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AUSTRALIAN

MUSEUM

VOLUME 34 PART 1

MAY 2001

CONTENTS

BAEHR, M. & HUDSON, P.

New species and a new genus of the subfamily Pogoninae from

South Australia (Coleoptera: Carabidae).

CLARKE, P. A.

The significance of whales to the Aboriginal people of southern South Australia.

CLARKE, P. A.

Review — Warrabarna Kaurna! Reclaiming an Australian Language

JAMES, B. H., DONNELLAN, S. C. & HUTCHINSON, M.N.

Taxonomic revision of the Australian lizard Pygopus nigriceps (Squamata: Gekkonoidea).

KEAR, B. P.

Elasmosaur (Reptilia: Plesiosauria) basicranial remains from the

Early Cretaceous of Queensiand.

McARTHUR, A.

Obituary — Patricia Marietje Thomas

MEGAW, J. V. S.

Australian Aboriginal Cultures Gallery ‘The Speaking Land’: a review article.

PLEDGE, N. S.

Obituary —- June Mavis Scrymgour

REID, A. L.

Sepia hedleyi Berry, 1918 (Cephalopoda: Sepiidae): a complete description and

clarification of the status of S. dannevigi Berry, 1918 and S. rex (Iredale, 1926).

STEWART, P. J. & STRATHERN, A.

Timbu Wara figures from Pangia, Papua New Guinea.

UNMACK, P. J.

Corrected identifications and lectotype designation for Porochilus

argenteus (Zietz) (Osteichthyes: Siluriformes: Plotosidae).

WATTS, C.H. S.

A new species of Australian Canthydrus Sharp with a key to the Australian

species of Noteridae (Coleoptera).

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

A new genus and six new species of Dytiscidae (Coleoptera) from underground

waters in the Yilgarn palaeodrainage system of Western Australia.

Volume 34(1) was published on 30 May 2001.

Volume 34(2) was published on 24 December 2001.

ISSN 0376—2750

19-36

135-138

37-52

127-134

53-56

115-126

139-141

79-98

65~78

57-59

61-64

99-114

NEW SPECIES AND A NEW GENUS OF THE SUBFAMILY POGONINAE

FROM SOUTH AUSTRALIA (COLEOPTERA : CARABIDAE)

M. BAEHR & P. HUDSON

Summary

A new genus and a new species, Syrdenoidius spinipes gen. nov., sp. nov., and three new species of

the genus Pogonus Dejean from salt lakes in South Australia are described: Pogonus matthewsi, P.

vicunus, and P. perovalis. The first two species are closely related to P. grossi Moore from Lake

Eyre and P. saskiae Baehr from Lake Gairdner and Island Lagoon and together they form a complex

of four very closely related species. Pogonus perovalis sp. nov. is outstanding in its body shape and

absence of wings, but may be remotley related to this complex. The new genus Syrdenoidius is

highly similar in shape and structure to the Palaearctic genus Syrdenus Chaudoir, though it differs

mainly in the asetose prosternum, the markedly spinose tibiae, the absence of wings, and a very

unusual stylomere 2. Syrdenoidius spinipes sp. nov. and Pogonus perovalis sp. nov. are outstanding

in their absence of wings and loss of flying ability, attributes that are extremely rare within

Pogoninae. Most of the new species were collected by excavation of their burrows on the lake

surface, or by pitfall trapping.

NEW SPECIES AND A NEW GENUS OF THE SUBFAMILY POGONINAE FROM

SOUTH AUSTRALIA (COLEOPTERA: CARABIDAE)

M. BAEHR & P. HUDSON

BAEHR, M. & HUDSON, P. 2001. New species and a new genus of the subfamily Pogoninae

from South Australia (Coleoptera: Carabidae). Records of the South Australian Museum 34(1):

1-18.

A new genus and species, Syrdenoidius spinipes gen. nov., sp. nov., and three new species

of the genus Pogonus Dejean from salt lakes in South Australia are described: Pogonus

matthewsi, P. vicinus, and P. perovalis. The first two species are closely related to P. grossi

Moore from Lake Eyre and P. saskiae Baehr from Lake Gairdner and Island Lagoon and

together they form a complex of four very closely related species. Pogonus perovalis sp. nov.

is outstanding in its body shape and absence of wings, but may be remotely related to this

complex. The new genus Syrdenoidius is highly similar in shape and structure to the Palaearctic

genus Syrdenus Chaudoir, though it differs mainly in the asetose prosternum, the markedly

spinose tibiae, the absence of wings, and a very unusual female stylomere 2. Syrdenoidius

spinipes sp. nov. and Pogonus perovalis sp. nov. are outstanding in their absence of wings and

loss of flying ability, attributes that are extremely rare within Pogoninae. Most of the new

species were collected by excavation of their burrows on the lake surface, or by pitfall trapping.

Martin Baehr, Zoologische Staatssammlung, Miinchhausenstr. 21, D-81247 Miinchen,

Germany and Peter Hudson, Department of Environmental Biology, University of Adelaide,

South Australia 5005, Australia. Manuscript received 24 July 2000.

INTRODUCTION

Most halophile pogonine species occupy

habitats such as seashores and inland salt lakes

and most are strong fliers that readily come to

lights. Such habitats have been favoured

collecting localities in other parts of the world for

a very long time and there is a general belief that

the taxonomy of the subfamily is well known.

Nevertheless, several new species have been

recently described throughout the world

(Kryzhanovskij & Michailov 1971, Morvan 1973,

Kryzhanovskij 1990, Bousquet & Laplante 1997,

Baehr & Sciaky (in press), Hudson 2000). At

present, there are 11 pogonine genera worldwide

containing about 78 taxa. Pogonus s. str. is by far

the most speciose genus with 49 described taxa

(Lorenz 1998).

In Australia, the number of described pogonines

was surprisingly low for a very long time

(Chaudoir 1871, 1878; Sloane 1895). Moore

(1977) described new species from Lake Eyre and

since then a number of additional species have

been detected and described (Baehr 1984, 1997,

1999; Moore 1991, Hudson 2000) resulting in a

total of 12 species of Pogonus known from

Australia. It is now evident that all parts of

Australia suitable for pogonines are colonised by

at least one or another species. The salt lake belt

from interior Western Australia through the

southern half of South Australia to northwestern

Victoria (and probably westernmost New South

Wales) is the stronghold of pogonines in

Australia, with many closely related but also some

unusual species occurring there.

Four new species are described herein,

including a representative of an unusual new

genus. Although the pogonine fauna of Australia

comprises predominantly the genus Pogonus s.

str., it is one of the most speciose in the world

with high morphological diversity and includes

some very strangely shaped species.

The majority of the Australian pogonines are

depigmented, many are extremely elongate and

depressed (‘syrdeniform’), characters which are

evidence of an obligatory nocturnal and at the

same time psammophile life. Apart from this, very

little is known about habits, ecological

preferences, and life histories of most of the

Australian pogonine species, since most

collections have been made using light traps. It is

the scope of this paper, among other things, to

increase ecological information about the

Australian pogonines.

2 M. BAEHR & P. HUDSON

The present paper was prepared when the senior

author (MB) worked through the unidentified

carabid material in the South Australian Museum,

Adelaide, and separated a number of pogonine

specimens that turned out to belong to

undescribed species. They had been mostly

collected by the junior author (PH) during his

ample collecting work on salt lakes throughout

South Australia. Personal contact between MB

and PH during MB’s stay at the Museum revealed

that PH had a number of additional, freshly caught

specimens that were subsequently sent to MB for

identification. Since PH has collected many taxa

using pitfall traps and by excavation of burrows,

both authors agreed to collaborate in a joint paper,

with MB mainly responsible for the taxonomic

section, and PH for the ecological part.

Authorship of the new taxa is in joint names.

Some records of known species from the SAMA

collection and those made by MB during a recent

collecting trip through northwestern Victoria and

southern South Australia are also included, in

cases where they enlarge the known range or add

any other information.

MEASUREMENTS

Measurements were taken using a stereo

microscope with an ocular micrometer. Length has

been measured from apical margin of labrum to

apex of elytra; measurements, therefore, may

slightly differ from those of other authors. Length

of pronotum was taken at the longest distance

(which is not always along midline!), and width

between the posterior lateral angles.

LOCATION OF MATERIAL

The holotypes of the new species are located in

the South Australian Museum, Adelaide (SAMA),

paratypes of all species are kept in the working

collection of the senior author (CBM) at the

Zoologische Staatssammlung Miinchen. Paratypes

of Syrdenoidius spinipes are also kept in the

Australian National Insect Collection, Canberra

(ANIC).

TAXONOMIC PRINCIPLES

Whereas one of the three new taxa of the genus

Pogonus is very specialised in shape and structure

and certainly merits full specific status, two taxa

belong to a group of depigmented, elongate,

depressed, syrdeniform species that so far include

Pogonus grossi Moore and P. saskiae Baehr from

saline inland habitats in South Australia. The four

taxa now known to exist in this group are very

similar and certainly they are closely related. They

have been collected at four different localities

throughout the salt lake belt of South Australia,

and could be either subspecies of one widespread

species, or closely related species. We have decided

to rank them as species rather than subspecies for

two reasons. Firstly there appear to be consistent,

though minor differences, between them (see

Tables 1 and 2). Secondly there are heuristic

TABLE 1. Some distinguishing characters within the Pogonus grossi-group

grossi vicinus

(L. Eyre) (L. Frome)

body size large large

pronotum wide wide

basal angle slightly excised not excised

basal angle not dentiform dentiform

lateral margin

evenly convex

rather straight

anteriorly convex

widest in front

of anterior third

rather depressed

slightly convex

barely punctate

rather shallow

lateral margin widest behind

anterior third

elytra rather depressed

intervals slightly convex

striae distinctly punctate

striae rather deeply impressed

microreticulation _—_ distinct distinct

matthewsi

(Pernatty Lag.)

large

wide

not excised

dentiform

evenly convex

widest at

anterior third

convex

strongly convex

distinctly punctate

deeply impressed

very distinct

saskiae

(Island Lag.)

small

narrow

excised

not dentiform

evenly convex

widest at

anterior third

depressed

slightly convex

distinctly punctate

rather deeply impressed

distinct

NEW POGONINES FROM SOUTH AUSTRALIA 3

TABLE 2. Summary of measurements and ratios for all species of the Pogonus grossi-group. For better recognition

of the species the measurements and ratios for all species of the P. grossi-group are compiled in the following table.

body ratio ratio width ratio ratio ratio

length width/length base/apex width length/width width

(mm) pronotum pronotum pronotum/head elytra elytra/pronotum

grossi 5.2-6.4 1.18-1.21 0.91-0.93 1.13-1.15 1.85-1.91 1.27-1.35

vicinus 5.1-6.0 1.13-1.16 0.98-1.01 1.16-1.21 1.72-1.77 1.30-1.34

matthewsi 5.6-6.2 1.15-1.17 0.96-1.02 1.12-1.16 1.86-1.90 1.26-1.28

saskiae* = 4.65-4.90(4.15) 1.07-1.11(1.21) 0.87-0.88(0.85) 1.03-1.10(1.12) 1.96-1.98(1.90) —_—‘1.29-1.40

* In Pogonus saskiae Baehr one extraordinary small specimen has some rather different proportions of pronotum

and elytra (see Baehr 1997) that are added in brackets.

reasons for doing so since definition of subspecies

is more difficult than that of species and would

require more information about gene flow or hybrid

zones. Further investigations should show which

concept is better applicable to this difficult group.

TAXONOMY

There is no need for an _ extensive

characterization of the subfamily Pogoninae (or

tribe Pogonini, according to the preferred system).

A list of synonymies including the most important

citations, and an extensive diagnosis of the

subfamily was recently given by Bousquet and

Laplante (1997) that should be used by readers

who want to receive more general information

about the group. A summary of the presently

known genera and species has been recently

provided by Lorenz (1998). More information

about the Australian Pogonus species can be

obtained from Moore (1977), and the subsequent

descriptions of additional species by Moore

(1991), Baehr (1984, 1997, 1999), and Hudson

(2000).

ADDITIONAL RECORDS OF DESCRIBED SPECIES

Pogonus grossi Moore

Moore 1977: 63; Moore et al. 1987: 146; Baehr

1997: 3; 1999: 147.

This species is only known from the type

locality Lake Eyre (Moore et al. 1987). In the

SAMA collection there is a large series of

specimens from Madigan Gulf and Prescott Gulf

(both Lake Eyre North). Thus, P. grossi seems to

have colonised a large part of the shores of this

lake.

Pogonus hypharpagioides Sloane

(Fig. 1)

Sloane 1895: 126; Moore 1977: 63; Moore et al.

1987: 146; Baehr 1997: 2, 3; 1999: 147.

In addition to its type locality, Lake Callabonna,

P. hypharpagioides has previously been recorded

from Lake Eyre (Moore et al. 1987) and Lake

Frome (Baehr 1997). Collections made by the

authors show this species is widely distributed

throughout the salt lake belt from northwestern

Victoria to the west coast of South Australia,

including some near-coastal localities (see Fig. 1).

A variety of techniques have been employed in

collecting this species; many specimens were

excavated from burrows, others were collected at

light, in pitfall traps or while running on the

surface at night.

Pogonus variabilis Moore

Moore 1991: 31; Baehr 1997: 2; 1999: 147.

In the collection of SAMA there is a series of

specimens caught by N. B. Tindale and P. Aitken

in 1963 at Normanton and on Mornington Island,

both northwestern Queensland, that apparently

escaped the attention of Moore when he described

this species in 1991. They are evidence of the

wide range of this species along the coasts of

tropical Australia. Characteristically, they have

been collected at light.

DESCRIPTIONS OF NEw SPECIES

Pogonus matthewsi sp. nov.

(Figs 2, 5, 8, 19)

Types

Holotype: male, P. Hudson. 3 Sept. 1996,

4 M. BAEHR & P. HUDSON

Kilometres

FIGURE 1. Distribution of Pogonus hypharpagioides Sloane, previous records (@), new records (@).

Pernatty Lagoon (sth), SA 31.6625°S 137.2375°E

(SAMA). Paratypes: 5 females, same data (CBM,

SAMA).

Diagnosis

Moderately small, elongate, uniformly reddish

‘syrdeniform’ species, distinguished from related

species mainly by its convex elytra with distinct

microreticulation and markedly convex intervals,

and its comparatively stout legs with wide

metafemur and strongly depressed tibiae. Further

distinguishing characters are in Tables 1 and 2.

Description

Measurements: Length: 5.6-—6.2 mm; width:

1.9-2.0 mm. Ratios: Width/Mength of pronotum:

1.15-1.17; width base/apex of pronotum: 0.96—

1.02; width of pronotum/width of head: 1.12-

1.16; length/width of elytra: 1.86-1.90; width

elytra/pronotum:1.26—1.28. Elytral ratios are only

based on two specimens since all are weakly

sclerotised and most have their elytra opened and

somewhat distorted.

Colour: All known specimens freshly hatched,

hence mature colouration somewhat uncertain.

Upper and lower surfaces including mouth parts,

antennae, and legs yellow. Apex of mandibles dark.

Head (Fig. 2): Rather large and wide, fairly

convex, slightly narrower than pronotum. Eyes

rather small, depressed, laterally not much

projecting, posteriorly slightly enclosed. Labrum

short and wide, apex gently emarginate.

Mandibles rather elongate, apex fairly incurved.

Palpi of moderate size, penultimate palpomere of

labial palpus fairly slender, slightly bowed.

Submentum quadrisetose, tooth of mentum wide,

slightly cleft, bidentate. Lacinia with elongate

spines. Clypeus and frons in middle convex,

frontal furrows fairly short, sinuate, rather

shallow, posteriorly ending in an irregularly

shaped impression. Impressions with irregular

wrinkles. Antennae short, well short of reaching

base of pronotum, median antennomeres little

longer than wide, sparsely pilose from middle of

3 antennomere, more densely pilose from 5"

antennomere. Surface rather glossy, impunctate,

NEW POGONINES FROM SOUTH AUSTRALIA 5

FIGURES 2-4. Habitus. 2. Pogonus matthewsi sp. nov. 3. Pogonus vicinus sp. nov. 4. Pogonus perovalis sp. nov.

Lengths: 5.6 mm; 6.0 mm; 5.5 mm.

microreticulation rather superficial, approximately

isodiametric.

Pronotum (Fig. 2): Slightly wider than long,

moderately convex, not cordiform, widest at

anterior third slightly behind position of anterior

lateral seta. Base about as wide as apex. Apex

well produced beyond anterior angles, convex, in

middle emarginate. Anterior angles very sharply

rounded off, almost rectangular. Lateral border

evenly convex throughout, with a very short

concavity just in front of basal angles. Marginal

channel narrow, border slightly upturned. Basal

angles rectangular, dentiform, base convex,

laterally rather oblique. Both apex and middle of

base unmargined. Anterior transverse sulcus fairly

distinct, median line very shallow, attaining base

though not apex, basal transverse sulcus shallow

though distinct. Basal grooves circular, merging

into marginal channel. Anterior lateral seta

situated in front of anterior third and of widest

diameter, posterior marginal seta arising from

basal angle. Both apex and base rugosely

punctate-striolate. Disk very finely punctate or

almost impunctate, at least laterally with more or

less distinct, somewhat isodiametric

microreticulation, with very faint, irregularly

transverse strioles, glossy.

Elytra (Fig. 2): Elongate, narrow, rather

parallel, dorsally remarkably convex, widest at

middle. Humeri very obtusely angulate. Lateral

margin straight over most of its distance, little

narrowed towards humeri, contiguous at sutural

angle. Marginal channel extremely narrow, margin

not explanate. All striae complete, well impressed,

coarsely punctate. Eighth stria not bowed away

from margin. Basal margin evenly curved into

short scutellar stria medially of 1“ stria. Intervals

rather convex. Third interval with 4, rarely

6 M. BAEHR & P. HUDSON

unilaterally 3 setiferous punctures, the anterior

and median ones adjacent to 3" stria, situated

about at basal third and at middle, the third and

fourth ones in posterior third attached to 2" stria.

11-12 widely spaced submarginal punctures in a

more or less uninterrupted row attached to 8"

stria. Scutellar pore and seta present, at base of 1*

stria. Intervals impunctate, with very conspicuous,

isodiametric microreticulation. Wings fully

developed.

Lower surface: Prosternum not pilose.

Metepisternum rather elongate, slightly more than

half longer than wide at apex. Terminal sternum

in male bisetose, in female quadrisetose.

Legs: Rather short and stout, metafemur

comparatively stout, tibiae strongly depressed,

with sparse and weak spination. Tibial spurs

moderately elongate. Tarsi rather short, 1*

tarsomere of metatarsus much shorter than both

following tarsomeres. Metatrochanter in both

sexes short, wide, a third of length of metafemur,

apex obtusely rounded. Tarsomeres 1-3 of male

protarsus slightly asymmetrically widened on

median side, 1“ tarsomere biseriately squamose,

2™ tarsomere with few squamae on median border

only.

Male genitalia (Fig. 5): Genital ring rather

regularly triangular, little sclerotised, because the

holotype is freshly hatched. Aedeagus small, short

and compact, laterally depressed, slightly

asymmetric, lower surface regularly though gently

curved, apex rather short, obtuse at tip, slightly

turned to right side. Internal sac with a coiled,

circular, sclerotised plate near base, and with a

distinctly denticulate, triangular fold near apex.

Right paramere narrow, slightly shorter than left,

with 3 elongate apical setae. Left paramere large,

convexly tapering to apex, with 2 elongate and 1

short apical setae, the short one situated below the

longer ones.

Female genitalia (Fig. 8): Stylomere 1 without

setae at apex. Stylomere 2 elongate, curved, with

1 or 2 small ventral ensiform setae near base, and

2 (attached) nematiform setae originating from a

groove near apex.

Variation: Only some variation of relative

length of elytra and width of pronotum noted.

Distribution (Fig. 19)

So far recorded only from Pernatty Lagoon,

slightly west of Lake Torrens, South Australia.

Habits

Specimens were collected following excavation

of their burrows. They were widely distributed on

parts of the lake surface lacking a salt crust.

Etymology

The name is a patronym in honour of Eric

Matthews, Curator of Coleoptera at the South

Australian Museum who kindly gave access to the

rich collections of pogonines.

Relationships

The species belongs to the grossi-complex, and

it appears to be most closely related to P. grossi

Moore and P. vicinus sp. nov.

Pogonus vicinus sp. nov.

(Figs 3, 6, 19)

Types

Holotype: male, P. Hudson 9 July 1997 SEG

Exp. Lake Frome 30.6667°S 139.6028°E

(SAMA). Paratypes: 1 male, P. Hudson Jul 1997

pit trap, Lake Frome, SA 30.6667°S 139.6028°E

(CBM); 1 male, G. Medlin 18 May, 1997 Lake

Frome, SA 30.6667°S 139.6028°E (SAMA).

Diagnosis

Moderately small, elongate, uniformly reddish

‘syrdeniform’ species, distinguished from related

species mainly by its rather short and wide elytra

with shallow, barely punctate striae, and the rather

straight though anteriorly abrupt convex lateral

margins of pronotum. Distinguishing characters

are summarised in Tables 1 and 2.

Description

Measurements: Length: 5.1-6.0 mm; width:

1.85—2.10 mm. Ratios: Width/length of pronotum:

1.13-1.16; width base/apex of pronotum: 0.98—

1.01; width of pronotum/width of head: 1.16-

1.21; length/width of elytra: 1.72—1.77; width

elytra/pronotum: 1.30—1.44.

Colour: Upper and lower surfaces including

mouth parts, antennae, and legs dark yellow. Apex

of mandibles dark.

Head (Fig. 3): Rather large and wide, fairly

convex, slightly narrower than pronotum. Eyes

rather small, fairly depressed, laterally not much

projecting, posteriorly slightly enclosed. Labrum

short and wide, apex gently emarginate.

Mandibles moderate, apex fairly incurved. Palpi

of moderate size, penultimate palpomere of labial

palpus fairly slender, slightly bowed. Submentum

quadrisetose, tooth of mentum wide, slightly cleft,

bidentate. Lacinia with elongate spines. Clypeus

and frons convex in middle, frontal furrows fairly

NEW POGONINES FROM SOUTH AUSTRALIA 7

6

7

FIGURES 5-7. Male genitalia: Aedeagus, left side, parameres, genital ring. 5. Pogonus matthewsi sp. nov. 6.

Pogonus vicinus sp. nov. 7. Pogonus perovalis sp. nov. Scales: 0.5 mm.

8 M. BAEHR & P. HUDSON

short, sinuate, fairly shallow, posteriorly ending in

an irregular impression. Antennae rather short, not

attaining base of pronotum, median antennomeres

about a third longer than wide, sparsely pilose

from middle of 3% antennomere, more densely

pilose from 5" antennomere. Surface glossy,

impunctate, without microreticulation.

Pronotum (Fig. 3): Slightly wider than long,

rather depressed, not cordiform, widest at anterior

third slightly behind position of anterior lateral

seta. Base about as wide as apex. Apex slightly

produced beyond anterior angles, convex,

emarginate in middle. Anterior angles very sharply

rounded off, almost rectangular. Lateral border

anteriorly convex or even almost oblique, then

oblique and straight, just in front of basal angles

incurved and forming a very small concavity.

Marginal channel narrow, posteriorly slightly

widened, border slightly upturned. Basal angles

rectangular, very faintly dentiform, base convex,

laterally rather oblique. Both apex and middle of

base unmargined. Anterior transverse sulcus fairly

distinct, median line very shallow, attaining base

though not apex, basal transverse sulcus

moderately shallow, distinct. Basal grooves

circular, merging into marginal channel. Anterior

lateral seta situated in front of anterior third and

point of greatest width, posterior marginal seta

arising from basal angle. Apex and base rugosely

punctate-striolate. Disk impunctate, without

microreticulation, with very faint, irregularly

transverse strioles, glossy.

Elytra (Fig. 3): Comparatively short and wide

for group, moderately narrow, not parallel,

dorsally rather depressed, widest at middle.

Humeri very obtusely angulate or almost rounded.

Lateral margin gently convex over most of its

distance, slightly narrowed towards humeri,

contiguous at sutural angle. Marginal channel

extremely narrow, margin not explanate. All striae

complete, not much impressed, almost

impunctate. Eighth stria not bowed away from

margin. Basal margin evenly curved into short

scutellar stria medially of 1 stria. Intervals

depressed. Third interval with 4, rarely unilaterally

3 or 5 setiferous punctures, the anterior and

median ones adjacent to 3" stria, situated about at

basal third and at middle, the third and fourth ones

in posterior third attached to 2™ stria. 10-11

widely spaced submarginal punctures in a more or

less interrupted row attached to 8" stria. Scutellar

pore and seta present, at base of 1‘ stria. Intervals

impunctate, with fairly superficial, isodiametric

microreticulation. Wings fully developed.

Lower surface: Prosternum not pilose.

Metepisternum rather elongate, about a half longer

than wide at apex. Terminal sternum in males

bisetose, unknown in females.

Legs: Rather short and stout, metafemur

moderately stout, tibiae fairly well depressed, with

sparse and weak spination. Tibial spurs

moderately elongate. Tarsi rather short, 1"

tarsomere of metatarsus much shorter than both

following tarsomeres. Metatrochanter in males

short, wide, a third of length of metafemur, apex

obtusely rounded. Tarsomeres 1-3 of male

protarsus slightly asymmetrically widened on

median side, 1“ tarsomere biseriately squamose,

2™ tarsomere with few squamae on median border

only.

Male genitalia (Fig. 6): Genital ring rather

regularly triangular. Aedeagus small, short and

compact, laterally depressed, slightly asymmetric,

lower surface regularly though gently curved,

apex rather short, obtuse at tip, slightly turned to

right side. Internal sac with a coiled, circular,

sclerotised plate near base, and with a distinctly

denticulate, triangular fold near apex. Right

paramere narrow, slightly shorter than left, with 3

elongate apical setae. Left paramere large,

comparatively elongate, tapering to apex, with 3

elongate apical setae.

Female genitalia: Unknown.

Variation: Only some variation of relative width

of pronotum noted.

Distribution (Fig. 19)

So far recorded only from Lake Frome,

southeast of Lake Eyre, South Australia.

Habits

Excavated from burrows in the lake surface and

collected by pitfall trapping.

Etymology

The name refers to the strong similarity of this

species to the related P. grossi Moore and P.

matthewsi sp. nov.

Relationships

The species belongs to the grossi-complex,

where it apparently takes a position near P. grossi

Moore and P. matthewsi sp. nov.

Pogonus perovalis sp. nov.

(Figs 4, 7, 9, 10, 19)

Types

Holotype: male, P. Hudson (SEG Exp 95) Jul

NEW POGONINES FROM SOUTH AUSTRALIA 9

FIGURES 8-9. Female stylomeres 1 and 2. 8. Pogonus

matthewsi sp. nov. 9. Pogonus perovalis sp. nov.

Scales: 0.1 mm.

1995 pit traps Lake Gairdner, SA 32.3014°S

135.8867°E (SAMA). Paratypes: 2 males, 1

female, same data (CBM, SAMA); 1 female, P.

Hudson 3 Sep 1996 Island Lagoon, SA 31.4139°S

136.9292°E (SAMA); | female, Remko Leijs 30

Jun 1996 Lake Hart, SA 31.2278°S 136.3792°E

(SAMA).

Diagnosis

Rather small, convex, apparently depigmented

species, at once distinguished by the oval-shaped,

convex elytra, convex, rather cordiform pronotum,

and elongate antennae.

Description

Measurements: Length: 4.35-5.70 mm; width:

1.60—2.10 mm. Ratios: Width/length of pronotum:

1.03-1.12; width base/apex of pronotum: 0.88—

0.96; width of pronotum/width of head: 1.10—

1.18; length/width of elytra: 1.49-1.53; width

elytra/pronotum:1.49—1.56. Elytral ratios are only

based on a few specimens since all are weakly

sclerotised and most have their elytra opened and

somewhat distorted.

Colour: Apparently depigmented, though all

known specimens freshly hatched, hence

colouration somewhat uncertain. All specimens

very light yellowish. Only apex of mandibles dark.

Head (Fig. 4): Fairly large, rather depressed,

slightly narrower than pronotum. Eyes large,

rather convex, laterally well projecting, posteriorly

not enclosed. Labrum short and wide, apex

straight. Mandibles very elongate, somewhat

decussate, apex incurved. Palpi elongate, apical

palpomeres somewhat incised, slightly subulate.

Penultimate palpomere of labial palpus slender,

slightly bowed. Submentum bisetose, tooth of

mentum wide, slightly cleft, bidentate. Lacinia

with elongate spines. Clypeus and frons convex in

middle, frontal furrows fairly elongate, rather

sinuate, fairly shallow, at posterior end forming

an irregular impression. Irregular wrinkles within

and between impressions. Antennae very elongate,

surpassing base of pronotum by almost 4

antennomeres, median antennomeres > 3 x as long

as wide, pilose from middle of 3 antennomere.

Surface glossy, impunctate, without any

microreticulation.

Pronotum (Fig. 4): Slightly wider than long,

convex, rather cordiform, widest at anterior third

approximately at position of anterior lateral seta.

Base slightly narrower than apex. Apex produced

beyond anterior angles, feebly convex. Anterior

angles sharply rounded off. Lateral border evenly

convex in anterior two thirds, then gently concave

to basal angles. Marginal channel extremely

narrow, border slightly upturned, forming a sharp

edge. Basal angles rectangular, base slightly

convex. Apex and base unmargined. Anterior

transverse sulcus barely indicated, median line

very shallow, attaining base though not apex,

basal transverse sulcus very shallow. Basal

grooves shallow, circular, merging into marginal

channel. Anterior lateral seta situated at anterior

third, at point of greatest width, posterior marginal

seta arising from basal angle. Both apex and base

rugosely punctate-striolate. Disk impunctate and

without microreticulation, highly glossy, with

some very faint, transverse strioles.

Elytra (Fig. 4): Short, markedly oval-shaped,

convex, widest at or slightly behind middle.

Humeri angulate, more or less rectangular. Lateral

margin evenly convex throughout. Marginal

channel narrow, margin not explanate. All striae

complete, well impressed, punctate. Eighth stria

originating at basal fifth, slightly bowed away

from margin. Basal margin evenly curved into

short scutellar stria medially of 1* stria. Intervals

depressed to very feebly convex. Third interval

with 3 setiferous punctures, the anterior and

median ones adjacent to 3" stria, situated at about

basal third and at middle, the third puncture in

posterior third attached to 2™ stria. Nine, rarely 10

10 M. BAEHR & P. HUDSON

10

FIGURE 10. Pogonus perovalis sp. nov. Right protibia

and tarsus, dorsolateral view.

widely spaced submarginal punctures in an

uninterrupted row attached to 8" stria. Scutellar

pore and seta present, at base of 1“ stria. Intervals

impunctate, though with very distinct, absolutely

isodiametric microreticulation. Wings absent.

Lower surface: Prosternum not pilose.

Metepisternum very short, shorter than wide at

apex. Terminal sternum in male bisetose, in

female quadrisetose.

Legs (Fig. 10): Remarkably slender and

elongate, especially femora and tibiae. Tibiae not

at all depressed, almost circular, with sparse and

weak spination. All tibial spurs elongate. Tarsi

rather short, 1% tarsomere of metatarsus shorter

than both following tarsomeres. Metatrochanter in

both sexes short, wide, a quarter of length of

metafemur, apex obtusely rounded. Tarsomeres 1—

3 of male protarsus slightly asymmetrically

widened on median side, 1 tarsomere biseriately

squamose, 2™ tarsomere with few squamae on

median border only.

Male genitalia (Fig. 7): Genital ring rather

regularly triangular. Aedeagus comparatively

large, fairly elongate, laterally depressed, slightly

asymmetric, lower surface almost straight, in

apical third markedly concave. Apex laterally very

depressed, scuriform, slightly turned to right side.

Internal sac with a coiled, circular, sclerotised

plate near base, and with a distinctly denticulate,

triangular fold near apex. Right paramere very

slender and elongate, slightly shorter than left,

with one very elongate and two shorter apical

setae, the elongate situated in middle. Left

paramere large, comparatively elongate, tapering

to apex, with 1 very elongate apical seta and a

short, delicate seta below.

Female genitalia (Fig. 9): Stylomere 1

apparently without setae at apex. Stylomere 2

moderately elongate, barely curved, with wide,

obtuse apex, without ventral ensiform setae, with

2 (attached) nematiform setae originating from a

groove very close to apex.

Variation: Apart from some differences in size,

little variation noted.

Distribution (Fig. 19)

Recorded from Lake Gairdner, Island Lagoon,

and Lake Hart, all South Australia.

Habits

Collected in pitfall traps on the lake surface.

Etymology

The name refers to the markedly ovate elytra of

this flightless species.

Relationships

Phylogenetically rather isolated species, but

may be remotely related to the P. grossi species

complex.

Syrdenoidius gen. nov.

Type species: Syrdenoidius spinipes sp. nov., by

monotypy.

Diagnosis

Genus of subfamily Pogoninae. In many

respects highly similar to the Palaearctic genus

Syrdenus Chaudoir, e.g. narrow, elongate,

depressed body shape; absence’ of

microreticulation on elytra; markedly angulate,

even dentate humeri; wide 9" elytral interval;

short, quadrate antennomeres. However, there are

some striking pecularities in Syrdenoidius: asetose

prosternum; presence of several rows of

remarkably elongate spines on all tibiae

(particularly conspicuous on protibia), that occupy

almost the whole length of the tibiae; prolongation

of 8" elytral stria to humerus; absence of scutellar

NEW POGONINES FROM SOUTH AUSTRALIA 11

striole and pore; absence of hind wings, and

quadrate metepisternum; very elongate, forceps-

like mandibles; very unusual female genitalia with

extremely short, straight stylomere 2.

Etymology

The name refers to the strong similarity in

external morphology to the genus Syrdenus.

Relationships

Whereas the similarity of some characters in

Syrdenus and Syrdenoidius is probably

coincidental, several could point to a closer

relationship of both genera. This is especially true

for the wide 9" elytral interval that bears on its

median margin the marginal pores and setae that

are, therefore, far removed from the lateral margin

of the elytra. On the other hand, elongate,

depressed body shape is an adaptation common

within several pogonine genera, not only in

Syrdenus, but also in Pogonus proper and in

Pogonistes. We think that the genus Syrdenoidius

is a highly isolated one though it possibly has

closest affinities with Syrdenus. Whilst Syrdenus is

so far decidedly a Palaearctic genus there is,

however, an as yet undescribed species known

from West Bengal, northeastern India, that

demonstrates that the genus has a wider distribution

than commonly believed. If the genus Syrdenus is a

relict of the former Tethys fauna, then a relationship

with the fauna of western and northwestern Australia

might well be possible since it has been argued that

this area formerly had contact with the Tethys or

was even part of its southern shore (Eric Matthews,

1998, pers. com.). This question may be settled in

the future by more intensive sampling along the

coast and on the inland lakes of Western Australia.

Syrdenoidius spinipes sp. nov.

(Figs 11-19)

Types

Holotype: male, S. AUST. L. Gairdner 32°07‘S

135°53‘E 14 July 95 P. Hudson (SAMA).

Paratypes: 6 males, 13 females, same data (ANIC,

CBM, SAMA); 1 female, P. Hudson 14 July 1995

SEG Exp. 95 Lake Gairdner 32°07‘47"S

135°53‘19"E (SAMA).

Diagnosis

Medium-sized, very elongate, narrow,

depressed, depigmented species with large,

convex head, large though depressed eyes, very

elongate mandibles, almost regularly trapezoidal

pronotum that is widest at apex, non-

microreticulate surface, and remarkably spinose

tibiae.

Description

Measurements: Length: 5.4-6.3 mm; width:

1.50-1.65 mm. Ratios: Width/length of pronotum:

1.03-1.08; width base/apex of pronotum: 0.73—

0.77; width of pronotum/width of head: 1.04—

1.07; length/width of elytra: 2.06—-2.12; width

elytra/pronotum:1,12-1.16.

Colour: Upper and lower surface, including

mouth parts, antennae, and legs light reddish,

elytra very slightly lighter. Apex of mandibles

darker.

Head (Figs 11, 16-18): Large, convex, slightly

narrower than pronotum. Eyes large though

remarkably depressed, laterally barely projecting,

posteriorly not enclosed. Lower rim of eye

forming a sharp edge that slightly surpasses the

eye laterally. Labrum short and wide, apex fairly

emarginate. Mandibles very elongate, somewhat

decussate, apex strongly incurved. Palpi elongate,

apical palpomeres somewhat incised, slightly

subulate. Penultimate palpomere of labial palpus

very slender, slightly bowed. Submentum

sexsetose, tooth of mentum wide, deeply cleft,

markedly bidentate (Fig. 16). Lacinia with very

elongate spines. Ligula bisetose, paraglossae

slightly surpassing glossa (Fig. 17). Clypeus and

frons convex in middle, frontal furrows elongate,

slightly sinuate, beginning on clypeus. Frons

behind frontal furrows with a wide, circular,

shallow impression on either side, irregular

wrinkles between impressions. Antennae short,

wide, not attaining base of pronotum, median

antennomeres almost as wide as long, sparsely

pilose from middle of 3 antennomere, more

densely pilose from 5™ antennomere. Surface

glossy, impunctate, without any microreticulation.

Pronotum (Figs 11, 18): Barely wider than long,

remarkably trapezoidal, widest at apex. Surface

moderately convex. Apex little produced beyond

anterior angles, in middle with very shallow,

triangular excision. Anterior angles broadly

rounded. Lateral margin almost straight, oblique,

slightly rounded only near apex, faintly incurved

towards base. Border somewhat irregular, bearing

several slight notches. Basal angles obtusely

rounded, base almost straight. Marginal channel

extremely narrow, margin forming a fairly sharp

ridge. Apex unmargined, base coarsely margined,

margin narrowly surrounding basal angles and

meeting lower margin of epipleura, this margin

12 M. BAEHR & P. HUDSON

15

FIGURES 11-15. Syrdenoidius spinipes gen. nov., sp. nov. 11. Habitus. Length 5.6mm. 12. Female stylomeres 1

and 2. Scale 0.1mm. 13. Right protibia and tarsus, dorsal view. 14. right protibia, dorsolateral view. Scale: 0.5 mm.

15. Male genitalia: Aedeagus, left side, parameres, genital ring. Scale 0.5mm.

NEW POGONINES FROM SOUTH AUSTRALIA 13

| |

wy

16

17

FIGURES 16-18. Syrdenoidius spinipes gen. nov. sp. nov. 16. Mentum and submentum, ventral view. Scale: 0.5

mm 17. Ligula, ventral view. Scale: 0.1 mm. 18. Base of left elytron. Scale 0.5 mm.

sometimes even visible from above (Fig. 18).

Anterior transverse sulcus barely indicated,

median line very shallow, deepened near base,

attaining apex and base. Basal transverse sulcus

shallow though distinct. Base coarsely striolate,

basal grooves rather shallow, irregular. Anterior

marginal seta inserted at anterior fourth, posterior

seta situated just in front of basal angle, anterior

and posterior setae slightly removed from margin.

Disk impunctate and without any

microreticulation, highly glossy, though laterally

with some transverse striolations.

Elytra (Figs 11, 18): Very narrow and elongate,

parallel-sided, markedly depressed. Humeri very

sharply projecting, markedly angulate. Lateral

margin gently oblique or slightly convex in basal

fourth, then parallel, only in apical fourth convex

again. Marginal channel narrow throughout.

Apical angles broadly rounded, apex in middle

slightly incised. No internal plica visible from

outside. Striae complete, deep, not punctate, but

somewhat irregular, sometimes interrupted or with

anastomoses. Eighth stria bowed away from

margin in anterior fourth, laterally leaving a wide

space, though anteriorly attaining humerus. Basal

margin evenly curved into 1™ stria, no scutellar

stria present. Scutellar pore and seta absent.

Intervals convex, though their surface rather

uneven. Third interval with 3 setiferous punctures

adjoining 3" stria, the first situated in anterior

third, the second in middle, the third in apical

third, but their position rather variable.

Submarginal punctures far removed from margin,

adjoining lateral margin of 8" stria, consisting of

4 punctures in anterior group and 6 punctures in

posterior group, with a wide space between them.

Also with 1 puncture at end of 5" interval and 2™

interval. Surface impunctate, without

microreticulation, highly glossy. Wings almost

completely reduced.

Lower surface: Prosternum not pilose.

Metepisternum short, barely longer than wide at

apex. Terminal sternum in male bisetose, in

female quadrisetose.

Legs (Figs 13, 14): Moderately elongate.

Protibia and mesotibia with 4 rows of remarkably

stout and elongate spines all along dorsal, lateral,

and latero-ventral surfaces. Metatibia less spinose.

Apical tibial spurs of protibia and mesotibia

remarkably elongate, much longer than 1*

tarsomere. Tarsi rather short, 1 tarsomere of

metatarsus shorter than both following tarsomeres.

Tarsi remarkably hirsute. Tarsomeres 1-3 of male

protarsus slightly asymmetrically widened on

median side, 1° and 2™ tarsomeres with few

squamae on median border. Metatrochanter in

both sexes less than half as long as metafemur,

obtuse at apex.

