as well as numerous

other undoubtedly animal fossils, with a wide

range of forms and some with problematical

affinities, became world famous as the Ediacara

Fauna—the earliest animal fossils known in the

geological record. In his final years Mawson was

a strong supporter of the investigation of the

Ediacara fossils and on-going research was carried

out in his old department after his death by Martin

Glaessner and Mary Wade.

Mawson’s vision of unravelling and ordering the

Precambrian rocks of the Flinders Ranges had been

largely realised but success in finding fossils (other

than stromatolites), which he felt must lie in the

thick sedimentary sequence, had eluded him. While

David's (as he believed) misguided forays into the

Adelaide region had annoyed him, the success of

one of his own students in finding the earliest fossil

animals was a vindication of his own long

dedication to teaching and researching in the

northern ranges, as well as being a fitting

justification of David’s equally dedicated search for

evidence of life in the distant past. More recent

finds of problematica similar to David’s in North

America have revived interest in the nature of these

supposed ‘fossils’, with an indication that there

may be an organic component in some of them.

Postscript

David’s death was a significant event in the

development of Australian geology. It also came

at a time which marked a watershed in the life and

career of Douglas Mawson. At the age of fifty-

two his direct involvement with Antarctic

exploration had ended and he began his most

sustained period of research into South Australian

geology. Between 1935 and 1952 (the year he

retired), Mawson published 30 significant papers,

over half of them concerned with the Precambrian

rocks of the Flinders Ranges. (This compares with

a similar number of papers over a period twice as

long between 1904 and 1934 on a wide variety of

topics). The death of Walter Howchin in 1937,

three years after David, saw the approach of the

end of the old order of Australian geologists, and

in South Australia, as elsewhere in Australia, a

new generation of native-born and trained

geologists had emerged, all of them graduates of

Mawson’s department and including, in addition

to R. C. Sprigg, many others, notably E. A. Rudd,

L. W. Parkin, Paul Hossfeld and A. Kleeman.

Many of Mawson’s later papers were co-authored

with junior members of his department or former

students.

The lives of T. W. Edgeworth David and

Douglas Mawson spanned the years when geology

became established as a profession in Australia.

David was the pioneer in the professionalising of

the science, his department at Sydney, in the early

years of the century, producing a galaxy of

talented men and women from which Mawson

was to emerge as the shining star, not least

because of his international reputation as an

Antarctic explorer. David was also the first, and

highly successful, networker in Australian geology

and Mawson, a generation later, by establishing

his own equally effective department, extended

the net and through his effort and inspiration,

created an influence which is still potent at the

close of the twentieth century.

ACKNOWLEDGMENTS

I am indebted to Professor David Branagan of The

University of Sydney for his critical review of the

manuscript, and for the benefit of his extensive

knowledge of David; in particular, for suggesting

alternative interpretations of aspects of what was not

always a smooth relationship.

I wish to thank the following for their valuable help

during various stages of the preparation of this paper:

Mr Mark Pharaoh, Assistant Curator of the Mawson

Antarctic Collection, Urrbrae House, The University of

Adelaide; Mr Ken Smith, Archivist, The Fisher Library,

The University of Sydney; Mr Ben McHenry and Mr

Simon Langsford of the South Australian Museum; and

Mr Gerald Buttfield of the Department of Geology and

Geophysics, The University of Adelaide.

My thanks also to Dr Allan Pring, Curator of

Minerals, The South Australian Museum and Dr Robin

Oliver, Department of Geology and Geophysics, The

University of Adelaide, for their encouragement and

continuing interest in my Mawson researches.

70 D. W. CORBETT

REFERENCES

BRANAGAN, D. F. (ed.) 1973. ‘Rocks, Fossils, Profs:

Geological Sciences in the University of Sydney

1866-1973’. Science Press, Sydney. 184 pp.

BRANAGAN, D. F. 1981. Putting geology on the map.

Historical Records of Australian Science. 5(2): 31-

BRANAGAN, D. F. 1985. Earthy Sciences in ‘Ever

Reaping Something New—A Science Centenary’.

Ed. D. Branagan & G. Holland. The University of

Sydney, Sydney.

CAREY, S. W. 1990. ‘Sir Edgeworth David Memorial

Oration: “Knight Errant of Science”.’ Australasian

Institute of Mining and Metallurgy. Parkville,

Victoria. xiv, 74 pp.

CORBETT, D. W. 1998. Douglas Mawson: the

geologist as explorer. Records of the South

Australian Museum. 30(2): 107-136.

DAVID, T. W. E. 1897. Evidence of glacial action in

the Carboniferous and Hawkesbury Series, New

South Wales. Quarterly Journal of the Geological

Society of London 43: 190-196.

DAVID, T. W. E. & W. HOWCHIN. 1897. Notes on

the glacial features of Inman Valley, Yankalilla and

Cape Jervis districts. Transactions of the Royal

Society of South Australia. 21: 61-67.

DAVID, T. W. E. (with R. HELMS & E. F. PITTMAN)

1901. Geological notes on Kosciusko, with special

reference to evidence of glacial action. Proceedings

of the Linnean Society of New South Wales 26: 26-—

TA.

DAVID, T. W. E. & R. PRIESTLEY, 1914. Geology

Vol. 1. Reports on the scientific investigations, the

British Antarctic Expedition, 1907-08.

DAVID, T. W. E. 1922. Occurrence of remains of small

crustacea in the Proterozoic? or Lower Cambrian?

rocks of Reynella near Adelaide. Transactions of the

Royal Society of South Australia. 46, 6-8.

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. 52: 191-209.

DAVID, T. W. E. 1932. ‘Explanatory notes to

accompany a new geological map of the

Commonwealth of Australia’. Australasian Medical

Publishing Company, Sydney.

DAVID, T. W. E. & R. J. TILLYARD. 1936. ‘Memoir

on fossils of the late Pre-Cambrian (Newer

Proterozoic) from the Adelaide Series, South

Australia’. Angus and Robertson, Sydney. 122 pp,

pls I-XIII.

DAVID, T. W. E. 1950. ‘The Geology of the

Commonwealth of Australia (edited and much

supplemented by W. R. BROWNE)’. 3 vols. Edward

Arnold, London.

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.

INNES, M. & H. DUFF. 1990. ‘Mawson’s Papers’. The

Mawson Institute for Antarctic Research, The

University of Adelaide.

JACKA, F. & E. JACKA. (Eds.). 1991. ‘Mawson’s

Antarctic Diaries’. Allen & Unwin, Sydney.

MACLEOD, R. 1988. ‘Full of Honour and Gain to

Science’: Munitions Production, Technical

Intelligence and the Wartime Career of Sir Douglas

Mawson, F.R.S. Historical Records of Australian

Science. 7(2): 189-204.

MAWSON, D. & T. H. LABY. 1904. 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.

MAWSON, D. 1905. The geology of the New Hebrides.

Proceedings of the Linnean Society of New South

Wales. 3: 400-485.

MAWSON, D. 1906. On certain new mineral species

associated with carnotite in the radioactive orebody

Olary. Transactions of the Royal Society of South

Australia. 30: 188-193.

MAWSON, D. 1912. Geological investigations in the

Broken Hill area. Memoir of the Royal Society of

South Australia. 2(4): 211-319.

MAWSON, D. 1916. Petrology of rock collections from

the mainland of south Victoria Land. Reports of the

British Antarctic Expedition, 1907-09, Geology. 2:

201-237.

MAWSON, D. 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.

MAWSON, D. 1935. Sir Tannatt William Edgeworth

David 1858-1934. Obituary notices of the Royal

Society of London. 4: 493-501.

SHROCK, R. R. & W. H. TWENHOFEL. 1953.

‘Principles of Invertebrate Paleontology’. McGraw

Hill, New York.

SPRIGG, R. C. 1989. ‘Geology is Fun’. Published by

the author, Adelaide.

WALCOTT, C. D. 1914. Pre-Cambrian Algonkian algal

flora. Smithsonian Miscellaneous Collections. 64(2):

77-118.

WARD, L. K. 1925. Notes on the geological structure

of Central Australia. Transactions of the Royal

Society of South Australia. 49: 61-84.

The David—Mawson correspondence quoted in this

paper is to be found in the Mawson Papers housed in

the Urrbrae House Historic Precinct, Waite Campus,

The University of Adelaide. It is catalogued under

Control Number 24 DM, Inventory Number 4 (see Innes

and Duff 1990 pp. 5/8,5/9.)

The Mawson-David correspondence is housed in the

David archive at the Fisher Library, The University of

Sydney.

NOTE ON THE ORIGIN OF FRESHWATER CRAYFISH OCCURING ON

KANGAROO ISLAND.

BY W. ZEIDLER

Summary

ZEIDLER, W. (2000). Two species of freshwater crayfish occur on Kangaroo Island. The common

yabbie Cherax destructor Clark, 1936 and the marron, Cherax tenuimanus (Smith, 1912).

NOTE ON THE ORIGIN OF FRESHWATER CRAYFISH OCCURRING

ON KANGAROO ISLAND, SOUTH AUSTRALIA

Two species of freshwater crayfish occur on

Kangaroo Island. The common yabbie Cherax

destructor Clark, 1936 and the marron, Cherax

tenuimanus (Smith, 1912).

The marron, a riverine species native to south

Western Australia, is an obvious introduction.

Marron are highly desirable as an aquaculture

species (Morrissy 1984) and were introduced to

Queensland and northern New South Wales in

1979 (Austin 1985). They were introduced to

Kangaroo Island, for aquaculture, in the early

1980s but soon escaped and by the mid 1980s

became established in the nearby Ravine des

Casoars. Marron have since spread, or have been

translocated, so that they are now also well

established in Rocky River and the De Mole River

with some patchy occurrences in Middle River

and Cygnet River (Nicholls 1995) and probably

also occur in other permanent water on the Island.

The origin of Cherax destructor is more

difficult to determine. In 1979 and 1981 I

conducted extensive surveys of Kangaroo Island

in order to determine the presence of freshwater

crayfish and to sample for other freshwater

invertebrates. I found populations of Cherax

destructor in most freshwater streams on the

eastern half of the Island, extending as far west as

the upper reaches of the South-West River

(Zeidler 1982). Although I did not take

quantitative samples at the time I noted that C.

destructor was less abundant as I travelled west of

Cygnet River suggesting that the species was

spreading west or had reached some geographical

barrier at the South-West River. There seemed no

reason for its absence in apparently suitable

habitats in the Flinders Chase region. At the time

I assumed that the Kangaroo Island population of

C. destructor had been introduced or had become

isolated when the Island separated from the

mainland. The species is relatively common on

the Fleurieu Peninsula.

Campbell et al. (1994) who conducted genetic

studies of Cherax destructor and C. albidus from

South Australia and western Victoria, also

examined a Kangaroo Island population from

Stunsail Boom River. They found that the

Kangaroo Island population lacked the genetic

divergence characteristic of most of the mainland

populations. A lack of genetic divergence is

typical of introduced populations and they

concluded that the crayfish populating Kangaroo

Island are not endemic but have been introduced.

Recently I came across some _ old

correspondence, addressed to Herbert Hale, dated

27 Feb. 1946 which states that some 20 years

previous yabbies were introduced to ‘Cygnet

River about one mile up from the bridge on the

Kingscote-Hog Bay main road’ from ‘the Torrens

at West Marden’. The same correspondence states

that more specimens were introduced to Cygnet

River in about 1938 and that some were also put

in the Middle River.

Despite the introduction of yabbies to Cygnet

River in about 1926 none were caught almost 20

years later (‘we have tried in the Cygnet River on

several occasions, to see if we could catch

yabbies, but have not had any luck’) indicating

that the yabbies had difficulty becoming

established. The fact that yabbies are now

abundant in Cygnet River, and are easily caught,

suggests that they were not present prior to the

above mentioned introduction. Their spread west

seems to have been a slow process.

In 1986 I received a number of specimens of

both C. tenuimanus and C. destructor caught in

the Ravine des Casoars from the park ranger at

Flinders Chase, Mr Terry Dennis. This was quite

alarming because apart from confirming the

establishment of the marron in this once crayfish-

free environment, it also confirmed that the

common yabbie, Cherax destructor, had also been

introduced as both were absent in 1981. Judging

by the topography of Flinders Chase and the

apparent slow rate at which C. destructor became

established in the eastern part of the Island a

deliberate introduction to Flinders Chase of this

species seems the only logical conclusion.

Without a base-line knowledge of the

freshwater fauna of Kangaroo Island prior to the

introduction of freshwater crayfish it is impossible

to ascertain the ecological implications of such

introductions. It is interesting to note that C.

destructor, a very successful and invasive species,

had difficulty in becoming established in Cygnet

River.

Despite local knowledge of the existence of

freshwater crayfish on Kangaroo Island since

about 1926 the only record in the literature up

until 1982 (Zeidler 1982) was that of Riek (1969)

who reported a species of Geocharax from

Kangaroo Island. Unfortunately he gave no

precise locality data and the record seemed

72 W. ZEIDLER

dubious. A search of collections in all Australian

museums failed to confirm the presence of this

genus on the Island (Gross et al. 1979) and no

specimens, or evidence (remains) of this genus

were found during my surveys of 1979 and 1981.

I am thus reasonably confident that Riek’s record

is erroneous. Most likely it was based on a

specimen labelled ‘KI’ referring to King Island,

Bass Strait, which is a known habitat for species

of this genus.

REFERENCES

AUSTIN, C. M. 1985. Introduction of the yabbie,

Cherax destructor (Decapoda: Parastacidae) into

Southwestern Australia. Western Australian

Naturalist 16: 78-82.

CAMPBELL, N. J. H., GEDDES, M. C. & ADAMS,

M. 1994. Genetic variation in the yabbies, Cherax

destructor and C. albidus (Crustacea: Decapoda:

Parastacidae), indicates the presence of a single,

highly sub-structured species. Australian Journal of

Zoology 42: 745-760.

CLARK, E. 1936. The freshwater and land crayfishes

of Australia. Memoirs of the National Museum 10:

5-58, pls 1-11.

GROSS, G. F., LEE D. C. & ZEIDLER, W. 1979.

Invertebrates. Pp. 129-137 in ‘Natural History of

Kangaroo Island’. Ed. M. J. Tyler, C. R. Twidale and

J. K. Ling. Royal Society of South Australia:

Adelaide.

MORRISSY, N. M. 1984. Crayfish research and

industry activities in Australia. Freshwater Crayfish

5: 534-544.

NICHOLLS, S. 1995. Factors affecting

temnocephalid distribution and abundance on

Kangaroo Island. Unpublished Honours Thesis,

Department of Environmental Biology, University

of Adelaide.

RIEK, E. F. 1969. The Australian freshwater crayfish

(Crustacea: Decapoda: Parastacidae), with

descriptions of new species. Australian Journal of

Zoology 17: 855-918.

SMITH, G. 1912. The freshwater crayfishes of

Australia. Proceedings of the Zoological Society of

London 10: 144-171, pls 14-27.

ZEIDLER, W. 1982. South Australian freshwater

crayfish. South Australian Naturalist 56: 36-42.

Wolfgang ZEIDLER, South Australian Museum, North Terrace, Adelaide, 5000. Records of the South Australian

Museum 33(1): 71-72, 2000.

SOUTH

AUSTRALIAN

MUSEUM

AUGUST 2000

ISSN 0376-2750

CONTENTS:

ARTICLES

J s"G; J PRIDEAUX.

Simosthenurus newtonae sp. nov., a widespread sthenurine kangaroo (Diprotodontia:

Macropodidae) from the Pleistocene of southern and eastern Australia.

17 -C.H.S. WATTS & A. PINDER

Two new species of Antiporus from Western Australia (Coleoptera: Dytiscidae).

2 RoC NELSON

Why Flutes on Boomerangs and Throwing Sticks?

2 (OAs SY AMIEIES)

Revision of Australian Chaetarthria Stephens (Coleoptera: Hydrophilidae).

33. P.G. JONES

Words to objects: origins of ethnography in colonial South Australia.

49 D.W. CORBETT

A staunch but testing friendship: Douglas Mawson and T. W. Edgeworth David.

NOTE

7 WW ZEIDEER

Note on the origin of freshwater crayfish occurring on Kangaroo Island,

South Australia

Published by the South Australian Museum,

North Terrace, Adelaide, South Australia 5000.

OlF

RIECOIRIDS

SOUTH

AUSTRALIAN

MUSEUM

VOLUME 33 PART 2

OCTOBER 2000

ISSN 0376-2750

CONTENTS:

ARTICLES

73. R.M.KRISTENSEN & B.S. MACKNESS

101

First record of the marine tardigrade genus Bati/lipes (Arthrotardigrada: Batillipedidae)

from South Australia with a description of a new species.

C. Hes: WATTS

Three new species of Tiporus Watts (Coleoptera: Dytiscidae) with redescriptions of

the other species in the genus.

E. GENTILI

The Paracymus of Australia (Coleoptera; Hydrophilidae).

M. BAEHR

A new species of the genus Lestignathus Erichson from Tasmania, with a note on the

Tasmanian species of Mecyclothorax Sharp (Insecta: Coleoptera: Carabidae: Licininae,

Psydrinae).

C.H.S. WATTS & W. F. HUMPHREYS

Six new species of Nirridessus Watts and Humphreys and Tjirtudessus Watts and

Humphreys (Coleoptera: Dytiscidae) from underground waters in Australia.

Published by the South Australian Museum,

North Terrace, Adelaide, South Australia 5000.

FIRST RECORD OF THE MARINE TARDIGRADE GENUS BATILLIPES

(ARTHROTARDIGRADA: BATILLIPEDIDAR) FROM SOUTH SUTRALIA

WITH A DESCRIPTION OF A NEW SPEICIES.

BY R.M. KRISTENSEN AND B.S. MACKNESS

Summary

KRISTENSEN, R.M. AND MACKNESS, B.S. (2000). A new species of marine tardigrade, Batillipes

lesteri sp. nov. is described from beach sand collected at below low tide at Henley Beach, Adelaide,

South Australia. Eighteen specimens including both sexes, four-toed larvae and juveniles were

recovered. The new species differs from all other members of the Batillipedidae by its combination

of toe patterns, fourth lateral projections caudal apparatus. It is the first member of the genus to be

described from South Australia and only the third species to be recorded from the Southern

Hemisphere.

FIRST RECORD OF THE MARINE TARDIGRADE GENUS BATILLIPES

(ARTHROTARDIGRADA: BATILLIPEDIDAE) FROM SOUTH AUSTRALIA

WITH A DESCRIPTION OF A NEW SPECIES

R. M. KRISTENSEN AND B. S. MACKNESS

Depication: To the late Alan Bird who provided the first illuminating

studies on South Australian tardigrades.

KRISTENSEN, R. M. & MACKNESS, B. S. 2000. First record of the marine tardigrade genus

Batillipes (Arthrotardigrada: Batillipedidae) from South Australia with a description of a new

species. Records of the South Australian Museum 33(2): 73-87.

A new species of marine tardigrade, Batillipes lesteri sp. nov. is described from beach sand

collected at below low tide at Henley Beach, Adelaide, South Australia. Eighteen specimens

including both sexes, four-toed larvae and juveniles were recovered. The new species differs

from all other members of the Batillipedidae by its combination of toe patterns, fourth lateral

projections and caudal apparatus. It is the first member of the genus to be described from

South Australia and only the third species to be recorded from the Southern Hemisphere.

Reinhardt M. Kristensen, Invertebrate Zoology Department, University of Copenhagen,

Universitetparken 15, DK-2100, Copenhagen, Denmark. Brian S. Mackness* School of

Biological Sciences, James Cook University, Townsville, Qld, 4811. *Current address: PO Box

560 Beerwah, Queensland, Australia, 4519 (email: megalania@ compuserve.com). Manuscript

received | December 1999.

The interstitial heterotardigrade genus Batillipes

was first described by Richters (1909) based on

specimens of B. mirus from Kieler Bay, in the

Baltic Sea. Eight years later, Hay (1917) described

B. caudatus obtained from algae from jetties at

Beaufort, North Carolina. This American species

was later incorrectly synonymised by Marcus

(1929) with B. mirus. Marcus (1946) described a

third species, B. pennaki from the Atlantic coasts

of North and South America. Since that time many

more new species have been discovered and

named: B. similis Schulz, 1955; B. carnonensis

Fize, 1957; B. littoralis, B. phreaticus Renaud-

Debyser, 1959; B. friaufi Riggin, 1962; B.

annulatus, De Zio 1962; B. bullacaudatus

McGinty and Higgins, 1968; B. gilmartini

McGinty, 1969; B. dicrocercus Pollock, 1970; B.

acaudatus, B.tubernatis Pollock, 1971; B.

noerrevangi, B. roscoffensis Kristensen, 1978; B.

adriaticus Grimaldi de Zio et al., 1979; B.

africanus, B. marcelli Morone De Lucia et al.,

1988; B. tridentatus Pollock, 1989; B. crassipes

Tchesunov and Mokievsky, 1995; B.

philippinensis, B. longispinosus and B. orientalis

Chang and Rho, 1997a,b. Most recently, Rho et

al. 1999 described a new species B. rotundiculus

and provided a key to eight batillipedid

tardigrades from Korea.

Mackness (1999) recorded the first Australian

members of Batillipes from beaches in Victoria

but was unable to identify the animals to species

level due to poor preservation. This paper presents

the first record of the genus from South Australia

and describes a new species based on 18

specimens collected subtidally from a beach in

South Australia. Furthermore, the paper describes

its life cycle including the four-toed larva,

juveniles, young adults as well as their sexual

dimorphism.

MATERIALS AND METHODS

Two sand samples (2 x 750 ml approx.) were

collected on 6 November, 1995 by one of us (BM)

at Henley Beach, Adelaide, South Australia (34°

55'S, 138° 30'E). One sample (Al) was taken at

the low tide level and the other (A2), was taken at

one metre water depth (subtidal). Tardigrades

were only obtained from the subtidal sample. The

tardigrades were sorted out alive at the

Queensland Museum by RMK, two days after they

were received. Each sample was fresh-water

‘shocked’ following the procedure set out by

Kristensen and Higgins (1984). This involved

soaking the sediments in fresh water for about 20

74 R. M. KRISTENSEN & B. S. MACKNESS

seconds and then swirling them around. This

osmotically incapacitated the tardigrades which

were collected after the heavier material had

settled by decanting off the liquid through a 63

um mesh filter. The meiofauna was sorted using a

binocular microscope (40-80 x magnifications)

and then examined using phase contrast

microscopes (1000 x magnifications). A few

tardigrades were removed and placed in sea water

where they quickly recovered. All drawings were

made using camera-lucida techniques on live

animals. The reason for using live material for

illustrations is that the lipoid eyes and the hard

structures in the pharyngeal disappear in all

known permanent mounts.

Live tardigrades adhering with their suction

discs to the cover glass were then preserved in 2%

buffered formalin added under the cover glass.

Permanent mounts were made by adding

specimens to a glycerine solution (4%) under

cover glasses. After two days dehydration, the

cover glasses were ringed with Glyceel.

Measurements were only made on permanent

mounts and taken to the nearest micron using an

ocular micrometer. Comparisons were made with

reference collections held in the Zoological

Museum of the University of Copenhagen

(ZMUC) and with original type descriptions from

the literature. Measurements for 12 specimens are

provided in Tables 1-3.

SYSTEMATICS

Order Arthrotardigrada Marcus, 1927

Family BATILLIPEDIDAE Ramazzotti, 1962

Revised family diagnosis

Arthrotardigrade with large median cirrus present

and secondary clava dome-shaped. Lateral cirrus

and primary clava with a common pedestal. Internal

cirrus and median cirrus with well-developed

cirrophorus, external cirrus with indistinct

cirrophorus. All cephalic cirri without scapus and

flagellum. With four toes (in larvae) or six toes (in

adults) of different lengths, with adhesive or

suction disc at terminus of toe stalk. Claws absent.

Cuticular seminal receptacles absent.

Discussion

This family had originally included the genera

Batillipes and Orzeliscus Marcus. In this paper

the family Orzeliscidae is considered as a sister

group of the family Halechiniscidae and not the

Batillipedidae. The Batillipidedae is therefore

currently regarded as generically monotypic.

Batillipes Richters, 1909

Generic diagnosis

As Batillipedidae is monogeneric, the generic

diagnosis is the same as that for the family.

Type species: Batillipes mirus Richters, 1909 by

monotypy.

Type locality: Kieler Bay, Baltic Sea.

Discussion

The original description of B. mirus was written

in German and perhaps this is the reason that this

excellent early description has been overlooked.

Batillipes mirus is a very large arthrotardigrade up

to 720 um (mean = 400-600 pm) with spade-

shaped suction discs, similar to those of B.

tubernatis illustrated by McKirdy (1975). The type

material of B. mirus was collected subtidally (20

m water depth) and not in sandy beaches as nearly

all later records for B. mirus worldwide. In the

comprehensive review by McKirdy (1975) of the

genus Batillipes, six species were examined

carefully. The American ‘B. mirus’ is a middle-

sized batillipedid (about 160 um) with ovoid or

round suction discs. It is very clear that these

animals are not the same as the type species. It

may well be that all tidal animals called ‘B. mirus’

are in fact B. caudatus described by Hay (1917)

from North Carolina. This species was later

incorrectly synonymised by Marcus (1929) with

B. mirus. The cosmopolitan distribution (Table 4)

of B. mirus must therefore be considered doubtful

and new, worldwide samplings are necessary.

Batillipes lesteri n. sp.

(Figs 1-3, Tables 1-3)

Material examined

6 females, 5 males, 4 juveniles and 3 four-toed

larvae collected sublittorally (1 m in water depth

from the mean low tide), medium coarse sand

from Henley Beach, Adelaide, South Australia.

The sand samples were collected by B.M. on 6

November 1995. Holotype and seven paratypes

will be deposited in the Queensland Museum. The

other paratypes will be deposited in the Zoological

Museum of the University of Copenhagen

(ZMUC). Gnly 14 specimens were measured. In

this paper, only the measurements of 12

specimens are given.

TARDIGRADE GENUS BATILLIPES 75

FIGURE 1. Batillipes lesteri n. sp. Holotypic female, ventral view. Scale bar equals 50 um. Abbreviations: an,

anus; bl, body length; bp, buccal projection; br, brain; bt, buccal tube; bw, body width; ca, caudal appendage; cE,

cirrus E; ec, external cirrus; ey, eye; go, gonopore; ic, internal cirrus; Ib,, the base of leg IV sensory structure; Is,-

Is,, leg I to leg III spines; Is,, the spine of leg IV sensory structure; mc, median cirrus; mg, midgut; mo, mouth; 00,

oocyte; ov, ovary; pc, primary clava; pl, placoid; pr,—pr,, lateral projections; sc, secondary clava; sl, stylet length;

ss, stylet support; sv, seminal vesicle; te, testis; to,-to,, toe 1 to toe 6.

716

R. M. KRISTENSEN & B.S. MACKNESS

TABLE 1. Morphometry of Batillipes lesteri n. sp. Females. All measurements to the nearest micron.

Specimen

Holotype

Female

(Specimen 1)

Female 2

large eyes

(Specimen 4)

Female 3

Part. destroyed

Female 4

Immature

(Specimen 5)

Body length

Body width

Buccal tube

Stylet length

Stylet support

Placoid

Median cirrus

Internal cirrus

External cirri

Lateral cirri

Primary clavae

Second. clavae

Cirri E

Leg IV base

Leg IV spine

Leg III spine

Leg II spine

Leg I spine

Buccal project.

Projection 2

Projection 3

Projection 4

Caudal apparatus

apparatus

(width x length)

Leg 1 (toe 1)

(toe 2)

(toe 3)

(toe 4)

(toe 5)

(toe 6)

Leg IV (toe 1)

(toe 2)

(toe 3)

(toe 4)

(toe 5)

(toe 6)

Gonopore/Anus

Ovary

178.2 (178)

72.8 (73)

25.0 (25)

23.1 (23)

6.0 (6)

7.2 (7)

27.7 (28)

29.7 (30)

23.2 (23)

37.8 (38)

14.5 (15)

5.6 (6)

23:2 (23)

23.1 (23)

16.5 (17)

13.0 (13)

12.5 (13)

10.0 (10)

9.9 (10)

12.513)

13.2'(13)

15.3 (15)

21.0 x 26.4

(21 x 26) bilobed,

each with 5 spikes

9.5 (10)

5.9 (6)

12.5 (13)

7.9 (8)

17.1 7)

10.5 (11)

13.8 (14)

19.8 (20)

8.5 (9)

13.2 (13)

22.4 (22)

13.4 (13)

11.0 (11)

3 eggs

6-6-6-6

142.2 (142)

65.8 (66)

23.7 (24)

22:5' (23)

6.0 (6)

24.4 (24)

25.3 (25)

17.8 (18)

35.6 (36)

14.9 (15)

5.1 (5)

21.5 (22)

19.2 (49)

15.0 (15)

12.1 (12)

11.9 (12)

10.5 (11)

7.7 (8)

8.1 (8)

10.3 (10)

15,2:(15)

16.8 x 19.2

(17 x 19) bilobed,

each with 3 spikes

9.5 (10)

5.7 (6)

14.1 (14)

7.8 (8)

17.2;07)

9.8 (10)

12.9 (13)

17.0 (17)

7.5 (8)

11.7 (12)

18.2 (18)

10.6 (11)

11.2 (11)

2 eggs

6-6-6-6

162.5 (163)

63.8 (64)

22.0 (22)

20.8 (21)

5.8 (6)

7.0 (7)

25.2;(25)

25.5 (26)

18.9 (19)

35.3 (35)

12.7 (13)

$2 (5)

24.4 (24)

18.3 (18)

15.8 (16)

10.2 (10)

10.1 (10)

9.7 (10)

8.2 (8)

11.4 (12)

12.3.2)

13.1 (13)

20.0 x 15.0

(20 x 15) bilobed,

each with 3 spikes

9.5 (10)

5.6 (6)

12.8 (13)

7.9 (8)

16.1 (16)

9.8 (10)

12.4 (12)

18.4 (18)

7.8 (8)

11,3) (11)

21.5 @2)

11.8 (12)

10.2 (10)

2 eggs

6-6-6-6

(Specimen 6)

115.8 (116)

49.7 (50)

20.2 (20)

19.5 (20)

4.8 (5)

6.0 (6)

19.2 (19)

213421)

13.8 (14)

29.4 (29)

12,2:(12)

6.1 (6)

21.0:@1)

13.4 (13)

12.2 (12)

10.4 (10)

10.0 (10)

9.5 (10)

7.5 (8)

8.9 (9)

9.5 (10)

1107-12)

16.0 x 12.9

(16 x 13) bilobed,

each with 4 spikes

9.5 (10)

§.2.(5)

12.0 (12)

15.0 (15)

10.0 (10)

10.8 (11)

15.2 (15)

5.8 (6)

10.2 (10)

16.0 (16)

9.8 (10)

rosette not present

immature

6-6-6-6

Toe formula

Diagnosis

Middle-sized Batillipes with large lipoid eyes,

swollen tips on lateral, internal and median cirri as

well as on fourth leg spine; enlarged fourth leg spine

with a van der Land body separating the cirrophore

from spinous part of the leg sense organ. The

prominent lateral projection between legs III and IV

is two-pointed in adults and juveniles, one-pointed

in larvae. The caudal apparatus is an ala-like

structure with thin cuticular fibres often covered with

detritus. This structure varies extremely from simple

bilobed to highly furcated (4-10 spikes). Slightly

sexually dimorphic (females larger than males);

females with wider gonopore-anus distance than

males, and females with two-valved cuticular

structures associated with the rosette gonopore

system. Uneven 3" (short) and 4" (long) toe on

fourth leg, all toes with ovoid suction discs.

TARDIGRADE GENUS BATILLIPES 77

50 um

Is, ca

FIGURE 2. Batillipes lesteri n. sp. Allotypic male, dorsal view. Scale bar equals 50 um. Abbreviations as for Fig. 1.

78 R. M. KRISTENSEN & B. S. MACKNESS

TABLE 2. Morphometry of Batillipes lesteri n. sp. Males. All measurements to the nearest micron.

Specimen Allotypic Male 2 Male 3 Male 4

male vent./gonopore lateral Imm. Testis

(Specimen 2) (Specimen 7) (Specimen 8) (Specimen 9)

Body length 133.7 (134) 174.2 (174) 129.0 (130) 112.5 (113)

Body width 66.8 (67) 135 (713) lateral view 47.8 (48)

Buccal tube 21.7 (22) 21.8 (22) 18.8 (19) 17.1.7)

Stylet length 19.8 (20) 21.0 (21) 17.5 (18) 15.3 (15)

Stylet support 5.9 (6) 6.2 (6) 5.36) 5,2 (5)

Placoid 6.6 (7) tel) TO.) 6.0 (6)

Median cirrus 23.0 (23) 28.1 (28) 22.8 (23) 19.8 (20)

Internal cirrus 23.1 (23) 28.9 (29) 23.0 (23) 22.2 (22)

External cirri 19.8 (20) 19.6 (20) 15.4 (15) 14.9 (15)

Lateral cirri 31.5 (32) 38.9 (39) 32.5 (33) 33.1 G3)

Primary clavae 13.8 (14) 15.0 (15) 13.0 (13) U7 C12)

Second. clavae §.2 (5) 53 (5) $1 (5) 4.9 (5)

Cirri E PAGAN) 22.5 (23) 21.0 (21) 20.2 (20)

Leg IV base 18.3 (18) 17.8 (18) 13.5 (14) 15.6 (16)

Leg IV spine 15.8 (16) 18.0 (18) 12.8 (13) 13.3: 13)

Leg III spine 10.5 (11) 13.2:(13) 10.2 (10) 9.8 (10)

Leg II spine 10.5 (11) 12.1 12) 9.8 (10) 9.4 (9)

Leg I spine 9.5 (10) 12.0 (12) 8.9 (9) 9.1 (9)

Buccal project. 8.0 (8) 9.5 (10) 6.2 (6) 7.27)

Projection 2 9.8 (10) 10.8 (11) 7.5 (8) 9.5 (10)

Projection 3 10.3 (10) 12.2;(12) 8.0 (8) 10.2 (10)

Projection 4 11.8 (12) 15.8 (16) 13.4 (13) 12.9 (13)

Caudal 16.2 x 17.8 15.8 x 16.0 11-5 x 13.0 15.0:x 13.2

apparatus (16 x 18) bilobed, (16 x 16) bilobed, (12 x 13) bilobed, (15 x 13) bilobed,

(width x length) each with 2 spikes each with 2 spikes each with 2 spikes each with 2 spikes

Leg 1 (toe 1) 9.9 (10) 10.1 (10) 8.5 (9) V2)

(toe 2) 5.9 (6) 6.0 (6) 5.0 (5) 4.9 (5)

(toe 3) 12.5 (13) 1§.3°:C15) 13.8 (14) 11.9 (12)

(toe 4) 7.9 (8) 9.2 (9) 8.2 (8) 6.1 (6)

(toe 5) L7G) 18.5 (19) 15:0 (15) 15-2 (15)

(toe 6) 9.2.9) 11.6 (12) 9.8 (10) 8.0 (8)

Leg IV (toe 1) 125/13) 14.1 (14) 10.2 (10) 10.1 (10)

(toe 2) 18.4 (18) 19.4 (19) 15.8 (16) 14.7 (15)

(toe 3) 8.5 (9) 7.5 (8) 6.2 (6) 5.2 (5)

(toe 4) 11.5 (12) 12.6 (13) 10.1 (10) 10.0 (10)

(toe 5) 21.0 (21) 21.7 (22) 18.2 (18) 15.3 (15)

(toe 6) 112d) 11.8 (12) 9.4 (9) 9.2 (9)

Gonopore/Anus 3:3 (3) 4.2 (4) 3.0 G) 2.9 (3)

Testis mature sperm mature sperm without seminal without seminal

vesicles vesicles

Toe formula 6-6-6-6 6-6-6-6 6-6-6-6 6-6-6-6

DESCRIPTION large cirrophore and swollen tip (28 um). Internal

cirri also with large cirrophores and swollen tips

Holotype (30 um) directed anteriorly. External cirri horn-

Adult female (Fig. 1) with body 178 pm long

measured excluding caudal appendage and 73 pm

wide between legs II and III. Head distinguished

from body by a constriction beneath lateral cirri

and primary clavae. Head width 45 ym between

bases of lateral cirri. Median cirrus unpaired, with

shaped (23 um long) with indistinct cirrophores.

Primary clavae moderately long (15 um), thick

and tube-shaped. Primary clava and lateral cirrus

with a common pedestal (cirrophore of lateral

cirrus). Primary clava with a thick cuticular

annulus (van der Land body) inside base. Lateral

TARDIGRADE GENUS BATILLIPES

TABLE 3. Morphometry of Batillipes lesteri n. sp. Larva and Juveniles. All measurements to the nearest micron.

(width x length) each with one spike

Leg 1 (toe 1) 5.2 (5) 5.2 (5)

(toe 2) 0 0

(toe 3) 11.2 (11) 12.1 (12)

(toe 4) 5.2 (5) 6.8 (7)

(toe 5) 0 0

(toe 6) 7.2 (7) 8.2 (8)

Leg IV (toe 1) 7.9 (8) 7.2 (7)

(toe 2) 14.2 (14) 14.0 (14)

(toe 3) 0 0

(toe 4) 0 0

(toe 5) 11.2 (11) 10.1 (10)

(toe 6) 8 .2 (8) 8.0 (8)

Gonopore/Anus lacking lacking

Toe formula 4-4-4-4 4-4-4-4

with blunt tip

Specimen Four-toed Four-toed Juvenile 2 Juvenile 3

larva (Fig. 3) larva 2 (Third instar) (Fourth instar)

(First instar) (First instar) (Specimen 11) (Specimen 12)

(Specimen 3) (Specimen 10)

Body length 76.5 (77) 78.5 (79) 85.8 (86) 105.2 (105)

Body width 29.7 (30) 27.9 (28) 29.7 (30) 43.8 (44)

Buccal tube 16.5 (17) 15.7 (16) simplex 18.2 (18)

Stylet length 13.8 (14) 15.2 (15) simplex 16.5 (17)

Stylet support 3.9 (4) 3.8 (4) simplex 5.0 (5)

Placoid 4.6 (5) 4.2 (4) simplex 6.3 (6)

Median cirrus 19.5 (20) 18.2 (18) 20.2 (20) 19.1 (19)

Internal cirrus 20.4 (20) 18.9 (19) 21.1 (21) 20.3 (20)

External cirri 10.5 (11) 10.2 (10) 13.2 (13) 14.4 (14)

Lateral cirri 29.7 (30) 28.9 (29) 29.7 (30) 30.7 (31)

Primary clavae 9.5 (10) 9.0 (9) 9.9 (10) 12.6 (13)

Second. clavae 2.6 (3) 3.7 (4) 3.3 (3) 5.0 (5)

Cirri E 19.8 (20) 19.0 (19) 19.1 (19) 20.1 (20)

Leg IV base 9.2 (9) 10.2 (10) 8.5 (9) 12.8 (13)

Leg IV spine 9.2 (9) 9.8 (10) 8.7 (9) 13.0 (13)

Leg III spine 9.9 (10) 9.5 (10) 9.2 (9) 9.3 (9)

Leg II spine 9.9 (10) 9.2 (9) 9.0 (9) 9.0 (9)

Leg I spine 8.5 (9) 8.0 (8) 8.7 (9) 8.7 (9)

Buccal project. 5.9 (6) 5.0 (5) 5.9 (6) 5.8 (6)

Projection 2 6.6 (7) 6.2 (6) 5.9 (6) 6.2 (6)

Projection 3 7.2 (7) 74 (7) 7.9 (8) 6.7 (7)

Projection 4 8.5 (9) 10.0 (10) 9.9 (10) 9.1 (9)

Caudal 6.2 x 4.6 8.2 x 10.5 6.8 x 5.2 11.2 x 12.7

apparatus (6 x 5) bilobed, (8 x 11) bilobed (7 x 5) bilobed, (11 x 12) bilobed,

each with two each with two

small spikes spikes

5.2 (5) 8.2 (8)

0 4.9 (5)

7.9 (8) 13.7 (14)

5.9 (6) 7.0 (7)

9.9 (10) 15.0 (15)

7.6 (8) 9.6 (10)

9.9 (10) 9.1 (9)

13.2 (13) 15.8 (16)

4.6 (5) 5.7 (6)

7.9 (8) 9.2 (10)

14.5 (15) 16.4 (16)

9.2 (9) 9.0 (9)

gonopore lacking/

anus present

5-6-6-6

gonopore lacking/

anus present

6-6-6-6

cirri long and tapered terminating in prominent

swelling (38 um). Secondary clavae (base 6 um)

located between internal and external cirri at

frontal edge of head. Two large lipoid eyespots

present only in live animal consisting of one very

large ball-shaped lipoid droplet. The hyalin

spherical structure is seen on the inside of the

external brain lobe (protocerebrum). It is supposed

they are of lipid composition based on their

solubility in alcohol and glycerol. Pharyngeal bulb

subcircular (18 um x 21 um) located between legs

I. In optical cross-section, bulb is trilobate, each

lobe with a calcium carbonate encrusted placoid

(7 um). Buccal tube straight and moderately long

(25 um) with (3 pm) width. Buccal tube extends

inside pharyngeal bulb attaching placoids via three

80 R. M. KRISTENSEN & B. S. MACKNESS

apophyses. Ventral mouth as in characteristic

pouting in typical Batillipes form (sensu

Kristensen 1978). Stylet supports straight (6 um)

with support knob showing slight deflection

anteriorly. Support knobs linked on large furca of

stylet. Stylets length 23 um.

Spines present on all legs. Spine of legs I

located more distally than all other spines. Spines

on legs I-IV located proximally to body.

Increasing size of spines from legs I-IV, 10.0 um,

12.5 pm, 13.0 um, 16.5 pm + 23.1 um

respectively. Fourth leg spine very long with large

base (cirrophore) and a cuticular annulus (van der

Land body) separating base from spine which is

the true sense organ. Total length of whole fourth

leg sensory structure 49.6 ym. First and fourth leg

spines with swollen tips as in cephalic cirri. Cirri

E moderately long (23 pm) and sharply pointed

with distinct cirrophore. Ventral tongue-shaped

projections in front of leg I (10 um), leg IT (13 pm

long) and leg III (13 ym long). Lateral projection

in front of leg IV bifurcate (15 pm long). Caudal

apparatus ala-like and strongly furcated (2 x 5

spikes). Cuticle of caudal apparatus has fine hairs;

it is covered with bacteria and detritus. Gonopore

rosette-shaped with six identical segments. Behind

the female gonopore there is a two valved

cuticular structure. Gonopore-anus distance is 11

pm. The holotypic female is sexually mature, with

three large eggs.

Legs I-IV possessing toes of varying lengths.

On leg I, toe 5 is the longest (17 um) with toes

decreasing in size 3, 6, 1, 4 and 2 (13 um, 11 pum,

8 um and 6 um respectively). On legs IV, toes 2

and 3 are unequal in size and conform to toe

pattern D (see Table 5). Length of toes in

decreasing order of size is 5, 22 um; 2, 20 um; I,

14 um; 4, 13 um; 6, 11 um and 3, 9 pm. Toe

discs ovate with conspicuous brace.

Allotype

Adult male (Fig. 2) with body 134 pm long

measured excluding caudal appendage, and 67 um

wide between legs II and II. Male is similar to

female except for size and differences in shape of

projection in front of leg IV which is slightly

bilobate, and caudal appendage lobate with only

four spikes. Gonopore-anus distance is only 3 ym,

meaning that the gonopore nearly connects the

anterior branch of a three-lobed anal system. Male

gonopore located on a small ovoid papilla. The

allotype is sexually mature with two lateral

seminal vesicles filled with mature spermatozoa.

The allotype is drawn in dorsal perspective.

Dorsal transverse lines indicating segmental plates

disappear after fixation. The punctations of the

dorsal cuticle (see McKirdy 1975) are very coarse,

especially laterally, where the epicuticular pillars,

which form the punctations, can be seen using

light microscopy. Each lateral pillar is about 2 ym

high.

Four-toed larva

The first instar of all species of Batillipes is so

different from subsequent instars, that the term

‘larva’ (sensu Bertolani et al. 1984) is correct.

The paratypic larva illustrated (Fig. 3, Table 3)

has a body length of 77 ym and body width of 30

um. It shows typical ontogenetic body proportions

with a larger head region in relation to body, lacks

the gonopore, the anus is only a slit, and it is not

three-lobed. The larva has only four toes instead

of the six toes of the adult. Toe 2 and 5 are

missing on legs I-III and toes 3 and 4 are missing

on leg IV. The leg spines I-III are with a swollen

tip, but the leg spine IV is pointed. The projection

4 has only one spike and the caudal appendage is

only slightly bilobed.

Juvenile

The animals without gonopore, but with three-

lobed anus are called juveniles. At least two

instars can be recognized but they could be more

numerous. The second instar has the toe formula

5-5-5-6 and the third instar has 5-6-6-6. All sense

organs have the characteristics of the adult, but

the projection in front of leg IV and the caudal

apparatus vary a lot. The differences between

males and females can be seen in the size and

especially in the caudal apparatus.

Young adults

Animals with gonopores, but with immature

reproductive systems are called young females and

males. These animals are smaller in size, but are

still sexually dimorphic. The young males lack the

seminal vesicles with mature spermatozoa while

the young females have only small-size oocytes in

the ovaries.

Life Cycle

There is no doubt that B. lesteri was in the

middle of its reproductive cycle when the

specimens were collected in November, which is

the last month in the Australian Spring. It is

unusual to have so many four-toed larvae and

juveniles in populations of Batillipes. The first

instar, the four-toed larva, differed dramatically

from the juveniles and adults. It is difficult to

interpret just how many instars are involved

TARDIGRADE GENUS BATILLIPES 81

Pr,

FIGURE 3. Batillipes lesteri n. sp. Paratypic four-toed larva. Scale bar equals 50 ym. Abbreviations as for Fig. 1.

R. M. KRISTENSEN & B.S. MACKNESS

TABLE 4. Zoogeographic distribution of species of the genus Batillipes. (Southern hemisphere records in bold).

Baws

B. acaudatus — England (Pollock, 1971)

B. adriaticus — \taly — (Grimaldi De Zio et al., 1979)

africanus — Libéria (Morone De Lucia et al., 1988)

. annulatus — Italy (De Zio 1962, Grimaldi De Zio et al. 1979)

. bullacaudatus — USA (McGinty & Higgins 1968, Pollock 1970b; Lindgren 1971; McKirdy 1975); Scotland

(McIntyre & Murison 1973)

. carnonensis — France (Fize 1957); Italy (Grimaldi De Zio er al. 1980, 1983); USA (Fleeger 1978)

. crassipes — Russia (Tchesunov & Mokievsky 1995); Korea (Rho et al. 1999)

. dicrocercus — USA (Pollock 1970a, McKirdy 1975, Hummon 1994); Italy (Hummon 1994; Grimaldi De Zio et

al. 1980, D’ Addabbo Gallo et al. 1987, Matarrese et al. 1996 ); Poland (Hummon 1994)

B. friaufi — USA (Riggin 1962, McKirdy 1975, Gaugler & Nelson 1997)

gilmartini - USA (McGinty 1969. Pollock 1989)

B littoralis — France (Renaud-Debyser 1959, D’Hondt 1970, Renaud-Debyser & Salvat 1963); Italy (Grimaldi De

Bas

Zio et al . 1983, D’ Addabbo Gallo et al 1987, D’ Addabbo Gallo et al. 1999, Grimaldi De Zio et al. 1999)

B. longispinosus — Korea (Chang & Rho 1997a, Rho er al. 1999)

B. mirus — Germany (Richters 1909); Wales (Boaden 1963); Ireland (Boaden 1966); Scotland (Pollock 1971;

marcelli — Italy (Morone De Lucia et al. 1988)

McIntyre & Murison 1973); Norway (Tambs-Lyche 1939-40); Denmark (Fenchel et. al 1967, Fenchel

1969, Kristensen 1978); Finland (Purasjoki 1953, Karling 1954-1955); Sweden (Jaégersten 1952); North

Sea (Remane 1940, Freidrich 1963); Black Sea (Plesa 1963); Germany (Schmidt 1969) Bulgaria

(Valkanov 1950, 1954); Romania (Rudescu 1964); Russia (Petelina & Tchesunov 1983, Biserov 1991);

France (Baudoin 1952, Swedmark 1956 a,b, Renaud- Debyser 1956, Guérin 1960, Renaud-Mornant &

Jouin 1965; D’Hondt 1970,Renaud-Debyser & Salvat 1963); Italy (Papi 1952, D’Addabbo Gallo et al.

1987); Madagascar (Renaud-Mornant 1979); USA (Hay 1917, King 1962, McGinty & Higgins 1968,

Pollock 1970a, Lindgren 1971, McKirdy 1975, Meyer cited in Pollock 1989, Pollock 1989, Gaugler &

Romano 1995, Gaugler & Nelson 1997); Bahamas (Pollock 1970b); Malaysia (Renaud-Mornant &

Seréne 1967)

. noerrevangi — Denmark (Kristensen 1976, 1978)

. orientalis — Korea (Chang & Rho 1997, Rho et al. 1999))

. pennaki — Massachusetts, USA (Marcus 1946, Pollock 1970b, McKirdy 1975); Brazil (Marcus 1946); France

(Renaud-Debyser 1959,Renaud-Debyser & Salvat 1963); Italy (De Zio 1962, 1964, Grimaldi de Zio &

D’Addabbo Gallo 1975, Grimaldi De Zio et al. 1979, Bertolani et al. 1984, D’Addabbo Gallo et al.

1987); Spain (Villora-Mofeno & Grimaldi de Zio 1993); India (Rao & Ganapati 1968); Korea (Rho et al.

1999)

. philippinensis — Philippines (Chang & Rho 1997b); Korea (Rho et al. 1999)

phreaticus — France (Renaud-Debyser 1959, Renaud-Debyser & Salvat 1963); Germany (Riemann 1966);

England (Pollock 1971); Spain (Villora-Moreno & Grimaldi De Zio 1993); Italy (D’Addabbo et al.

1987).

roscoffensis — France (Kristensen 1978)

rotundiculus — Korea (Rho et al. 1999)

similis — Germany (Schulz 1955); Korea (Rho et al. 1999) France (Fize 1963); Italy (Grimaldi De Zio et al.

1980, D’ Addabbo Gallo ert al. 1999, Grimaldi De Zio et al. 1999)

tridentatus — Washington & California, USA (Pollock 1989); Korea (Rho et al. 1999)

tubernatis — Scotland (McIntyre & Eleftheriou 1968, Pollock 1971); Germany (Riemann 1966, Hummon 1994);

USA (McKirdy 1975)

before an animal is sexually matured but at least

four moults are present before the gonopore is

seen. The following instars have been recorded

for this species. First instar -the four toed larvae;

Second instar — juvenile with toe formula 5-5-5-6;

Third instar — juvenile with toe formula 5-6-6-6;

Fourth instar juvenile without gonopore but with

toe formula 6-6-6-6; Adult—with gonopore and

three-lobed anus.

Etymology

Named in honour of curator Dr Lester R. G.

Cannon, Queensland Museum, who facilitated our

collaboration.

TARDIGRADE GENUS BATILLIPES 83

TABLE S. Patterns of toe length on fourth foot of species of Batillipes. (Modified after Pollock 1970a)

A B

(Il & IV equal) (I & HI equal)

c D

(II & IV equal) (IIT & IV unequal)

. acaudatus

. adriaticus

. annulatus

. bullacaudatus

carnonensis

crassipes

dicrocercus

. gilmartini

longispinosus

marcelli

B. phreaticus

mirus

noerrevangi

orientalis

pennaki

Philippinensis

roscoffensis

. rotundiculus

. tridentatus

SYSTEMATIC DISCUSSION

Species of the genus Batillipes are amongst the

most studied of all marine tardigrades. Their

taxonomy remains problematic with relatively few

characters commonly used in systematic

investigations of the group (McKirdy 1975). Some

of the characters used (e.g. conformation of lateral

body projections, relative length and shape of

cephalic appendages), are directly affected by the

physical mounting process. Total body length and

width may vary due to cover slip pressure;

shrinkage can occur due to the mounting medium

and lengths of spines and cirri may be

miscalculated because of their orientation to the

observer. Internal features such as eyespots and

buccal apparatus can also be affected by mounting

media and cover slip pressure. Apart from these

mechanical problems, there are also a range of

other factors to contend with. The caudal

appendage of Batillipes has often been used in the

diagnosis of species but various authors have

shown this can vary ontogenetically (Grimaldi de

Zio and D’Addabbo Gallo 1975; Morone De

Lucia et al. 1988). Further ontogenetic variation

has been recorded by Kristensen (1978), McGinty

and Higgins (1968) and Villora-Morena and

Grimaldi de Zio (1993).

Comparative studies have also been made

difficult because of the inconsistent nomenclature

of toes and the lack of a full range of specimens

B. friaufi

B. littoralis

B. africanus

B. lesteri

B. tubernatis

B. similis

of both sexes, four-toed larvae and juveniles for

many species. In this study, we have used the

standard method of numbering toes on the fourth

leg with toe 1 being the closest cranially on the

right hand side and the toes then numbered

sequentially in a clockwise direction. This makes

the caudoproximal toe, toe 6 in our scheme (Table

5). Furthermore, the attachment of the toes to the

tarsus has systematic value in determining which

toe is missing in larvae and in juveniles. At leg 1,

the 2nd and 4th toes are ventral, 1st and 6th lateral

and 3rd and Sth dorsal on the tarsus. This

distribution is functionally correct: the shortest

toes are ventral and the longest dorsal.

The relative position of these toes, particularly

those on right leg IV has been used as an

important taxonomic indicator (Pollock 1970a).

Even here problems can arise if the legs are not

properly positioned or incorrectly illustrated. It is

therefore necessary to have a range of specimens,

of both sexes, including juveniles if any

meaningful taxonomic decisions are to be made.

Batillipes lesteri n. sp. differs from all other

species of Batillipes by having uneven 3" and 4"

toes on leg IV except for B. africanus, B. similis

and B. tubernatis. It differs from B. tubernatis by

having a caudal apparatus with fine cuticular hairs

and differs from B. africanus by the males having

bilobate caudal appendages with four spikes and

females having highly furcated (6-10 spikes)

caudal appendages. It differs from B. similis by

84 R. M. KRISTENSEN & B.S. MACKNESS

having different primary clava as well as caudal

and lateral processes.

Ontogenetic differences between other species

of Batillipes are seen in the toe formula. Larvae of

B. noerrevangi lack toe 2 (the smallest one of the

adult) and toe 6 on leg I. Larvae of B. lesteri also

lack toe 2 but instead of toe 6, they lack toe 5 (the

longest toe in the adult) on leg I.

Marine tardigrades also go through a life cycle

with a number of moults and will vary

morphologically at different stages of this cycle

including a process called cyclomorphosis

(Kristensen 1982) Furthermore the species may

vary in morphology according to differences in

salinity (Kristensen 1978). A proper analysis of

both sexual and ontogenetic variations is

necessary in many instances before new taxa

should be raised.

The new species of Batillipes is only the third

recorded for the Southern Hemisphere. A

zoogeographic analysis of the 24 species recorded

so far (Table 4) shows a singular distributional

bias towards the northern hemisphere. This is

probably more an artefact of where marine

tardigradologists have been working and

collecting rather than any zoogeographic pattern.

In this analysis, most Batillipes species are still

primarily known from their type localities with

only three species B. mirus, B. pennaki and B.

phreaticus having more cosmopolitan

distributions.

Several studies have shown that certain species

of Batillipes live sympatrically occupying specific

regions of the littoral environment (Renaud-

Debyser 1959, Schmidt 1969, Lindgren 1971).

Furthermore, Renaud-Debyser (1959) and Pollock

(1975) have demonstrated that Batillipes shows

tolerances and preferences to water content,

temperature, light and salinity. This predicates that

any understanding of the distribution of species of

Batillipes at any location must take all these

factors into account. Some beaches may have an

almost monotypic batilliped fauna while others

may support several species in different locations.

Further ecological studies and collecting of

Australian marine tardigrades is needed to clarify

whether these patterns hold true in southern

latitudes.

ACKNOWLEDGMENTS

The authors wish to thank the Australian Biological

Resources Study (ABRS), the whole curatorial staff and

especially Dr Lester R. G. Cannon of the Queensland

Museum for the provision of facilities and ABRS

Fellowship during the one year stay (1995-1996) of one

of us (RMK). Stine Elle and Aslak Jgrgensen (ZMUC)

undertook the illustrations, and their work was enabled

by a grant to RMK from the Carlsberg Foundation

(Grant No. 970345/30-488). Dr Susanna Grimaldi de

Zio, Dr Leland Pollock and Dr Woody Horning Jr

provided useful comments on the manuscript.

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THREE NEW SPECIES OF TIPORUS WATTS (COLEOPTERA:

DYTISCIDAE) WITH REDESCRIPTIONS OF THE OTHER SPECIES IN

THE GENUS.

BY C.H.S. WATTS

Summary

WATTS, C.H.S. (2000). Three new species of Tiporus Watts, 1985 (Coleoptera: Dytiscidae) are

described and figured: Tiporus georginae, T. lachlani and T. moriartyensis. The original

descriptions of the existing species were based on a small number of speciamens. Recent

collections have allowed these descriptions to be reviewed and, where necessary, corrected and/or

enlarged. A key is given to the 11 known species.

THREE NEW SPECIES OF TIPORUS WATTS (COLEOPTERA: DYTISCIDAE)

WITH REDESCRIPTIONS OF THE OTHER SPECIES IN THE GENUS

C.H.S. WATTS

WATTS, C. H. S. 2000. Three new species of Tiporus (Coleoptera; Dytiscidae) with

redescriptions of the other species in the genus. Records of the South Australian Museum

33(2): 89-99.

Three new species of Tiporus Watts, 1985 (Coleoptera: Dytiscidae) are described and

figured: Tiporus georginae, T. lachlani and T. moriartyensis. The original descriptions of the

existing species were based on a small number of specimens. Recent collections have allowed

these descriptions to be reviewed and, where necessary, corrected and/or enlarged. A key is

given to the 11 known species.

C. H. S. Watts, South Australian Museum, North Terrace, Adelaide, South Australia 5000.

Manuscript received 11 October 1999.

The genus Hypodes Watts, 1978, later changed

due to preoccupation to Tiporus Watts, 1985, was

erected for a group of small to moderate sized

Australian Hydroporini related to Antiporus

Sharp. Members of Tiporus are distributed across

coastal Northern Australia where they appear to

be restricted to rivers and streams, or to the small

pools which form in their beds in the dry season.

In these habitats some species, such as T.

undecimmaculatus and T. josepheni, are often

abundant.

The genus has not been reviewed since my

1978 paper. Many more specimens are now

available, among them the three new species

described below. The taxonomy of the genus is

based on the males, which have distinctive,

species-specific genitalia and secondary sexual

characters on the legs. Females are more difficult

taxonomically but the additional material has

allowed these to be tentatively keyed for the first

time. Unfortunately, in the case of T. josepheni, I

mis-associated the sexes in my original

description, which has led to confusion over the

identification of this species. The type material of

all species except TJ. collaris and T.

undecimmaculatus has been restudied.

The collections from which specimens were

examined are listed under the following

abbreviations:

ANIC Australian National Insect Collection,

Canberra

NIM Northern Territory Museum and Art

Gallery, Darwin

QM Queensland Museum, Brisbane

SAMA _ South Australian Museum, Adelaide

WAM __ Western Australian Museum, Perth

QDPIM Queensland Department of Primary

Industries, Mareeba

SYSTEMATICS

The genus Tiporus Watts belongs to that section

of Australian Hydroporini with pseudotetramerous

protarsi and densely rugose/punctate bodies

including the legs. It can be distinguished from

the other genera in this section, Antiporus Sharp

and Sekaliporus Watts, by the form of the humeral

angle of the elytron and the three-segmented

protarsi in the male (see Watts 1997 for more

details).

Within the genus there are two morphological

groupings, one group characterised by: lack of

pronotal ridges/grooves, moderate to strong

dorsal colour pattern, a tendency for the edge of

the elytron to bulge or extend outwards near the

tip, protibia of the male with a small tubercle

on the outside near the base, and the other

group characterised by: pronotal ridges/grooves,

dark colouring with only vague reddish patches,

edges of the elytron serrated but not extended

and the male protibia having a broad spine in

the middle or towards the apex on the outside.

To what degree these groupings reflect

phylogenetic relationships has yet to be

determined.

C. H.S. WATTS

Key To Tiporus WATTS

Disc of pronotum with a ridge/groove

running backwards parallel to and at a

little distance from sides; edge of elytron

serrated in apical half, never with lateral

extension near tip; sides of pronotum dark,

at most diffusely lighter in colour....... q

Disc of pronotum smooth, at most with

weak ridges; edges of elytron usually

smooth but often with rounded or triangular

extension near tip; sides of pronotum

usually with well-defined light coloured

Female with edge of elytron near apex

with large, triangular extension reaching

beyond tip of elytron (Fig. 8); male without

such an extension, protibia with triangular

spine on outside near middle ................

Seay aan ameree ae T. josepheni (Watts)

Both sexes with elytron with at most a

modest bulge near tip or with the tip

extended; males with small tubercle on

outside of protibia at base................... 3

Larger (4.5 mm long); postcoxal lines

narrow, reaching metathorax; outer

metatibial spine in male greatly elongated,

about twice the length of the inner (Fig.

12); (female unknown) s:-s2-<2-..4e00:

inteetiesttcicsurees T. moriartyensis sp. nov.

Smaller (< 4.1 mm long); postcoxal lines

wide, often poorly defined towards front;

outer metatibial spine in male normal, not

greatly different in length to inner...... 4

Female with tip of elytron clearly extended

(Fig. 15); edge of pronotum weakly sinuate

in front; male with metatibia expanded on

INSIDE NEA APEX ee o.ces ees e see skceesettveeseces

Pin on teurra re T. tambreyi (Watts)

Tip of elytron not, or only weakly, extended

(edge of elytron may be moderately

expanded near, but not at, tip); sides of

pronotum with only a hint of sinuation;

male with normal metatibia ................ 5

Extreme front of elytron considerably

narrower than adjacent pronotum,

difference > width of mid segments of

antenna (Fig. 4): lateralextensiontoelytron

near tip absent or weak (Fig. 4); male with

very long dagger-shaped proclaw (Fig. 7)

PE et ct fois Ride ies T. georginae sp. nov.

Extreme front of elytron only slightly

narrower than adjacent pronotum,

difference <= width of mid segments of

antenna; edge of elytron with moderate

bulge near tip; males with either short

stout proclaws or with moderately long

but thin proclaws (Figs 13, 14).......... 6

Pronotum weakly ridged at sides; edge of

elytra straight immediately prior to it

meeting pronotum; male proclaw short,

thickened basally (Fig. 13 ) «0... eee

ee adie teeta T. denticularis (Watts)

Pronotum smooth; edge of elytra slightly

curved immediately prior to it meeting

pronotum; male proclaw elongate, thin

CHiN e teen, T. centralis (Watts)

Large (> 5.0 mm long); male proclaw

thick (Fig. 2), protibia stout, with basal

spine (Fig. 3); apical segment of protarsi

large, deeply bifid (Fig. 3); Pilbara region

Of WEA Teh cea T. lachlani sp. nov.

Smaller (< 5.1 mm long); males not as

ADOVE ssescscesvestysays catessersarscsseetecseesemnczses 8

Tip of elytron with small but distinct point

(except in some T. alastairi); elytral

serrations usually strong; male protibia

with prominent, triangular enlargement in

apical glial fgswretes treet rte seer ruses tse et e)

Tip of elytron rounded; elytral serrations

weak; male protibia with small spine/

tubercle on inside close to base ............

es evn ette eee ee ees cise T. collaris (Hope)

Viewed laterally, the edge of elytra curves

forward for short distance prior to meeting

pronotum; male protibia with large

triangular expansion on front edge near

apex; male proclaw squat ............. cee

Paty ueese assess eat suri oor ee. T. alastairi (Watts)

Viewed laterally, the edge of elytra not, or

only slightly, curved immediately prior to

meeting lateral edge of pronotum; male

protibia with small triangular expansion

on front edge in middle or towards apex;

mialemroclawethiniees 2...) ee. 10

Male with tip of central lobe of aedeagus

pointed (Fig. 20); protibia with triangular

expansion just beyond centre................

eee ee T. giuliani (Watts)

Male with tip of central lobe of aedeagus

broad, weakly tridentate (Fig. 16); protibia

with triangular expansion on front edge,

SPECIES OF TIPORUS 91

either in middle (N. Qld.) or towards apex

CNETAWLAS IER cere care ene men

Eee reo T. undecimmaculatus (Clark)

Tiporus georginae sp. nov.

Types

Holotype: male: ‘W. Aust. Mitchell Plateau.

14°40'S 125°44'E 23 Sept 1982. B. V. Timms’,

dissected and mounted on card, SAMA.

Paratypes: 5, “14°52’S 125°50’E WA ‘The

Crusher’ CALM Site 9/1 4 km S by W Mining

Camp Mitchell Plateau 2-6 June 1988 I. D.

Naumann’, ANIC; 4, “14°25’S 126°40°’E CALM

Site 4/3 14 km S by E Kalumburu Mission W. A.

3-6 June 1988 T. A. Weir’, ANIC (3), SAMA

(1). All labelled as collected at light in open

forest except for one in closed forest at CALM

site 4/3.

Description Figs 4-7

Length 3.5 — 4.0 mm. Elongate-oval, wider in

middle, front edge of elytron considerably

narrower than rear edge of pronotum (Fig. 4).

Pronotum smooth, without lateral ridges or

grooves; edge of elytron turns forward sharply just

before meeting edge of pronotum, very weakly

serrate towards rear, usually slightly expanded

near but not at tip, quite strongly acuminate.

Whole body strongly rugose/punctate. Dorsal

surface dark testaceous-black, front margin of

head, sides and parts of front and rear margins of

pronotum, three lateral spots and often some spots

inwards from these on elytron, testaceous. Ventral

surface dark testaceous-black, appendages

including pro- and mesocoxae, much lighter.

Pronotal process narrow, keeled, bluntly pointed.

Metacoxal lines relatively close in hind half,

widening to a bit more than two times their

narrowest width in front quarter.

Male. Protarsi three-segmented, with adhesive

setae ventrally, anterior parts of segments

expanded, small spine on hind apical corner of

second segment, third segment about 2.5 times

length of second, single claw extremely long,

expanded in middle (Fig. 6). Protibia with small

tubercle on anterior edge at base (Fig. 7).