Male genitalia (Fig. 15): Genital ring wide, in

basal half almost quadrangular, apically regularly

triangular, rather symmetric. Aedeagus short and

very compact, laterally depressed, slightly

asymmetric, lower surface barely curved, apex

very short, obtuse, barely turned laterally. Internal

sac with a complexly coiled, in parts heavily

sclerotised plate near base, and with a distinctly

denticulate, triangular fold near apex. Right

paramere moderately narrow, about a third shorter

than left, with 3 elongate apical setae. Left

paramere elongate, rather slender, abruptly

narrowed to the downcurved apex, with 3 elongate

apical setae.

14 M. BAEHR & P. HUDSON

Female genitalia (Fig. 12): Stylomere 1 2. — _ Elytra with distinct microreticulation,;

apparently without setae at apex. Stylomere 2 very humerus not so angulate (Figs 2-4);

short, straight, almost quadrate, with wide, convex protibia with a short row of moderate

apex, without ensiform setae, though with an spines on lateral surface only (Fig. 10);

elongate nematiform seta arising from apex. pronotum differently shaped, widest far

Variation: Very little variation noted. behind anterior angles (Figs 2-4). Widely

distributed in Australia... ccc

Distribution Agog) scat Radececdeahoongacetebogeete? Pogonus Dejean

Only recorded from Lake Gairdner, South

Australia.

Habits Key To THE AUSTRALIAN SPECIES OF THE GENUS

Excavated from burrows in the lake surface. Poconus DEJEAN

Etymology 1. — Body completely metallic green or

The name refers to the remarkably spinose LACK... 44.40 detovh Meatie Mett ste dhol. 2

tibiae of this species. — Body entirely or in parts testaceous ..... 5

2. — Colour shining black; punctures of elytra

very coarse, microreticulation in-

conspicuous. Northeastern Queensland,

coastal and inland ...... nigrescens Baehr

— Colour greenish metallic; punctures of

elytra finer, microreticulation conspicuous.

Key To THE GENERA OF AUSTRALIAN POGONINAE

1. — Elytra without microreticulation; humerus

sharply angulate (Figs 11, 18); protibia with

several rows of elongate spines on dorsal

and lateral surfaces that occupy almost the Southern half of Australia ............0.0. 3

whole length (Figs 13, 14); pronotum 3. — Sixth and 7" striae almost similar to inner

remarkably trapezoidal, widest at anterior 5 striae; flightless; elytral margins ovate.

angles (Fig. 4). Lake Gairdner, SA........... Lake Yindarlgooda, Western Australia .

Syrdenoidius gen. nov. (only S. spinipes sp. sresteseseeeneeseeseeeeteetsesersetees fennelli Hudson

nov.) — Sixth and 7" striae less marked than 5

ny

Fike Fat, a )

a wy se

-

FIGURE 19. Distribution of Pogonus matthewsi sp. nov. (W), P. vicinus sp. nov. (A), P. perovalis sp. nov. (@),

Syrdenoidius spinipes gen. nov., sp. nov. (*).

NEW POGONINES FROM SOUTH AUSTRALIA 15

inner striae; fully winged; elytral margins

more parallel sided ............::cceseeeeee 4

Smaller (body length 6.3-6.8 mm), body

narrower, more convex; pronotum

markedly sinuate in front of the acute

basal angles, these being 90° or less;

submarginal basal ridge of pronotum

scarcely indicated. Southern Australia from

Western Australia to Victoria along coast

satecntetaded@eshezitemretngs australis Chaudoir

Larger (body length 7.5-8.5 mm), body

wider, less convex; pronotum less sinuate,

basal angles more obtuse, these being c.

100°; submarginal basal ridge of pronotum

conspicuous. Coastal and inland from

southern half of Western Australia to

southeastern Queensland ............eseeeeee

WTtanedadelinwa dontetee cardiotrachelus Chaudoir

Bicoloured, head and pronotum distinctly

darker than elytra 00.0... cseccesseeeeseseeee 6

Completely testaceous or light reddish,

head and pronotum not perceptibly darker

than elytra (variable species under both

0111 0) (316) De ne 7

Large, convex species (body length 9-11

mm); pronotum not lobate, distinctly

sinuate in front of basal angles. Northern

Australia from northwestern Queensland

to the Kimberleys, coastal and along tidal

TIVETS ssisscesoussngesgapandvads variabilis Moore

Small, depressed species (body length c.

6.5 mm); pronotum lobate, barely sinuate

in frontofbasal angles. Inland saline habitats

in Western Australia and South Australia

watagecodat auch is iapweHhesseerseatdiacaee zietzi Sloane

Large species (body length >9 mm); either

pronotum distinctly sinuate in front of

basal angles and base about as wide as

apex, or pronotum rather quadrate with

characteristically bisinuate base, and elytra

wide, oval-shaped, and depressed with

wide, explanate lateral margin. Northern

Australia from northwestern Queensland

to Exmouth Gulf, coastal and along tidal

Smaller species (body length <8.5 mm);

pronotum distinctly sinuate or not in front

of basal angles; but when distinctly sinuate,

then base markedly narrower than apex;

when not sinuate, then elytra not wide,

oval-shaped, and depressed; and with

narrower, not explanate lateral margin.

Inland in southern half of Australia..... 9

8.

10.

11.

12.

Pronotum distinctly sinuate in front of

basal angles, lateral margin not explanate

at base; eyes convex, laterally markedly

protruding; elytra less wide and depressed,

lateral margin not explanate, striation

deeper and more complete, striae distinctly

punctate. Northern Australia from

northwestern Queensland to the

Kimberleys ............005 variabilis Moore

Pronotum rather quadrate, lateral margin

conspicuously explanate at base; eyes

depressed, laterally little protruded; elytra

wide and depressed, lateral margin

explanate, striation shallow and laterally

incomplete, striae more finely punctate.

Coast of northwestern Australia south of

Great Sandy Desert.......... sumlini Baehr

Pronotum distinctly sinuate in front of

basal angles and base markedly narrower

than apex; left paramere with 2 apical

setae, right paramere with a single apical

seta. South Australia, Lake Eyre Basin..

adhe ge lh Gan ditsl Lavt aacics) Raptcatceh cual gilesi Moore

Pronotum either not distinctly sinuate in

front of basal angles or base about as wide

as apex; left paramere with 3, right

paramere with 2 or 3 apical setae ...... 10

Large, convex species (body length 7.2-

8.4 mm); head large, pronotum laterally

markedly convex. South Australia,

northwestern Victoria 0.0... eeeeseeeeeteeee

hi ht tncatoute'cy dels hypharpagioides Sloane

Smaller, more depressed species (body

length <6.5 mm); head smaller, pronotum

laterally less CONVEX .........:.:ccesseeeseees 11

Pronotum rather quadrate, lateral margin

evenly curved from apex to base, widest in

middle; elytral striae shallow, only 3 inner

striae distinct. Interior of Western Australia

doudidaderrdioeledhgpeaday diplochaetoides Baehr

Pronotum more narrowed to base than to

apex, widestin anterior third, lateral margin

not so evenly curved; elytral striae deeper,

at least 5 inner striae distinct. South

AUStra liao). isctectagtdeitectues coseesgactoa tes 12

Elytra laterally and dorsally convex,

oviform; pronotum narrow, convex, with

elongate sinuation in front of basal angles

(Fig. 4); antenna elongate, median

antennomeres >3 x as long as wide. Lake

Gairdner, Island Lagoon, Lake Hart ......

Sac iiitaibtetbaabecgvenshadarsete perovalis sp. nov.

Elytra elongate, parallel, dorsally

16 M. BAEHR & P. HUDSON

depressed; pronotum wide, depressed, with

at most a short sinuation in front of basal

angles (Figs 2, 3); antenna short, median

antennomeres <1.5 x as long as wide ....

ablidiveblencasateshosnpeqesibesveasonedhgesdetengnea dines 13

Smaller species (body length4. 14.9 mm);

lateral margin of pronotum straight or

slightly concave in front of base, basal

angle almost 90°, distinctly projecting.

Lake Gairdner, Island Lagoon, Lake Hart

FantVsesshsseneseedeneaticeveraninscascand saskiae Baehr

— Larger species (body length 5.2-6.4 mm);

lateral margin of pronotum convex to base,

basal angle obtuse, or dentiform........ 14

14. —_ Elytraratherconvex, intervals remarkably

convex, microreticulation conspicuous.

Pernatty Lagoon....... matthewsi sp. nov.

— Elytra rather depressed, intervals but

slightly convex, microreticulation distinct

though not CONSPICUOUS ......... eee 15

Lateral margin of pronotum evenly convex,

basal angle not dentiform; elytra longer,

striae rather deep, distinctly punctate. Lake

VTS 0 ctebpretecs Becteverpeadtesst grossi Moore

— Lateral margin of pronotum rather straight,

in anterior third suddenly convex, basal

angle dentiform; elytra shorter, striae rather

shallow, barely punctate. Lake Frome ...

saidpusnanedindsvievapeterepegeeaptyertens vicinus Sp. NOV.

REMARKS

The collections included within this paper have

been made from lakes ranging up in size from

approximately 0.3 km? (‘Artaming Lake’, south of

Lake Acraman) to large systems such as Lake

Torrens, Lake Frome, Lake Gairdner, and Lake

Tyrell. Usually the collections were made when

the lakes were devoid of surface water. The

beetles were usually collected during the day by

excavating their burrows, the location of which

was indicated by a small mound of mud on the

lake surface. The beetles occupy a large part of

the dry lake surface that is free of salt crust. Some

collections have also been made using pitfall

traps, even on surfaces covered with a thin salt

crust and some specimens were detected while

running on the surface at night.

From excavated specimens and pit-trapping at

least two species are known to occur on Lake

Gairdner, Syrdenoidius spinipes and P. perovalis.

According to Moore (1977), at the shore of Lake

Eyre three species have been also found together

at light, namely P. hypharpagioides, P. gilesi, and

P. grossi. However, as on Lake Gairdner, the three

species represent rather different life style types,

namely a large, bulky one (P. hypharpagioides), a

medium sized one (P. gilesi, comparable to P.

perovalis), and an elongate, depressed one (P.

grossi, comparable to Syrdenoidius spinipes).

Hence, the species obviously occupy a special

niche, at least with respect to food preferences

which is demonstrated by their different head

sizes and mandible lengths and shapes.

If this and other recent papers on the Australian

pogonine fauna are any indication then it is most

likely that there are numerous other taxa awaiting

collection and description. With the existence of

flightless taxa there is an increased likelihood that

some species will be highly endemic. Clearly a

systematic survey of the salt lake fauna of

southern Australia is warranted, in the first

instance to establish the extent of the Australian

fauna and secondly for conservation purposes

since many salt lakes, especially those in Western

Australia, are sites of intense mining activity.

The ‘island-like’ nature of the Australian salt

lake habitats and the range in their size and degree

of geographic isolation make studies of their

fauna particularly interesting from an evolutionary

and genetic perspective (e.g. see Hudson and

Adams 1996). Desender and Serrano (1999) have

highlighted the value of using electrophoresis

techniques in the study of variation in Atlantic

and Mediterranean European populations of

Pogonus chalceus and similar such studies could

provide valuable insight to the Australian

pogonines.

Wing atrophy and flightlessness in beetles has

been documented by numerous authors

(Kavanaugh 1985, Roff 1990, and references

therein). In pogonines outside Australia, full wing

atrophy is only known in the Californian

Thalassotrechus barbarae (Horn), an intertidal

species which is outstanding also in other

morphological characters. Principally, all other

known pogonines possess fully developed wings

and are capable of flight. However, reduction in

wing size and lack of functional flight

musculature is known in some Pogonus chalceus

populations (Desender 1989, pers. com. 1999),

though the Mediterranean populations of that

species are capable of flight (Ravizza 1972).

Hence, from the viewpoint of wing development

the Australian pogonine fauna seems to be highly

unusual, because full reduction of wings is known

in three rather different Australian groups, the

NEW POGONINES FROM SOUTH AUSTRALIA 17

genus Syrdenoidius and the fully pigmented and

depigmented species groups of the genus

Pogonus.

It should be stressed, however, that in

Australia at least, there is a clear difference

between species inhabiting saline coastal habitats

and those that exclusively live on inland salt

lakes. Unlike in the Mediterranean, for example,

the Australian sea shores are inhabited only by

quite large, convex species of normal

‘pogoniform’ habitus (P. australis, P.

cardiotrachelus, P. nigrescens, P. sumlini, P.

variabilis), most of which are pigmented,

whereas elongate, depressed, depigmented

‘syrdeniform’ species are completely absent from

these environments. These latter species occur in

saline inland environments together with

pigmented species (P. cardiotrachelus, P.

fennelli), and convex, medium-sized or large,

either depigmented or partly depigmented

species (P. gilesi, P. hypharpagioides, P. zietzi).

Flightless species have been found only in inland

environments (P. fennelli, P. perovalis,

Syrdenoidius spinipes).

If we argue that the pigmented ‘pogoniform’

species, from the viewpoint of phylogeny, are most

plesiotypic, and the depigmented ‘syrdeniform’

species most apotypic, then the interior saline

environment in Australia has a very diverse, highly

evolved pogonine fauna, which is evidence of a

long term history of the fauna. In the rich pogonine

fauna of the Mediterranean region which likewise

includes normal ‘pogoniform’ species and

‘syrdeniform’ species the latter occur only at the

sea shore. The history of the Australian pogonine

fauna thus is strikingly different and once more it

demonstrates the island-like structure of the

Australian inland saline habitats that must have

been isolated for a long period.

ACKNOWLEDGMENTS

Lakes Gairdner and Frome are within the SA

National Parks and Reserves system and permits

Z23651-01 and Z23803-02 were issued for the

collections made from these lakes. The Scientific

Expedition Group is thanked for the opportunity to be

involved in field trips to the Gawler Ranges and

Balcanoona Station. Paul Fennell, Remko Leijs and

Jayne Skinner are thanked for their collecting efforts.

We thank Eric Matthews (SAMA) for the loan of most

of the material.

REFERENCES

BAEHR, M. 1984. Pogonus nigrescens sp. n. from

North Queensland (Coleoptera: Carabidae). Journal

of the Australian Entomological Society 23: 169-

171.

BAEHR, M. 1997. Two new species of the genus

Pogonus Nicolai from Australia (Insecta, Coleoptera,

Carabidae, Pogoninae). Spixiana 20: 1-6.

BAEHR, M. 1999. A new species of the genus Pogonus

Nicolai from Northwestern Australia (Insecta,

Coleoptera, Carabidae, Pogoninae). Spixiana 22:

143-147.

BAEHR, M. & R. SCIAKY (in press). A new species of

Pogonus Nicolai from Madagascar (Insecta,

Coleoptera, Carabidae, Pogoninae). Journal of

African Zoology.

BOUSQUET, Y. & S. LAPLANTE 1997. Taxonomic

review of the New World Pogonini (Coleoptera:

Carabidae). The Canadian Entomologist 129: 699-

731.

CHAUDOIR, M. de 1871. Essai monographique sur le

groupe des Pogonides. Annales de la Société

Entomologique de Belgique 14: 21-61.

CHAUDOIR M. de 1878. Descriptions de genres

nouveaux et d’espéces inédites de la famille des

Carabiques. Bulletin de la Société Impériale des

Naturalistes de Moscou 53: 1-80.

DESENDER K. 1989. Heritability of wing development

and body size in a carabid beetle, Pogonus chalceus

Marsham, and its evolutionary significance.

Oecologia 78: 512-520.

DESENDER, K. & J. SERRANO 1999. A genetic

comparison of Atlantic and Mediterranean

populations of a saltmarsh beetle. Belgian Journal of

Zoology 129: 83-94.

HUDSON, P. (2000). A new flightless species of

Pogonus Nicolai (Coleoptera: Carabidae) from

Western Australia. Australian Journal of

Entomology 39: 241-243.

HUDSON, P. & M. ADAMS 1996. Allozyme

characterisation of the salt lake spiders (Lycosa:

Lycosidae: Araneae) of southern Australia:

Systematic and population genetic implications.

Australian Journal of Zoology 44: 535-567.

KAVANAUGH, D. H. 1985. On wing atrophy in

carabid beetles (Coleoptera: Carabidae), with special

reference to Nearctic Nebria. Pp. 408-432 in

‘Taxonomy, phylogeny and zoogeography of beetles

and ants’. Ed. G. E. Ball. Dr. W. Junk Publishers:

Dordrecht.

KRYZHANOVSKIJ, O. L. 1990. New and little known

species of the tribe Pogonini (Coleoptera, Carabidae)

from Middle Asia and Kazakhstan. Trudy

Zoologicheskogo Instituta Akademiya Nauk SSSR

211: 3-8 (in Russian).

18 M. BAEHR & P. HUDSON

KRYZHANOVSKIJ, O. L. & V. A. MICHAILOV

1971. New and little known ground beetles

(Coleoptera, Carabidae) from Middle Asia.

Entomologicheskoe Obozrenie 50: 632-640 (in

Russian).

LORENZ, W. 1998. Systematic list of extant ground

beetles of the world (Insecta Coleoptera

‘Geadephaga’: Trachypachidae and Carabidae incl.

Paussinae, Cicindelinae, Rhysodinae). Tutzing: pp.

1-502 (printed by the author).

MOORE, B. P. 1977. New or little known Pogoninae

(Coleoptera: Carabidae) from Lake Eyre, South

Australia. Australian Entomological Magazine 4:

63-67.

MOORE, B. P. 1991. A new species of Pogonus

Nicolai (Coleoptera: Carabidae) from northern

Australia. Australian Entomological Magazine 18:

31-34.

MOORE, B. P., WEIR, T. A. & J. E. PYKE. 1987.

Rhysodidae and Carabidae. In: Zoological Catalogue

of Australia 4: 17-320. Australian Government

Publishing Service, Canberra.

MORVAN, P. 1973. Nouveaux Coléoptéres Carabiques

d’Iran. Bulletin du Museum National d'Histoire

Naturelle. 3e séries, Zoologie, no 110: 169-186.

RAVIZZA, C. 1972. I Pogonus (s.1.) dei litorali italiani

(Coleoptera, Carabidae). Bollettino del Museo Civico

di Storia Naturale di Venezia 22-23: 7-65.

ROFF, D.A. 1990. The evolution of flightlessness in

insects. Ecological Monographs 60(4): 389-421.

SLOANE, T. G. 1895. Notes on Carabidae from Lake

Callabonna, Central Australia. Transactions of the

Royal Society of South Australia 19: 124-137.

APPENDIX

Checklist of the recorded species of Pogoninae from Australia

Pogonus australis Chaudoir

Pogonus cardiotrachelus Chaudoir

Pogonus diplochaetoides Baehr

Pogonus fenelli Hudson

Pogonus gilesi Moore

Pogonus grossi Moore

Pogonus hypharpagioides Sloane

Pogonus matthewsi sp. nov.

Pogonus nigrescens Baehr

Pogonus perovalis sp. nov.

Pogonus saskiae Baehr

Pogonus sumlini Baehr

Pogonus variabilis Moore

Pogonus vicinus sp. nov.

Pogonus zietzi Sloane

Syrdenoidius spinipes gen. nov., sp. nov.

s.WA, VIC

s.WA, VIC, s.QLD

c.WA

s.WA

c.SA

c.SA

c.SA, w. VIC

c.SA

ne.QLD

c.SA

c.SA

n.WA

n.QLD, n.NT, n. WA

c.SA

c.WA, c.SA

c.SA

THE SIGNIFICANCE OF WHALES TO THE ABORIGINAL PEOPLE

OF SOUTHERN SOUTH AUSTRALIA

PHILIP A. CLARKE

CLARKE, P. A. 2001. The significance of whales to the Aboriginal people of southern South

Australia. Records of the South Australian Museum 34(1): 19-35.

Aboriginal people in the coastal region of South Australia had a broad relationship with

whales. This is reflected in their hunting and gathering economy, mythology and totemism, and

with their historical interaction with the first Europeans who arrived to hunt the whale. This

paper provides a cultural geographical view of Aboriginal associations with whales.

P. A. Clarke, Science Division, South Australian Museum, North Terrace, Adelaide, South

Australia 5000. Manuscript submitted 4 November 1999.

INTRODUCTION

The importance of whales to both Aboriginal

people and Europeans had an impact upon the

nature of their relationship on the frontier of

British expansion into South Australia in the early

nineteenth century. In earlier times, coastal

Aboriginal people feasted upon stranded whales,

which also had significance as totemic ancestors

to some groups. In the Lower Murray region,

whales were the mythological source of their

ability to make fire. After official European

settlement in 1836, many Aboriginal people were

involved directly and indirectly in whaling

activities. To Europeans, whales were the basis of

an industry that they hoped would help the Colony

of South Australia to develop and prosper. In the

developing Australian colonies, the whaling

establishments were in need of manual labour,

which Aboriginal people sometimes supplied. The

first Europeans recognised the skill of Indigenous

hunters, sometimes employing them as harpooners

and whale spotters. As a region, southern South

Australia is defined in this paper as the whole of

the coastal zone and the surrounding temperate

parts of the state.

SOURCES

The present study is part of a series of papers

by the author that considers the historical and

mythological relationships between Aboriginal

people and the southern South Australian

landscape (Clarke 1995, 1996, 1997, 1998, 1999a,

1999b). Europeans recorded the information upon

which it is based from Aboriginal people who had

lived in the southern districts prior to and during

the early phases of European colonisation in

South Australia. The material available consists of

the observations from Europeans involved in the

colonial process. The German missionaries,

Christian G. Teichelmann, Clamor W. Schiirmann

and Heinrich A. E. Meyer actively recorded the

culture of the local Aboriginal people. Their

publications were essentially studies of Aboriginal

language and religion (see Teichelmann &

Schurmann 1840; Teichelmann 1841; Meyer

1843, 1846; Schurmann 1844, 1846). The records

of William D. Wyatt (1879) from the late 1830s

came from his official investigations into the

admissibility of Aboriginal evidence in court.!

Richard Penney was involved in the welfare of

Aboriginal people in the Encounter Bay district.

Later sources, such as George Taplin (Journals;

1874; 1879), acknowledged the first recorders,

such as Meyer, as major sources of primary data.

These observers had a practical interest in

Aboriginal religion and traditions, in spite of the

fact that their records were compiled before the

development of anthropological theory. From

them the early relationship between Europeans

and Aboriginal people can be interpreted.

During the twentieth century, with the

development of scientific methods with obtaining

anthropological data, much additional

' Quarterly Report (1 October to 31 December 1837) from Wyatt to the Colonial Secretary, dated 1 January 1838 (Colonial Secretary Reports, 1838/

3 & 1838/69, Public Records Office, Adelaide).

20 P. A. CLARKE

ethnographic material was collected from southern

South Australia. Norman B. Tindale (1937, 1938,

1974, 1986, 1987), the Curator of Anthropology

at the South Australian Museum, published the

results of his fieldwork with the descendants of

Aboriginal ‘tribes’ in southern South Australia.”

The social anthropologists, Alfred R. Brown

(1918), Ronald M. Berndt and Catherine H.

Berndt (Berndt 1940; Berndt & Berndt 1993)

recorded Aboriginal culture and traditions in this

region. The twentieth century ethnographic

sources are essentially attempts to record a pre-

European culture, as it would have been lived by

the generations preceding that of their informants.

The popular accounts of southern South

Australian mythology by Charles P. Mountford

and Ainslie Roberts were chiefly based on the

published records of Tindale. A more detailed

analysis of the chief historical sources used in this

paper is given elsewhere (Clarke 1994: 63-81,

417-425; 1995: 145-146; 1999a: 52-53).

PRE-EUROPEAN PRACTICES

The coastline of South Australia is part of the

migration route of the southern right whale

(Eubalaena australis) from the Southern Ocean.

Occasional strandings of whales were events that

attracted many Aboriginal people for feasting.

Daisy Bates recorded that the Wanmaring local

group at the Head of the Bight in western South

Australia had ‘occasional gorges in large fish

food, dead whales and other large sea creatures

being found after some great storm’.? Aboriginal

people considered that they had an active role in

some strandings. For example, songs were used

by ‘strong men’ of the coastal Lower Murray

groups to ‘cause’ whale strandings (Tindale 1974:

18, 23-24, 80).* People from surrounding areas

converged on the dead whale, being called by the

aroma of a stranding, if not the messages sent out.

In the Aboriginal languages around Adelaide and

Encounter Bay the whales of all species were

collectively called kondoli.6 Tindale records

another term for them that referred to their water

blowing. Tindale says:

The whales which frequented the shores off the

mouth of the Murray River and the rocks at

Encounter Bay where they often came close in shore

were called winkulare, literally the ‘whistlers’ or

‘blowers’ from the intransitive verb winkulun

‘whistle’ because of their ‘blowing’. Magical spells

designed to entice the whales ashore were practised

by the ‘clever’ men.°®

For the Lower South East of South Australia,

there is a transcription of a whale song in the

“‘Booandik’ language (Smith 1880: 139). This was

translated as “The whale is come. And thrown up

on land.’ These lines were repeated over and over,

perhaps to induce a stranding.’ Evidence from

elsewhere in Australia, such as the rock art around

Sydney in New South Wales, suggests that coastal

Aboriginal people had a keen interest in whales.®

Aboriginal people of the whale descent group

at Encounter Bay believed that some of them

could ‘sing’ whales in towards the shore or out,

by standing on a rock and singing some

‘wordless chant’.? Tindale (1937: 107, 112)

recorded a whale song, in the Ramindjeri dialect

Tindale’s journals, field notebooks, and the ‘Milerum’ manuscript are also important ethnographic sources. This material is housed in the

Anthropology Archives, S.A. Museum.

‘The Passing of the Australian Native. The Wanmaring Group (Head of the Great Australian Bight).’ D. M. Bates manuscript collection at the Barr-

Smith Library in Box 12/Section XIII/Part 6b.

Aboriginal people claimed in the early 1980s that an elderly Ngarrindjeri person living at Point McLeay, who had recently died, knew how to ‘sing

a song to bring in whales’. Also in the early 1980s, some elderly Ngarrindjeri informants knew that there was a myth concerning whales and fire,

but had forgotten the details (audiotape of the Cameron family talking at Kingston South East with P. G. Jones, S. J. Hemming & P. A. Clarke, 23

April 1983. Anthropology Archives, S.A. Museum).

In the Adelaide area, whales were termed ‘con-dol-ley’ (Williams 1839 [1840: 295]), ‘kondolli’ (Teichelmann & Schiirmann 1840 2; 12) and

‘condollee’ (Stephens 1889: 498). For the Lower Murray dialects, the name for whale is recorded as ‘kondarli’ by G. Taplin (1879 [1874]; 1879)

and F. Taplin (Point McLeay Letter Books, 1879-84, p.98), ‘kondolle’ (Wyatt 1879: 170), ‘kandarli’ (Howitt 1904), ‘kondolly’ (M. Wilson in

‘conventional mission script’ [Tindale 1930-52: 161-165]), ‘kondolly’, ‘kondal’ (Tindale 1934-37), ‘kondoli’ by Tindale (1937), ‘kondali’

(Mountford & Berndt 1941), ‘kondole’ by Mountford and Roberts (1969) and ‘kondili’ by Berndt and Berndt (1993). This paper uses ‘Kondoli’ as

a standard term for the Whale Ancestor.

Tindale (‘Milerum' manuscript, stage A, no.3. Anthropology Archives, S.A. Museum).

After European settlement, Aboriginal people of the South East of South Australia also had a popular song about the ‘whale fishery’ at Rivoli Bay

(Stewart n.d, [1977: 65-66]).

Whales are major elements in the rock art in the Sydney area. At least one site appears to show a feast around a stranded whale (Campbell 1899:

46-47, pl. 24, fig. 1; Stanbury & Clegg 1990: 22-23, 54, 80-83, 113, 119). In September 1790, Tench (1793 [1996: 134]) records the feasting of a

stranded whale which was ‘in the most disgusting state of putrefaction’ in the Sydney region. Kingston (1876: 277-278, 327) gives a related

account from north of Sydney.

The Mail newspaper, 19 August 1932. There are recent accounts recorded by Bell (1998: 320) concerning Aboriginal people formerly using whales

for transport, although these are unsupported by the historical record.

SIGNIFICANCE OF WHALES 21

of the northern end of Encounter Bay, to send

whales away.'° This was sung by a man of the

kondoli nga:tji, or whale descent group, in order

to assist a female whale and calf escape the

shallow waters of Encounter Bay.!' In this

account, other people who were ‘evil-minded’,

desired the whales to be stranded so that

kraipunuk or oil could be collected for ‘spear

poison’, perhaps as a sorcery material used

against people who had the whale as their

totemic familiar or nga: tji. Tindale stated:

A man who had the whale as totem would not eat

whale. If a whale became stranded on the coast, he

would give permission for others to use it as food,

but he himself would merely rub himself with the oil

which it produced. If he saw a whale floundering off

the shore, he would sing a magical song, telling the

whale to avoid the shallows and escape to the sea.'*

Tindale stated that to decorate themselves the

men of the Warki and Tangani groups of the

Lower Murray region would be ruddled with red

ochre and whale oil.'’ Elsewhere in southern

South Australia, it is also likely that whales were

considered to be totemic beings, although apart

from the Lower Murray region, the available early

ethnographic data is slim.

The whale was associated with the Latalindjera

descent group, which received its name from the

place, Latang, at Hindmarsh River near Victor

Harbor at Encounter Bay (Berndt & Berndt 1993:

311). To the south of Encounter Bay,

Kondilindjarung (‘whale, place-name marker’)

was a death place for whales, where one or two

often beached themselves during the winter

months (Berndt & Berndt 1993: 81, 103, 320,

326-327, 342, 565). It is situated on the

Younghusband Peninsula on the seaside, south

east of the Murray Mouth but north west of

Noonamena. It was claimed that Kondilindjarung

was where the people of this group and their

associated beings, the whales, attempted to return

before death. The Coorong ocean beach was

renowned for its whale remains. Tindale records

that:

On the ocean beach one might be the inhabitant of

another world, composed of sand, sea, spume, and

giant whale bones. The scattered bones of blue

whales lie here and there along the beach, for this is

a graveyard where the great circumpolar current

impinges upon the shores of Australia and casts up

its burden of flotsam.'*

According to the Berndts, Kondilindjarung is in

the Kondilindjera descent group territory, believed

to be an offshoot of the Ramindjerar, but who

spoke Tangani instead. The relationship between

the Kondilindjera and the Latalindjera descent

group described by Tindale above is uncertain."

The presence of the Kondilindjera on the south

eastern side of the Murray Mouth may well have

been the result of a post-European adjustment in

territory, as Taplin (1874 [1879: 2]; 1879: 34)

recorded the Kondarlinyeri as a ‘clan’ based on

the north western side of the Mouth.

Kondarlinyeri was said to mean ‘belonging to

whales’ (Taplin 1879-84: 98). Brown (1918: 252)

provided evidence that suggests that there were no

‘Kandarlinyeri’. He stated ‘I was told by the

natives that Kondarlinderi (the place of the

whales) is merely the name of a part of the

country occupied by Pankinderar clan, and that

there is no clan of this name (Brown 1918: 252).’

During his fieldwork, Brown (1918: 240)

considered it too late to obtain detailed

information on the totemism of the Yaraldi

people. Nevertheless he claimed:

The men and women of a clan might eat, and did

eat, their totem, if it were edible, but they were

careful to destroy all the remains (bones etc.), lest

they should fall into the hands of an enemy and be

used for evil magic. Some part of the totemic animal

was in some instances used as a badge of the clan.

Thus the Liwurinderar used to carry pelican skins

on their spears when they went out to fight (Brown

1918: 241).

‘0 See Tindale (1931-34 (1); 252-253) for notes of ‘Kondoli Tungar or Whale Song of the Ramindjeri tribe at Encounter Bay, Song 14’.

"' Tindale inscribes a draft typescript version of his paper with ‘much of the song data given verbatim as dictated by Milerum’. Here it is stated that

the whales were swimming in ‘white sand water’. Other details include that the whales were ‘mother’ and ‘son’ and that whale ‘slime’ or oil was

required to rub on their bodies with red ochre as a base for poison (copy in E. H. Davies papers, AA309, Accession no.1, Anthropology Archives,

S.A. Museum).

"Tindale, The Advertiser newspaper, 14 May 1936. The symbolic importance of whales may be reflected in the rock art of the Sydney area, which

depicts people inside whales. Stanbury & Clegg (1990: 22-23) suggest that this might be explained by the eastern seaboard practice of sick people

lying inside the body of a stranded whale.

"Tindale (‘Milerum’ manuscript, folder 1, draft A. Anthropology Archives, S.A. Museum).

4 Tindale, The Advertiser newspaper, 12 May 1936.

'S Tindale (Lower Murray Totems Chart, miscellaneous papers, Anthropology Archives, S.A. Museum) links together the descent groups of

Kondolinyeri (Taplin 1874 [1879]), Kondarlinyeri (Taplin 1879), Kandarl-inyeri (Howitt 1904), and Karagarindjeri [equivalent to Kondolindjeri]

(Tindale ms). He does not list the Latalindjera.

22 P. A. CLARKE

This account is consistent with Meyer’s claim

that the nga:tji was a ‘friend’, ‘countryman’ or

‘protector’ (Meyer 1843: 86; 1846 [1879: 198])

and Taplin’s definition of it as a ‘tribal symbol’ or

‘tutelary genius’ (Taplin 1874 [1879: 1, 63-64,

134]). The Berndts also record that some people

were able to eat their nga:tji. Regarded as the

flesh of the Kondilindjera, the members of this

Lower Murray group alone could make the first

cut on a beached whale carcass (Berndt & Berndt

1993: 81, 103, 320, 326-327, 342, 565). Other

groups would wait until they had eaten.'® The

spirit of Kondoli departed through

Kondilindjarung, leaving its meat for its

‘brothers’. In this area, Aboriginal people

possessed a pre-European system giving certain

groups rights and privileges to the resources of

the sea (see Peterson & Rigsby 1998).

The Kondilindjera responded to a whale

stranding by sending messengers with invitations

to neighbouring groups to come in for a feast on

whale blubber, pailpuli, and meat, mami."

Normally, the coastal districts were sparsely

populated during the wintertime (Tindale 1938: 21;

1974: 61-62; Clarke 1994: 182-183). The cutting

of the flesh and blubber was performed with flint

knives, maki. Aboriginal people cooked the fat

before it was eaten and the oil was rubbed on their

bodies for protection against the weather. The

witjeri, leaves of the pigface (Carpobrotus rossi),

were used as a relish with the meat. Ceremonies

were held during the nights of the feast. The site of

the stranding was not left until all the blubber was

eaten. Aboriginal people used the ear bones as

drinking containers and water storage vessels.'*

Aboriginal midden material collected from Moana,

a sand dune area near the mouth of Pedlar Creek

south of Adelaide, by Tindale in the 1920s contains

sperm whale teeth and dolphin vertebrae.'? This

suggests that Aboriginal people had eaten beached

whales and dolphins. Also, in the Rivoli Bay area,

south of Encounter Bay, whales were also

sometimes stranded. Aboriginal people called this

place Weirintjam or Wilitjam, which Tindale

claimed meant ‘place of whales’ from the

Potaruwutj word ‘weirintj’, a whale.”” It was a

place noted by Tindale’s informants for the whales

that became stranded there, providing big feasts for

the local people.

It is unlikely that Aboriginal people in South

Australia were able to use their watercraft, built

for calm water conditions, in killing small

whales.?! For marine animals stuck in shallow

water or stranded on land, killing was a more

straightforward task. Nevertheless, hunting across

the open sea was limited. For instance, Ramindjeri

people swam or travelled on rafts to West Island,

which is less than a kilometre off the coast south

west of Victor Harbor, to kill sea lions (Neophoca

cinerea) and fur seals (Arctocephalus fosteri)

(Tindale 1941: 241). They also travelled to Pullen

Island on rafts in calm weather for sealing.”” Most

distant islands, such as Kangaroo Island, were

places beyond the range of Aboriginal hunters

immediately prior to European settlement (Clarke

1996, 1998). There is no evidence to suggest that

Aboriginal people actively hunted whales or

dolphins.

WHALES IN ABORIGINAL MYTHOLOGY

The earliest written account of whale mythology

in South Australia was by C. Teichelmann in

1840, presumably recounted by Aboriginal people

who lived in the Adelaide area (see Amery 1998:

219, 224). He records:

When the lark and the whale were men, they fought

against each other. The lark speared the whale twice

in the neck. The whale, finding itself sorely

wounded, made its escape, jumped from pain into

the sea, became a whale and spouted through the

two wounds water to heal them; but in vain, till this

day.

‘6 Bates (1985: 197-198) records that the mammang borungur or whale totem people of Cape Leewin in the South West of Western Australia also ate

the flesh of stranded whales, in spite of the fact that this was not the practice of people from other totems.

7 Wyatt (1879: 170) claimed that Encounter Bay people also called whale blubber ‘kondolle’.

South Australian Museum specimen A49445; container made from a whale ear bone, collected from Pelican Point on the Coorong, donated by Mrs

O. T. Cleggett, 1940s.

© Ross (1984; 19-41) provides an overview of the archaeological investigations based at Moana. This material would be less than 6,000 years before

present, when the coastline of present-day Adelaide was largely formed. The sperm whale teeth found in Moana ‘Site C’ may have come from a

multiple stranding, as they are from at least two individuals, one young and one old specimen (C. Kemper, pers. com.).

20 _N. B. Tindale (Geographic Place Names Cards & Potaruwutj Vocabulary Cards, Anthropology Archives, S.A. Museum), Tindale thought that the

variation in the place name, Wilitjam, is probably relatively modern.

In contrast, Aboriginal hunters in northern Australia had more access to the sea through their use of dugouts and outriggers gained through contact

with Macassans and Torres Strait Islanders, For a description of their watercraft, see Haddon (1913), Tindale (1926: 103-112) & Baker (1988).

2 See also Tindale (Wunindjeri clan entry in ‘Clan Data’ folder in ‘Milerum’ manuscripts, stage 2 Anthropology Archives, S.A. Museum).

23 South Australian Colonist newspaper, 7 July 1840,

SIGNIFICANCE OF WHALES 23

The same corpus of whale mythology appears

to have been shared by the Adelaide and

Encounter Bay peoples, perhaps reflecting

Aboriginal movements around the time of

European settlement in the early nineteenth

century. In a Ramindjeri account of the myth,

whales are connected with the origin of fire

(Meyer 1879: 202-204). Here, it was believed that

once long ago, all the Ramindjeri people gathered

to dance at Mootaparinga (Cut Hill, along

Hindmarsh River).** They did not have a fire, so

they had to dance all day and it was hot. Their

perspiration dripped down and became the large

ponds there, and hills and valleys formed through

the buckling of the ground caused by the stamping

of their feet. Eventually, they sent for Kondoli, a

large and powerful man who possessed fire. He

came, but hid his fire. This made the Ramindjeri

angry. Another Ancestor, Riballi, threw a spear at

Kondoli, hitting him in the neck.** The commotion

this caused transformed most of the people there

into different animals, such as fish and birds.

Kondoli himself rushed into the sea and ever after

blew water out of his wound. Riballi took

Kondoli’s fire and placed it in a grasstree

(Xanthorrhoea species), where it can be removed

by using the dried flower stems as fire-sticks.°

The series of accounts recorded by Tindale in

the twentieth century provide details that were not

part of Meyer’s account. In May 1934, Tindale

received a Ramindjeri version of the Kondoli

myth from Frank Blackmoor (Tindale 1930-52:

161-165; 1934-37: 181-184).?” Here, Kondoli

was a big man who possessed fire that fell from

his body as he danced. He came to participate at a

Ramindjeri ceremony. Kuroldambal the Owl

argued with Krilbalil (= Krilbali) the Lark about

who was to spear Kondoli to get the fire.

Kuroldambal, who had large eyes, wanted

Krilbalil’s spear to use, as his eyes were too small

to see properly. But when the dancing was close

enough, Krilbalil grabbed his spear and struck

Kondoli at the back of the head, making the fire

fall out. Krilbalil grabbed the fire and placed it

amongst grasstrees, causing the scrub to burn.

People now had access to fire. Kondoli dived into

the ground, eventually coming out and going into

the sea.

Tindale recorded a version of the ‘Story of

Kondoli’ from Milerum, a Tangani speaker, in

May 1936 (Tindale 1934-37 vol.2: 232-235). In

this account a ‘tribe’ that was part Mereldi

(Murray River people), part Ramindjeri and

Yoltindjeri (= Joltindjeri) planned to have a

molkaldi type ceremony at which all participants

would perform. Messengers were sent out in all

directions and people gathered at Mutabaringga

(= Mootaparinga), a site on the largest bend of the

Hindmarsh River. The dancing area was at a flat

on top of a range that had a depression. Kondoli,

who was a big man, was camped at Brown Hill.

He carried the flint with which fire could be

made.** A ‘river man’ (Murray River person)

called Ratunangi, the Robin Redbreast Bird, was

there. Seven or eight Teiwuri (Brown Treecreeper)

women from a local group, who greatly desired

Ratunangi, grabbed him.” Relatives among the

‘river people’ grabbed their weapons and a fight

with spears and waddies (clubs) took place.

Firesticks were also thrown through the air.

To escape the fight, Kondoli fled towards the

sea down Hindmarsh Valley, carrying his flint. He

also had pieces of granite, which he dropped as

boulders along the creek. At the site of Gay

Bridge, Port Elliot (Kandeining) he took a boulder

out into the sea. After returning and heading a

little further inland he rested before diving into

the sea at Kantjoar, where there is a swamp.”

pL

26

Colonists used Mootaparinga (or Murtaparri) in the 1830s as the name for the mouth of Hindmarsh River at Encounter Bay (Wade cited Gouger

1838: 25; Mann cited Gouger 1838: 39, 42; Teichelmann & Schiirmann 1840 2: 75). In the Adelaide language Mootaparinga reputedly means

‘brackish water’ (Robinson 1975; 44). If so, then it appears to be a term from a language north west of the Lower Murray. Adelaide terms, such as

murta [‘excrements of animals’), parri [‘river’] and -ngga [grammatical ‘in’] (Teichelmann & Schiirmann 1840 1: 22; pt 2: 25, 38, 76) appear

relevant, possibly translating as ‘animal manure water, place of’. Tindale (‘Myths’ folder in ‘Milerum’ manuscript collection, Anthropology

Archives, S.A. Museum) says ‘Mutabaringga’ is known by Europeans as Cut Hill.

The ‘Riballi’ of Meyer is probably his rendering of ‘Krilbali’, the brown skylark.

The use of fire is a major element of the Waiyungari mythology (Clarke 1999b). Bell (1989: 321-322, 427) speculates that whales are of special

significance to Aboriginal women due to the link with fire and through associating the protective nature of whales over their calves with human

child-care responsibilities.

Yaraldi man, Mark Wilson (Thalrum or Thralrum), wrote the original text using conventional mission script. Tindale then annotated it by working

directly with Frank Blackmoor, a Yaraldi man of the Piltindjeri descent group. Although Tindale refers to the Skylark as ‘Krilbalil’, 1 prefer the

form ‘Krilbali’, as written by other authors and as recognised by some contemporary Ngarrindjeri people (see foot note 44).

The flints were struck whilst being held in a dry pad of fungus, kumpalatingi (Mountford & Berndt 1941: 342-344).

Tindale (1934-37; 233) records that as a result of what happened then, ‘today you always see him [Ratunangi, Robin Redbreast] with seven or

eight wives’,

According to Tindale (Geographic Place Names Cards, Anthropology Archives, South Australian Museum), Kantjoar is a site in the Ngurunderi

myth were he urinated (kaindji =: urine).

24

From here, Kondoli went west along the shore,

sometimes walking and other times swimming in

water for safety. He walked between Granite

Island and the shore, looking back to see if the

others were following him. When Kondoli

reached Wunangg, the mouth of the Inman River,

he saw his friend Yamakawi rushing down the

Inman towards him.*! The ‘wild men’ were

chasing Yamakawi for his firestick, made of

ngalaji (dried flower stem of the grasstree), that

he carried. Although Kondoli had eluded his first

pursuers, he had to flee again, this time with

Yamakawi. Kondoli went a little way into the sea

and turned into a whale. There is a reef there

today and the spray that rises with the waves is

the whale spout, which represents smoke.

Yamakawi went into the sea at Narailkang, near

Waitpinga, and turned into a shark with big

teeth.*? For some time, people on the land had no

fire as Yamakawi had taken away the ngalaji.

Kondoli did not go far away as he had left

something back at his camp. The people ‘sang’

(charmed) him into the shore, where they jumped

on him and took his flint so that they could make

fire. They also grabbed Yamakawi’s firestick,

which had been used as a back fin.

In another of Tindale’s records, Kondoli the

Whale man chased the Shark man who had stolen

Kondoli’s fire-making equipment, wintjimi,

comprised of flint and pyrites.*7 This occurred at

Spring Mount, which was called Mutabaringga,

said to mean ‘sacred magic doctor’s place’. The

hill represents Kondoli, who later became a whale.

A big hole at the summit was said to be the ‘blow

hole’ of the whale. Kondoli chased the Shark man

to Brown Hill, called Kondolanangg.** From here

Kondoli in anger rushed towards the sea in pursuit

of the thief. Kondoli was transformed into a whale

P. A. CLARKE

at Elliot Beach, called Kandeining, which was

translated by Tindale’s informants as ‘from

here’.*> The Shark man was transformed into the

whale’s enemy, the shark, who retains the fire

flints as teeth in his mouth. Tindale linked this

account to the name of the south western spur of

the Brown Hill, Nangge-we:ke, which he claimed

means the ‘stolen sun’, referring to the stealing of

fire.*° In what appears to be a related account,

Tindale stated:

One of the myths of the area tells how there was

contention between a man of the whale totem and

another, the shark, for possession of the fire flints,

without which the whale could not use his pyrites.

At the climax of the story the men became animals

and the shark fled taking with him the flint stones

which were transmuted into his teeth.*”

These versions place emphasis on the origin of

fire relating to conflict between Ancestors,

involving the Whale and Shark.

Tindale recorded other elements of the ‘Fire

myth’ in the Warki language of the Goolwa area.

Mutabaringga or Cut Hill was one of the homes

of the whale man, Kondoli.*® According to

Tindale it was derived from muturi (sacred), bari

(river) and -ingga (at).*? Kondolinggara or Mount

Jagged was where Karilbali (= Krilbali) fought

with Kondoli for the possession of fire, wounding

him. The place-name was reputedly derived from

kondoli (whale) and nggaran (to assault or

wound).*° Kondolanang (= Kondolanangg or

Brown Hill), where Kondoli emerged from the

ground, was said to mean ‘refuge of whales’,

derived from kondoli (whale) and nangare (refuge

or shelter).41 In another version, Kondoli, after

being wounded by Karilbali:

set fire to the country. Hiding first at this place

[Kondolanang] he [Kondoli] fled to Kandeining

31 According to Tindale (1934-37: 232-233), Yamakawi was related to Kondoli, their hunting grounds adjoined. The quartz stones that Yamakawi

left behind when he fled were used for making the jags on spears. Some of these were given to Yamakawi, after he became a shark, for his use as

teeth (Tindale 1934-37: 234). Tindale writes ‘Yamakawi' as ‘Jamakawi'’.

Tindale (‘Myths’ folder in ‘Milerum’ manuscript collection, Anthropology Archives, S.A. Museum) claims that Yamakawi was the White Pointer

Shark being who lived at Inman Hill.

Tindale (Geographic Place Names Cards & ‘Milerum’ manuscript, stage A, no.3. Anthropology Archives, S.A. Museum). This is presumably a

Ramindjeri version of the myth. Tindale claims that Mootaparinga is erroneously ascribed to Hindmarsh River by Cockburn (1908: 58).

See also Tindale (Wunindjeri clan entry in ‘Clan Data’ folder in ‘Milerum’ manuscripts, stage 2. Anthropology Archives, S.A. Museum).

N.B. Tindale (Geographic Place Names Cards, Anthropology Archives, S.A. Museum). Informants not named.

This place name is the Nangawooka of Tite, which Tindale (Geographic Place Names Cards, Anthropology Archives, S.A. Museum) claims

Cockburn (1908: 63-64]) incorrectly translates as ‘place of springs’. Informants not named. See also Tindale (Wunindjeri clan entry in ‘Clan Data’

folder in ‘Milerum' manuscripts, stage 2, Anthropology Archives, S.A. Museum),

N. B. Tindale (no date) ‘Strike-a-lights, Fire Flints’. Tindale collection, miscellaneous papers, Anthropology Archives, S.A. Museum.

Data recorded by Tindale (‘Aboriginal Placename Cards’, Warki Language, AA338, Anthropology Archives, S.A. Museum) from the Yaraldi

informant, Albert Karlowan (Karloan) and the Tangani informant, Clarence Long (Milerum).

The derivation of ‘muturi’ (sacred) here seems less likely than as ‘murta’ (animal excrement) [see footnote 23].

Data recorded by Tindale (‘Aboriginal Placename Cards’, Warki Language, AA338, Anthropology Archives, S.A. Museum). Informants not named.

Data recorded by Tindale (‘Aboriginal Placename Cards’, Warki Language, AA338, Anthropology Archives, S.A. Museum) from the Yaraldi

informant, Mark Wilson (Thalrum or Thralrum) and the Tangani informant, Clarence Long (Milerum).

SIGNIFICANCE OF WHALES 25

where he entered the sea as a whale. There a shark

being stole his fire flints. A wide spread story.”

After entering the sea, the fire flints were then

stolen by Ngarakkani (= Ngarankani) the Shark

being and the property of fire making was

transferred to grasstrees and wood.*?

In 1934 Tindale recorded an account, ‘Story of

the Stealing of Fire by the Whale’ from Mark

Wilson in the Yaraldi language.“ Here, all the

birds and animals, who were then human, met for

a ceremony at Mutabaringga. Only Kondoli the

Whale possessed fire; he guarded it jealously.

Krilbali the Lark and another unnamed bird

quarrelled over who should steal it. Kondoli was

speared in the head by Krilbali and leaped into a

cave on the hill before the fire could be taken

from him. The Whale emerged in the deep water

at Victor Harbor that is close to the beach and

extends to Granite Island. The steam coming from

Kondoli’s wound was like that coming out of an

earth cooking-oven, showing that there was fire

inside. The whale still blows its steam out and

stranded whales are quite hot, compared with

other marine animals which are cold. In the sea,

the Shark stole fire from the Whale and gave it to

the birds, who were unable to control it and set

fire to the country. The raging fire entered the

trees, which means that they would now readily

burn if set alight. Fire also entered the flints on

the ground. Many of the birds got burnt, such as

Tuta the Scarlet Robin being who had his breast

burnt bright red. After the fire died, people

discovered how to make fire from splitting

grasstree sticks and rubbing them together and

also by striking flints against ironstone pieces.

The anthropologists, Ronald and Catherine

Berndt, also provided a record of the Kondoli

mythology in the Encounter Bay district (Berndt

& Berndt 1993: 16, 118, 235-236, 341-342, 450—-

451).4° The Hindmarsh River area at Victor

Harbor is associated with the Whale Ancestor and

the origin of making fire with flints, maki. The

flints were obtained from the Ramindjeri hills

people. In the Berndts’ version, many people

attended a dance meeting held at Kondilinar,

which is inland. When Kondoli the Whale

Ancestor danced, sparks came from inside him,

which excited the onlookers who were at that time

without fire. Krilbali the Brown Skylark and

Retjurukuru the Wagtail planned to spear Kondoli

whilst he was dancing, to get his fire. Krilbali

speared him deep in the back of his neck. Then

Krilbali grabbed some of the fire that gushed out

of the wound and fled with it. The people

assembled there laughed, but Kondoli, Ngarankani

(Shark), Mulori (Stingray) and Pungari (Seal)

jumped into the water and came out at Latang,

near the coast at the mouth of Hindmarsh River.”

The Berndts (1993: 311) stated that Latang meant

‘place of spearing’.*7 The Whale stayed at

Balgolin (Pultung, Victor Harbor).

One of the Berndt’s chief informants for the

Kondoli mythology was the Yaraldi speaker,

Albert Karloan, from the Manangka descent

group. He told the Berndts that the spear thrown

at the Whale Ancestor had entered the back of the

neck, penetrating as far as the jaw. It was the jaw

that produced the sparks. Karloan claimed that

when the Whale was speared, the fire that fell on

the ground at Kondilinar turned to flints, maki,

which could be used to make fire. Flints could be

also found where the Skylark had dropped them

whilst running all over the country. The method

of the transference of the fire from the Whale

Ancestor to humanity is different from Meyer’s

earlier account, when a fire-stick made from a

grasstree flower stem was used.

The Mountford and Roberts (1969: 40-41)

version has details that appear as poetic

adjustments to the main recordings. Fire was

needed here because the dancers required light to

dance at night. The large and powerful Kondoli

was the sole owner, but hid the fire. After Kondoli

was speared, the people at the ceremony were

transformed into animals, such as kangaroos,

possums, birds and fish. Kondoli, as the largest

man, became the largest animal, the whale.

Ngarrindjeri writer, David Unaipon, wrote a story

“Data recorded by Tindale (‘Aboriginal Placename Cards’, Warki Language, AA338, Anthropology Archives, S.A. Museum). Informants not named.

The derivation of Karilbali is here and on other cards recorded as ‘magpie-lark being’ instead of ‘lark’, but this clearly an error. See end note 44.

“Data recorded by Tindale (‘Aboriginal Placename Cards’, Warki Language, AA338, Anthropology Archives, S.A. Museum). Informants not named.

“ Tindale (1930-52: 48, 91, 272-273). In this account, Tindale incorrectly identifies Krilbali as the magpie lark (Grallina cyanoleuca). From other

sources, including contemporary Aboriginal informants, this term refers to the skylark (Alauda arvensis) (Brown 1918: 242; Berndt & Berndt 1993:

235-236, 311, 450-451, 461-462).

4S The Berndts write ‘Kondoli’ as ‘Kondili’.

“* Many of these Ancestors became ngavtji of Ramindjeri clans surrounding Victor Harbor (Berndt & Berndt 1993: 311). Ngarankani, shark, was the

nga:tji of the Ngarakerindjera descent group, based at King Point. In the Tjirbruki mythology he was speared and become the gummy shark

(Tindale 1987: 9). Krilbali, brown skylark, was the nga:tji of Krilbalindjera based near Kondilinar. Pangari, seal, was the nga:tji of the

Ratalwerindjera at Middleton.

“7 Berndts (1993: 311) claimed that Latang was equivalent to ‘Yalla-doola’ as listed by Wyatt (1879: 179).

26 P. A. CLARKE

about Wondangar the Whale and Goon na Ghun

the Starfish Ancestors.*® Although some

Ngarrindjeri words are used in the text, the

inclusion of a lyrebird, which is a species not

found in South Australia, suggests it was intended

as a generalised eastern states account.

There are whale mythologies in other parts of

southern South Australia that bear some

resemblance to the Kondoli myth. The existence

of the Whale Ancestor as a land-based person

appears widespread. Schurmann recorded in the

Adelaide Plains region that:

Nganno — travelled far and wide, seeking the

murderers of his son, Gurltatakko, and while

travelling named the country as we know it today.

After killing the murderers he went home but his

tribesmen on seeing him panicked and ran in fear

into the sea where they were transformed into sea

creatures. He told them not to do it but they

responded ‘I am a shark’, ‘I am a whale’ etc.

Nganno turned himself into a sea monster at the end

(Schurmann Journals, 21 August 1839).

It is interesting to note that many of the terms

and place-names used in the Lower Murray

versions above are derived from the Aboriginal

vocabulary of Adelaide. In the Tjirbruki

mythology of Gulf St Vincent and western

Encounter Bay, some of the Ancestors ran into the

sea to become species of shark (Tindale 1987: 9).

The language used in this account recorded by

Tindale is a mixture of Adelaide and Lower

Murray vocabulary.

Stealing the ability to make fire is a common

theme in the mythology of Aboriginal Australia.

There are other accounts that appear structurally

similar to the Kondoli myths discussed above, but

do not involve whales as one of the primary

Ancestors. For example, in a version of a Western

Desert myth recorded by Tindale, the Turkey

Bustard Ancestor kept fire to himself and fled

south towards the Southern Ocean at Eucla to

place the flints under the water.” The Hawk Men

rescued the flints, which are accessible today only

at low tide on calm days. Similarly, Smith (1880:

18-20) records that the Booandik people between

Mount Gambier and MacDonnell Bay in the

South East of South Australia believed that fire

was once the sole property of Mar the Cockatoo,

and was hidden on his head. At a meeting, people

argued over how to find out more about Mar’s

fire. Several people attempted to spy on Mar

making the fire. As a result, Tatkanna the Robin

Red Breast had his chest singed red from the heat

of Mar’s fire. When fire was at last caught in a

grasstree stick, it spread to the grass and dry

underwood where it started a bushfire. Mar rushed

to where the others were camped and a fight

started. One of those there, a large man named

Kounterbull, received a deep spear wound in the

back of his neck. He rushed into the sea where, as

a whale, he was afterwards to be seen spouting

water from his wound. In Western Victoria,

Crows had fire and the Fairy Wren and Hawk

stole it (Dawson 1881: 54). In the West Coast

district of South Australia, Aboriginal people

possibly perceived whales as having a more

sinister character. In the Mirniny language, whales

were called muburn kailgaburdi, with muburn (or

moburn) recorded as ‘devil’.*° Tindale claimed

that there ‘are several versions of the [fire] story

extending as far to the west as Perth and to the

Tanganekald on the Coorong.’*! He cites the

record of Grey (1839: 76) from the South West of

Western Australia that lists maad-jit-teeyl as a

‘magic stone’ of the shark. Tindale also provided

an account from the Murray Basin where a small

species of hawk stole an ember from the shark as

he was entering the sea and placed it for safe

keeping in a grasstree.*? The similarity of the

events in widespread accounts of this myth, along

with the variety of language chosen by the

informants to communicate them, make it difficult

to determine the origin and range of the whale

myths.

The Kondoli mythology of the Lower Murray

has important cultural insights into both methods

of fire-making. The use of fire-sticks in what is

referred to as the rotation method is widespread

across Australia. Tindale describes this process in

the southern regions:

Fire making is practised using a dry grasstree flower

stalk, the ‘male’, twirled into a split section of the

same stalk, in which usually a notch has been cut.

This is the ‘female’. Tinder is placed below the

“ Unaipon, no date (Davis et al. 1990: 33-52).

“ —N.B, Tindale (no date) ‘Strike-a-lights, Fire Flints’. Tindale collection, miscellaneous papers, Anthropology Archives, S.A. Museum.

so D. M. Bates, ‘Native Vocabulary, Eucla District’, Barr Smith Library, University of Adelaide (box XII, 2A 4-5). According to Bates the shark

appears to have been called by a related term, kailga-kailga. Similarly, the porpoise was kailga-ailga and the trapdoor spider was kailga wurdi.

51 Data recorded by Tindale (‘Aboriginal Placename Cards’, Warki Language, AA338, Anthropology Archives, S.A. Museum). The informants are not

named. Tindale (1930-52: 273) lists Grey 1839 as a record relating to the Kondoli fire myth.

%2_N.B. Tindale (no date) ‘Strike-a-lights, Fire Flints’. Tindale collection, miscellaneous papers, Anthropology Archives, S.A. Museum.

SIGNIFICANCE OF WHALES 27

notch which supplements the hot powdered pith

engendered by the rotary process. When a spark

appears it is deftly transferred to a ball of dry, teased

grass or other easily ignitable matter,~

In the same reference, Tindale added that fine

stone grit was sometimes placed in the female part

of the apparatus to help fire-making by increasing

friction. The use of percussion methods to make

fire, generally by striking flint, together with

‘ironstone’ or iron pyrites, has been recorded in

some parts of South Australia. In the Coorong

region, Tindale’s Tangani informant, Milerum,

told him that flint, marti, was used for fire-making

and came by way of trade from south of Cape

Jaffa. The flint was used by striking it across

iron pyrites, baruke, that came from the Mount

Lofty Ranges at a place called Whale Hill, just to

the north of the western boundary of the Tangani

speakers. Powdered sun dried kangaroo dung,

fungus, bark and sandalwood scrapings were used

as tinder (narn) and kept wrapped in possum fur.

Although this method of fire-making was rarely

recorded, Charles P. Mountford and Ronald M.

Berndt (1941: 343-344) use the Karloan version

of the Kondoli myth as proof that the percussion

method existed in Australia before Europeans

arrived. The Tjirbruki mythology of the Adelaide

plains also mentions the use of iron pyrites in

making fire (Tindale 1987).

EUROPEAN SETTLEMENT

In the first few months of the official British

settlement of South Australia in 1836, Encounter

Bay came under scrutiny as a possible site for the

capital of South Australia. The large harbour and

the proximity of a supposed navigable entrance to

the Murray River were major factors in its favour.

Nevertheless, it was Encounter Bay’s suitability

for whaling activities that started settlement there.

Captain John Hart, who was familiar with the

South Australian coast through his whaling and

sealing activities during the early 1830s, was

consulted by the South Australian Colonisation

Commissioners in London (Blacket 1911: 431).%

Whaling was the first official industry in the

Colony of South Australia. Two ‘shore’ whaling

stations were set up in Encounter Bay in 1837 -—a

group from Sydney headed by Captain

Blenkinsopp and the other controlled by the South

Australia Company.*° The former was situated at

Police Point (now Victor Harbor), and the latter at

the foot of the ridge connecting Rosetta Head

(now The Bluff) with the hills. Other stations were

later set up at The Nob (now Port Elliot) and on

Granite Island. In the case of the whaling

establishments at Police Point and Granite Island,

these were considered to be of ‘lesser importance’

and soon abandoned (Newland 1921: 16).

Maritime archaeological and historical studies

have produced a list of whaling stations across the

coast line of South Australia (Angas 1847a, text

for plate XVI; Hosking 1973, legend: 2;

Kostoglou & McCarthy 1991, Table 1, p.67).

There were large whaling stations situated in

Encounter Bay at Rosetta Head (1837 to 1855)

and Freeman Nob (about 1840). On Kangaroo

Island there were establishments at Hog Bay

(1841 to 1844) and D’Estrees Bay (1843 to 1844).

There were also smaller whaling stations at the

mouth of the Onkaparinga River (from 1841 to

about 1843) and at Fishery Cove near Cape Jervis

(from 1842 to the early 1850s). Several

establishments were present on the West Coast of

South Australia at Thistle Island (1838 to 1839),

Fowler Bay (1840s), Saint Peter Island (1840s),

Streaky Bay (1843 to 1846) and Trial Bay (1845).

In the South East of South Australia, there was

the Rivoli Bay Station (1830s intermittently to the

1880s).

Two species of whale were commercially

hunted in southern South Australia, the sperm or

cachalot whale (Physeter macrocephalus) and the

southern right whale (Eubalaena australis).*’

There were two distinct seasons for European

whaling, referred to as ‘in-shore’ and ‘off-shore’.

The ‘in-shore season’ was during winter, with the

southern right whales caught as they travelled

along the coast of south eastern Australia from

Tasmania (Fig. 1). This was known as ‘bay

whaling’. The ‘off-shore season’ aimed to catch

*° _N. B. Tindale (no date) ‘Strike-a-lights, Fire Flints’. Tindale collection, miscellaneous papers, Anthropology Archives, S.A. Museum.

‘4 N, B. Tindale (no date) ‘Strike-a-lights, Fire Flints’. Tindale collection, miscellaneous papers, Anthropology Archives, S.A. Museum.

5° In 1831, Captain Hart left a sealer at Baudin Rocks in the South East of South Australia (Kostoglou & McCarthy 1991: 63). After official British

settlement in 1836, Hart became a successful merchant and politician in South Australia. He was Premier of the colony on three occasions (Blacket

1911: 431-432).

‘¢ ‘Report on Whaling in South Australia’ (1841 [reprinted in Proceedings of the Royal Geographical Society of Australasia, South Australian

Branch, 1921: 15—20]) and Cameron (1979: 23). Newland (1936) gives a fictional account of whaling in the region in ‘Paving the Way.’

For descriptions of South Australian whaling practices refer to ‘Report on Whaling in South Australia’ (1841 [1921]), Hosking (1973, explanatory

note: 1), Glover & Ling (1976), Whitelock (1985; 61, 64), Kostoglou & McCarthy (1991: 1-2) and Judd et al (1992: 13-14).

P. A. CLARKE

28

‘gTeYM IYBII UJOYINOS oY} JO sajNoI UONRIZTUL JoWUIM ZIQISSOd ey “| AUNOI

poy ulpneg \ a SWY8Z = WI]

\ .

_ ayes

\ ‘Ss

\ SS

\ SS

\ se N

uea2Q UJaYINOS

BINSUIYIdg DYIOA

p

Binsuluad 243

nN

SIGNIFICANCE OF WHALES 29

whales as they were returning from the west about

320 kilometres off the coast beyond the

continental shelf. Sperm whales were generally

only caught offshore, although, as with other

whale species, they occasionally stranded along

the coast. The ‘off-shore’ whaling was more

expensive than ‘bay whaling’, as deep-sea

activities required bigger boats. Whitelock (1985:

63) states that from 1836 to the end of 1840, 536

barrels of southern right whale oil, 158 barrels of

sperm oil, and 655 bundles of whale bone were

exported from Encounter Bay to London via

Hobart.

Impact OF WHALING Upon ABORIGINAL PEOPLE

European whalers operating in South Australian

waters utilised Aboriginal labour, being cheaply

available when required (see Clarke 1996, 1998).

For instance, it was reported in an Adelaide

newspaper in 1839 that:

We are rejoiced to hear that the fishery at Encounter

Bay is proceeding most prosperously, nearly 300

tons of oil having been already obtained. A fact

connected with it has been communicated to us

which is of an interesting and not unimportant

nature. It appears that a boat is employed in the

fishery which is entirely manned with natives. They

take their part in the occupation equally with the

white men, and are found to be not less expert than

they. If the aboriginal inhabitants are competent to

this laborious species of employment, what should

prevent them being rendered efficient in many other

paths of industry.*8

Furthermore, in 1846 Aboriginal people

manned a boat run by Barton at Encounter Bay.

In the 1830s, the Kangaroo Island sealers knew an

elderly Aboriginal man called Kondoy (Condoy),

who was described as being a ‘tribal chief? and

apparently lived at Cape Jervis. It is possible that

this man’s name referred to the whale. Since

Kondoy was a senior member of his descent

group, he may have taken on the identity of their

main nga:tji, a whale. The sealers were part of a

seafaring population originating in various

Northern Hemisphere and Pacific countries. They

and their Aboriginal wives were engaged in

various enterprises that were reliant on passing

vessels, such as gathering salt, growing vegetables

and the hunting wallabies, kangaroos and seals

for their skins. On the frontier, the whalers and

sealers were opportunistic. They were also the

first direct contact that Aboriginal people had with

Europeans. For example, in May or June 1838,

whalers on the Elizabeth were blown ashore at the

north end of Rivoli Bay (Stewart n.d. [1977: 78-

79, 89]). They set up a station there, which

attracted Aboriginal people looking for food. On

at least one occasion these whalers rowed out in a

boat to Penguin Rock with Aboriginal women,

who caught for them the fat birds after which the

islet was named. Similarly, Davies (cited Tindale

1986: 235) recorded a song in the Wirangu

language that records the careening of a whaling

boat at Denial Bay in the West Coast of South

Australia.

From the outset, the whaling station at

Encounter Bay attracted large numbers of

Aboriginal people. Gouger (1838: 53) claims that

the establishment of the whale fishery at

Encounter Bay had made this area a favourite

residence of Aboriginal people during the whaling

season (presumably referring to the winter time).

Penney noted this seasonal movement in

November 1840, when he observed Aboriginal

people leaving Encounter Bay and heading east.

Penney reports:

I found the natives going or preparing to go to the

Lower Murray. The natives of the Lower Murray

tribes usually return at the end of the Whaling

season, accompanied by some of the Encounter Bay

tribe, to their stations on this river ...°!

Leigh (1839: 163-164) describes how upon the

capture of a whale, the local Aboriginal people

came to the stations for feasts. A painting by

George French Angas shows whalebones being

used as the framework of an Aboriginal hut at the

Encounter Bay whaling stations (Fig. 2). Cameron

(1979: 4) suggests that the whalers encouraged

local Aboriginal people to camp around the

whaling station by distributing whale-meat and

rum, so that the white men had access to

Aboriginal women. In the 1840s, Richard Penney

and David Wark treated Aboriginal people

ise

The Southern Australian newspaper, 1 August 1839.

The South Australian Register newspaper, 8 August 1846. Aboriginal involvement in the whaling industry is noted elsewhere in Australia during

the 1840s, with Aboriginal whalers present at Twofold Bay in southern New South Wales (Davidson & Davidson 1988: 25-34).

© See Thomas (1925: 45-46) and Clarke (1998; 39). Also see The Advertiser newspaper, 27 December 1886. An elderly Aboriginal man named

‘Konday’ is a character in Cawthome’s ‘Kangaroo Islanders’ (Cawthome 1854 [1926: 110-111)).

*! Transcript of a letter from Penney to the Governor of South Australia, 7 January 1841. (Group 787; 1841, no.8 — AA309, Stirling collection,

Anthropology Archives, S.A. Museum),

30 P, A. CLARKE

FIGURE 2. Aboriginal people living in camps on the edge of the Encounter Bay whaling stations used whale bones

to make their shelters (G. French Angas watercolour, 1844 [lithograph version published in Angas 1847b, plate

LV]I)).

suffering from small pox and venereal diseases at

Encounter Bay (Jenkin 1979: 46-49). Penney was

concerned about the ill treatment of Aboriginal

people by whalers. Of the Encounter Bay people,

he said:

There are many individuals of this tribe, who have

been content, whilst there was any employment, to

live for months at the fisheries, and to do the work

of whalers; and one was, the last season, rated

amongst the crew of the chief headsman, Thomas,

being considered as a good a hand as many of the

white people.

The German missionary, Heinrich A.E. Meyer,

operated a school for Aboriginal people near the

whaling station. The Lower Murray people living

there were among the colony’s first Aboriginal

‘fringe dwellers’, although at this stage they were

essentially opportunistic rather than dispossessed.

At the Encounter Bay whale fishery, it is

claimed that Aboriginal men were employed to

cart blubber to pots where Aboriginal women

boiled it up in vats (Cameron 1979: 39).%

Payment was usually made in gin and tobacco.

The whalers employed at least one Aboriginal

man, named Charlie Warmer, as a ‘spotter’. He

was a member of the Kondoli clan and it was said

that Warner could also sing the whales to come in

or to go out. The whalers believed in his abilities,

paying him with rations. On one occasion, Warner

did not receive his payment through an oversight.

Sustie Wilson, an Aboriginal whaler, claimed:

So Charlie Warner ran out to a rock near the sea,

and began his chanting. A huge whale which was

lying in the bay vanished in a few seconds. The

Australian Register newspaper, 26 June 1841).

The origin of the historical source for this reference is not clear.

SS The Mail, 19 August 1932.

Penney, The South Australian Register newspaper, 21 November 1840.

Penney mentions a ‘Rey. Mr Myers of Encounter Bay’ [= H.A.E. Meyer] who was ‘improving the natives’ and studying their language (The South

SIGNIFICANCE OF WHALES 31

whaling crew dashed out, but could not even find

the wake which is usually left by the whales. They

returned, and went to old Charlie, and give him his

rations. He said, ‘Now you catch him. You go back

same place this afternoon. You catch him all right.’

The same afternoon, they found the whale in the

same place. I often saw him bring whales into the

bay, as well.

Aboriginal men, such as One Arm Charley,

Peter and Encounter Bay Bob, had experience in

the whaling and sealing industries, dating back to

before official European settlement (see Clarke

1991: 98-100; 1996: 56-59; 1998: 37). After

1836, when European colonists arrived at

Encounter Bay, these men became important

agents employed by the Europeans as trackers,

guides and translators.

In the second half of the nineteenth century,

some Aboriginal men were still engaged as whalers

(Taplin Journals, 5 May & 22 July 1861). In 1932

an Aboriginal whaler, Sustie Wilson, claimed that

many of the whalers from the ‘Encounter Bay tribe’

were much better than the Europeans. He was

reported to have said that this ‘was because they

had been throwing spears all their lives, and took to

harpooning naturally.’*® Throughout the whaling

industry, the use of Indigenous harpooners seems

to have been the practice with some teams. During

the operation of the whale fishery on Granite

Island, Aboriginal people were employed to take

wood and drinking water out to the island along a

connecting reef during low tide (Hodge 1932: 124).

Whaling was hazardous employment. For example,

when Sustie Wilson was still quite young and

working in a whaling crew, their boat was dragged

about nineteen kilometres out to sea by a huge

whale. Sustie claimed that it ‘took two days and

two nights to row back ... it was hard going too,

especially when the wind was against us. Not many

of the young men of today could have done it.’”

Transactions between Europeans and

Aboriginal people did not always occur freely.

During the 1837 season, a European named

© The Mail newspaper, 19 August 1932.

over the massacre of the Maria wreck survivors.

°8 The Mail newspaper, 19 August 1932.

Driscoll was travelling on a ‘native pad’ overland

from Encounter Bay to Adelaide when he was

murdered by ‘waddy’ (club) near Hindmarsh

Valley by his Aboriginal guide, Reppindjeri,

known by Europeans as ‘Elic’ (Cameron 1979: 4,

28, 39-42; Jenkin 1979: 52-55; Castles & Harris

1987: 11-13).”' This had resulted from a quarrel

over Driscoll’s apparent refusal to pay for sex

with one of Elic’s two Aboriginal women. The

slain man was a whaler with Captain Blenkinsopp.

Although the Aboriginal people involved kept the

circumstances quiet, a local Aboriginal woman,

Kalinga, who Europeans called ‘Sarah’, leaked the

facts. She was the wife of a Kangaroo Island

sealer and whaler, Walker. Reppindjeri was

arrested by Walker and imprisoned by authorities

for some months on the South Australian moored

near the Bluff, before he allegedly escaped and

disappeared.

Some Aboriginal people in the Lower Murray

area apparently considered that there was an

advantage in working with European colonists. At

Encounter Bay in the 1840s Penney stated that:

The Encounter Bay natives have been properly

brought up, they have never been accustomed to get

anything, without working for it, and this has not

only made them more industrious, but also made

them pay that attention to the instruments and

proceedings of Europeans, that renders them almost

equal to them as general labourers.”

This integration into a European pattern may

well have been due in part to the disruption of

hunting and gathering practices through European

occupation of the land. Tindale blames early

settlement by the whalers for loss of knowledge

of indigenous place-names amongst Aboriginal

people. He states:

Since occupation of much of the western head

waters of the two rivers [Hindmarsh and Inman] had

ceased after the early visits of white whalers, the

names of places there never learned [by his

Aboriginal informants].”

See Pullen’s account (The South Australian Register newspaper, 15 August 1840) of the investigations, involving the assistance of these three men,

* In the fiction of ‘Moby Dick or The Whale’, the main harpooner, Queequeg, was a Maori man (Melville 1851).

70 The Mail newspaper, 19 August 1932.

The ‘native pad’ was a feature of the Aboriginal landscape that Europeans started to use. Meyer (1843: 52) refers to an Aboriginal song about ‘a

fine road ... winding between the hills’ between Encounter Bay and Willunga. This was possibly the trail that Watts Newland and his party were

taken on when two Aboriginal people guided them from Adelaide to Encounter Bay in 1839 (Robinson 1975: 21). In the case of the Encounter Bay

track, this was originally just over half a metre wide, and so its use by Europeans was initially confined to foot and horse traffic (Sweetman 1928

[1988: 4]). Eventually, however, it was widened for use by coaches.

72 The Examiner newspaper, 28 January 1843.

ne

Tindale (Wunindjeri clan entry in ‘Clan Data’ folder in ‘Milerum’ manuscripts, stage 2. Anthropology Archives, S.A. Museum).

32 P. A. CLARKE

The pre-European culture of the northern parts

of the Lower Murray region was, by the twentieth

century, poorly known due to early European

settlement there (see Clarke 1994: 58, 227-231,

241-242).

At Encounter Bay, Aboriginal people adapted

to the new situation, quickly gaining skills in a

wide range of European practices. Penney

claimed:

that many of them can use the axe and the saw, and

have been employed in cutting posts and rails for

fencing, and two or three can drive bullock-drays;

they are all of them very useful in tracking cattle

and horses — they are very good hands in a boat, and

in diving — and they have rendered great services to

owners of the wrecks at Encounter Bay.”

When a drowning occurred in the Encounter

Bay and Lower Murray region, Aboriginal people

invariably assisted in the recovery of the bodies

(Linn 1988: 60).7> Although Meyer ran an

Aboriginal School at Victor Harbor (from 1840 to

about 1846), Aboriginal people were relatively

free to work for European colonists.”° Encounter

Bay people were frequent visitors to the Adelaide

region in the 1840s.”” Even with the establishment

of the Point McLeay Aboriginal Mission in 1859,

Aboriginal people in the Lower Murray region

still moved around fairly freely.

Whaling declined in the early 1840s as gaslight

in the Northern Hemisphere challenged the use of

candles and oil lamps (Blainey 1977: 115). The use

of whale oil declined further when kerosene from

American oil wells became available (Hosking

1973, chap.2: 9, 32-35; Whitelock 1985: 66). The

decreasing value of whale products led to the South

Australia Company pulling out in 1842 (Cameron

1979: 23). By this time, European activities had

already denuded much of the hills behind the

whaling stations at Encounter Bay (‘Baleineau’,

cited in Whitelock 1985: 65). Seasons of whaling

continued sporadically up to the 1870s. In about

1870, whaling recommenced at Encounter Bay,

after stopping in the 1860s (Hodge 1932: 127). For

two seasons there was an Aboriginal crew on the

whaling boats. Aboriginal labour became crucial to

the colony at times when European labour was too

expensive and scarce. Aboriginal people were

The South Australian newspaper, 21 November 1840.

considered suitable for employment that demanded

heavy use of the body. Nevertheless, in 1871 only

one whale was killed and towed to the Bluff

Station (Whitelock 1985: 66). In the early twentieth

century, Aboriginal people in the Lower Murray

area were able, on at least one occasion, to actively

hunt a whale. They killed a 5.6 metre female whale

trapped near Rabbit Island in the Coorong system.

Parts of it were dragged into a boat, cut up and

distributed among the people at the Point McLeay

Mission (Ely 1980).

DISCUSSION

Versions of the Kondoli mythology establish

that the main players were Ancestors and

associated spirit familiars of various descent

groups in the Encounter Bay area of the Lower

Murray cultural region. Here, the whale

mythology, although existing in many different

forms, was used by Aboriginal people to explain

the origin and making of fire. The involvement of

particular bird ancestors and the shark vary widely

across the corpus of this mythology. There are

also parallels between these myths and others that

do not involve the whale. In the coastal regions of

southern Australia, the observation of whales

spouting and the recognition of their warm

bloodedness appear to have been major elements

in the mythology. There is insufficient data for

areas of coastal South Australia outside the Lower

Murray to determine the totemic significance of

whales, but there is an indication that whale

strandings provided other coastal Aboriginal

groups with occasional feasts of food. The records

of who had access to the meat of stranded whales

indicate that Aboriginal people had a system of

determining ownership of food resources

originating from the sea.

From the early nineteenth century, Aboriginal

people were being incorporated into the world

economy through their participation in the marine

industries controlled from the Northern

Hemisphere. European colonists used indigenous

people as a cheap labour force. Aboriginal people

in the Encounter Bay and Adelaide areas, through

75 On 12 December 1837, Aboriginal people dragged the two surviving members of Captain Blenkinsopp’s exploration party from the Murray Mouth

in a whaleboat (The South Australian Register newspaper, 20 January 1838; Gouger 1838: 42-45; Hahn 1838-1839 [1964: 130]; Hosking 1973,

chap.1: 21-22; Cameron 1979: 4, 28). In June 1838 Aboriginal people from east of the Murray Mouth assist the survivors of the wrecked ‘Fanny’

to return to Victor Harbor (The South Australian Register newspaper, 25 August & 8 September 1838).

7° Meyer left the Encounter Bay district in 1848 and settled at Bethany in the Barossa Valley (Cameron 1979: 48).

7 The Observer newspaper, 27 April 1844.

SIGNIFICANCE OF WHALES 33

their interaction with the sealers and whalers,

were the first groups in South Australia to gain

extensive experience of Europeans. Thus whaling

at Encounter Bay provided the background to

Aboriginal and European interaction during the

British colonisation of South Australia.

ACKNOWLEDGMENTS

This paper is based on material in the author’s Ph.D

thesis, which was supervised by Chris Anderson, Peter

Smailes and Kingsley Garbett. Terry Arnott commented

upon a draft of this paper.

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Limits, and Proper Names.’ Australian National

University Press, Canberra.

TINDALE, N. B. 1986. Anthropology. Pp.235—49 in C.

R. Twidale, M. J. Tyler & M. Davies (eds.). ‘Ideas

and Endeavours—The Natural Sciences in South

Australia.” Royal Society of South Australia,

Adelaide.

TINDALE, N. B. 1987. The wanderings of Tjirbruki: a

tale of the Kaurna people of Adelaide. Records of the

South Australian Museum. 20: 5-13.

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

of the Aborigines of the Adelaide district...’

MacDougall, Adelaide. Reprinted in the South

Australian Colonist newspaper, 7 July 1840. Also

pp. 57-70 in T. A. Parkhouse (ed). 1926. ‘Reprints

and Papers Relating to the Autochthones of

Australia.” Parkhouse, Woodville.

WHITELOCK, D. 1985. ‘Conquest to Conservation.

History of Human Impact on the South Australian

Environment.’ Wakefield Press, Adelaide.

WYATT, W. 1879. Some account of ... the Adelaide

and Encounter Bay tribes. Pp.157-81 in J. D. Woods

(ed.) ‘The Native Tribes of South Australia.’ E. S.

Wiggs & Sons, Adelaide.