Mesotarsi as for female. Metatibia with a slight

thickening on inside near apex. Apical ventrite

with small, sharp, apical keel/spine. Median lobe

of aedeagus broad in centre, very narrow in apical

quarter (Fig. 5).

Female. Protarsi five-segmented, first three

moderately expanded with adhesive setae

ventrally, apical segment cylindrical. Mesotarsi

much narrower.

Remarks

Resembling T. centralis and T. denticularis in

size and colour but with the mis-match of the

humeral angle of the elytron and the posterolateral

angle of the pronotum more noticeable (Fig. 4).

Some specimens of T. centralis have this feature

to a degree but the extremely long, dagger-like

male proclaw in T. georginae (Figs 6, 7) readily

separates male specimens from this and other

Tiporus. The aedeagus is distinctive (Fig. 5). The

species is known only from the Kimberley region

where it is broadly sympatric with T. centralis.

Distribution

The Kimberley region of Western Australia.

Tiporus moriartyensis sp. nov.

Types

Holotype: male, ‘Moriarty ck N.T. Reardon 8/

82’, SAMA.

Description Figs 9-12

Length 4.5 mm. Elongate-oval, widest in

middle. Pronotum narrower than elytra. Dorsal

surface dark brown-black, rear of head, lateral

margins and rear margin of pronotum, three

small lateral spots and two larger spots near

suture on elytron testaceous. Ventral surface

mottled dark and light testaceous, appendages

including pro- and mesocoxae much lighter.

Whole body covered with rugose punctures,

punctures smaller and less rugose than in other

Tiporus species. Disc of pronotum smooth,

without lateral ridges or grooves. Side of elytron

curves forward immediately before meeting edge

of pronotum, very weakly serrate towards tip,

apex obliquely truncated. Pronotal process

relatively broad, keeled, tip blunt. Metacoxal

lines close, diverging to about two times their

minimum width towards front, reaching

mesosternum.

Male. Protarsi three segmented, segments

expanded in front, with dense adhesive setae

ventrally, second segment with small tooth on

hind margin, apical segment 2.5 times length of

second; claw short, rapidly narrowing to about

half its width in middle (Fig. 10). Protibia with

weak tubercle/spine near base on anterior edge.

Metatibia with inner apical spine, twisted,

flattened and broadened slightly at tip, about two

92 C.H.S. WATTS

FIGURES 1-15. 1-3, dorsal view of median lobe of aedeagus, apical view of proclaw and anterior view of protibia

and tarsi of T. lachlani; 5-7, ditto T. georginae; 9-11, ditto T. moriartyensis; 4, dorsal outline of T. georginae; 8,

dorsal outline of elytron of T. josepheni; 12, metafemur of T. moriartyensis; 13, apical view of proclaw and third

tarsal segment of 7. denticulatus; 14, ditto T. centralis; 15, dorsal outline of elytron of T. tambreyi.

SPECIES OF TIPORUS 93

times length of outer, which is simple (Fig. 12).

Apex of last abdominal ventrite broadly depressed

in middle, rear edge with two small spines in

middle. Median lobe of aedeagus broad in middle,

abruptly narrowing near apex (Fig. 9).

Female. Not known.

Remarks

Although only one specimen is available this is

enough to show that 7. moriartyensis is a very

distinctive species. It belongs to the group of

species lacking lateral pronotal ridges but is

noticeably larger and less strongly punctured than

other members. No other Tiporus has the inner

apical spine on the metatibia modified as in this

species. The female is unknown but it is likely

that this character is restricted to the male.

Distribution

Known only from the type locality—Moriarty

Creek, 160°4'S 129°12'E—in the Northern

Territory.

Tiporus lachlani sp. nov.

Types

Holotype: male, ‘Wooramel R. WA 25°47.52'S

115°17.44'E 26.8.94 S.A. Halse’, SAMA.

Paratypes: 1, female, as for holotype, SAMA;

1, male, ‘Gregory Gorge Fortescue R W.A. 2

xii.74 coll. K. F. Walker’, SAMA.

Description Figs 1-3

Length 5.6 mm. Elongate-oval, widest behind

middle, outline strongly indented at junction of

pronotum and elytra. Lateral edge of elytra not

curved forward immediately prior to meeting

pronotum. Pronotum with well-developed lateral

ridge/groove on each side, edge of elytron weakly

serrated towards tip, tip weakly pointed. Dark

testaceous, rear edge of head, lateral edges of

pronotum, small patches at side of elytron near

apex and appendages slightly lighter. Densely

rugose/punctate throughout. Pronotal process

narrow, keeled, bluntly pointed. Metacoxal lines

raised, relatively wide apart, strongly diverging in

anterior half to about 2.5 times their narrowest

width, raised lines not reaching metasternum.

Epipleuron evenly narrows until close to apex

where it abruptly ends.

Male. Protarsi three-segmented, anterior

portions greatly expanded and densely covered

with adhesive setae ventrally, third segment as

long as first two; claw shorter than third segment,

squat, broad but narrowing to sharp point (Figs 2,

3). Protibia relatively short, thick, with strong

tubercle on outside at base (Fig. 3). Apical ventrite

with short well-marked keel near apex. Median

lobe of aedeagus relatively narrow, parallel-sided

(Fig. 1), ending in pick-like beak (not visible

dorsally).

Female. Protarsi five-segmented, first three

segments weakly expanded anteriorly, densely

covered with adhesive setae ventrally, fourth

segment very small, apical segment thin,

cylindrical, shorter than third. Inner apical edge of

metatarsal segments extended, extension

accentuated by a number of strong spines.

Remarks

Tiporus lachlani differs from other Tiporus

with ridged/grooved pronotum by its large size

and strongly narrowed pronotal-elytral junction, as

well as in male characters.

Distribution

The Pilbara/Gascoyne region of Western

Australia.

REDESCRIPTIONS OF THE OTHER SPECIES IN THE

Genus, AFTER WATTS (1978)

(LisTED IN ALPHABETICAL ORDER)

Tiporus alastairi (Watts, 1978)

Description Fig. 23

Length 4.0 — 5.2 mm. Oval, convex. Dark

red-brown, underside lighter. Strongly and

densely rugose-punctate all over. Pronotum

with a distinct raised ridge parallel to and a

little distance inwards from each side, area

just inside ridge depressed, ridge and

depression strongest anteriorly, weak

posteriorly. Elytron weakly margined, weakly

serrated towards apex, usually sharply pointed

(rounded in male holotype). Edge of elytron

curves forward for short distance immediately

prior to meeting edge of pronotum.

Prothoracic process narrow, strongly convex,

roundly pointed at apex, only slightly

constricted between procoxae. Metacoxal

lines strongly raised, well separated, quite

strongly diverging in anterior two-thirds, not

quite reaching mesosternum.

Male. Protarsus three-segmented. Anterior sides

of segments of protarsus moderately expanded, a

little expanded on mesotarsus. Single claw on

94 C. H.S. WATTS

protarsus short, broad, weakly curved,

dorsoventrally flattened, anterior edge deeply

notched just beyond middle. Protibia with a large

sharp tooth on outside just anterior to middle.

Apical sternite strongly carinate for a short

distance in middle near apex. See Fig. 23 for

aedeagus.

Female. Protarsus five-segmented. Anterior

sides of pro-and mesotarsi less expanded than in

male. Apical abdominal sternite with a short sharp

ridge in middle near apex.

Remarks

Females are difficult to separate from those of

T. undecimmaculatus or T. giuliani but are larger

than most specimens of these species and the edge

of the elytron curves forward for a short distance

immediately before it meets the edge of the

pronotum whereas it is straight or almost so in T.

giuliani and T. undecimmaculatus. The same

applies to the males but in addition these have a

much shorter and broader proclaw and a much

narrower central lobe to the aedeagus than in these

species. The species is so far known only from the

north of Western Australia.

Distribution

Western Australia

2, Derby, SAMA; 3, 4 km W King Cascade,

15°38S 125°15'E, ANIC; 1, Kings Sound, ANIC.

Tiporus centralis (Watts, 1978)

Description Fig. 24

Length 3.2 — 3.9 mm. Oval, convex. Black;

sides of pronotum and appendages reddish, sides

and portions of middle of pronotum and base of

elytron with vague red patches. Strongly and

densely rugose-punctate throughout. Reticulate.

Pronotum not flanged. Elytron with a weak

shallow stria near the suture, edge curves forward

slightly immediately before meeting pronotum,

edge weakly to moderately expanded for a short

distance near, but not at, tip which is pointed.

Prothoracic process narrow, ridged in midline,

apex rounded, weakly constricted between

procoxae. Metacoxal lines strongly raised to

metasternum, quite strongly divergent in anterior

two-thirds.

Male. Protarsus three-segmented. Anterior sides

of segments of protarsus moderately expanded,

mesotarsus slightly expanded. Single claw on

protarsus long, sharply curved near base, apical two-

thirds straight. Protibia with a small tooth on outside

near base. Tip of apical abdominal sternite broadly

grooved in midline. See Fig. 24 for aedeagus

Female. Protarsi five-segmented; basal three

segments of roughly similar size, weakly

expanded anteriorly, ventral surface covered in

adhesive setae; fourth segment very small;

apical segment narrow, cylindrical, about same

length as third segment; claws weakly

developed.

Remarks

Tiporus centralis is very similar to T.

denticularis and T. georginae—see discussion

under those species.

Originally known only from the male holotype,

the species is now known to be reasonably

common in the Northern Territory and into

Western Australia. It seems to have a more inland

distribution than either 7. collaris or T.

undecimmaculatus, the other common Northern

Territory species.

Distribution

Northern Territory

2, 46 km SSW Borroloola, ANIC; 1, 48 km SW

by S Borroloola, ANIC; 3, 45 km W Borroloola,

SAMA; 1, Davenport Ranges, 40 km NE Murray

Downs Station, SAMA; 1, Elkedra Homestead,

NTM; 6, Gosse River, Murchison Ranges, NTM;

2, Kakadu Highway, 31 km from Pine Creek,

SAMA; 1, 35 mi N Larrimah, SAMA; 2, Moriarty

Creek, SAMA; 1, Skull Creek (Victoria River),

SAMA; 10, Victoria River, SAMA.

Western Australia

1, Stumpy Creek ‘Glenroy’, 16°55’S 125°34’E,

SAMA.

Tiporus collaris (Hope, 1842)

Description Figs 18, 21

Length 3.3 — 4.1 mm. Oval, convex. Black;

undersides and appendages a little lighter.

Strongly and densely rugose-punctate all over.

Reticulate. Pronotum widely flanged at sides,

lacking raised ridges between disc and flange

except very slightly anteriorly. Elytron weakly

margined, moderately serrate towards apex which

is rounded, base with vague longitudinal

impressions. Prothoracic process narrow, strongly

convex, roundly pointed at apex, not constricted

between procoxae. Metacoxal lines raised, slightly

to moderately diverging anteriorly.

Male. Protarsus three-segmented. Anterior sides

SPECIES OF TIPORUS 95

22 23 24

FIGURES 16-24. 16, dorsal and lateral views of median lobe of aedeagus of T. undecimmaculatus; 17, ditto T.

tambreyi; 18, ditto (lateral view) T. collaris (holotype); 19, ditto T. josepheni; 20, ditto T. giuliani; 21, ditto (dorsal

view) T. collaris (holotype); 22, ditto T. denticularis; 23, ditto T. alastairi; 24, ditto T. centralis. From Watts

(1978).

96 C.H.S. WATTS

of segments of protarsus moderately expanded,

those on mesotarsus a little expanded. Single claw

on protarsus thick and curved, strongly toothed on

underside near base. Protibia with a small tooth

on outside near base. Tip of apical abdominal

sternite moderately tuberculate in middle. See Figs

18, 21 for aedeagus.

Female. Protarsus five-segmented. Anterior

sides of segments of protarsus a little less

expanded than in male. Protarsus with two simple

claws. Apical abdominal sternite simple.

Remarks

Tiporus collaris is very similar to T. giuliani

and to T. undecimmaculatus. Males are readily

distinguished from both these species by the basal

position of the tubercle on the protibia rather than

the central or more apical position in 7. giuliani

and T. undecimmaculatus. In addition, T. collaris

is slightly more rounded, has slightly weaker

pronotal ridges/grooves and has rounded rather

than sharply pointed tips to the elytra.

The species occurs in small temporary streams

in stony country across the north of the Northern

Territory and Western Australia. Like T.

undecimmaculatus, with which it is often

collected, it appears to be more coastal than T.

centralis.

Distribution

Northern Territory

8, 5 km SE Mt Borradaile, SAMA; 4, 6 km SE

Mt Borradaile, SAMA; 1, 19 km E by S Mt

Borradaile, ANIC; 1, Canon Hill, Kakadu

National Park, SAMA; 1, Cooper Creek near Mt

Borradaile, SAMA; 17, 1 km W Gubara, Kakadu

National Park, SAMA; 1, 20 km SSW Jabiru,

SAMA; 1, Kambolgie Creek, SAMA; 4, 10 km E

by N Mt Cahill, ANIC; 1, Nawurlandja, Kakadu

National Park, SAMA; 1, 6 km SW by S Oenpelli,

SAMA.

Western Australia

1, Carson Escarpment, 14°49'S 126°49'E,

ANIC; 1, Drysdale River, 15°02'S 126°55’E,

ANIC; 2, Kimberley, SAMA; 4, Mitchell Plateau,

14°40'S 125°44'E, SAMA; 1, Upper Camp Creek,

14°49'S 125°S1'E, SAMA.

Tiporus denticulatus (Watts, 1978)

Description Fig. 22

Length 3.4 — 3.6 mm. Oval, convex. Dark red-

brown; sides, base and portions of anterior of

pronotum, some small patches on elytron and

appendages yellowish. Strongly and densely

rugose-punctate throughout. Pronotum not

flanged, with a weak ridge with deep groove on

inside, parallel to and some distance inwards from

sides. Apex of elytron pointed. Epipleuron with a

small triangular expansion just before apex.

Prothoracic process relatively broad, ridged in

midline, apex rounded, moderately constricted

between procoxae. Metacoxal lines straight in

posterior half, rapidly diverging in anterior half,

not reaching metasternum.

Male. Protarsus three-segmented. Anterior sides

of segments of protarsus moderately expanded.

Mesotarsus a little expanded. Single claw on

protarsus short, flat, spindle-shaped, rapidly

narrowing near apex on inside. Protibia with small

tooth on outside near base. Tip of apical sternite

with small weak ridge in midline. See Fig. 22 for

aedeagus.

Female. Protarsus five-segmented, basal

segments somewhat expanded on inside.

Remarks

Although the species has a weak ridge/groove

on the side of the pronotum, the testaceous margin

of the pronotum, smooth elytron edge, lateral

expansion of the epipleuron near the tip and the

basal position of the tubercle on the male protibia,

ally it to the 7. josepheni group of species.

Tiporus denticularis is very similar to T. centralis

and females are difficult to separate, but they

differ from this species by the greater

development of the pronotal ridge/groove and by

the extreme front edge of the elytron immediately

before it meets the edge of the pronotum being

slightly curved in 7. centralis but straight in T.

denticularis.

Originally only known from the Cairns-

Cooktown region of North Queensland the

additional specimens extend its known range into

the Northern Territory.

Distribution

Queensland

1, 34 km NW Chillagoe, SAMA; 8, 70 km SW

Greenvale, SAMA; 7, Lakeland Downs, SAMA;

3, Laura, SAMA; 4, Mary Creek, 16°33'S 12°5'E,

ANIC; 2, 11 km WSW Petford, QDPIM; 2, Walsh

River near Chillagoe, QDPIM; 1, Windsor

Tableland, QDPIM.

Northern Territory

1, Bullita Outstation, 16°07’S 130°25’E, NTM;

1, Victoria River, SAMA.

SPECIES OF TIPORUS 97

Tiporus giuliani (Watts, 1978)

Description Fig. 20

Length 3.6 — 4.2 mm. Oval, convex. Black;

appendages, some vague reddish areas at base of

pronotum and base and sides of elytron, lighter.

Moderately strongly rugose-punctate throughout.

Pronotum with a raised ridge parallel to and a

little distant inwards from each side, area just

inside ridge depressed, ridge and depression

strongest anteriorly, weak posteriorly. Elytron

weakly serrated towards apex which is pointed.

Prothoracic process narrow, strongly convex,

bluntly pointed at apex, only slightly constricted

between procoxae. Metacoxal lines relatively

close, strongly diverging in central third, weakly

in anterior and posterior third, reaching

metasternum.

Male. Protarsus three-segmented. Anterior sides

of segments of protarsus moderately expanded,

weakly so on mesotarsus. Single claw on protarsus

short, narrow, with slight notch on underside in

middle. Protibia moderately strongly toothed on

outside in middle. Apical sternite strongly carinate

for a short distance in middle near apex. See Fig.

20 for aedeagus.

Female. Protarsus five-segmented. Pro- and

mesotarsi less expanded than in male. Apical

abdominal sternite very weakly carinate in middle

near apex.

Remarks

Tiporus giuliani is close to T.

undecimmaculatus and T. collaris. It can be

separated from T. collaris by its stronger pronotal

ridges/grooves, sharply pointed elytral tips,

stronger colour pattern and the central position of

the spine on the male protibia.

The only clear separation from T.

undecimmaculatus is the shape of the tip of

the median lobe of the aedeagus which is

broad with three small spines at the tip in T.

undecimmaculatus (Fig. 16) and narrow and

bluntly pointed in 7. giuliani (Fig. 20). In the

Northern Territory and Western Australia the

more apical position of the spine on the male

protibia and the more pronounced dorsal

colour pattern will separate it from T.

undecimmaculatus. However specimens of T.

undecimmaculatus from Queensland resemble

T. giuliani both in the central position of the

spine on the male protibia and in the well

developed dorsal colour pattern.

A less common species than some and possibly

more inland than coastal in distribution.

Distribution

Northern Territory

4, Kakadu Highway, 31 km from Pine Creek,

SAMA.

Western Australia

1, Beverley Springs, WAM; 2, Duncan

Highway, WAM; 2, near Dampier Downs, WAM;

4, Logues Springs, 102 km SE by E Broome,

ANIC; 1, 163 km SE by S Broome, ANIC.

Tiporus josepheni (Watts, 1978).

Description Figs 8, 19

Length 3.5 — 4.1 mm. Oval, convex. Black,

sides of pronotum narrowly vaguely reddish,

appendages dark red-brown. Ventral surface

moderately and densely rugose-punctate,

dorsal surface closely and moderately

punctate. Pronotum not laterally flanged or

ridged. Tip of elytron weakly pointed.

Prothoracic process narrow, strongly convex,

only slightly constricted between procoxae,

apex rounded, midline virtually impunctate.

Metacoxal lines well separated, moderately

diverging posteriorly and in middle,

subparallel in anterior quarter, reaching

metasternum.

Male. Protarsus three-segmented. Anterior

sides of segments of protarsus strongly

expanded. Mesotarsus a little expanded. Single

claw on protarsus short, flat, narrowing abruptly

on inside near apex, base with a large rounded

expansion beneath. Protibia with small tooth on

outside in middle. Tip of apical sternite

depressed slightly in midline. See Fig. 19 for

aedeagus.

Female. Protarsus five-segmented, robust, basal

segments somewhat expanded on inside. Elytron

with very well developed lateral subapical spine

(Fig. 8).

Remarks

Originally described from Beverley Springs and

Wittenoom Gorge in the Kimberley and Pilbarra

regions of Western Australia respectively, the

species is now known to be much more

widespread, occurring right across coastal

Northern Australia. It is particularly common in

North Queensland. It seems to favour larger and

more permanent rivers than other Tiporus.

Unfortunately I mis-associated the sexes in my

original description. (The female paratype in

ANIC belongs to T. tambreyi.)

98 C. H.S. WATTS

Distribution

Northern Territory

2, Cooper Creek near Mt Borradaile, SAMA; 7,

Magela Creek, SAMA; 2, Magela Creek, 12 km E

Jabiru, SAMA; 1, Nourlangie Creek, 20 km SSW

Jabiru, SAMA.

Queensland

1, Barron River, QDPIM; 52, 8 km N

Bluewater, SAMA; 1, 25 km N Coen, SAMA; 2,

2 km NW Daintree, QDPIM; 2, Emu Creek 5 km

W Petford, QDPIM; 8, Eubenangee Swamp near

Babinda, SAMA; 2, Helenvale, SAMA; 18,

Lakeland Downs, SAMA; 1, Laura, SAMA; 2,

Mazlin Creek Atherton area, QDPIM; 1, 2 km S

Mt Molloy, SAMA; 2, 20 km S Townsville,

SAMA; 1, 37 km S Townsville, SAMA; 7, Walsh

River 34 km NW Chillagoe, SAMA.

Western Australia

1, Beverley Springs, WAM; 1, Stumpy Creek

‘Glenroy’ 16°55'S 125°34'E, SAMA.

Tiporus tambreyi (Watts, 1978)

Description Fig. 17

Length 3.5 — 4.4 mm. Oblong oval, strongly

convex, pronotum a little constricted in anterior

quarter. Black, extreme anterior of head, sides

of pronotum narrowly, a basal spot and extreme

lateral markings on elytron and appendages

reddish. Moderately strongly and very densely

punctate all over, punctures on pronotum and

elytron with short setae. Reticulation strongest

anteriorly. Pronotum and elytron weakly

margined, margins of elytron weakly serrated

towards apex, extreme apex not margined.

Prothoracic process lanceolate, quite strongly

expanded behind procoxae, narrow between

procoxae, weakly flanged, strongly carinate.

Metacoxal lines strongly raised, rapidly

diverging in anterior half, subparallel

posteriorly.

Male. Anterior sides of segments of protarsus

moderately expanded, posterior sides

unexpanded. Basal segment of mesotarsus

weakly expanded. Single claw of protarsus

greatly expanded in basal half, flattened

dorsoventrally. Metafemur robust, curved.

Metatibia angularly thickened on inside near

apex. Protibia with a small tooth on outside close

to base. Last abdominal sternite widely and

weakly grooved in midline with a small knob in

middle of groove at extreme apex, edges of

sternite at apex raised for some distance either

side of groove. See Fig.17 for aedeagus.

Female. Smaller, more densely punctate.

Protarsus asymmetrically expanded as in male but

not to same extent. With small triangular

extension at tip of elytron, diverging from each

other when elytra closed (Fig. 15).

Remarks

A relatively large species known only from the

Pilbara region of Western Australia, readily

distinguished from all other Tiporus by the

expanded male metatibia and the triangularly

extended elytral tips in the female. The sides of

the pronotum straighten slightly in front giving

the pronotum a slightly sinuate outline.

Distribution

Western Australia

1, 17 km N by E Cane River Homestead, ANIC;

15, Gregory Gorge, Fortescue River, SAMA; 16,

Millstream, ANIC; 1, Millstream, Palm Pool area,

WAM; 9, 1 km N Millstream, ANIC; 4, 3 km NW

by W Millstream, ANIC; 1, Tambrey, WAM; 1,

Wittenoom Gorge, ANIC.

Tiporus undecimmaculatus (Clark, 1862)

Description Fig. 16

Length 3.4 — 4.6 mm. Oval, convex. Black or

dark red-brown; underside and appendages a little

lighter, base of pronotum and sides of elytron with

vague red patches in many. Strongly and densely

rugose-punctate all over. Reticulate. Pronotum

with a distinct raised ridge parallel to and a little

distance from each side, area just inside ridge

depressed, ridge and depression strongest

anteriorly, weak posteriorly. Elytron weakly

margined, strongly convex, rounded towards apex

which is pointed. Prothoracic process narrow,

strongly convex, roundly pointed at apex, little if

at all constricted between procoxae. Metacoxal

lines raised, slightly to moderately diverging

anteriorly.

Male. Anterior sides of segments of protarsus

moderately expanded, those on mesotarsus a little

expanded. Single claw on protarsus weakly

curved, slightly thickened and with a small tooth

on underside near base and another just before the

middle. Protibia with a small sharp tooth on

outside in apical half. Tip of apical abdominal

sternite weakly to strongly tuberculate in middle.

See Fig. 16 for aedeagus.

Female. Anterior sides of segments of protarsus

SPECIES OF TIPORUS 99

a little less expanded than in the male. Protarsus

with two simple claws. Apical abdominal sternite

simple or slightly tuberculate at tip.

Remarks

Tiporus undecimmaculatus is a widespread and

variable species very similar to T. collaris and T.

giuliani—see notes under those species. Two

forms are distinguishable. The nominal form,

from the Northern Territory and Western

Australia, is usually dark with the dorsal colour

pattern subdued or absent and has the spine on the

male protibia towards the apex. The other form,

from north Queensland, has a well-developed

dorsal colour pattern, the spine on the male

protibia close to the centre of the protibia, the

central lobe of the aedeagus a bit broader and also

tends to be larger.

Tiporus undecimmaculatus is common in small

temporary streams in stony country in coastal

areas across Northern Australia in similar habitats

to T. collaris but is usually more abundant than

that species. A more coastal species than 7.

centralis.

Distribution

Western Australia

1, Bigge Island, 14°29'S 125°10'E. SAMA; 7,

Careening Bay, 15°06S 125°00E, SAMA; 9,

Carson Escarpment, 14°49'S 126°49'E, ANIC; 1,

Cave Spring, 15°32'S 128°50'E, WAM, ANIC; 1,

Duncan Highway, WAM; 3, 14 km S by E

Kalumburu Mission, ANIC; 2, 4 km W King

Cascade, 15°38'S 125°15'E, ANIC; 3, 4 km S by

W Mining Camp, Mitchell Plateau, 14°52'S

125°50'E, ANIC; 7, Mitchell Plateau, 14°40'S

12°44'E, SAMA.

Northern Territory

3, 6 km SE Mt Borradaile, SAMA; 6, 19 km E

by S Mt Borradaile, ANIC, NTM; 137, 5 km SE

Mt Borradaile, SAMA; 2, Bukkita outstation,

16°07'S 130°25'E, NTM; 1, 7 km NW by N

Cahills Crossing, Kakadu National Park, SAMA;

1, Canon Hill, Kakadu National Park, SAMA; 1,

4 mi S Coolibah, WAM; 3, Gosse River,

Murchison Ranges, NTM; 13, 1 km W Gubara,

Kakadu National Park, SAMA; 2, Kambolgie

Creek, SAMA; 2, Magela Creek, SAMA; 2, 19

km E by N Mt Cahill, ANIC; 5, 10 km E by N Mt

Cahill, ANIC; 1, 15 km E by N Mt Cahill, ANIC;

15, 40 km NE Murray Downs Station, SAMA; 4,

6 km SW by S Oenpelli, ANIC, SAMA; 5,

Nawurlandja, Kakadu National Park, SAMA; 1,

Nourlangie Creek, 20 km S Jabiru SAMA.

Queensland

2, Cairns, SAMA; 1, Charters Towers, SAMA;

1, 25 km N Coen, SAMA; 1, Helenvale, SAMA;

15, 14 km W Herberton, SAMA; 21, Lakeland

Downs, SAMA; 4, Laura, SAMA; 1, 12 km N

Laura, SAMA; 1, MclIlwraith Ranges Weather

Station, SAMA; 1, 17 km up Mt Lewis Road,

QDPIM; 6, Mt Spec, ANIC; 8, 10 mi W Paluma,

SAMA; 1, 20 km W Petford, SAMA; 8, 15 km W

Petford, QDPIM.

ACKNOWLEDGMENTS

The curators of the collections listed earlier are

thanked for allowing me to examine specimens in their

care. The illustrations were most ably done by Mr R.

Gutteridge and Ms Debbie Churches helped finalise the

manuscript. All are thanked for their support and help.

REFERENCES

WATTS, C. H. S. 1978. A revision of the Australian

Dytiscidae (Coleoptera). Australian Journal of

Zoology, Supplementary series No 57: 1-166.

WATTS, C. H. S. 1985. A faunal assessment of

Australian Hydradephaga. Proceedings of the

Academy of Natural Sciences Philadelphia 137: 22—

28.

WATTS, C. H. S. 1997. A new genus and species of

Australian Dytiscidae (Coleoptera). Records of the

South Australian Museum 29: 121-123

THE PARACYMUS OF AUSTRALIA (COLEOPTERA, HYDROPHILIDAE).

BY ELIO GENTILI

Summary

GENTILI, E. (2000). The described Australian Paracymus were hitherto placed in six species. Of

these three are to be considered synonyms of P. pygmaeus (MacLeay, 1871): nitidiusculus (Broun,

1880), metallescens Fauvel, 1883, and desolatus Woolridge, 1976 (new synonymy). ‘To the

remaining valid species P. pygmaeus (MacLeay, 1871), spenceri Blackburn, 1896, and gigas

Gentili, 1996 another six new species are now added: P. cariceti, wattsi, opasus, australiae, weiri,

and ovum. Lectotypes are designated for P. pygmaeus, nitidiusculus, and metallescens. Each valid

species is described, discussed, mapped (with a detailed list of localities), figured (aedeagi, outlines)

and keyed.

THE PARACYMUS OF AUSTRALIA (COLEOPTERA, HYDROPHILIDAE)

ELIO GENTILI

GENTILI, E. 2000. The Paracymus of Australia (Coleoptera, Hydrophilidae). Records of the

South Australian Museum 33(2): 101-122.

The described Australian Paracymus were hitherto placed in six species. Of these three are

to be considered synonyms of P. pygmaeus (MacLeay, 1871): nitidiusculus (Broun, 1880),

metallescens Fauvel, 1883, and desolatus Wooldridge, 1976 (new synonymy). To the remaining

valid species P. pygmaeus (MacLeay, 1871), spenceri Blackburn, 1896, and gigas Gentili,

1996 another six new species are now added: P. cariceti, wattsi, opacus, australiae, weiri, and

ovum. Lectotypes are designated for P. pygmaeus, nitidiusculus, and metallescens. Each valid

species is described, discussed, mapped (with a detailed list of localities), figured (aedeagi,

outlines) and keyed.

E. Gentili, via San Gottardo 37, Rasa di Varese, 21030 Italy. Manuscript received 3 April

2000.

INTRODUCTION

This work is intended to continue on with the

project of A. F. Newton and M. K. Thayer to

improve our knowledge of the Staphyliniformia

of Australia. After the studies on the hydrophilid

genera Notohydrus (Gentili 1992) and

Paranacaena (Gentili 1993) the genus Paracymus

Thomson, 1867 is now considered.

Hansen (1991) placed Paracymus in the tribe

Anacaenini, giving a description of the genus. The

features basically characterising Paracymus

among Anacaenini appear to be the prosternum

longitudinally carinate and the presence of a

sutural stria on the elytra (see also d’Orchymont

1942).

Gentili (1993) dealt with the nine Australian

taxa treated in the literature under the genus

Paracymus, placing four of them (indi Blackburn,

1888, sublineatus Blackburn, 1888, horni

Blackburn, 1896, and eremita Blackburn, 1896)

in Paranacaena Blackburn, 1888 and one

(nigerrimus Blackburn, 1891) in Chaetarthria

Stephens, 1835, leaving in Paracymus the

remaining four (pygmaeus McLeay, 1871,

nitidiusculus Broun, 1880, spenceri Blackburn,

1896, and desolatus Wooldridge, 1976). Four

other taxa might be added to the list: Paracymus

phalacroides (Wollaston, 1867); Wooldridge

1978 wrote: ‘two females from Australia seem to

belong here. If these...specimens are really P.

phalacroides, they probably are the result of

accidental introduction, because they do not seem

to be established in...the country’; Paracymus

metallescens Fauvel, 1883 from New Caledonia;

Anacaena tepida Winterbourn, 1970 treated as

Paracymus in a manuscript of R. Ordish on New

Zealand Hydrophilidae, and Paracymus gigas

Gentili, 1996.

MATERIAL AND METHODS

More than 6 000 specimens were studied from

the Institutes and Museums listed below under

‘Acronyms’, nearly 5 000 of them belonging to the

Australian National Insect Collection. Specimens

were examined with a Beck Kassel CBS

stereoscopic microscope; the figured aedeagi were

mounted in di-methyl-hydanthoin-formaldehyde

(DMHF) on transparent plastic card, studied and

drawn with a GALILEO LG transmitted light

microscope equipped with a projection device.

Acronyms

AMS -— Australian Museum, Sydney.

ANIC — Australian National Insect Collection,

CSIRO, Canberra.

CASF — California Academy of Sciences, San

Francisco.

FMNH -— Field Museum Natural History, Chicago.

ISNB_ -— Institut Royal des Sciences Naturelles de

Belgique, Bruxelles.

MSNV — Museo Civico di Storia Naturale,

Verona.

NHML — Natural History Museum, London.

NMW — Naturhistorisches Museum, Wien.

SAMA — South Australian Museum, Adelaide.

USNM —- United States National Museum,

Washington.

102

DESCRIPTIONS

Each valid Australian species is discussed

below according to the following scheme: (1)

References; the dates of description and of the

quoted papers conform to Hansen (1999). (2)

Type material; I searched for the available types

of the described taxa (valid species and

synonyms), designating lectotypes and

paralectotypes where necessary. (3) Description;

many characters are considered: measures, outline,

upperside, underside, palps, antennae, legs and

aedeagus. (4) Discussion; the species are

compared with other similar species or with the

proposed synonyms. (5) Material examined; the

States of the Australian continent are listed

alphabetically; the localities within each State are

also listed alphabetically. (6) Biology; the

biological notes are derived from the label data.

1. Paracymus pygmaeus (MacLeay, 1871)

Cyclonotum pygmaeum MacLeay, 1871: 133;

White in Masters, 1871: 5.

Coelostoma pygmaeum (MacLeay): Zaitzev 1908,

404.

Paracymus pygmaeus (MacLeay): Blackburn

1888: 820; 1894: 203; Knisch 1924: 167;

d’Orchymont 1937: 154, 157; McKeown 1948:

99; Wooldridge 1976: 459-461; Matthews 1982:

55; Hansen 1999: 113.

Hydrobius nitidiusculus Broun, 1880: 78.

Paracymus nitidiusculus (Broun): Sharp 1884:

467; Blackburn 1888: 820-821.

Paracymus metallescens Fauvel, 1883: 352;

Knisch 1924: 166; d’Orchymont 1926: 376 (?

synonym of pygmaeus).

Paracymus desolatus Wooldridge, 1976: 458-

459; Hansen 1999: 110. New synonymy.

Types

Lectotype male (1.7 x 1.3 mm): Queensland:

Gayndah, W. McLeay, 1871, ANIC. A single pin

bears the following cards or labels: 1. insect and

its abdomen; 2. aedeagus and spiculum gastrale in

DMHF; 3. Round amaranth label; 4. Gayndah; 5.

Cyclonotum pygmaeum, Macl. Gayndah (by

hand); 6. On permanent loan from Macleay

Museum University of Sydney; 7. Syntype (red

label); 8. Lectotype male designated by E. Gentili

1991 (red label); 9. Lectotype male Cyclonotum

pygmaeus Macl. Det. R. G. Ordish.

E. GENTILI

Paralectotypes: Queensland: Gayndah, W.

McLeay 1871 (4), ANIC, AMS. The previously

mentioned manuscript of R. G. Ordish says: ‘For

access to syntypes I am indebted to Mr T. A.

Weir, CSIRO Canberra and to Dr G. A. Holloway

of the Australian Museum, Sydney. I am advised

that the material collected from Gayndah was

shared between the Macleay Museum and the

Australian Museum and that a specimen in the

latter has subsequently been labelled as the

Holotype, seemingly on the basis of an original

name label (McKeown 1948). As the writer points

out, ‘types in the Macleay Museum collection are

not specially indicated as such’. Validity aside,

there are two obstacles to this recognition of a

holotype. Firstly, all three mounts bear original

determination labels in Macleay’s hand, and

secondly the mount bearing the holotype label (K

19573) has two specimens on it. There are five

syntypes from which a subsequent author could

choose a lectotype and I have done this by

selecting the best preserved male’. The manuscript

of Ordish was not published, but his work is so

reliable that I follow his statements, designating

as lectotype the specimen chosen by him, seen

also by me by courtesy of T. A. Weir.

Synonyms

(1) Hydrobius nitidiusculus Broun, 1880.

Lectotype female: New Zealand, Tairua, Broun

1880, NHML. A single pin bears the following

cards and labels: 1. Insect; 2. 144; 3. Type

(Circular label red-bordered); 4. Tairua; 5. New

Zealand, Broun Coll., Brit. Mus. 1922-182; 6.

Paracymus nitidiusculus; 7. female Lectotypus,

Hydrobius nitidiusculus Broun, E. Gentili 1992

(red label). I did not see other types, but Broun

certainly described the species based on more

specimens, as in the description he states: ‘I

obtained the specimens now before me at Tairua

and Whangarei Heads’. Blackburn (1888) wrote:

‘Appears to be common in South Australia... I

have taken it in Western Victoria also’. But in

1894 he wrote: ‘Paracymus (Cyclonotum)

pygmaeus, Macl. I have recently received

examples (compared with the type) of this insect

from Mr. Lea. They seem certainly identical with

Paracymus (Hydrobius) nitidiusculus, Brown.

Macleay’s is the older name’. After examination

of the type, I agree with Blackburn.

(2) Paracymus metallescens Fauvel, 1883.

Lectotype male (2.2 x 1.2 mm): New Caledonia,

Tonghoué, Savés, ISNB. A single pin bears the

following cards and labels: 1. Insect and its

abdomen; 2. Aedeagus in DMHF; 3. Coll.

PARACYMUS OF AUSTRALIA 103

R.I.Sc.N.B., Nouvelle Calédonie, Tonghoué 9me,

Rec. Savés, ex Coll. Fauvel (pink card); 4. Coll. et

det. A. Fauvel, Paracymus metallescens Fvl.,

R.LSc.N.B. 17.479; 5. Syntype; 6. Lectotypus

male, Paracymus metallescens Fauv., E. Gentili

1991. Three paralectotypes are also present in the

ISNB, from the following localities: 1) Marais de

Vanse Vata, juilliet 8me, Nouméa, Rec. Savés; 2)

Kanala, Rec. Coste; 3) Koné, Rec. Atkinson.

Fauvel (1883) wrote: ‘Aussi en Australie’; and

d’Orchymont 1926 stated: ‘Paracymus

metallescens Fauvel, 1883 from New Caledonia is

perhaps the same as P. pygmaeus W. S. MacLeay,

1871, from Australia, but no material from the

original country could be seen’. Comparison with

the Australian insects, including the aedeagus

(Fig. 3), leads me to synonymise the two taxa.

(3) Paracymus desolatus Wooldridge, 1976.

Holotype male (2.1 x 1.2 mm): Western

Australia, Winjana Gorge, Ross & Cavagnaro

17.10.1962, CASF. A single pin bears the

following cards and labels: 1. Insect with

semiextracted aedeagus; 2. W. Australia: Winjana

Gorge 100 m X.17.62; 3. Collectors E. S. Ross D.

Cavagnaro; 4. male; 5. HOLOTYPE Paracymus

desolatus Wooldridge (red label); 6. California

Academy of Sciences Type No. 12007. Paratypes:

the description quotes 143 males 149 females

(including the allotype) from Northern Territory,

Queensland, Western Australia, housed in CASF,

NHML, SAMA, USNM. I here synonymise P.

desolatus with P. pygmaeus after a long effort to

isolate the true characteristics of desolatus. These

might be: 1) pronotal punctation fine and widely

separate, elytral punctation closer and more

impressed; in pygmaeus pronotal and elytral

FIGURES 1-17. Aedeagi. 1, Paracymus pygmaeus (MacLeay, 1871), lectotype, dorsal view; 2, Idem, ventral view;

3, Paracymus metallescens Fauvel, 1883, lectotype, ventral view; 4, Paracymus pygmaeus (Kosciusko NP) showing

the preapical collar and the connected membrane; 5, Paracymus pygmaeus (Warburton): as in Fig. 4; 6, Paracymus

spenceri Blackburn, 1896, lectotype, ventral view; 7, Idem, dorsal view; 8, Idem, spiculum gastrale; 9, Paracymus

cariceti n. sp., holotype, ventral view; 10, Paracymus australiae n. sp., holotype, ventral view; 11, Paracymus

gigas Gentili, 1996, holotype, ventral view; 12, Idem, dorsal view; 13, Paracymus weiri n. sp., holotype, ventral

view; 14, Idem, dorsal view; 15; Paracymus wattsi n. sp., holotype, ventral view; 16, Paracymus opacus nN. sp.,

holotype, dorsal view; 17, Paracymus ovum n. sp., holotype, ventral view.

104

punctation nearly equal. This character is

conspicuous at 100 x but in a great number of

sympatric specimens presents a high variability,

from insects nearly impunctate to coarsely

punctate on the whole dorsal surface; 2) the last

segment of male protarsi less widened and hooked

than in pygmaeus. I examined the holotype but

was not able to distinguish the pattern of its

protarsi from that of pygmaeus; 3) the

mesofemora pubescent only on basal triangle,

covering only 1/3 of anterior edge; in pygmaeus

more pubescent, covering nearly 1/2 of anterior

edge. In many cases this distinction is in my

opinion impossible to see; 4) penis rapidly

narrowing and becoming parallel-sided for about

one-third its length; narrowly triangular in

pygmaeus. Really the penis is narrowly triangular,

due to a membranous expansion, from the base to

a preapical collar in both forms. Sometimes the

membranous expansion is scarcely visible or

contracted, possibly due to effects of preservation.

E. GENTILI

(4) Anacaena tepida Winterbourn, 1970 is

treated as a synonym of P. pygmaeus by R. Ordish

in the above mentioned manuscript. But no author

records it from Australia. Therefore I think it is

not necessary to treat this species here.

Description

Length 1.6-3.0 mm; width 0.9-1.6 mm.

Elongate oval, slightly convex (Figs 18, 19).

Head black, evenly punctured, surface shining

between punctures, occipital region alutaceous;

transverse sutures distinct, coronal suture

obsolete. Pronotum dark, a little paler at sides,

black to reddish-brown, with olive-green

reflections in some specimens; punctation finer

than on head, obvious at sides, faint or absent on

disc. Elytra uniform black to reddish-brown, with

green reflections in some specimens, shining

between punctures, punctures shallow like those

of pronotum or shallower, but evenly distributed,

with sparse setae laterally; parasutural furrow

FIGURES 18-21. 18, Outline of Paracymus pygmaeus (MacLeay, 1871); 19, Profile of Paracymus pygmaeus; 20,

Outline of Paracymus wattsi n. sp.; 21, Profile of Paracymus opacus n. sp.

PARACYMUS OF AUSTRALIA 105

extended from apex nearly to anterior third of

elytra. Under side dark; prosternum broadly

keeled in middle and with a projection on

anterior border; mesosternum with transverse

ridge, a median keel anterior and posterior to it

extending to both borders. First visible (third)

abdominal ventrite with a median keel

throughout its length. Palps pale yellow except

for darkened apices; antennae pale yellow, eight-

segmented. Profemur ventrally pubescent on

proximal third; male protarsus with last segment

shorter and broader than in female with two short,

ventral, blunt spines; tarsal segments 14 (male

protarsus) with a single ventral spine; protarsal

claws of male less evenly curved than in female

but equal in length. Ventral pubescence of

mesofemur confined to proximal half or third and

not reaching posterior border. Metafemur glabrous

with weak longitudinal scratch-like impressions.

Aedeagus (Figs 1-5) with tegmen pointed at base,

parameres blunt at apex and scarcely longer than

tegmen. Penis blunt at apex, with a collar at

beginning of the apical third; from collar a

membranous expansion reaches base of parameres

describing a triangle; sometimes this expansion is

scarcely conspicuous, possibly due to the methods

of preservation (e.g. ethyl alcohol). Gonopore

subapical and ventral.

Discussion

This is the most widespread Paracymus in

Australia. Some of its characters, such as the

dorsal punctation and the extension of the hairy

surface on the mesofemora, show great variability.

It is easy to separate from other species by the

shape of its aedeagus: tegmen pointed at base

(difference from weiri and ovum), scarcely shorter

than parameres (difference from opacus, gigas,

weiri, australiae, cariceti),; penis simply pointed

(difference from cariceti, opacus, gigas), with a

preapical collar (as in spenceri, but the collar of

spenceri lacks the membranous expansion). Other

distinctive characters are the last protarsal

segment of males, which is broader than in any

other Australian Paracymus, and the scarce

pubescence of the profemora.

Material examined (Fig. 22)

Australian Capital Territory: Black Mt., I. F.

B. Common 12.11.1964 (1), 16.11.64 (4), 22-

26.12.64 (1), 6.1.65 (1), 14.11.65 (1), 14.12.65

(6), 20.12.65 (6), 29.12.65 (9), 6.1.66 (2), 10.1.66

(2), 11.1.66 (2), 18.1.66 (1), 20.1.66 (1), 17.3.66

(2), 21.3.66 (2), 22.3.66 (1), light trap, ANIC;

Black Mt., M. S. Upton 15.10.1965 (7), 8.12.65

(1); 15:12:65" (3), 16:12:65 (1);. 22:12:66 (3),

24.1.67 (6), 16.10.67 (2), 30.11.67 (1), 12.12.67

(1), 10.1.68 (2),14.1.68 (1), 17.1.68 (2), 29.1.68

(1), 1.2.68 (2), 2.2.68 (1), 13.2.68 (2), 28.2.68 (1),

4.3.68 (1), 25.3.68 (2), 26.3.68 (5), 27.3.68 (2),

28.3.68 (3), 29.3.68 (3), 12-16.4.68 (1), 22.4.68

(1), light trap, ANIC; Black Mt., Z. Liepa

20.5.1966, ex rotting wood in creek (1); Black

Mt., E. B. Britton 25.11.1964 (1) at light, ANIC;

Black Mtn Reserve, S. Misko 4.12.1970 (2) light

trap, ANIC; Black Mt. 600 m, 35°16'S 149°06'E,

Weir Dressler & Lawrence 12.1986 (1) flight

intercept window/trough trap, ANIC; Black Mt.,

Bywater & Clayton 23.1.1967 (1) from nest of

Buff-tailed Thornbill, ANIC; Canberra, H. & A.

Howden 2-3.XII.1986 (1) black light, ANIC;

Cotter River, E. J. Pook 20.12.1965 (5) ANIC;

Lake Burley Griffin, Z. Liepa 19.11.1965 (1)

ANIC; Narrabundah Orchard, 21.11.1966 (1)

ANIC; Paddy’s River 1 mi S of Cotter Dam, S.

Misko 17.4.1969 (2) ANIC; Piccadilly Circus

1240 m, 35°22'S 148°48'E, Lawrence Weir &

Johnson 5.1984, 9.84, 12.84 (6) flight intercept

window/trough trap ANIC; Piccadilly Circus, C.

Reid 10.12.1984 (1) powerline clearing, ANIC;

Piccadilly Circus 6 km NE, Wombat Ck, 35°19'S

148°47'E, 750 m, Weir Lawrence & Johnson 8.

1985 (1) flight intercept window/trough trap,

ANIC; Snowy Flat Ck, Mt Gingera 0.5 km NE,

35°35'S 148°47'E, A. A. Calder 28.6.1988, ANIC.

New South Wales: Albury, E. F. Riek 26.1.1963

(2) ANIC; Araluen, Apple Tree Ck, W. & S. Allen

6.12.1975 (1) ANIC; Berry, C. Watts 1.1967 (2)

SAMA; Blue Mountains, Foulcon Bridge, 500 m,

G. Wewalka 15.1.1993 (8) NMW, MSNV; Blue

Mts, H .J. Cox (2) ANIC; Braidwood 15 km NW,

Shoalhaven R., C. Reid 19.12.1984 (1) on Acacia

spp., ANIC; Broken Head Nat. Reserve, Byron

Bay 8 km S, Common & Edwards 23.11.1976 (2)

ANIC; Canberra Coast Rd, Manar Ck, Britton &

Misko 18.5.1967 (2) ANIC; Canberra Coast Rd,

Cabbage Tree Ck, 7.7.1965 (1) ANIC; Casino 4

mi W, E. B. Britton at light (1) ANIC; Chichester

St. Forest, Allyn R. Park, J. T. Doyen 8.XI.1982

(2), T. Weir 10-11.1981 ANIC; Chiswick nr

Armidale, B. Clydesdale 12.1965 (5), 2.66 (2),

6.12.67 (1) ANIC; Clarence R., Brisbane, Coates,

Griffith (6) SAMA; Cooma, Duboulay (3) ANIC;

Coonabarabran 9 km W, 533 m, 31°17S 149°11'E,

Common & Edwards 2.12.1974 (1) ANIC

Coonabarabran 9 km NNE, Newe 11 Hwy, E.

Britton 24.10.1980 Pilliga scrub (2) ANIC;

Coonabarabran 14 km W, nr Timor Rock, J.

Doyen 4.11.1982 (3) ANIC; Culcairn, E. W.

Ferguson (4) ANIC; Darling R., Bourke 20 mi

106 E. GENTILI

FIGURE 22. Distribution of the studied specimens of Paracymus pygmaeus (MacLeay, 1871).

SSW, on bank, G. F. Gross 26.12.1973 at light (1)

SAMA; Delagate 14 km W, C. Watts 4.11.1997

(2) SAMA; Deniliquin, V. R. Squires 10.2.1966

light trap (1) ANIC; Dungong 35 km N,

Chichester State Forest, Telegherry Forest Park,

300 m, Pollock & Reichert 18.12.1990 UV light

along river (14) NMW, MSNV; Fowlers Gap Res.

Stn, 31°05'S 141°42'E, Cardale & Naumann

29.11—2.12.1981 (18) at light, ANIC; Gilgandra,

C. Watts 19.11.1992 (5) SAMA; Griffith, 34°17'S

146°03'E, R. Kohout 10.5.1972 (2) ANIC; Hay 23

mi. E, Britton & Misko 23.10.1967 at light (10)

ANIC; Hay 8 km W, 34°30'S 144°S1'E, Britton,

Misko & Pullen 14.12.1970 (10) river bank, at

light, ANIC; Hay 37 km E, E. G. Matthews

10.3.1971 (5) SAMA; Jenolan Caves, vicinity, J.

C. Wiburd (4) SAMA; Jindabyne 13 km NE, Kara

Ck 940 m, T. A. Weir 16.3.1979 (8) ANIC,

MSNV; Jindabyne 12 km NNE, The Lake Ck

1160 m, T. A. Weir 16.3.1979 (114+10 not

prepared) ANIC, MSNV; Jindabyne 6 km NW,

Thredbo R. 500 m, T. A. Weir 15.3.1969 (1)

ANIC; Jindabyne 10 km W, Wollindibby Ck 1060

m, T. A. Weir 15.3.1969 (2) ANIC; Khancoban,

below Khancoban Dam 300 m, 36°13'S 148°06'E,

dry sclerophyll forest, Newton & Thayer

13.2.1987 UV blacklight along river (43) FMNH;

Kiandra, Alpine Ck, E. Britton 27.1.1966 (2)

ANIC; Kosciusko NP, Diggers Ck 1510 m, T. A.

Weir 14.3.1969 (2) ANIC; Kosciusko NP, Betts

Ck 1740 m, 36°25'S 148°22'E, alpine meadow,

Newton & Thayer 14.2.1987 ex Sphagnum etc. at

stream edge and in bog (2) FMNH; Kosciusko

NP, Leather Barrel Ck 980 m, 36°32'S 148°12'E,

wet sclerophyll forest, Newton & Thayer

19.12.1986 flood debris ex large log jam, large

forest stream (2) FMNH; Kosciusko NP, Sawpit

Ck 1200 m, T. A. Weir 14.3.1969 (7) ANIC,

MSNV; Kyogle 345 m, Newton & Thayer

2.3.1980 blacklight (1) USNM; Lake Bathurst, C.

Reid 17.12.1984, shore mud & on grassland (1)

ANIC; Macksville, Wachtel 12.1990 (1f) SAMA;

Merindee Lakes Pk, Trust Caravan, G.P.Gross

26.12.1973 at light (6) SAMA; Moonbi Lookout

760 m, 30°58'S 151°06'E, Common & Edwards

11.12.1974 (1) ANIC; Moruya 8 km SE, Congo

35°58'S 150°09'E, M. S. Upton 8 & 14.2.1981 (2)

ANIC; Mossgiel 11 km N, Willandra Bridge

PARACYMUS OF AUSTRALIA 107

33°16'S 144°34'E, dry swamp, Britton, Misko &

Pullen 21.12.1970 at light (21) ANIC; Mt

Kaputar, 2000 ft, C. W. Frazier 4.9.1964, at light

(2) ANIC; Mt Keira, 1000 ft, Britton & Misko

7.3.1967 at light (1) ANIC; Mudgee 4 mi SW,

32°36'S 149°35'E, Britton & Misko 18.11.1968

(1) ANIC; Namoi R., Narrabri 30°19'S 149°47'E,

J. A. L. Watson 10.3.1969 at light (1) ANIC;

Nelson Bay 6 mi ESE, Britton & Misko

21.11.1967 at light (6) ANIC; Queanbeyan 2.7 km

NE, 670 m, I. F. B. Common 11.11.1976 (1)

ANIC; Sydney 100 m, H. P. Chandler 7.3.1943

(2) FMNH; Tamworth, Lea (2) SAMA; Uralla 6

km N, CSIRO Research Stn ‘Chiswick’, A. G.

Furnival 10.1.1972 (10), 20.11.72 (2), 30.11.73

(4) ANIC; Valery, A. P. M. Eucalyptus plantation,

Mc Mullens Block, R. McInnes 10.1.1967 (1)

light trap, ANIC; Wahroonga, H. J. Carter (4)

ANIC; Warrumbungle NP, Siding Spring Mtn, C.

Reid 23.XI.1985, Eucalyptus forest at light (8)

ANIC; Wee Jasper, E. F. Riek 20.4.1961 (4)

ANIC; Wentworth Fs, 5.1.1895 (1) SAMA;

Wingello 3 km SW, S. Misko 20.2.1974 (4) ex

pool, ANIC; Wingham Scrub, 31°52'S 152°22'E,

rainforest edge, Britton & Misko 3.1.1970 (3) at

light, ANIC; Woy Woy, Pearl Beach, C. Reid

1.12.1984 at light (1) ANIC. Northern Territory:

Adelaide R., at Daly R. Road Crossing, 13°29'S

131°06'E, E. B. Britton 9.11.1972 (12) ANIC;

Adelaide R., 10 km on Daly R. Road, E. F. Riek

25.10.1972 (1) ANIC; Adelaide R., 13°15'S

131°06'E, M.S.Upton 17.10.1972 (14) ANIC,

MSNV; Adelaide R. 27 km N, Coomalie Ck,

Gross & Forrest 28.9.1977 at light (16) SAMA;

Alice Springs 6 mi SE, Emily Gap, Britton, Upton

& Mc Innes 17.2.1966 (1) ANIC; Alice Springs 9

km N, Todd R., 23°38'S 133°53'E, M. S. Upton

10.10.1978 (1) ANIC; Alroy Downs HS 15 km

SW, 19°24'S 135°58'E, Key & Balderson

10.4.1976 (7) ANIC; Barrow Ck 2 mi S, Britton,

Upton & McInnes 13.2.1966 (4) ANIC; Boko Hill

1 km N, SW of Borroloola, 16°26'S 136°0O1'E,

Key & Balderson 14.4.1976 (2) ANIC; Borroloola

2 km SSE, McArthur R., 16°05'S 134°19'E, J. E.

Feehan 19-20.4.1976 (1) ANIC; Borroloola 4.5

km W, T. Reardon 8.1982 (1) SAMA; Borroloola

11 km SW, Goose Lagoon, 16°10'S 136°15'E, J.

E. Feehan 17.4.1976 at light (9), M. S. Upton

31.10.1975 (1) ANIC; Borroloola 12 km NNE,

15°538'S-13622VE,. M.S. Upton 1.11.1975 13)

ANIC; Borroloola 22 km WSW, 16°08'S

136°06'E, J. E. Feehan 16.4.1976 at light (30), M.

S. Upton 2.11.1975 (11) ANIC; Borroloola 30 km

NE, Batten Point, 15°54'S 136°32'E, J. E. Feehan

18.4.1976 at light (9) ANIC; Borroloola 31 km

WSW, Batten Ck, 16°10'S 136°03'E, J. E. Feehan

15.4.1976 at light (7) ANIC; Borroloola 33 km

SW, Caranbirini W.H., 16°16'S 136°OS'E, J. E.

Feehan 21.4.1976 at light (10), M. S. Upton

3.11.1975 (10) ANIC; Borroloola 45 km SW,

Surprise Ck, 16°25'S 136°0S'E, M.S.Upton

5.11.1975 (1) ANIC; Borroloola 46 km SSW,

16°28'S 136°09'E, J. E. Feehan 23.4.1976 (4), M.

S. Upton 28.10.1975 (1) ANIC; Borroloola 48 km

SW, McArthur R., 16°27'S 136°05'E, J. E. Feehan

13.4.1976 at light (4), M. S. Upton 29.10.1975 (1)

ANIC; Borroloola 54 km S, Cattle Ck, 16°32'S

136°10'E, M. S. Upton 27.10.1975 (1) ANIC;

Borroloola 80 km SW, McArthur R., 16°39'S

135°51'E, M. S. Upton 13.5.1973 (10) ANIC,

MSNYV; Burrell’s Ck, Stuart H’ way, D. H. Colless

25.11.1972 at light (1) ANIC; Cahills Crossing, E

Alligator River, 12°26'S 132°58'E, E. B. Britton

311051972, 321231972. at light (5), E: G.

Matthews 29.5.1973 at light (10) ANIC, SAMA;

Cahills Crossing 1 km N, E Alligator R., 12°23'S

132°57'E, Upton & Feehan 7.6.1973 (7), E.

Britton 31.10.1972 at light (2) ANIC; Cahills

Crossing 5 km NNW, E Alligator R., 12°23'S

132°57'E, E:-B: -Brittom 5:11, 1972 :@1)2 Ez -G:

Matthews 28.5.1973 (10), Upton & Feehan

8.6.1973 (4), A. H. Watson 8.6.1973 (1) ANIC,

SAMA; Cahills Crossing 7 km NW, E Alligator

R., 12°23'S 132°56'E, E. B. Britton 4.11.1972 at

light (10), E. G. Matthews 27.5.1973 (10) ANIC,

SAMA; Cape Crawford 8 km ESE, Bessie

Springs, 16°40'S 135°51'E, J. E. Feehan 12.4.1976

at light (13) ANIC; M. S. Upton 26.10.1975 (11),

J. E. Feehan 12.4.1976 at light (3) ANIC; Cape

Crawford 14 km NW, 16°34'S 135°41'E, M. S.

Upton 6.11.1975 (15) ANIC; Cape Crawford 14

km S, Mc Arthur R., 16°47'S 135°45'E, M. S.

Upton 25.10.1975 (9), J. E. Feehan 11.4.1976 (16)

ANIC; Colyer Lagoon, October Ck, Gross &

Forrest 26—27.9.1977 at light (3) SAMA; Curtin

Springs HS., Thurmer & Lacis 17.8.1978 (1)

SAMA; Daly R., J. C. Lesoeuf 12.7.1971 (5)

ANIC; Daly River 10 mi E, B. K. Head 28.6.1972

at light (41) SAMA; Daly R. Mission, J.

Hutchinson 6.6.1974, 8.10.1974 at light (2)

ANIC; Darwin, F. J. Gay 24.4.1966 (2) ANIC;

Darwin, B. Malkin 25.3.1945 (1) FMNH; nr

Darwin, Coastal Plains Rsrch Station CSIRO, E.

C. B. Langfield 6.6.1966 at light (17), 30.5.1966

at light (8) ANIC, MSNV; Darwin 24 km S,

Howard Springs, 12°28'S 131°03'E, E. B. Britton

10.11.1972 rainforest, at light (3), J. A. L. Watson

27.1.1968 at light (1) ANIC; Darwin 30 km SSE,

Berry Springs, 12°41'S 130°58'E, E. B. Britton

11.11.1972 at light (2) ANIC; Darwin 50 km S,

108 E. GENTILI

Coomalie Ck, G. F. Gross 28.9.1977 (1) SAMA;

Darwin 52 km S, Livingstone Field, Stuart

Highway, 12°44'S 132°05'E, E. B. Britton

9.11.1972 at light (16) ANIC; East Point, nr

Darwin, 12°28'S 130°50'E, E. B. Britton

12.11.1972 in flowers of Hybiscus tiliaceus (1)

ANIC; Elliott 15 km SW, L. Woods, Gross &

Forrest 5.10.1977 at light (25) SAMA; Groote

Eylandt, N. B. Tindale, in moss and lichens (1)

SAMA; Jabiru, C. Watts 22.3.1998 (6) SAMA,

MSNV; Kakadu NP, Upper S Alligator R.,

13°35'S 132°36'E, P. S. Cranston 4—5.6.1988 light

trap (8) ANIC; Katherine, L. P. Kelsey 16.8.1973

at light (1), E. G. Matthews 6-10.2.1968 (1)

ANIC; Katherine Gorge, M. J. Muller 26.10.1975

light trap (2) ANIC; Katherine 3 km SSW,

14°30'S 132°15'E, T. Weir 12.11.1979 (4) ANIC;

Katherine 25 km NE, Katherine R., Gross &

Forrest 3—4.10.1977 at light (24) SAMA;

Koongarra 12°52'S 132°50'E, M. S. Upton 6—

10.3.1973 (4) ANIC; Mataranka 5 km E, Roper

R., Gross & Forrest 27.9.1977 at light (8) SAMA;

Mataranka 19 km SSE, Elsey Ck, 15°0S'S

133°07'E, M. S. Upton 14.5.1973 (9) ANIC; Mt

Borradaile 19 km E, Cooper Ck 12°06'S 133°04'E,

E. B. Britton 2.11.1972 (4), E. G. Matthews

31.5.1973 at light (25), M .S. Upton 9.11.1972

(1), 5.6.1973 (8) ANIC, SAMA; Mt Cahill 6 km

E, 12°52'S 132°46'E, Nourlangie Ck, M. S. Upton

18.11.1972 (2) ANIC; Mt Cahill 8 km N,

Nourlangie Ck, 12°48'S 132°42'E, E. B. Britton

26.10.1972 (14), id., D. H. Colless (1), M. S.

Upton 19.11.1972 (3), Upton & Feehan 16.6.73

(23), E. G. Matthews 21.5.73 (6) at light ANIC,

SAMA, MSNV; Mt Cahill 8 km E, Nourlangie

Ck, 12°52'S 132°47E, E. B. Britton 27.10.1972

(58), id., mud at edge of waterhole (7), E. G.

Matthews 22.5.1973 at light (9) ANIC, SAMA,

MSNV; Mt Cahill 10 km E, 12°51'S 132°47'E, E.

G. Matthews 21.5.1973 (2) ANIC; Mt Cahill 12

km NNW, 12°46'S 132°39'E, E. B. Britton

25.10.1972 at light (1), Matthews & Upton

20.5.1973 (20), Upton & Feehan 15.6.1973 (3)

ANIC, SAMA; Mt Cahill 15 km E, Koongarra,

12°52'S 132°50'E, M. S. Upton 15.11.1972 (11),

Upton & Feehan 12.6.1973 (9) ANIC; Mt Cahill

15 km E by N, 12°50’N 132°51'E, E. B. Britton

29.10.1972 at light (4), D. Colless 30.10.1972 by

sweeping (8) ANIC; Mt Cahill 16 km E by N,

12°50'S 132°51'E, Upton & Feehan 13.6.1973

(14) ANIC, SAMA; Mt Cahill 19 km NE,

Baroalba Ck Springs, 12°47'S 132°51'E, E. B.

Britton 28.10.1972 (43), M. S. Upton 16.11.1972

(8) ANIC; Mt Cahill 19 km WSW, Jim Jim Ck,

12°57'S 132°33'E, E. B. Britton 24.10.1972 at

light (8), Upton & Feehan 17.6.1973 (6) ANIC,

SAMA; Mt Cahill 30 km WSW, 12°58'S

132°26'E, E. G. Matthews 19.5.1973 (1) ANIC;

Mt Cahill 46 km WSW, S Alligator River,

13°03'S 132°19'E, Matthews & Upton 20.5.1973

(7) SAMA, ANIC; Mudginbarry HS. 2 km N,

Magela Ck, 12°35'S 132°52'E, M. S. Upton

14.11.1972 (9) ANIC; Mudginbarry HS. 9 km

SSE, Magela Ck, 12°40'S 132°54'E, E. B. Britton

6.11.1972 at light (20) ANIC; Mudginbarry HS. 9

km N, 12°31'S 132°54'E, Upton & Feehan, E. G.

Matthews 26.5.1973 (12) ANIC, Upton & Feehan

10.6.1973 (9) ANIC, SAMA, E. B. Britton

30.10.1972 (16) ANIC; Nabarlek Dam, 15 km S

of Nimbuwah Rock, 12°20'S 133°19'E, E. G.

Matthews 2.6.1973 at light (10) ANIC, SAMA;

Nimbuwah Rock 11 km S, Cooper Ck, 12°17'S

133°20'E, E. B. Britton 1.12.1972 at light (12),

Feehan & Upton 3.6.1973 (3) ANIC; Oenpelli 6

km SW, 12°22'S 133°01'E, E. G. Matthews

30.5.1973 at light (2), Upton & Feehan 6.6.1973

(5) ANIC, SAMA; Oenpelli 18 km E, 12°17'S

133°13'E, Matthews & Upton 1.6.1973 (6) ANIC;

Pine Ck, C. Watts 5.5.1963 (2) SAMA; Port

Keats, 14°06'S 129°33'E, M. Mendum 19.8.1968

(1) ANIC; Renner Springs 4.8 km S, N.

McFarland 8.3.1966 UV light (11) SAMA;

Timber Ck 4 mi W, N. McFarland 14.4.1966 UV

light (67) SAMA; Tindal, 14°31'S 132°22'E, W. J.

M. Vestjens 1—20.12.1967 light trap (50) ANIC,

MSNYV; Victoria River Downs 6.4 km SSW, L. P.

Kelsey 14-17.7.1973 (4) ANIC; Victoria River

Downs 4 mi. WSW, Irrigation Farm, L. P. Kelsey

13.9.1973 (2) ANIC; Victoria River Downs 8 km

WSW, L. P. Kelsey 14.8.1973 at light (10) ANIC;

Yuendumu, C. Watts 3.1965 (3) SAMA; Wildman

River Lagoon, 12°58'S 132°00'E, E. B. Britton

24.10.1972 (6) ANIC; Woolwonga Fauna Res.,

Dreaming Water, E. F. Riek 20.10.1972 (2) ANIC,

MSNV. Queensland: Annan Falls 1 km W,

15°31'S 145°14'E, E. B. Britton 26.5.1976 (7)

ANIC; Archers Ck, Mt Garnet Rd., J. G. Brooks

28.12.1964 (2) ANIC; Archers Ck 2220’, J. G.