TAXONOMIC REVISION OF THE AUSTRALIAN LIZARD PYGOPUS

NIGRICEPS (SQUAMATA : GEKKONOIDEA)

BRIAN H. JAMES, STEPHEN C. DONNELLAN & MARK N. HUTCHINSON

Summary

Allozyme electrophoresis and analysis of scalation, colour pattern and body proportions were used

to define species boundaries in the widespread, arid adapted legless lizard Pygopus nigriceps. Three

species are recognised. Pygopus nigriceps (Fischer, 1882) is confined to the sandy deserts of central

and western Australia. Pygopus schraderi Boulenger, 1913 inhabits mainly heavier soils and rocky

substrates in the arid and semi-arid zones of eastern Australia. The third species is described a new,

and occurs in the wet-dry tropics of northern Australia.

TAXONOMIC REVISION OF THE AUSTRALIAN LIZARD PYGOPUS NIGRICEPS

(SQUAMATA: GEKKONOIDEA)

BRIAN H. JAMES, STEPHEN C. DONNELLAN & MARK N. HUTCHINSON

JAMES, B. H., DONNELLAN, S. C. and HUTCHINSON, M. N. 2001. Taxonomic revision of

the Australian lizard Pygopus nigriceps (Squamata: Gekkonoidea). Records of the South

Australian Museum 34(1): 37-52.

Allozyme electrophoresis and analysis of scalation, colour pattern and body proportions

were used to define species boundaries in the widespread, arid adapted legless lizard Pygopus

nigriceps. Three species are recognised. Pygopus nigriceps (Fischer, 1882) is confined to the

sandy deserts of central and western Australia. Pygopus schraderi Boulenger, 1913 inhabits

mainly heavier soils and rocky substrates in the arid and semi-arid zones of eastern Australia.

The third species is described as new, and occurs in the wet-dry tropics of northern Australia.

B. H. James, S. C. Donnellan, and M. N. Hutchinson. South Australian Museum, North

Terrace, Adelaide, South Australia 5000. Manuscript received 11 May 2000.

Pygopods are a lineage of snake-like lizards

endemic to Australia and New Guinea with

evolutionary affinities to the geckoes (Kluge

1987; King 1990; Donnellan et al. 1999). There

are 35 species of pygopods recognised, placed by

most recent authors into eight genera (Kluge

1974, 1976; Wilson & Knowles 1988; Storr et al.

1990; Ehmann 1992; Cogger 1996). The genus

Pygopus Merrem is considered (Kluge 1974;

Cogger 1996) to comprise two species, Pygopus

lepidopodus (Lacépéde) which occupies the

southwestern, southern and eastern margins of

Australia, and Pygopus nigriceps (Fischer) which

is widespread throughout the more xeric parts of

Australia.

Kluge (1974) found considerable phenotypic

divergence between populations of P. nigriceps

from the western and eastern parts of Australia.

They were separated by a narrow zone of

character discontinuity located approximately

along 135°E. In addition, he found that

specimens from north of 18°S in Western

Australia (WA) and the Northern Territory (NT)

tended to be more like the eastern than the

western form but Kluge had insufficient

specimens to be able to identify the northern

extension of the character discontinuity. He

suggested that two subspecies be recognised, a

western P. n. nigriceps and an eastern P. n.

schraderi Boulenger, and that the northern

specimens be treated as a taxonomically separate

unnamed, problematical set.

Kluge’s separation of P. nigriceps into two

subspecies was continued by Storr et al. (1990)

and Cogger (1996) who both ascribed the northern

WA and NT specimens to P. n. schraderi. Wilson

and Knowles (1988) suggested that northern

populations were an intermediate form that could

be tentatively regarded as undescribed. Ehmann

(1992, 1995) provided two different

interpretations for the status of P. nigriceps. In

1992, he suggested there were two separate

species, P. nigriceps whose range he gave

approximately as west of 135°E, including

northern WA and the NT, and P. schraderi, whose

range he gave approximately as east of 135°E

including Cape York Peninsula. However, by

1995 he had altered this view and suggested there

were two subspecies, P. n. nigriceps and P. n.

schraderi. Their ranges were as he had suggested

previously for P. nigriceps and P. schraderi,

except that populations from northern WA and

the NT were assigned to P. n. schraderi while

those from Cape York Peninsula were not

assigned to either subspecies.

In the present study, we have employed

multilocus allozyme electrophoresis and

morphology to examine the taxonomic status of

P. nigriceps and to clarify species boundaries.

This revision explicitly invokes the evolutionary

species concept (Simpson 1951), the significant

feature of which is that a species is a lineage

evolving separately from others and with its own

unitary evolutionary history and fate.

38 B. H. JAMES, S.C. DONNELLAN & M. N. HUTCHINSON

MATERIALS AND METHODS

Specimens

For the allozyme analysis, frozen liver samples

from the Australian Biological Tissue Collection

(ABTC) at the South Australian Museum,

Adelaide were used. The samples had been

dissected from fresh specimens and stored

continuously at —80°C. In all, liver samples from

43 specimens of P. nigriceps were available for

analysis. In addition, liver samples from nine

specimens of P. lepidopodus, chosen to cover the

known range, were used to provide a sister lineage

for the P. nigriceps species-complex, and as a

check on the authenticity of the P. nigriceps

samples. An incidental aim was to gain

preliminary data on possible cryptic speciation,

given the extensive range and known colour

variations in P. lepidopodus.

There was the potential for a very large

number of P. nigriceps specimens to be

available for morphological analysis from the

Australian state museum collections. We

concentrated on specimens from north of the

Tropic of Capricorn (23°26'30"S). All

specimens whose liver tissues were examined

in the allozyme study were examined for

morphological characters, provided that the

body was available for analysis, as were all

specimens in the South Australian Museum

collection and selected specimens from the

other state museums. Museum registration

numbers for all specimens examined

electrophoretically and/or morphologically (n =

260) are given in the Appendix. Institutional

acronyms follow Leviton et al. (1985).

Allozyme electrophoresis

Allozyme electrophoresis was conducted with

liver homogenates on cellulose acetate gels

(Cellogel, Chemetron) according to the methods

of Richardson et al. (1986). Proteins and enzyme

products of 35 presumed loci were scored. The

enzymes and other products stained, abbreviations

and Enzyme Commission Numbers are: aspartate

aminotransferase (AAT, EC 2.6.1.1), aconitate

hydratase (ACOH, EC 4.2.1.3), aminoacylase

(ACYC, EC 3.5.1.14), adenosine deaminase

(ADA, EC 3.5.4.4), alcohol dehydrogenase (ADH,

EC 1.1.1.1), carbonate dehydratase (CA, EC

4.2.1.1), diaphorase (DIA, EC 1.6.99.?), enolase

(ENO, EC 4.2.1.11), esterase (EST, EC 3.1.1.7),

fructose-bisphosphatase (FBP, EC 3.1.3.11),

fumarate hydratase (FUMH, EC 4.2.1.2),

glyceraldehyde-3-phosphate dehydrogenase

(GAPDH, EC 1.2.1.12), guanine deaminase

(GDA, EC 3.5.4.3), (S)-2-hydroxy-acid oxidase

(GOX, EC 1.1.3.15), glycerol-3-phosphate

dehydrogenase (G3PDH, EC _ 1.1.1.8),

glucose-6-phosphate isomerase (GPI, EC 5.3.1.9),

B-glucuronidase (BGLUR, EC 3.2.1.31), L-iditol

dehydrogenase (IDDH, EC 1.1.1.14), isocitrate

dehydrogenase (IDH, EC 1.1.1.42), cytosol

aminopeptidase (LAP, EC 3.4.11.1), L-lactate

dehydrogenase (LDH, EC = 1.1.1.27),

lactoglutathione lyase (LGL, EC 4.4.1.5), malate

dehydrogenase (MDH, EC 1.1.1.37), ‘malic’

enzyme (MDHP, EC 1.1.1.40),

mannose-6-phosphate isomerase (MPI, EC

5.3.1.8), nucleoside-diphosphate kinase (NDPK,

EC 2.7.4.6), dipeptidase (PEP-A, EC 3.4.13.?),

tripeptide aminopeptidase (PEP-B, EC 3.4.11.?),

proline dipeptidase (PEP-D, EC 3.4.13.?),

phosphogluconate dehydrogenase (PGDH, EC

1.1.1.44), phosphoglucomutase (PGM, EC

5.4.2.2), superoxide dismutase (SOD, EC

1.15.1.1) and triose-phosphate isomerase (TPI, EC

5.3.1.1). For the genetic analysis, geographically

proximate specimens of a single genetic type that

is, where there were no fixed allelic differences,

were pooled to form Operational Taxonomic Units

(OTUs). On this basis, 25 OTUs of P. nigriceps

were designated. Each of the P. lepidopodus

specimens was treated as a separate OTU. The

OTU localities are shown in Fig. 1 and

composition in the Appendix. Evolutionary

distances between OTUs were estimated with

Cavalli-Sforza chord distances (Cavalli-Sforza &

Edwards 1967) calculated with BIOSYS-1

(Swofford & Selander 1981). The Neighbour-

joining (NJ) algorithm, implemented in PHYLIP

version 3.5c (Felsenstein 1993) was used to build

trees from these distances. The maximum

parsimony (MP) criterion optimality was also used

to recover phylogenetic trees with each locus

considered as a character and each allele as an

unordered character state. Polymorphic loci were

encoded as uncertainties using this option for

multistate characters in PAUP* version 4.0b3.

(Swofford 1999). The data were bootstrapped to

assess confidence for individual nodes.

Morphology

Kluge (1974) assembled a large character set

for morphological analysis of the pygopods. Many

of these characters are not applicable to the genus

Pygopus and some are difficult to score in that

they depend upon arbitrary starting points or

locations on the body. Initially, we selected those

REVISION OF PYGOPUS NIGRICEPS 39

FIGURE 1. Map showing collection localities for the 34 Pygopus OTUs examined electrophoretically. See

Appendix | for a key to the OTU labels. [_] = ‘eastern’ group, © = ‘northern’ group, Il = ‘western’ group, & = P.

lepidopodus.

characters reported by Kluge as varying

significantly within Pygopus and were either

quantitative in nature, or if qualitative, defined by

unambiguous reference points. We added several

new characters that seemed likely to be useful for

taxonomic purposes. Measurements were made

with vernier callipers to the nearest mm.

The characters and their abbreviations used in

the morphological study are listed below. Where

characters were defined by Kluge (1976), they are

indicated by *; characters modified from those

used by Kluge are indicated by #.

Subnostril scale*_ (SNS). The nostril may

contact the first supralabial scale (0) or a thin

strip of the nasal scale may separate the nostril

from the supralabial (1). Kluge’s ‘subnostril

scale’ refers to the portion of the nasal

underlying the nostril.

Ventral_scales* (VS). The number of ventral

scales between the posterior edge of the mental

and the vent, including the preanal scale.

Orbital _scales*_(OS). The number of scales

along the anterior margin of the bony portion

of the orbit between the anteriormost and

posteriormost enlarged supraciliary scales.

Dorsal scale keeling* (DSK). Keels absent (0),

scales weekly keeled (1), or scales moderately

keeled (2). The keeled scales were always

unicarinate.

Dorsal scale row keeling (DSRK). The number

of dorsal rows of scales exhibiting keeling.

Preanal pores* (PP). The total number of

preanal pores.

Dorsal interorbital pattern* (DIP). Pattern on

the dorsal surface of the head between the

orbits. Varies from no significant pigmentation

to a mottled appearance, not distinctly different

from the snout, to the presence of brown or

black pigmentation forming a faint to strongly

contrasting dark bar. Scored as no significant

pigmentation, not distinctly different from

snout (0), mottled appearance not distinctly

different from snout (1), brown or black

pigmentation present as a faint bar (2), or

40 B. H. JAMES, S.C. DONNELLAN & M. N. HUTCHINSON

TABLE 1. Allele frequencies expressed as a percentage for 34 OTUs of Pygopus at 35 loci. Alleles are designated alphabetically, with ‘a’ being the most

cathodally migrating allele. Where enzymes are encoded by more than one locus, the loci are designated numerically in order of increasing mobility.

Sample sizes are given at the head of each column, except when fewer individuals were successfully typed. In the latter case sample sizes are indicated by

the number in superscript beside the first allelic frequency entry for a locus. Where allele frequencies are not given, the OTU is fixed for the allele. The

following loci were invariant: Ca, BGlur, Ldh-1, Ldh-2, Lap, Mdh and Tpi.

‘eastern’ ‘northern’

Locus WIN WID COR COO ARC CAL MID IRO- RMI CAP TCK TAN GLE FIN) MAC YUL

1 2 1 1 1 1 1 1 2 3 3 1 1 1 2 2

Aat a a a a a a a a a a d(17) a a c(50) c(25) —c(25)

(33) a(50) a(75) a(75)

b(17)

a(33)

Acoh-l b b b b b b b b b b a a b(50) b(50)_c(50) bb

a(50) a(S0) = a(S0)

Acoh-2. ¢ c d(50) cc c b b c(50) c a b(83) ob b o bl (50)

c(50) b(S50) a(17) b(50)

Acyc b b b b c(50) (50) b 2 c(75) d b b b b b b

b(50) (50) b(25)

Ada a a a a a a a a a b b b b b b b

Adh a a a a a b(50) a a a a a a a a a a

a(50)

Dia b b b a a b b(50) b b a c c c c c c

a(50)

Eno b b(S0) b b b b b b b b b(67) b b b b b

a(50) a(33)

Est c(50) b b b b b b b b b(83) b b b b b b

b(S0) a(17)

Fbp a a a a a a a a a a a a a a b(25) a

a(75)

Fumh b b b b b b b b b b b b b b b b(75)

a(25)

Gapdh b b b b - - b b b b b - b b b b

Gda a a a a a a a a a b(17) b b b b b b

a(83)

Gox b b b b b c(50) (50) c b (83) c(83) oc ce, c c(25) c¢

6650) a(50) bd7) ——-b(17) b(75)

G3pdh_ b b b b b b b b b a b b b b ~ b

Gpi a a a a - - a a a a a a a a a a

Iddh ¢ C c c c c ¢ c c c f f e fS50q - (50)

e(50) e(50)

£(50)

(50)

bG3)

a(67)

(8)

£60)

b(86)

a(14)

b

NIN

b

REVISION OF PYGOPUS NIGRICEPS

CAN BAR

a(25)

P. lepidopodus

YLE YAL ESP HAM STO

1 1 1 1 1 1 1

a a a a a c(50)

a(50)

b c c c c(50) a c

b(50)

c(S0) e(50) e(50) d(S0) d c d

b(S0) (50) (50) (50)

b b b b b b b

b - b b b b (50)

b(S0)

a a a a b a a

c - d d d d d

b c b c c a c

b b b b(50) b b b

a(50)

a b(50) a a a a b

a(50)

b b b b b b b

b a a a a a -

b b b b b b b

- b c b b c(50) —

b(50)

b - b b c a -

a a a a a a a

f c c(50) b (50) d -

b(50) b(S0)

c(50)

b(50)

41

42 B. H. JAMES, S. C. DONNELLAN & M. N. HUTCHINSON

TABLE 1. (cont.)

‘eastern’

‘northern’

Locus WIN WID COR COO ARC CAL MID IRO_ RMI CAP TCK TAN GLE FIN) MAC YUL

1 2 1 1 1 1 1 1 2

Mpi c (75) (50) b b b b b b

(25) (50)

Ndpk b b b b b b b b b

Pep-A a a a a a a a a

Pep-Bl a a a a a a a a

Pep-B2. (50) _b b d e b d d d

Pep-D (50) e&50) e(50) e e e e e e

Pgm a a a a a a a a a

Sod c c c c c c c c c

brown or black pigmentation present as a

strong bar (3).

Dorsal nuchal pattern* (DNP). The pattern on

the dorsal surface of the head and neck

immediately posterior to the parietal region

varies in a similar way to the interorbital

region. Scored as no significant pigmentation

not distinctly different from snout (0), mottled

appearance not distinctly different from snout

(1), brown or black pigmentation present as a

faint band (2), or brown or black pigmentation

present as a strong band (3).

Orbital patch (OP). Dorsoventrally orientated

darkly pigmented patch around and/or below

the orbit extending to the supralabial scales and

sometimes to the infralabial scales. Scored as

absent (0), faint (1), moderately intense (2) or

intense (3).

Narial patch (NP). Dorsoventrally oriented

darkly pigmented patch or streak around and/or

3 3 1 1 1 2 2

a a a a a al a

b b b b b b b

b b b b b b b

d b b(50) b b b b

a(50)

b b b b b b b

b(75) b c b b (50)! b

a(25) b(50)

a b a a a al a

c(50) c f(50) c c(50) f(50) c

b(33) (50) b(50) (25)

a(17) b(25)

e(67) ~—-g(17)- £650) g (50) g(S0) ff (25)

(33) f(83) b(S0) £50) (50) (75)

a a a a a c(25) a

a(75)

b c(83) oc c c c c

b(17)

scales and sometimes to the infralabial scales.

Scored as absent (0), faint (1), moderately

intense (2) or intense (3).

Contrast of lateral head pattern (LHP). Relative

intensities of the orbital and narial patches may

vary. Whereas some specimens or species show

an equally intense development of both

patches, others may have a noticeably weak

expression of the narial patch (even absence)

compared to the orbital patch. The converse

(narial patch more strongly developed than

orbital patch) was not observed. Scored as

intensities of both patches equal (1), differing

by one on the NP and OP scores (2), differing

by two (3), or differing by three (4).

Snout-vent length* (SVL). The horizontal

distance between the median anterior-most

extreme of the snout and the median posterior-

most extreme of the middle preanal scale.

These characters were scored for each specimen

below the nostril extending to the supralabial from which liver samples had been taken for

REVISION OF PYGOPUS NIGRICEPS

‘western’

ILL OLY MAB LAV NIN CAN BAR BOO YLE

1 3 7 2 1 1 2 1 1

- a a - a - a a a

- b c7) =»b b b b b b

b(93)

b b b(86) b b - b b b

a(14)

b b b b c(50) b b e(50) b

b(50) b(50)

b b b® b b b b b b

c(25) (33) b* b b - b b b

B75) b(67)

a c(83) a b(25) (50) - c(25) (50) a

a(17) a(75) _—_a(S0) b(75) —_a(S0)

c c f(21) c¢ c - c c c

c(79)

- f h(7)—-g(50)_ f g g(25) (50) f

£(86) — e(25) £(50) (50)

e7) (25) e(25)

a a a a a a b(SO0) a a

a(50)

c c c c c c c c c

43

P. lepidopodus

YAL ESP HAM STO SYD WAL KAR RM2 CIN

1 1 1 1 1 1 1 1

- a a a b(50) - b(50) a a

a(50) a(50)

- a a a a a a a a

b b b b a - - b a

c(50) a a a

t £(50) b f f e b £(50)

b(50) (50)

a a a a a a a a a

a a a a a a a a a

allozyme electrophoresis, except for three

‘western’ specimens for which vouchers were not

available for examination (ABTC 6588, 31799,

31812). For statistical comparisons of the

morphological characters except SVL, the

significance of differences between taxa was

assessed with the non-parametric Mann-Whitney

U test. For comparisons of SVL, unpaired

Student’s f tests were used after testing for

departures from a normal distribution for each

taxon (Shapiro-Wilks test) and pairwise tests of

equality of variances (Levene test). All tests were

two-tailed with an a set at 0.05 and were carried

out with STATISTICA (Statsoft Inc. 1997).

RESULTS

Table 1 shows the allelic profiles of the 25

OTUs of P. nigriceps and nine OTUs of P.

lepidopodus for the 35 loci. These data were

converted into matrices of percentages of loci

showing fixed allelic differences (FD) and

Cavalli-Sforza chord distances between OTUs

(not shown). A fixed allelic difference occurs at a

locus when the two samples under comparison

share no alleles (Richardson et al. 1986).

Richardson et al. (1986) argued that percentage of

loci showing fixed allelic differences is an

appropriate genetic distance metric for species

boundary studies and is relatively unaffected by

small sample size. We present a phenogram

constructed from Cavalli-Sforza chord distances

between OTUs by NJ (Fig. 2A). A heuristic search

under MP found 77 335 equally most

parsimonious trees of length 53 steps. A strict

consensus of the equally most parsimonious trees

with bootstrap proportions from 10 000

pseudoreplicates is presented in Fig. 2B. For the

NJ and MP analyses adjacent populations between

which there were no fixed allelic differences were

pooled and allele frequencies recalculated, making

44 B. H. JAMES, S.C. DONNELLAN & M. N. HUTCHINSON

RMI eastern

CAP northern

FIN GLE ILE MAC YUL

MAB OLY

LAV NIN

BAR BOO YLE

CAN TAN TCK

western

STO P. lepidopodus

eastern

ARC

coo

‘WID COR

78 CAP

BAR BOO YLE

MAB OLY

FIN GLE ILE MAC YUL

CAN TAN TCK

LAV NIN

northern

western

syD P. lepidopodus

FIGURE 2. (A) A Neighbour-joining phenogram

constructed from Cavalli-Sforza chord distances

between OTUs; and (B) a strict consensus tree of more

than 2000 equally most parsimonious trees found with a

heuristic search. Bootstrap proportions >70% from 2000

pseudoreplicates are indicated to the left of relevant

nodes.

a total of 14 OTUs within P. nigriceps for the

final analyses.

The allozyme data (Fig. 2 and Table 1) show

that the P. nigriceps OTUs are genetically more

similar to one another than they are to any of the

P. lepidopodus OTUs. Monophyly of the P.

nigriceps OTUs received strong support (78%)

from bootstrapping (Fig. 2B). The P. lepidopodus

OTUs exhibit considerable genetic heterogeneity

suggesting that more study is warranted.

Within P. nigriceps, both the NJ and the MP

analyses are concordant in showing the presence

of three lineages herein designated ‘eastern’,

‘northern’, and ‘western’. However, strong

bootstrap support (>70%) was found only for the

monophyly of the ‘eastern’ lineage (Fig. 2B). The

primary split between the P. nigriceps OTUs

occurs at a minimum of 14% FD and broadly

distinguishes two groups, an ‘eastern’ and a

‘western’ group. These two groups show

diagnostic allozyme differences at four loci (Ada,

Dia, Gda and Iddh) and other differences that are

diagnostic for some of the OTUs between each

group at eight loci (Aat-2, Acoh-1, Acoh-2, Acyc,

Fumh, Pep-A, Pep-B1 and Pep-D). There is

genetic divergence within both the ‘eastern’ and

‘western’ groups with seven loci (Acoh-2, Acyc,

Dia, Gox, Mpi, Pep-B2 and Pep-D) displaying

fixed allelic differences between one or more

OTUs in the ‘eastern’ group and six loci (Acoh-1/,

Acoh-2, Eno, Iddh, Pep-BI and Pep-D) displaying

fixed allelic differences between one or more

OTUs in the ‘western’ group. The greatest

intragroup genetic divergence, at an average of

10% FD, is between the OTUs WIN, WID and

COR from central Queensland (Qld) and the

‘eastern’ group OTUs situated in the corridor

between the eastern and western deserts. This may

reflect the well-known climatic-ecological barrier

of the Simpson Desert on patterns of speciation

(Cracraft 1986, 1991).

None of the OTUs of the ‘eastern’ and

‘western’ groups were found in strict sympatry.

However, ‘eastern’ and ‘western’ groups were

found in regional sympatry, i.e. within 30-80 km

of each other, in the Coober Pedy/Mabel Creek

area (OTUs COO and MAB). Fixed allelic

differences at six loci (Ada, Dia, Gda, Iddh, Pep-

A and Pep-B2) were found between the two

OTUs. Given the null hypothesis that two OTUs

are sampled from a single population of eight

individuals, the probability of not observing a

heterozygote at the six loci showing fixed allelic

differences can be derived from the Hardy-

Weinberg equation as (1-[2 x 0.125 x 0.875])** =

7.035 x 10° (see Richardson et al. 1986 for a

fuller explanation of this approach). Hence, the

null hypothesis can be rejected and a reasonable

alternative hypothesis is that two species are

present in the Coober Pedy/Mabel Creek area.

A second split among the P. nigriceps OTUs

occurs at a minimum of 21% FD and distinguishes

REVISION OF PYGOPUS NIGRICEPS

the ‘eastern’ and ‘western’ groups from a single

‘northern’ OTU, CAP. There are diagnostic

allozyme differences between the ‘eastern’ and

‘northern’ groups at six loci (Acoh-2, Acyc, Ada,

G3pdh, Pep-B1, and Sod) and between the

‘western’ and ‘northern’ groups at five loci (Acyc,

Dia, G3pdh, Iddh and Mpi). In addition, there are

other differences that are diagnostic for some of

the OTUs between the ‘northern’ and other two

groups at 11 loci (Acoh-1, Acoh-2, Dia, Gda,

Iddh, Mpi, Pep-A, Pep-Bl, Pep-B2, Pep-D and

Sod).

The magnitude of the genetic differences

encountered, the evidence of separate evolutionary

histories of the ‘eastern’, ‘western’ and ‘northern’

groups and direct evidence of lack of gene flow

between two of the lineages is sufficient to reject

the null hypothesis of a single species within P.

nigriceps. Consequently, we considered the

groups as three separate species for the purposes

of morphological examination. The ‘eastern’ and

‘western’ groups correspond roughly in

45

geographic location with Kluge’s (1974)

suggested ranges for P. n. schraderi and P. n.

nigriceps respectively. The ‘northern’ group

comprises only one population in the samples

available for electrophoresis thus preventing any

delineation of its geographical distribution on

allozyme data alone. These three groups proved to

have a distinctive suite of morphological

characters as outlined in the following analysis.

A summary of the variation in morphological

characters is shown in Table 2 for the three

groups. The data in Table 2 show that a suite of

morphological features varies concordantly with

the electrophoretic data. This table therefore

provides the basic morphological characters for

separating specimens into the ‘eastern’, ‘northern’

and ‘western’ groups.

The ‘western’ group is easily separated from

the other two groups in that: the nostril is

separated from the first supralabial scale by the

nasal (‘subnasal scale’ present); dorsal scale

keeling is absent; the mean number of ventral

TABLE 2. Morphological statistics for specimens of Pygopus for which allozyme data were collected. E = ‘eastern’

group, N = ‘northern’ group, W = ‘western’ group. See text for character abbreviations. X = mean, S = standard

deviation, R = range, n = sample size, * P for two-tailed Mann-Whitney U or unpaired Student’s f tests, tests for

normal distribution" and equality of variances’ were not significant. See Materials and Methods for the definitions

of characters.

Univariate Statistics

Character Group E (n=11) N (n=3)

SNS X (S) 0 (0) 0 (0)

VS x (5) 110.3 (4.6) 109.7 (1.2)

R 105-122 109-111

OS X (S) 9.18 (1.17) 10.7 (1.5)

R 7-11 9-12

DSK X (S) 2(0) 1(0)

DSRK x (S) 12.4 (0.8) 7.67 (1.58)

R 12-14 6-9

PP xX (S) 13.0 (1.2) 13.3 (0.6)

R 12-15 13-14

DIP x (S) 0.45 (0.52) 0.33 (0.58)

R 0-1 0-1

DNP X (S) 2.09 (0.83) 1.33 (1.15)

R 0-3 0-2

OP x (S) 1.91 (0.70) 2.67 (0.58)

R 1-3 2-3

NP X (S) 1.82 (0.60) 0

R 1-3 -

LHP x (S) 1.09 (0.30) 3.67 (0.58)

R 1-2 3-4

SVL x (S) 147.6 (23.1)" = 136.0 (7.9)"

R 96-178 130-145

Pairwise Statistical

Comparisons*

W (n=26) E/N E/wW N/W

1 (0) 0.88 <0.001 <0.001

130.8 (5.9) 1.0 <0.001 <0.001

114-140

11.8 (1.6) 0.17 <0.001 0.35

10-16

0(0) 0.005 >0.001 <0.001

0 (0) 0.005 >0.001 <0.001

10.2 (1.1) 0.45 <0.001 <0.001

8-12

2.32 (.80) 0.78 <0.001 0.004

0-3

2.92 (0.28) 0.29 0.001 0.002

2-3

2.44 (0.71) 0.37 0.09 0.89

1-3

1.2 (1.04) 0.005 0.02 0.06

0-3

2.24 (0.88) 0.005 <0.001 0.019

14

156.1 (33.2)" 0.42" 0.45” 0.31

79-204

46 B. H. JAMES, S.C. DONNELLAN & M. N. HUTCHINSON

scales is greater than 120; the mean number of

preanal pores is less than 12; there is usually a

well-developed blackish interorbital bar.

The ‘eastern’ group can be separated from the

‘northern’ group in that: The dorsal scales are

moderately keeled with keeling occurring over ten

or more rows in the ‘eastern’ group but weakly

keeled over fewer than ten rows in the ‘northern’

group. The orbital and narial patches are present

and are of moderate and equal intensities in the

‘eastern’ group, while in the ‘northern’ group the

orbital patch is more strongly expressed than the

nasal patch in any individual, and the nasal patch

may be completely absent.

These characters were then used to separate the

museum specimens for which tissues had not been

available for allozyme electrophoresis. The great

majority of specimens were readily separated (n =

251) but some difficulty was experienced in

separating six ‘eastern’ and ‘northern’ specimens

that had ‘washed out’ body patterns together with

indeterminate scale keeling. The consistent

allozyme and morphological differences between

the three groups is sufficient for each group to be

given species status. The ‘western’ group is P.

nigriceps, the ‘eastern’ group P. schraderi and the

‘northern’ group P. steelescotti sp. nov. A map

showing the geographic distribution of the three

species is shown in Fig. 3. Note that the

distributions of P. nigriceps and P. schraderi

overlap in south-western Queensland (Qld),

eastern NT and central and eastern SA; and those

of P. nigriceps and P. steelescotti sp. nov. overlap

in northern NT and WA; and all three species are

found in eastern NT.

Once separated into the three species, the

characters used in the morphological study on the

electrophoresed specimens were scored. The

results are summarised in the species descriptions

given below. An additional character of Kluge,

tail length, was added. It was measured as the

horizontal distance between the posterior-most

extreme of the middle preanal scale and the tip of

the tail. Measurements were made only on those

few specimens with complete and unregenerated

tails.

FIGURE 3. Map showing geographic distribution of specimens of Pygopus examined for morphological characters.

W@ = P. nigriceps, © = P. steelescotti, | = P. schraderi.

REVISION OF PYGOPUS NIGRICEPS 47

The P. nigriceps species complex conforms to

Kluge’s (1976) definition of the genus Pygopus.

The dorsal surface of the head is covered with

large and small scales, there are between three

and five postmental scales, 21 or more midbody

scale rows, nine or more preanal pores and the

dorsal body scales are often keeled. The three

species of the complex differ from the only other

species in the genus, P. lepidopodus, in that the P.

nigriceps complex has immaculate ventral

surfaces (versus boldly variegated with black),

smooth to moderately keeled dorsal scales (versus

strongly keeled) and there is no dorsal snout

pattern.

Pygopus nigriceps (Fischer, 1882)

Cryptodelma nigriceps Fischer, J. G. (1882), p.

290. Holotype SMNS 2259, Nickol Bay, WA.

Type lost, believed destroyed during World War

II (Schliiter and Hallermann 1997).

Delma (Cryptodelma) baileyi Giinther, A. (1897),

p. 170. Holotype BMNH specimen, now lost

(Kluge 1974), from the neighbourhood of Cue,

WA.

Pygopus territorianus Wells, R. W. and

Wellington, C. R. (1985), p. 16. Holotype AMS

R56823, near Tennant Creek, NT.

Notes

Both of the names that could be applied to the

‘western’ species are based on type specimens that

are now lost. Fortunately, both have fairly precise

type localities, which place them well outside the

known distribution of the other two species.

Cryptodelma nigriceps was described and figured

by Fischer (1882) who proposed the new genus

for this species, which appeared to combine the

smooth scalation of Delma with preanal pores as

in Pygopus. Fischer acknowledged that his very

small specimen (SVL 64 mm) was almost

certainly a juvenile, and its SVL is smaller than

any we measured, although we did not extensively

sample juveniles. The illustrations clearly depict

the V-shaped preanal pore row (11 pores), large

hindlimb flaps and head scalation (form of the

two frontal shields and double row of loreal

scales), a combination of features confined to

Pygopus (sensu Kluge 1974). The dark head and

nape markings are described and figured as

confluent, unusual in specimens of the ‘western’

species but not unknown (e.g. SAMA R22932

from Barradale, WA).

Giinther (1897) described his new species,

Delma (Cryptodelma) baileyi, from an immature

specimen (SVL 90 mm), noting that it was very

close to Fischer’s nigriceps but differed in its

lower midbody scale count (22, versus 26 or 28).

Fischer, however, implicitly included the ventrals

in his midbody count, making his specimen’s

likely dorsal count 24 or 26, 24 being a frequent

count in WA specimens. As with C. nigriceps,

Giinther’s specimen had smooth scales and 11

preanal pores, both characteristic features of the

‘western’ group. Giinther’s illustration shows

distinct black interorbital and nuchal colouring,

with blackish narial and orbital patches extending

on to the lower labials again features typical of

the ‘western’ species.

Wells and Wellington (1985) proposed Pygopus

territorianus as a new species from the Northern

Territory, but provided little justification. They

stated that this species could be ‘readily diagnosed

by consulting existing descriptive references’ and

the figure of a typical specimen (possibly the

holotype) in Swanson (1976; Pl 34). They also

added that the species lacks the distinctive keeling

of the body scales seen in Pygopus klugei (q.v. =

P. schraderi) and lacks the colour pattern

(‘reticulated patterning’) of P. schraderi. The

holotype is a specimen of the western form, and

we therefore regard P. ferritorianus as a junior

synonym of C. nigriceps. The body scales are

completely smooth, the nostril is separated from

the first upper labial and there are 11 preanal

pores. The ventral count is relatively low (120)

for the western form, but still within the sample

range, and the colour pattern is markedly ‘faded’,

with little black pigment remaining on top of the

head, a trend in some populations of all of the P.

nigriceps complex. The tail shows the strong

pattern of dark-edged scales typically present in

the ‘western’ form.

Neotype

To stabilise the name, a neotype, WAM

R102063, has been selected for Cryptodelma

nigriceps, from the same geographical area as the

lost type (see notes). It was collected on the Yule

River, WA (20°40'S, 118°21'E) by D. Robinson in

1990.

Diagnosis

A large pygopod (SVL up to 227 mm.) differing

from other Pygopus in having smooth dorsal

scales, the nostril entirely contained within the

nasal, usually 120 or more ventral scales, and

fewer than 14 preanal pores.

48 B. H. JAMES, S. C. DONNELLAN & M. N. HUTCHINSON

FIGURE 4. Representative head patterns of three

species of Pygopus: (A) P. nigriceps (SAMA R48749);

(B) P. steelescotti (NTM R20546); (C) P. schraderi

(SAMA R46370).

Description

Ventral scales 114-143 (X = 130.0, S=5.9, n=

116); orbital scales 9-16 (K = 11.6, S=1.4,n=

118); nostril always completely enclosed by the

nasal, separated from first supralabial scale (n =

119); preanal pores 8-14 (X = 10.8, S= 1.1, n=

117); snout-vent length 73-227 mm (X = 157.6, S

= 33.7, n = 118); tail length ranges from 104% of

SVL in smallest juveniles to 165% of SVL in

largest specimens; no dorsal scale keeling (n =

118).

Colouration

In preservative, the basic head, body and tail

colour is light tan to reddish brown. Scales, except

for ventral scales, are speckled with brown or

black pigmentation. Head, body and tail patterns

are caused by the relative intensities and widths of

the pigmented scale margins or by individual paler

and darker speckled scales. Head patterns are

formed with brown to black pigmentation and

body and tail patterns by light to dark brown

pigmentation. Ventral surfaces are immaculate

with the scales a creamy-white or silver colour.

There is a dark brown or black pigmented bar

present on the dorsal surface of the head between

the orbits which extends as a teardrop-shaped

orbital patch descending to the supralabial scales

and sometimes to the infralabial scales (Fig. 4A).

A further area of dark pigmentation, but less

intense, is found around the nostrils, again

extending to the supralabial scales and sometimes

to the infralabial scales. A brown to black nuchal

band extends to the ear openings. There is no

distinct body patterning. The tail is marked, often

strongly, in the form of dark brown backward

pointing chevrons that converge on the vertebral

line. The smallest juveniles (SVL about 75 mm)

are sandy-yellow with low levels of scale

pigmentation except for the heavily black

pigmented and contrasting head pattern. There is

no distinct body pattern in juveniles and only

weak light-brown chevrons on the tail. Coloration

in life is shown in photographs in Storr et al.

(1990: Pl 20, Figs 3-4), Glasby et al. (1993: Pl 4,

Fig. 12), Cogger (1996: p. 297).

Distribution

West of 136°E and the far south-western corner

of Queensland (Qld) (Fig. 3).

Pygopus schraderi Boulenger, 1913

Pygopus schraderi Boulenger, G. A. (1913), p.

564. Holotype BMNH 1946.8.27.2 (formerly

1913.7.28.2), collected at Milparinha [sic =

Milparinka], NSW by P. Schrader.

Pygopus klugei Wells, R. W. and Wellington, C.

R. (1985), p. 16. Holotype AMS field series

28686 (now registered as AMS R116980), 6.2 km

S of Big Warrambool, NSW.

Notes

The holotype specimen is in good condition,

with the colour pattern still readily discernible. It

has an equal development of the orbital and narial

REVISION OF PYGOPUS NIGRICEPS 49

patches, but the interorbital area is only weakly

pigmented. Although very young (SVL 65 mm), it

has distinct low keels on the dorsal scales. There

are 14 preanal pores. Boulenger (1913) gave the

ventral count as only 97 pairs (enlarged scales

only); the ventral count using our (Kluge’s)

method is 109. The characteristics of the type are

completely concordant with the ‘eastern’ species

in the complex.

Wells and Wellington’s (1985) Pygopus klugei

is a junior synonym of P. schraderi. The holotype

specimen has 12 dorsal scale rows with distinct

keels, there are 102 ventrals and the nostril

contacts the first supralabial. There is a complex

colour pattern on head body and tail, including

well developed eye and nostril patches. Wells and

Wellington were clearly influenced by the

purported lack of keeling of the holotype of P.

schraderi, a persistent misapprehension due to the

rudimentary keeling in juveniles, including the

holotype. Two illustrations said by Wells and

Wellington to be P. schraderi (Cogger 1983;

plates 104 and 495) show, respectively, a juvenile

P. nigriceps and an adult P. schraderi.

Diagnosis

A large pygopod (SVL up to 198 mm) having

unicarinate dorsal scale keeling extending over 10

or more rows, dark teardrop-shaped patches under

both the orbits and nostrils, fewer than 120

ventrals, usually 13 or more preanal pores, nostril

always in contact with the first supralabial scale

and often strongly marked body and tail

patterning.

Description

Ventral scales 100-122 (X = 109.4, S=4.9,n=

85); orbital scales 6-13 (XK = 9.87, S = 1.27, n =

85); nostril always in contact with first supralabial

scale (n = 92); preanal pores 11-17 (XK = 13.8, S=

1.4, n = 85); snout-vent length 70-198 mm (X =

142.6, S = 29.4, n = 91); tail length ranges from

99% of SVL for smallest juveniles to 170% of

SVL for the largest specimens; dorsal scale rows

with either weak or moderate keeling, usually

moderate (80%), (X = 1.88, S = 0.28, n = 86)

extending over 9-14 dorsal rows (X = 11.7, S =

1.8, n = 86).

Colouration

In preservative, the scales are pigmented in a

manner similar to P. nigriceps. The background

colour of the head, body and tail can vary from

light tan to dark grey. The head is usually of

mottled appearance. Head patterning is similar to

that of P. nigriceps except that there is no

pigmented band on the dorsal surface of the head

between the orbits, the relative intensities of the

pigmented bands below the orbits and nostrils

are approximately equal and the nuchal band is

frequently indistinct (Fig. 4B). Often, there is a

strongly marked body pattern in the form of a

longitudinal series of light or dark brown

uniformly pigmented scales on the dorsal surface

and upper flanks giving the impression of

discrete broken lines. Many individuals have an

almost continuous stripe along the vertebral line.

On the lower flanks is a similar series of discrete

broken lines caused by a scattering of creamy-

white scales. The tail patterning is in the form of

narrow crossbands (occasionally posteriorly

pointing chevrons converging on the vertebral

line). The body patterning is caused by

individual scales of differing levels of

pigmentation whilst the tail patterning is caused

by scale margin pigmentation. The intensity of

patterning is very variable with many specimens

being strongly marked, others having a light tan

overall colour and a washed-out appearance,

whilst still others are heavily pigmented all over.

Body patterning does not appear to be

geographically correlated. The smallest juveniles

are similar in size and colouration to those of P.

nigriceps. Head patterns are pronounced and are

brown or black. There is no distinct body pattern

and only weak light brown crossbands or

chevrons on the tail. There is weak dorsal scale

keeling.

Colouration in life is shown in photographs in

Wilson and Knowles (1988: Fig. 263), Swan

(1990: p. 47), Ehmann (1992: p. 102).

Distribution

East of 135°E, south of 17°S and west of the

Great Dividing Range in New South Wales and

Victoria (Fig. 3).

Pygopus steelescotti sp. nov.

Holotype

NTM R20546, collected at Cape Crawford, NT

(16°34.3'S, 135°57.9'E) by P. Horner, 1994.

Diagnosis

A large pygopod (SVL up to 185 mm) similar

to P. schraderi except for weaker unicarinate

dorsal scale keeling usually extending over less

than 10 rows and a less complex color pattern,

50 B. H. JAMES, S.C. DONNELLAN & M. N. HUTCHINSON

with a dark orbital patch but no or very weak

nasal patch.

Description

Ventral scales 103-125 (XK = 114.2, S=4.8, n=

43); orbital scales 7-13 (X = 9.86, S = 1.10, n =

43); nostril always in contact with first supralabial

scale (n = 44); preanal pores 12-17 (X = 14.0, S=

1.0, n = 43); snout-vent length 79-185 mm (X =

139.0, S = 24.2, n = 43); tail length ranges from

100% of SVL for smallest juveniles to 152% of

SVL for the largest specimens; dorsal scale rows

keeling either absent or weak, usually weak

(86%), (X = 0.86, S = 0.35, n = 44) extending over

0-13 dorsal rows (X = 7.1, S = 3.46, n = 44).

Colouration

In preservative, the background head, body and

tail colour is light tan to sandy-yellow with

patterns similar to those found on the less heavily

patterned individuals of P. schraderi. There is

often no pigmentation around and below the

nostrils; if it is present it is usually faint in

intensity compared to the teardrop below the

orbits (Fig. 4C). Many specimens are light tan in

colour and have a washed out appearance and,

unlike P. schraderi, there are no strongly

patterned or darkly coloured specimens. The

smallest juveniles are similar to those of P.

schraderi differing only in the head pattern and in

having no (or extremely weak) dorsal scale

keeling. Colouration in life is shown in

photographs in Wilson and Knowles (1988: Fig.

264) and Ehmann (1992: p. 101).

Etymology

The species is named in honour of the late Dr

Colin Steele-Scott, a keen supporter of the South

Australian Museum.

Distribution

North of 22°S in NT, Qld and WA (Fig. 3).

DISCUSSION

When considered as a single species, the

concept of P. nigriceps that existed prior to this

study was of an extremely successful, ecologically

generalised arid zone lizard, adapted to a broad

habitat range. The three species that we now

recognise are each more restricted ecologically, as

well as geographically. Pygopus nigriceps (s.s.) is

primarily a sandy desert species, at least in Central

Australia. Pygopus schraderi occupies a range of

habitats, and while it has been recorded from

sandy terrain (e.g. far eastern S.A.) it is typically

found on rocky hillsides or clayey or stony flats.

Pygopus steelescotti is less well-known with

respect to habitat selection but differs again in

being confined to the wet-dry tropical belt across

northern Australia.

All three species in the complex may be strictly

nocturnal, relatively unusual in pygopods, most of

which engage in significant diurnal activity (Shea

1993). Field experience with both P. nigriceps

and P. schraderi in South Australia indicates that

specimens are only seen actively foraging at night,

and pitfall trapped individuals are taken only

overnight, not by day (M. Hutchinson pers. obs.).

Of the three species, P. nigriceps is the most

easily identified, based on its completely smooth

dorsal scalation, the nostril completely contained

by the nasal, and the high ventral and low preanal

pore counts. By contrast, the other two species are

very similar, exacerbated by the tendency of

geographically proximate populations of P.

schraderi to be the most weakly patterned and

therefore most similar to P. steelescotti. The

similarity of P. schraderi and P. steelescotti is

such that specimens from the potential area of

contact along the southern margin of the wet-dry

tropics in the NT and Qld should be carefully

checked. At present, P. steelescotti seems

consistently identifiable by the lower number of

keeled dorsal scale rows (usually nine or fewer),

weakly developed keeling on dorsal scale rows

and differential development of the dark nasal

(absent or weak) and orbital (moderately well

developed) patches.

Further taxonomic work will be useful to help

in defining the variation and degree of sympatry

of the three dark-headed species of Pygopus.

Taxonomic work is also further warranted on the

northeastern Qld populations of P. lepidopodus.

Based on our samples, the Cooktown specimen

appears to be completely distinct from the more

southerly populations, which themselves show

some heterogeneity in morphology and

electrophoretic markers. The status of the eastern

Australian P. J. squamiceps Gray is yet to be

properly assessed.

ACKNOWLEDGMENTS

We thank B. Miller for assistance in the initial stages

of this study, C. James, R. Murphy, T. Schwaner for

collection of specimens and J. Armstrong for assistance

with figure preparation.

REVISION OF PYGOPUS NIGRICEPS 51

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1990. ‘Lizards of Western Australia III. Geckos and

Pygopods’. Western Australian Museum: Perth.

SWAN, G. 1990. ‘A Field Guide to the Snakes and

Lizards of New South Wales’. Three Sisters

Publications: Winmalee, NSW.

SWANSON, S. 1976 ‘Lizards of Australia’. Angus and

Robertson: Sydney.

SWOFFORD, D. L. 1999. PAUP*: phylogenetic

analysis using parsimony. Version 4.0b2a. Sinauer

Associates, Inc.: Sunderland Massachusetts, USA.

SWOFFORD, D.L. & SELANDER, R.B. 1981.

BIOSYS-1. A computer program for the analysis of

allelic variation in genetics. Journal of Heredity 72:

281-283.

WELLS, R. W. & WELLINGTON, C. R. 1985. A

classification of the Amphibia and Reptilia of

Australia. Australian Journal of Herpetology,

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WILSON, S.K. & KNOWLES, D. G. 1988. ‘Australia’s

Reptiles: A Photographic Reference to the Terrestrial

Reptiles of Australia’. Collins: Sydney.

52 B. H. JAMES, S.C. DONNELLAN & M. N. HUTCHINSON

APPENDIX

Museum registration numbers for Pygopus specimens

examined. The superscript " denotes specimens which

were used in the electrophoretic study. The superscript *

denotes specimens used in the electrophoretic study

where the body was not available for morphological

analysis. Abbreviations in bold upper case refer to

OTUs designated in the allozyme electrophoresis

analysis. Institutional acronyms (underlined) follow

Leviton et al. (1985) and ABTC = Australian Biological

Tissue Collection, South Australian Museum.

Pygopus nigriceps: AMS R60245-6. NTM CAMRS31,

NTM R30, R157-8, R291-3, R321, R444, R829, R905,

R1427-8, R1630, R2209, R2216, R2356, R2532,

R3228, R3316, R5314, R5907, R6387, R7092, R9530,

R9691-6, R9701, R9703, R9808, R14021,

R15153°(TAN—30 km SW Sangster’s Bore, Tanami

Desert, NT), R17571, R18058, R18079,

R20670°(FIN—Finke Gorge National Park, NT). QM

J28440, J33389. SAMA ABTC6588"*(MAB—Mabel

Creek Station, SA), ABTC31799"*(MAC—MacDonnell

Ranges, NT), ABTC31812(MAC), R600, R876,

R4791, R8116, R12911, R15548, R16279, R16768,

R17459, R19251, R20711°(OLY—Olympic Dam, SA),

R21029/33"(OLY), R21035, R22932-3°(BAR—8 km S

Barradale, WA), R23197°(LAV—134 km ENE

Laverton, WA), R23908°(LAV), R26198-200°(MAB),

R26202°(MAB), R26222°(MAB), R26385, R26646,

R26686°(MAB), R28546°(BOO—near Boologooroo

Homestead, WA), R33824, R34003°(NIN—Ninghan

Homestead, WA), R36152°(YUL—Yulara, NT),

R36169*(YUL), R37139, R38785-6°(TCK—Tennant

Ck, NT), R38824*(TCK), R38841°(GLE—near Glen

Helen, NT ), R42026°(LL—21 km WSW Illintjtja,

SA), R45264, R48608, R48668, R48749, R48765,

R48792, R48830, R48928, R49156, R49310. WAM

R5328, R5329, R13350, R19239, R26071-2, R30930,

R36327, R40238, R52727, R64002, R64704, R69525,

R73631, R73842, R75119, R75143, R79010-1,

R81513, R82599, R83574, R94761, R95028,

R94957°(CAN—Canning Stock Route, WA), R95670,

R102055, R102061, R102063°(YLE—Yule River,

WA), R103677.

Pygopus schraderi: AMS R6691, R8974—-5, R26083,

R65966. NTM R31, R8526. QM J11029, J1266, J2917,

J2918, J5092, J5238, J7282, J7288, 37473, J7489,

J7919, J8008, J8436, J8882, J9116, J9241, J13010,

J13028, J13560, J21444, J21965, J22714, J23319,

J23677, J24988, J25386, J33390, J33391, J33392,

J33393, J33394, J33395, J35362, J37060, J40323,

J40701, J40717, 344420, J44711, 344942, 346952,

J48459, J51198, J51527, J51717, J52546, J52547,

J52778, J52863, J54617, J54618, J54900, J55109,

J57197, J57228, J58079, J5921, J59359, J59931,

J61825. SAMA R2234, R4996, R5051, R5386, R5397,

R5626, R5893, R9860, R11752, R16712,

R23120°(RM1—Mt Remarkable National Park, SA),

R23269"(RM1), R28389°(IRO—25 km NW Iron Knob,

SA), R28927, R30403°(COO—25 km NNW Coober

Pedy, SA), R40948°(MID—Middleback Homestead,

SA), R42131, R42750°(WIN—15 km S Winton, Qld),

R42947°/63°(WID—85 km W Windorah, Qld),

R44808°(COR—Cordillo Downs, SA),

R46247°(ARC—17 km NNE Arckaringa, SA),

R46370°(CAL—Callana Station, SA), R46786,

R48303, R49061, R49087. BMNH 1946.8.27-2.

Pygopus steelescotti: AMS R13233, R17981, R26579,

R80336, R133267. NTM R99, R370, R821, R828,

R830, R2270, R2271, R3790, R4730, R5033, R5280,

R5303, R6423, R6677, R6704, R6777, R11247,

R12416, R20513°(CAP—Cape Crawford, NT),

R20545*-6"(CAP), R20588, R22333. OM J39061,

J52746. SAMA R3510, R8117. WAM R23792,

R56304, R70079, R70085, R70339, R75537-8,

R83195, R83575, R87307, R99201, R101361.

P. lepidopodus: QM J47145*(CTN—Shipton’s Flat,

Qld). SAMA R33291°(SYD—Terry Hills, NSW),

R20865’°(STO—Stony Point, SA), R23629°(RM2—Mt

Remarkable National Park, SA), R25341°(HAM—

Hambidge Conservation Park, SA), R25689°(YAL—

Yalata, SA), R30270°(ESP—Esperance, WA). WAM

R77939°(WAL—Walpole-Nornalup National Park,

WA), R90119°(KAR—Karridale, WA).

OBITUARY PATRICIA MARIETJE THOMAS BSC, MSC AO 13 APRIL 1915

-— 16 DECEMBER 1999

ARCHIE MCARTHUR

Summary

Patricia Marietje Thomas, Pat to her many friends, died peacefully in Adelaide on 16 December,

1999 bringing to an end an outstanding career in parasitology. The elder of two daughters, Pat was

born in Melbourne a year after her famous father, Sir Douglas Mawson, returned from the Antarctic

but while he was in America. Pat was raised and educated in Adelaide, receiving her Batchelor of

Science degree from the University of Adelaide in 1936, and her Masters degree two years later.

She found employment in the Zoology Department as a part-time teacher in practical classes and

researcher under Professor T. Harvey Johnston. After marrying marine biologist Ifor Thomas in

1947, and bearing three sons, Pat returned to academic life in 1950, spending most of it, apart from

a short period overseas, in the Zoology Department until her retirement in 1980.

OBITUARY

PATRICIA MARIETJE THOMAS BSc, MSc, AO

13 April 1915 - 16 December 1999

Pat Thomas at Coolgubbin Camp, Connie Sue Hwy, Great Victoria Desert,

June 1974.

Patricia Marietje Thomas, Pat to her many

friends, died peacefully in Adelaide on 16

December, 1999, bringing to an end an

outstanding career in parasitology. The elder of

two daughters, Pat was born in Melbourne a year

after her famous father, Sir Douglas Mawson,

returned from the Antarctic but while he was in

America. Pat was raised and educated in Adelaide,

receiving her Batchelor of Science degree from

the University of Adelaide in 1936, and her

Masters degree two years later. She found

employment in the Zoology Department as a part-

time teacher in practical classes and researcher

under Professor T. Harvey Johnston. After

marrying marine biologist Ifor Thomas in 1947,

and bearing three sons, Pat returned to academic

life in 1950, spending most of it, apart from a

short period overseas, in the Zoology Department

until her retirement in 1980.

From 1981 until failing health slowed her

down in 1995, Pat transferred her activities to

the South Australian Museum, where, as an

Honorary Curator, she greatly enlarged the

helminth collection to a position of pre-eminence

in Australia, and successfully pressed for its

recognition as the Australian Helminthological

Collection. With this achievement, Pat then

worked to find a successor to herself, to continue

the work, and in 1994 Dr Sylvie Pichelin was

appointed as the first Curator of Helminths. Ian

Beveridge, a younger colleague of Pat’s, has

listed her many professional achievements in

Obituaries, Royal Society of South Australia

(2000).

54 A. McARTHUR

Pat was a worker, a helper, quick-witted and

sharp as a tack, could show kindness and

sensitivity when needed, and abhorred

irresponsibility. In the following, a little about her

personal life has been compiled after talking to

some of her friends.

Philippa Horton (Collection Manager, Birds,

South Australian Museum). ‘I first met Pat

Thomas at Adelaide University in 1974 during

Zoology 1 practical classes for which Pat was a

demonstrator. As a shy student I was in awe of

her wealth of experience and knowledge, and

was somewhat terrified of her forthright and no-

nonsense manner. During one particular class—

the dissection of a rat—a fellow student and I

were having trouble in locating an anatomical

feature. On being called for help, Pat blustered

over, tackled our rat with forceps and scalpel,

and severed one of its arteries. The body cavity

filled with blood, obliterating anything we might

have seen, and rendering the three of us helpless

with laughter. From then on any feelings of terror

vanished and were replaced with warm affection.

One other occasion which stands out in my

memory showed the warmth which was not

always obvious in Pat’s personality. I was

pregnant with my first child and, at morning tea at

the Museum, expressed the hope that it wasn’t

going to be twins. Pat described her own

experience of giving birth to a son only THEN to

find out there was another about to follow. She

would have been thrilled with just one but was

absolutely delighted to have two beautiful babies.

She was filled with maternal love as she spoke

and her words were a great comfort to a new

mother-to-be. Nurturing twins must call upon

extra reserves of physical stamina and strength of

character: I was sure Pat had both in abundance.’

Mrs Anne Hardy, Rostrevor. ‘I began to work

as a technologist in the Zoology Department,

University of Adelaide, for 25 shillings per week

in 1940. Then the Mawsons were living

temporarily in Ruthven Mansions in Pulteney

Street. My job was caring for the fish, newts,

snails etc. used in the study of parasitic flatworms

and often at weekends Pat did it to save me

coming in to feed them.

“When the Department moved to the newly

constructed Benham Building which overlooked

the river on one side and the Jubilee Oval on the

other, Pat had the task of packing the huge

collection of ‘preserves’ (specimens from the two

1929-31 British, Australian, New Zealand

Antarctic Expeditions) a challenge which she

excelled in.

‘Once when the supply of dead rabbits for

dissection turned up at the wrong time, it was

necessary to skin them. Pat’s technique was

magnificent; a body held in one hand appeared to

almost leap out of its fur. During the war, Pat

took up marine biology and often accompanied

groups to Port Willunga to study the reef. Her

organising abilities were meticulous. As the war

ended, undergraduate numbers ballooned out and

the Professor deputed Pat and me, then a post-

graduate student, to give repeat lectures. Pat had

an engaging way with words.

‘Occasionally Professor T. Harvey Johnston

took us collecting near Tailem Bend and on one

occasion, students were allowed to swim off a log

in the river. One of the students got herself heavily

entwined by the long strands of floating weed and

submerged. When her panicky face surfaced, it

was Pat who wrenched her out of danger. Typical

of Pat, on having saved her life, she gave her a

sharp rebuke for having done the wrong thing.

One of my vivid memories was at the Spring

School at the Marine Research establishment at

Cronulla, NSW. Students at the school were

housed a mile or more from the laboratories and

used a bus to go to lectures unless there was a

better offer, which in Pat’s case, there soon was.

As I waited for the bus, a motor bike with the

school’s tutor skilfully tackled the sandy track,

and riding pillion was a passenger with bright red

hair and skirts blowing and a face of blissful

exhilaration as she clutched her future spouse Ifor

Thomas.

‘After her marriage, Pat had developed

domestic enthusiasm. When I returned to teaching

in the 60s and 70s in Zoology demonstration

classes, she had won a shirt ironing contest at the

Royal Adelaide Show, and with typical efficiency,

produced three sons in fifteen months, sewed,

organised and was soon back with her nematodes.

I recall that names suggested for the twins were

Son and MawSon. Later, Pat followed her mother

as President of the Mothers and Babies Health

Association.

‘Pat and I took part in many of the

ornithological excursions, led by Joan Paton, to

the Simpson Desert, the Great Victoria Desert,

Kakadu and many more. On one such, Pat

complained about Rex Ellis’s dog which took a

dislike to her and protected the essential

communal spade from Pat’s urgent grasp.’

Mrs Medindie.

Anne Dow, ‘On an

OBITUARY — PARTICIA MARIETJE THOMAS 55

ornithological excursion to Angorichina with Pat

and Joan Paton, Pat overheard someone

complaining about the food. Anne remembers

Pat’s quick response “ I never complain about

camp food, after all it goes in one end and out the

other”.

‘Around the camp fire Pat had a delightful

singing voice.’

Mrs Elizabeth Simpson, Stonyfell. “The two

Mawson girls and we three Cleland girls were

great friends and spent much of our lives in the

same pursuits. Pat and I were contemporaries at

the University and many were the parties and

excursions which we attended. I knew that if Pat

was in attendance my enjoyment and performance

would be heightened and all would be well.

‘On many occasions in the bush after dinner I

remember Pat with a Tilley lamp dissecting and

searching for helminths. On one of Sir Douglas’

geology excursions crossing a field, Pat and I

noticed a cow giving birth to a calf. Although the

cow was feeding placidly, the calf’s head was

protruding. Blushing hotly, for matters of that

nature were a little taboo those days, Pat and I

approached Sir Douglas and drew his attention to

the phenomenon. Instead of leading the students

in a different direction, he drew everyone’s

attention, in his loudest voice, to the plight of the

cow. A minute later a calf was standing with

wobbly legs alongside mother.

‘Pat was a stimulating companion. She did not

suffer fools gladly, even at times, her best friends.

There were few of us who had not felt her critical

tongue but this in no way detracted from the deep

love and respect we held for her.’

Order of Australia Association, South

Australian Branch Newsletter No 97; In

Memoriam. Awarded AO, Queen’s Birthday

1994, for service to the science of zoology in both

research and teaching and the development of the

Australian Helminthological Collection.

Slim Sommerville (retired zoologist, University

of Adelaide) remembers how driving in the

country with Pat was so appalling because she

insisted on stopping to examine carcasses of dead

animals. He also remembers being invited to

dinner at the Mawson home in 1954. Sir Douglas

sat tall at the head of the table and when kangaroo

tail soup was served, he exclaimed ‘What’s this!

Pll have none of it’. This surprised Slim knowing

of the horrible food on which Sir Douglas had

survived in Antarctica.

Dene Cordes (Department for the

Environment), and Noel Lothian (formerly

Director, Adelaide Botanic Gardens) were

associated with Pat through the Board of the

National Parks Commissioners and the Wildlife

Advisory Council. There she was a particularly

astute member whose lifelong interest in

environmental matters enabled her to contribute

greatly. She was interested in remote areas such as

the Koonalda Caves which she urged should

become a National Park, and this has since been

achieved.

Sylvie Pichelin (Former Curator of Helminths,

South Australian Museum). ‘I first met Pat in

1995—the same day as the interview for the

position of Curator of Helminths at the South

Australian Museum. Both events made quite an

impression on me. Although I had corresponded

with Pat about the Collection, I did not know her

personally. I knew of her impressive publication

record but at the same time had heard tales of

sherry in beakers next to kangaroo guts. I quickly

learnt that Pat liked to call a spade a spade—a

trait that was appreciated and helped me grasp the

size and importance of the Helminth Collection.

Pat was also very kind to me on my first visit and

took me out to a delightful lunch. There are few

like Pat—her death marks the closure of an era of

parasitology in Australia.’

Cath Kemper (Curator of Mammals, South

Australian Museum). ‘Pat was always very

interested in expanding the helminth collection so

when I started to study (and dissect) cetaceans in

the late 1980s she was keen for us to collect

parasites and hand them on to her. Several times

she came out to Bolivar herself and assisted with

dissections. It was great to work with her because

she always “hoed in” and nothing was too much

trouble. I remember one day when she arrived at

Bolivar before us and, not being able to get in and

not wanting to do nothing, she set to work

weeding the patio in front of the kitchen! It looked

lovely and tidy when we arrived. I always had a

great admiration for her because it would not have

been easy to have been a female scientist earlier

in her life. Science was something that men did

and the few women involved were often

considered odd. Pat was able to combine family

and work so was just like everyone else, only

much more determined than most!’

Archie McArthur (Honorary Research

Associate, South Australian Museum). ‘When I

56 A. McARTHUR

started on the ant collection at the Museum it

was Pat who “showed me the ropes”; she always

found time to help me solve problems. She was

often one of the first to start work in the

mornings on the Invertebrate Floor of the Natural

Science Building of the South Australian

Museum, where as an Honorary Research

Associate she built up the biggest collection of

helminths from Australian animals. In addition

to dissecting organs and extracting specimens,

she spent months at a computer, cataloguing this

vast collection. Occasionally, Joan Paton called

in around noon and the three of us had lunch

together in Pat’s lab where she kept a plentiful

supply of sweet vermouth. This she served in

little beakers and even though we knew that the

same beakers had contained a kangaroo’s liver or

a salmon’s gut only minutes before, we

appreciated her generosity and the purifying

effects of alcohol.

“When her father Sir Douglas Mawson was

teaching, Pat was often required as chaperone or

cook on his many field trips. About one such

expedition, Reg Sprigg reminisced in Geology is

Fun (1989) “I relished an experience at the

deserted Old Myrtle Springs Station ruins. We

arrived out of Copley just before sunset. We

carried little in the way of lamps so the race was

on to beat the fading light. Patricia in

commendably practical fashion grabbed a loaf of

bread, placed it on the boards of a weather beaten

and long abandoned old couch and got to work.

Said Mawson “Whatever are you doing there

Pat?” “Cutting the bread. Can’t you see?” “But

Pat, you can never tell what dirty old swagman

has been sleeping on that. We could all get

syphilis.” Pat, never to be panicked, turned her

head and nonchalantly answered, “You don’t get

syphilis through the mouth, Daddy.” Father for

once was stuck for words.’

REFERENCES

BEVERIDGE, I. 2000. Obituary. Patricia Marietje

Thomas, BSc, MSc, AO. 13.iv.1915 — 16.xii.1999.

Transactions of the Royal Society of South Australia

124: 56-60.

SPRIGG, R. 1989. ‘Geology is Fun. Recollections.’

Gillingham: Adelaide. x + 349 pp.

Archie McArthur, Honorary Research Associate, Terrestrial Invertebrate Section, South Australian Museum, North

Terrace, Adelaide, South Australia 5000. Records of the South Australian Museum 34(1): 53-56.

CORRECTED IDENTIFICATIONS AND LECTOTYPE DESIGNATION FOR

POROCHILUS ARGENTEUS (ZIETZ) (OSTEICHTHYES :

SILURIFORMES: PLOTOSIDAE)

PETER J. UNMACK

Summary

Australian plotosid catfishes are poorly understood (Merrick & Scmida 1984; Allen 1989) and,

among other problems, the taxonomy of Porochilus argenteus (Zietz) is confused. The present note

is intended to clarify the identification of the syntypes of P. argenteus and to designate a lectotype.

Museum abbreviations follow Leviton et al. (1985).

CORRECTED IDENTIFICATIONS AND LECTOTYPE DESIGNATION FOR POROCHILUS

ARGENTEUS (ZIETZ) (OSTEICHTHYES: SILURIFORMES: PLOTOSIDAE)

Australian plotosid catfishes are poorly

understood (Merrick & Schmida 1984; Allen

1989) and, among other problems, the taxonomy

of Porochilus argenteus (Zietz) is confused. The

present note is intended to clarify the

identification of the syntypes of P. argenteus and

to designate a lectotype. Museum abbreviations

follow Leviton et al. (1985).

Zietz’s (1896b: 410, Plate 16, Fig. 7)

description of Plotosus (= Porochilus) argenteus

was based primarily on a single, 138 mm syntype

from Cooper Creek near Innamincka, South

Australia. Glover (1976) incorrectly listed this

specimen (SAMA F1090) as the holotype. Zietz

included other specimens but scarcely mentioned

them. The specimen emphasized is clearly P.

argenteus as recognized today by Allen (1989).

Upon re-examination, it fits the description well,

having a concave dorsal profile between snout and

dorsal fin and a smooth inner face of the pectoral

spine. Dorsal-fin ray counts were difficult to

obtain as the fin is in poor condition; pectoral-fin

ray counts were also difficult, although an

approximate value of I-6 was obtained (cf. I-7 in

Zietz [1896b]). It is the only plotosid listed from

the Horn Expedition as ‘type’ in the original

SAMA register. A second syntype, probably

SAMA F1082 (Table 1), was discussed by Zietz

(1896b: 411) as follows: ‘A larger specimen of

about eight inches total length, from Finke River,

Hermannsburg, is of darkish colour; but this is

most likely caused by having been dried

formerly.’ Additional syntypes are not specified,

but localities were listed as Barcoo River, Finke

River at Idracowra, and Palm and Ilpilla creeks,

all in the Northern Territory. Extant syntypes are

shown in Table 1.

Several lines of evidence identify all specimens

except SAMA F1090 as Neosilurus hyrtlii

Steindachner. Specimens of Porochilus argenteus

are generally ligher coloured, whereas N. hyrtlii

are usually dark. Zietz (1896a, b) did not record

N. hyrtlii in other collections of the Horn

Expedition, yet they are clearly present in the

Finke River (Unmack in press). Specimens

examined from the Horn Expedition (BMNH,

NMV, SAMA) all are N. hyrtlii based on the

following characters: convex or straight dorsal

profile; rough inner side of pectoral spine; higher

pectoral-fin ray counts. (Dorsal-fin ray and

pectoral-fin ray counts could not be obtained from

some specimens due to the poor condition of

those fins.) (Table 1). All additional plotosid

specimens examined (AMS, AMNH, BMNH,

NTM, SAMA, NMV, WAM) from Finke River

not associated with the Horn Expedition are N.

hyrtlii. Finally, all SAMA P. argenteus syntypes

(Table 1) were not registered until 1928 and,

except for F1090, had been reidentified before

cataloging as N. hyrtlii. Records of P. argenteus

from the Finke River system (Zietz 1908; Glover

& Sim 1978; Glover 1982; Allen 1989; Glover

1990; Larson & Martin 1990; Davis 1996; Kerle

& Fleming 1996) are thus based on Zietz’s record

or on misidentifications of N. hyrtlii. The single

specimen on which Zietz’s (1896b) description

was largely based (SAMA F1090) is hereby

designated as the lectotype since it is the only

specimen in the type series identified as that

species; all remaining paralectotypes are

reidentified as N. hyrtlii. Porochilus argenteus

does not occur naturally in the Finke River system,

where the only known plotosid is N. hyrtlii

(Unmack in press).

ACKNOWLEDGMENTS

Thanks to all museums and collection staff, especially

T. Sim (SAMA) and to A. C. Gill for examining BMNH

specimens on my behalf. Thanks to W. L. Minckley and

W.N. Eschmeyer for comments on earlier drafts.

REFERENCES

ALLEN, G. R. 1989. ‘Freshwater Fishes of Australia’.

T. F. H. Publications: New Jersey.

DAVIS, J. A. 1996, Aquatic ecosystems in central

Australia: comparison of recent records of fishes and

invertebrates with those of the Horn Expedition. Pp.

282-286 in ‘Exploring Central Australia: Society, the

Environment and the 1894 Horn Expedition’. Eds S.

R. Morton & D. J. Mulvaney. Surrey Beatty & Sons:

Chipping Norton, New South Wales.

GILLBANK, L. & MAROSKE, S. 1996. Behind the

botany of the Horn Expedition: Ferdinand Mueller’s

documentation of the larpintine flora. Pp. 209-224 in

‘Exploring Central Australia: Society, the

Environment and the 1894 Horn Expedition’. Eds. S.

P. J. UNMACK

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LECTOTYPE FOR POROCHILUS ARGENTEUS 59

R. Morton & D. J. Mulvaney. Surrey Beatty & Sons:

Chipping Norton, New South Wales.

GLOVER, C. J. M. 1976. Vertebrate type-specimens in

the South Australian Museum. I. Fishes. Records of

the South Australian Museum 17: 169-175.

GLOVER, C. J. M. 1982. Adaptations of fishes in arid

Australia. Pp. 241-246 in ‘Evolution of the Flora

and Fauna of Arid Australia’. Eds W. R. Barker &

P. J. M. Greenslade. Peacock Publications:

Adelaide.

GLOVER, C. J. M. 1990. Fishes. Pp. 189-198 in

‘Natural History of the North-East Deserts’. Eds M.

J. Tyler, C. R. Twidale, M. Davies & C. B. Wells.

Royal Society of South Australia: Adelaide.

GLOVER, C. J. M. & SIM, T. C. 1978. Studies on

Central Australian fish: a progress report. South

Australian Naturalist 52: 35-44.

KERLE, J. A. & FLEMING, M. R. 1996. A history of

vertebrate fauna observations in central Australia:

their value for conservation. Pp. 341-366 in

‘Exploring Central Australia: Society, the

Environment and the 1894 Horn Expedition’. Eds S.

R. Morton & D. J. Mulvaney. Surrey Beatty & Sons:

Chipping Norton, New South Wales.

LARSON, H. K. & MARTIN, K. C. 1990. ‘Freshwater

Fishes of the Northern Territory’. Northern Territory

Museum of Arts and Sciences: Darwin.

LEVITON, A. E., GIBBS, R. H. Jr., HEAL, E. &

DAWSON, C. E. 1985. Standards in herpetology and

ichthyology: Part I. Standard symbolic codes for

institutional resouce collections in herpetology and

ichthyology. Copeia 1985: 802-832.

MERRICK, J. R. & SCHMIDA, G. E. 1984. ‘Australian

Freshwater Fishes: Biology and Management’.

Griffith Press Ltd.: Netley.

UNMACK, P. J. in press. Fish persistence and fluvial

geomorphology in central Australia. Journal of Arid

Environments.

ZIETZ, A. H. C. 1896a. Pisces. Pp. 176-180, pl. 16 in

‘Report on the work of the Horn Scientific

Expedition to Central Australia, Part 2, Zoology.’ Ed.

B. Spencer: London.

ZIETZ, A. H. C. 1896b. Description of an additional

new species of fish from the Finke and Barcoo

Rivers. Pp. 410-411 in ‘Report on the work of the

Horn Scientific Expedition to Central Australia, Part

2, Zoology.’ Ed. B. Spencer: London.

ZIETZ, A. H. C. 1908. A synopsis of the fishes of South

Australia. Part 1. Transactions and Proceedings of

the Royal Society of South Australia. 32: 288-293.

Peter J. UNMACK, Biology Department, Arizona State University, Tempe, AZ 85287-1501, USA,

peter.unmack@asu.edu. Records of the South Australian Museum 34(1): 57-59, 2001. Manuscript received 28

March 2000.

IRIECOJRIDS

SOUTH |

AUSTRALIAN

MUSEUM

VOLUME

MAY 2001

34 PART 1

ISSN 0376-2750

CONTENTS:

37

55

a7,

ARTICLES

M. BAEHR & P. HUDSON

New species and a new genus of the subfamily Pogoninae from South Australia

(Coleoptera: Carabidae).

PA CLARKE

The significance of whales to the Aboriginal people of southern South Australia.

B. H. JAMES, S. C. DONNELLAN & M. N. HUTCHINSON

Taxonomic revision of the Australian lizard Pygopus nigriceps (Squamata: Gekkonoidea).

NOTES

A. McARTHUR

Obituary — Patricia Marietje Thomas

P. J. UNMACK

Corrected identifications and lectotype designation for Porochilus argenteus (Zietz)

(Osteichthyes: Siluriformes: Plotosidae).

Published by the South Australian Museum,

North Terrace, Adelaide, South Australia 5000.

» RECORDS

> OF

SOUTH

AUSTRALIAN

MUSEUM

VOLUME 34 PAIRT 2

DECEMBEIR 2001

A NEW SPECIES OF AUSTRALIAN CANTHYDRUS SHARP WITH A KEY

TO THE AUSTRALIAN SPECIES OF NOTERIDAE (COLEOPTERA)

C. H. S. WATTS

Summary

A new species of Canthydrus, C. ephemeralis sp. nov. from Northern Australia is described and

figured. It is distinguished from the other known Australian Canthydrus, C. bovillae Blackburn,

1889, by size and dorsal colour pattern as well as a number of structural characters. A key to the

genera and species of the Australian Noteridae is included.

A NEW SPECIES OF AUSTRALIAN CANTHYDRUS SHARP WITH A KEY TO THE

AUSTRALIAN SPECIES OF NOTERIDAE (COLEOPTERA)

C. H. S. WATTS

WATTS, C. H. S. 2001. A new species of Australian Canthydrus Sharp with a key to the

Australian species of Noteridae (Coleoptera). Records of the South Australian Museum 34(2):

61-64.

A new species of Canthydrus, C. ephemeralis sp. nov., from Northern Australia is described

and figured. It is distinguished from the other known Australian Canthydrus, C. bovillae

Blackburn, 1889, by size and dorsal colour pattern as well as a number of structural characters.

A key to the genera and species of Australian Noteridae is included.

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

Manuscript received 22 November 2000.

Australia has a limited fauna of aquatic beetles

of the family Noteridae consisting of only five

described species in four genera; Canthydrus

Sharp, 1882, Hydrocanthus Say, 1823,

Neohydrocoptus Sato, 1972, and Notomicrus

Sharp, 1882. All are essentially tropical although

Notomicrus tenellus Clark, 1863 occurs in

decreasing numbers down the east coast almost to

the Victorian / South Australian border. Although

all but Neohydrocoptus are extremely streamlined,

they do not inhabit running water as their body

form might suggest, but are found in still water in

swamps, embayments and dams amongst

emergent vegetation. All species are common,

Neohydrocoptus subfasciatus Sharp, 1882, less so

than the others. Most of the species fly readily to

light.

In this paper I describe a distinctive new

Canthydrus which is seasonally common in the

escarpment country of coastal Northern Territory

and adjacent Western Australia. All specimens are

from the collection of the South Australian

Museum.

SYSTEMATICS

KEY TO THE GENERA AND SPECIES OF AUSTRALIAN

NOTERIDAE

(AFTER PEDERZANI 1995)

Fore tibia expanded beyond base of tarsi

and with a strong hooked spur at the outer

apical angle... cc. seeqeesesenssantesesescvenees 2

— Fore tibia not expanded beyond base of

tarsi, with several weak apical spines .. 5

Prosternal process not broader than long,

its apex about 2 to 2.5x as wide as its

breadth between postcoxae; < 3 mm long

CONANYAIUS coscsciscotseiecserssnetascsstizenstosteds 3

— Prosternal process broader than long, its

apex very broad, at least 2.5 to 3x as wide

as its breadth between postcoxae; > 4 mm

long Hyd rocanthu .....c1ccccscesssessceeesees “

Elytron with a pale spot towards apex, as

well as a medial one (Fig. 7), metatarsi

stout (Fig. 3), metatibial spines of equal

length (Fig. 3) .. C. ephemeralis sp. nov.

— Elytron without apical pale spot (Fig. 5),

metatarsi more elongate (Fig. 1), metatibial

spines unequal in length (Fig. 1) ...........

Lavea cage tsuaeanMecnersigaes C. bovillae Blackburn

Uniformly reddish; weak to moderately

impressed row of punctures adjacent to

the suture of the elytron .......... cesses

Ie) OS H. waterhousei Blackburn

— Uniformly black; lacking sutural row of

punctures ...... H. australasiae Wehncke

Lateral margins of metasternal plate not

bordered by a lateral ridge; < 2 mm long

gta}iescdafetecniante Notomicrus tenellus Clark

— Lateral margins of metasternal plate with

a lateral ridge; > 2 mm long...............0++

wets Neohydrocoptus subfasciatus Sharp

Canthydrus ephemeralis new species.

(Figs 3, 4, 6, 7, 9)

Types

Holotype: male: ‘NT 5 km SE Mt Borradaile

stn. 8/10/98 C. Watts’.

62 C.H. S. WATTS

8 9 10

FIGURES 1-10. Canthydrus bovillae; 1, metaleg; 2, tip of elytron (ventral view); 5, habitas drawing; 8, dorsal

view of central lobe of aedeagus; 10, lateral view of central lobe of aedeagus. Canthydrus ephemeralis; 3, metaleg;

4, tip of elytron (ventral view); 6, lateral view of central lobe of aedeagus; 7, habitas drawing; 9, dorsal view of

central lobe of aedeagus.

NEW SPECIES OF AUSTRALIAN CANTHYDRUS 63

Paratypes: 29, ‘NT 5 km SE Mt Borradaile stn.

27.5.99 C. Watts’; 14, ‘NT Mt Borradaile stn.

26.5.99 C. Watts’; 33, ‘NT 1 km W Gubara

Kakadu NP 17.3.98 C. H. S. Watts’; 25, ‘NT

5 km SE Mt Borradaile stn. 8/10/98 C. Watts’; 2,

“6 km SE Mt Borradaile NT 8/10/98 C. Watts’; 5,

‘NT 1 km W Gubara 17/3/98 C. H. S. Watts’; 3,

‘NT 1 km W Gubara Kakadu NP 29/1/99 C.

Watts’; 1, ‘Darwin NT 13.5.63 CW’; 23, ‘W.

AUST. Ck on Phillips Range 16° 53’S, 125° 48’E

4 Oct 1982 B. V. Timms’; 4, ‘W. AUST Dawn

Ck 15° 57’°S, 126° S51’E 5 Oct 1982 B. V.

Timms’. All specimens in the collection of the

South Australian Museum, except for 10 from

1 km west of Gubara which have been deposited

in the Australian National Insect Collection,

Canberra.

Description (number examined, 141)

Habitus. Length, 2.5-2.9 mm long. Elongate

oval, deep bodied, moderately accumulate behind;

shiny black, labrum and sides of pronotum

testaceous, elytron with two yellow/white lateral

spots, one in the middle, the other three-quarters

the way to the apex (Fig. 7), appendages dark

testaceous.

Dorsal surface. Smooth, virtually impunctate

except for serial punctures which are traceable but

weak, covered with fine reticulation, meshes

small, regular, much smaller than eye facet. Head

relatively broad, eyes large, antennae relatively

short, segments 8 to 10 somewhat thicker, apical

segment twice the length of penultimate.

Pronotum with thin well-impressed line a little

behind anterior margin, weaker towards sides,

lateral margin with somewhat more strongly

impressed line delineating a distinct beading.

Elytron with well-marked lateral flange/beading.

Ventral surface. Smooth, virtually impunctate,

covered with fine reticulation similar to dorsal

surface. Apical segment of labial palpus large,

oval, bifid at tip. Pronotal process wide, flat,

strongly setose, lateral margins weakly beaded,

approximately parallel-sided, narrower between

procoxae, hind angles extended. Elytron epipleura

very broad in anterior quarter, then rapidly

narrowing, very narrow in apical half, absent near

tip; apical ligula a smooth curve (Fig. 4). Raised

midsection of meso- and metasterna broad, flat,

widening towards rear, hind edge strongly

concave, outer hind angle with three to four strong

setae, covered with strong setae similar to those

on pronotal process. Protibia with strong spine,

approximately half length of tibia. Metaleg stout,

apical spines on metatibia equal in length (Fig. 3).

Hind margins of ventrites with row of setae,

stronger laterally.

Male. External characters as for female. Central

lobe of aedeagus with the apical ‘overturn’ which

covers a broad medial groove less than half total

length of lobe (Figs 6, 9).