Brooks 18.4.1974 at light (9) SAMA; Barkley

Hwy 7 mi N, on Burketown Rd., J. A. Forrest

23.9.1977 burnt out area, some regrowth, at light

(1) SAMA; Boulia 42 km NNW, 22°35'S

139°43'E, M. S. Upton 11.5.1973 (1) ANIC;

Bowen, A. Simson (1) SAMA; Brisbane (4)

SAMA; Brisbane 50 km S, Canungra, Pollock &

Reichert 11.1.1991, black light (105) NMW,

MSNV; Cape York Pen., Old Strathgordon,

H’stead W of Musgrave, Walford & Huggins

24.11.1983 (9) ANIC; Cardstone, J. G. Brooks

14.11.1966 (3), K. Hyde 17-23.2.1966 (13)

PARACYMUS OF AUSTRALIA

ANIC; Cardwell Range, J. G. Brooks 30.9.1967

(8) ANIC; Charleville S, M. S. Upton 9.5.1973

(1) ANIC; Cairns Distr., A. M. Lea, at light (6)

SAMA; Chillagoe Ck, campsite, Ellis & Hawkins

8.8.1967 (1) ANIC; Cooktown, Airport Rd.,

roadside swamp, 15°28'S 145°11'E, E. B. Britton

24.5.1976 (9) ANIC; Cooktown 3 km S, Keatings

Gap, 15°30'S 145°15'E, Common & Edwards

16.5.1977 (6) ANIC; Cooktown 21 km W,

15°25'S 145°03'E, Common & Edwards

17.5.1977 (5) ANIC; Cooktown 25 mi N, Mc Ivor

R., S. R. Curtis 6.5.1970 (37) ANIC; Cooktown

75 km, Cooktown Rd., Boggy Ck, E. B. Britton

26.4.1976 (8) ANIC; Cunnamulla, A. Hardcastle

(2) SAMA; Dalby, F. H. Hobler, Griffith Coll. (1)

SAMA; Dalrymple 300 m, 30 km N Charters

Towers (2) NMW; Eidsvold N, Burnett R.,

24°46'S 152°25'E, Holloway & Misko 10.1.1970

at light (2) ANIC; Forty Mile Scrub N. P., Mt

Garnet 52 km SW, 18°05'S 144°52'E, Weir &

Calder 21.7.1986 (24), 55 km S, 18°06'S

144°SO'E, J. Balderson 29-30.11.1981 (1) ANIC;

Funnel Ck, 21°47'S 148°5S'E, Britton & Misko

12.12.1968 at light (7) ANIC; Gladstone 23 km

SE, Calliope R., 23°50'S 151°13'E, S. Misko

23.1.1970 (11) ANIC; Green Hills, J. G. Brooks

19.12.1967 (3) ANIC; Herberton 7 mi SW,

17°27'S 145°27'E, Britton & Misko 6.12.1968 at

light (8) ANIC; Hope Vale Mission 7 km N,

15°14'S 145°07'E, T. Weir 4.10.1980 (1) ANIC;

Ingham, K. L. Harley 24.2.1960 (2), 30.3.1960 (2)

ANIC; Ingham 23 mi SSE, 18°58'S 146°16'E,

Britton & Misko 9.12.1968 at light (69) ANIC;

Iron Range, 12°42'S 143°18'E, J. G. & J. A. G.

Brooks 15.5.1971 at light (1) ANIC; Julatten,

Bushy Ck, 16°37'S 145°21'E, E. B. Britton

3.12.1968 from gravel at water’s edge (12) ANIC;

Kelly, St George R., Cooktown Rd, 16°29'S

144°47'E, E. B. Britton 22.5.1976 (1) ANIC;

Kennedy Forest Rd, rainforest 18°13'S 145°47'E,

Britton & Misko 8.12.1968 (1) ANIC; Kingaroy

24 mi SW, 26°44'S 151°31'E, Britton & Misko

21.11.1968 (1) ANIC; Kuranda, Barron Falls, J.

G. Brooks 12.12.1964 (2) ANIC; Lake Barrine,

18-22.9.1965, E. Britton (2) ANIC; Laura, C.

Watts 18.7.1982 (3), 2.8.1974 (1) SAMA; Laura

73 km NW, Hann R., 15°12'S 143°52'E, Weir &

Calder 27.6.1986 (25) ANIC; Longreach 31 km

NW, Darr R., 23°13'S 144°04'E, M. S. Upton

10.5.1973 (8), 22.10.1975 Darr R. (3) ANIC; L’tle

Mulgrave R., J. G. Brooks 16.12.1967 (11) ANIC;

MacDonald N.P., Mt Tamborine, J. & E. Doyen

26.11.1982 (1) ANIC; Mackay W, Finch Hatton

Ck, 21°08'S 148°38'E, S. Misko 29.11.1968 (5)

ANIC; Mackay 50 mi W, Broken River, Misko &

109

Britton 29.11.1968 rainforest, at light (2), S.

Misko 30.11.1968 (6) ANIC; Mareeba, K. & E.

Carnaby 22.5.1976 (1) ANIC; Mareeba 24 km N,

16°47'S 145°22'E, J. Balderson 24—25.11.1981 (1)

ANIC; Mary Ck, 16°33'S 145°12'E, Britton &

Misko 4.12.1968 at light (108) ANIC; Mary Ck,

22 km N of Mt Molloy, J. G. Brooks 14.3.1970 at

light (20) ANIC; Miriam Vale 21 mi S, 24°38'S

151°34'E, Britton & Misko 14.12.1968 (14)

ANIC; Monto 22 km NW, Coominglah St. For., J.

T. Doyen 23.12.1982 (4) ANIC; Mornington

Island Mission, Aitken & Tindale 7—-11.5.1963 at

light (4) SAMA; Mourangee nr Edungalba, 50 mi

SW of Rockampton, E. Adams 24.11.1968 at light

(3), Britton & Misko 26.11.1968 at light (2)

ANIC; Mt Baird 3.5 km SW, 15°10'S 145°07'E,

A. Calder 3-5.5.1981 (7) ANIC; Mt Baldy nr

Atherton, Forest Res. No. 194, 4000’, rainforest,

Britton & Misko 5.12.1968 (13) ANIC; Mt

Carbine 35 km NNW, J. T. Doyen 13.12.1982 (2)

ANIC; Mt Cook N. P., 15°29'S 145°16'E, A.

Calder 10-12.5.1981 (1) ANIC; Mt Coolum,

26°35'S 153°0S5'E, Britton & Misko 15.12.1968 at

light (25) ANIC; Mt Inkerman 2 mi SW, 19°45'S

147°30'E, Britton & Misko 11.12.1968 mud, lily

ponds (4) ANIC; Mt Lewis, ca. 3000’, rainforest,

Britton & Misko at light tin working site,

3.12.1968 (2) ANIC; Mt Molloy 17.7 km N,

Station Ck 427 m, J. G. Brooks 21.12.1970 (1)

ANIC; Mt Tozer 3 km ENE, 12°44'S 143°14'E,

Weir & Calder 28.6-4.7.1986 (23), J. C. Cardale

(1) ANIC; Mt Tozer 11 km ENE, 12°43'S

143°18'E, Weir & Calder 11-16.7.1986 (5) ANIC;

Mt Webb 3 km NE, 15°03'S 145°09'E, A. Calder

30.4-3.5.1981 (3) ANIC; Nettle Ck, J. G. Brooks

20.8.1969 (2) ANIC; Normanton, Tindale &

Aitken 4.5.1963 at light (3) SAMA; Paluma 2 km

W, Ewan Rd 800 m, 19°06'S 146°34'E, J. G.

Brooks 22.2.1972 at light (1) ANIC; Paluma 9 km

W, J. G. Brooks 4—13.12.1973 at light (1) ANIC;

Pentland, J. C. Lesoeuf 18.7.1975 (1) ANIC;

Pistol Gap, Byfield, 22°50'S 150°40'E, Britton,

Holloway & Misko 10.1.1970, dry sclerophyll, at

light (4) ANIC; Proserpine 8 mi NE, Brandy Ck,

20°20°5S 148°41'E, Britton & Misko 11.12.1968

at light (2) ANIC; Ravenshoe 17.7 km W, Archers

Ck, J. G. Brooks 13.4.1974 (18) ANIC; Reedy St

George R., Cooktown Rd, E. B. Britton 22.5.1976

(1) ANIC; Stanthorpe 9 mi. S, Conardoo, Fletcher

28°46'S 151°51'E, Britton & Misko 20.11.1968

(6) ANIC; Station Ck, J. G. Brooks 14.4.1970 (2)

ANIC; Stuart R., Hale & Tindale 1-2.1927 (2)

SAMA; Summit 4 km W, Cunninghams Gap N.P.,

J. Doyen 27—28.11.1982 (1) ANIC; Townsville,

B. Malkin 1—2.1945 (2) FMNH; Townsville, P.

110

Ferrar 23-30.5.1968 light trap (1) ANIC;

Townsville 10 m, G. Wewalka 17.1.1993 (2)

NMW; Townsville 5 km N, at Town Common,

19°15'S 146°48'E, S. Misko 19.1.1970 at light (1)

ANIC; Wenlock R., Xing Portland Roads Road,

13°06'S 142°56'E, Weir & Calder 17.7.1986 (1)

ANIC; Woodstock 7 km S, Lansdown Station,

19°40'S 146°S51'E, R. A. Barrett 16.1.1974 (4)

ANIC. South Australia: Barossa, B. J. Burton (2)

SAMA; Devon Downs, S. A. Museum Exped. (1)

SAMA; Donovans 6.4 km NW, Ponds Cave,

Aitken & Tindale 28.1.1965 (13) SAMA; Eyre

Pen., McKeckives Spr., White Flat Rd, Bishop &

Diener 14.12.1976 (8) SAMA; Eyre Pen., Strm nr

Epsom Sp., White Flat Rd., Bishop & Diener

14.12.1976 (6) SAMA; Eyre Pen., Todd R., White

Flat Rd., Bishop & Diener 14.12.1976 (9) SAMA;

Eyre Pen., Woolshed Ck, Bishop & Diener

13.12.1976 (2) SAMA; Fairview Cons. Pk, J. A.

Forrest 1.4.1982 at light (12) SAMA; Fairview

Wildlife Res., 36°49'S 140°24'E, Matthews &

Forrest 23.3.1981 at light (2) SAMA; Finness R.,

R. Malcolm 4.1976 (2) SAMA; Flinders Ranges,

Arkaba Ck, E. G. Matthews 5.3.1973 (1) SAMA;

Lake Fox edge, P. J. M. Greenslade 18.11.1978,

ex litter samples (1) SAMA; Lake George,

37°20'S 140°10'E, Roffey & Mitchell 13.10.1972

(1) ANIC; Monarto Sth, P. McQuillan 19.1.1973

UV light (2) SAMA; Mosquito Ck SE mouth at

Haks Lagoon, Thurmer & Gackle 23.4.1979 in

water (1) SAMA; Mt Crawford Forrest, C. Watts

10.11.1996 (3) SAMA; Mt Gambier 27 km NE,

nr Linwood Pk., swamp in Pine forest, J. A.

Forrest 26.3.1982 (4) SAMA; Mt Gambier, Valley

Lake, K. F. Walker 1.1.1975 (11) SAMA; Mt

Remarkable NP, Mambray Ck, J. A. Forrest

7.5.1981, E. G. Matthews 17.1.1982 (7) SAMA;

Murbko, R. Murray, G. F. Gross 20.2.1973 (2)

SAMA; Murray R., R. J. Burton (3) SAMA;

Mylor, Scout Jamboree 20.12.1973-6.1.74, el.

light (1) SAMA; Nanam’s Well 15 km SW,

Scorpion Springs C.P., Museum Party 14.12.1983,

at light (2) SAMA; New Kalamurina St.,

Warburton R., Matthews & Houston 9.3.1972 (1)

SAMA; Olary 24 km WNW, 32°17'S 140°19'E,

Britton, Misko & Pullen 20.12.1970 at light (3)

ANIC; Oodnadatta, Blackburn (1) SAMA; Penola

Cons. Pk, Penola 14 km W, J. A. Forrest

24.3.1982 (1) SAMA; Penola W nr Calectasia NP,

Baker Range Drain, J. A. Forrest 24.3.1982 (12)

SAMA, MSNV; Port Lincoln, Blackburn (5)

SAMA; Robe 10 km S, C. Watts 1.1983 (2)

SAMA; Rudall 2 km S, 33°41'S 136°16'E,

Britton, Misko & Pullen 18-19.12.1970 (1)

ANIC; Salt Creek 17 mi SE, Gross & Aitken

E. GENTILI

14.1.1962 at light (3) SAMA; Tintinara 15 mi E,

Jimmy’s Well, Aitken & Tindale 3.2.1965 (2)

SAMA; Yorke Pen., 8 km WSW Carritin Hs., S

end of Formby Lisy, N. McFarland 4.11.1965 (1)

SAMA. Tasmania: Barrow Ck, Mt Barrow 6 km

NW, 41°21S 147°22'E, E. & S. Britton 3.2.1973

(1) ANIC; Forest Reefs, Griffith Coll. (4) SAMA;

Frankford, A. M. Lea (2) SAMA; George, C. E.

Cole 3.11.1917 (2) SAMA; Hobart, base Mt

Wellington, L. Hill 26.1.1979 ex moss & grass (1)

ANIC; Kelso (1) SAMA; Kempton Water Tray, L.

Hill 28.11.1985 (1) ANIC; Launceston, F. M.

Littler (2) SAMA; Orford 4 km W, 42°34'S

147°50'E, J. C. Cardale 27.1.1983 at light (1)

ANIC; Tooms R., 460 m, 42°13'S 147°46'E, L.

Hill 19.4.1981 (4 ff) ANIC, MSNV. Victoria:

Alexandra 25 km S, Cathedral Range Nat. Park,

Blackwood flat Campground, 445 m, Pollock &

Reichert 5.12.1990 black light (1f) NMW;

Ballarat, W. W. Froggatt (1) ANIC; Baw Baw

Alpine Res., Neulines Mill 1.2 km NW, 1145 m,

37°51'S 146°15'E, wet sclerophyll & Nothofagus

cunninghami, Newton & Thayer 29.1.1987

berlese leaf and log litter forest floor (1f) FMC;

Billabong, Yara Glen, A. Fletcher 20.4.1976 (1)

SAMA; Birchip V., J. C. Goudie (1f) ANIC;

Dartmooor 5 km NE, C. Watts 11.10.1997 (1)

SAMA; Dimboola, Caravan Park, S. Misko

18.11.1973 light trap (5) ANIC; East Pomborneit,

24 km ESE Campdown, temporary pond, P. S.

Lake 5.X.1978 (5) ANIC; Healesville, Goudie &

Lea 11.? (1), C. Watts 12.1968 (5) SAMA,

MSNV; Kawarren 3 km N, Otway Ranges, Gross

& Aitken 15.1.1962 at light (14) SAMA; Kerang,

R. Blackwood 4.1935 (1) ANIC; Lake Hattah (4)

ANIC; Lake Hattah, G. W. Anderson 24—

25.10.1967 (2), 28.11.1967 (4), 9-15.3.1969 (2),

9.12.1969 (4) light trap ANIC; Lake Learmonth,

E. F. Riek 15.12.1966 (2ff) ANIC; Lerderderg

R., 3.8 km WNW Blackwood, A. J. Boulton 27.6

& 25.11.1982 (3) ANIC; Lilydale, A. Fletcher

2.7.1976 (1) SAMA; Lorne, Cressy Ck, N. B.

Tindale 21.1.1963 (5) SAMA; Mirrantwa 10 km

NE, C. Watts 12.10.1997 (11) SAMA; Mt

Buffalo, Blackburn (5) ANIC, SAMA; Mt Buffalo

NP, 1310 m, alpine bog, Sphagnum moss, Newton

& Thayer 18-19.1.1980 (1) FMNH; Noojee 6 km

N, C. Watts 8.11.1997 (2) SAMA; Orbost 12 km

SW, C. Watts 5.11.1997 (5) SAMA; Otway NP,

Binn Rd. 450 m, Cape Horn 58 km N, 38°42'S

143°34'E, wet sclerophyll forest, Newton &

Thayer 24.1.1987 (1) FMNH; Ovens R.,

Wangaratta, Newton & Thayer 9.1.1980 black

light (1f) ANIC; Ovens R., Porepunkah, 300 m,

36°42'S 146°55'E, mixed dry sclerophyll and

PARACYMUS OF AUSTRALIA 111

exotic trees, Newton & Thayer 12.2.1987 UV

blacklight nr River (1f) FMNH; nr Porepunkah,

A. Newton 18.1.1980 UV light (1) ANIC;

Portland 30 km W, C. Watts 10.10.1997 (2)

SAMA; Shepperton 13 km SE, S. Misko

22.11.1973 (2ff) ANIC; Tangil River E, C. Watts

8.11.1997 (8) SAMA; Violet Town 14 km NW,

Rd to Shepperton, S. Misko 22.XI.1973 (5)

ANIC; Yarra River, Healesville 4,5 km SW, 80

m, 37°41'S 145°29'E, dry sclerophyll forest,

Newton & Thayer 6.2.1987 UV black light along

river (195) FMNH; Warburton, A. Newton 13-

17.1.1980 UV light (1f) ANIC; Warburton 12 km

E, 215 m, Eucalyptus forest, leaf litter stream

edge, Newton & Thayer 12—16.1.1980 (1) FMNH;

Wyperfield Nat. Park, Frew’s Plain 35°37'S

142°0O1'E, S. Misko 15-17.XI.1973 at light (14)

ANIC, MSNV; Wyperfield Nat. Park, Lowan

Treck 35°35'S 142°0S'E, S. Misko 16.XI.1973

light trap (21) ANIC, MSNV; Wyperfield Nat.

Park, Ranger’s House, 35°37'S 142°01'E, Misko

& Anderson 17.XI.1973 light trap (4) ANIC;

Wyperfield NP, Common-Upton 5.11.1966 (1)

ANIC. Western Australia: Albany, K. & E.

Camaby 15.12.1976 (5) ANIC; Albany 48 km N,

Porongorup NP, J. Kethley 24.12.1976 soil litter

und grasses (1) FMNH; Appleton, F. H. Uther

Baker, 4.1.1966 (10) ANIC; Armadale 15 mi SSE,

Pipehead Dam, M. S. Upton 26.1.1967 (7) ANIC;

Armadale, D.E. 7.1961 (1) SAMA; Beverley, A.

M. Lea 1870 (5) SAMA; Bunbury, K. & E.

Carnaby 31.12.1971 at light (14) ANIC; Bunbury,

Whitlock (7) ANIC; Capel, E. Britton 29.10.1965

(6) ANIC; Carson Escarpment, 14°49'S 126°49'E,

Common & Upton 9-15.8.1975 (11) ANIC,

MSNV; Collie 16 mi N, Common & Upton

7.4.1968 (1) ANIC; Coodanup nr Mandurah, T. E.

Bellas 12.1979-1.1980 (1) ANIC; Darling Rgs.,

A. M. Lea (1) SAMA; Deepdene, Karridale, M. S.

Upton 18.1.1967 (10) ANIC; Denmark, Walpole

Rd, roadside pool, E. Britton 21.9.1965, 24.9.65

(2) ANIC; Denmark 14 mi E, Parry’s Inlet turnoff,

35°O1'S 117°09'E, E. Britton 9.11.1969 roadside

pond (45) ANIC; Donnybrook, A. M. Lea 1870

(3) SAMA; Drysdale R., 15°02'S 126°55'E,

Common & Upton 3-8.8.1975 (10) ANIC;

Drysdale R., 14°39'S 126°57'E, Common &

Upton 18-21.8.1975 (10) ANIC, MSNV;

Dunsborough 20 m, J. B. Kethley 11.1976 at light

(1) FMNH; Esperance 20 km E, 33°50'S

122°06'E, J. F. Lawrence 8.11.1977 at light (16)

ANIC; Esperance 101 km E, Thomas R., 33°51'S

121°53'E, Britton, Taylor & Upton 20.11.1969 at

light, beach dunes (24) ANIC; Fitzroy R., K. & E.

Carnaby 16.4.1976 at light (7) ANIC; Fitzroy

Crossing, K. & E. Carnaby 18.4.1976 at light (5)

ANIC; Fremantle 10 km S, C. Watts 24.10.1996

(1) SAMA; Fremantle, North Lake, H. Demarz

30.1.1954 (2) FMNH; Geraldton 11 km N, N.

McFarland 12.11.1972 UV light (3) ANIC; Hyden

E, K. & E. Carnaby 5.2.1977 (1) ANIC; Julimar

St. Forest, E. Matthews 9.10.1967 (2) ANIC;

Kimberley E, Benn R., Helms (6) SAMA;

Kimberley E, Upp. Ord R., Helms (1) Griffith

Coll., SAMA; Kuliba, Ravensthorpe-Hopetoun, E.

Britton 21.9.1965 (1) ANIC; Kununurra nr.

Wyndham, Kimberley Research Station, 15°28'S

128°06'E, 27.X1.1956 (31) ANIC, MSNV;

Kununurra, E. G. Matthews 13-22.2.1968 (1)

ANIC; Kununurra 100 mi E, J. A. Mahon

27.3.1966, light trap (1) ANIC; Margaret R.,

roadside pond, E. Britton 29.9.1965 (1) ANIC;

Mitchell Plateau, 14°40'S 125°44'E, B. V. Timms

23.9.1982 (1) SAMA; Mitchell Plateau, King Ed.

R. crossing, Rd., J. B. Kethley 14.10.1976 at light

(1) FMNH; Mitchell Plateau, Mining Camp,

14°49'S 125°50'E, Naumann & Cardale 9-

19.5.1983 at light (7) ANIC; id., Rentz &

Balderson 9-19.5.1983 on light sheet (5) ANIC;

id. 3 km NW, 14°48'S 125°49'E, Rentz &

Balderson 15.5.1983, airstrip (4) ANIC; id. 4 km

S, 14°52'S 125°50'E, Rentz & Balderson

13.5.1983, crusher at light (17) ANIC; id. 10 km

NW, 14°45'S 125°47'E, Rentz & Balderson 11-

17.5.1983 (2) ANIC; Mowen 10 mi E, Margaret

River, 33°57'S 115°34'E, E. Britton 14.11.1969

roadside pond (10) ANIC; Mt Arid 23 km NW,

Thomas River 33°51'S 123°00'E, J. F. Lawrence

4-7.11.1977 (14) ANIC; Mt Chudalup St. Pk,

North Cliffs 16 km S, J. Kethley 4.12.1976 wet

moss on sand over seepage area (1) FMNH;

Nannup, Augusta Rd., E. Britton 28.9.1965 (1)

ANIC; Osmington 5 mi N, nr Margaret R.,

33°57'S 115°04'E, E. B. Britton 15.11.1969 edge

of roadside pool (3), under bark of recently felled

trees (1) ANIC; Pago Mission 3 mi SE, J. B.

Kethley 26.10.1976 at light (3), 27.10.1976 small

pool (6) FMNH; Perth, Floreat Park, M. S. Upton

21.1.1967 (1) ANIC; Perth E, Orange Grove

Caravan Pk Gosnells, Reid & Gullan 5.1.1986 at

light (2) ANIC; Perth S, Yule Brook Univers.

Res., C. Reid & P. J. Gullan 7-8.1.1986 at light

(23) ANIC; Perth 30 km N, C. Watts 14.10.1996

(3) SAMA; Picton Junction, swamp nr Ferguson

R., Britton & Uther Baker 30.11.1965 (16) ANIC;

Pinjarra, Lea 1870 (10) SAMA; Pinjarra 8 mi E,

South Dandalup R., 32°35'S 115°53’E, E. Britton

17.11.1969 clear brook flowing over stones (42)

ANIC, MSNV; Pinjarra 6 km S, C. Watts

6.10.1996 (9) SAMA; Walpole Rd, Nornalup

112

Beach Rd, S. & J. Peck 20—26.6.1980, flight

intercept traps (2) ANIC; Walsh Pt 5 km W,

14°34'S 125°48'E, Rentz & Balderson 10.5.1983

(2) ANIC; Wilga, K. & E. Carnaby 11.1973,

8.12.74 etc. (33) ANIC; William Bay, stream on

road to, E. Britton 24.9.1965 (3) ANIC;

Wyndham, K. & E. Carnaby 20.4.1976 at light (1)

ANIC; Yanchep NP, Yanchep 9 mi N, Common

& Upton 12.4.1968 (1) ANIC.

Biology

From label data I extracted the following

indications about the habits of P. pygmaeus. Most

specimens are collected in flight: at light, light

trap, on light sheet, UV light, black light, flight

intercept traps, window-trough traps. Another

method is collecting along edges, shores or banks

of brooks, creeks, rivers, pools, bogs, swamps,

waterholes, ponds and lakes: in mud, sand, gravel,

aquatic plants such as lilies, Sphagnum, moss, and

lichens; more rarely in open water. Also, litter is a

source of specimens: wet rotting wood, bark of

fallen trees, leaves on forest floor, soil and

grasses, floating debris. Exceptional captures are:

from nest of buff-tailed thornbill; in flowers of

Hybiscus tiliaceus; on Acacia spp.

2. Paracymus spenceri Blackburn, 1896

Paracymus spenceri Blackburn 1896: 256-257;

d’Orchymont 1942: 59-60; Wooldridge 1976:

454-455; Hansen 1999: 114.

Paranacaena spenceri (Blackburn): Knisch 1924:

168.

Types

Lectotype male: Northern Territory: Paisley

Bluff, Reedy Creek, Blackburn Coll. NHML. A

single pin bears the following cards and labels: 1.

Insect (and) I, 5480, Reedy Cr., male; 2. Aedeagus

(on transparent card); 3. Type (round label with red

border); 4. Australia, Blackburn Coll., B.M. 1910 —

236; 5. Paracymus spenceri Blackb.; 6. Paranacaena

spenceri Blackb., LECTOTYPUS male, E. Gentili

1991, Aedeagus in DMHF soluble in distilled water.

Paralectotype: Northern Territory: Paisley Bluff,

Reedy Creek SAMA. A single pin bears: 1. Insect

(dorsally glued, and) 5480, Reedy Creek; 2.

Paracymus spenceri Blackb., co-type; 3. Paracymus

spenceri Blackb., Paralectotypus, E. Gentili 1992

(red label); 4. S. A. Museum specimen (red label).

The Blackburn description records four typical

specimens from Paisley Bluff, Reedy Creek; I did

not see other types.

E. GENTILI

Description

Length 1.5-2.3 mm; width 0.8-1.3 mm.

Elongate oval, slightly convex (Figs 18, 19). Head

black, sometimes with violet reflection, evenly

punctured, surface shining between punctures,

occipital region alutaceous; only lateral branches

of Y-suture scarcely conspicuous. Pronotum dark

as head, becoming yellow to mahogany at sides;

punctation thicker than on head, surface shining

between punctures. Elytra mahogany or red-

brown, lighter near apical region; punctation as on

pronotum but less evident, without serial

arrangement; some specimens nevertheless have

traces of serial punctures due to partial

transparency of elytra; parasutural furrow in apical

2/3 of elytra or a bit less. Underside dark;

prosternum with longitudinal keel; mesosternum

carinate with an arrow-like keel; first visible

abdominal ventrite laterally shorter than second,

centrally with thin longitudinal keel. Maxillary

palpi yellow, tips darker; antennae eight-

segmented, yellow with darker club. Profemora

ventrally pubescent on basal 3/4 (or 2/3), male

protarsi with a blunt spine under last segment;

mesofemora pubescent near base; metafemora

without hydrofuge hairs. Aedeagus as in Figs 6-8,

tegmen nearly as long as parameres, with a

cardioid form; parameres progressively narrowing

but slightly expanded laterally at their tip; penis

roughly triangular, provided with a subapical

collar.

Discussion

Easy to distinguish among Australian

Paracymus by the contrasting colour of the

pronotum/elytra. The male protarsus is thinner,

and the profemoral pubescence thicker than in

pygmaeus. The shape of the aedeagus is near to

that of pygmaeus, but the outline of the tegmen

and parameres is more curved, and the preapical

collar lacks any membranous expansion.

Material examined (Fig. 23)

Australian Capital Territory: Black

Mountain, I. F. B. Common 25-26.12.1964 light

trap (1), 6.1.1966 light trap (1), M. S. Upton

22.12.1966 light trap (1) ANIC. New South

Wales: Dorrigo NP, E end, Blackbutt Track 710

m, subtropical rainforest, Newton & Thayer

28.2-5.3.1980 in and under rotting fruits of

Endiandra introsa (1) USNM; Nightcap NP, Mt

Nardi, Newton Drive 700 m, warm-temperate

rainforest, 28°33'S 153°17'E, Newton & Thayer

4.1.1987 Berlese leaf and log litter, forest floor

(5) FMNH; Uki 18 km W, Mebbin St. For., J.

PARACYMUS OF AUSTRALIA 113

FIGURE 23. Distribution of the studied specimens of Paracymus spenceri Blackburn, 1896.

Doyen 23-24.11.1982 (1) ANIC. Northern

Territory: Alice Springs 80 mi NW, C. Watts

2.1968 (5) SAMA; Alice Springs 5 mi N, Wigley

Waterhole, Britton Upton & McInnes 16.2.1966

(2) ANIC; Alice Springs 6 mi SE, Emily Gap,

Britton Upton & McInnes 17.2.1966 (2) ANIC;

Alice Springs 9 km N, Todd R., 23°38'S

133°53'E, M. S. Upton 10.10.1978 (3) ANIC;

Alice Springs 33 km WNW, 23°36'S 133°34'E,

M. S. Upton 30.9.1978 (3) ANIC, MSNV; Alice

Springs 99 km NE, Ongeva Ck, 23°01'S

134°29'E, M. S. Upton 13.10.1978 (1) ANIC;

Batchelor 1 km SE, N. McFarland 12.4.1966 UV

light (3) SAMA; Borroloola 11 km SW, Goose

Lagoon, 16°10'S 136°15'E, J. E. Feehan

17.4.1976 at light (4) ANIC, MSNV; Borroloola

22 km WSW, 16°08'S 136°06'E, J. E. Feehan

16.4.1986 at light (1) ANIC; Borroloola 30 km

NE, Batten Point, 15°54'S 136°32'E, J. E. Feehan

18.4.1976 (1) ANIC; Borroloola 31 km WSW,

Batten Ck, 16°10'S 136°03'E, J. E. Feehan

15.4.1976 at light (1) ANIC; Borroloola 45 km

SW, Surprise Ck, 16°25'S 136°0S'E, J. E. Feehan

14.4.1976 at light (1) ANIC; Borroloola 48 km

SW, McArthur R., 16°27'S 136°0S'E, J. E.

Feehan 13.4.1976 at light (1) ANIC; Edith Falls,

J. C. Lesoeuf 8.7.1971 (1) ANIC; Mt Cahill 8

km N, Nourlangie Ck, 12°48'S 132°42'E, Upton

& Feehan 16.6.1973 (1) ANIC; Renner Springs

4.6 km S, N. McFarland 8.3.1966 UV light (1)

SAMA; Timber Ck 4 mi W, N. McFarland

14.4.1966 (2) SAMA; Wauchope 30 km N,

20°22'S 134°14'E, M. S. Upton 13.10.1972 (5)

ANIC, MSNV. Queensland: Archer R., C. Watts

28.10.1984 (1) SAMA; Barkley Hwy 7 mi N, on

Burketown Rd, J. A. Forrest 23.9.1977 burnt out

area some regrowth, at light (1) SAMA; Cairns,

Edge Hill, J. G. Brooks 11.11.1967 (1) ANIC;

Cardstone, K. Hyde 9-13.3.1966 (3) ANIC;

Cooktown Rd 75 km, Boggy Ck, E. B. Britton

26.4.1976 (3) ANIC; Eidsvold N, Burnett R.,

24°46'S 152°25'E, Britton. Holloway & Misko

10.1.1970 at light (2) ANIC; Forty Mile Scrub

Nat. Pk, Mt Garnet 52 km SW, 18°0S'S

144°52'E, Weir & Calder 21.7.1986 (2) ANIC;

Funnel Ck 21°47'S 148°5S'E, Britton & Misko

12.12.1968 at light (4) ANIC, MSNV; Ingham

23 mi SSE, Crystal Ck, 18°58'S 146°16'E,

114

Britton & Misko 9.12.1968 at light (2); Laura, C.

Watts 18.7.1982 (2) SAMA; Monto 22 km NW,

Coominglah St. For., J. T. Doyen 23.12.1982 (1)

ANIC; Mornington Island Mission, Aitken &

Tindale 11.5.1963 at light (1) SAMA;

Normanton, Tindale & Aitken 4.5.1963 at light

(1) SAMA; Pentland, J. C. Lesoeuf 12.7.1975

(1) ANIC; Station Ck, J. G. Brooks 19.4.1970

(3) ANIC; Townsville, P. Ferrar 19.11.1967 light

trap (1) ANIC; Tully 40 km W, E. F. Riek

31.5.1971 (3) ANIC, MSNV. South Australia:

Chambers Gorge, C. Watts 9.1983 (5) SAMA;

Enorama Ck, Amena Valley Rd, Nth. Flinders,

Suter Mitchell & Marchant 26.7.1976 (1)

SAMA; Flinders Ra., Arkaba Ck, E. G.

Matthews 5.3.1973 (10) SAMA, MSNV;

Flinders Ra., Blinman 12 km NW, I & II springs,

J. A. Forrest 10.5.1981 (2) SAMA; Flinders Ra.,

Brachina Ck nr. Heysen Hill, J. A. Forrest

9.5.1981 (1) SAMA; Flinders Ra., Eringunda V.,

E. G. Matthews 6.3.1973 (2) SAMA; Flinders

Ra., Parachilna Gorge, C. Watts 10.1981 (1)

ANIC; Flinders Ra., Wooltana HS 6 km SW,

Munyallina Ck, J. A. Forrest 12.5.1981 (1)

SAMA; Manu Ra., Angatja homestead 5 km N,

damp litter, L. E. Watrous 11.5.1983 (3) FMNH;

Mt Remarkable, L. E. Watrous 29.4.1983 black

light (1) FMNH; Musgrave Ra., Ernabella 13 km

N, L. E. Watrous 7.5.1983 (12) FMNH;

Musgrave Ra., Rock Hole, SSE Mt Woodruffe,

L. E. Watrous 8.5.1983 black light (7) FMNH;

Owieandana, N Flinders Ra., Hale & Tindale (2)

SAMA; Parachilna, Flinders Ra., Griffith Coll.,

(1) SAMA; Quorn 30 km NNW, Buckaringa

Gorge, C. ‘Reid 18:12°51985 (2). ANIC;

Woodendina Ck, Nirrana, Flinders, Bishop &

Diener 16.12.1976 (4) SAMA, MSNV. Victoria:

Turtons Track, Otway Ranges, Gross & Aitken

17.1.1962 (1) SAMA. Western Australia:

Beverley, Griffith Coll. (7) SAMA, MSNV;

Cane River HS. 17 km N, 21°56'S 115°39'E,

Upton & Mitchell 27.4.1971 (5) ANIC, MSNV;

Carson Escarpment, 14°49'S 126°49'E, Common

& Upton 9-15.8.1975 (7) ANIC, MSNV;

Dampier, Dampier Is., Intercourse, pool standing

water, J. B. Kethley 2.10.1976 (2) FMNH;

Derby, W. D. Dodd (1) SAMA; Kimberley Bore,

K. & E. Carnaby 13.3.1980 at light (1) ANIC;

Kimberley W, Cape Bertholet 8 km S, 17°19'S

122°10'E, D. H. Colless 19.4.1977 (1) ANIC;

Millstream, 21°35'S 117°04E, E. B. Britton

28.10.1970 at light, open eucalypt paperbark

woodland (2) ANIC; 28.10.1970 at light, open

eucalypt-paperbark woodland kangaroo grazed

grass (1) ANIC; 28.10.1970 Crystal Pool (4)

E. GENTILI

ANIC; 29.10.1970 shallow stream (18) ANIC;

29.10.1970 at light spinifex-eucalypt junction

(22) ANIC; 30.10.1970 eucalypt-spinifex (2)

ANIC; 31.10.1970 (10) ANIC; 1.11.1970 gravel

at margin of Crossing Pool (24) ANIC;

2.11.1970 from gravel at edge of pool at pipe

crossing (82) ANIC, MSNV; 1.11.1970 at light,

eucalypt-paperbark woodland (1) ANIC;

2.11.1970 palm-eucalypt-melaleuca assocn, at

light (6) ANIC; 3.11.1970 shallow weed-grown

pool (2) ANIC; 3.11.1970 at light (28) ANIC;

4.11.1970 Crossing Pool, at light (1) ANIC;

5.11.1970 waterside gravel in pipe, Crossing

Pool (16) ANIC; 5.11.1970 eucalypt-spinifex, at

light (2) ANIC; 5.11.1970 Crystal Pool, at light

(14) ANIC; 7.11.1970 mouth of Dawson’s Ck

(17) ANIC; 8.11.1970 Deep Reach, at light (3)

ANIC; Millstream, Crossing Pool, D. H. Colless

21.10.1970 (2) ANIC; Millstream HS, E. F. Riek

4.4.1971 (5) ANIC; Millstream HS. % km WNW,

21°35'S 117°04'E, M. S. Upton 21.10.1970 (4),

Upton & Mitchell 14.4.1971 (24) ANIC, MSNV;

Millstream 1 km NE, 21°35'S 117°04'E, M. S.

Upton 24.10.1970 (14) ANIC; Millstream 1 km

NNE, M. S. Upton 3-4.4.1971 (14) ANIC;

Millstream 1 km N, M. S. Upton 9-10.4.1971

(3) ANIC; Millstream 2 km ENE, M. S. Upton

21.10.1970 (1) ANIC; Millstream HS. 3 km NW,

21°34'S 117°03'E, Upton & Mitchell 22.4.1971

(7) ANIC; Millstream HS. 5 km SE, 21°37'S

117°06'E, Upton & Mitchell 17.4.1971 (1)

ANIC; Millstream 8 mi ENE, D. H. Colless

20.10.1970 (5) ANIC; Millstream 15 km E,

21°35'S 117°12'E, M. S. Upton 20.10.1970 (8)

ANIC; Minilya R., 23°49'S 114°00'E, Upton &

Mitchell 29.3.1971 (14) ANIC, MSNV; Mitchell

Plateau, Mining Camp, 14°49'S 125°S0'E, Rentz

& Balderson 9-19.5.1983 (1) ANIC; id., 4 km S,

14°52'S 125°S0'E, Rentz & Balderson 13.5.1983

(1) ANIC; Mt Magnet, K. & E. Carnaby 28-

29.9.1978 (1) ANIC; Newman 13 km E, 23°15'S

119°52'E, E. B. Britton 12.11.1970 (8) ANIC;

Walsh Pt 8 km SW, Escarpment, 14°37'S

125°48'E, Rentz & Balderson 10—-17.5.1983 (1)

ANIC; Wittenoom 13 km ESE, 22°18'S

118°27'E, E. B. Britton 11.11.1970 (1) ANIC.

Biology

The label data are scarcer than for pygmaeus,

but similar. The species is collected most

frequently at light (also UV light, black light),

then at edge of standing or flowing water, e.g. in

gravel, and occasionally in damp leaf and log litter

on the forest floor. An exceptional capture is in

and under rotting fruits of Endiandra introsa.

PARACYMUS OF AUSTRALIA 115

3. Paracymus gigas Gentili, 1996

Paracymus gigas Gentili 1996: 178; Hansen

1999: 111.

Types (Fig. 24)

Holotype male: Western Australia: Mitchell

Plateau, Amax Warrender Rd 3 mi W, J. B.

Kethley 21.10.1976, spring pool drainage, FMNH.

Description

Length 3.0 mm, width 1.8 mm. Convex, widely

oval, black. Head 1.5 x as wide as long, black,

densely punctured, interspaces once to twice

width of punctures; anterior margin of labrum

uniformly curved; eye/interocular space ratio

nearly 0.7. Pronotum black with testaceous lateral

margins, sides strongly curved; densely punctured

as head, between punctures slightly shagreened.

Scutellum black, trapezoidal, elongate, slightly

protruding anteriorly. Elytra black, pale near

lateral borders, with normally developed

parasutural furrows, punctures somewhat denser

than on pronotum and head, shagreened between

punctures. Postlabium rectangular, flat and

smooth; antennae 9-jointed; maxillary palpi long

0.6 x width of head; eyes large below. Prosternum

with longitudinal keel; mesosternum with arrow-

head-like keel; metasternum raised in middle, with

conspicuous triangular dimple in raised area;

abdominal ventrites pubescent. Profemora

ventrally pubescent, mesofemora so only on and

near trochanters, metafemora glabrous. Aedeagus

as in Figs 11, 12: tegmen pointed at base, longer

than parameres, ventrally protruding and covering

base of penis with a jut; parameres blunt at apex

and interiorly scythed; penis progressively

constricted from base to apex, this slightly

expanded as a button.

Discussion

Larger than all described Australian Paracymus,

this species is immediately identified by its long

maxillary palps, the 9-segmented antennae, and

the metasternal dimple. The length of the palps

and the shape of the aedeagus (e.g. tegmen with a

FIGURE 24. Distribution of the studied specimens of Paracymus wattsi n. sp. (Ad), cariceti n. sp. (MB), gigas

Gentili, 1996 (@).

116

ventral jut covering the base of the penis) might

suggest that it belongs to the genus Notohydrus;

but the longitudinally keeled prosternum suggests

the genus Paracymus.

4. Paracymus cariceti n. sp.

Types (Fig. 24)

Holotype male (1.7 x 1.1 mm): Tasmania:

42°32'S 146°30'E, Map 8212.593.923, Adj.

Misery Creek 780 m, 21.2.1980, L. Hill coll.,

short sedge and rush turfs, ANIC, CSIRO.

Paratype: Tasmania: male (without aedeagus,

previously lost): Tasmania, Launceston, SAMA.

Another specimen in SAMA, a female, labelled

‘Launceston, Tas., Jan., C. Watts’, might belong

to this species, but apart from the slightly greater

size it has the first abdominal segment with a

longitudinal keel, lacking in the two typical males.

Description

Length 1.7-1.8 mm; width 1.0-1.1 mm. Widely

oval, slightly convex. Dorsally black, head with

metallic reflections, pronotal borders and elytral

apex darkly testaceous. Labrum centrally hollow;

anterior margin of clypeus straight; head, under

100 x magnification, slightly shagreened, with

scarcely visible punctures; interocular space 3.6 x

eye width. Pronotum smooth, with sparse

punctures clearly visible at 100 x magnification,

space between them nearly as large as punctures.

Scutellum triangular, equilateral, smooth, with

scarce and fine punctures. Elytra smooth, with a

more or less faint shagreen at 100 x, with sparse

punctures as on pronotum; parasutural furrow in

apical 2/3 or a bit less. Underside testaceous, head

black. Postlabium nearly rectangular, flat, smooth

and shining; gula smooth. Prosternum carinate

with a high longitudinal keel; mesosternum

anteriorly provided with transverse tooth, and

posteriorly with longitudinal keel; metasternum

concealed under recumbent hairs. Five visible

abdominal ventrites, glabrous and smooth, the

first being the shorter, the last the longer; first

ventrite without a longitudinal keel but with

scarcely visible tubercle along central line; if

specimen from Launceston previously recorded

belongs to this species, female has a keel on first

ventrite. Palps short and stout; profemora

pubescent along anterior margin; mesofemora

pubescent to knees; metafemora with scarce hairs

near trochanters. Aedeagal length (Fig. 9) between

1/3 and 1/4 of body; tegmen narrow at its base,

much shorter than parameres; these acute and

E. GENTILI

longer than penis; penis basally swelling, then

rod-like, anchor-shaped at tip, without any collar.

Discussion

This Tasmanian species differs from other

Australian Paracymus chiefly by its aedeagus,

which has the tip of the penis anchor-shaped at

the apex, and the apex of the parameres acute.

Other unique characters are the mesofemur which

is completely pubescent on its ventral face, and

the first ventrite having a tubercle instead of a

longitudinal keel. Size and colour are similar to

small P. pygmaeus; but it differs from this species

in the aedeagus, which has a shorter tegmen, and

the features mentioned; in its shorter and wider

body shape; in its shorter and stouter palps; and in

having the first abdominal segment not keeled.

Biology

The name cariceti comes from a label note:

short sedge and rush turfs.

5. Paracymus ovum n. sp.

Types (Fig. 25)

Holotype male (2.8 x 1.45 mm): Kalbarri Nat.

Pk, Northampton 54 mi N, Common & Upton

19.4.1968, ANIC; paratypes (3): same data as the

holotype, ANIC, SAMA, MSNV.

Description

Length 2.5—2.8 mm; width 1.35-1.45 mm;

elongate oval, slightly convex (Figs 18, 19). Head

black, irregularly punctured, interspaces

alutaceous or smooth, as large or larger than

punctures, these less strong and deep than in

weiri,; labrum nearly concealed under head; eyes

together as wide as nearly 4/7 of interocular space;

Y-suture conspicuous only as periocular groove.

Pronotum black with thin testaceous lateral

border; punctures irregularly distributed as on

head, a little more faint, interspaces alutaceous

(chiefly on borders) or smooth (chiefly on disc);

anterior corners clearly produced. Scutellum

triangular, longer than wide. Elytra black with thin

testaceous border; punctured as on pronotum,

partly alutaceous; parasutural furrow in apical 2/3

or a bit less. Underside dark; postlabium smooth

with small punctures; prosternum roof-like,

keeled; mesosternal keel anteriorly like an arrow-

head; metasternum posteriorly with glabrous area

surrounded by tufts of hairs; first abdominal

ventrite centrally covered with flat longitudinal

keel, enlarging anteriorly. Palps yellow, second

PARACYMUS OF AUSTRALIA

joint enlarged, last joint more slender than in

weiri; antennae 8-segmented, yellow with a dark

club. Profemora widely hairy, mesofemora only

on basal third, metafemora glabrous. Aedeagus

(Fig. 17) concave at base, tegmen nearly as long

as parameres, these blunt at apex; penis rod-like

with small subapical expansion or collar.

Discussion

Only the shape of the aedeagus differentiates

ovum from all other Australian Paracymus; other

differences are combinations of characters. It is

similar to weiri, but differs in having a more

slender, nearly parallel-sided body (Figs 18, 19);

an upperside partly alutaceous; a flat keel on the

entire first ventrite; a different aedeagus: tegmen

shorter (in weiri it is longer than parameres, in

ovum subequal), and penis less stout.

6. Paracymus wattsi n. sp.

Paracymus phalacroides Wooldridge 1978: 129;

Hansen 1999: 112.

Types (Fig. 24)

Holotype male (1.7 x 1.0 mm): New South

Wales: Barrington, NSW, 17.8.1997 C. Watts,

SAMA; aedeagus preserved in DMHF. Paratypes:

New South Wales, same data (2 females) SAMA;

Batemans Bay 2 km N, C. Watts 18.4.1997 (5

males 3 females) SAMA, MSNV; Blue

Mountains, Faulcon Bridge 500 m, G. Wewalka

15.1.1993 (lmale 2 females) NMW, MSNV;

Cabbage Tree Creek, Nelligan 20 km W, C. Watts

30.11.1995 (Imale) SAMA; Dorrigo, W. Heron

(1 female) ANIC [Paracymus phalacroides? det.

D. P. Wooldridge]; Failford 8 km N, C. Watts

18.8.1997 (2 males 1 female) SAMA, MSNV;

Valery, A. P. M. Eucalyptus plantation,

McMullen’s Block, 30°24'S 152°57'E, light trap,

R. S. McInnes 10.1.1967 (4) ANIC, MSNV;

Windsor, A. M. Lea 1871, Griffith Coll. (17)

SAMA, MSNV. Northern Territory: Jabiru 10

km SW, C. Watts 22.3.1998 (1) SAMA; Gubara 1

km W, Kakadu NP, C. Watts 17.3.1998 (1 male)

SAMA. Western Australia: Fremantle 10 km S,

C. Watts 24.10.1996 (1) SAMA.

Description

Length 1.6-1.9 mm; width 1.0-1.1 mm. Body

convex, short oval (Figs 20, 21), entirely black,

only tips of palps and legs reddish. Head shining,

smooth, only occipital region alutaceous, evenly

punctured, intervals between punctures large,

117

nearly twice as large as the punctures. In frontal

view one eye measures nearly 1/6 of interocular

space. Pronotum black, shining, smooth, evenly

punctured as on head; at 100 x bottoms of

punctures are umbilicate or jutting. Anterior

margin of pronotum protruding in centre and

laterally; lateral margin arched so that posteriorly

pronotum enlarges. Scutellum triangular, slightly

elongate, with curved sides, faintly punctured.

Elytra short, their maximum width at anterior

third; punctures denser than on pronotum, at 100

x umbilicate or rugose, evenly distributed; surface

slightly alutaceous between punctures; parasutural

furrow on apical 2/3 or a bit less. Underside black.

Labrum arched, slightly hollow at centre;

postlabium flat or convex, alutaceous at 100 x

with scarce and faint punctures. Gula alutaceous,

with two furrows and two deep punctures.

Prosternum tectiform, keeled; mesosternum with

longitudinal keel anteriorly expanded;

metasternum punctured and pubescent, elevated in

centre, slightly hollow and posteriorly produced.

Five visible abdominal sternites, micropunctured,

not keeled; pygidium with a dozen stiff and short

setae directed posteriorly and centrally. Profemora

excavated, granulose and hairy on basal 2/3;

mesofemora hairy on basal 3/4; metafemora

smooth, with sparse hairs. Tibiae shorter than

femora, spiny at their margins; tarsi not expanded

in male, claws hooked. Aedeagus (Fig. 15) nearly

1/4 as long as body; tegmen shorter than

parameres, narrow and pointed at base; parameres

slender, with rounded apices; penis conical,

pointed, without any collar.

Discussion

Characters of the species distinguishing it from

all Australian Paracymus are the umbilicate

punctures, the copious hydrofugal hairs at the base

of the mesofemora (only cariceti has more), the

lack of any keel on the first ventrite (as in cariceti,

which however has a tubercle instead of a keel),

and the aedeagus (penis triangular to the apex,

tegmen abruptly pointed). From pygmaeus it is

also separated by the small size, the short and

convex body (length:width 1.7 in pygmaeus; 1.6

in wattsi). With opacus, australiae and weiri it

constitutes a group of short and convex

Paracymus (Figs 20, 21). Wooldridge 1978

records two females of this species (SAMA; now

1 in ANIC) as possibly belonging to the

palaearctic P. phalacroides (Wollaston, 1867).

But the punctures of phalacroides are geminate,

not umbilicate as in wattsi; the mesosternal keel

of phalacroides is very low, in wattsi normally

118 E. GENTILI

shaped; and the bases of the parameres are

internally excavated in phalacroides, swollen in

wattsi.

Biology

One label has the note: light trap. C. Watts

captured his specimens ‘in thick grassy vegetation

at edge of water in large swampy areas’ (personal

communication).

7. Paracymus opacus n. sp.

Types (Fig. 25)

Holotype male (1.7 x 1.0 mm): Queensland,

Bentinck Is. Minakuri, Aitken & Tindale

23.5.1960 at light, SAMA. Paratypes:

Queensland: Bentinck Is. Minakuri, Aitken &

Tindale 23.5.1960 at light (4) SAMA, MSNV;

Paluma Dam Road, J. G. Brooks 13.1.1968 (1)

ANIC; Seaforth 1 km NW, 20°53'S 148°57'E, A.

Gillison 18.11.1981 Berlesate, Melaleuca

woodland (1) ANIC; Stewart R., W. D. Dodd (3)

SAMA, MSNV; Townsville 20 km N, Bushland

Beach, A. J. Watts 6—-11.2.1998 (2) SAMA.

Description

Length 1.6—2.0 mm; width 0.95—1.2 mm. Body

convex, short, oval (Figs 20, 21), entirely black.

Upper body entirely alutaceous, satiny,

impunctate. Head grey-black, satiny like pronotum

and elytra, branches of Y-suture conspicuous.

Pronotum enlarged posteriorly. Scutellum

triangular, impunctate. Elytra with parasutural

furrow in apical 2/3 or a bit less. Underside black

or dark brown; postlabium flat, alutaceous,

impunctate; prosternum short, roof-like, with a

longitudinal keel; mesosternal keel short, ending

anteriorly close to a high transverse ridge;

metasternum hairy, elevated in middle as a flat

triangle posteriorly acute. First abdominal ventrite

with a median keel, rising from a central process

protruding anteriorly. Antennae 8-segmented;

basal 2/3 of profemur densely pubescent, protibiae

and protarsi short and thick, last tarsal segment

with a small denticle in males; claws short and

hooked. Aedeagus (Fig. 16) with a very long

tegmen, penis rod-like, anchor-shaped at tip,

parameres provided with a triangular membrane,

progressively expanded from base to apex.

Discussion

Among Australian Paracymus, opacus differs

in having a strongly alutaceous and impunctate

upperside, a flat hairy triangular prominence on

the metasternal surface, and a very unusual

aedeagus: tegmen more than twice as long as the

length of the parameres, which are expanded

towards the penis by a membrane. It is very near

P. wattsi in body form and size, but is easily

separated by the preceding characters.

Biology

Four specimens captured at light, one from

forest floor litter, with Berlese funnel.

8. Paracymus australiae n.sp.

Types (Fig. 25)

Holotype male (2.3 x 1.4 mm): Northern

Territory: Oenpelli 6 km SW by S, 12°22'S

133°O1'E, E. G. Matthews 30.5.1973 at light,

ANIC. Paratypes: Northern Territory: Baroalba

Spring, 12°47'S 132°51'E, 20.11.1972 (1) ANIC;

Borroloola 46 km SSW, 16°28'S 136°09'E, J. E.

Feehan 23.4.1976 (6) ANIC, SAMA, MSNV;

Borroloola 48 km SW by S, McArthur R., 16°27'S

136°OS'E, J. E. Feehan 13.4.1976 at light (1)

ANIC; Cahills Crossing 1 km S, E Alligator R.,

12°26'S 132°58'E, E. B. Britton 3.11.1972 at light

(1) ANIC; Cape Crawford 8 km ESE, Bessie

Spring, 16°40'S 135°S1'E, M. S. Upton

26.10.1975 (1) ANIC; Mt Cahill 8 km E,

Nourlangie Ck, 12°52'S 132°47'E, E. B. Britton

27.10.1972 at light (1) ANIC; Mt Cahill 15 km E

by N, 12°50'S 132°51'E, E. B. Britton 29.10.1972

at light (4) ANIC, SAMA, MSNV; Mt Cahill 15

km E Koongarra, 12°52'S 132°50'E, M. S. Upton

15.11.1972 (5), 6-10.3.1973 (3) ANIC, SAMA,

MNSV; Mudginbarry HS 9 km N by E, 12°31'S

132°54'E, Upton & Feehan 10.6.1973 (3) ANIC,

SAMA, MSNV; Oenpelli 6 km SW by S, 12°22'S

133°01'E, Upton & Feehan 6.6.1973 (1) ANIC.

Description

Length 2.0-2.5 mm; width 1.35—1.5 mm; short,

oval, slightly convex. Upper side black with dark

testaceous contour zone; head, pronotum and elytra

coarsely punctate, space between punctures nearly

as large as punctures; each puncture larger than

those of P. pygmaeus. Head entirely black,

anteriorly slightly hollow in centre to receive

labium; sutures not conspicuous; eyes large,

together nearly 6/10 width of interocular space.

Pronotum anteriorly slightly curved, with notably

prominent corners, much larger at base (ratio hind

width : fore width 1.65); black with brown sides,

evenly punctured as on elytra. Elytra, observed

from above, nearly as long as wide (ratio elytral

PARACYMUS OF AUSTRALIA

FIGURE 25. Distribution of the studied specimens of Paracymus opacus n. sp. (A), australiae n. sp. (MB), weiri n.

sp. (@), and ovum n. sp. ().

length : elytral width 1.07); parasutural furrow in

apical 3/4 or a bit less; black with brown sides and

apex. Underside dark, brown to reddish; labrum of

male with two small specula, postlabium flat, slight

and shining; prosternum roof-like, with

longitudinal keel; a fine mesosternal keel anteriorly

reaches a robust crescent-shaped ridge;

metasternum hairy, two reliefs provided with tufts

of setae delimiting a central glabrous area; first

abdominal ventrite keeled on anterior 1/2—1/3.

Palps yellow, antennae 8-segmented; profemora

hairy on basal 3/4, male protarsi with a robust seta

or tooth; mesofemora with hydrofuge hairs on a

large postero-basal area; metafemora glabrous.

Aedeagus (Fig. 10) long, nearly 3/10 of body

length, tegmen constricted near base, bluntly

pointed, longer than parameres, these blunt at

apices, penis nearly triangular, gradually narrower

from base to apex, without any collar.

Discussion

Among the Australian Paracymus, some

features are exclusive to australiae: the great

length of the parasutural furrow, the coarse

dorsal punctation with large punctures, the male

specula, the two hairy metasternal reliefs, and

the first ventrite, which is keeled only anteriorly.

This species is distinguishable from pygmaeus

also by the stocky body shape, and by the

aedeagus being more slender, straightened

basally, with penis gradually narrowing, not

suddenly restricted in the apical zone,

unprovided with a collar.

Biology

Some specimens were captured at light, others

in standing or flowing water.

9. Paracymus weiri n. sp.

Types (Fig. 25)

Holotype male (2.9 x 1.7 mm): Western

Australia: Carson Escarpment, 14°49'S 126°49'E,

Common & Upton 9-15.8.1975 ANIC. Paratypes:

Western Australia: Carson Escarpment, 14°49'S

120

126°49'E, Common & Upton 9-15.8.1975 (9)

ANIC, SAMA, MSNV.

Description

Length 2.3-3.0 mm, width 1.5—1.8 mm; short

oval, convex. Head black, evenly punctured,

interspaces smooth, nearly as large as punctures;

anteriorly a central hollow receives labrum; eyes

slightly protuberant, their width together nearly 2/

3 of interocular space; Y-suture reduced to an

incomplete periocular groove. Pronotum and

elytra black with lateral margins and apex

testaceous-reddish, punctured as on head or more

deeply; elytra with parasutural furrow in apical 2/

3 or a bit less; scutellum triangular, slightly

elongate, punctured. Underside dark. Postlabium

flat and smooth; prosternum evidently keeled;

mesosternum with a longitudinal keel ending

anteriorly as an arrow-head; metasternum glabrous

in centre, the glabrous area delimited by two hairy

lines converging posteriorly. First abdominal

ventrite with a high keel almost reaching posterior

margin. Palps and antennae (8-segmented) yellow

with dark tips; legs dark; profemur pubescent

nearly to the knees, last segment of male protarsi

with two little teeth; mesofemora pubescent for

nearly 2/3 of posterior margin. Aedeagus as in

Figs 13, 14: tegmen concave at base, longer than

parameres; these blunt at apices; penis conical,

stout, provided with a preapical expansion or

collar.

Discussion

Considering the body shape, P. weiri belongs to

the group containing wattsi, opacus, and

australiae (Figs 20, 21); the outline of the

aedeagus recalls P. ovum. The body shape and

upper punctation are similar to P. australiae, but

weiri is larger in size and not provided with male

specula; it has the first abdominal ventrite with a

high keel and a very different aedeagus: base of

tegmen concave (in australiae bluntly pointed),

penis abruptly expanded before the apex (in

australiae uniformly conical).

A Key To THE AUSTRALIAN PARACYMUS

1. — Elytralcolour mahogany ortestaceous brown;

pronotum black. Aedeagus as in Figs 6-8:

tegmen pointed at base, penis witha simple

preapical collar, external border of

parameres evenly curved. Elongate oval

(Figs 18, 19). Length 1.5—2.3 mm. .........

Sssesdolissadevevsessseoessiussey spenceri Blackburn

E. GENTILI

— Pronotum and elytra black. Aedeagus with

different combination of characters: base

of tegmen concave, or penis without a

simple preapical collar, or parameres

externally straight ss:.cc<cccsesscessoeasecseeesees 2

2. — Male protarsus swollen, last segment

evidently thicker than in female, proclaws

spatulate, inner one stronger than outer.

Aedeagus as in Figs 1—5: base of tegmen

pointed, penis with a preapical collar

originating a basal membrane, parameres

externally straight. Elongate oval (Figs

18; 19): Length 1.6—3.0 mm «.Jescccscsse

Helper teens pygmaeus (McLeay)

— Male protarsus not or only a bit swollen.

Aedeagus without a preapical collar

originating a membrane ....................+ 3

Upper side impunctate and strongly

alutaceous, satiny. Aedeagus as in Fig. 16:

a very long tegmen (more than 2 x the

length of parameres), penis rod-like, anchor

shaped at apex, parameres internally

originating as a membrane. Short oval,

convex (Figs 20, 21). Length 1.6—2.0 mm

Sofogre Pease saa rat estes cutee certs opacus N. sp.

— Upperside punctured, smooth or moderately

alutaceous. Tegmen shorter, parameres

without any membrane .................00.. 4

Dorsal punctures (magnification 100 x)

umbilicate or with ajutin the bottom. First

abdominal ventrite not keeled nor

tuberculate. Short, widely oval, convex

(Figs 20, 21). Aedeagus as in Fig. 15:

tegmen sharply pointed, penis conical,

without any collar. Length 1.6-1.9 mm.

sogsnnsveceperedswateth ecerseseteraveessects wattsi n. sp.

— Dorsal punctures simple, not umbilicate.

First abdominal ventrite keeled or

tuberculate. Tegmen never sharply pointed

Antennae 9-segmented. Maxillary palps

long, 0.6 x width of head. Dorsal surface

slightly shagreened. Metasternum with a

conspicuous triangular dimple. Aedeagus

as in Figs 11, 12: tegmen provided with a

central jut covering the base of the penis.

Length 3.0 mmics.c.00.3500.+ gigas Gentili

— Antennae 8-segmented. Maxillary palps

shorter than 0.5 x the width of head.

Metasternum without any dimple. Tegmen

simple, without any jut anteriorly

PLOMUGIN Serester..srcctssets-ccraocnconrscentesrantees 6

PARACYMUS OF AUSTRALIA 121

Parasutural furrow covering nearly 3/4 of

elytral length. Upper side coarsely

punctured with large punctures. Male

labrum with small specula. Only anterior

1/2-1/3 of the first ventrite keeled.

Aedeagus as in Fig. 10: base of tegmen

bluntly pointed, penis simply conical.

Length 2.0-2.5 mm ..... australiae n. sp.

— Parasutural furrow covering nearly 2/3 of

elytral length. Labrum without specula.

First ventrite not or differentlykeeled. Apex

of penis anchor-shaped ..............0.::006 ui

Whole ventral surface of mesofemora

pubescent. First ventrite tuberculate along

the median line. Aedeagus bluntly pointed

at base, penis with apical anchor, apices of

parameres acute (Fig. 9). Length 1.7—1.8

TIM os sstsat de cesscsvccsevsvvuesseens: cariceti n. sp.

— Atleast apical 1/3 of mesofemora glabrous.

First ventrite keeled. Aedeagus concave at

base, penis with preapical expansion,

apices of parameres blunt ............0.2. 8

First ventrite completely carinate with a flat

longitudinal keel. Dorsal punctures more

feeble. Elongate oval (Figs 18, 19).

Aedeagus as in Fig. 17: tegmen nearly as

long as parameres, penis more thin, rod-

like, with feeble preapical expansions.

Length 2.5=2.8) Mim scence ovum N. sp.

— First ventrite with a high keel not covering

whole segment. Dorsal punctures stronger.

Short oval (Figs 20, 21). Aedeagus as in

Figs 13, 14: tegmen longer than parameres,

penis more stout, subconical, with strong

preapical expansions. Length 2.3-3.0 mm

Rene peri ceneersscet eee ee weiri N. sp.

ACKNOWLEDGMENTS

I wish to thank Manuela Caccia (Milano) for the

drawings, Tom Weir of the CSIRO (Canberra) for the

maps, Chris Watts of the South Australian Museum

(Adelaide) for information, useful suggestions and

communication of rare specimens, R. G. Ordish

(Wellington), E. G. Matthews (Adelaide), F. Hebauer

(Grafling) for advice, and the following persons for

their kindness in procuring types or other material: E.

G. Matthews (SAMA), T. Weir (ANIC), R. Brett and D.

Kavanaugh (CASF), A. Newton and M. Thayer

(FMNH), K. Desender (ISNB), S. J. Hine (NHML), M.

Jach (NMW), P. Spangler (USNM).

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WOLLASTON, T. V. 1867. Coleoptera Hesperidum,

being an enumeration of the Coleopterous insects of

the Cape Verde Archipelago. J. V. Voorst: London,

pp. 1-285.

ZAITZEV, F. A. 1908. Catalogue des Coléoptéres

aquatiques des familles Dryopidae, Georyssidae,

Cyathoceridae, Heteroceridae et Hydrophilidae.

Horae Societatis Entomologicae Rossicae 38: 283-

420.

A NEW SPECIES OF THE GENUS LESTIGNATHUS ERICHSON FROM

TASMANIA WITH A NOTE ON THE TASMANIAN SPECIES OF

MECYCLOTHORAX SHARP (INSECTA: COLEOPTERA: CARABIDAE:

LICININAE, PSYDRINAE).

BY MARTIN BAEHR

Summary

BAEHR, M. (2000). Lestignathus pieperi, sp. nov. is described from Mt Field, southwestern

Tasmania. It is distinguished inter alia from the three known species of the genus Lestignathus by

presence of only one setiferous puncture on the 3" interval. A key to all species of the genus is

added.

A NEW SPECIES OF THE GENUS LESTIGNATHUS ERICHSON FROM TASMANIA

WITH A NOTE ON THE TASMANIAN SPECIES OF MECYCLOTHORAX SHARP

(INSECTA: COLEOPTERA: CARABIDAE: LICININAE, PSYDRINAE)

MARTIN BAEHR

BAEHR, M. 2000. A new species of the genus Lestignathus Erichson from Tasmania, with a

note on the Tasmanian species of Mecyclothorax Sharp (Insecta: Coleoptera: Carabidae:

Licininae, Psydrinae). Records of the South Australian Museum 33(2): 123-126.

Lestignathus pieperi, sp. nov. is described from Mt Field, southwestern Tasmania. It is

distinguished inter alia from the three known species of the genus Lestignathus by presence of

only one setiferous puncture on the 3" interval. A key to all species of the genus is added.

Recent collections revealed that in Tasmania two species of Mecyclothorax occur, namely

the well-recorded M. ambiguus (Erichson), and M. punctipennis (Macleay) that is widespread

on the mainland but not previously recorded from Tasmania.