Remarks

Canthydrus ephemeralis is most readily

separated from C. bovillae Blackburn, 1889 by its

smaller size (2.5-2.9 mm long against 2.9—

3.4mm) and dorsal colour pattern. Canthydrus

bovillae, in general, is less strongly coloured with

some specimens nearly completely black

(identification from descriptions and specimens

identified by Blackburn in the South Australian

Museum). Well-coloured specimens (Fig. 5) differ

from C. ephemeralis (Fig. 7) in having the front

of the head testaceous rather than dark; the

testaceous areas at the sides of the pronotum are

restricted to the front, whereas in C. ephemeralis

they extend along the whole side; two light-

coloured areas at the base of the elytra (often

indistinct) are lacking in C. ephemeralis; the

absence of a subapical spot on the elytron which

is present in C. ephemeralis (both species have a

light coloured lateral spot in about the middle of

the elytron). Other differences between the two

species are: the shape of the ligula near the apex

of the elytra—it has a pronounced indentation in

C. bovillae (Fig. 2) which is lacking in C.

ephemeralis (Fig. 4); the hind legs in C.

ephemeralis (Fig. 3) are more robust than those of

C. bovillae (Fig. 1); the metatibial spines in C.

ephemeralis are of equal length (Fig. 3) whereas

the inner one is longer in C. bovillae (Fig. 1); the

meso-metasternal central plate is a bit broader in

C. bovillae; the central lobe of the aedeagus has

the medial groove covered for most of its length

(Figs 8, 10) rather than about half as in C.

ephemeralis (Figs 6, 9).

Distribution

Known only from the localities listed above

under Types.

Habitat

All the collections of C. ephemeralis where the

habitat details are known have been from small,

temporary, wet-season, low gradient streams

flowing off escarpments in the coastal Northern

Territory and northern Western Australia. The

substrate of these streams is clean sand or rock

with accumulations of dead leaves and other

debris in places. The adults forage in the open

64 C.H.S. WATTS

sandy areas, at times in areas that may have water

only after rain and are dry a few hours later. In

contrast, C. bovillae lives in the more permanent

flood-plain billabongs and is not found in the

seasonal streams of the escarpment.

Etymology

In reference to its ephemeral appearance in

streams that only flow during the wet season.

REFERENCES

BLACKBURN, T. 1889. Notes on Australian

Coleoptera, with descriptions of new species.

Proceedings of the Linnean Society of New South

Wales IV: 445-482.

PEDERZANI, F. 1995. Keys to the identification of the

genera and subgenera of adult Dytiscidae (sensu lato)

of the world (Coleoptera; Dytiscidae). Atti

Dell’Accademia Roveretana Degli Agiati, a. 244

series VII, volume IV, B, pp. 5-83.

TIMBU WARA FIGURES FROM PANGIA, PAPUA NEW GUINEA

PAMELA J. STEWART & ANDREW STRATHERN

Summary

This paper discusses wickerwork figures known as Timbu Wara, formerly made in the Pangia area

of the Southern Highlands Province of Papua New Guinea. The paper sets the historical context of

the described figures. The figures were associated with a fertility cult periodically celebrated with

large sacrifices of pigs. Wickerwork figures of this same general kind are known also from the

neighbouring Kewa and Enga areas. After the cult was abandoned in the 1960s people began

making the figures for sale to tourists.

TIMBU WARA FIGURES FROM PANGIA, PAPUA NEW GUINEA

PAMELA J. STEWART & ANDREW STRATHERN

STEWART, P. J. & STRATHERN, A. 2001. Timbu Wara figures from Pangia, Papua New

Guinea. Records of the South Australian Museum 34(2): 65-77.

This paper discusses wickerwork figures known as Timbu Wara, formerly made in the

Pangia area of the Southern Highlands Province of Papua New Guinea. The paper sets the

historical context of the described figures. The figures were associated with a fertility cult

periodically celebrated with large sacrifices of pigs. Wickerwork figures of this same general

kind are known also from the neighbouring Kewa and Enga areas. After the cult was abandoned

in the 1960s people began making the figures for sale to tourists.

P. J. Stewart (Research Associate) & A. Strathern (Mellon Professor), Department of

Anthropology, University of Pittsburgh, PA, USA 15260 [Pamjan+@pitt.edu]. Manuscript

received 18 April 2001.

INTRODUCTION

The wickerwork ritual objects that we describe

here were manufactured in the Pangia area of the

Southern Highlands Province of Papua New

Guinea (Fig. 1). The name Pangia was originally

given to a single small locality south of Mount

Jalibu in the Wiru language area. The Wiru

language is related to a set of neighbouring

languages in the Highlands which includes those

of the Enga, Huli, Mendi and Kewa peoples

(Wurm 1964). Among these languages, Wiru

appears as a Family-level isolate, less closely

related to the others than they are related among

themselves (Wurm & Hattori 1982).

The first non-New Guinean outsiders to go

through the wider area were the gold mining

prospectors, Michael and Danny Leahy, during the

1930s. In the late 1950s government officers

selected Pangia for their station after the first

patrol outward from Ialibu. Catholic and Lutheran

missionaries arrived shortly thereafter and set up

stations at Yaraparoi and Tiripini in the close

vicinity of the Pangia government station. Other

missions followed—the Evangelical Bible

Mission (at Mele) and the Wesleyans (at Takuru).

Also, a linguist with the Summer Institute of

Linguistics, Harland Kerr, came to live in the area

at Poloko (Borona) where he studied the language

and social practices of the people (Kerr 1975).

Relatively little has been published about the

Pangia people. Jeffrey Clark, who worked in

Takuru village, has written a number of articles

about the impact of colonialism and Christianity

on the people and his book Steel to Stone, which

brings together and synthesizes the content of

these articles, has been posthumously published

(2000). (For further materials on the Wiru see

Strathern & Stewart 1999a, 1999b, 2000; Stewart

& Strathern 1999a, n.d.).

The Wiru are horticulturalists and pig-keepers

and their staple crop is the sweet potato. The area

has little primary forest and is mostly grassland

and regrowth, dotted by limestone outcropping

and sinkholes. This landscape has figured greatly

in local notions of sorcery and spirit beings that

were thought to live around settlements (Stewart

& Strathern 1999b). In the past, long houses were

built, through the initiative of male leaders, for

important pig-kill ceremonies (Strathern &

Stewart 1999a).

Pangia is encircled on three sides by mountain

ranges. During the 1960s and much of the 1970s,

travel into and out of the area was difficult. This

remained so until the building of the Ialibu—

Pangia road during the 1970s. Further, the Wiru

traditionally had only minor trading and exchange

links with non-Wiru peoples. Thus the impact of

the colonial government and Christian

missionaries was more extreme in a number of

ways than in other regions where there had been

more historical contact and interaction with other

Papua New Guineans (Clark 2000; Strathern

1984; Stewart & Strathern n.d.). In particular,

colonial control came to Pangia late, some 30

years after first contact in areas to the north such

as Jalibu and Hagen, and the Administration

attempted to bring about a swift set of social

transformations and bring to the area the same

‘developments’ as were established elsewhere.

P. J. STEWART & A. STRATHERN

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TIMBU WARA FIGURES, PAPUA NEW GUINEA 67

Prior to 1960, the Wiru had little experience in

interacting with the colonial power. Non-New

Guinean outsiders were said to have perhaps come

from rivers (uele nekenea) because of their white

color; hence their resemblance to spirit beings

known to the Wiru people already. They were also

referred to as ‘red men’ and ‘spirits from the sky’.

Non-New Guinean outsiders were seen as

desirable and powerful entities since they were

thought to have abundant wealth items. One way

in which the Wiru adjusted to their presence was

to make them feel, to some extent, as though they

were insiders, thereby ‘domesticating’ them into

the local transactional framework. They were

identified as the wealth people, the people with

possessions, as elsewhere in the Highlands also.

This engendered a desire in the Wiru to obtain

these possessions by whatever trading means were

accessible to them. But the colonial powers had

an agenda of ‘development’ that was not in

harmony with the ‘traditional’ means of sociality

familiar to the Wiru.

The colonial administration required people to

work as labourers on road projects and

government buildings and to seek permission to

hold pig-kills which might disrupt road work

schedules. There was a great push to bring Pangia

up to the ‘development’ level of other, longer-

contacted areas of the Highlands. Thus the

Administration reduced the time that the people

had to adjust to these outside pressures which had

come into their lives. Since the missions had come

into the area on the back of the colonial

government, the Wiru in some ways conflated the

two and their expectations of what would

eventually be obtained by participating in

‘development’ projects and mission activities was

very great indeed, escalating with every passing

year.

Initially, the Administration regarded various

‘traditional’ practices as a hindrance to

‘development’ in the area and some of the

missions labelled ‘traditional’ cult practices as

satanic. This was particularly true of the

Evangelical Bible Mission and the Wesleyans.

The Wiru had accepted Christianity into their

pantheon of cults as a powerful new practice that

might assist them in obtaining the good health

and wealth that they thought the non-New

Guinean outsiders possessed. Christian missions

did bring schools into the area, which many even

today remember as a positive change. In 1999, a

middle-aged Pangia man from Wiliame village

told us that he had appreciative memories of the

missionaries, saying that ‘they brought schools,

roads and teaching about hygiene’ which he felt

allowed him to have a better life and his children

to become educated and earn money.

But some missionary workers, notably

indigenous evangelists, also ordered the smashing

of cult stones and the burning of cult houses—a

practice also followed in other areas of the

Southern Highlands such as among the Duna

(Strathem & Stewart 1998; Stewart & Strathern

2000). Before the coming of the missions, the

Wiru held pig-kills in association with a number

of religious cults: Timbu, Tapa and Aroa Ipono

(Strathern & Stewart 1999b; Stewart & Strathern

1999a). These cults all aimed at bringing health

and fertility to the people and involved the

construction of specific ritual objects and

buildings. The Tapa cult centered on ancestral

stones belonging to subgroups in each settlement

area, and had to do with sacrifices of pigs made

on occasions of sickness. The Aroa Ipono

belonged to a widely spread complex of such cult

forms, also found in the Western Highlands

Province.

THE TiMBU CULT

The Timbu cult involved the construction of a

specific cult house, which contained a central post

(tungi) onto which were attached the shoulder and

jaw bones of many pigs that had been killed over

the years prior to the culmination of the festival.

The cult’s cycle of enactment was approximately

five to eight years, ending with a great feast in

which hundreds of pigs, the raising of which

involved innumerable hours of female labor,

would be killed. Prior to the pig-kill and

ceremonial exchange marking the end of the cult

cycle, the cult participants would dance around

the tungi post holding the timbu wara.

Subsequently, the tungi post would be taken out

from the cult house to the ceremonial ground

where the pig-kill and ceremonial exchanges took

place. This would be in the area of a main village

(tumbea ta) where cult houses of various kinds

were located.

The Timbu cult was aimed at restoring

ecological balance and fertility to humans, pigs

and the land. The cult looked outward and brought

together clans from other neighbouring districts,

reinforcing continued exchange relations. It also

may have functioned to redress exchange

imbalances and outstanding debts that arose from

warfare deaths (Dosedla 1984). The term timbu

means ‘sky’, so the cult may have had a

68 P. J. STEWART & A. STRATHERN

connection also with mythical Sky Beings who

are seen aS important in other Highlands

cosmologies. A comparable cult complex known

as Timp is found in the Mendi area (Lederman

1986).

The cult involved various male age and status

grades. Prior to the cult’s culminating pig-kill, the

participants would receive lengths of sugar cane

that demarcated the quantity of pork they would

subsequently receive. Although only males were

allowed into the cult house, females participated

in the production of the foodstuffs for the festival

and the pigs for the feast. They also had the highly

significant role of carrying back to their respective

settlements the pork received at the ceremonial

distribution.

Central to the whole process by which a Timbu

performance was organised was the sacrifice of

large numbers of pigs over a period of time, so as

to provide bones to festoon the central pole.

Informants agreed that marsupial bones would

also be attached to the pole; some declared that

the bones of rats and eels were also used. There

were probably local variations in this regard. The

overall purpose was clearly to show how many

animals had been sacrificed to the spirit. When

the pole was covered with bones, it was said that

‘it is ripe’ (nondokako), that is ready for the

climactic phase of the cult. Men and boys were

the ones who entered the tall house in which the

‘bones pole’ was kept. Older men were in an inner

circle, with younger men at their back. A ritual

specialist made spells for health and fertility,

which were said to come up from his stomach, his

tepe. For each cult house there was a single leader

who held the knowledge of these spells and was

therefore said to be the man in charge of the cult

site (yopikango). Such a position of leadership

and knowledge might be passed from father to

son. But the knowledge could also be passed on

between specific lines of descent within the larger

residential complex of the settlement.

Joint participation in the cult was a mark of

unity; separate performances later would mark

social processes of differentiation over time. Two

performances might be held in the same place,

then a shift would occur. Clearly the Timbu

represented the ongoing social life of a named

group and its diachronic extension and alteration.

The exclusion of women and young children

from the inner cult area does not mean that the

Timbu was an exclusive ‘men’s cult’. The whole

purpose of concealing the post and festooning it

with bones was so it could be brought out later

and displayed to the community at large, when the

final dance for the ‘coming out’ of the pole was

staged. One senior informant said that ‘after this

the young men could find wives for themselves’.

The parts of pigs that were taken inside the cult

house for consumption were the tail-joints

(regularly used in ritual contexts elsewhere in the

Highlands, such as among the Duna of the Lake

Kopiago area) and the sides. The inner parts,

including the entrails, and the heads of the pigs

were given to the women to eat. A woman would

dance up to a ‘fence’ and hold up a pig to be

killed. The men rushed at it calling out and

clubbing it as it squealed. The women then stood

at the fence and received their portions of the pig

to cook. When the ‘bones pole’ was finally buried

at the end of the performance, both men and

women were said to have participated in this final

ritual act.

The Timbu spirit was said to be male—ali, ‘a

man’. The bringing out of the pole was therefore

like the bringing out of a male initiate from a

condition of secrecy into one of display. Prior to

this moment of display in the ritual process,

women and children should not see the pole

because if they did so, their legs, arms and vaginas

would suffer from sores described as ‘scabies’.

That is, if they saw the Timbu when it was

‘unripe’, the result would be the opposite of the

effect of showing the spirit to them when it was

‘ripe’, like a mature, healthy man.

The Timbu also presided over marsupials. If

hunters failed to find marsupials in the forest they

would attribute this to the spirit. Hence the point

of hanging trophies on the pole as a sign of

hunting success—these were both a return to the

spirit for granting access to the game and a

demonstration to the community of this access.

The spirit was said to dislike pigs’ heads

because pigs spoil people’s gardens. If the Timbu

saw pigs’ heads it too would spoil gardens in the

same way, it was declared. We see here a sign of

the latent aggressiveness believed to be inherent

in the spirit.

Other taboos were observed. The participants

made small flutes and blew these to warn women

and children not to look when cult paraphernalia

were being carried into the house. The flutes were

blown also when pork was cooked and divided

out in the cult place, to prevent sickness in people

and pigs. These flute sounds were said to imitate

the calls of two marsupials known as tekelepo and

wapenge, which came ‘along with the Timbu cult

itself’, it was said. The marsupials’ call of tu-li,

tu-li inspired the flute-blowers to imitate them.

The spirit’s connection with the forest and its

TIMBU WARA FIGURES, PAPUA NEW GUINEA 69

game is reaffirmed in this detail, while the act of

blowing the flutes links the cult to a wide

spectrum of such practices in the Highlands region

as a whole.

Finally, the cult’s forest associations are shown

in the practice of roofing the cult house with wild

pandanus leaves and black palm thatch, into

which pig bones were pushed. White and rusty

brown forest leaves were used to decorate the

house; these also appear as decorations on houses

for spirit cults in the Hagen area north of Pangia.

We are dealing here with the fluid transmission of

cult elements from place to place, in which forest

associations are a constant feature.

After the performance was over, the pole and

all pig bones were carefully buried. At the burial

site the participants planted a kendo, a cordyline

shrub, as a long-lasting marker, surrounding the

shrub with boards. (This would make them like

the pokla mbo which marked moka events in the

Hagen area.) It seems likely that the kendo marked

the place where the spirit and its power was now

located in a dormant mode. The actual dance for

the culmination of the ritual activities also

involved wearing tall cordyline sprigs as bustles

at the back. Pairs of men danced facing each other

with their bustles bouncing at their backs, singing

songs to the Timbu. The dance was called polo

pendeko, ‘the bending dance’. Men cut a

pandanus leaf and blew musical notes on it as

they performed it.

The timbu wara wickerwork figures which are

the focus of this paper were made to be worn on

the heads of the male dancers who participated in

the climactic phase of the cult. We quote here

initially two statements in 1967 by senior

knowledgeable male informants of two different

settlement areas in the southern part of the Wiru

speaking area:

We make a figure of a man with rope and

decorate it. When we kill the pigs we carry it on

our heads. We put tall cordyline sprigs in our

rear bustles and we carry bows and arrows.

(Longai of Tunda village)

We made a wara, like a man, and placed it on

our heads. This was a good custom; it was our

own. If this figure was not made, pigs, men,

women, and children would all become sick. We

killed a pig [as a sacrifice before constructing

the figures] and we made a ‘picture’ [yomini in

Wiru] of the spirit. One special kind of rope and

one special liana were used to make the wara

and we carried them on our heads at the time of

cooking pigs. Women could not hold these wara,

only men and their sons ...[After the

performance was over] some wara were buried,

but some were kept. Out of fifteen, maybe ten

were buried but five were kept. For the next

performance we made new ones and finally

buried the remaining old ones. But now all the

wara have been thrown away. The Lutheran

Mission men cut up the wara with their axes.

(Kuluwa of Marapini village)

Another old ritual expert noted:

In the past we made gardens of taro, bananas,

sweet potato, and they all grew big. Now that we

have given up the Timbu our crops are small. In

the past pigs and men were large because of the

Timbu. Now they are small. (Wipai of Tunda

village)

In spite of this tone of regret, shared by many

others at the time, the Timbu cult had vanished

from the Wiru area by 1967. In several instances

it was reported that the yopikango of the cult had

himself turned in its sacred stones to the missions

or had declared that it would not be performed

again. Since he was the authorized expert for the

group, his decision could not be challenged. Such

men were afraid of the messages of Hell and sin

that the missions brought with them. When asked

if the people could themselves have cut up the

wara and destroyed them, informants

unequivocally said ‘No, we could not have done

this, because we were sorry (ela toka) for the

wara.’

In Tunda village in 1967, an effort was made to

collect a number of these wickerwork figures used

in the Timbu. Tunda village had a road into it,

making access somewhat easier although the road

was often barely passable to vehicles. The people

were converted to the Lutheran and Catholic

forms of Christianity, and the settlement itself was

artificially structured through the actions of the

government bringing previously separate small

settlements together (Strathern 1984; Stewart &

Strathern n.d.). Tunda was, however, a traditional

tumbea ta or ‘big place’, functioning as a ritual

and political centre for surrounding hamlets. Some

villagers at this time still possessed the knowledge

of how to make the wickerwork figures called

timbu wara, even though the Timbu cult was no

longer practised.

The Lutheran Mission in the area was very

active, as was the Catholic mission. Tunda was

divided between these two. Hageners from the

Western Highlands were working with the

Lutherans at this time, and these Hageners

boasted that they were involved in removing the

ritual cults of the local people. But the actual

time period since the arrival of Christianity, and

the changes associated with it, was short; thus,

70 P. J. STEWART & A. STRATHERN

the people still retained a great deal of

knowledge about previous religious activities

such as the Timbu cult, although they were

sometimes reluctant to discuss these in any

detail.

THE TimBu WaRA AND CHRISTIAN MISSIONS

Timbu wara (‘the spears of the Timbu spirit’)

were very striking wickerwork figures known

from only a few areas of the Southern Highlands.

As indicated above, these figures were made

specifically for a particular cult performance and

then buried or allowed naturally to decay. They

would not have been kept indefinitely, so there

were no heirloom examples that one could

purchase for a museum collection. Even if some

had been retained in the past, the missionaries had

ordered the people to destroy or burn any

remaining items from their pre-Christian religious

practices. Therefore, a request had to be made to

the people to manufacture some of these figures

for a collection to be sent to the Museum of

Archaeology and Anthropology at the University

of Cambridge.

Several local youths agreed to make them for a

set fee. They were sons of men who had the right

to wear such items in the past and there was no

bar to learning from their fathers how to make

them. In the past, these items were not made by

the ritual experts, whose power lay in their

knowledge of spells for the cult.

When the completed figures were presented

they were placed vertically on top of the heads

of the craftsmen to show how they had been

worn in their ritual context. They were shaped

like a human figure but they represented the

Timbu spirit. At the time of presenting the

figures, their makers carried them in this

displayed manner through the village, which was

close by a government road. At the moment that

this parade was occurring, a local fundamentalist

missionary from the Evangelical Bible Mission

at Mele was walking along the road from

Mamuane village, where he wanted to set up a

mission station. The missionary’s baggage

carriers pointed at the men with the timbu wara

figures and began to yell out, ‘Satan, Satan,

Satan figures are coming!’ They were told that

these were not Satan figures but rather artifacts

that were commissioned to be made for a

collection and not for any religious purpose, but

these men were very afraid and went off. The

missionary himself was rather upset and also

went off without any further discussion of the

objects.

Ironically, this missionary later did

subsequently set up a church in Mamuane village

with the stated aim of bringing into congruence

Christian ideology and aspects of indigenous

culture. But in the course of doing this it was

reported that he made a number of transactions in

which the people felt they had not obtained

enough in return for services rendered and they

took revenge by burning the mission station down,

forcing the missionary to leave.

The timbu wara figures were, as we have noted,

objects made to celebrate the action of dancing

for the Timbu spirit during the cult performance.

The central ritual objects of the cult were items

(which could include, for example, a tree fungus

which was hard and stone-like and was called

timbu kapa) that were kept in the specially

constructed cult house, and pigs were sacrificed to

these items. One of the senior men, who had

himself been a custodian of the cult, explained

further some of the ritual actions that were

performed:

Inside the house they dug a hole and they made

man [magical spells] over it. They used a pearl

shell to scoop out this hole, speaking a spell

over it also. Inside the cult house they planted

tree saplings. They killed pigs and filled blood

from them into a bamboo tube. They also took

tree oil and poured it along with the blood into

the hole. They took a special leaf, timbu

yombolu, and buried it in the hole with the oil

and blood. For every action they made a special

man, including the house-building. When we had

killed the pigs we made a special net bag,

tetaleme ka, like a woman’s net bag, made from

rope, which we carried outside. The axes we

used to cut this rope in the bush and to cut the

wood for the house were all bespelled in

advance. We used four different sorts of wood

for the actual pole on which we hung the pigs’

bones and also the bones of eels and marsupials.

These woods were tungi, lepa, walea, and

pokota, (Kuluwa of Marapini village)

This quotation gives some idea of the

complexity and diversity of the ritual acts

performed inside the cult house. The action of

giving tree oil to the ground also parallels the

Hagen male spirit cult sequence in which

Campnosperma oil is poured into the ‘spirit’s eye’

(see Stewart & Strathern 2001: 99-112). The wara

in a sense marked the successful performance of

these other acts: the creation of the spirit as ‘a

man’ (ali), representing the regeneration or

revivification of the community itself. The fact

TIMBU WARA FIGURES, PAPUA NEW GUINEA 71

that wara figures, like the ‘bones pole’, were all

buried does not mean that they were casually

discarded. Burying items in the ground is a way of

preserving and containing their force, as is

abundantly testified from all over the Highlands.

Since the pole was buried at a site where a kendo

or cordyline was planted as a marker of group

identity, we may reasonably suppose that the wara

FIGURE 2. These two boys are displaying decorations they have been involved in making to demonstrate the

methods of manufacture of timbu wara. The boy on the left wears one figure attached to his head and holds another

in his hand. The boy on the right wears an alipo wig with a set of bird plumes mounted on bamboo pieces above it

as a headdress. The alipo wig style belongs especially to the Ialibu area north of Pangia. The house in the

background is a newly made house for cooking food, beside a large dwelling house.

72 P. J. STEWART & A. STRATHERN

were buried in the same place. Elsewhere in the

Highlands, ritual items were left to decay or were

buried, for example the gerua boards or painted

plaques found from the Wahgi area eastwards to

the Siane (O’Hanlon 1989: 102ff.).

At this historical moment, in the 1960s, the

FIGURE 3. A close-up of the timbu wara figures.

fundamentalist Christians from one of the

missions were saying that these timbu wara

objects were ‘satanic’ and should be destroyed.

But the Lutherans and Catholics of the village

were not making this a particular issue. The

Catholic priests of the Capuchin order in the area

TIMBU WARA FIGURES, PAPUA NEW GUINEA 73

had even used these objects to decorate their own THE TIMBU FIGURES

houses and encouraged people to make them as

tourist objects so as to obtain some income. By Figures 2—4 show the designs of the two figures

suggesting the objects be used as tourist items that were constructed for sale in 1967 in Tunda

they were also deflecting the people away from village. The two boys were those assigned to carry

associating them with the ritual itself. the objects and who had been involved in their

: : es : . i ~

FIGURE 4. Rear view of the timbu wara figure attached to the bast lining of an alipo wig, along with feather

decorations mounted on rattan.

74 P. J. STEWART & A. STRATHERN

making. Both figures were flat constructions with

protrusions representing arms and legs; they were

coloured with earth pigments. One also had a

rudimentary head on which a headdress was

mounted. This figure had a central strip of white

running from its head to its genital area. It is likely

that this strip marked the presence of vital

‘grease’, of fluid in the body. The second figure

had a softwood plaque marked alternately in dark

and light colours to resemble the pearl shell

omament hung on the body. It may be thought to

have signified the idea of the body as wealth, a

significant notion in Pangia. It may also have

represented a notched horbill beak, a favourite

design also painted on men’s noses at pig-kills

(kaila timini is the name of this design); each of

the timbu wara in the South Australian Museum

(A.66227, -8) has a real hornbill beak attached as

the head (Figure 5).

COMPARATIVE MATERIALS

The timbu wara figures are one of a category of

objects that are found also in other areas of New

Guinea. Heinz Dosedla (1984) has written a

review article on this category of woven figures

that are made from flexible materials. He

discusses items from the Southern Highlands

Province, including the Enga, Kewa and Wiru

peoples. His classification, based on a survey of

the literature available to him, includes a set of

figures that are in the shapes of animals, 100-

180 cm in length and decorated with earth

pigments; and a second set of anthropomorphic

figures, 60-90 cm in height, unpainted, and with

an enclosed cavity. For both types, a spiral

weaving technique was used in their manufacture,

in which plant materials were bent and wound

around a supporting frame that was made from

stiffer materials, such as bark rope in some

instances.

Dosedla states that among the East Kewa of the

Kagua District, who had trading relations with

both the Enga and Wiru peoples, the woven

figures were significant in the cult known as

Rimbu, which ensured fertility through sacrifices

to the dead. The cult included the use of flat

woven figures whose construction had been

supervised by special ritual experts. These figures

resembled those that the Wiru used, but these

Kewa figures were in the shape of animals and

they were worn by dancers on their lower chests.

The figures were said to represent spirits of dead

FIGURE 5. Two Timbu wara figures in the South Australian Museum (A.66227, -8), each with hornbill beak

attached. Purchased from Stephen Kellner of Sydney in 1977. Kellner purchased from a private collector c.1972.

Photo courtesy South Australian Museum.

TIMBU WARA FIGURES, PAPUA NEW GUINEA 75

members of the clan. The East Kewa Rimbu cult

also employed the second category of woven

figures—the hollow anthropomorphic ones—

which were used as freestanding figures in

conjunction with cult stones.

The Rimbu cult festival developed over a

number of years with several stages of activity

and involved two separate groups of men, the

‘red’ and the ‘black’ sections. These two colours

are symbolically associated elsewhere in the

Highlands with dualistic schemes of classification.

The ‘red’ group took control of the

anthropomorphic figures and the cult stones,

while the ‘black’ group took control of the

animal-shaped figures. The latter were displayed

and carried at the end of the ritual by cult

participants.

Dosedla also describes wickerwork figures from

the Enga area. One type called yupin was kept in a

special spirit house (yainanda, war house) along

with cult stones. The yupin figure, seen as ‘male’

and marked by its prominent genitals, was thought

to have intercourse with the rounded cult stones,

bringing fertility to the land, pigs and humans.

(See Wiessner & Tumu 1998: 199, 204, 207 for

further details on Enga yupin or yupini figures.)

Neich (1976) provides a valuable survey of

yupin figures derived from information provided

by an Apostolic Church missionary, H. M. Reah,

who lived in the Yandapu Enga area for some ten

years in the 1960s and 1970s. Reah’s account

indicates that these figures were considered very

powerful; that they were appealed to in times of

drought; that their keepers would ntually place

them on top of round flat female stones with holes

in the centre; and that groups tended to have their

own yupin figures. But several groups might come

together on an inter-local basis to ask the yupin

for health and prosperity in times of sickness.

Neich says that yupin figures had to be made

from special vines (as was true for the timbu wara

also); that their manufacture was commissioned

and the maker rewarded handsomely with shells

and axes; and that women and children were not

allowed to see them. If a woman saw one, her

next child would be born deformed. If a child saw

one, he or she would fail to grow.

Here, the logic of time reversal was operating.

Seen ‘out of time’, the yupin would harm, not

help, the community. Most valuably, Neich

provides information from a number of Enga

areas, including those that are closer to the Kewa,

on variant designs and names of yupin. One was

from Margarima and was said to represent

Tatagali-Wabe or Tali-eli, a major spirit appealed

to when gardens did not grow well. It is

interesting to note that this is the name Glasse

gave to a supreme spirit among the Huli people

(Glasse 1965). The name yupin itself may mean

‘root or base man’, that is ‘the source of things’.

All yupin figures are cylindrical, with a hollow

area inside of them, in contrast with the flat timbu

wara. While the Enga yupin practices and ideas

are clearly cognate with the Wiru wara customs,

the Wiru obviously developed their own ways of

thinking about these objects.

Dosedla also briefly describes the Wiru timbu

wara figures, which he says were painted with

concentric rings of white and yellow with a

background of red ochre. Such concentric rings

are a significant motif for the Wiru. They appear

on decorations women made around their navels

for dances and in the bands of earth colours

painted around the Female Spirit (Aroa Ipono)

cult stones (Strathern & Stewart 1999b). He notes

that they appeared only as the flat type, and not as

animal but anthropomorphic forms. He suggests

that the Wiru merged two types (the one seen in

the East Kewa Rimbu cult and the other from the

Enga area) into one type; that is, the Wiru figures

were flat, not three-dimensional, but represented

humans, not animals.

RITUAL AND CHANGE

As Dosedla points out, the Wiru timbu wara

figures belong to a wider geographical area in

which similar figures were made for cult contexts.

Dosedla traces the provenance of these figures

and analyses their distribution in cultural—

geographical terms. For our purposes here, it is

important to ask how the Wiru figures were used

in the wider contexts of cult and political activity;

and also why they so rapidly ceased to be made if

they held ritual importance, and what

consequences ensued,

The name of these figures is significant. Wara,

‘spear’, refers to the context of warfare and inter-

group conflict. The cult performers looked out on

the spectators in order to demonstrate their

political power and solidarity. Wara can also be

used to mean ‘wealth’, the ability to ‘kill’ objects

by paying for them with wealth items, or to obtain

wives by the payment of bride price. As the

figures represented the human form, they could

ambiguously take onto themselves an aura both of

the dead ancestors and of the living men who

carried them in the festival. Since, among the

Wiru, the Tapa cult dealt with internal problems

716 P. J. STEWART & A. STRATHERN

of sickness centered on male ancestors and the

Female Spirit cult was concerned with general

fertility and alliance, the space occupied by the

Timbu cult clearly had to do with the outward-

looking political strength of settlements.

The decline of the Timbu cult thus predictably

accompanied the decline in warfare between

parishes and hastened the demise of the pre-

colonial patterns of inter-parish competition and

display, although aspects of these continued in the

pig-kills held periodically (Strathern & Stewart

1999a). In addition, as we have seen, some

fundamentalist Christian missions in the 1960s

were able successfully enough to label cult

practices as satanic, and the timbu wara figures

were prominent and easily visible targets for

disapproval, whereas cult stones could more easily

be hidden and used still for healing rituals or

small sacrifices. The timbu wara figures thus

disappeared, only to reappear briefly as ‘tourist’

objects, once emptied of their meaning and

dislocated from their political and ritual contexts.

Their decline is one of the markers of the success

of colonial hegemony, with a trajectory from

prime ritual object to tourist object to forgotten

piece of knowledge.

Tracing the decline of these sacred figures

through time, we have first to note that the major

cult leaders were often those who surrendered

their ritual power to the Christian missions in

order to get the new ritual power that was on

offer. In other words, they themselves in some

sense began the process of desacralisation.

Mission helpers then took this further by cutting

the effigies into pieces, something their custodians

themselves would not do because of ela. The

same mission authorities, along with colonial

government officers, completed the process by

suggesting the items be made for sale, relying on

people’s desire for money.

The making of the figures for the Cambridge

museum in 1967 pre-dated this process. These

figures closely paralleled traditional designs.

Those made later, for sale to tourists, showed

variations. They were bigger, designed for display

on hotel walls; more uniform in color and design;

were turned out repeatedly; or were made in the

shape of crosses or abstract designs. After

Independence in 1975 and the departure of many

expatriate missionaries and government officers,

the people did not persist with this craft for long.

Indeed, the figures were soon in over-supply: the

patrol officer’s space in the Pangia Station

overflowed with them at one time, jumbled up

and tangled together in disarray because of

insufficient demand for them. As with other

matters, the people became disappointed with

their returns and ceased production.

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WURM, S. A & HATTORI, S. 1982. ‘Language Atlas

of the Pacific Area’. Australian Academy of the

Humanities and the Japanese Academy.

SEPIA HEDLEYI BERRY, 1918 (CEPHALODA: SEPIIDAE): A COMPLETE

DESCRIPTION AND CLARIFICATION OF THE STATUS OF S.

DANNECIGI BERRY, 1918 AND S. REX (IREDALE, 1926)

A. L. REID

Summary

Following the examination of both qualitative and quantitative morphological characters, Sepia

dannevigi Berry, 1918 from off Kangaroo Island, South Australia and Sepia rex (Iredale, 1926)

from eastern Australia (Manly Beach, NSW) are synonymised with Sepia hedleyi Berry, 1918, also

from Kangaroo Island. These three species have never been satisfactorily delimited in the literature,

so this synonomy is long overdue. A full description of Sepia hedleyi is provided.

SEPIA HEDLEYI BERRY, 1918 (CEPHALOPODA: SEPIIDAE): A COMPLETE

DESCRIPTION AND CLARIFICATION OF THE STATUS OF

S. DANNEVIGI BERRY, 1918 AND S. REX (IREDALE, 1926)

A. L. REID

REID, A. L. 2001. Sepia hedleyi Berry, 1918 (Cephalopoda: Sepiidae): a complete description

and clarification of the status of S. dannevigi Berry, 1918 and S. rex (Iredale, 1926). Records

of the South Australian Museum 34(2): 79-97.

Following the examination of both qualitative and quantitative morphological characters,

Sepia dannevigi Berry, 1918 from off Kangaroo Island, South Australia and Sepia rex (Iredale,

1926) from eastern Australia (Manly Beach, NSW) are synonomised with Sepia hedleyi Berry,

1918, also from off Kangaroo Island. These three species have never been satisfactorily

delimited in the literature, so this synonomy is long overdue. A full redescription of Sepia

hedleyi is provided.

Amanda L. Reid, 6 Sturt Place,

Bulli,

NSW, 2516, Australia (email:

mandy.reid@ optusnet.com.au). Manuscript received 24 April 2001.

The cuttlefish Sepia hedleyi was described by

Berry (1918) on the basis of a number of specimens

collected from the Great Australian Bight, off the

coast of South Australia. While the original

description is detailed and relatively well

illustrated, the cuttlebones of all the specimens

were missing or damaged, and its structure was

estimated in a reconstruction of the ventral side of

the bone from fragments obtained from a single

female specimen. In the same publication, another

species from the same collection locality, S.

dannevigi, was described on the basis of three

female specimens, again with damaged cuttlebones.

Perhaps (in part) because the cuttlebones of

these species were so incompletely known,

another species, S. rex, was described by Iredale

in 1926 on the basis of a cuttlebone collected on

Manly Beach, Sydney (33°48'S 151°17'E). Iredale

(1926) provided no justification for the erection

of this new species: no comparison was made

with other cuttlefish species. In addition, he

placed S. rex in a new genus, Decorisepia, on the

basis of its ‘remarkable’ rounded spine and the

absence of an inner cone and ventral sulcus.

Strangely, the two latter traits are clearly visible in

his illustration accompanying the description

(Iredale 1926: pl. xxii, figs 9-10) and are obvious

on the type specimen. Following the

comprehensive revision of the Sepiidae by Adam

and Rees (1966), the validity of Decorisepia (and

many other sepiid genera) has been questioned,

and few workers follow this scheme, preferring to

retain most sepiids within a single genus Sepia

until more is known about the phylogenetic

relationships among species. The generic

classification of Adam and Rees (1966) is the one

followed in this paper.

In a later work, Adam (1979) placed two other

species, Decorisepia cottesloensis Cotton, 1929

and D. jaenschi Cotton, 1931, both known only

from cuttlebones, in synonomy with S. rex.

However, the description of S. rex in Adam

(1979) is clearly that of yet another species, S.

opipara (Iredale, 1926), so Adam’s synonomy is

not valid. After examining the type cuttlebones of

all these species, Lu (1998) confirmed that D.

cottesloensis and D. jaenschi are indeed synonyms

of S. rex, so that part of the story, at least, has

been resolved.

The three names, S. dannevigi, S. hedleyi and S.

rex, have persisted in the literature until now,

though they have never been satisfactorily

differentiated from each other. Lu (1998) gave a

diagnosis for each species based on his

examination of the type specimens and

comparison with other material housed in the

Museum Victoria collections. The only difference

he noted between S. hedleyi and S. rex is that the

cuttlebone striae are nearly straight in S. hedleyi

(based on Berry’s figure of the reconstructed

cuttlebone), but rounded in S. rex. He also noted

the similarity between S. dannevigi and S. hedleyi

and suggested they may be conspecific.

The aim of the present study is to clarify the

status and identity of the three putative species, S.

dannevigi, S. hedleyi and S. rex.

80 A. L. REID

MATERIALS AND METHODS

This work was based on museum material. The

non-type material examined is listed in Appendix

1. Material examined in detail for both qualitative

and quantitative characters is listed in the Material

examined section of the species description below.

Australian institutional abbreviations used are:

AM, Australian Museum, Sydney; MV, Museum

Victoria, Melbourne; SAM, South Australian

Museum, Adelaide.

To determine whether S. dannevigi, S. hedleyi

and S. rex are conspecific, the type specimens and

their original descriptions were compared. In

addition, specimens from close to the type locality

of S. rex (Manly Beach, NSW, 33°48'S 151°17'E)

were compared with specimens from southern

Australia. Because the S. rex type specimen is a

beachwashed cuttlebone, it may have drifted from

any distance before reaching Manly Beach, so the

selection of specimens for comparison is not

entirely satisfactory, but unavoidable.

Identification of these specimens could only be

based on the appearance of the cuttlebones and

their comparison with the type specimen.

Unfortunately, apart from the type specimens,

no other material was available for study from the

type locality of S. dannevigi and S. hedleyi, both

from the ‘Investigator St. Area, south of Kangaroo

Island, South Australia’ (Berry 1918: 263, 266).

(Herein lies another difficulty: the Investigator

Strait is actually north of Kangaroo Island, and no

latitudes and longitudes are given in Berry’s

paper, so the exact type locality is unknown.)

Some additional specimens were examined from

sites that were as close as possible to Kangaroo

Island, though some distance away (see Fig. 1).

Apart from the appearance of the cuttlebone (see

Results to follow), these specimens conformed in

all other respects to Berry’s (1918) descriptions

and type specimens of both S. hedleyi and S.

dannevigi (the two of which appear to be

indistinguishable).

In addition to comparing qualitative characters,

the statistics package ‘Systat’ (Systat

Incorporated) was used to compare differences in

morphometric characters between the populations

from eastern Australia (ten males and ten females)

and those from southern Australia (seven males

and nine females). Sexes were treated separately.

FIGURE 1. Distribution of Sepia hedleyi Berry, 1918 based on specimens examined in this study (solid triangles).

The open square indicates the approximate type locality of S. dannevigi and S. hedleyi (see Materials and Methods),

and the open triangle the type locality of S. rex. Arrows indicate the collection sites for specimens that were

examined in detail for both morphological and morphometric characters.

REVISION OF SEPIA HEDLEYI 81

In addition to preserved specimens, the

measurements given in Berry’s (1918) paper for

S. hedleyi were included in the analyses. The

measurements of one female specimen (E4377)

were excluded for reasons given in the Results

section below. Slopes and intercepts of regression

equations were compared statistically between the

eastern and southern Australian specimens for

those characters showing a significant correlation

with body size. For soft parts, mantle length was

used as a size indicator. For cuttlebone

measurements, cuttlebone length was used.

Measurements and indices used throughout this

paper are primarily those given in Roper and Voss

(1983), using dorsal mantle length (ML) as a size

standard. Some additional measurements are used,

and these with the definitions listed by Roper and

Voss (1983) are given in Appendix 2. In the

species description to follow, parts of the club

and arm sucker rims are described using the

terminology of Nixon and Dilly (1977) while

nomenclature for the radula follows Nixon (1995).