Martin Baehr, Zoologische Staatssammlung, Miinchhausenstr. 21, D-81247 Miinchen,

Germany. Manuscript received 16 May 2000.

INTRODUCTION

Through the kindness of Dr Harald Pieper

(Kiel) I received a sample of Tasmanian Carabidae

for identification that he collected mainly in

western Tasmania during a trip in December 1998.

This sample included, inter alia, a new species of

the licinine genus Lestignathus Erichson, as well

as the first Tasmanian record of the psydrine

species Mecyclothorax punctipennis (Macleay).

By courtesy of Dr Eric Matthews, Adelaide, I had

the opportunity to compare types or material of all

described species of Lestignathus stored in the

collection of the South Australian Museum.

So far, the genus Lestignathus Erichson

includes three rather different species (Sloane

1920, Moore et al. 1987) that all occur in

Tasmania, namely the common and widespread,

large Lestignathus cursor Erichson, and the

apparently much rarer, smaller species L. foveatus

Sloane and L. simsoni Bates. The new species

differs from all described species in several

respects, hence description of the single known

specimen seems advisable, the more so as all

species of Lestignathus, except for L. cursor,

apparently are rare or very rare, and additional

material is unlikely to be detected soon.

MEASUREMENTS

Measurements have been made under a stereo

microscope by use of an ocular micrometer.

Length has been measured from apex of labrum to

apex of elytra. Length of pronotum was taken

from the most advanced tip of anterior angles to

the most advanced part of base. Width of base

was taken at position of the posterior marginal

setae. Measurements, therefore, may slightly differ

from those of Sloane (1920).

LocaTION OF TYPES

To facilitate further study of the genus

Lestignathus, the holotype of L. pieperi sp. nov. is

presented to the South Australian Museum.

Therefore, the types of three of the four recorded

species of the genus Lestignathus are assembled

in that collection which also holds historical

material of the fourth species.

Lestignathus pieperi sp. nov.

(Figs 1-2)

Holotype: f, TAS, Mt Field Lyrebird Nat. Walk

30.11.1998 leg. H. Pieper (SAMA).

Diagnosis

Distinguished from all other species of the

genus Lestignathus by presence of only 1

setiferous puncture on 3" elytral interval. Further

distinguished from L. cursor Erichson by much

lesser size; from both, L. foveatus Sloane and L.

simsoni Bates, by less cordate, anteriorly much

124

FIGURE 1. Lestignathus pieperi sp. nov. Habitus.

Length: 7.55 mm.

Narrower pronotum, almost completely depressed

elytral intervals, and narrower head which is less

than half as wide as pronotum; and from latter

species also by slightly lesser size.

Description

Measurements. Length: 7.55 mm; width: 3.0

mm. Ratios: width/length of pronotum: 1.06;

width base/apex of pronotum: 1.37; width

pronotum/head: 2.08; length/width of elytra: 1.59;

width elytra/pronotum: 1.52.

M. BAEHR

Colour. Piceous-black, pronotum and elytra

with narrow reddish margins; labrum and

mouthparts red, clypeus reddish-piceous, basal

and apical antennomeres reddish, median

antennomeres piceous, femora and tibiae in

middle reddish-piceous, basally and apically

lighter, tarsi light reddish. Lower surface black.

Head. Very small in comparison to prothorax.

Eyes large though laterally little produced, with

small orbits. Clypeus bisetose, anterior central part

of clypeus membranous. Labrum medially deeply,

symmetrically v-shaped, excised for about a third

of its length, quadrisetose. Mentum with an

indistinct, slightly triangular tooth, ligula bisetose,

glossa and paraglossae of about equal length.

Lacinia with elongate tooth at end, median border

with dense fringe of stiff setae. Palpi rather

slender and elongate, apical palpomeres

thickened, both terminal palpomeres extremely

sparsely pilose. Both mandibles bidentate, though

lower tooth of right mandible more acute than that

of left mandible, therefore apical excision in right

mandible about quadrate, in left mandible more

semicircular. Clypeofrontal suture very shallow,

straight. Frons convex, near clypeal suture with

shallow, rather irregularly shaped impression on

either side. Both supraorbital pores very large.

Frons impunctate, with distinct, isodiametric

microreticulation. Antenna slender and elongate,

surpassing anterior third of elytra, median

antennomeres > 4 x as long as wide, two basal

antennomeres glabrous, 3 antennomere sparsely

pilose, the following antennomeres densely pilose.

Prothorax. Slightly wider than long, laterally

fairly convex, more than twice as wide as head,

widest slightly in front of middle. Apex in middle

deeply excised, anterior angles prominent, at apex

rounded off. Lateral margin in basal half almost

straight. Base distinctly concave in middle, basal

angles widely rounded off. Apex and lateral

borders with narrow though distinct margin, base

in middle not margined. Median line distinct

though shallow, almost complete. Anterior and

posterior transverse impressions barely indicated.

Basal impressions large, wide, with an elongate,

linear impression in median part. Disk rather

depressed, near basal angles widely explanate,

rather even. Disk without any wrinkles or

punctures, with distinct, more or less isodiametric

microreticulation. Anterior marginal seta situated

slightly behind anterior third, well in front of

widest diameter, slightly removed from margin.

Posterior marginal seta situated a short distance in

front of basal angle, close to margin.

Elytra. Elongate-ovalish, widest about at

LESTIGNATHUS AND MECYCLOTHORAX

FIGURE 2. Lestignathus pieperi sp. nov. Female stylomere

2 and base of stylomere 1. Scale line: 0.25 mm.

middle, surface moderately convex, considerably

wider than prothorax. Humeri barely projecting,

basal and lateral margins meeting without any

angle. Lateral margin evenly rounded to apex,

slightly incurved at the very tip. Striation

complete, striae well impressed, very faintly

crenulate, intervals gently convex. Scutellar stria

elongate, scutellar pore present. 3 interval with a

setiferous puncture in about middle, puncture

attached to 2™ stria. Marginal series consisting of

13-14 large punctures that are more widely

spaced in middle. Two additional punctures

situated preapically and apically at 7" stria.

Intervals impunctate, with highly superficial,

transverse microreticulation, rather glossy. Wings

reduced.

Lower surface. Impunctate. Metepisternum

about as long as wide. Terminal abdominal

sternite in female with 4 setae on either side.

Legs. Slender and elongate. Structure of male

anterior tarsus unknown. Metatibia almost

straight. Metatarsus very slender. 5" tarsomeres of

all legs slender, lower surface setulose.

Male genitalia. Unknown.

Female genitalia (Fig. 2). Both stylomeres

markedly depressed, foliaceous. Stylomere 2 short

and wide, triangular, with short, acute apex,

laterally with 2 very short latero-ventral ensiform

setae, mediodorsally with a moderately short

dorso-medial ensiform seta, on median rim near

apex with a nematiform seta originating in a large

groove. In middle of the groove with a small

tubercle. Apex of stylomere 1 without any setae.

Lateral plate conspicuously triangular at apex,

with a 2-3 elongate and some very short

nematiform setae at apical rim.

125

Variation. Unknown.

Distribution. Mt Field, southwestern Tasmania.

Known only from type locality.

Collecting circumstances. Collected on the

ground in temperate rain forest. The holotype was

captured together with Lestignathus cursor

Erichson.

Etymology. The name is a patronym in honour

of the collector Dr Harald Pieper.

Relationships

The sparsely setulose 3 antennomere and the

presence of both marginal pores on the pronotum

place this species in the genus Lestignathus

Erichson, although externally it resembles species

of the related genus Lacordairia Castelnau.

Because the male genitalia of this species are

not yet known and two of the three other species

of Lestignathus are apparently very rare, nothing

can be said about relationships of this species,

which is unique within the genus by the presence

of a single elytral puncture only. Certainly, all

species of the genus differ remarkably in certain

external characters.

Key To THE Species OF LESTIGNATHUS ERICHSON

Size larger, body length >12.5 mm; elytra

with 2 non-foveate punctures, at apex not

deeply sinuate .............. cursor Erichson

— Size smaller, body length <10 mm; elytra

either with foveate punctures, or with 1

puncture only, or at apex deeply sinuate

Each elytron with 3-4 foveate punctures ..

BB otters choca Perea foveatus Sloane

— Each elytron with at most 2 non-foveate

PUMCUULOS <5 eis epes Seesrscests case ccosceesacereorsas 3

Size larger, body length>9 mm; eachelytron

with 2 setiferous punctures, apex of elytra

deeply sinuate ................ simsoni Bates

— Size smaller, body length c. 7.5 mm; each

elytron with 1 setiferous puncture, apex of

elytra barely sinuate ...... pieperi sp. nov.

Mecyclothorax punctipennis (Macleay)

Moore 1984: 162; Moore et al. 1987: 149.

The material collected recently by H. Pieper in

Tasmania includes a male specimen of

Mecyclothorax punctipennis (Macleay) besides

126 M. BAEHR

specimens of the well known M. ambiguus _ species which is common and widespread on the

(Erichson). The identity of M. punctipennis has mainland (Moore et al.1987).

been confirmed by dissection of the male genitalia

and by comparsion with genitalia of the

Tasmanian M. ambiguus, moreover by comparison ACKNOWLEDGMENTS

with the figures in Moore (1984). Data for this My sincere thanks are due to Dr H. Pieper (Kiel) for

specimen are: TAS, Rocky Cape NP, 22.11.1998, kindly submitting his interesting material for

leg. H. Pieper. identification, and to Dr E. Matthews (Adelaide) for the

This is the first Tasmanian record of this _ kind loan of types and material for comparison.

REFERENCES

MOORE, B. P. 1984. Taxonomic notes on some Rhysodidae and Carabidae. In: Zoological Catalogue

Australian Mecyclothorax Sharp (Coleoptera: of Australia, 4: 17-320. Australian Government

Carabidae: Psydrinae) and description of new Publishing Service: Canberra.

species. Journal of the Australian Entomological SLOANE, T. G. 1920. The Carabidae of Tasmania.

Society 23: 161-166. Proceedings of the Linnean Society of New South

MOORE, B. P., WEIR, T. A. & J. E. PYKE. 1987. Wales 45: 113-178.

SIX NEW SPECIES OF NIRRIDESSUS WATTS AND HUMPHREYS AND

TJIRTUDESSUS WATTS AND HUMPHREYS

(COLEOPTERA:DYTISCIDAE) FROM UNDERGROUND WATERS IN

AUSTRALIA.

BY C.H.S. WATTS AND W.F. HUMPHREYS

Summary

WATTS, C.H.S. AND HUMPHREYS, W.F. (2000). Six new species (Nirridessus bigbellensis, N.

cueensis, N. hinkleri, N. morgani, Tjirtudessus hahni and T. magnificus) of stygobiontic beetles of

the family Dytiscidae, subfamily Hydroporinae, tribe Bidessini, from relatively shallow, calcrete

aquifiers in Western Australia, are described and figured, The species are members of a diverse ,

recently discovered, relictual stygofauna, predominantly of Crustacea and Oligochaeta, inhabiting

calcretes lying along palaeodrainage channels. The two genera occur in palaeochannel deposits on

either side of the divide beteween the inland and Indian Ocean drainages. Each calcrete area

contains a distinct assemblage of beetles and the fauna occurs in both fresh and saline groundwater.

The physicochemical properties of the groundwater, and the palaeogeography and hydrology of the

region are discussed in some detail.

SIX NEW SPECIES OF NIRRIDESSUS WATTS AND HUMPHREYS AND

TJIRTUDESSUS WATTS AND HUMPHREYS (COLEOPTERA: DYTISCIDAE)

FROM UNDERGROUND WATERS IN AUSTRALIA

C. H. S. WATTS AND W. F. HUMPHREYS

WATTS, C. H. S. and HUMPHREYS, W. F. 2000. Six new species of Nirridessus Watts and

Humphreys and Tjirtudessus Watts and Humphreys (Coleoptera: Dytiscidae) from underground

waters in Australia. Records of the South Australian Museum 33(2): 127-144.

Six new species (Nirridessus bigbellensis, N. cueensis, N. hinkleri, N. morgani, Tjirtudessus

hahni and T. magnificus) of stygobiontic beetles of the family Dytiscidae, subfamily

Hydroporinae, tribe Bidessini, from relatively shallow, calcrete aquifers in Western Australia,

are described and figured. The species are members of a diverse, recently discovered, relictual

stygofauna, predominantly of Crustacea and Oligochaeta, inhabiting calcretes lying along

palaeodrainage channels. The two genera occur in palaeochannel deposits on either side of the

divide between the inland and Indian Ocean drainages. Each calcrete area contains a distinct

assemblage of beetles and the fauna occurs in both fresh and saline groundwater. The physico-

chemical properties of the groundwater, and the palaeogeography and hydrology of the region

are discussed in some detail.

C. H. S. Watts, South Australian Museum, North Terrace, Adelaide, South Australia 5000. W.

F. Humphreys, Western Australian Museum, Francis Street, Perth, Western Australia 6000.

Manuscript received 11 October 1999.

Stygobiotic water beetles have been reported

from a number of widely scattered localities

around the world (Spangler 1986) but it was not

until 1999 that they were first reported from

Australia (Watts and Humphreys 1999). These

belonged to the Dytiscidae and were discovered

living in relatively shallow calcrete aquifers in the

Lake Way/Lake Carey palaeodrainage system of

Western Australia. Five species in three genera in

the tribe Bidessini were involved.

These groundwater calcretes are restricted to

the arid parts of Australia (Fig. 40) where the

potential evaporation exceeds rainfall by more

than an order of magnitude (detailed in

Humphreys 1999; Watts and Humphreys

1999).

In May 1999 one of us (W.F.H.) spent some

time investigating other areas of inland

palaeodrainage in Western Australia with

groundwater calcretes and discovered additional

stygofaunas which included dytiscids.

In this paper we describe the six species of

Dytiscidae collected and record the existence of a

further two, all belonging to two of the original

genera: Tjirtudessus and Nirridessus. We also

give details of the physico-chemical properties of

the water in the aquifers and the palaeogeography

and hydrology of the region.

METHODS

About 120 sites were sampled, comprising

pastoral wells and boreholes constructed for water

abstraction, groundwater investigation and

mineral exploration in a number of fractured rock,

alluvial and groundwater calcrete aquifers from

the central Yilgarn Craton of Western Australia

(Fig. 38). The beetles and associated fauna were

taken by plankton nets hauled through the water

column of the bores and wells, or sometimes from

baited traps.

Physico-chemical parameters in the water were

determined either in situ, or in a sample taken

near the surface using a bailer, using electronic

instruments—pH using a WTW pH 320 meter

with a SenTix 97T pH combined electrode with

integrated temperature sensor and redox probe,

and dissolved oxygen using a WTW Oxi 320

meter and a CellOx 325 oxygen sensor

(Wissenschaftlich-Technisch Werkstatten GmbH,

Weilheim, Germany). Conductivity was measured

with a TPS Model LC 84 conductivity meter (TPI

Electronics, Springwood, Queensland, Australia).

All were calibrated as specified using the

recommended standards. In some samples the

salinity was determined using a _ hand

refractometer (Atago S-10e).

128 C.H.S. WATTS & W. F. HUMPHREYS

Abbreviations used

BES Prefix for field numbers, WAM

Biospeleology.

OB Observation bore.

PAT Prefix, observation bore number, Austin

Downs Borefield, Big Bell Mine.

SB Prefix, piezometer number in Hinkler

Well calcrete to the west of Lake Way.

Prefix, piezometer number in Hinkler

Well calcrete to the west of Lake Way.

SAMA South Australian Museum, Adelaide.

WAM Western Australian Museum, Perth.

TPB

SYSTEMATICS

Key To AUSTRALIAN SPECIES OF STYGOBIONIC

BIDESSINI

Body length <1.2 mm, dorsal surface strongly

reticulate, legs stout, without swimming-

hairs on fore and midlegs .................

Kintingka kurutjutu Watts & Humphreys

— Body length >1.2 mm, dorsal surface with

weak to moderate reticulation, legs normal,

all legs with swimming hairs ............... 2

Pronotum usually wider than elytra (Fig. 1);

third and fourth (apical) segments of labial

palpi subequal in length (Figs 23-25);

setae on hind edge of mesofemur not

greatly different in robustness from those

on mesotrochanter (Figs 10, 19,20).Length

> 3-0 MMe ceee-ssvee eens (Tjirtudessus) 9

— Pronotum same width or narrower than elytra

(Figs 3-6); third segments of labial palpi

half to two thirds length of apical (Figs

26-32); setae on hind edge of mesofemur

near base much more robust than those on

mesotrochanter or elsewhere on femur

(Figs 7-9). Length 1.0-3.2 mm .............

Bye ae ln oer ance ea (Nirridessus) 3

Pronotal plicae well marked...............0... 4

— Pronotal plicae difficult to trace.............. 3

Length of metatarsal segments 1 & 2 >

segments 3 & 4; eye remnant present;

parameres with long apical lobe ............

Prete N. pulpa Watts & Humphreys

— Length of metatarsal segments 1 & 2=<

segments 3 & 4; without eye remnant;

parameres with small apical lobe (Fig. 33)

PP Rete heed N. morgani sp. nov.

Length => 3.0 mm; eye remnant present

(Fig. 5); group of six spines close to base

of mesofemur on hind edge (Fig. 11) ....

arate Sleyederasssacsresss N. bigbellensis sp.nov.

— Length =< 2.5 mm; with or without eye

remnant; spines on mesofemur spread out

along hind edge or, if restricted to base,

four or fewer (Figs 8, 9, 13,15). .......... 6

Pro- and mesotarsi broad (Fig. 4); segments

two and three of antennae similar in shape;

apical lobe of paramere overlies part of

rest of paramere (Fig. 34) ...0...2..0...20s000.

Shonedbeacecdosreoohoncconcg N. cueensis sp.nov.

— Pro- and mesotarsi only weakly expanded

(Fig. 3); segments two and three of

antennae different in shape, second broad

and rounded, third narrow and triangular;

apical lobe of paramere well separated

from rest of paramere (Fig. 35)............ 7

Elytron without subsutural row of punctures;

metatrochanters bluntly pointed (Fig. 14);

length approximately 1.5 mm; median lobe

of aedeagus truncated (Fig. 35) .............

sqéssasevavserecvauuneieeaioees N. hinkleri sp.nov.

— Elytron with row of large subsutural

punctures; metatrochanters rounded;

length up to 2.5mm; median lobe of

ACUEAIS DOM imen es fro Le. eee 8

Length 2.2—2.3 mm; with eye remnant......

.. N. windarraensis Watts & Humphreys

— Length 1.3-1.5 mm; without eye remnant

Pees N. lapostaae Watts & Humphreys

Pro- and mesotarsi strongly expanded (Fig.

1); apical five segments of antennae

noticeably thinner than others (Fig. 1)...

SADR aie M a anaer T. magnificus sp. nov.

— Pro-andmesotarsi only moderately expanded

(Fig. 2); apical five segments of antennae

not narrower than others (Fig. 2) ....... 10

10. — Length > 4.0mm; median lobe of aedeagus

twisted, tip knobbed (Fig. 37); without eye

remnant; pronotum at its base a little

narrower than elytra (Fig. 2) «0.00...

ssa ssicscteavech saaescressesvescess T. hahni sp. nov.

— Length < 4.0mm; median lobe of aedeagus

not twisted, tip pointed; with small eye

remnant; pronotum at its base wider than

SLY (Care coer eee ee ce nn ee

NIRRIDESSUS AND TJIRTUDESSUS 129

FIGURES 1-4. 1, Dorsal view of Tjirtudessus magnificus; 2, ditto T. hahni; 3, ditto Nirridessus hinkleri; 4, ditto N.

cueensis; 5, ditto N. bigbellensis; 6, ditto N. morgani. Scale bar = 1mm.

Tjirtudessus Watts & Humphreys, 1999 bore, 27°16'10"S, 117°59'23" E, 12/5/99, coll W.

F. Humphreys & H. J. Hahn’, in spirit, WAM,

registration number 26840.

Types Paratype: f. BES 7066, same data as for

Holotype: m. ‘BES 7040, Old Cue water supply _ holotype except 13/5/99, SAMA.

Tjirtudessus magnificus sp. nov.

130

Description (number examined, 2) Figs 1, 19, 20,

25, 36,

Habitus: Length 4.7-4.8 mm.; relatively flat,

strongly constricted at junction of pronotum/

elytra; uniformly light testaceous; hindwing

vestigial, about half length of elytron (Fig 1).

Head: Large, nearly as wide as elytra; smooth,

reticulation very fine, punctures sparse, weak;

subparallel in posterior half; sides with dark

suture in middle near anterior edge. Antenna

relatively stout, basal two segments cylindrical,

third segment longer and narrower at base, next

three subequal, next four progressively thinner,

apical segment a bit longer and much narrower

than penultimate, each segment with some very

small setae on inside apically (Fig. 1). Maxillary

palpus thin, elongate, apical segment large, a little

shorter than segments one to three combined,

three long setae on outer side and some sensillae

towards tip, tip truncated. Labial palpus moderate,

apical two segments subequal, tip weakly bifid,

penultimate segment with small papilla near tip

bearing two setae (Fig. 25).

Pronotum: Very broad, wider than elytra (Fig.

1); anterolateral angles projecting strongly

forward; base quite strongly narrowed,

posterolateral angles acute; smooth, with sparse,

very weak punctures and a row of stronger

punctures along front margin; basal plicae weak,

reaching to about half way along pronotum,

slightly excavated inwards; with row of long setae

laterally, denser towards front.

Elytra: Not fused, lacking inner ridges;

elongate, widest behind middle, smooth, quite

densely and evenly covered with very small

punctures, row of widely spaced larger punctures

close to inner edge; lacking setiferous

micropunctures; row of long setae near lateral

edge, a few additional larger punctures with long

setae, more frequent towards sides. Epipleuron

broad in anterior fifth, then rapidly narrowing to

be virtually absent over rest of elytron.

Ventral surface: Prothoracic process relatively

broad, strongly narrowed between coxae, not

reaching mesothorax, apical half spatulate,

strongly arched in lateral view with highest point

(viewed ventrally) between coxae. Mesocoxae in

contact at midline. Metathorax sharply triangular

in front in midline, wings very narrow, broadly

rounded in midline behind. Metacoxal plates

large, metacoxal lines short, weak, widely spaced,

reaching to about halfway to metasternum,

diverging in anterior two-thirds; moderately

covered with small setae-bearing punctures;

closely adpressed to first abdominal ventrite. First

C.H.S. WATTS & W. F. HUMPHREYS

and second ventrites fused, sutural lines distinct,

ventrites three to five mobile, moderately covered

with small seta-bearing punctures, ventrites three

and four with a long central seta or bunch of long

setae.

Legs: Protibia relatively narrow, inner edge

straight, outer edge bowed, widest past middle

where it is about three times its basal width;

protarsi greatly expanded, first segment very

broadly oval, second segment broad about one

third length of first, third segment as long as first

but much narrower and very deeply bifid, fourth

segment very small and hidden within lobes of

third segment, apical segment narrow,

cylindrical, about length of third, segments one

to three with very dense covering of adhesive

setae; claws short and simple. Mesotrochanter

rounded with row of setae on inner edge;

mesofemur with row of 10-12 relatively weak

setae along hind edge in basal half (Fig.19);

mesotarsi similar to protarsi. Metatrochanter

weakly pointed (Fig. 20); metafemur elongate,

lacking spines; metatibia strongly curved,

widening towards apex; metatarsi elongate, basal

segment longest, apical segment a little longer

than fourth, segments one and two in

combination about as long as others; claws weak.

Male: Antennae a little stouter; pro- and

mesotarsi a little stouter. Median lobe of aedeagus

narrow, narrowing rapidly in apical quarter;

paramere broad, apical segment with pronounced,

narrow, apical lobe (Fig. 36).

Etymology

The species is named in reference to its

appearance.

Remarks

A very large broad flat Tjirtudessus readily

recognised by the large round first segment of the

pro- and mesotarsi of both sexes, and the

distinctive antennae with the apical segments

noticeably narrower than the middle segments.

Tjirtudessus hahni sp. nov.

Types

Holotype: m. ‘BES 7197, mineral exploration

bore, 26°41'16"S, 120°17'52"E, 21/5/99, coll. W.

F. Humphreys & H. J. Hahn’, slide mounted,

WAM, registration number 26887.

Paratypes: 4; same data as holotype, most

incomplete, 2 slide mounted, SAMA, 2 in spirit,

WAM, registration numbers 26841, 26842.

NIRRIDESSUS AND TJIRTUDESSUS 131

Description (number examined, 5) Figs 2, 21,22, uniformly light testaceous; hindwing vestigial,

24, 37. about one-third length of elytron (Fig. 2).

Habitus: Length 4.8 mm.; relatively flat, strongly Head: Large, smooth, reticulation very fine,

constricted at junction of pronotum/elytra; punctures sparse, weak; subparallel in posterior

13 14

—_

PPEPVIBP py

FIGURES 7- 32. 7, Ventral view of mesotrochanter and mesofemur of Nirridessus pulpa; 8, ditto N. windarraensis;

9, ditto N. lapostaae; 10, ditto Tjirtudessus eberhardi; 11-12, ventral views of mesotrochanter and mesofemur and

metatrochanter and metafemur of N. bigbellensis; 13-14, ditto N. hinkleri; 15-16, ditto N. cueensis; 17-18, ditto N.

morgani; 19-20 ditto T. magnificus; 21-22, ditto T. hahni; 23, labial palpus of T. eberhardi; 24, ditto T. hahni; 25,

ditto T. magnificus; 26, ditto N. bigbellensis; 27, ditto N. hinkleri; 28, ditto N. morgani; 29, ditto N. cueensis; 30,

ditto N. windarraensis; 31, ditto N. lapostaae; 32, ditto N. pulpa.

132

half; sides with dark suture in middle near anterior

edge. Antenna stout, basal two segments largest,

next six segments narrower at base, subequal,

segment six the widest, apical segment twice as

long as penultimate (Fig. 2); each segment with

some very small setae on inside apically.

Maxillary palpus relatively thin, elongate; apical

segment largest, some sensillae towards tip, tip

weakly bifid. Labial palpus not particularly thin;

apical two segments subequal; tip weakly bifid;

penultimate segment with two setae near apex

arising from slight bulge (Fig. 24).

Pronotum: Broad, a little narrower than elytra

(Fig. 2); anterolateral angles projecting strongly

forward; base quite strongly narrowed;

posterolateral angles acute; smooth, punctures

sparse, very small, a row of stronger punctures

along front margin; basal plicae short, very weak,

only visible in some lights; row of long setae

laterally, denser towards front.

Elytra: Not fused; lacking inner ridges;

elongate, widest behind middle. Elytron smooth,

quite densely and evenly covered with small

punctures each with a small seta, row of widely

spaced larger punctures close to inner edge; some

setiferous micropunctures at base, near suture and

near apex; row of long setae near lateral edge, a

few additional long setae, more frequent towards

sides. Epipleuron broad in anterior fifth, then

rapidly narrowing to be virtually absent over rest

of elytron.

Ventral surface: Prothoracic process relatively

broad, strongly narrowed between coxae, not

reaching mesothorax, apical half spatulate, tip

rounded, strongly arched in lateral view with

highest point (viewed ventrally) between coxae.

Mesocoxae meet. Metathorax sharply triangular in

front in midline, wings very narrow, broadly

rounded in midline behind. Metacoxal plates

large; metacoxal lines short, weak, widely spaced,

almost obsolete; punctures very sparse, very weak;

closely adpressed to first abdominal ventrite. First

and second ventrites fused, sutural line virtually

obliterated, ventrites three to five mobile,

moderately covered with small seta-bearing

punctures, ventrites three and four with a long

central seta or bunch of long setae.

Legs: Protibia moderately broad, inner edge

straight, outer edge bowed, widest past middle

where it is about three times its basal width;

protarsi weakly expanded, second segment about

half length of first, third segment equal in length

to first, fourth segment very small and hidden

within deeply lobed third segment, apical segment

long, thin, segments one to three with dense

C.H. S. WATTS & W. F. HUMPHREYS

covering of adhesive setae, claws relatively strong,

simple. Mesotrochanter parallel-sided broadly

triangular at apex with a few setae on inner edge,

mesofemur with a row of five to six relatively

weak setae along hind edge in basal half (Fig. 21);

mesotarsi more elongate than _ protarsi.

Metatrochanter curved on outer edge straight on

inner edge (Fig. 22), metafemur elongate, sinuate

(Fig. 22), metatibia weakly curved, widening

towards apex; metatarsi elongate, basal segment

longest, apical segment a little longer than fourth,

segments one and two in combination about as

long as others; claws weak.

Male: Antennae and tarsi as in female. Median

lobe of aedeagus narrow, twisted slightly,

knobbed at apex; paramere broad, two-segmented,

apical segment with pronounced, narrow, apical

lobe (Fig. 37).

Etymology

This species is named after Hans Jurgen Hahn

who assisted with describing the physicochemical

environment occupied by the stygofauna. The type

locality is an excellent example of this kind of

habitat.

Remarks

T. hahni resembles T. magnificus in its large

size and broad body but differs in its much less

expanded pro- and mesotarsi and in its more

normal antennae which have segments 5-10

subequal in size. It can be separated from both T.

magnificus and T. eberhardi by the median lobe

of the aedeagus which is noticeably twisted and

has a rounded knob at the tip and by its pronotum

which, unlike in those species, is a little narrower

than the elytra.

Nirridessus Watts & Humphreys, 1999.

Nirridessus hinkleri sp. nov.

Types

Holotype: m. ‘BES 7134, SB 32/1, 26°52'31"S,

120°12'05"E, 17/5/99, coll. W. F. Humphreys &

H. J. Hahn’, slide mounted, WAM, registration

number 26843.

Paratypes: 58, same data as holotype, 29 WAM,

registration number 26844, 28 SAMA; 3, ‘BES

7130, TPB25/4, 26°52'50"S, 120°09'44"E, 17/5/

99’, 1 WAM, registration number 26845, 2

SAMA; 3, ‘BES 7218, 26°51°37”S, 120°18’05”E,

22/5/99’, 2 WAM, registration numbers 26846,

26847, 1 SAMA; 34, ‘BES 7137, SB32/1,

NIRRIDESSUS AND TJIRTUDESSUS

26°52'31"S 120°18'05"E’, 12 WAM, registration

number 26848, 15 SAMA; 3, ‘BES 7136,

26°52'31"S 120°12'05"E’, WAM, registration

numbers 26849, 26850, 26851; 1, ‘BES 7166,

26°41'14"S 120°18'09"E’, WAM, registration

number 26852; 2, BES 7228, 1 WAM, registration

number 26853, 1 SAMA. All collected by W. F.

Humphreys & H. J. Hahn.

Additional specimen

The following partial specimen probably

belongs to this species. BES 7222 Hinkler calcrete

east, 26°51'36"S, 120°18'05"E, 22/5/99, coll. W.

F. Humphreys & H. J. Hahn. WAM registration

number 26854.

Description (number examined, 106) Figs 3, 13,

14, 27, 35.

Habitus: Length 1.4-1.8 mm.; relatively flat,

elongate, pronotum narrowing at base; uniformly

very light testaceous; hindwing vestigial, reduced

to about one half length of elytron.

Head: Broad, parallel sided in basal half,

rapidly narrowing forward of area where eye

would be; a short dark suture at each side in

middle at edge; reticulation moderate; punctures

sparse, weak, row of setiferous punctures running

backwards from above antennal base. Antenna

moderately stout (Fig. 3), basal segment parallel

sided, second segment rounded, third much

narrower than second, fourth much shorter, then

approximately the same size until penultimate,

apical segment thinner and about twice as long as

penultimate, a few small setae near apex of each

segment. Tip of last segment of maxillary palpus

very weakly bifid, a few small setae towards tip,

penultimate segment much shorter than apical,

with small papilla bearing two setae near apex

(Fig. 27).

Pronotum: A little narrower than elytra (Fig. 3),

broad in front, narrowing behind, strongly

extended forward at anterolateral angles,

posterolateral angles acute; punctures very sparse,

weak, a few larger punctures towards front edge;

moderately reticulate; basal plicae weak, straight,

reaching about half way along pronotum; row of

long, thin setae in front half at edges and on

forward extensions.

Elytra: Not fused but tightly locked; lacking

inner ridges; widest in middle; punctures very

fine, very sparse, each with a small seta, a few

punctures with longer setae; moderately reticulate;

moderately covered with micropunctures at base,

near suture and apex; sides of elytra quite strongly

vertical, with row of long thin setae at edge,

133

denser towards front. Epipleuron present in

anterior quarter, absent in apical half.

Ventral surface: Pronotal process arched in

lateral view, highest point (viewed ventrally)

between coxae, apical half roughly parallel-sided,

bluntly pointed, narrowing between coxae, not

reaching metathorax. Mesocoxae in contact in

midline. Metathorax with a few very small

punctures; quite sharply triangular in midline in

front; wings very narrow, subobsolete; narrowing

to a broad point behind in midline. Metacoxal

lines, well separated, weakly diverging, reaching

to about halfway to mesosternum; sparsely

punctate; adpressed to first abdominal ventrite.

Metacoxal plates and first and second ventrites

fused but sutures evident, other ventrites free,

sternites three and four with central group of

setae, otherwise virtually without setae; virtually

impunctate.