The beak was described following Clarke (1986).

Diagrammatic illustrations of measurements and

terminology used for key structures are shown in

Reid (2000).

Measurements were made either using dial

callipers, or an eyepiece micrometer inserted in a

stereo microscope. All measurements are

expressed in millimetres (mm). Measurements and

counts for individual specimens and ranges of arm

length indices, arm sucker diameter indices and

arm sucker counts are presented in tables; ranges

for all other characters appear in the text. In the

species description and tables, the range of values

for each character is expressed as: minimum—

mean—maximum (standard deviation, SD). Values

for each sex are given separately. Measurements

were not taken from the Sepia dannevigi (AM

C148249) or Sepia hedleyi (AM C148252)

holotype specimens because of the poor state of

preservation of these animals.

Measurements for structures that were clearly

distorted or broken were not attempted; these, in

addition to missing and unknown values, appear

as a dash (—) in the appendices. Ranges for

specific character traits given with the species

description do not, therefore, always refer to the

total number of specimens examined.

For examination of arm and club sucker rims,

suckers were removed from the middle of

designated arms and the tentacular club, mounted

in glycerine jelly and viewed using a compound

microscope. Radulae and beaks were dissected

from the buccal mass and soaked for

approximately 30 minutes in a warm, saturated

potassium hydroxide solution; then radulae were

cleaned using forceps and a fine brush. Radulae

were mounted in glycerol and the new, unused

portion was examined. All characters refer to both

sexes unless stated otherwise.

The species description was generated using

DELTA (DEscription Language for TAxonomy)

software (Dallwitz, 1980; Dallwitz et al., 1993;

Partridge et al., 1993).

RESULTS

With a single exception, only minor differences

were found between the type specimens and

descriptions of S. dannevigi Berry, 1918, S.

hedleyi Berry, 1918 and S. rex (Iredale, 1926).

The exception is in the shape of the cuttlebone

striae of S. hedleyi (Berry’s figure pl. 1xii, fig. 2,

reproduced here as Fig. 2A), which are nearly

straight compared with those of S. rex (Fig. 2C,

D) which are an inverted U-shape (Fig. 2D).

Unfortunately, the cuttlebone upon which Berry’s

illustration is based could not be found in the

Australian Museum collection (Ian Loch, personal

communication), so the accuracy of the

reconstruction of the cuttlebone could not be

checked. Assuming the shape of the striae as

illustrated is true to the original specimen (a

possibility that seems likely given that this section

of the cuttlebone fragment is largely intact), a

striking resemblance between this cuttlebone and

that of another southern Australian species, S.

cultrata Hoyle, 1885, is apparent (compare Fig.

2A and 2B). Females of S. hedleyi and S. cultrata

are not easy to distinguish in most respects,

however, the club suckers of S. cultrata are in 5-6

transverse rows and larger than those of S.

hedleyi, which are in 9-12 rows. The tentacles

were missing from the specimen Berry (1918)

used for the illustration of the cuttlebone, so it is

possible that this specimen may have been

misidentified, and is actually S. cultrata. Both

species inhabit similar depths and are often

trawled together, so I believe this possibility is a

real one. Given the doubt about the identity of the

specimen (E4377, Berry’s register [507]), its

measurements were excluded from consideration

in this study.

Morphometrics

Nine characters for males (MW, VML, FuL,

HW, AS3, ASI4, CIL, ClSv and CbL) and 19

characters for females (MW, AMH, VML, HW,

82

A. L. REID

TABLE 1. Morphological parameters showing differences in eastern and southern populations for females.

Regression data Y = a + bX, where Y = dependent variable, a = intercept, b = slope, X = ML, Sig. = significant

difference between the regression lines of eastern (E) and southern (S) Australian populations (Pop.) with respect to

intercept, N = number of specimens, r?= proportion of total variation accounted for by regression.

Y Pop. N r? a b Sig.

VML E 10 0.922 0.46 0.88 P<0.001

S 8 0.882 18.65 0.61

AS3 E 10 0.508 0.23 0.02 P <0.05

S 5 0.833 -0.41 0.02

CIL E 10 0.748 —9.83 0.27 P<0.001

S -11.91 0.35

4 0.980

ED, AL1, AL2, AL3, AL4, AS2, AS3, AS4, CIL,

ClSd, GiL, CbL, CbW, CbB and LoL) were

significantly correlated with mantle length in both

populations. Therefore, the slopes and intercepts

of the regression equations could be compared

statistically between the eastern and southern

populations to determine whether there are any

differences in these quantitative characters.

For males, no significant differences in either

the slopes or the intercepts of the regression

equations were found, suggesting that (for the

characters that could be compared) the southern

and eastern populations do not differ. Three

characters differed between the eastern and

southern populations in females: VML, AS3 and

CIL (Table 1). To determine whether these

differences are significant, or perhaps the result of

geographic variation across the range of a single

species, residual variables calculated for each of

these characters were regressed against latitude

and longitude of each specimen collection site

following the methods detailed in Reid (1991).

Residual values for VML and CIL were correlated

with both latitude and longitude, indicating a

strong geographic component in the difference

between the eastern and southern populations

(Table 2). The character AS3 was not correlated

with either latitude or longitude and represents the

only significant difference between the two

populations in quantitative characters. A

difference in this trait alone, the diameter of the

suckers on the third arm of females, does not

warrant further detailed investigation and is of not

sufficient magnitude to warrant distinct species

recognition.

One further difference was noted in the number

of club suckers, with CIRC for the southern

Australian population 9-10 (Appendix 3) and 11-

12 for the eastern Australian population

(Appendices 4 and 5), though the S. hedleyi

holotype clearly has 12 suckers in transverse rows

as noted by Berry (1918). While this does not

appear to be a significant difference between the

two populations, and given the difficulty in

determining the exact number of club suckers,

particularly in species where the suckers are small,

it may still be worth investigating this character

when more material from southern Australia

becomes available.

From all available evidence, I conclude from

the examination of both qualitative and

quantitative characters that all specimens currently

referred to as S. dannevigi, S. hedleyi and S. rex

belong to a single species. Unless specimens are

found that match all parts of Berry’s (1918)

description of S. hedleyi, including the shape of

TABLE 2. Latitudinal and longitudinal regression of residual variables pooled from female specimens from eastern

and southern Australia tested by Y = a + bX where Y = the predicted dependent variable, a = y intercept, b = slope,

X is either latitude or longitude, N = number of specimens, r*= proportion of total variation accounted for by

regression, N/S = not significant.

X = Latitude X = Longitude

Variable N r a b Sig. Yr? a b Sig.

VML 28 0.511 54.40 -1.59 P<0.05 0.360 42.09 0.29 P<0.05

AS3 25 0.217 -1.84 0.06 N/S 0.032 0.75 -0.01 N/S

CIL 28 0.670 -39.79 1.71 P<0.01 0.354 23.61 —0.16

P < 0.05

REVISION OF SEPIA HEDLEYI 83

the cuttlebone striae, I believe his illustration of

the cuttlebone of S. hedleyi is in fact the

cuttlebone of a specimen of a different species,

probably S. cultrata. Sepia hedleyi has date

precedence (ICZN, 1999: Art. 23.1) with respect

to S. rex, and page precedence (ICZN, 1999: Art.

69A.10) with respect to S. dannevigi, so the latter

two species are placed in synonomy with S.

hedleyi in the description to follow.

Counts and indices for individual specimens

from southern Australia are given in Appendix 3,

and from eastern Australia in Appendices 4 and 5.

Ranges for arm length indices, arm sucker

diameter indices and arm sucker counts for

southern Australian specimens are shown in

Appendix 6. Measurements included in the

description refer only to southern Australian

specimens.

TAXONOMY

Sepia hedleyi Berry

(Figs 1(in part)-7, Appendices 3, 6)

Sepia hedleyi Berry, 1918: 258-264, pls 71-72. —

Lu, 1998: 169, fig. 10.

Sepia dannevigi Berry, 1918: 264-268, pls 73-

74, figs 1-2. — Lu, 1998: 168, fig. 8.

Decorisepia rex Iredale, 1926: 193. — Lu, 1998:

180-181, fig. 22.

Material examined

Holotype. Sepia dannevigi, 1F (61.0 mm ML),

Investigator St. Area, south of Kangaroo Is, South

Australia, Jan-Feb 1912, FIS ‘Endeavour’ E2466

(AM C148249),.

Holotype. Sepia hedleyi, 1M (56.9mm ML),

Investigator St. Area, south of Kangaroo Island,

South Australia Jan-Feb, 1912, FIS ‘Endeavour’

E2464 (AM C148252).

Holotype. Sepia rex, 1 cuttlebone (107.0 mm

CbL), Manly Beach, NSW [33°48'S 151°17'E]

(AM C127593).

Other material examined. Australia: New

South Wales: 1M (84.3 mm ML), 2F (83.1,

103.8 mm ML), 32°27'S 152°54'E, 244-242 m, 30

Jan 1982, coll. C. C. Lu & R. Tait (MV F77179);

1F (108.8 mm ML), 33°40'S 151°50'E, 210-204

m, 25 Jan 1982, coll. C. C. Lu & R. Tait (MV

F77138); 5M (75.9-100.5 mm ML), 9F (72.6—

107.7 mm ML), 33°42'S 151°51'E, 300-293 m, 25

Jan 1982, coll. C. C. Lu & R. Tait (MV F77136).

South Australia: 3F (93.1-107.6 mm ML), 25.5

C D

FIGURE 2. A, Sepia hedleyi Berry, 1918, reconstructed

cuttlebone, pl. Ixxii, fig. 2 Berry, 1918, female [507],

91.0 mm ML (E4377) (specimen possibly S. cultrata

Hoyle, 1885); B, Sepia cultrata Hoyle, 1885,

cuttlebone, ventral view, female, 67 mm ML (MV

F66203); C, Sepia rex, cuttlebone, dorsal view,

holotype, 107.0 mm CbL (AM C127593); D, Sepia rex,

cuttlebone, ventral view, specimen as in C. Scale bars;

A, B = 10 mm; C, D = 20 mm.

84 A. L. REID

ORL. |,

we Ola Clare

BE-Cene

FIGURE 3. Sepia hedleyi Berry, 1918. A, funnel organ, male, 47.2 mm ML (SAM D19241); B, funnel-locking

(right) and mantle-locking (left) cartilage, male, 47.2 mm ML (SAM D19241); C, sucker rim arm 1, portion of

toothed half, male, 55.0 mm ML (SAM D19243); D, sucker rim, portion of non-toothed half, specimen as in C; E,

portion of hectocotylised arm, male, 75.7 mm ML (SAM D19241) (D, dorsal; V, ventral). Scale bars; A = 5 mm; B

= 2mm; C, D=0.05 mm; E=2.0 mm.

REVISION OF SEPIA HEDLEYI 85

miles [41 km] SW of Cape Buffon, 37°58'S

139°45'E, 300 fm [549 mJ, May 1981, coll. J.

Sealey (SAM D19240). Western Australia: 2M

(47.2, 75.7 mm ML), Great Australian Bight, 115

nm SW Eucla, 33°18'S 127°40'E, 180 m, 16 Jan

1989, coll. W. Zeidler & K. Gowlett-Holmes

(SAM D19241); 2F (67.9, 68.6mm ML), Great

Australian Bight, 110 nm SW of Eucla, 33°19'S

127°50'E, 250 m, 16 Jan 1989, coll. W. Zeidler &

K. Gowlett-Holmes (SAM D19242); 1M

(55.0mm ML), Great Australian Bight, 115 nm

SW of Eucla, 33°19'S 127°S50'E, 180 m, 16 Jan

1989, coll. W. Zeidler & K. Gowlett-Holmes

(SAM D19243).

Diagnosis

Male and female arms subequal in length; arm

suckers tetraserial throughout. Hectocotylus

present, left ventral arm of males modified: 6-8

rows of normal suckers proximally, 9-10 rows of

reduced suckers, remaining suckers normal to arm

tip; suckers in two dorsal series smaller than those

in two ventral series, dorsal and ventral series

widely spaced, those in ventral two series aligned

in a single row. Club with 9-12 suckers in

transverse rows, all similar sized, small; dorsal

and ventral protective membranes not fused at

base of club; swimming keel extends well beyond

carpus along stalk. Cuttlebone acute anteriorly and

posteriorly, with median rib dorsally; spine

without keels; anterior striae inverted U-shape;

inner cone limbs uniform width, narrow, V-

shaped posteriorly.

Description

Counts and indices for individual specimens are

given in Appendix 3; ranges for arm length

indices, arm sucker diameter indices and arm

sucker counts are shown in Appendix 6.

Small to moderate-sized species; ML males

47.2-10.7-83.0 (SD, 13.9); females 67.9-87.8-

107.6 (SD, 13.9). Mantle broad, oval; MWI males

41.9-49.8-55.6 (SD, 5.4); females 44.2-51.6—

59.8 (SD, 5.9); dorsal anterior margin triangular,

acute; extending anteriorly beyond eyes; AMHI

males 11.9-14.2-15.5 (SD, 2.0); females 12.8-

14.6-17.8 (SD, 2.0). Ventral mantle margin

emarginate, without distinct lateral angles; VMLI

males 80.7-86.0—-93.7 (SD, 4.4); females 75.3-

83.1-88.4 (SD, 4.5); posterior gland and gland

pore absent. Fins widest in posterior third; FWI

males 7.2—9.7-11.7 (SD, 1.3); females 4.4-8.6—

11.6 (SD, 2.4); anterior origin posterior to mantle

margin; Flla males 4.5-5.0—5.3 (SD, 0.4); females

2.0-3.7-5.7 (SD, 1.5); rounded posteriorly; with

narrow gap between; FIIp males 6.4—7.3-8.0 (SD,

0.8); females 4.7—7.9-12.4 (SD, 3.3). Funnel

short, robust, broad; extends to level of anterior

rim of eye; FuLI males 27.7—33.2-40.5 (SD, 5.5);

females 25.6—30.0-34.6 (SD, 3.3). Funnel free

portion approximately half funnel length; FFul

males 14.5-17.3-20.1 (SD, 2.8); females 12.6-

15.3-19.3 (SD, 2.8). Funnel organ dorsal elements

inverted V-shape with small anterior papilla;

ventral elements oval with acute anterior tips (Fig.

3A). Mantle-locking cartilage curved, with

semicircular ridge; funnel-locking cartilage with

depression that corresponds to ridge (Fig. 3B).

Head short; HLI males 17.6—23.2—29.7 (SD, 4.8);

females 13.4—-25.8-40.8 (SD, 9.4); broad,

narrower than mantle; HWI males 33.8-37.8-44.4

(SD, 3.7); females 31.6-41.3-46.6 (SD, 5.1). Eyes

moderate size; EDI males 12.7—14.2-16.5 (SD,

2.1); females 8.9-10.8-12.7 (SD, 1.6); ventral

eyelids present.

Male and female arms subequal in length

(Appendix 6). Arm length index (ALD of longest

arms in males (ALI4) 33.8-40.2-54.4 (SD, 7.2);

ALI of longest arms in females (ALI4) 34.7-42.0-—

51.3 (SD, 4.8). Protective membranes in both

sexes narrow. Non-hectocotylised arms normal,

not thickened. Distal arm tips in both sexes not

markedly attenuate. Arm sucker arrangement same

in both sexes: arm suckers tetraserial. Male non-

hectocotylised arm suckers normal in size (not

greatly enlarged); smaller than female arm suckers

in size (Appendix 6). Chitinous rims of arm

suckers with elongate rectangular teeth on distal

half of inner ring (Fig. 3C), teeth absent on

proximal half of ring (Fig. 3D); infundibulum

with 7-8 rows of hexagonal processes, inner 4-5

(variable) rows with elongate rounded pegs, pegs

becoming smaller towards periphery of sucker;

peripheral sucker rim processes radially arranged,

elongate, without pegs.

Arm sucker counts range from 106 to 248;

females with higher average counts than males

(Appendix 6).

Hectocotylus present, left ventral arm modified;

sucker size normal proximally, reduced medially,

then normal to arm tip; from proximal to distal

end of arm, 6-8 rows of normal suckers; 9-10

rows of reduced suckers (Fig. 3E). Suckers in two

dorsal series smaller than those in two ventral

series; reduced suckers much smaller than normal

arm suckers: ASIn4 1.32—1.46—1.69 (SD, 0.20) v.

ASInI4m 0.26—0.36—0.42 (SD, 0.09). Oral surface

of modified region wide, swollen, fleshy, with

transversely grooved ridges. Suckers in two dorsal

and two ventral series displaced laterally, suckers

86 A. L. REID

FIGURE 4. Sepia hedleyi Berry, 1918. A, club, male, 75.7 mm ML (SAM D19241); B, club sucker rim, portion of

toothed half, male, 75.7 mm ML (SAM D19241); C, club sucker rim portion of non-toothed half, specimen as in B;

D, upper beak, lateral view, female, 68.6 mm ML (SAM D19242) (stippling indicates extent of dark brown highly

sclerotised portion of beak); E, lower beak anteriolateral view, specimen as in D; F, lower beak, ventral view,

specimen as in D; G, radula, male, 75.7 mm ML (SAM D19241). Scale bars; A = 1.0 mm; B, C = 0.05 mm; D-F =

2.0 mm; G = 0.2 mm.

REVISION OF SEPIA HEDLEYI 87

in two ventral series aligned in a single row (Fig.

3E). Hectocotylised arm not markedly attenuate

distally.

Tentacular club longer in females than males;

CILI males 9.9-13.9--17.2 (SD, 2.8); females

16.2-—20.0—24.2 (SD, 3.4). Club crescent-shaped;

moderate length; sucker-bearing face flattened.

Club with 9-12 suckers in transverse rows; 32-42

suckers in longitudinal series. Suckers all similar

size; small (Fig. 4A). Distal tip of club with pair

of slightly larger suckers partially covered by a

thick, fleshy flap. CISI males 0.40—0.45—0.53 (SD,

0.07); females 0.40-0.46—0.52 (SD, 0.05); dorsal

and ventral marginal longitudinal series of suckers

similar in size; CISId males 0.40—0.42-0.42 (SD,

0.01); females 0.40-0.46-0.52 (SD, 0.05); CISIv

males 0.29-0.37-0.42 (SD, 0.07); females 0.32-

0.41—0.52 (SD, 0.10). Sucker dentition: half inner

ring circumference in both sexes with elongate-

rectangular teeth (Fig. 4B), remaining half with

blunt projections (Fig. 4C); infundibulum with 5—

7 rows of hexagonal processes, innermost with

elongate rounded pegs, pegs smaller towards

periphery of sucker; at periphery, processes

smaller, elongate-rectangular, without pegs

(similar to arm suckers). Swimming keel of club

extends well beyond carpus (Fig. 4A). Dorsal and

ventral protective membranes not fused at base of

club; joined to stalk; dorsal and ventral

membranes same length; extend beyond carpus

along stalk; approximately equal width; dorsal

membrane forms shallow cleft at junction with

stalk.

Gills with 29-30 lamellae per demibranch;

GiLC males 29-29-29 (SD, 0); females 29-29-—

30 (SD, 0.7). Gill length: GiLI males 24.9-30.0—

35.0 (SD, 5.0); females 29.5—33.9-38.2 (SD, 3.6).

Buccal membrane without suckers. Upper beak

(Fig. 4D) rostrum pointed, short, length

approximately equal to width, cutting edge

GO

FIGURE 5. Sepia hedleyi Berry, 1918. Male reproductive tract (testis not shown), 55.0 mm ML (SAM D19243)

(AAG, appendix of accessory gland; AG, accessory gland; CC, ciliated canal; DDC, distal deferent canal; GO,

genital orifice; MG, mucilaginous gland; SG, spermatophoric gland; SS, spermatophoric sac (containing

spermatophores); VD, vas deferens). Scale bar = 3 mm.

88 A. L. REID

slightly curved; hood high above crest

posteriorly; crest curved, lateral wall shallowly

indented posteriorly; wings and hood narrow and

short; jaw angle approximately 90 degrees; hood

and crest dark brown. Lower beak (Fig. 4E, F)

rostrum protrudes only slightly, cutting edge

straight; hood low on crest; crest straight, no

indentation on lateral wall edge; lateral wall edge

angled posteriorly, not perpendicular to crest;

hood and wings, width broad; hood notch

shallow, broad; wings widely spaced; crest

narrow; rostrum pigmented dark brown, wings

dark brown on inner margin only, rest of wing

light brown, crest dark brown. Radula (Fig. 4G)

homodont; rhachidian teeth with truncate bases,

slender, triangular, sides straight; first lateral

teeth similar length and width to rhachidian

teeth, asymmetrical with mesocone slightly

displaced toward centre of radula; second laterals

slightly longer than first, not distinctly curved on

lateral margin, with broad heels; marginal teeth

much longer than second lateral teeth, elongate

with long tapered and curved mesocone.

Digestive tract: (not illustrated) paired salivary

glands approximately one-third length of buccal

mass; paired digestive glands large, located close

together, with narrow, elongate triangular lobes

posteriorly, ducts connect digestive glands near

midline with caecum, ducts with branched

attached pancreatic tissue; oesophagus runs

dorsally along median junction of digestive

glands, joins sac-like stomach immediately

posterior to digestive glands; caecum disc-like,

grooved in a blunt V-shape anteriorly, surface

lining finely pleated; intestine undifferentiated;

ink sac very large, elongate; anal flaps well

developed.

Male reproductive tract (Fig. 5): testis on left

posterior side of viscero-pericardial coelom; at

distal end, convoluted vas deferens opens into

broad, cone-shaped mucilaginous gland, then

narrower, curved, spermatophoric gland. Close to

junction with lobe-shaped accessory gland and

gland appendix, delicate ciliated canal joins

spermatophoric gland; distal deferent canal

connects appendix of accessory gland to

spermatophore storage sac; genital orifice opens

dorsal to left gill in anterior end of mantle cavity.

Spermatophores (Fig. 6): cement body unipartite

(not divided into distinct regions); flask-shaped,

rounded posteriorly, connects to sperm reservoir

via narrow duct; tapers abruptly following

junction with middle tunic, which commences

towards basal half of cement body; ejaculatory

apparatus coiled, extends into oral dilation of

spermatophore. Spermatophores 5.8—7.9 mm long;

0.2-0.4 mm wide; SpLI 10.4—LL5-12.3 (SD, 0.9);

SpWI 3.4-4.4-5.1 (SD, 0.8). Buccal membrane in

females extends ventrally with spermatheca.

Female reproductive tract: (not illustrated)

ovary hangs from dorsal wall of posterior viscero-

pericardial coelom. Oviduct thin-walled,

continuous with body cavity; distally with

thickened, glandular walls (oviducal glands).

Nidamental glands in mature animals occupy large

portion of ventral side of mantle cavity. Accessory

nidamental glands anterior to nidamental glands.

Eggs spherical; 2.5—3.1 mm diameter; EgDI 2.5-

2.8-3.2 (SD, 0.4).

Cuttlebone length approximately equal to

mantle length. Subdermal cartilaginous layer

FIGURE 6. Sepia hedleyi Berry, 1918. A,

spermatophore, oral end, male, 85.6 mm (MV F56762)

(CB, cement body; EA, ejaculatory apparatus); B,

enlargement of cement body, male, 90.0 mm ML (MV

F57291). Scale bars; A = 0.15 mm; B = 0,10 mm,

REVISION OF SEPIA HEDLEYI 89

between cuttlebone and skin absent. Cuttlebone

outline oblong (Fig. 7); CbL males 46.2-50.6—

55.0 (SD, 6.2); females 69.3-94.2-107 (SD,

16.9); CbWI males 34.7-36.4-38.1 (SD, 2.4);

females 33.4—35.2-37.8 (SD, 2.0); not strongly

convex in lateral view; CbBI males 9.6—10.0-10.4

(SD, 0.5); females 9.9-10.9-11.9 (SD, 1.1). Bone

acuminate, acute anteriorly (Fig. 2C, D);

acuminate, acute posteriorly (Fig. 2C, D and Fig.

7A, B); not strongly recurved ventrally. Dorsal

surface creamy white; evenly convex; entire

surface calcified with very fine granulose

sculpture; spine and extreme posterior tip of bone

covered with smooth glaze-like substance. Dorsal

median rib present (Fig. 2C and Fig. 7A); distinct;

sides approximately parallel, broaden slightly

anteriorly; bordered laterally by distinct grooves;

lateral ribs present, indistinct. Chitin surrounds

entire margin of cuttlebone. Spine present; short,

pointed; SLI females 4.7—4.9-5.1 (SD, 0.3);

straight, parallel to bone; keel(s) absent;

cuttlebone smooth between spine and outer cone;

ventral notch at base of spine absent. Dorso-

posterior end of cuttlebone without median

longitudinal ridge anterior to spine. Striated zone

flat; extends laterally to inner cone; not separated

from outer cone by smooth marginal zones; StZI

males 61.3-63.2-65.1 (SD, 2.7); females 64.1—

65.5-66.8 (SD, 1.3). Last loculus convex; LoLI

males 32.2-34.1-35.9 (SD, 2.7); females 28.7—

30.8-33.6 (SD, 2.6); at midline half length of

striated zone. LoL/StZ(%) males 49.4-54.0-58.7

(SD, 6.5); females 43.7-45.3-46.9 (SD, 2.2).

Loculus ends at striated zone, does not extend

posteriorly on each side of striated zone. Sulcus

extends entire length of cuttlebone; shallow,

narrow; not flanked by rounded ribs. Last loculus

with shallow median indentation, not very

pronounced. Anterior striae inverted U-shaped.

Limbs of inner cone extend anteriorly to end of

striated zone; inner cone lateral limbs not

separated from outer cone by two distinct smooth

zones. Inner cone limbs uniform width, narrow,

V-shaped posteriorly; not raised to form ledge

FIGURE 7. Sepia hedleyi Berry, 1918. A, cuttlebone, dorsal view, female, 93.1 mm ML (MV _ F57307); B,

cuttlebone, ventral view, specimen as in A. Scale bars = 10 mm.

90 A. L. REID

posteriorly; thickened; shiny; without calcareous

ribs radiating into outer cone. Outer cone present;

calcified; moderate width; narrow anteriorly,

broadens posteriorly; posteriolateral wall without,

or with weak, indentation in both sexes; lateral

limbs not flared ventro-laterally; limbs forming

thin rim ventral to spine.

Body papillae present; dorsal mantle with

longitudinal row of ridge-like papillae along each

side, close to base of each fin; up to six fin

papillae (approximately) in each row. Ventral

mantle without ridges; head and arm papillae

absent.

Ground colour (alcohol preserved specimens)

pale buff pinkish-brown; arms without markings.

Paired dorsal eye spots absent; fins pale. Fins

without markings at base. Ventral pigment

present, pale. Ridges orange-pink in colour.

Distribution

Australia: Queensland, from off the southern

Great Barrier Reef, 22°35.3'S 153°46.7'E around

southern Australia to Western Australia, SW of

Shark Bay, 27°07'S 112°49'E. Depth range 47-

1092 m (average collection depth 218 m).

ACKNOWLEDGMENTS

Thanks to Wolfgang Zeidler from the South

Australian Museum, Adelaide and Ian Loch from the

Australian Museum, Sydney for the loan of specimens. I

would also like to acknowledge C. C. Lu for initiating

this study of the Australian sepiids and for providing

helpful advice. I also thank Dermot Henry from the

Museum Victoria for providing facilities at the Museum

Victoria, Melbourne; and the referees for their helpful

comments.

REFERENCES

ADAM, W. 1979. The Sepiidae (Cephalopoda,

Decapoda) in the collections of the Western

Australian Museum. Records of the Western

Australian Museum 7(2): 111-212.

ADAM, W. & REES, W. J. 1966. A review of the

cephalopod family Sepiidae. Scientific Reports of the

John Murray Expedition 1933-1934 11(1): 1-165.

BERRY, S. S. 1918. Report on the Cephalopoda

obtained by the F.I.S. “Endeavour” in the Great

Australian Bight and other Southern Australian

localities. Biological Results of the Fishing

Experiments carried on by the FIS “Endeavour”,

1909-1914 4(5): 203-296.

CLARKE, M. R. 1986. ‘Handbook for the Identification

of Cephalopod Beaks’. Oxford Science Publications:

New York.

COTTON, B. C. 1929. Contributions to the fauna of

Rottnest Island No. 4. Western Australian Sepiidae.

Journal of the Royal Society of Western Australia

15: 87-94.

COTTON, B. C. 1931. Cuttlebones from Robe with

description of a new species. South Australian

Naturalist 12(3): 39-41.

DALLWITZ, M. J. 1980. A general system for coding

taxonomic descriptions. Taxon 29: 41-46.

DALLWITZ, M. J., PAINE T, A. & ZURCHER, E. J.

1993. ‘User’s Guide to the DELTA System: a

General System for Processing Taxonomic

Descriptions’. CSIRO Division of Entomology:

Canberra.

ICZN 1999. ‘International Code of Zoological

Nomenclature. 4th Edition’. International

Commission on Zoological Nomenclature: London.

IREDALE, T. 1926. The cuttle-fish “bones” of the

Sydney beaches (Phylum Mollusca — Class

Cephalopoda). Australian Zoologist 4: 186-196.

LU, C. C. 1998. A synopsis of Sepiidae in Australian

waters. Pp. 159-190 in ‘Systematics and

Biogeography of Cephalopods’. Vol. 586. Eds N. A.

Voss, M. Vecchione, R. B. Toll & M. J. Sweeney.

Smithsonian Institution Press: Washington DC.

NIXON, M. 1995. A nomenclature for the radula of the

Cephalopoda (Mollusca) — living and fossil. Journal

of Zoology. London 236: 73-81.

NIXON, M. & DILLY, P. N. 1977. Sucker surfaces and

prey capture. Pp. 447-511 in ‘The Biology of

Cephalopods’. Symposia. Vol. 38. Eds M. Nixon &

J. B. Messenger. Zoological Society of London:

London.

PARTRIDGE, T. R., DALLWITZ, M. J. & WATSON,

L. 1993. ‘A Primer for the DELTA System’. CSIRO

Division of Entomology: Canberra.

REID, A. 1991. Taxonomic review of the Australian

Rossiinae (Cephalopoda: Sepiolidae), with a

description of a new species, Neorossia leptodons,

and redescription of N. caroli (Joubin, 1902).

Bulletin of Marine Science 49(3): 748-831.

REID, A. L. 2000. Australian cuttlefishes

(Cephalopoda: Sepiidae): the ‘doratosepion’ species

complex. Invertebrate Taxonomy 14: 1-76.

ROPER, C. F. E. & VOSS, G. L. 1983. Guidelines for

taxonomic descriptions of cephalopod species.

Memoirs of the National Museum of Victoria 44:

48-63.

REVISION OF SEPIA HEDLEYI 9]

APPENDIX 1. Complete list of non-type material examined (not all specimens were sexed and are cited only as

number in each specimen lot).

Australia: Queensland: 1, off GBR, 22°35.3'S

153°46.7'E, 350-345 m, 4 Nov 1985, coll. FV ‘Soela’

(MV F57306); 1F, off Brisbane, 26°40'S 153°41.9'E,

380 m, 2 Aug 1982, coll. M. Potter on ‘Iron Summer’

(MV F57294); 2, 27°11.48'S 153°43.9'E, 230-210 m,

14 Dec 1982, coll. G. Smith on ‘Iron Summer’ (MV

F57299); 1, W Moreton I., 27°35'S 153°5S0'E, 210 m, 15

Dec 1982, coll. G. Smith on ‘Iron Summer’ (MV

F57309); many 27°46'S 153°5S1'E, 205 m, 23 Mar 1983,

coll. C. C. Lu on ‘Iron Summer’ (MV F57308); 1M, 1F,

27°58'S 153°49'E, 220 m, 23 Mar 1983, coll. ‘Iron

Summer’ (MV F57300); many, off N Stradbroke L.,

27°58'S 153°51.5'E, 183 m, 28 Jul 1982, coll. ‘Iron

Summer’ (MV F89852); 1M, 9F, 32°23'S 152°49'E, 278

m, 30 Jan 1982, coll. C. C. Lu & R. Tait on FV ‘Soela’

(MV F77180); 10M, 6F, off NSW, 32°23'S 152°59'E,

278 m, 30 Jan 1982, coll. C. C. Lu & R. Tait on FV

‘Soela’ (MV F77191); 3M, 6F, 32°23'S 152°59'E, 278

m, 30 Jan 1982, C. C. Lu & R. Tait on FV ‘Soela’ (MV

F77182); 7M, 5F, 32°24'S 152°56'E, 246-244 m, 30 Jan

82, coll. C. C. Lu & R. Tait on FV ‘Soela’ (MV

F77084). New South Wales: 4, off Port Stephens,

32°24'S 152°56'E, 240 m, 30 Jan 1982, coll. FV ‘Soela’

(AM C152609); 2M, 10F, 32°27'S 152°54'E, 244-242

m, 30 Jan 1982, coll. FV ‘Soela’ (MV F77178); 1M,

2F, 32°27'S 152°54'E, 244-242 m, 30 Jan 1982, coll. C.

C. Lu & R. Tait (MV F77179); 14, off Port Stephens,

32°50—-52'S 152°42-41'E, 550 m, 6 Dec 1978, coll. K. J.

Graham, W. B. Rudman & P. H. Colman on FRV

‘Kapala’ (AM C152641); 2, 8 mls E of Port Stephens,

101 m, 18 Aug 1975, coll. K. J. Graham & C, Short on

FRV ‘Kapala’ (AM C100805); many, off Newcastle,

33°01'S 152°01'E-33°03'S 151°58'E, 121 m, 28 Oct

1993, coll. FRV ‘Kapala’ (MV F89853); 17, 33° 06'S

124°33'E, 28 Nov 1981, coll. FV ‘Soela’ (MV F89854);

4, off Sydney 33°33-36'S 151°59-57'E, 373-366 m, 21

Dec 1976, coll. K. J. Graham & P. H. Colman on FRV

‘Kapala’ (AM C105862); 1F, 33°40'S 151°5S0'E, 210-

204 m, 25 Jan 1982, coll. C. C. Lu & R. Tait (MV

F77138); many, 33°42'S 151°51'E, 300-293 m, 25 Jan

1982, C. C. Lu & R. Tait (MV F77136); many, off

Sydney, 33°45-34'S 151°39-40'E, 143 m, 5 Dec 1978,

coll. FRV ‘Kapala’ (AM C152596); many, E of

Cronulla, 34°05-11'S 151°19-14'E, 132 m, 22 Apr

1975, coll. FRV ‘Kapala’ (AM C152594); many, E of

Wollongong, 34°19-20'S 151°19-18'E, 161 m, 12 Dec

1978, coll. K. J. Graham, W. B. Rudman & P. H.

Colman on FRV ‘Kapala’ (AM C152646); many,

Wreck Bay, 35°13'S 150°41'E-35°12'S 150°44'E, 60-47

m, 20 Oct 1993, coll. FRV ‘Kapala’ (MV F89855);

many, off NSW, 36°20'S 150°20'E, 130 m, coll. FV

‘Soela’ (MV F77164); 1, 34°22'S 151°23'E, 26 Mar

1981, coll. FV ‘Soela’ (MV F89856); 1F, 36°26'S

150°12'E, 104 m, 21 Apr 1981, coll. FV ‘Soela’ (MV

F77207); 2, off Tathra 36°35'S 150°11'E-36°39'S

150°10'E, 117-115 m, 8 Dec 1993, coll. K. Graham on

FRV ‘Kapala’ (MV F89857); 2M, IF, 25 km SSE

Merimbula, 36°56.5'S 150°11.8'E-36°54.1'S 150°11.9'E,

115 m, 5 Aug 1993, coll. RV ‘Southern Surveyor’ (MV

F89858); 1F, NE Twofold Bay, 37°05'S 149°54'E, 29

Dec 1960, coll. R. Slack-Smith (MV F89859); several,

off Eden, 131.7 m, FRV ‘Kapala’ (AM C174072); 5M,

8F, E of Disaster Bay, 37°24'S 150°17.S'E, 172-162 m,

2 Aug 1993, coll. FRV ‘Southern Surveyor’ (MV

F89860). Victoria: 4, 38°01.7'S 150°04.9'E, 3 Feb

1985, coll. FV ‘Soela’ (MV F89861); 5, 38°02'S

150°0S'E, 4 May 1984, coll. FV ‘Soela’ (MV F51887);

1M, 3F, 38°06'S 149°55'E, 270-267 m, 18 Jan 1982,

coll. C. C. Lu & R. Tait (MV F77112); 4, 38°09.1'S

149°54.0'E, 3 Feb 1985, coll. FV ‘Soela’ (MV F89862);

4, 38°13'S 149°43'E, 4 May 1984, coll. FV ‘Soela’ (MV

F51888); 3, 38°15.4'S 149°19.5'E, coll. FV ‘Soela’ (MV

F89863); 2, 38°34.25'S 148°32.25'E, 24 Aug 1982, coll.

MSL (MV F89864); 1F, E Bass Strait 38°34.3'S

148°16.1'E, 86 m, 6 Jun 1984, coll. P. Moulton, MSL

(MV F89865); IF, 18 mls S of Cape Nelson, 38°44'S

143°33'E, 152.9 m, 26 Aug 1975, coll. FV ‘Sarda’ (MV

F56764); 1F, off Portland, 38°50'-S1'S 141°46'-55'E,

54.8 m, 5 Mar 1980, coll. M. Gomon on ‘Halcyon’

(MV F30332); 3, 38°50'S 141°46'E, 6 Mar 1980, coll.

‘Halcyon’ (MV F30843); 1M, IF, eastern Bass Strait,

38°53.05'S 148°24.2'E, 126-101 m, 8 Feb 1981, coll.

RV ‘Hai Kung’ (MV F57310); IF, 38°55.6'S

148°27.7'E, 210-150 m, 27 Aug 1994, coll. FRV

‘Southern Surveyor’ (MV F89866); many, 39°16'S

145°05.08'E-38°77'S 145°07.64'E, 65.9 m, 3 Feb 1981,

coll, M. F. Gomon et al. on RV ‘Hai Kung’ (MV

F89867); 1 juv., E Bass Strait, 39°28.4'S 148°41.8'E,

110 m, 28 Mar 1979, coll. HMAS ‘Kimbla’ (MV

F57292); many, central Bass Strait, 39°44.55'S

143°33.82', 78.7 m, coll. C. C. Lu on ‘Hai Kung’ (MV

F57296). Tasmania: 1M, S of Flinders I., 40°43.79'S

148°32.7'E, 58.6-54.9 m, 7 Feb 1981, coll. FV ‘Hai

Kung’ (MV F57311); 1F, SE Tasmania, 42°42.8'S

148°24.4'E, 446 m, 26 Jun 1984, coll. FV ‘Soela’ (MV

F52265); 4, 42°38'S 148°24'E, 23 Jun 1984, coll. FV

‘Soela’ (MV F52100); 3, 42°42.8'S 148°24'E, 26 Jun

1984, FV ‘Soela’ (MV F51908); 6, 42°43.7'S

148°22.3'E, 15 Jun 1984, coll, FV ‘Soela’ (MV

F52099); 1F, off Tasmania, 43°38.9'S 147°49.4'E, 160

m, 16 Feb 1976, coll. K. Nesis on RV ‘Dmitry

Mendeleev’ (MV F57307). South Australia: 4F, 27 nm

SW of Beachport, 37°50'S 139°46'E, 1092 m, 24 Oct

1981, coll. M. F. Gomon & R. Wilson on ‘Halcyon’

(MV F52793); 1F, off Beachport, 37°51'S 139°48'E,

437-400 m, 24 Oct 1981, coll. M. F. Gomon & R.

Wilson on ‘Halcyon’ (MV F51369); 2F, off Beachport,

180 fm [329 m], Apr 1981 (SAM D19239). Western

92 A. L. REID

Australia: 1M, 1F, SW of Shark Bay, 27°07'S

112°49'E, 248-238 m, 3 Mar 1981, coll. M. F. Gomon

on ‘Hai-Kung’ (MV F56762); 1F, Great Australian

Bight, 12.0 km S of Middini Beach, 33°17.4'S

127°44.5'E-33°17.1'S 127°31.25'E, 180-167 m, 14 Feb

1990, coll. R. Poole on FV ‘Comet’ (MV F89868); 1F,

Great Australian Bight, 33°20'S 128°10'E-33°22'S

128°08'E, 200 m, 29 Sep 1980 (MV F56763).

REVISION OF SEPIA HEDLEYI 93

APPENDIX 2. Description of measurements and counts. Definitions largely follow Roper and Voss (1983). New or

modified definitions are indicated by an asterisk (*). Indices (shown in square brackets) are calculated by expressing

each measure as a percentage of mantle length or, for cuttlebone characters, cuttlebone length (unless otherwise

specified).

Arm Length - AL: length of each designated (ie 1,2 etc)

arm measured from first basal (proximal-most)

sucker to distal tip of arm (Arm 1, dorsal; 2, dorso-

lateral; 3, ventro-lateral; 4, ventral) [ALI].

Anterior Mantle to Head length *: AMH: dorsal length

of mantle measured from anterior-most point of

mantle to intersection of transverse line joining

dorso-lateral mantle margin [AMHI].