Legs: Protibia relatively thin, about four times

as broad at apex than at base; protarsi weakly

expanded, second segment about a quarter the

length of first, fourth segment very small, hidden

within deeply bilobed third segment, adhesive

setae sparse, weak; claws weak. Mesotrochanter

relatively large, rounded with a few setae on inner

edge; mesofemur with three to four strong spines

on hind edge restricted to basal half but not all

grouped near base (Fig. 13); mesotarsi less

strongly expanded than protarsi. Metatrochanter

large, outer edge curved, inner edge straight,

weakly pointed, well separated from femur at apex

(Fig. 14); metafemur relatively narrow, anterior

edge weakly sinuate, impunctate, without spines;

metatibia strongly curved, thickening apically;

metatarsal segments elongate, progressively

smaller towards apical segment which is a little

longer than penultimate, combined length of basal

two segments approximately equal to other three;

claws weak, outer one slightly smaller than inner.

Male: Appendages and legs as for female.

Median lobe of aedeagus moderately broad,

concave above, tip broadly rounded with small

central point; parameres moderately broad, two

segmented, apical segment with pronounced

narrow apical portion (Fig. 35).

Etymology

The name pertains to the Hinkler Well

Catchment in which the classic study of the

hydrogeochemistry of ground water calcretes was

undertaken (Mann and Deutscher 1978).

Remarks

This is a small species with a distinctive

134

aedeagus. The pointed rather than rounded

metatrochanters and acute rather than rectangular

posterolateral angles to the pronotum, will

separate it from the similar sized N. lapostaae.

Nirridessus cueensis sp. nov.

Types

Holotype: m. ‘BES 7040, Old Cue water supply

bores, 27°16'11"S, 117°59'23"E, 12/5/99, coll W.

F. Humphreys & H. J. Hahn’, slide mounted,

WAM, registration number 26856.

Paratypes: 6, same data as holotype, 2 SAMA,

4 WAM, registration numbers 26857, 26858,

26859, 26860; 8, same data as holotype except

for, ‘BES 7067, 13/5/99’, 4, SAMA 4, WAM,

registration numbers 26861, 26862, 26863,

26864; 2, same data as holotype except for ‘BES

7038’, WAM, registration numbers 26865, 26866.

Description (number examined, 18) Figs 4, 15,

16, 29, 34.

Habitus: Length 2.1-2.4 mm.; relatively flat,

elongate, pronotum a little narrower than elytra,

constricted at base; uniformly very light

testaceous; hindwing vestigial, reduced to about

one half length of elytron.

Head: Broad, parallel sided in basal half,

rapidly narrowing forward of area where eye

would be; a short dark suture at each side in

middle at edge; reticulation very weak; punctures

sparse, weak, row of setiferous punctures running

backwards from above antenna base. Antenna

stout, robust, basal segment cylindrical, second

more elongate, third smaller narrowing to base,

then progressively widening until penultimate,

apical segment thinner and slightly longer, a few

small setae near apex of each segment. Last

segment of maxillary palpus relatively broad, tip

weakly bifid, a few small setae towards tip.

Penultimate segment of labial palpus with strong

cone-like projection near apex bearing two setae,

approximately two thirds length of apical (Fig.

29).

Pronotum: Same width or a bit narrower than

elytra, broad in front, narrowing quite markedly

behind, strongly extended forward at anterolateral

angles, posterolateral angles rectangular;

punctures sparse, weak; weakly reticulate; two

fine, basal plicae weakly impressed, straight,

reaching about a half way along pronotum; row of

long, thin setae in front half at edges and on

forward extensions.

Elytra: Not fused but tightly locked, lacking

C. H.S. WATTS & W. F. HUMPHREYS

inner ridges; widest in middle; punctures very

fine, sparse, each with a small seta, a few

punctures with longer setae; a few micropunctures

near apex; sides of elytra quite strongly vertical;

with row of long thin setae at edge, denser

towards front. Epipleuron broad in front,

narrowing quite rapidly to level of first sternite,

then thin to apex, difficult to differentiate from

disc.

Ventral surface: Pronotal process arched in

lateral view, highest point (viewed ventrally)

between coxae, apical half broadly spatulate,

narrowing between coxae, not reaching

metathorax. Mesocoxae in contact in midline.

Metathorax with a few very small punctures; quite

sharply triangular in midline in front; wings very

narrow, subobsolete; rounded behind. Metacoxal

plates with weakly raised central portion;

metacoxal lines weak, well separated, diverging in

anterior third, reaching to about halfway to

mesosternum; sparsely punctate; adpressed to first

abdominal ventrite. Metacoxal plates and first and

second ventrites fused but sutures evident, other

ventrites free, ventrites three and four with central

group of setae, otherwise virtually without setae;

virtually impunctate.

Legs: Protibia about three times as broad at

apex than at base; protarsi moderately expanded,

second segment half length of first, third

segment deeply bilobed, a little longer than first,

fourth segment very small and hidden within

bilobed third segment, adhesive setae moderately

dense; claws moderate. Mesotrochanter rounded

at tip with some setae on inner edge (Fig. 15);

mesofemur with three to five strong spines on

hind edge grouped together near base (Fig.15);

mesotarsi similar to protarsi. Metatrochanter

large, outer edge rounded, inner edge straight,

weakly separated from femur at apex (Fig.16);

metafemur relatively narrow, anterior edge

weakly sinuate, impunctate, without spines;

metatibia strongly curved, thickening apically;

metatarsal segments elongate, progressively

smaller towards apical segment which is a little

longer than penultimate, combined length of

basal two segments approximately equal to other

three; claws weak, outer one slightly smaller than

other.

Male: Appendages and legs as above. Median

lobe of aedeagus moderately broad, concave

above, narrowing rapidly close to tip, small brush

of setae dorsally near tip; parameres moderately

broad, apical segment with pronounced narrow

apical portion which partly overlaps basal portion

(Fig. 34).

NIRRIDESSUS AND TJIRTUDESSUS 135

Etymology

This species is named after the type locality.

Remarks

Nirridessus cueensis seems close to N.

lapostaae but is larger, its pro- and mesotarsi are

much broader and its metatrochanters are not as

rounded. The spines/strong setae on the hind edge

of the mesofemur are grouped together near the

base whereas in N. lapostaae they are more spread

out along the basal half of the femur. The

aedeagus differs from most Nirridessus by having

a group of setae near the apex. The parameres are

unique in Nirridessus in having the apical lobe

overlapping part of the rest of the paramere and

with strong spines on the inner edge (Fig. 34).

Nirridessus morgani sp. nov.

Types

Holotype: m. ‘BES 7192, Sample 2 Site 284,

FIGURES 33-37. 33, Paramere, dorsal view of central lobe and lateral view of central lobe of aedeagus of

Nirridessus morgani; 34, ditto N. cueensis; 35, ditto N. hinkleri; 36, ditto, Tjirtudessus magnificus; 37, ditto T.

hahni.

136

mineral exploration bore, 26°41'15"S,

120°21'10"E; 21/5/99, coll W. F. Humphreys &

H. J. Hahn’, in spirit, WAM, registration number

26867.

Paratypes: 6, as for holotype, 4 SAMA, 2

WAM, registration numbers 26868, 26869; 5,

‘BES 7171, Site 262, mineral exploration bore,

Lake Way, 26°68'74"S 120°35'31"E, 19/5/99’, 1

SAMA, 4 WAM, registration numbers 26870,

26871, 26872, 26873.

Description (number examined, 10) Figs 6, 17,

18, 28, 33.

Habitus: Length 2.1-2.2 mm; relatively flat but

deep bodied, elongate, pronotum weakly

narrowing at base; uniformly very light testaceous;

hindwing vestigial, reduced to about one half

length of elytron.

Head: Relatively narrow, parallel sided in basal

half, rapidly narrowing forward of area where eye

would be; a short dark suture at each side in

middle at edge; very weak reticulation; punctures

sparse, weak, row of setiferous punctures running

backwards from above antenna base; weakly

reticulate. Antenna relatively stout, basal two

segments broad, third narrower, fourth similar to

third, then approximately equal in size until

penultimate, apical segment a bit longer but same

width, a few small setae near apex of each

segment. Tip of last segment of maxillary palpus

weakly bifid, a few small setae towards tip.

Penultimate segment of labial palpus shorter than

apical, with two setae on slight bulge near apex

(Fig. 28).

Pronotum: Much narrower than elytra, narrower

behind, strongly extended forward at anterolateral

angles, posterolateral angles square; punctures

very sparse weak, a few larger punctures towards

front edge; two strongly raised basal plicae,

straight, reaching about half way along pronotum;

row of long, thin setae in front half at edges and

on forward extensions.

Elytra: Fused but may open slightly in

preparations, lacking inner ridges; sides

subparallel in middle; punctures very fine, sparse,

each with a small seta, a few punctures with

longer setae; moderately covered with

micropunctures particularly at base, near suture

and apex; sides of elytra quite strongly vertical;

with row of long thin setae at edge, denser

towards front. Epipleuron broad in front quarter,

narrowing quite rapidly to middle, absent in apical

half.

Ventral surface: Pronotal process arched in

lateral view, highest point (viewed ventrally)

C.H.S. WATTS & W. F. HUMPHREYS

between coxae, apical half parallel sided, tip

weakly and bluntly pointed, narrowing between

coxae, not reaching metathorax. Mesocoxae in

contact in midline. Metathorax with a few small

punctures; broadly triangular in midline in front;

wings very narrow; narrowing to blunt point

behind. Metacoxal plate with weakly raised

central portion; metacoxal lines well separated,

progressively diverging, reaching to about halfway

to mesosternum; sparsely punctate; adpressed to

first abdominal ventrite. Metacoxal plates and first

and second ventrites fused but sutures evident,

other ventrites free, ventrites three and four with

central group of setae, otherwise virtually without

setae; virtually impunctate.

Legs: Protibia about three times as broad at

apex than at base; protarsi weakly expanded,

second segment not much shorter than first, the

fourth segment very small and hidden within

deeply bilobed third segment, adhesive setae

sparse; claws weak. Mesotrochanter sharply

pointed, without setae on inner edge; mesofemur

with two strong setae on hind margin at base (Fig.

17); mesotarsi slightly less strongly expanded than

protarsi. Metatrochanter large, completely

exposed, pointed, close to metafemur at apex (Fig.

18); metafemur relatively narrow, anterior edge

weakly sinuate, impunctate, without spines;

metatibia curved, thickening apically; metatarsal

segments elongate, progressively smaller towards

apical segment which is a little longer than

penultimate, combined length of basal two

segments approximately equal to other three;

claws weak, outer one slightly smaller than other.

Male: Appendages and legs as for female.

Median lobe of aedeagus moderately broad,

concave above, narrowing to tip, without setae;

parameres moderately broad, apical segment

relatively long with small apical lobe (Fig. 33).

Etymology

This species is named after Kevin Morgan for

his understanding of the hydrology and

geochemistry of the palaeodrainage channels in

Western Australia (Morgan 1993).

Remarks

The well marked pronotal plicae and pointed

metatrochanters suggest a relationship with N.

pulpa. The lack of an eye remnant and thinner

antennae will separate it from this species. The

distinctive, small, apical lobe on the paramere and

the group of strong spines on the hind edge of the

mesofemur reduced to two will separate it from

all other Nirridessus.

NIRRIDESSUS AND TJIRTUDESSUS 137

Nirridessus bigbellensis sp. nov.

Types

Holotype: f. ‘BES 7050, borefield monitoring

bore PAT 7, Austin Downs Pastoral Station,

27°24'48"S 117°42'40"E, 12/5/99’, coll. W. H.

Humphreys & H. J. Hahn’, slide mounted, WAM,

registration number 26874.

Paratype: f. ( partial specimen) as for holotype,

slide mounted, SAMA.

Description (number examined, 2) Figs 5, 11, 12,

26.

Habitus: Length 3.0-3.2 mm.; relatively flat

and broad, pronotum weakly narrowing at base;

uniformly very light testaceous; hindwing

vestigial, reduced to about two-thirds length of

elytron.

Head: Broad, parallel sided in basal half,

rapidly narrowing forward of area where eye

would be; a small oval area delineated by dark

sutures at each side in middle at edge; very weak

reticulation; punctures sparse, weak, row of

setiferous punctures running backwards from

above antennal base. Antenna relatively thin, basal

two segments broadest, third as long as second

but narrower, fourth shorter, then approximately

equal in size until apical segment which is longer

and thinner than penultimate, a few small setae

near apex of each segment. Maxillary palpus

relatively thin, apical segment as long as other

three combined, tip of last segment truncated, a

few small setae towards tip. Apical segment of

labial palpus twice length of penultimate which

has two setae on slight bulge near apex (Fig. 26).

Pronotum: Broad, narrower than elytra, strongly

constricted near base, strongly extended forward

at anterolateral angles, posterolateral angles acute;

punctures very sparse, weak, a few larger

punctures towards front edge; basal plicae

subobsolete; row of long, thin setae in front half

at edges and on forward extensions.

Elytra. Not fused, lacking inner ridges; sides

rounded; punctures small, very sparse, setiferous,

a few punctures with longer setae; some

micropunctures near apex; row of long thin setae

laterally, denser towards front. Epipleuron broad

in front fifth, narrowing quite rapidly to middle,

absent in apical half.

Ventral surface: Pronotal process arched in

lateral view, highest point (viewed ventrally)

between coxae, apical half parallel sided, tip

weakly and bluntly pointed, narrowing between

coxae, not reaching metathorax. Mesocoxae in

contact in midline. Metathorax with a few small

punctures; broadly triangular in midline in front;

wings narrow; rounded behind in midline.

Metacoxal plates sparsely punctate; metacoxal

lines, relatively close, weakly diverging, reaching

to about two thirds of way to mesosternum;

adpressed to first abdominal ventrite. Metacoxal

plates and first and second ventrites fused, other

ventrites free, ventrites three and four with central

group of setae, otherwise virtually without setae;

virtually impunctate.

Legs: Protibia elongate about three times as

broad at apex than at base; protarsi weakly

expanded, second segment not much shorter than

first, the fourth segment very small and hidden

within deeply bilobed third segment, adhesive

setae moderate; claws moderately strong.

Mesotrochanter parallel sided, apex rounded with

a few setae on inner edge; mesofemur with six

strong setae on hind margin at base (Fig. 11);

mesotarsi more elongate than _protarsi.

Metatrochanter large, inner edge curved, outer

edge straight, rounded at tip, well separated from

femur at apex (Fig. 12); metafemur narrow,

anterior edge weakly sinuate, impunctate, without

spines; metatibia strongly curved, thickening

apically; metatarsal segments elongate,

progressively shorter towards apical segment

which is a little longer than penultimate,

combined length of basal two segments

approximately equal to other three; claws weak.

Male: Not known.

Etymology

The name pertains to Big Bell Mine which

draws its water from the aquifer in which the

species lives.

Remarks

At over three millimetres in length the largest

Nirridessus so far known. It also is more rounded

in outline and less flattened than most. The thin

maxillary and labial palpi, narrow metafemur and

strongly acute posterolateral angles of the

pronotum also set it apart.

ADDITIONAL SPECIMENS

Taxon 1

One small (length 1.5 mm) female specimen

was collected at Lake Violet. Although close to N.

hinkleri it probably represents a distinct species.

More specimens, including males are needed to

confirm this.

‘BES 7160, observation bore for Pump 5,

138 C.H.S. WATTS & W. F. HUMPHREYS

Wiluna Gold Lake Violet Borefield, 26°41'08"S,

120°13'05"E 8/5/99, coll. W. F. Humphreys & H.

J. Hahn’, slide mounted, WAM, registration

number 26875.

Taxon 2

Eight partial specimens of a large species

(approximately 4.0mm long.), together consisting

of pronotum, abdomen, elytra and male genitalia,

but lacking head and all appendages, were

collected at Lake Way. These represent a

distinctive species of uncertain generic placement.

‘BES 7222, Sample 3 Site 289 Hinkler calcrete

east, unequipped water bore, 26°51'36"S,

120°18'05"E, 22/5/99, coll. W. F. Humphreys &

H. J. Hahn’, 4 in spirit, WAM, registration

numbers 26876, 26877, 26878, 26879. 4 slide

mounted, SAMA.

Larvae

Larvae of two very different taxa were collected

at Austin Downs (type 1) and Lake Violet (type

WESTERN

AUSTRALIA

2). Although bidessine they differ considerably

from the two larval taxa described in our earlier

paper (Watts and Humphreys 1999). In

comparison to epigean bidessine larvae we would

consider the four larval taxa to be generically

distinct. None appear to belong to the very small

Kintingka kurutjutu Watts and Humphreys. This

conclusion is in conflict with the adult taxonomy.

Association of larvae and adults and more

collecting will be needed to resolve this.

Type 1: 1, ‘BES 7021, borefield monitoring

bore PAT 2, Austin Downs Pastoral Station ,

27°23'44"S, 117°42'25"E, 11/5/99’, slide mounted,

WAM, registration number 26880; 1, ‘BES 7050,

borefield monitoring bore PAT 7, Austin Downs

Pastoral Station , 27°24'48"S, 117°2'40"E, 12/5/

99’, in spirit, SAMA; 1, ‘BES 7055 borefield

monitoring bore PAT 1, Austin Downs Pastoral

Station , 27°23'19"S, 117°43'33"E, 12/5/99’, slide

mounted, SAMA.

Type 2: 10, ‘BES 7148, observation bore for

Leonora ac

Lake

Carey

100 km

FIGURE 38. The distribution of stygal Dytiscidae in the Yilgarn Region. Groundwater calcrete deposits (shaded)

are shown in the palaeodrainage channels (solid lines). The dotted line denotes the surface drainage divide (Beard

198); inland drainage to the east, Indian Ocean drainage to the west. The numerals denote discrete areas of calcrete

(Table 1): 1, Paroo; 2, Austin Downs; 3, Cue; 4, Lake Violet; 5, North-east Lake Way; 6, Hinkler Well; 7, Mount

Windarra.

NIRRIDESSUS AND TJIRTUDESSUS

139

TABLE 1. The distribution of stygal species of dytiscids amongst discrete calcrete bodies in the Yilgarn district of

Western Australia.

Genus

Calcrete body

Tjirtudessus W & H

Western drainage

Austin Downs -

Cue 4magnificus sp. nov.

Eastern drainage

Paroo eberhardi W & H!

Lake Violet =

NE Lake Way hahni sp. nov.

Hinkler Well

Mount Windarra lapostaae W & H

Nirridessus W & H Kintingka W & H

bigbellensis sp. nov. -

‘cueensis sp. nov. -

*pulpa W & H "kurutjutu W & H

undescribed sp. =

morgani Sp. nov. =

hinkleri sp. nov. -

3windarraensis W & H . os

'W & H= Watts & Humphreys 1999; ? sympatric; * sympatric; ‘sympatric.

Pump 1, Wiluna Gold Lake Violet Borefield ,

26°40'30"S, 120°13'55"E, 18/5/99’. 4 WAM,

registration numbers 26881, 26882, 16883, 16884,

6 SAMA, slide mounted and in spirit; 1, same

data but ‘BES 7231, WAM, registration number

26885; 1, same data but ‘BES 7242’, WAM,

registration number 26886. All collected by W. F.

Humphreys and H. J. Hahn.

DIscuSssION

Distribution

Dytiscid specimens were collected from six

separate calcrete deposits: 1, Austin Downs

(borefield for Big Bell Gold Mine); 2, Cue

(former borefield for the town of Cue); 3, Paroo

(pastoral station, detailed in Watts and Humphreys

1999); 4, Lake Violet at the northern end of Lake

Way contains the borefield for Wiluna Gold Mine;

5, the Hinkler Well calcrete, the

hydrogeochemistry of which is detailed in Mann

and Deutscher (1978); and 6, the northeastern side

of Lake Way adjacent to the Lakeway Uranium

prospect (DCE 1981) (Fig. 38). Sites 1 and 2 drain

towards the Indian Ocean whilst the remainder are

to the east of the drainage divide in the Carey

palaeodrainage system (sensu Morgan 1993)

which drains to the interior of the continent. The

distribution of the taxa by calcrete body is shown

in Table 1.

No dytiscids or other large stygofauna were

taken from open, hand dug, pastoral wells even if

they were adjacent to narrow bores containing

stygal dytiscids and other stygofauna. Despite

sampling widely in non-calcrete aquifers no

dytiscids were collected in other than calcrete

aquifers, which confirms the conclusion of our

earlier, much more restricted sampling, that the

beetle and other larger stygofauna is restricted to

aquifers in areas of calcrete (W. F. Humphreys

unpublished; Watts and Humphreys 1999). In

addition, this study extends the range of waters

inhabited by stygal dytiscids, and other

stygofauna, to saline water with a salinity of at

least 20 g I.

These discoveries significantly extend the

known range of both Tjirtudessus and Nirridessus

from the inland draining Lake Way/Lake Carey

palaeodrainage channel to the seemingly never-

connected Murchison palaeodrainage system that

drains westward to the Indian Ocean. No

Dytiscidae were discovered in stygofauna-rich

calcrete aquifers in the Pilbara region. Since only

about 9% of the major calcrete deposits in the

palaeodrainage channels of Western Australia

alone (Humphreys 1999) have been sampled for

stygofauna it is likely that additional subterranean

Dytiscidae remain to be discovered in Australia.

Associated fauna

The dytiscids were collected amongst a diverse

stygofauna comprising mainly phreodrillid

Oligochaeta, bathynellids (Syncarida), Ostracoda,

cyclopoid and_ harpacticoid Copepoda,

crangonyctoid and ceinid Amphipoda and

Haloniscus (Oniscoidea: Isopoda).

Water quality

Fresh to saline groundwaters occur widely in

calcretes and may be reached as close as two

metres from the surface to more than 100 m below

the surface. The calcrete aquifer itself may vary

up to 30 m in thickness (Barnett and Commander

140

Depth (m)

15 20 25

C.H.S. WATTS & W. F. HUMPHREYS

@ Temperature

@ Salinity

30 35 40 45

Temperature (°C) or Salinity (g L-1)

Depth (m)

0 5 10

15) 20 25 30

Temperature (°C) or Salinity (g L-1)

FIGURE 39. Physiochemical profiles in some aquifers from which dytiscid beetles were collected. Upper, site 286

NE Lake Way; lower, site 289 Hinkler East.

1985), but those sampled in the Yilgarn are

typically thin and the groundwater is close to the

surface (Table 2). The area is arid and rainfall (c.

200 mm per year) is irregular predominately as

episodic heavy falls (Sanders 1973) which

recharge the calcrete aquifers through the porous,

often karstic (Barnett and Commander 1985),

surfaces. Indeed, groundwater calcretes are

carbonate deposits forming near the water table in

arid lands as a result of concentration processes

by near-surface evaporation and they are

associated with slow moving groundwater that

fluctuates widely in depth (Jacobson and Arakel

1986; Morgan 1993). In consequence the

groundwater table varies quite widely between

storm events and markedly in salinity. The

dytiscids were collected from water of variable

quality (Table 2) with samples collected from

fresh water (salinity < 1000 mg I') as well as

saline conditions (salinity > 20 000 mg I'). At

some locations the water was well stratified with

marked changes in the physico-chemical

conditions with depth, while at others the vertical

profile was less marked (Fig. 39; Table 2).

Considerable seasonal variation occurs in the

salinity of at least some of the habitats (foot note

to table 2) but the effect of this on the distribution

of the stygofauna is unknown. Marked seasonal

changes in numbers have been reported for stygal

dytiscids inhabiting alluvial aquifers near rivers

(Ordish 1976), and it is known that changes to the

water table and the direction of groundwater flow

have profound effects on the location of such

populations (Richoux and Reygrobellet 1986).

Palaeogeography and hydrology

While much is still to be learnt of these stygal

dytiscids, it is worthwhile at this stage to consider

the palaeogeographic and hydrological setting in

which this fauna, and that reported by Watts and

Humphreys (1999), occurs.

The Western Shield of Western Australia is

divided from north to south by a drainage divide

that separates the rivers, some still active,

draining to the Indian Ocean, from those

draining to the east and which are now largely

inactive and disorganised (Beard 1998). The

present drainage, now mostly a palaeodrainage

system, formed during the Mesozoic when the

Western Shield was attached to Antarctica

(Beard 1998; van de Graaf et al. 1977; Morgan

1993). The sediments’ infilling the

palaeochannels are mostly Eocene or later but

the age of the calcretes is unknown. Morgan

NIRRIDESSUS AND TJIRTUDESSUS 141

TABLE 2. Physico-chemical characteristics of sites from which stygal dytiscids were recorded.

Site Sample Salinity Temperature Dissolved O, pH 7Depthto Depth of

depth (m) gL A % sat mg L" water (m) water(m)

PAT2 — Austin Downs - 52 - - - - 4.8 6

PATI — Austin Downs _ 8! - — - - 8 4.8

Old Cue town bore - S) 25 50.5 3.9 - 4 0.5

GSWAS surface- Paroo ~ 1.38 26.7 59.7 4.5 Si. 3.6 34

GSWAS deep- Paroo - = - 49 ag - - -

GSWA6 surface- Paroo - 0.77 24.8 59.7 4.6 15 5 36.6

GSWAB6 deep- Paroo - 0.79 24.7 43.5 3.4 18 - _

GSWAIS5 small- Paroo - 0.53 26 59.2 4.5 8.4 3 38

GSWAI5 small- Paroo - 0.6 25.3 38.6 3 7.9 - -

SB 32/1 — Hinkler west - 1.66 26.9 79.2 6 7.7 4.8 2

TPB 25/4 — Hinkler west — 2 - = - = 4.8 34

OB Pump 1 — Lake Violet — 2.36 24.6 88.7 / 8.2 4.8 5.6

262 — NE Lake Way - 4.46 25.7 86.1 6.6 7.6 4 1.6

286 — NE Lake Way 0 19.8 24.2 23 1.8 - 2: 20

286 0.5 20.1 24.4 - - - - -

286 1 20.7 24.6 - ~ — = =

286 2 23 24.2 - - - - -

286 4 32.6 23.7 - - - - -

286 8 40.5 23:2 - - - - -

288 — Hinkler east 0 1.46 24.2 68.6 5.4 8.3 2.4 19

288 0.5 2.43 24.5 - - - - -

288 1 1.43 24.5 - - = = —

288 2 1.42 24.5 - - - - -

288 4 1.42 24.5 - - - - -

288 7 1.42 24.5 - - - - -

289 — Hinkler east 0 1.56 24.4 74.4 5.9 8 2.4 19

289 0:5 1.7 24.6 - - - = -

289 1 2.04 24.5 - - - - -

289 2 2.1 24.5 — = = - a

289 4 2.12 24.5 - - - - -

289 8 2.15 24.5 - - - - -

' PATI and PAT2 have annual salinity variation between 5-9 g I! and 5-22 g I respectively as determined from borefield monitoring by Big

Bell Mine. ? Depth approximate. *To base of calcrete (Sanders 1972).

(1993) considered it likely that they formed from

the start of the Oligocene following the onset of

the continental aridity but they have probably

been remobilized and redeposited. As this

process is continuing it is not possible to date

the calcretes using standard radiometric methods.

The Yilgarn Craton covers 750,000 km? of

southwestern Australia between latitudes 34° and

25° S and has mostly not been submerged by the

sea since the beginning of the Mesozoic. The

northeastern half is semi-arid with unreliable

rainfall that may fall throughout the year.

The central watershed traverses a palaeosurface

barely modified since the Cretaceous (Beard

1998). This central watershed is of fairly uniform

elevation traversed only by a few minor gaps or

low points that may be indicative of a change to

the drainage patterns in the distant past. One of

these is on Killara Station where a col, 50-100 m

below the level of the adjacent watershed,

separates the Murchison and the Carey (Lake

Way-Lake Carey in Watts and Humphreys 1999)

palaeodrainage systems (Beard 1998) which

together encompass the entire distribution of the

Australian stygal dytiscids known to this time.

While it is tempting to invoke some significance

of this gap to the biogeography of the stygofauna,

this minor gap may merely reflect the trend of less

resistant Proterozoic rocks (Beard 1998).

The collection sites at Cue and Lake Austin

142 C.H.S. WATTS & W. F. HUMPHREYS

FIGURE 40. The distribution of groundwater calcrete aquifers in Western Australia (from Humphreys 1999).

Modern and palaeo-drainage (respectively, continuous and dashed lines) and calcrete areas (black) are shown.

Derived from data in Geological Survey (1989), drawn by Julianne Waldock on a base map provided by Philip

Commander (further details in Humphreys 1999).

NIRRIDESSUS AND TJIRTUDESSUS 143

(containing Tjirtudessus magnificus, Nirridessus

cueensis and Nirridessus bigbellensis) are

separated from those to the east (containing

Tjirtudessus hahni, Nirridessus hinkleri and

Nirridessus morgani plus the five species in Watts

and Humphreys 1999) by the continental water

divide (Beard 1998) between the inland drainage

and that draining to the Indian Ocean. As the

water divide is also the approximate water divide

for the palaeodrainage systems that date from at

least the Cretaceous no subterranean hydrological

connection is likely to have occurred. The

presence of congeneric stygal species on either

side of this water divide suggests that the fauna

colonised groundwater from a widespread epigean

ancestor, possibly driven by the increasing aridity

of central Australia since the Eocene.

Groundwater calcretes stretch through arid

central Australia as far as the border of the

Northern Territory and Queensland (Fig. 40;

Humphreys 1999). Similar calcretes occur on the

Pilbara Craton which comprises much of the rest

of the Western Shield of Australia, and there

also, each discrete calcrete area sampled so far

contains a distinct stygofauna (Poore and

Humphreys; W. F. Humphreys and S. M.

Eberhard, unpublished; S. M. Eberhard,

unpublished; S. M. Eberhard and W. F.

Humphreys unpublished). However, no dytiscids

have been taken from the Pilbara Craton.

Each of the seven discrete, but sometimes

adjacent, calcrete areas sampled from the

palaeodrainage lines on the Yilgarn Craton (herein

and Watts and Humphreys 1999) contains a

distinct assemblage of dytiscids (and probably

other stygofauna). The data represented here

contains samples from only two of 42 major

calcretes areas in the upper Murchison catchment,

and five out of 18 major calcrete areas in the

Carey palaeodrainage system. In Western

Australia alone there are about 210 major calcrete

areas divided amongst about five major drainage

systems (Fig. 40)

Each calcrete sampled in the Yilgarn (and

Pilbara) containing stygofauna has a unique

stygofaunal community and as only a small

proportion of the available calcrete areas have

been sampled, this suggests that there is

considerable biodiversity to be unearthed amongst

the dytiscids and in the arid zone stygofauna

generally.

Changes to the water table and the direction of

groundwater flow may have profound effects on

the location of populations of stygal dytiscids

(Richoux and Reygrobellet 1986) and unique

stygal assemblages, including Dytiscidae, may be

lost if groundwater pollution occurs (Uéno

1996). As the stygofaunas are unique with

circumscribed distributions and they occur in

systems of potential or actual resource

developments, they present a real challenge for

innovative environmental management.

Furthermore, these calcrete aquifers are

sufficiently replicated and contain a diversity of

fauna sufficient to test independently theories

and processes that gave rise to the vicariance

within these systems.

While much work is needed to start to

understand distribution patterns, the faunistic

distinctiveness of the groundwater calcrete

aquifers is consistent with the evolution of the

hydrogeological system in the palaeodrainage

channels as interpreted by Morgan (1993). In

essence, Morgan argues that in the palaeorivers

north of latitude 30°S separate geochemical

systems develop associated with the formation

of each salt lake (playa) along a

palaeodrainage system. In the groundwater

there is a well defined change in common ion

ratios developed with increasing salinity

marked especially by a relative increase in

chloride and sulphate with respect to other

ions—this may contribute to the heterogeneity

in stygofauna distribution within a given

calcrete area reported by Poore and Humphreys

(1998). As it is related to the rate of movement

of the groundwater, this increase in salinity

and relative chloride/sulphate content is both

spatial and temporal because the change takes

place between widely separate intake and

outflow locations. This hydrochemical trend

commences at the headwaters of each recharge

system, such as a large alluvial fan, and

completes its cycle at the evaporation outlet

marked by the lower boundary of the calcrete

with a salt lake. The main channel calcretes are

formed at the downstream end of an individual

hydrochemical system and immediately

upstream of an evaporation outflow area

forming a salt lake (Morgan 1993). Several

similar hydrochemical cycles may occur along a

single palaeodrainage system.

The marked age and stability of the

palaeodrainage systems themselves, coupled with

the repeated cycles of fresh to hypersaline (>200

g I') groundwaters along the length of each

palaeodrainage system would effectively isolate

each stygal assemblage within the region where

groundwater characteristics are suitable for their

development.

ACKNOWLEDGMENTS

We would like to thank R. Gutteridge (Adelaide) for

the dytiscid illustrations and librarians M. Trifitt

(WAM), M. Anthony, and J. Evans (SAMA). Access

and local information was provided by D. and J. Ford,

Paroo Station, and by the Environmental Officers at Big

Bell Mine (P. Horwood) and Wiluna Gold (R. Bird).

Kevin Morgan generously discussed the hydrology of

the area and provided a copy of an unpublished ms. We

C.H.S. WATTS & W. F. HUMPHREYS

appreciate the comments on the non-dytiscid fauna by J.

Bradbury (Amphipoda), A. Pinder (Oligochaeta), P.

Marmonier (Ostracoda), G. Pesce (Copepoda), H. K.

Schminke (Syncarida), G. D. F. Wilson and S. Taiti

(Isopoda). We thank Dr H. J. Hahn (Universitat

Koblenz— Landau, Germany) for his stimulating

assistance during the field work in 1999 and which was

enabled by a BankWest Landscape Conservation Visa

Card Trust Fund Grant to WFH.

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