Arm Sucker Count *: ASC: total number of suckers on

each designated arm (eg.ASC2).

Arm Sucker diameter: AS: diameter of largest normal

sucker on each designated (ie 1,2 etc) arm [ASIn];

ASl4m* diameter of smallest sucker on modified

portion of left ventral arm of males [ASInI4m].

Cuttlebone Breadth*: CbB: greatest dorso-ventral width

of cuttlebone.

Cuttlebone Length: CbL: dorsal length of cuttlebone

along midline, including spine.

Cuttlebone Width: CbW: greatest lateral width of

cuttlebone [CbWI].

Club Length: CIL: length of tentacular club measured

from proximal-most basal suckers (carpus) to distal

tip of club [CILT].

Club Row Count: CIRC: number of suckers in

transverse rows on tentacular club.

Club Sucker diameter: CIS: diameter of largest sucker

on tentacular club [CISTI].

Club Sucker dorsal *: ClSd: diameter of largest

tentacular club sucker in dorsal-most (closest to

swimming keel) longitudinal row [CISId].

Club Sucker ventral *: ClSv: diameter of largest

tentacular club sucker in ventral-most (opposite

swimming keel) longitudinal row [CISIv].

Eye Diameter: ED: diameter of eye [EDI].

Egg Length *: EgL: length of egg [EgLI].

Egg Width *: EgW: width of egg [EgWI).

Free Funnel length: FFu: the length of the funnel from

the anterior funnel opening to the point of its dorsal

attachment to the head [FFul].

Fin Insertion anterior *: Fla: anterior origin of fin

measured from mantle margin to anterior-most

junction of fin and mantle [FIIa].

Fin Insertion posterior*: FIp: measured between

posterior junctions of fins with mantle [FIIp].

Funnel Length: FuL: the length of the funnel from the

anterior funnel opening to the posterior margin

measured along the ventral midline [FuLI].

Fin Width: FW: greatest width of single fin [FWI].

Gill Lamellae Count: GiLC: number of lamellae on

outer demibranch including the terminal lamella.

Gill Length *: GiL: length of gill [GiL]).

Head Length: HL: dorsal length of head measured from

point of fusion of dorsal arms to anterior tip of

nuchal cartilage [HLI).

Head Width: HW: greatest width of head at level of

eyes [HWI].

Loculus Length *: LoL: length of the last loculus

(ventral anterior smooth zone of the cuttlebone)

[LoL].

Mantle Length: ML: dorsal mantle length. Measured

from anterior-most point of mantle to posterior

apex of mantle.

Mantle Width: MW: greatest straight-line ventral width

of mantle [MWI].

Spine Length *: SL: length of spine [SLI].

Spermatophore Length: SpL: length of spermatophore

[SpLI].

Spermatophore Width: SpW: greatest width of

spermatophore. Spermatophore width index is

expressed as a percentage of spermatophore length

[SpwI).

Striated Zone length: StZ: length of striated zone of

cuttlebone [StZI].

Transverse Row Count: TrRC: number of suckers in

longitudinal series on tentacular club (counted

from proximal-most basal suckers (carpus) to distal

tip of club).

Ventral Mantle Length: VML: length of ventral mantle

measured from anterior mantle margin at ventral

midline, to posterior apex of mantle [VMLI].

94 A. L. REID

APPENDIX 3. Measurements (mm), counts and indices of Sepia hedleyi Berry, 1918 from southern Australia.

Museum SAM SAM SAM SAM SAM SAM SAM SAM

Reg. no. D19241 D19243 D19241 D19242 = Di9242 D19240 D19240 D19240

SEX M M M F E F F F

ML 47.2 55.0 75.7 67.9 68.6 93.1 95.5 107.6

MWI 55.1 55.6 51.8 55.5 45.5 54.7 59.8 55.0

AMHI 11.9 15.5 15.3 14.7 13.0 12.8 14.8 17.8

VMLI 85.0 80.7 82.8 85.6 84.8 81.2 81.8 75.3

FWI 11.7 10.2 9.4 7.2 8.2 7.8 9.1 4.4

Fila 5.1 4.5 5.3 2.8 3.2 4.9 3.7 2.0

FIIp 6.4 8.0 74 5.6 12.4 10.3 4.7 6.7

FuLI 40.3 34.5 27.7 34.6 34.3 25.6 30.4 28.8

FFul 20.1 14.5 17.2 14.7 16.8 19.3 12.6 13.0

HLI 25.6 27.5 29.7 23.9 13.4 40.8 30.3 35.8

HWI 38.1 44.4 35.7 44.0 45.0 40.5 46.6 43.5

EDI 13.3 16.5 12.7 10.0 12.1 12.7 8.9 10.2

ALI 33.9 34.5 35.7 33%] 34.3 37.6 49.2 42.3

ALI2 33.9 30.9 29.1 31.7 35.7 39.2 42.4 42.8

ALI3 38.1 33.6 29.1 37.6 34.3 41.4 44.0 43.7

ALI4 43.4 40.9 34.3 39.0 42.3 41.9 51.3 44.6

ASIn1 1.69 1.36 1.32 1.47 1.53 172 1.68 1.77

ASIn2 1.48 1.36 1.59 1.47 1.53 2.15 1.68 1.77

ASIn3 1.48 1.36 1.59 1.47 1.75 2.15 1.78 E77,

ASIn4 1.69 1.36 1.32 1.33 1.56 2.15 1.78 1.77

ASC1 118 112 116 148 144 106 106 144

ASC2 140 152 148 148 156 130 175 176

ASC3 148 132 144 200 172 157 164 172

ASC4 180 208 164 240 248 193 212 224

ASInl4m 0.42 0.40 0.26 - ~ - ~ -

CILI 17.2 - 13.2 18.6 16.2 - 21.2 24.2

CIRC 10 10 10 10 10 - 9 10

TrRC 42 - 32 36 32 - 40 34

CISI 0.42 0.40 0.53 0.40 0.47 - 0.52 0.46

CISId 0.42 0.40 0.42 0.40 0.47 - 0.52 0.46

CISIv 0.42 0.40 0.29 0.32 0.32 - 0.52 0.46

GiLC 29 29 29 29 30 - - -

GiLI 35.0 24.9 30.1 29.5 38.2 - 33.6 34.5

SpLI 12.3 11.6 10.4 ~ - - - -

SpWI 3.45 4.69 5.06 ~ - - - -

EgDI - - - - - 2.7 322, 2.5

CbL 46.2 55.0 - - 69.3 100.5 99.8 107.0

CbWI 38.1 34.7 - - 37.8 35.5 33.4 33.9

CbBI 10.4 9.6 - ~ 11.8 9.9 10.2 11.9

SLI - - - - - 4.7 Se -

StZI 61.3 65.1 - - 66.8 65.6 64.1 -

LoLI 35.9 32.2 - - - 28.7 30.1 33.6

LoL/StZ (%) 58.7 49.4 ~ - - 43.7 46.9 -

REVISION OF SEPIA HEDLEYI 95

APPENDIX 4. Measurements (mm), counts and indices of 10 male Sepia hedleyi Berry, 1918 from eastern

Australia.

Museum MV MV AM AM AM MV MV MV MV AM

Reg.no. F77136 F77179 C152623 C152623 C152623 F77136 F77136 F77136 F77136 C152623

ML 75.9 84.3 84.8 85.7 86.1 87.0 90.9 91.7 100.5 103.5

MWI 53.0 49.2 50.7 48.4 49.2 54.6 51.7 54.1 48.1 46.6

AMHI 16.2 15.1 12.5 13.0 15.0 14.4 13.3 12.5 11.9 14.9

VMLI 88.8 88.3 89.2 91.4 87.8 90.9 90.4 88.3 81.9 88.3

FWI 9.6 11.4 9.4 10.5 8.2 10.5 12.7 9.8 5.6 10.9

Fila 4.2 3.3 3.8 4.8 4.8 6.6 3.7 3.8 7.0 4.4

FIlIp 5.8 3.0 5.8 5.6 Ted 75 3.5 8.2 4.5 8.6

FuLI 32.9 31.2 30.4 31.0 31.1 36.6 30.3 31.7 32.7 30.7

FFul 14.5 15.4 15.2 ATT 15.2 18.4 15.4 20.7 16.8 13.0

HLI 30.4 23.3 317. 27.3 31.6 30.9 23:7 30.9 30.3 26.5

HWI 38.3 38.7 35.1 36.6 36.5 40.9 40.9 38.8 - 34.3

EDI 15.5 11.0 13.2 14.0 11.0 12.0 13.1 15.7 12.5 12.3

ALT 38.2 33.2 33.6 32.8 26.7 37.4 33.0 33.8 27.9 26.1

ALI2 34.3 32.0 30.7 31.5 25.6 37.4 34.1 31.6 27.9 32.9

ALI3 36.9 32.0 34.2 32.7 29.0 36.8 31.9 28.4 31.4 33.3

ALI4 40.8 37.4 36.6 30.2 30.8 40.2 34.1 37.6 40.6 32.8

ASIn1 1.54 1.59 1.42 1.27 1.21 1.64 1.43 1.42 1.42 1.22

ASIn2 1.45 1.48 1.13 1.33 1.02 1.49 1.43 1.56 1.42 1.28

ASIn3 1.37 1.48 1.49 1.46 1.30 1.64 1.43 1.56 1.42 1.39

ASIn4 1.45 1.48 1.27 1.25 1.11 1.64 1.43 1.42 1.42 1.16

ASC1 96 93 72 89 76 93 89 86 92 119

ASC2 136 128 105 106 93 126 124 120 135 121

ASC3 139 142 121 112 101 124 124 142 144 123

ASC4 204 202 171 151 102 185 178 196 - 140

ASInl4m 0.47 0.37 0.25 0.28 0.29 0.36 0.40 0.36 0.39 0.20

CILI 13.7 15.5 15:3 17.4 14.2 16.1 15.8 15.5 15.8 19.8

CIRC 12 11 11 12 11 12 12 12 12 12

TrRC 36 32 31 31 36 34 35 33 35 39

CISI 0.51 0.52 0.65 0.53 0.64 0.60 0.43 0.43 0.52 0.68

ClSId 0.51 0.66 0.65 0.46 0.64 0.45 0.35 0.35 0.52 0.53

CISIv 0.38 0.52 0.35 0.42 0.35 0.45 0.43 0.39 0.39 0.48

GiLC 30 31 34 32 27 33 32 32 32 32

GiLI 30.3 26.0 27.9 30.9 33.7 33.6 26.7 29.8 - 22.2

SpLI 10.9 8.7 - - - 9.7 9.1 9:2 8.5 -

Spwl 3.15 3.01 - - - 4.62 3.15 4.62 4.55 -

CbL 75.8 85.1 84.8 - - 86.5 92.7 90.0 100.2 101.6

CbWI 36.1 35.6 32.0 - - 34.9 32.9 34.7 31.2 31.3

CbBI 10.0 10.5 10.0 - - 9.5 9.2 9.9 10.0 9.0

SLI - 4.3 4.1 - - - 2.8 3.0 3.9 2.6

StZI 63.6 60.2 59.0 - - 46.6 66.1 61.7 65.0 64.5

LoLI 37.6 33.0 32.5 - - 34.0 28.5 31.1 31.8 29.4

LoL/StZ (%) 59.1 54.9 55.2 - - 73.0 43.1 50.5 49.0 45.6

96 A. L. REID

APPENDIX 5. Measurements (mm), counts and indices of 10 female Sepia hedleyi Berry, 1918 from eastern

Australia.

Museum MV MV MV MV MV MV MV MV MV MV

Reg. no. F77179 ~=F77136 = F77136—s-F77136 —s-F77136 =~-F77136 =F77136 =F77136 =F77179_ ~——F77138

ML 83.1 89.0 91.7 94.7 99.1 99.4 99.8 102.1 103.8 108.8

MwWI 51.9 52.2 52.1 51.3 S34 46.7 51.9 58.1 47.1 51.5

AMHI 16.1 12.9 12.3 14.8 15.9 13.6 13.9 14.8 13.2 14.8

VMLI 85.0 91.0 91.1 89.2 86.0 90.0 89.7 87.9 86.6 88.8

FWI 12.0 10.1 11.7 11.9 11.6 12.4 10.5 11.0 9.6 74

Fila 4.5 7.0 4.1 4.9 3.6 4.0 4.2 3.1 4.6 6.3

FiIp 13.7 8.4 4.3 6.7 6.6 7A 5.1 2.8 4.5 7.2

FuLI 30.7 38.3 34.4 32.5 32.2 33.7 33.6 32.6 27:3 35.2

FFul 16.8 15.7 17.4 18.5 18.2 18.1 16.5 16.7 15.4 13.3

HLI 27.0 21.3 37.1 26.1 27.7 36.2 39.1 275. 26.8 26.4

HWI 39.4 36.4 39.8 38.5 39.9 41.6 40.7 39.5 38.8 36.5

EDI 12.4 12.4 14.1 11.7 14.3 13.6 12.4 16.4 13.0 12.8

ALTI1 33.7 32.6 38.2 35.9 41.4 342. 36.1 41.1 34.3 31.3

ALI2 34.9 30.3 34.9 32.7 40.4 34.2 36.1 38.2 32.8 35.8

ALI3 36.1 34.8 39.3 34.3 40.4 41.2 41.1 42.6 36.6 32.2

ALI4 39.7 42.7 37.1 39.6 46.4 46.3 38.1 44.1 40.5 41.4

ASIn1 1.56 1.46 1.34 1.58 1.84 1.70 1.56 1.40 1.42 1.43

ASIn2 1.56 1.61 1.48 1.52 1.84 1.70 1.56 1.66 1.45 1.49

ASIn3 1.56 1.46 1.48 1.48 1.84 1.63 1.69 1.66 1.36 1.55

ASIn4 1.72 1.53 1.48 1.52 1.71 1.70 1.69 1.66 1.45 1.55

ASC1 106 112 94 110 120 96 106 102 98 92

ASC2 138 133 158 166 156 156 162 170 142 138

ASC3 150 142 148 192 168 162 154 168 162 138

ASC4 178 162 192 227 204 206 214 110 214 171

CILI 15.4 15.7 15.0 15.2 18.3 16.3 18.5 16.7 15.2 18.4

CIRC 12 12 12 11 12 12 12 12 12 12

TrRC 27 35 47 29 38 30 32 36 30 38

CISI 0.67 0.51 0.43 0.57 0.32 0.71 0.45 0.64 0.72 0.53

ClSId 0.49 0.44 0.43 0.57 0.32 0.58 0.39 0.64 0.75 0.53

ClSIv 0.42 0.40 0.43 0.57 0.32 0.52 0.42 0.44 0.54 0.41

GiLC 28 33 32 30 35 28 31 28 33 30

GiLI 24.5 23.0 25.8 25.3 32.0 30.5 27.5 30.7 27.2 31.6

EgDI 3.9 4.1 5.0 323 4.4 4.8 3.4 3.9 4.8 3.3

CbL 82.1 87.0 89.0 99.7 98.0 99.2 96.6 105.2 104.9 108.0

CbWI 34.0 35.3 34.2 32.1 36.8 34.6 37.9 37.0 35.3 34.3

CbBI 10.0 - 10.1 11.2 9.4 10.9 10.1 9.5 10.0 9.6

SLI 3.3 - ~ 3.3 3.2 - 3.0 4.6 49 -

StZI 65.0 - 79.4 79.4 67.3 65.4 67.7 63.1 63.5 64.6

LoLI 22.4 - 20.6 20.6 21.1 20.3 27.3 30.6 30.9 35.4

LoL/StZ (%) 34.5 - 25.9 25.9 31.4 31.0 40.4 48.5 48.6 54.7

REVISION OF SEPIA HEDLEYI 97

APPENDIX 6. Sepia hedleyi Berry, 1918; ranges of arm length indices (ALI), arm sucker diameter indices (ASIn)

and arm sucker counts (ASC) of mature males and females from southern Australia; N = number of specimens,

min. = minimum, max. = maximum, SD = standard deviation.

Males Females

N min. mean max. SD N min. mean max. SD

ALI1 7 25.3 34.9 $5.7 9.8 8 30.6 374 49.2 6.2

ALI2 7 25.9 L5 45.6 6.8 8 27.6 35.8 42.8 5.6

ALI3 7 29.1 33.5 44.3 5.8 8 30.6 33.7 44.0 5.2

ALI4 7 33.8 40.2 54.4 7.2 8 34.7 42.0 $1.3 4.8

ASIn1 3 1.32 146 1.69 0.20 5 1.47 1.63 1.77 0.13

ASIn2 3 1.36 148 1.59 0.11 5 1.47 172 2.15 0.27

ASIn3 3 1.36 148 1.59 0.11 5 1.47 178 2.15 0.24

ASIn4 3 1.32 1.46 1.69 0.20 5 1.33 L72 2.15 0.30

ASC1 3 112 U5 118 3 5 106 130 148 22

ASC2 3 140 147 152 6 5 130 157 176 19

ASC3 3 132 141 148 8 5 157 1733 200 16

ASC4 3 164 184 208 22 5 193 223 248 22

A NEW GENUS AND SIX NEW SPECIES OF DYTISCIDAE

(COLEOPTERA) FROM UNDERGROUND WATERS IN THE YILGARN

PALAEODRAINAGE SYSTEM OF WESTERN AUSTRALIA

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

Summary

A new genus and six new species of stygobitic beetles of the family Dytiscidae, subfamily

Hydroporinae, from relatively shallow calcrete aquifers in Western Australia, are described and

figured. The new genus (Nirripirti) and species (N. hinzea) are in the tribe Hydroporini, whlst the

remaining species belong in the tribe Bidessini, namely, Nirridessus challaensis, N. masonensis, N.

fridaywellensis, N. pinnaclesensis and Tjirtudessus raesidensis. The species are members of a rich,

recently discovered, relictual stygofauna, predominantly of Crustacea and Oligochaeta, inhabiting

calcretes lying along palaeodrainage channels. The new genus represents a significant taxonomic

extension of the Australian stygobitic Dytiscidae, being the first Australian stygobitic member of its

tribe. The new species represent a significant geographic extension of Australian stygobitic

Dytiscidae into a new palaeodrainage channel unconnected with any previously examined. Each

calcrete area examined contained a distinct assemblage of beetles often with two species in

sympatry.

A NEW GENUS AND SIX NEW SPECIES OF DYTISCIDAE (COLEOPTERA)

FROM UNDERGROUND WATERS IN THE YILGARN PALAEODRAINAGE

SYSTEM OF WESTERN AUSTRALIA

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

WATTS, C. H. S. & HUMPHREYS, W. F. 2001. A new genus and six new species of

Dytiscidae (Coleoptera) from underground waters in the Yilgarn palaeodrainage system of

Western Australia. Records of the South Australian Museum 34(2): 99-114.

A new genus and six new species of stygobitic beetles of the family Dytiscidae, subfamily

Hydroporinae, from relatively shallow calcrete aquifers in Western Australia, are described

and figured. The new genus (Nirripirti) and species (N. hinzeae) are in the tribe Hydroporini,

whilst the remaining species belong in the tribe Bidessini, namely, Nirridessus challaensis, N.

masonensis, N. fridaywellensis, N. pinnaclesensis and Tjirtudessus raesidensis. The species

are members of a rich, recently discovered, relictual stygofauna, predominantly of Crustacea

and Oligochaeta, inhabiting calcretes lying along palaeodrainage channels. The new genus

represents a significant taxonomic extension of the Australian stygobitic Dytiscidae, being the

first Australian stygobitic member of its tribe. The new species represent a significant

geographic extension of Australian stygobitic Dytiscidae into a new palaeodrainage channel

unconnected with any previously examined. Each calcrete area examined contained a distinct

assemblage of beetles often with two species in sympatry.

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 1 February 2001.

The presence of an extensive stygofauna

consisting of Crustacea, Oligochaeta and beetles

of the family Dytiscidae living in near-surface

aquifers of the Western Desert areas of Western

Australia has recently been documented by us

(Watts & Humphreys 1999, 2000). In this paper

we report on significant geographic and

taxonomic extensions to the known fauna of

Dytiscidae.

In our two previous papers we described

eleven species of Dytiscidae belonging to three

genera all in the subfamily Hydroporinae. One

genus, Kintingka, known from only one female

specimen, could not unequivocally be placed in

a tribe but the other two genera possessed the

defining synapomorphy for the _ tribe

Bidessini—two-segmented parameres. In the

most recent collection reported on here most

specimens again belonged in the same two

Bidessine genera, Tjirtudessus and Nirridessus.

A series of relatively large specimens from one

location, however, had single segmented

parameres and other morphological characters

in both adults and associated larvae which place

them in the tribe Hydroporini, the first

underground members of this tribe to be

recorded from Australia, although they are a

major component of the stygodytiscidae world-

wide (Spangler 1986).

The Raeside palaeodrainage system (Fig. 40),

from which some of the latest specimens were

collected, is unconnected to the drainage systems

from which we earlier reported underground

Dytiscidae (Watts & Humphreys 2000: Fig. 38);

this represents a significant extension of the

geographical, as well as the taxonomic range, of

the Yilgarn stygofauna. As for the systems

reported in our earlier papers, numerous

specimens of Crustacea—bathynellids,

harpacticoid and cyclopoid copepods, ostracods

and oniscid isopods—and some Oligochaeta and

Hydracarina were collected. As before, the beetles

and larger stygofauna were restricted to aquifers

in areas of calcrete, as is the stygofauna found in

the northern parts of the Western Shield (Poore &

Humphreys 1998; Humphreys 1999 in press). But,

unlike our previous observation that the

stygofauna was found only in narrow bores, thus

seemingly restricted to ‘closed’ aquifers (Watts &

Humphreys 2000), in this recent survey

stygofauna were found to be common in formed-

wells dug for water as well as in narrow bore-

holes drilled for geological purposes, water

monitoring, water pumping or aquifer assessment.

100

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

METHODS

The collection methods and measurements of

physicochemical parameters in the water were the

same as in our previous papers (Watts &

Humphreys 1999, 2000).

Abbreviations used:

BES

Prefix for field numbers, WAM

Biospeleology.

SAMA South Australian Museum, Adelaide.

WAM Western Australian Museum, Perth.

SYSTEMATICS

KEY TO AUSTRALIAN SPECIES OF STYGOBITIC

DyTISCIDAE

Profemur with small peg-like structure

close to base; metatibia approximately the

same width throughout 0.0.0... eceeeeeeee

sadseceytiaeereaesese Nirripirti hinzeae sp. nov.

Profemur lacking such a structure;

metatibia narrow at base then strongly

expanding towards apex ........eeeeeeees 2

Body length approximately 1.0 mm; legs

stout, without swimming-hairs on fore-

and Mmidlegs .0...... ses eeeesesseeeeeeeeeeeseeseees

Kintingka kurutjutu Watts and Humphreys

Body length > 1.2 mm; legs normal, all

with swimming-haifS ............eeeeeeeeees 3

Third and fourth (apical) segments of labial

palpi subequal in length; setae on hind

edge of mesofemur not greatly stronger

than those on mesotrochanter; length

> 3.0 mm (Tjirtudessus) ........:000c0+ 14

Third segment of labial palpi half to two-

thirds length of apical; setae on hind edge

of mesofemur near base much stronger

than those on mesotrochanter or elsewhere

on femur; length 1.3-3.2 mm.

(NE TIdESSUS) saicisg cs onayde eben ceiset Oe car incsvnns 4

Pronotal plicae strong, well marked ....5

Pronotal plicae weak, difficult to trace ..

Length of first two segments of metatarsi

> rest; eye remnant present; parameres

with long apical lobe .............eeeeeeeeeee

$ahedegzerss N. pulpa Watts and Humphreys

Length of first two segments of metatarsi

=< rest; without eye remnant; parameres

10.

11.

12.

with small apical lobe ...........ee eee

BS N. morgani Watts and Humphreys

Length => 2.9 mm; eye remnant present;

group of six spines close to base of

mesofemur on hind edge ..............:000

... N. bigbellensis Watts and Humphreys

Length < 2.6 mm; with or without eye

remnant; spines on mesofemur spread out

along hind edge or if restricted to base four

OL TOWEL Nase Siacangiaieoncebcenadp onaavontstyeaieees 7

Row of larger punctures adjacent to suture;

metatrochanters rounded ..............00 8

Without sutural punctures,

metatrochanters bluntly pointed .......... 9

Length 2.2—2.3 mm; with eye remnant ..

N. windarraensis Watts and Humphreys

Length 1.3-1.5 mm; without eye remnant

sisees N. lapostaae Watts and Humphreys

Apical segment of antenna approximately

the same size as penultimate, segments 3

and 4 subequal; pro- and mesotarsi strongly

expanded, approximately equal in size ..

srectaeteas Nirridessus challaensis sp. nov.

Apical segment of antennae approximately

1.5x length of penultimate, segment 3 >

segment 4; protarsus moderately or weakly

expanded, much more so than mesotarsi

bs lvedaneventton ingotnstdsivacttdenpoeettvangt naiteeBbaycenegt 10

Mesofemur with four to five spines spread

out along hind edge in basal half...........

evsiedest sed Nirridessus masonensis sp. nov.

Mesofemur with two to four strong spines

clustered on hind edge close to base.. 11

Mesofemur with four spines near base;

segments 2 and 3 of antennae similar in

size, apical segment approximately 1.5x

penultimate in length........... Nirridessus

cueensis Watts and Humphreys

Mesofemur with two to three strong spines

on hind edge near base; segment 2 of

antenna large and oval, segment 3 much

smaller and thinner, apical segment

approaching 2x length of penultimate ...

ssvareuastav@ivaverercaizseteatheltestesdeasttesspettedt 12

Mesofemur with two strong spines on

hind edge near base; apical segment of

paramere with two finger-like projections

wheats Nirridessus pinnaclesensis sp. Nov

Mesofemur with three strong spines on

hind edge near base; apical segment of

paramere with one finger-like projection

Wire sa Mecuegin toaua depended daevapaerueap bubedadet vatahe 13

NEW UNDERGROUND DYTISCIDAE, WESTERN AUSTRALIA 101

Apical lobe of paramere overlapping rest

of apical segment; central lobe pointed;

protarsus weak, similar in size to mesotarsi;

segment 3 of antenna >> segment 4 in

lengthNirridessus fridaywellensis sp.nov.

— Apical lobe of paramere well separated

from rest of apical segment; central lobe

bluntattip; segment 3 of antenna> segment

4 in length; protarsus moderately

expanded, mesotarsus less SO «1.0.4...

Nirridessus hinkleri Watts and Humphreys

14. — Pro- and mesotarsi strongly expanded;

apical four segments of antennae

noticeably thinner than others................

Lashes tavererdetySpacB ye M caetudbgedesnsed Tjirtudessus

magnificus Watts and Humphreys

— Pro- and mesotarsi only moderately

expanded; apical four segments of

antennae not narrower than others..... 15

15. — Length >3.8 mm; pronotum a little

narrower than elytra ...........::eeeeeeeeee 16

— Length <3.8 mm; pronotum wider than

GL YU As 0. .tchercaserBettendecserrenzes Tjirtudessus

eberhardi Watts and Humphreys

Metatrochanters rounded at tip; central

lobe of aedeagus straight, tip pointed;

with small eye remnant ...........0:::cceeee

endeas Tjirtudessus raesideensis sp. nov.

— Metatrochanters pointed at tip; central lobe

of aedeagus twisted, tip knobbed; without

SYS TEMMANE: ves sesessecgesassgetevestincesegeseg heres

Tjirtudessus hahni Watts and Humphreys

16. —

Tjirtudessus Watts and Humphreys, 1999

Tjirtudessus raesideensis sp. nov.

(Figs 1--7)

Types

Holotype: m. ‘BES 8354, Salt Well, Lake

Mason Station, 27°32'24"S, 119°37'27"E, 24/6/00,

coll W. F. Humphreys, S. Hinze’ in spirit, WAM

27601.

Paratypes: 32, as for holotype, 19 SAMA, 13

WAM 27602-27615; 22, ditto except ‘BES

8355’, 10 SAMA, 12 WAM 27616-27627; 2,

ditto except ‘BES 8350’, SAMA.

Description (number examined, 57)

Habitus. Length 3.5-4.0 mm; relatively flat,

weakly constricted at junction of pronotum/

elytra; uniformly light testaceous; hindwing

vestigial, about half length of elytron.

Head. Narrower than elytra; smooth,

reticulation very weak, punctures sparse, very

small; subparallel in posterior half, widest just

behind eye remnant; eye remnant reduced to

narrowly-oval structure. Antenna relatively stout;

basal two segments cylindrical, third segment as

long but narrower and narrowing towards base,

next seven subequal but becoming progressively

shorter, apical segment a bit longer than

penultimate; each segment, except segment 1,

with some very small setae on inside apically (Fig.

7). Maxillary palpus elongate, apical segment a

little shorter than segments 1 to 3 combined,

oblique row of long setae on outer side, tip

truncated. Labial palpus elongate, apical two

segments subequal, tip weakly bifid, penultimate

segment with two setae near tip (Fig. 6).

Pronotum. Almost as wide as elytra;

anteriolateral 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 moderately marked, straight,

reaching to about halfway 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, sparsely

covered with very small punctures, row of widely

spaced larger punctures close to inner edge;

numerous setiferous micropunctures towards apex

and near base; 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 strongly

narrowed between coxae, not reaching

mesosternum, apical half narrow, almost parallel-

sided, strongly arched in lateral view with highest

point (viewed ventrally) between coxae.

Mesocoxae in contact at midline. Metasternum

sharply triangular in front in midline, wings very

narrow, broadly rounded in midline behind.

Metacoxal plates large, metacoxal lines

moderately widely spaced, reaching to about

halfway to metasternum, evenly diverging; a few

small setae-bearing punctures towards midline;

closely adpressed to first abdominal ventrite. First

and second ventrites fused, sutural lines distinct

towards midline, becoming indistinct laterally,

ventrites 3 to 5 mobile, sparsely covered with

small seta-bearing punctures, ventrites 3 and 4

with a long central seta or bunch of long setae.

Legs. Protibia relatively narrow, inner edge

102

1 2 3

Sess

Ca

5

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

FIGURES 1-7. Tjirtudessus raesideensis: 1, lateral view of central lobe of aedeagus; 2, ditto dorsal view; 3,

paramere; 4, mesotrochanter and mesofemur; 5, metatrochanter and metafemur; 6, labial palpus; 7, dorsal view.

Scale bar represents 1 mm.

straight, outer edge bowed, widest past middle

where it is about 3x its basal width; protarsi

expanded, first segment broad narrowing at base,

second segment a little narrower 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 1 to

3 with very dense covering of adhesive setae;

claws short and simple. Mesotrochanter elongate

with row of setae on inner edge; mesofemur with

row of eight to nine relatively weak setae along

hind edge in basal half (Fig. 4); mesotarsi similar

to protarsi. Metatrochanter tip rounded (Fig. 5);

metafemur elongate, lacking spines; metatibia

strongly curved, widening towards apex;

metatarsi elongate, basal segment longest, apical

segment much longer than fourth, segments 1

and 2 in combination about as long as others;

claws weak.

Male. Antennae a little stouter; pro- and

mesotarsi a little stouter. Median lobe of aedeagus

variable in width along shaft, tip bluntly pointed;

paramere broad, apical segment with long, narrow,

apical lobe well separated from rest of segment

(Figs 1-3).

Etymology

Named after the palaeodrainage system in

which it was collected.

Remarks

In size and in the relatively narrow pronotum,

T. raesideensis most nearly resembles T. hahni

but differs from this species in the more gracile

fore- and midlegs, thinner antennae, more

rounded tips to the metatrochanters and the

presence of a small eye remnant which is absent

in T. hahni. The aedeagi of the two species are

quite different. It is smaller than T. magnificus

with a narrower prothorax and much weaker pro-

and mesotarsi. Tjirtudessus eberhardi is a little

smaller with a broad pronotum and different

aedeagus.

NEW UNDERGROUND DYTISCIDAE, WESTERN AUSTRALIA 103

Nirridessus Watts and Humphreys, 1999

Nirridessus challaensis sp. nov.

(Figs 8-14)

Types

Holotype: m. ‘BES 8319, Nyung well, Challa

Station, 27°59'18"S, 118°31'03"E, 22/6/00, coll

W. F. Humphreys, S. Hinze’, slide mounted, in

spirit, WAM 27668.

Paratypes: 1, as for holotype, WAM 27669; 3,

‘BES 8346, Nyung well, Challa Station,

27°59'18"S, 118°31'03"E, 23/6/00, coll W. F.

Humphreys, S. Hinze’, 2 SAMA, 1 WAM 27671.

Description (number examined, 5)

Habitus. Length 2.3-2.5 mm; relatively flat,

moderately constricted at junction of pronotum/

elytra; uniformly light testaceous; hindwing

vestigial, about half length of elytron.

Head. Large, nearly as wide as elytra; smooth,

weakly reticulate, a few small scattered punctures

more numerous near antennal bases; subparallel

in posterior half; eye remnant reduced to small

12

triangular to oval area beneath head. Antenna

relatively stout; basal segment cylindrical, second

weakly ovoid, third segment about same length

but narrower at base, fourth slightly smaller, next

six subequal, apical segment a bit longer and

slightly narrower than penultimate; each segment,

except segments 1 and 4, with some very small

setae on inside apically (Fig. 14). Maxillary

palpus, elongate, apical segment large, about

length of segments 1 to 3 combined, an oblique

row of long setae on outer side near tip, tip

truncated. Labial palpus moderate, apical segment

longer that penultimate, tip weakly bifid,

penultimate segment with small papilla near tip

bearing two setae (Fig. 13).

Pronotum. Broad, as wide as elytra;

anteriolateral angles projecting strongly forward;

base moderately strongly narrowed, posterolateral

angles acute; smooth, with sparse, very weak

punctures more numerous towards the front

margin; basal plicae very weak, slanting inwards,

reaching to about halfway along pronotum,

slightly excavated inwards; with row of long setae

laterally, denser towards front.

la ‘R

( \)

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14 {

FIGURES 8-14. Nirridessus challaensis: 8, lateral view of central lobe of aedeagus; 9, ditto dorsal view; 10,

paramere; 11, mesotrochanter and mesofemur; 12, metatrochanter and metafemur; 13, labial palpus; 14, dorsal

view. Scale bar represents 1 mm.

104

Elytra. Not fused but tightly closed, lacking

inner ridges; elongate, widest behind middle,

smooth, sparsely covered with scattered punctures

larger laterally, row of small serial punctures in

centre of elytron; setiferous micropunctures in

basal fifth; a few scattered punctures with long

setae, more frequent towards sides and apex.

Epipleuron rather narrow in anterior fifth, then

rapidly narrowing to be virtually absent over rest

of elytron.

Ventral surface. Prothoracic process strongly

narrowed between coxae, not reaching

metasternum, apical half spatulate, strongly

arched in lateral view with highest point (viewed

ventrally) between coxae. Mesocoxae nearly in

contact at midline. Metasternum sharply

triangular in front in midline, wings very narrow,

broadly rounded in midline behind. Metacoxal

plates large, metacoxal lines weak, widely

spaced, reaching to about halfway to

metasternum, diverging in anterior two-thirds;

virtually impunctate except for a few towards

midline; closely adpressed to first abdominal

ventrite. First and second ventrites fused, sutural

lines distinct, ventrites 3 to 5 mobile, virtually

impunctate except for some small seta-bearing

punctures towards anterior edge of ventrites,

ventrites 3 and 4 with a long central seta or

bunch of long setae.

Legs. Protibia relatively narrow, edges straight,

widest near apex where it is about 3x its basal

width; protarsi moderately expanded, first

segment broadly oval, second segment as broad,

about one-third length of first, third segment as

long as first but 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 1 to 3

with dense covering of adhesive setae; claws short

and simple. Mesotrochanter oval with row of fine

setae on inner edge towards apex; mesofemur with

row of five to six relatively weak setae along hind

edge in basal half (Fig. 11); mesotarsi similar to

protarsi. Metatrochanter weakly pointed (Fig. 12);

metafemur elongate, lacking spines; metatibia

curved, widening towards apex; metatarsi

elongate, basal segment longest, apical segment a

little longer than fourth, segments 1 and 2 in

combination about as long as others; claws weak.

Male. Antennae a little stouter; pro- and

mesotarsi a little stouter. Median lobe of aedeagus

quite broad, weakly narrowing in apical quarter,

tip blunt; paramere broad, apical segment with

broad apical lobe folded over basal portion of

apical segment (Figs 8-10).

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

Etymology

Named after Challa Pastoral Station, the

location of the calcrete area in which it occurs.

Remarks

Nirridessus challaensis is a moderately large

species without a row of large punctures adjacent

to the suture. It can be recognised by the row of

five relatively weak (for Nirridessus) spines on

the mesofemur, the relatively broad pro- and

mesotarsi, and by the antenna which has

segments—including segments 3, 4 and the apical

one—of approximately the same length.

Nirridessus fridaywellensis sp. nov.

(Figs 15-21)

Types

Holotype: m. ‘BES 8379, Bore at Shearing

Quarters, Depot Springs Station,

27°55'50"S, 120°04'45"E, 26/6/00, coll W. F.

Humphreys, S. Hinze’ in spirit, WAM 27628.

Paratypes: 2, as for holotype, 1 SAMA, 1

WAM 27629, slide mounted; 4, ‘BES 8401, Site

425, Depot Springs, 28°03'00"S, 120°02'21"E,

28/6/00, coll W. F. Humphreys, S. Hinze’, 2

SAMA, 2 WAM 27630-27631; 3,. ‘BES 8406,

Friday Well, Depot Springs, 28°03'36"S,

120°04'03"E, 28/6/00, coll W. F. Humphreys, S.

Hinze’, 2 SAMA, 1 WAM 27632, slide mounted.

Additional specimen: 1, f., ‘BES 8374 Gums

Well Depot Spring’ SAMA. This specimen may

also belong to this species but a male is needed to

confirm the identification.

Description (number examined, 10)

Habitus. Length 1.7—1.8 mm; relatively flat,

strongly constricted at junction of pronotum/

elytra; uniformly light testaceous; hindwing

vestigial, about quarter length of elytron.

Head. Relatively small, narrower than elytra;

smooth, weakly reticulate, almost impunctate;

subparallel in posterior half; sides with dark

suture in middle near anterior edge. Antenna

relatively stout; basal segment cylindrical, second

large, oval, third segment almost as long, much

narrower, narrowing towards base, next six

subequal, apical segment about twice as long as

penultimate; each segment, other than 1 and 4,

with some very small setae on inside apically (Fig.

21). Maxillary palpus thin, elongate, apical

segment large, a little shorter than segments 1 to 3

combined, oblique row of long setae on outer side

near tip, tip truncated. Labial palpus moderate,

NEW UNDERGROUND DYTISCIDAE, WESTERN AUSTRALIA

19

105

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21

FIGURES 15-21. Nirridessus fridaywellensis: 15, lateral view of central lobe of aedeagus; 16, ditto dorsal view;

17, paramere; 18, mesotrochanter and mesofemur; 19, metatrochanter and metafemur; 20, labial palpus; 21, dorsal

view. Scale bar represents 1 mm.

apical segment longer than penultimate, tip

weakly bifid, penultimate segment with small

papilla near tip bearing two setae (Fig. 20).

Pronotum. Narrower than elytra; anteriolateral

angles projecting strongly forward; base strongly

narrowed, posterolateral angles acute; smooth,

with very sparse, very weak punctures more

numerous towards the front margin; moderately

reticulate; basal plicae absent; with row of long

setae laterally, denser towards front.

Elytra. Not fused, lacking inner ridges;

elongate, widest behind middle, smooth, evenly

covered with sparse very small punctures, central

row of serial punctures hard to trace; some

setiferous micropunctures near apex and along

sutural line, very sparse or lacking at base; row of

long setae near lateral edge, a few additional

larger punctures with long setae, more frequent

towards sides; moderately reticulate. Epipleuron

broad in anterior fifth, then rapidly narrowing to

be virtually absent over rest of elytron.

Ventral surface. Prothoracic process strongly

narrowed between coxae, not reaching

metasternum, apical half spatulate, strongly arched

in lateral view with highest point (viewed

ventrally) between coxae. Mesocoxae in contact at

midline. Metasternum sharply triangular in front

in midline, wings very narrow, broadly rounded in

midline behind. Metacoxal plates large, metacoxal

lines absent; virtually impunctate; moderately

reticulate; closely adpressed to first abdominal

ventrite. First and second ventrites fused, sutural

lines distinct, ventrites 3 to 5 mobile, virtually

impunctate except for ventrites 3 and 4 with a

long central seta or bunch of long setae.

Legs. Protibia relatively narrow, edges straight,

evenly diverging, widest at apex where it is about

4x its basal width; protarsi little expanded, first

segment oval, second segment as wide as about

one-half length of first, third segment as long as

first and as wide, deeply bifid, fourth segment

very small and hidden within lobes of third

106

segment, apical segment narrow, cylindrical,

about twice length of third, segments 1 to 3 with a

few adhesive setae; claws short and simple.

Mesotrochanter elongate oval with a few fine

setae on inner edge; mesofemur with three strong